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29
Bulletin of the
British Museum (Natural History)
Entomology series Vol 41 1980
British Museum (Natural History)
cLondon 1981
Dates of publication of the parts
No 1 28 August 1980
No 2 25 September 1980
No 3 .... 30 October 1980
No 4 18 December 1980
No 5 18 December 1980
ISSN 0524-6431
Printed in Great Britain by The Whitefriars Press Ltd, London and Tonbridge
^
GENERAL
2
Contents
Entomology Volume 41
Page
No 1 A revision of the western Palaearctic species of the genus Meteorus
(Hymenoptera : Braconidae)
T. Huddleston 1
No 2 A revision of the Old World Polymorphanisini (Trichoptera :
Hydropsychidae)
P. C. Barnard 59
No 3 A review of the genera of Neotropical Encyrtidae (Hymenoptera:
Chalcidoidea)
John S. Noyes 107
No 4 A revision of the British species of Dendrocerus Ratzeburg
(Hymenoptera: Ceraphronoidea) with a review of their biology as
aphid hyperparasites
N. D. M. Fergusson 255
No 5 A revision of the Pacific species of Conocephalus Thunberg
(Orthoptera : Tettigoniidae)
Linda M. Pitkin 315
<
GENERAL
-20CTI980
Bulletin of the \^C£
British Museum (Natural History)
A revision of the western Palaearctic
species of the genus Meteorus
(Hymenoptera : Braconidae)
T. Huddleston
Entomology series Vol 41 No 1 28 August 1980
The Bulletin of the British Museum (Natural History), instituted in 1949, is issued in four
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World List abbreviation : Bull. Br. Mus. nat. Hist. (Ent.)
Trustees of the British Museum (Natural History), 1980
ISSN 0524-6431 Entomology series
Vol 41 No 1 pp 1-58
British Museum (Natural History)
Cromwell Road
London SW7 5BD Issued 28 August 1980
A revision of the western Palaearctic species of the\ t,8*
genus Meteorus (Hymenoptera: Braconidae)
T. Huddleston
• - k <
Department of Entomology, British Museum (Natural History), Cromwell Road, London SW7
5BD
Contents
Synopsis ............. 1
Introduction ............ 1
Depositories ............ 2
Taxonomic history . . . . . . . . . . .2
Taxonomic characters ........... 4
Biology ............. 6
List of host records ........... 7
Meteorus Haliday ........... 9
Synonymic list of species . . . . . . . . . .10
Key to species (females) . . . . . . . . . .11
Descriptions of species .......... 20
Nominadubia ............ 54
Acknowledgements . . . . . . . . . . .55
References ............. 55
Index .57
Synopsis
The species of the genus Meteorus from the western Palaearctic zoogeographical region are revised, keyed
and illustrated. Thirty-six of the 121 nominal species discussed are considered to be valid and 3 are new.
Seventy-one names are placed in synonymy, 44 for the first time. Seventy-seven primary types have been
examined and of these 35 lectotypes are herein designated. Fourteen species are considered nomina dubia.
About 3500 specimens have been examined.
Introduction
The taxonomy of the parasitic Hymenoptera is the least known of any large group of insects and,
until recent years, the Braconidae have received even less attention than most other groups of
Parasitica. There is therefore a pressing need for revisionary work in most parts of the family.
Moreover this is no mere academic exercise; many species of Braconidae use as hosts insects which
are important economically as pests of crops. The need for programmed 'biological' control is
already becoming more urgent as the effectiveness of chemical methods of pest control declines.
The use of parasitoid insects in the control of pest species has so far met with only sporadic
success. For biological control to be effective an intimate knowledge of the biology of the insects
involved is essential and this in turn is founded upon an accurate knowledge of their taxonomy.
This paper is concerned with the taxonomy of the western Palaearctic species of Meteorus, all of
which appear to be primary parasites of the larvae of either Lepidoptera or Coleoptera.
It was originally intended to include in this revision the species of the whole of the Palaearctic
region. No material was available for study from the U.S.S.R. or from China, however, nor was
type-material available of the species described from the U.S.S.R. (M. baicalensis Telenga, M.
flagellatus Alexeev, M. politutele Shenefelt [replacement name for politus Telenga, not
Provancher], M.pospelowi Telenga). These areas have therefore been omitted from consideration.
Bull.Br.Mus.nat.Hist.(Ent.)41(l): 1-58 Issued 28 August, 1980
2 T. HUDDLESTON
All other areas of the Palaearctic region have been included but the bulk of the material available
for study came from NW. Europe.
MNHU, Berlin
NMB, Berne
IRSNB, Brussels
HNHM, Budapest
NMI, Dublin
IP, Eberswalde
RNH, Leiden
BMNH
ZI, Lund
NMV, Melbourne
ZSBS, Munich
EH, Munich
MZ, Naples
CNC, Ottawa
UM, Oxford
MNHN, Paris
ZC, Plovdiv
MSC, Reading
USDA, Sevres
NR, Stockholm
HC, Stockholm
MZS, Turin
UDE, Uppsala
NM, Vienna
IFF, Vienna
AU, Wageningen
USNM, Washington.
MAS, Yatabe
Depositories
Museum fur Naturkunde der Humboldt-Universitat, Berlin.
Naturhistorisches Museum Bern, Berne.
Institut Royal des Sciences Naturelles de Belgique, Brussels.
Hungarian Natural History Museum, Budapest.
National Museum of Ireland, Dublin.
Institut fur Pflanzenschutzforschung, Eberswalde.
Rijksmuseum van Natuurlijke Historic, Leiden.
British Museum (Natural History), London.
Zoologiska Institution, Lund.
National Museum of Victoria, Melbourne.
Zoologische Sammlung des Bayerischen Staates, Munich.
E. Haeselbarth collection, Munich.
Museo Zoologia, Naples.
Canadian National Collection, Biosystematics Research Institute, Ottawa.
University Museum, Oxford.
Museum National d'Histoire Naturelle, Paris.
A. Zaykov collection, Plovdiv.
M.R. Shaw collection, Reading.
USDA, European Parasite Laboratory, Sevres.
Naturhistoriska Riksmuseet, Stockholm.
K.-J. Hedqvist collection, Stockholm.
Museo ed Istituto di Zoologia Sistematica, Universita di Torino, Turin.
University Department of Entomology, Uppsala.
Naturhistorisches Museum, Vienna.
Institut fiir Forstentomologie und Forstschutz, Vienna.
Agricultural University, Wageningen.
[U.S. National Museum] National Museum of Natural History, Washington, D.C.
Plant Disease Herbarium and Insect Museum, National Institute of Agricultural
Sciences, Yatabe.
Taxonomic history
Meteorus Haliday (1835: 24) was first used as a subgenus of Perilitus Nees von Esenbeck, as it
was then defined, to accommodate those species having three submarginal (cubital) cells in the
forewing. Haliday dealt with 17 species of Meteorus of which only the 13 comprising his 'section B'
are now considered to belong in Meteorus s.str.; the remainder are placed in the genus Zele Curtis
[Zemiotes Foerster]. In section B, Haliday described seven species as new to science. Haliday's
collection is now housed in the NMI, Dublin where A. W. Stelfox in 1948 collated the Meteorus
and transferred them to their present cabinet from the boxes in which Haliday had left them.
Stelfox, in his unpublished notes on the collection, stated 'no specimen bears the name of one of his
[Haliday's] own species, though several carry his Ms labels with Wesmael's names; also two have
his Ms labels bearing "nomina nuda" or possibly "pet names" which are nowhere referred to in his
Ms lists. He has therefore left us no "types"!'. Furthermore, the four species which were least
adequately described (delator, jaculator, profligator and vexator) are no longer present in the
collection; the interpretation of these species must therefore rest entirely upon their descriptions.
Nees von Esenbeck (1811) described five species of 'Bracon' which he later transferred to section
II of his genus Perilitus (corresponding to Meteorus s.l.), adding descriptions of four more species.
Nees' collection was destroyed during the 1939-45 war (Griffiths, 1964: 829) and there is
therefore no original material of his species. In most cases, however, there are long-established and
generally accepted usages of Nees' names and the identification of his species causes but little
difficulty. It has not therefore been considered necessary to designate neotypes.
REVISION OF THE WESTERN PALAEARCTIC METEOR US 3
Wesmael (1835) described 16 species ot'Meteorus in the genus Perilitus. Wesmael's collection is
well-preserved in Brussels and the identity of the species he described can therefore be established
beyond doubt. Some slight difficulty arises in distinguishing valid syntype specimens from amongst
Wesmael's material because the number of specimens labelled by Wesmael as belonging to a
particular species is generally more than the number of specimens of that species which he recorded
as being in his possession at the time of the description. Wesmael stated, for instance, that he
possessed one female ofqffinis but three specimens in the collection are so labelled by him. Again, a
specimen (abdominator) labelled fragilis in Wesmael's handwriting has a second label bearing the
date 1838 and could not therefore have been part of the original material of fragilis. The determina-
tion label is indistinguishable, however, from those on the rest of the series labelled fragilis which
includes other misidentified specimens. I suspect therefore that Wesmael labelled his collection long
after the publication of the descriptions of his species, probably after 1838, and that he included
subsequently captured material which he had not examined with the meticulous care displayed in
his original publication. I do not consider it at all likely that Wesmael had before him mixed series
when preparing his descriptions because these are far too precise to have been based on badly
sorted material.
Ratzeburg (1844; 1848; 1852) described (in the genus Perilitus) nine new species of Meteorus.
Ratzeburg's descriptions are deficient even by the standards of his contemporaries and so some of
his species are impossible to interpret without types. Ratzeburg's collection was largely destroyed
during the 1939-45 war and Konigsmann (1964) has shown that the remnant of the collection
which survives contains specimens which can be accepted as type-material of only two of the
species of Meteorus which Ratzeburg described.
Ruthe (1862) described 20 new species of Meteorus in a posthumous paper which was usefully
annotated by Reinhard. I have been able to examine all of Ruthe's original material most of which
is housed in the BMNH. Ruthe placed too much emphasis on minor differences in colour and
consequently only one of his species remains valid.
Marshall (1887; 1891; 1898; 1899) gave a comprehensive interpretation of the work of Haliday
and Wesmael and to a lesser extent of other continental workers. He deprived himself, however, of
the use of many valuable characters by his method of mounting his specimens face down on
squares of card. Marshall described two species of Meteorus as new, only one of which is presently
considered valid, and brought forward a description of a third (Capron in Marshall, 1887).
Thomson (1895), in typically idiosyncratic fashion, put forward 10 new names in Meteorus of
which five were re-interpretations of names used by Haliday or by Ruthe and five proposed as
undoubtedly new to science. Despite his procedural eccentricities Thomson's work was superior to
that of his predecessors (save only Wesmael) and, indeed, to that of most of his successors.
Thomson's collection is well preserved in Lund and I have had the opportunity of examining the
material. Four of Thomson's species remain valid. The presence on a pin of a species label in
Thomson's handwriting should not be taken as indicating that that specimen was so labelled by
Thomson. Labels which were Thomson's drawer labels were transferred to the specimen which stood
first in the series above the label when the collection was moved from Thomson's cabinets into more
modern cabinets (R. Danielsson, pers. comm.).
Schmiedeknecht (1897) published a key to the European species of Meteorus incorporating
descriptions of three new species. Schmiedeknecht appears to have based his work almost entirely
upon earlier descriptions and this several times led him into misinterpretation. One of his species
remains valid.
Morley (1908) reiterated much of Marshall's work on the British species of the genus; his
contribution to the subject was insignificant.
Lyle (1914) published some useful data on the biology of some species of Meteorus; this work
needs to be carefully interpreted because Lyle's identification of the parasite species was sometimes
inaccurate. I have been able to examine Lyle's collection, now housed in the BMNH, but his
material is deprived of much of its value by Lyle's method of mounting insects on large squares of
thick card. Lyle described one species which I have placed in synonymy in this paper.
Muesebeck (1923) monographed the Nearctic species of Meteorus s.l. and emphasized the value
of several important characters which had not been fully exploited in previous work on the genus.
4 T. HUDDLESTON
Muesebeck (1939; 1954) described two Palaearctic species of Meteorus, one of which is now
considered valid.
Fischer (1956-70) has contributed much to the literature on the genus and though he has mis-
interpreted some of the older species he has established some useful synonymy and has designated
a neotype for M. vexator.
Tobias (1976) has keyed the species of Meteorus which are represented in the fauna of the
Caucasus.
Nineteenth-century hymenopterists were handicapped in their understanding of the limits of
species by their practice of working with relatively few specimens of each species. Many so-called
species were founded upon one or two specimens which differed but little from already-described
taxa. Then too, the reliance on colour differences to distinguish species has been a fertile source of
misinterpretation; colour in Meteorus is the least stable characteristic and in no case is colour alone
sufficient to distinguish species in the absence of more stable morphological differences. There have
been only two keys to major parts of the Palaearctic Meteorus fauna (Marshall, 1891;
Schmiedeknecht, 1897). Both of these works contained a plethora of putative species because of the
authors' acceptance of differences in colour as indicating different species. Also both largely
derived their concepts of species from published descriptions which were often imprecise or
misleading. I have based my interpretation of the species upon the original material wherever
possible together with much other material of the species involved and I have arrived at my concept
of these species in the light of the characters discussed below.
Taxonomic characters
The antennal segments vary slightly within a species in number, shape and relative length, but
nevertheless provide characteristics of considerable usefulness if the possibilities of variation are
borne in mind. The antennae exhibit marked sexual dimorphism in most species; within a given
species those of the female are almost always shorter than those of the male, not only because they
have fewer segments on average but also because the segments are themselves shorter.
The shape of the head is generally extremely stable within a species. The shape of the temples as
seen from above, the lateral profile of the face, the frons and the vertex and the shape in face view
of the vertex, genae and clypeus often furnish diagnostic characters of great value. The length of the
malar space and the breadth of the face in relation to its height are also of considerable
significance; both are related largely to the size and disposition of the eyes.
The degree of convergence of the eyes is an extremely useful character for the differentiation of
species in Meteorus. Its full significance does not appear to have been noticed by previous workers.
When the distance between the tops of the eyes (at their point of intersection with the apparent
'edge' of the head) is twice or more the breadth of the face (the minimum distance between the eyes)
I have called the eyes 'strongly convergent' (Fig. 4), otherwise 'not strongly convergent'. More
subtle differences in convergence will only be appreciated with the aid of the various figures. For
the proper appreciation of this character the head must be examined directly from in front so that
the bases of the hind ocelli are upon the 'horizon'. This character is subject to only minor variation
in most species though there is considerable sexual dimorphism, the eyes of the male usually being
much less convergent than those of the female. The size of the eyes is also often of considerable
importance in distinguishing species, particularly their length in dorsal view in relation to the length
of the temple.
The size of the ocelli varies slightly within a species but is nonetheless of considerable importance
at species-group level. Closely related species generally have ocelli of approximately the same size
so that this character is of limited use at species level. The comparative size of the ocelli is
expressed as the greatest diameter of a hind ocellus compared with its least distance from the
border of the nearest eye. In the text this is expressed in the form OO (ocellar-ocular distance) =OD
(ocellar diameter). I have used the term 'small ocelli' where the ocellar-ocular distance is at least 2-5
times ocellar diameters, and the term 'large ocelli' where the ocellar-ocular distance is at most 2-0
times ocellar diameters.
REVISION OF THE WESTERN PALAEARCTIC METEOR US 5
The amount of twist of the mandibles and their size is constant within a species. The mandibles
of different species, however, show considerable differences which are related to other
characteristics of the ocelli, the tarsal claws and the head shape as expressed in couplet one of the
key to species. The mandibles of some species are small, delicate and strongly twisted so that the
lower tooth is behind the upper when viewed from in front, i.e. at right angles to the base of the
mandible (Fig. 38). Mandibles of this type tend to be constricted in the middle. Other species have
mandibles which are not twisted (i.e. the teeth are in the same plane as the base of the mandible)
and these are long and broad and not markedly constricted in the middle (Figs 1 2, 29). Species do
occur in which the mandibles are intermediate in type between these two extremes but nonetheless
the mandibular characteristics are most useful in the differentiation of species and do not appear to
have been fully exploited by previous workers.
The presence or absence of paired pits in the dorsal surface of tergite one is a useful character for
the separation of groups of species in Meteorus. Some care is needed for the correct evaluation of
this character; in several species without pits there are in some specimens convolutions of the
sculpture on the dorsal surface of tergite one which give the appearance of pits at first glance.
Where pits are present they are distinct even if small and the ventral borders of the tergite are never
joined beneath. In nearly all species which do not have pits in the dorsal surface of tergite one the
ventral borders of the tergite are in contact to a greater or lesser extent. Muesebeck (1923) brought
to notice the fundamental importance of the degree of contiguity of the ventral borders of tergite
one as a character for the separation of species in Meteorus. Nixon (1941) made use of this
character in his review of the African species of Meteorus but it has been largely ignored in more
recent work on the genus. There is little or no variation in this structure at the infraspecific level in
those species lacking pits and it is therefore a stable and most useful character. M. pulchricornis, as
I have defined that species, is exceptional in that most specimens have no dorsal pits in tergite one
and the ventral borders of the tergite are joined while a few specimens do have true pits and, in
these, the ventral borders of the tergite are narrowly separated. Some care is required in tracing the
borders of tergite one; their course is not always easy to discern because the lateral faces of the
tergite often bear longitudinal striations which can be mistaken for the edge of the tergite. Then too,
the sternite at the base of segment one of the gaster is sometimes heavily chitinized and sculptured
and this can be confused with the ventral part of the tergite on a superficial examination.
Ovipositor length varies within a species but the range of variation is well defined and fairly small
and ovipositor length is therefore a useful character for the differentiation of species in Meteorus.
Previous workers have generally expressed ovipositor length in relation to the length of the gaster.
This is not entirely satisfactory because the gaster is a telescopic structure which can therefore vary
in length appreciably within a species. I have expressed ovipositor length as a multiple of the length
of tergite one of the gaster. The ovipositor is measured from the base to the tip and not merely the
portion of it which, in dorsal view, can be seen projecting beyond the end of the gaster - the
exserted part. In many specimens the ovipositor base is concealed by the overlapping terminal
tergites of the gaster and so I have taken the apex of the hypopygium as providing the most reliable
indication of the position of the ovipositor base. Apart from the differences in length between
species the ovipositor shows few structural features which can be used in the differentiation of
species and the hypopygium none.
The presence or absence of lobes on the tarsal claws is a useful character for the separation of
groups of species in Meteorus (Figs 34, 35, 58). This character does not appear to have been
noticed by previous workers. There is little variation in claw shape within a species but differences
in claw shape between closely related species though often significant are generally rather too subtle
to be of use in a key. When examining tarsal claws care must be taken that they are seen in full
lateral view, the presence of even a large lobe may otherwise be overlooked.
The venation in Meteorus is extremely labile and therefore furnishes far fewer diagnostic features
than has often been supposed.
In general the sculptural characteristics of Meteorus exhibit a rather wide degree of infraspecific
variation. In most species, however, the sculpture of the propodeum and of the sternaulus seem to
be rather more stable than that elsewhere and can be of considerable help in distinguishing species.
The sculpture of the hind coxa is also often of considerable value in diagnosis of species; in those
6 T. HUDDLESTON
species with strongly rugose hind coxae it is rare to find specimens in which the hind coxae are
smooth and in those which typically have a smooth, punctate hind coxa it is unusual to find
individuals whose coxae bear rugosity and very rarely is this strongly developed. Even where the
sculpture varies there tends to be a characteristic pattern of sculptural elements which can often aid
in forming a 'picture' of a species. Tergite one, for instance, tends to have a characteristic pattern in
each species though specimens occur more or less frequently in which the usually predominant
sculptural element is superseded by that which is normally subsidiary.
Size differences in Meteorus are often of little taxonomic significance. The size of a parasitoid
insect is governed to a certain extent by the size of the host in which it develops. Insects belonging
to a species which uses a variety of hosts therefore often differ considerably in their body size. A
host species which itself exhibits considerable size differences affects its parasites in the same way.
The morphological terms used in this paper are defined in Richards (1977).
Biology
In general little detailed work has been done on the biology of Meteorus and, naturally, most of the
attention which has been given to the subject has been focused on the species of potential
usefulness in biological control programmes. Tobias (1976) and Marsh (1979) cite the relevant
literature. The work of Madel (1963) on the morphology and biology of colon (asfragilis) is an ex-
ceptionally scholarly piece of work and should serve as a model for such studies. Achtelig (1974)
gives a fine study of the male genital anatomy in oculatus (as pachypus).
Most species of Meteorus are solitary parasites but M. rubens and M . heliophilus are gregarious
(Marshall, 1887; Lyle, 1914; Tobias, 1976). The mature larvae of some species spin a cocoon
suspended by a thread and it is from this habit that the name of the genus is derived. Among the
exceptions to this are the gregarious species, which spin their cocoons together in loose heaps, and
several species which parasitize wood-boring beetles and which, as far as is known, form stalkless
cocoons within the beetles' burrows (Mason, 1973). Lyle (1914) suggested that species of Meteorus
overwinter as eggs or as immature larvae within the body of the host or as mature larvae within the
cocoon; on the evidence so far the last alternative seems the least likely. Rearing data suggest that
at least some species of Meteorus overwinter in different host species from those which they use in
their summer generations. There is no evidence that any species of the genus overwinters as an
adult though the possibility remains. Some species of Meteorus attack Coleoptera whilst others
attack Lepidoptera; there is no evidence that any species of the genus utilizes both groups as hosts.
In general, the species which parasitize Coleoptera are those with small eyes and ocelli, simple
tarsal claws and stout mandibles though two species, cespitator and micropterus, have these
characteristics but are parasitic on larvae of Lepidoptera. The Coleoptera from which Meteorus
have been recorded are all species in which the larvae live in concealment, boring in wood, in bark
or in fungus. The relatively small eyes and ocelli of these parasites is presumably a reflection of
their use of concealed hosts and the consequent necessity for them to rely on non-visual means of
detection. M. cespitator and micropterus have been recorded from hosts the larvae of which are
often at least partially concealed. The species of Meteorus which parasitize Lepidoptera generally
have large eyes and ocelli, lobed tarsal claws and slender, strongly twisted mandibles. M . consimilis
has some of these characteristics, however, and has been recorded (in the literature) only from
Scolytus multistriatus (Marsham). The majority of the Lepidoptera from which Meteorus species
have been recorded are those with larvae which do not live in concealment though tortricid larvae
are generally partly concealed and most noctuid larvae expose themselves only after dark. The
larvae of some species of Lepidoptera rely on camouflage for their protection, those of Them
variata (Denis & Schiffermiiller), for instance, are exceptionally well camouflaged and only
Meteorus lionotus, which has exceptionally large eyes and ocelli, has been recorded as parasitizing
.hem. The species of Meteorus which parasitize Lepidoptera therefore have large eyes and ocelli,
Dresumably because they are able to use sight in locating their hosts.
M. micropterus, consimilis and abdominator are similar in appearance; they are dark, heavily
culptured insects with short, dark wings and short, thick legs and antennae. The morphological
REVISION OF THE WESTERN PALAEARCTIC METEOR US 7
similarities of these species probably indicate that they have a common mode of host-searching
rather than indicating any close phylogenetic affinity. M. micropterus has been recorded from
Lepidoptera the larvae of which live at the base of dense herbage such as grass tussocks; it
probably hunts on foot and its stout build will enable it the better to push its way into suitable host-
habitats. No doubt consimilis also reaches its host by pushing through crevices in bark.
M. corax, sulcatus and nixoni form a distinctive group of species of which corax and sulcatus
are known to parasitize the larvae of cerambycids. The three species have in common a stout basal
abdominal segment and a long ovipositor; this is presumably because they need to oviposit through
wood.
M. affinis has been recorded from hosts belonging to families of Lepidoptera which are not
closely related taxonomically . The larvae of both host-species, however, have a similar mode of life
feeding at the base of moss on walls or on tree-stumps.
I have examined specimens of ictericus and obsoletus bred only from tortricids. Both species are
rather small and slender and have long ovipositors the better to reach their host larvae which are
typically loosely concealed in rolled leaves, catkins, buds and so on. There is a single record of
ictericus (as lophyriphagus) having been bred from a sawfly. I believe this to have been a fortuitous
occurrence because of the abnormal abundance of the larvae of that sawfly at that particular time,
or an erroneous host-identification.
In the literature on Meteorus there are many records of host-parasite relationships. Much of this
information is of doubtful value because of the uncertainty about the accuracy of the identification
of the parasites owing to the lack of reliable keys for their determination. With few exceptions I
have cited host data only from the specimens I have examined. In order to confirm all records in
the literature it would be necessary to redetermine the material on which those records were based,
a quite impracticable task. As far as possible I have put into their present combinations the host
names which I have cited. This information is summarized in the following list.
List of host records
HOSTS SPECIES OF Meteorus
COLEOPTERA
BIPHYLLIDAE
Biphyllus lunatus (Fabricius) vexator
SCOLYTIDAE
Scolytus multistriatus (Marsham) consimilis
? Scolytus triarmatus Eggers obfuscatus
CERAMBYCIDAE
Monochamus saltuarius Gebler corax
Monochamus sartor (Fabricius) corax
Semanotus undatus (L.) sulcatus
Molorchus umbellatarum (von Schreber) sulcatus
MELANDRYIDAE
Orchesia micans (Panzer) longicaudis, obfuscatus
TENEBRIONIDAE
Eledenoprius armatus Panzer longicaudis
Corticeus longulus (Gyllenhal) punctifrons
CISIDAE
Cis boleti (Scopoli) profligator
LEPIDOPTERA
HEPIALIDAE
Hepialus humuli (L.) micropterus
Hepialusfusconebulosa (De Geer) micropterus
ZYGAENIDAE
Zygaena filipendulae (L.) unicolor
T. HUDDLESTON
HOSTS
TINEIDAE
Infurcitinea argentimaculella (Stainton)
Nemapogon granella (L.)
? Monopis rusticella (Hiibner)
Trichophaga tapetzella (L.)
GELECHIIDAE
Pectinophora gossypiella (Saunders)
TORTRICIDAE
Archips podana (Scopoli)
Epiphyas postvittana (Walker)
Adoxophyes orana (Fischer von Roslerstamm)
Tortrix viridana (L.)
Croesia bergmanniana (L.)
Acleris hastiana (L.)
Epinotia sordidana (Hiibner)
Gypsonoma dealbana (Frolich)
PYRALIDAE
Scoparia sp.
Ephestia cautella (Walker)
LYCAENIDAE
Thecla betulae (L.)
NYMPHALIDAE
Ladoga Camilla (L.)
Cynthia cardui (L.)
Charaxes jasius jasius (L.)
SATYRIDAE
Maniola jurtina (L.)
LASIOCAMPIDAE
Poecilocampa populi (L.)
Malacosoma neustria (L.)
Lasiocampa quercus (L.)
Macrothylacia rubi (L.)
Dendrolimus pint (L.)
GEOMETRIDAE
Idaea muricata (Hufnagel)
Eulithis testata (L.)
Thera obeliscata (Hiibner)
Thera variata (Denis & Schiffermiiller)
Thera juniper ata (L.)
Operophtera brumata (L.)
Eupithecia nanata (Hiibner)
A nticollix sparsata (L.)
Apeira syringaria (L.)
Ennomos quercinaria (Hufnagel)
Agriopis leucophaearia (Denis & Schiffermiiller)
Agriopis aurantiaria (Hiibner)
THAUMETOPOEIDAE
Thaumetopoeia processionea (L.)
LYMANTRIIDAE
Dasychira pudibunda (L.)
Euproctis chrysorrhoea (L.)
Leucoma salicis (L.)
Lymantria dispar (L.)
ARCTIIDAE
Setina irrorella (L.)
NOLIDAE
Nola cuculatella (L.)
Si'FCi
ajfinis
cespitator
cespitator
cespitator
graciliventris
ictericus
ictericus
ictericus
ictericus, obsoletus
ictericus
ictericus
ictericus
obsoletus
affinis
graciliventris
pulchricornis
colon
rubens
pulchricornis
versicolor
pulchricornis
versicolor
versicolor
versicolor
versicolor
rubens
versicolor
lionotus
lionotus
lionotus
pulchricornis
pulchricornis
colon
melanostictus
melanostictus
pulchricornis
pulchricornis
versicolor
versicolor
versicolor
versicolor
pulchricornis
abscissus
colon, pulchricornis
REVISION OF THE WESTERN PALAEARCTIC METEORUS
HOSTS
NOCTUIDAE
Agrotis exclamationis (L.)
Agrotis ipsilon (Hufnagel)
Noctua pronuba (L.)
Noctuafimbriata (Schreber)
Lycophotia porphyrea (Denis & Schiffermiiller)
Diarsia brunnea (Denis & SchifTermiiller)
Xestia triangulum (Hufnagel)
Xestia xanthographa (Denis & Schiffermiiller)
A narta myrtilli (L.)
Polia nebulosa (Hufnagel)
Mamestra brassicae (L.)
Lacanobia oleracea (L.)
Orthosia stabilis (Denis & SchifTermiiller)
Mythimna sp.
Cleoceris viminalis (Fabricius)
Lithophane ornitopus (Hufnagel)
Eupsilia transversa (Hufnagel)
Agrochola lota (Clerck)
Thalpophila matura (Hufnagel)
Euplexia lucipara (L.)
Ipimorpha retusa (L.)
Cosmia trapezina (L.)
Spodoptera exigua (Hiibner)
Bena prasinana (L.)
HYMENOPTERA
DIPRIONIDAE
Neodiprion sertifer (Geoffrey in Fourcroy)
rubens
rubens
rubens
heliophilus
gyrator, pulchricornis, versicolor
gyrator
heliophilus
gyrator
versicolor
heliophilus
rubens
gyrator
heliophilus
gyrator
gyrator
heliophilus
pulchricornis
gyrator
gyrator
gyrator
gyrator
gyrator
rubens
colon
ictericus
METEORUS Haliday
Meteorus Haliday, 1835: 24. Type-species: Meteorus pendulator (Latreille), by subsequent designation
(Haliday in Westwood, 1839: 61, Synopsis).
Saprotichus Holmgren, 1868: 430. Type-species: Saprotichus chinensis Holmgren, by subsequent designation
(Viereck, 1914:130).
DIAGNOSIS. Meteorus is a Euphorine (as defined, for instance, by van Achterberg, 1974) which can
be differentiated by the following combination of characters.
Notaulices always present, generally distinct; first segment of gaster narrow at base and broadened apically,
never cylindrical, the spiracles at midpoint of segment or slightly behind, occasionally in front but never at base;
hair rows on tergites restricted to a single subapical row; ovipositor always exserted; forewing with three
submarginal cells, the second generally higher than long (Fig. 13), generally quadrangular, often slightly
narrowed anteriorly and rarely triangular, never petiolate; radial cell of hindwing narrowed to apex or parallel-
sided never apically broadened or divided.
Meteorus has been recorded from all zoogeographical regions.
Meteorus was placed in a subfamily, Meteorinae, distinct from though related to the Euphorinae
until Muesebeck (1923: 4) pointed out the artificial nature of this division and (1935: 5) placed
Meteorus within the Euphorinae. This placement of the genus has been generally accepted by
workers on Braconidae and no modern authority on the group retains Meteorus as distinct at more
than tribal level. Support for this placement of the genus has come from the work of Capek (1970) on
larval morphology and biology, and from that of van Achterberg (1974; 1979) and Tobias (1966;
1 976) on adult morphology.
Mason (1973) lifted from synonymy the genus Zemiotes, previously regarded as a part of
Meteorus, and postulated a relationship between Zemiotes and Zele. I have accepted Mason's
division of Meteorus s.l. into two genera but have followed van Achterberg's (1979) placement of the
10
T. HUDDLESTON
two genera together in the Meteorini. Further, van Achterberg (1979) has shown that the name Zele
is properly applied to the species hitherto placed in Zemiotes. He has lifted from synonymy the name
Homolobus Foerster to accommodate the species traditionally placed in Zele. The two genera in the
Meteorini may be distinguished as follows.
Radial cell of hindwing narrowed to apex or parallel-sided, never divided, the radial vein not angled; hairs on
tergites restricted to single apical rows; tarsal claws sometimes without a basal lobe; tergite one sometimes
without dorsal pits .... . METEORUS UaMay
Radial cell of hindwing widened to apex, sometimes divided, radial vein always angled; hairs on tergites not
in single rows, covering at least apical half of tergite; tarsal claws always strongly bent and with large
basal lobe; tergite one always with dorsal pits ZELE Curtis
Synonymic list of species
abdominator (Nees)
brunnipes Ruthe syn. n.
brevipesalis Shenefelt syn. n.
abscissas Thomson
affinis (Wesmael)
punctiventris Ruthe syn. n.
ruthei Schmiedeknecht syn. n.
voloscensis Fischer syn. n.
Brevicauda Thomson
thuringiacus Schmiedeknecht syn. n.
mongolicus Fahringer syn. n.
cespitator (Thunberg)
atrator (Curtis)
similator (Nees)
ambiguus Ruthe syn. n.
cinctellus (Spinola)
fuscipes (Wesmael) syn. n.
tenellus Marshall syn. n.
colon (Haliday)
fragilis (Wesmael) syn. n.
fasciatus (Ratzeburg)
luridus Ruthe syn. n.
consimilis (Nees)
brevipes (Wesmael)
albicornis Ruthe
corax Marshall
monochami Fischer
eadyi sp. n.
filator Haliday
laticeps (Wesmael) syn. n.
hodisensis Fischer syn. n.
graciliventris Muesebeck
gyrator (Thunberg)
scutellator (Nees)
parvulus Thomson syn. n.
heliophilus Fischer
hirsutipes sp. n.
ictericus (Nees)
minutus (Thunberg)
ephippium (Curtis)
xanthomelus (Wesmael)
confinis Ruthe syn. n.
fallax Ruthe syn. n.
pleuralis Ruthe syn. n.
liquis Ruthe syn. n.
consors Ruthe
crassicrus Thomson syn. n.
lophyriphagus Fahringer syn. n.
adoxophyesi Minamikawa syn. n.
jaculator (Haliday)
obscurellus Ruthe
tenuicornis Thomson syn. n.
lionotus Thomson
ruficoloratus Fischer syn. n.
longicaudis (Ratzeburg)
melanostictus C apron
niger Lyle syn. n.
micropterus (Haliday)
nixoni sp. n.
obfuscatus (Nees)
thoracicus (Curtis)
formosus (Wesmael)
fodori Papp syn. n.
obsoletus (Wesmael)
viridanae Johansson syn. n.
oculatus Ruthe
pachypus Schmiedeknecht syn. n.
pallipes (Wesmael)
nigritarsus Ruthe syn. n.
profligator (Haliday)
pulchricornis (Wesmael)
striatus Thomson syn. n.
thomsoni Marshall
japonicus Ashmead
nipponensis Viereck
macedonicus Fischer syn. n.
graeffei Fischer
tuberculifer Fischer syn. n.
puncttfrons Thomson
rubens (Nees)
leviventris (Wesmael)
islandicus Ruthe
medianus Ruthe
scutatus Costa syn. n.
heteroneurus Thomson
szechuanensis Fahringer syn. n.
mesopotamicus Fischer syn. n.
REVISION OF THE WESTERN PALAEARCTIC METEOR US 1 1
sattcorniae Schmiedeknecht vexator (Haliday)
ocellatus Watanabe syn. n.
sulcatus Szephgeti .,
instgnis Muesebeck °
Fischer syn. n. N
(Ratzeburg)
/oscfcto Ruthe
j. tj r> *u
cfaZw/s Ruthe syn. n. .,./> t
, . „. , gracilis (Ratzeburg)
penmen Fischer syn. n. . . ,„ . s
. , ,„, n ' longicorms (Ratzeburg)
umcolor (Wesmael) ,f . . ,K T
, . . /T ¥ i \ palhdus (Nees)
chmensis (Holmgren) syn. n. , ... ,
• i /«/ i\ pendulator (Latreille)
versicolor (Wesmael) ^ , . \
,. , , /,., n rubnceps (Ratzeburg)
bimaculatus(Wesmad) « %x\ ,
, . . /T; . ruficeps (Nees)
brevicorms (Ratzeburg) % *L /- \
, , B . ru^is (De Geer)
decoloratus Ruthe v^. '
. , tl, . stenostigma Thomson
camp tolomae Watanabe syn. n. ,./r> . x
., . „. , wesmaeu (Boie)
ikonomovi Fischer syn. n.
hartigi Shenefelt syn. n.
Key to species (females)
I am unable to give a key to males. In Meteorus, as in many parasitic Hymenoptera, males of closely related
species are often impossible to separate. Also there are several species of which I have seen no males.
1 Ocelli small, OO (ocellar-ocular distance) at least 2 times OD (ocellar diameter), usually
more; tarsal claws simple (Figs 34, 35), without a distinct basal lobe but sometimes
swollen at the base or with a small tubercle on the inner side of the claw; mandibles large,
stout and not strongly twisted (Figs 12, 29); generally head full and rounded behind the
eyes and in front view rather square, the face broad and the eyes small (but
cf. Figs 2, 4, 5) ........ . . 2
Ocelli large, OO at most 2 times OD (except in cinctellus); tarsal claws with a well-developed
lobe (Fig. 58) (except in consimilis and in rubens); mandibles most usually small and
strongly twisted (Fig. 38); generally head contracted behind eyes and in front view rather
triangular in outline, being more or less contracted ventrally and the eyes
large .............. 19
2(1) Dorsal pits absent, the ventral borders of tergite one joined from near the base of the
segment to about its midpoint (Fig. 7) ........ 3
Dorsal pits present, the ventral borders of tergite one not touching, usually widely
separated ............. 5
3 (2) Flagellar segments obconical, mostly only as long as broad, only the first segment as much
as twice as long as broad; frons flattened and face strongly protuberant, forming a 'shelf
upon which the antennae are mounted (Fig. 1); propodeum strongly reticulate-
rugose and without distinct carinae .... micropterus (Haliday) (p. 39)
At least the basal flagellar segments cylindrical and not less than 3 times as long as broad;
face not strongly protuberant; propodeum only weakly rugose and with distinct carinae 4
4 (3) Antennae short, 21- to 24-segmented, the three preapical segments only slightly longer than
broad, submoniliform; basal segments of flagellum lighter in colour than apical segments;
eyes large, convergent (Fig. 2); ovipositor about 2-5 times length of tergite
one ........... filator (Haliday) (p. 30)
Antennae longer, 26- to 27-segmented, all segments at least twice as long as broad,
cylindrical; basal segments of the flagellum not distinctly lighter in colour than apical
segments; eyes smaller, very little convergent (Fig. 3); ovipositor about 1-5 times
length of tergite one ......... eadyi sp. n. (p. 29)
5 (2) Eyes strongly convergent, face at most as wide as high (Figs 4, 5) .... 6
Eyes not strongly convergent, face broader than high, usually about twice as broad as high
6 (5) Hind tibia strongly swollen, about as wide as femur (Fig. 6); tergite one longitudinally striate,
sometimes rugose medially; antennae 30- to 33-segmented; eye about 3 times as
long as temple ......... oculatus Ruthe (p. 42)
12
T. HUDDLESTON
Figs 1-9 1, Meteorus micropterus (Haliday), head, lateral view. 2, M.filator (Haliday), head, frontal
view. 3, M. eadyi sp. n., head, frontal view. 4, M. vexator (Haliday), head, frontal view. 5, 6, M.
oculatus Ruthe; (5) head, frontal view; (6) hind leg. 7, M.filator (Haliday), segment one of gaster,
ventral view. 8, M. nixoni sp. n., head, dorsal view. 9, M. sulcatus Szepligeti, head, dorsal view. All
figures x 43.
REVISION OF THE WESTERN PALAEARCTIC METEOR US 1 3
Hind tibia not strongly swollen, distinctly narrower than femur; tergite one reticulate-rugose,
occasionally with a longitudinal striate element laterally; antennae 20- to 24-
segmented; eye about twice length of temple . . . . vexator(Haliday)(p. 53)
7 (5) Tergite one stout, broad at base and with spiracles distinctly before the middle, dorsal pits
large (Fig. 11); tergite two usually strongly sculptured and with distinct thyridiae; hind
tibia usually swollen and basally lighter in colour; ovipositor always long, at least 3
times length of tergite one .......... 8
Tergite one slender at base and with spiracles at about mid-segment, dorsal pits smaller;
tergite two smooth, never strongly sculptured and thyridiae obsolescent; hind tibia
rarely swollen or basally lighter in colour; ovipositor sometimes distinctly less than
three times length of tergite one . . . . . . . . . 10
8 (7) Temples shorter than eyes (Fig. 8); clypeus wide, mandibles large, not twisted (Fig. 12)
nixoni sp. n. (p. 40)
Temples longer than eyes (Figs 9, 10); clypeus narrow, mandibles short, strongly twisted 9
9 (8) Propodeum rather flat in profile, not carinate, rugose; clypeus smooth-punctate; second
submarginal cell at least 1-5 times as high as wide (Fig. 13); antennae 31- to 36-
segmented ......... sulcatus Szepligeti (p. 49)
Propodeum divided by a strong medial transverse carina into dorsal and posterior planes
which are sharply angled one to the other; clypeus vertically carunculate; second
submarginal cell about as high as wide (Fig. 14); antennae 39- to 4 1-segmented
corax Marshall (p. 28)
10 (8) Ovipositor long, at least 3-0 times length of tergite one; sternaulus reticulate-rugose or
reticulate-foveolate, at least anteriorly; hind coxa generally rugose . . . 11
Ovipositor shorter, at most 2-5 times length of tergite one, if, rarely, slightly more
(salicorniae) then frons depressed and face strongly protuberant (Fig. 20); sternaulus
narrow, foveolate or, if broad and reticulate-rugose (abdominator), wings strongly
infumate; hind coxa generally smooth, punctate . . . . . . 13
11 (10) Head subcubic, temples longer than eyes (Fig. 15); tergite one predominantly coarsely
rugose with only a few striae laterally; a small species 2-5-3-0 mm in length
jaculator (Haliday) (p. 36)
Head more transverse, temples shorter than eyes or at most equal (Fig. 16); tergite one
striate and with a reticulate medial longitudinal band; generally larger species 4-5 mm
in length 12
12 (11) Eyes large, convergent (Fig. 17); malar space shorter than basal breadth of mandible, face
about 1-5 times as broad as high; antennae short, at most 27-segmented; face generally
almost completely rugulose, rarely completely punctate . cespitator (Thunberg) (p. 24)
Eyes smaller, less strongly convergent (Fig. 18); malar space about equal to basal breadth of
mandible; face about twice as broad as high; antennae longer, 29- to 32-segmented; face
densely punctate with at most a trace of rugulose sculpture beneath the base of the
antenna ........ ./ongicai«/is(Ratzeburg)(p. 38)
13 (10) Sternaulus broad and strongly reticulate-rugose, at least anteriorly; pronotum strongly
rugose; wings infumate, the second submarginal cell strongly narrowed anteriorly; ocelli
minute, OO = 3 • 5—4-0 times OD and head transverse (Fig. 1 9) abdominator (Nees) (p. 20)
Sternaulus narrow, foveolate; pronotum generally mostly smooth; wings hyaline, the second
submarginal cell only slightly narrowed anteriorly; ocelli usually larger, if minute
then head not distinctly transverse . . . . . . . . . 14
14 (13) Face strongly convex medially, frons depressed, occiput distinctly concave (Fig. 20)
propodeum strongly reticulate-rugose with no distinctly differentiated carinae
salicorniae Schmiedeknecht(p. 48)
Face evenly and only weakly convex, frons not depressed, occiput at most weakly
concave; propodeum with scattered rugae and with distinct carinae . . . 15
15(14) Clypeus narrower than face (Figs 22, 23, 24); mandibles at least slightly twisted . 16
Clypeus as broad as the face (Figs 25, 26); mandibles not twisted . . . . 18
16 (15) Antennae short, the segments in the apical half of the flagellum not longer than broad;
vertex rather flat in front view (Fig. 22); eyes larger; clypeus protuberant
obfiiscatus(Nees)(p. 41)
Antennae longer, at most one or two segments of flagellum not longer than broad; vertex
rounded in front view (Figs 23, 24); eyes smaller; clypeus rather flat . . . 17
14
Figs 10-19 10, 11, Meteorus corax Marshall; (10) head, dorsal view; (11) segment one of gaster, dorsal
view. 12, M. nixoni sp. n., mandible. 13, M. sulcatus Szepligeti, 2nd submarginal cell of forewing. 14,
M. corax Marshall, 2nd submarginal cell of forewing. 15, M.jaculator (Haliday), head, dorsal view.
16, 17, M. cespitator (Thunberg); (16) head, dorsal view; (17) head, frontal view. 18, M. longicaudis
(Ratzeburg), head, frontal view. 19, M. abdominator (Nees), head, dorsal view. All figures x 43.
29
Figs 20-29 20, Meteorus salicorniae Schmiedeknecht, head, dorsal view. 21, M. abdominator (Nees),
head, frontal view. 22, M. obfuscatus (Nees), head, frontal view. 23. M. punctifrons Thomson, head,
frontal view. 24, M. profligator (Haliday), head, frontal view. 25, M. hirsutipes sp. n., head, frontal
view. 26, 27, M. brevicauda Thomson; (26) head, frontal view; (27) head, dorsal view. 28, 29, M.
hirsutipes sp. n.; (28) head, dorsal view; (29) mandible. All figures x 43 except Fig. 29 x 107.
16 T.HUDDLESTON
17 (16) Frons between anterior ocellus and eye with a patch of large distinct punctures separated by
less than their own diameter; height of eye at most equal to breadth of face (Fig. 23); legs
conspicuously long, hind tibia and tarsus equal in length to thorax and gaster;
ovipositor short, down-curved ..... puncttfrons Thomson (p. 46)
Frons with smaller punctures separated by much more than their own diameter; height of
eye distinctly greater than breadth of face (Fig. 24); legs not conspicuously long, hind tibia
and tarsus together distinctly shorter than thorax and gaster; ovipositor slightly
longer, straight profligator (HaMay) (p. 44)
18 (15) Temples about as long as eyes, less strongly contracted (Fig. 27); ovipositor 1-5 times as
long as tergite one; legs short, stout; tarsal claws shorter, thicker (Fig. 34)
brevicauda Thomson (p. 23)
Temples distinctly shorter than eyes, strongly contracted (Fig. 28); ovipositor 2-5 times
length of tergite one; legs long, slender; tarsal claws very long, slender (Fig. 35)
hirsutipes sp. n. (p. 33)
19 (1) Dorsal pits absent or, occasionally, weakly indicated, the ventral borders of tergite one
touching or at most with a narrow gap between them ...... 20
Dorsal pits distinct, the ventral borders of tergite one not touching, usually widely
separated ............. 27
20 (19) Tergite one not joined beneath at the base of the segment and either joining shortly in
front of the middle of the segment or a small gap remaining (Figs. 39, 40) . . 21
Tergite joined beneath at the base of the segment (Fig. 41) 26
21 (20) Eyes strongly convergent; ocelli small, OO about 2- 5 times OD . cinctellus (Spinola) (p. 25)
Eyes not strongly convergent; ocelli larger, OO at most 2-0 times? OD , ' . . . 22
22 (21) Antennae short, 24- to 28-segmented; tarsal claws not strongly curved and with a very
small basal lobe or with none; mandible fairly large and moderately twisted
miens (Nees) (p. 47)
Antennae longer, usually wi.th 30 or more segments; tarsal claws strongly curved and with
a pronounced basal lobe; mandible small, strongly twisted "'.' . . . . 23
23 (22) Head strongly, narrowed behind eyes (Figs. 31, 33); length of eye 2-5-3-0 times length of
temple .* . ' '. '• ; . . 24
Head less strongly narrowed behind eyes (Figs; 36, 37); length of eye at most twice length
of temple ............. 25
24 (23) Ocelli very large, OO = 0-5 times OD (Fig. 31); head more strongly contracted behind
eyes .......... .lionotus Thomson (p. 37)
Ocelli not so large, OO= 1-0-1-5 times OD (Fig. 33); head less strongly contracted
behind eyes heliophilus Fischer (p. 32)
25 (23) Clypeus evenly and densely punctate with a dense pile of erect hairs; frons not tuberculate;
propodeum in lateral view rounded, strongly rugose dorsally and with no distinct
carinae; recurrent antefurcal or interstitial . . pulchricornis (Wesmael) (p. 45)
Clypeus with scattered punctures and a few scattered long hairs; frons with a blunt tubercle
in front of the fore ocellus (Fig. 44); propodeum in lateral view flattened and often with a
central longitudinal carina; recurrent always distinctly postfurcal (Fig. 45)
colon (Haliday) (p. 26)
26 (20) Eyes strongly protuberant (Fig. 43), large, temple generally strongly contracted, not
rounded; ovipositor thick, strongly constricted just before apex, generally short, 1 -5 times
length of tergite one, occasionally twice; propodeum short, wide, strongly
excavate medially versicolor (Wesmael) (p. 51)
Eyes not strongly protuberant (Fig. 42), smaller, temple not strongly contracted, rounded;
ovipositor slender, not strongly constricted at apex, generally longer, 2-5 times length of
tergite one but sometimes only twice; propodeum not conspicuously short and
wide and only weakly excavate medially .... obsoletus (Wesmael) (p. 41)
27 (19) Eyes strongly convergent or, if only moderately convergent (Fig. 52) then length of eye in
dorsal view 2-5-3-0 times length of temple; ovipositor long, always at least 2-5 times
length of tergite one 28
Eyes at most moderately convergent (Fig. 56); length of eye at most twice length of temple
(Figs 55, 57); ovipositor not more than twice length of tergite one ... 32
28 (27) Length of eye 4 times length of temple (Fig. 46); OO= 1-25-1-5 times OD; eyes strongly
convergent (Fig. 47) . . . . ". graciliventris Muesebeck (p. 31)
38
41
Figs 30-41 30, 31, Meteorus lionotus Thomson; (30) head, frontal view; (31) head, dorsal view. 32, 33,
M. heliophilus Fischer; (32) head, frontal view; (33) head, dorsal view. 34, M. brevicauda Thomson,
tarsal claws. 35, M. hirsutipes sp. n., tarsal claws. 36, M.pulchricornis (Wesmael), head, dorsal view.
37, M. rubens (Nees), head, dorsal view. 38, M. lionotus Thomson, mandible. 39, M. cinctellus
(Spinola), segment one of gaster, ventral view. 40, M. lionotus Thomson, segment one of gaster,
ventral view. 41, M. versicolor (Wesmael), segment one of gaster, ventral view. All figures x 43 except
Figs 34, 35 and 38 x 107.
18
Figs 42-53 42, Meteorus obsoletus (Wesmael), head, dorsal view. 43, M. versicolor (Wesmael), head,
dorsal view. 44, 45, M. colon (Haliday); (44) head, lateral view; (45) 2nd submarginal cell of forewing.
46, 47, M. graciliventris Muesebeck; (46) head, dorsal view; (47) head, frontal view. 48, M. tabidus
(Wesmael), head, dorsal view. 49, M. qffinis (Wesmael), head, dorsal view. 50, 51, M. pallipes
(Wesmael); (50) head, frontal view; (51) head, dorsal view. 52, 53, M. ictericus (Nees); (52) head,
frontal view; (53) head, dorsal view. All figures x 43.
REVISION OF THE WESTERN PALAEARCTIC METEORUS
19
Length of eye at most 3 times length of temple; if OO less than 2-0 times OD then eyes
only moderately convergent .......... 29
29 (28) Outer surface of hind coxa strongly rugose or reticulate-rugose; anteriorly the mesopleuron
beneath the sternaulus reticulate-rugose or reticulate-foveolate; tergite one
often rugose, at least centrally ......... 30
Outer surface of hind coxa smooth with at most some puncturation; sternaulus foveolate,
the sculpture often partly obsolescent so that the sternaulus is in part a smooth furrow;
mesopleuron smooth-punctate with no reticulate sculpture beneath the sternaulus;
tergite one longitudinally striate, usually without rugosity . . . . . 31
30 (29) Head in dorsal view (Fig. 48) contracted behind the eyes, not strongly rounded, the occiput
not concave; vertex always with a light yellow patch on the orbit opposite the hind ocellus;
mesopleuron beneath the sternaulus reticulate-rugose so that the sternaulus appears to be
a broad reticulate-rugose band; pronotum laterally strongly reticulate-
rugose ......... tabidus (Wesmael)(p. 50)
Head in dorsal view (Fig. 49) not contracted behind eyes, strongly rounded, the occiput
concave; vertex rarely pale-marked; mesopleuron beneath the sternaulus rarely reticulate-
rugose though often reticulate-foveolate anteriorly; pronotum laterally smooth
sometimes rugose, rarely reticulate ..... . q^Z/i/s(Wesmael)(p- 22)
31 (29) Eyes strongly convergent (Fig. 50); temples rounded, not strongly contracted (Fig. 51);
ocelli smaller OO = 2-0 times OD pa/flpes (Wesmael) (p. 43)
Eyes moderately convergent (Fig. 52); temples less rounded, strongly contracted (Fig. 53);
ocelli larger OO= 1-0-1 -5 times OD fcter/cas (Nees) (p. 34)
32 (27) Sternaulus broad, reticulate-rugose or, if narrow foveolate then length of malar space about
twice basal breadth of mandible; propodeum coarsely reticulate-rugose without a distinct
central longitudinal carina and frons not tuberculate; hind coxa generally
strongly rugose 33
54
58
Figs 54-58 54, 55, Meteorus consimilis (Nees); (54) head, frontal view; (55) head, dorsal view. 56, M.
gyrator (Thunberg), head, frontal view. 57, M. melanostictus Capron, head, dorsal view. 58, M.
ictericus (Nees), tarsal claw. All figures x 43 except Fig. 58 x 107.
20 T. HUDDLESTON
Sternaulus narrow foveolate and length of malar space not more than basal breadth of
mandible; propodeum weakly rugose and generally with a distinct central longitudinal
carina or the frons has a blunt tubercle in front of fore ocellus; hind coxa generally
smooth, punctate ........... 35
33 (32) Malar space long, almost twice basal breadth of mandible (Fig. 54); ovipositor at most
equal in length to tergite one, down-curved; second submarginal cell most usually
strongly narrowed anteriorly ;flagellum pale banded . . . consimtlts (Nees) (p. 27)
Malar space shorter, not more than basal breadth of mandible; ovipositor 1-5-2-0 times
length of tergite one, straight; second submarginal cell only slightly narrowed anteriorly;
basal segments of flagellum sometimes paler than apical segments but never
with a distinct pale band .......... 34
34 (33) Clypeus rather flat, finely and densely punctured and with a dense erect pile; hind coxa
almost completely rugose; pterostigma dark with a pale outer border
abscissas Thomson (p. 21)
Clypeus protuberant, more coarsely punctate-rugulose and with scattered long hairs; hind
coxa rugose only laterally; pterostigma uniformly pale . unicolor (Wesmael) (p. 51)
35 (32) Antenna longer, 30- to 34-segmented; frons with no tubercle but often with a weakly
impressed line in front of fore ocellus; propodeum finely rugose, often reticulate-rugose,
generally with a central longitudinal carina which, though weak, is distinct; medial
transverse carina sometimes present but never strongly raised gyrator (Thunberg) (p. 31)
Antenna shorter, 25- to 27-segmented; frons with a blunt tubercle in front of the fore ocellus
(Fig. 57); propodeum more coarsely rugose, generally with no central
longitudinal carina but with a strongly raised medial transverse carina
melanostictus C apron (p. 38)
Descriptions of species
Meteorus abdominator (Nees von Esenbeck)
(Figs 19, 21)
Perilitus abdominator Nees von Esenbeck, 1 8 1 1 : 24. Syntypes $ 9» GERMANY (destroyed).
[Perilitus brevipes Wesmael; Wesmael, 1838: 141. Misidentification: see brevipesalis below.]
Perilitus brunnipes Ruthe, 1862: 37. LECTOTYPE 9, GERMANY: Ruthe coll. (BMNH), here designated
[ examined j.Syn.n.
Meteorus brevipesalis Shenefelt, 1969: 55. Holotype 9> BELGIUM: Brussels, coll. Wesmael (IRSNB, Brussels)
[examined]. [Proposed as a new name for 'brevipes Wesmael, 1838'.] Syn. n.
I have based my interpretation of abdominator on specimens in Wesmael's collection labelled
'Perilitus abdominator NVE' in Wesmael's handwriting. Wesmael was the first revisor of
abdominator.
Wesmael (1835) described brevipes solely from males but later (1838) he published a description
of a single female which he considered to belong to brevipes. As pointed out by Marshall (1887:
110) this led Ruthe (1862: 34) to describe the true female of brevipes as albicornis though he
expressed doubt about the correctness of his action. There is one female specimen in Wesmael's
collection which agrees with Wesmael's description of the supposed female of brevipes and which
bears a label 'Perilitus brevipes mihi' in Wesmael's handwriting. This specimen comes within the
limits of variation of abdominator.
Ruthe (1862: 38) stated that his description of brunnipes was based on six female specimens. In
the collection of the BMNH there are six specimens from Ruthe's collection standing above the
name 'brunnipes', two of them bearing labels 'P. brunnipes M.' in Ruthe's handwriting. These
specimens are conspecific and agree with Ruthe's description of brunnipes. They come within the
limits of variation of abdominator. I have chosen as lectotype the specimen labelled 'BM Type
Hym. 3C750'. Fischer (1970Z>: 275) suppressed brunnipes as a synonym of Zemiotes deceptor
(Wesmael). The latter species has recently been synonymized with Z. albiditarsus (Curtis) (van
Achterberg, 1979). Van Achterberg (in litt.) states that in the MNHU, Berlin there are specimens of
albiditarsus identified as brunnipes. It seems probable that Fischer's interpretation of brunnipes
was based on these wrongly-identified specimens.
REVISION OF THE WESTERN PALAEARCTIC METEOR US 2 1
9- Antennae short, 22- 25-segmented, distal segments of flagellum only slightly longer than broad. Head
slightly broader than thorax; distinctly transverse; temples contracted but not strongly so. Ocelli small,
OO = 3-5-4-0 times OD. Eyes generally only slightly convergent, never strongly so. Face always transverse,
about twice as broad as high; protuberant; generally rather smooth-punctate. Malar space slightly shorter
than basal breadth of mandible. Tentorial pits deep. Clypeus strongly protuberant, divided from face by a
deep distinct furrow, smooth with relatively few large punctures. Mandible stout, twisted. Frons depressed,
usually completely smooth-punctulate but occasionally with a small patch of rugosity just above the antennal
sockets and also sometimes with scattered larger punctures along the margin of the eye. Vertex polished,
punctulate. Genae swollen so that the head is rather broad ventrally in front view (Fig. 21). Pronotum
expanded forwards and sideways, strongly rugose, sometimes reticulate-rugose except for a smooth polished
band dorsally. Mesonotum polished, punctate, the middle lobe often strongly so; notaulices deep, foveolate.
Sternaulus broad, reticulate-rugose with particularly strong vertical rugae; rest of mesopleurae smooth-
punctulate except for the subalar prominence. Propodeum coarsely rugose, often reticulate-rugose, at least in
part; longitudinal and transverse carinae present but often obscured by the rugosity; distinctly excavate
medio-posteriorly. Tergite one stout, with distinct dorsal pits; longitudinally striate dorsally, the lateral
surfaces rugose at the base and sometimes also striate. Sternite of segment one thickened in its basal half and
finely reticulate-rugose. Ovipositor 2-0-2-5 times length of tergite one; down-curved. Legs stout; hind coxae
smooth-punctate, sometimes rugose dorsally but this rugosity often weak. Tarsal claws not strongly bent but
swollen at the base. Wings short; second submarginal cell generally more or less strongly contracted
anteriorly.
Colour black; legs, tegulae and usually tergite 2 and tergite 3 at base testaceous; basal flagellar segments
often lighter in colour than rest, sometimes yellow. In darker specimens the legs and the tergites are darker
brownish testaceous and the flagellar segments are all dark. Hind legs often darker at the apex of the femur,
tibia and tarsus. Wings always at least slightly infumate, often densely so.
(j\ Same as female except antennae longer, 25- to 31-segmented, all flagellar segments generally at least
twice as long as broad; colour often darker.
MATERIAL EXAMINED
143 , , 71 ;\ Belgium, France, Germany, Great Britain, Hungary, Ireland, Italy, Netherlands, Poland,
Sweden, Switzerland.
HOSTS. No reared material examined.
REMARKS. M. abdominator is superficially similar to micropterus and consimilis; this question is touched
upon in the discussion of micropterus. M. abdominator is rather variable in sculptural characteristics and in
venation.
Meteorus abscissas Thomson
[Meteorus pulchricornis Wesmael; Ruthe, 1862: 40. Misidentification.]
Meteorus abscissus Thomson, 1895: 2156. LECTOTYPE 9, SWEDEN (ZI, Lund), here designated
[examined].
Ruthe stated that he possessed one specimen of this species which he referred to pulchricornis
though he expressed some doubt as to the correctness of his identification. There is a specimen
present in the collection of the BMNH bearing a label 'P.pulchricornis Wsm' in Ruthe's
handwriting which fits the description given by Ruthe of that species. Thomson (1895: 2156) stated
that Ruthe's pulchricornis differed from that of Wesmael and put forward a new name, abscissus,
for the former species. There are six specimens in Thomson's collection standing above the name
'abscissus' and these are conspecific with Ruthe's specimen of 'pulchricornis' and agree precisely
with what Thomson wrote about abscissus.
9- Antennae 30- to 33-segmented, thick at base tapering to apex, all flagellar segments distinctly longer than
broad. Head broad, temples strongly contracted behind eyes, eyes at most twice length of temple Ocelli large,
OO = 2 times OD. Eyes large, moderately convergent. Malar space about equal to basal breadth of mandible.
Face 1-5 times as wide as high, not strongly protuberant, with weak transverse rugae though sometimes
smooth, punctate laterally. Clypeus not strongly protuberant, sometimes flattened, densely and regularly
punctate and with a dense erect pile. Mandibles small, strongly twisted. Pronotum laterally strongly
reticulate-rugose; notaulices deep-foveolate, coalescing posteriorly to form a strongly rugose area. Sternaulus
broad, reticulate-rugose, rest of mesopleuron smooth-punctate except dorsally where it is reticulate-rugose.
Prepectal carina strongly raised. Propodeum broad, strongly convex, strongly reticulate-rugose dorsally and
22 T. HUDDLESTON
posteriorly. Tergite one long with distinct dorsal pits and shallow glymmae; dorsal surface finely striate;
lateral surface at the base smooth, shining. Ovipositor 1-5-2-0 times length of tergite one, straight. Hind
coxa strongly rugose, reticulate-rugose dorsally and sometimes laterally, often transversely rugose ventrally.
Tarsal claws with a distinct basal lobe.
Colour brownish testaceous, legs testaceous; propodeum and tergite one sometimes black wholly or in
part; thorax, head and hind coxa also sometimes black-marked.
tf. Same as 9 except that antenna longer, 34- to 36-segmented; tergite one narrower; sculpture of hind
coxa sometimes obsolescent.
MATERIAL EXAMINED
9 9, 4 c?. Austria: 1 tf, Sudtirol, Partschins, 20.vii.1966 (HaeselbartK) (EH, Munich). Hungary: 1 9,
Budapest, Harmashatarhegy, 6.vi.l972 (Papp) (HNHM, Budapest); 1 9, Harsbokorhegy, 15.viii.1952
(Bajari) (HNHM, Budapest). Ireland: 1 <•?, Co. Clare, Fanore, 10.vi.1974 (West), ex larva of S. irrorella
(BMNH). Italy: 2 <$, Riva s. Garda (HaeselbartK) (EH, Munich). Sweden: 1 $, Sk., Horna, 5.ix.l967
(Hedqvist) (HC, Stockholm); 6 9, Thomson coll. (ZI, Lund).
HOST. Setina irrorella (L.) (Lepidoptera: Arctiidae).
REMARKS. M. abscissus is structurally very close to pulchricornis but has a flatter clypeus and the
pronotum, sternaulus, propodeum and hind coxa are more strongly rugose; it is also a rather
stouter insect. M. pulchricornis generally has no true dorsal pits on tergite one but there is likely to
be considerable difficulty in the separation of those specimens of pulchricornis with pits and
specimens of abscissus.
Meteorus affinis ( Wesmael )
(Fig. 49)
Perilitus affinis Wesmael, 1835: 31. Holotype 9, BELGIUM: Brussells, coll. Wesmael (IRSNB Brussels)
[examined].
Meteorus punctiventris Ruthe, 1862: 25. Holotype 9, GERMANY: Ruthe collection (BMNH) [examined].
Syn. n.
Meteorus gracilis Ruthe, 1862: 31. Holotype 9> GERMANY: Ruthe coll. (BMNH) [examined]. [Junior
primary homonym of gracilis Ratzeburg, 1852: see ruthei below.]
Meteorus ruthei Schmiedeknecht, 1897: 205. [Replacement name for gracilis Ruthe, 1862.] Syn. n.
Meteorus voloscensis Fischer, 1959: 14. Holotype 9, YUGOSLAVIA: Volosca, coll. Graeffe (NM, Vienna)
[examined]. Syn. n.
Wesmael (1835: 31) stated that he possessed only a single female of the species. In Wesmael's
collection there are three specimens labelled 'Perilitus affinis mihi' in Wesmael's handwriting. Only
one of these specimens, which I consider to be the holotype, fits the description of affinis, therefore
Fischer (1970a: 52) has unnecessarily designated this specimen as lectotype of affinis.
Ruthe (1862: 27) stated that he possessed but one female specimen of punctiventris. In Ruthe's
collection there is one female labelled 'P. punctiventris m.' in Ruthe's handwriting; this specimen
agrees with Ruthe's description and I therefore accept it as the holotype of punctiventris.
Ruthe (1862: 31) stated that he possessed only a single female of gracilis. This specimen now
stands in the collection of the BMNH; it is in rather poor condition, being broken and embedded in
a mass of glue. The specimen bears a label 'P. gracilis m' in Ruthe's handwriting and agrees with
his description of that species. I regard this specimen as the holotype of gracilis; it is within the
limits of variation of affinis.
Fischer (1959: 14) postulated that his new species voloscensis was related to vexator and cited
differences in venation to support the differentiation of the two species. The large ocelli and lobed
tarsal claws of voloscensis show that it is not closely related to vexator, it does, however, come well
within the limits of variation of affinis.
9- Antennae 27- to 30-segmented, the penultimate five or so segments only about 1 -5 times as long as broad.
Head strongly rounded behind eyes, occiput rather concave, length of eye at least twice length of temple.
Ocelli large, OO= 1-5—2-0 times OD. Eyes large, protuberant, strongly convergent. Face about as broad as
high, not protuberant, rugulose. Clypeus smooth, shining with scattered punctures, not strongly protuberant.
Malar space short, much less than basal breadth of mandible. Mandible stout, moderately twisted. Pronotum
REVISION OF THE WESTERN PALAEARCTIC METEOR US 23
smooth laterally except for a median band of strong vertical rugae, sometimes rugose ventrolaterally, rarely
reticulate-rugose. Sternaulus foveolate, sometimes smooth posteriorly and, usually, anteriorly coalescing with
a patch of reticulate-foveolate sculpture on the mesopleuron beneath the sternaulus, occasionally there is
reticulate-rugose sculpture in this area and rarely the sculpture there is obsolete (this is characteristic of small
specimens); rest of mesopleurae shining punctulate except for a subalar patch which is reticulate-foveolate.
Propodeum rugose, the carinae not usually distinctly differentiated from the rugae, transverse carinae the
most well-developed. Tergite one narrow, about twice as long as apically broad, with distinct dorsal pits;
dorsal surface striate laterally and with a medial longitudinal band of reticulate sculpture; the lateral surface
of tergite one rugose anteriorly and striate posteriorly. Ovipositor long, 3-0-3-5 times length of tergite one,
slender. Legs long, slender; hind coxae rugose ventrolaterally; tarsal claws with small but distinct lobe.
Colour black; legs yellow except sometimes hind coxa black and tarsi infuscated; clypeus, mandibles,
prothorax centrally and tergite two often lighter in colour, sometimes yellow; head and mesothorax
occasionally also lighter in colour.
<$. Same as 9 except antennae longer, 30- to 33-segmented; eyes smaller and not strongly convergent;
temples sometimes longer and less contracted; sculpture of sternaulus and of hind coxa often obsolescent.
MATERIAL EXAMINED
46 9> 17 $. Bulgaria, Finland, France, Great Britain, Ireland, Netherlands, Sicily.
HOSTS. Infurcitinea argentimaculella (Stainton) (Lepidoptera: Tineidae) larvae feeding on moss on
wall. I have also examined a single male of this species reared from a larva feeding at the base of
moss on a dead tree stump and identified as either Gelechia sp. (Lepidoptera: Gelechiidae) or
Scoparia sp. (Lepidoptera: Pyralidae). Marshall (1887: 108) recorded Scoparia as the host of this
species (as punctiventris).
The specimens reared from Infurcitinea argentimaculella (5 9> 2 ^ England, Kent, Faversham,
9.vii.l977 (Bradford) (MSC, Reading)) are considerably smaller than average specimens ofaffinis.
If Marshall's record of Scoparia as the usual host of affinis is correct then the smaller size of the
parasites from Infurcitinea corresponds to the difference in size of the two hosts.
REMARKS. The smaller specimens of qffinis are difficult to separate from vexator. They have 24- to
26-segmented antennae, sternaulus and hind coxal sculpture obsolescent; in these characteristics
and in size they are close to vexator but they have the lobed claws and larger ocelli ofaffinis. The
reduction in antennae and in sculpture is no doubt a result of their reduction in size due to their
choice of a small host species.
Meteorus brevicauda Thomson
(Figs 26, 27, 34)
Meteorus brevicauda Thomson, 1895: 2165. LECTOTYPE 9, SWEDEN (ZI, Lund), here designated
[examined].
Meteorus thuringiacus Schmiedeknecht, 1897: 190. Syntypes 9, GERMANY (lost). Syn. n.
Meteorus mongolicus Fahringer, 1935: 10. Holotype 9» CHINA: S. Kansu (Hummel) (NR, Stockholm)
[examined]. Syn. n.
In Thomson's collection there are three conspecific specimens standing above the name brevicauda.
I have selected as lectotype of this species the specimen labelled by Fischer as holotype. The
lectotype has no locality label, the second specimen is labelled 'Satrop 3.8.81, the third 'Hbg'
(= H&lsingborg).
Thomson (1896: 2165) tentatively suggested that brevicauda and profligator might be the same
species. Schmiedeknecht (1897: 299) placed brevicauda in unquestioned synonymy with profligator
from which it was removed, in my view correctly, by Fischer (1959: 75). Having so disposed of
brevicauda, Schmiedeknecht was misled into describing that species as thuringiacus. The
description of thuringiacus agrees precisely with brevicauda and with no other species and I have
no doubt that the two are conspecific, although I have been unable to locate original material of
thuringiacus. I have examined one specimen labelled 'Meteorus thuringiacus Schmiedek.' in
Schmiedeknecht's handwriting but this specimen is a male and therefore could not have been part
of the syntype-series which was stated to be all female.
The holotype of mongolicus is well within the limits of variation of brevicauda.
24 T. HUDDLESTON
9. Antennae 24- to 27-segmented, all segments at least slightly longer than broad. Head rounded behind eyes
but not strongly contracted; length of temple about equal to length of eye in dorsal view. Ocelli small, OO = 3
times OD. Eyes not strongly protuberant, converging slightly. Malar space very short, distinctly less than
half basal breadth of mandible. Face about 2 times as broad as high, rather flat, laterally punctate, medially
reticulate-punctate with a trace of rugulose sculpture. Clypeus large, as wide as face, flat, polished and with
scattered large punctures. Tentorial pits large. Mandibles large, stout, not twisted. Pronotum projects in front
of mesonotum forming a 'neck', laterally smooth. Mesonotum short, polished, punctate; notaulices narrow,
foveolate. Sternaulus foveolate, rest of mesopleuron polished, punctate. Propodeum short with distinct
carinae between which are obsolescent rugae. Tergite one with distinct dorsal pits and glymmae, dorsal
surface striate laterally, the striae turning inwards distally and joining the reticulate medial area of the tergite.
Ovipositor short, about 1-5 times length of tergite one. Legs short, the hind coxa smooth, punctate; tarsal
claws with no basal lobe but slightly swollen at the base.
Colour black except antenna at base, clypeus, mandibles, prothorax ventrally testaceous, tegulae and legs
yellow.
$. Only one specimen examined and this lacks a head; rest of body same as 9-
MATERIAL EXAMINED
5 9, 1 cf . Austria: 1 9, Siidtirol, St. Peter, Ahrntal, 25.viii.1967 (HaeselbartK) (EH, Munich). Germany: 1
$, 'Schmiedeknecht coll.' (CNC, Ottawa). Poland: 1 9, Polanowice, 10 km N. of Wroclaw, 15.vii.1957
(Pulawski) (RNH, Leiden). Sweden: 2 9> Thomson coll. (paralectotypes of Meteorus brevicauda Thomson)
(ZI, Lund); 1 9, Vormland, Stollet, 30.vi.-19.vii.1975 (van Rossem) (RNH, Leiden).
HOSTS. No reared material available.
Meteorus cespitator (Thunberg)
(Figs 16, 17)
Ichneumon cespitator Thunberg, 1822: 269. Holotype 9, SWEDEN (UDE, Uppsala) [examined].
Zele atrator Curtis, 1832: folio 415. Syntypes, GREAT BRITAIN: 'Wilts, Durnford House' (NMV,
Melbourne) [not examined]. [Synonymized by Roman, 1912: 289.]
Perilitus similator Nees von Esenbeck, 1834: 41. Syntypes 9» GERMANY (destroyed). [Synonymized by
Haliday, 1835:32.]
Meteorus ambiguus Ruthe, 1862: 30. Holotype 9, GERMANY: Ruthe coll., 4.10.55 (BMNH) [examined].
Syn. n.
There is one specimen in Thunberg's collection standing above the name cespitator. I regard this
specimen as the holotype of the species. It is in poor condition but is nevertheless readily
identifiable. I have based my concept of atrator on specimens in the BMNH collection which have
been identified as that species by G. E. J. Nixon who examined Curtis' collection in 1948 and chose
a type-specimen but never validated this selection by publication.
Ruthe (1862: 31) stated that he possessed one female of ambiguus captured on 4 October, 1855.
A specimen bearing these data now stands in the collection of the BMNH; it agrees with the
description of ambiguus and also bears a label in Ruthe's handwriting: 'P. ambiguus m'. I regard
this specimen as the holotype of the species. It comes within the limits of variation of cespitator
except that the flagellar segments are all distinctly longer than broad.
9. Antennae short, 23- to 27-segmented, the apical twelve or so segments of the flagellum short, not or very
little longer than broad. Head transverse, temples shorter than eyes. Ocelli small, OO = 2-5-3-0 times OD.
Eyes large, protuberant, convergent but not strongly so (Fig. 1 7). Malar space distinctly shorter than basal
breadth of mandible. Face about 1-5 times as broad as high, protuberant, generally completely rugulose but
sometimes smooth, punctate laterally and occasionally, especially in smaller specimens, more or less
completely smooth, punctate. Clypeus not strongly protuberant but distinctly divided from face, smooth but
with large punctures, these are sometimes dense, particularly at the sides of the clypeus, so that they form a
reticulate-punctate pattern. Tentorial pits distinct. Mandibles large, stout, moderately twisted. Pronotum
projecting in front of the mesonotum, laterally rugose but this is sometimes reduced so that the area is
smooth, shining. Notaulices deep, foveolate, rest of mesonotum densely punctate. Sternaulus deep and, at
least medially, irregularly reticulate-rugose, generally with a reticulate-foveolate area beneath; the sculpture
of the Sternaulus is sometimes obsolescent in smaller specimens; rest of mesopleurae smooth, with minute
punctures. Propodeum irregularly rugose with carinae distinct in all but the most heavily sculptured
REVISION OF THE WESTERN PALAEARCTIC METEOR US 25
specimens in which the carinae merge with the rugose sculpture. Tergite one striate dorsally, the striae
turning inwards towards the apex of the tergite, occasionally joining together medially but generally
coalescing in a regularly reticulate-rugose, medial longitudinal area. Ovipositor long, 3-4 times length of
tergite one. Hind coxae rugose, generally strongly so but in smaller specimens the sculpture is often
obsolescent; tarsal claws long, slightly swollen at base but not lobed.
Colour black; clypeus, face, prothorax ventrally often orange-testaceous, legs testaceous often the hind
coxa basally and the hind tibia apically infuscate, occasionally the hind leg almost completely dark brown-
testaceous; middle tergites of gaster sometimes testaceous. Some specimens from older collections have a dull
reddish body colour which I suspect to be due to fading, possibly exacerbated by over-exposure to cyanide in
the killing-jar (see also discussion ofconsimilis and offilator).
$. No material available.
MATERIAL EXAMINED
73 9- France, Germany, Great Britain, Hungary, Ireland, Luxemburg, Netherlands, Norway, Sweden.
HOSTS. Nemapogon granella (L.), Trichophaga tapetzella L., Tineola sp., (Lepidoptera: Tineidae).
Van Burgst recorded (as jaculator) the specimens reared from T. tapetzella L.; I have examined
these specimens and consider them to belong to cespitator. I have also examined specimens
recorded as 'probably ex Monopis rusticella infesting barn owl pellets' [Monopis rusticella
(Hiibner) (Lepidoptera: Tineidae)].
REMARKS. I have examined one specimen which I think probably belongs to this species but in
which the eyes are less convergent and which therefore has a face almost twice as broad as high.
The face is also densely reticulate-punctate and facially the specimen therefore resembles
longicaudis. In all other structural features, however, it comes within the limits of variation of
cespitator.
Meteorus cinctellus (Spinola)
(Fig. 39)
Bracon cinctellus Spinola, 1808: 135. Holotype tf, ITALY (MZS, Turin) [examined].
Perilitus fuscipes Wesmael, 1835: 48. Holotype 9» BELGIUM: Brussels, coll. Wesmael (IRSNB, Brussels)
[examined]. Syn. n.
Meteorus tenellus Marshall, 1887: 125. LECTOTYPE 9, GREAT BRITAIN (BMNH), here designated
[examined]. Syn. n.
Two specimens (<$ and 9) presently stand above the name 'cinctellus' in Spinola's collection. The
original description was based on a single male specimen and I therefore consider the male in
Spinola's collection to be the holotype. M. cinctellus has been correctly interpreted by most authors
although many of them have incorrectly attributed authorship of the species to Nees, who first
described the female sex.
Wesmael stated that he possessed only one female specimen of fuscipes :; one such specimen
presently stands in Wesmael's collection. It is labelled 'Perilitus fuscipes mini' in Wesmael's
handwriting and agrees precisely with his description of the species except that the darker colours
are somewhat faded. I regard this specimen as the holotype of fuscipes; it comes within the limits of
variation of cinctellus.
Marshall (1887: 125) based his description of tenellus on six female specimens reared by Bignell
from Peronea [Acleris] hastiana (L.). Two female specimens from Marshall's collection stand
above the name tenellus in the BMNH collection. They are mounted together on a square card on
the underside of which is written the host data cited by Marshall. The two specimens are
conspecific and agree with the description of tenellus and I have no doubt that they are syntypes of
that species. I have chosen the right-hand specimen as lectotype and have so marked it. Marshall
distinguished tenellus from cinctellus by the relatively shorter length of the narrow proximal part of
the first segment. I have found that the proportions of the first tergite exhibit considerable variation
within a species and I therefore consider tenellus to come within the limits of variation of cinctellus.
9- Antennae 24- to 29-segmented: distal flagellar segments not much longer than broad. Head contracted
behind eyes, generally more or less rounded, eyes almost twice as long as temples; frons rather depressed
26 T. HUDDLESTON
with a blunt tubercle in front of the fore ocellus. Ocelli small, OO = 2-5 times OD. Eyes large, protuberant,
strongly convergent. Malar space short, clearly shorter than the basal breadth of the mandible. Face about as
high as broad, sometimes smooth, punctate but often at least partly rugose, the rugae joining to form a
medial tubercle just below the antenna! sockets. Clypeus narrow, at most as wide as face; strongly
protuberant; polished with large punctures. Tentorial pits deep. Mandible small, strongly twisted. Pronotum
generally rugose, occasionally smooth, at least in part. Notaulices thin, shallow, foveolate. Sternaulus deeply
impressed, foveolate, generally broadened anteriorly by a patch of reticulate-foveolate beneath; rest of
mesopleuron polished, punctulate except for a reticulate area beneath the subalar prominence; mesolcus
deep, foveolate. Propodeum regularly rugose, sometimes reticulate-rugose; generally with a strong medial
transverse carina, other carinae usually indistinct except occasionally when the rugosity is reduced.
Propodeum generally rather flat with a long dorsal surface. Tergite one long, slender in basal half; dorsal pits
absent, ventral borders generally touching at about middle of segment but occasionally open; spiracles
distinctly behind midpoint of segment; dorsal surface longitudinally striate, the striae generally running
towards the middle of the segment distally, often reticulate-rugose at mid-segment and occasionally also at
the base but more usually smooth at base. Ovipositor 2-0-2-5 times length of tergite one. Legs long, slender;
hind coxae rugose, often reticulate-rugose, at least on the dorsal surface. Tarsal claws small and difficult to
see but with a distinct basal lobe.
Colour black; legs and generally tergite 2 testaceous; often basal half of flagellum and head in part
brownish testaceous; mesonotum around notaulices often reddish brown; hind legs occasionally dark, at least
in part; wings generally slightly embrowned.
cf. Same as 9 except antenna longer, 28- to 30-segmented; ocelli slightly larger; eyes less strongly
convergent; colour often darker.
MATERIAL EXAMINED
281 9, 87 cf. Austria, Finland, France, Germany, Great Britain, Ireland, Italy, Netherlands, Portugal,
Sweden, Yugoslavia.
HOST. Acleris hastiana (L.) (Lepidoptera: Tortricidae).
REMARKS. The convergent eyes and small ocelli readily distinguish the female of cinctellus from
closely related species. The male of cinctellus is rather similar to that of colon; the legs and
antennae of cinctellus, however, are not as conspicuously long and slender as those of colon; the
temples of cinctellus are more rounded, the eyes less protuberant and the sternaulus more strongly
and more extensively sculptured.
Meteor us colon (Haliday)
(Figs 44, 45)
Perilitus (Meteorus) colon Haliday, 1835: 30. Syntype(s) 9» IRELAND (lost).
Perilitus fragilis Wesmael, 1835: 52. LECTOTYPE £, BELGIUM: Brussels (IRSNB, Brussels), here
designated [examined]. Syn. n.
Perilitus fasciatus Ratzeburg, 1844: 77. Syntypes 9> GERMANY (lost). [Synonymized with fragilis by Ruthe,
1862:55.]
Meteorus luridus Ruthe, 1862: 57. LECTOTYPE, 9, GERMANY: Ruthe collection (BMNH), here designated
[examined]. Syn. n.
I have based my interpretation of colon on a specimen from Stelfox's collection which was named
as that species by Stelfox. The specimen agrees precisely with Haliday's description. The
descriptions of colon and fragilis were both published in 1835; Dessart (1972) has shown, however,
that Haliday's names were published prior to those of Wesmael and they must therefore take
precedence. M. colon was placed in synonymy with fragilis by Reinhard (in Ruthe, 1862: 55).
Fischer (1965: 21) employed this synonymy but correctly used the name colon for the species with
fragilis as a synonym. Later, however, Fischer (1970£: 260) lifted/ragzfo from synonymy.
Wesmael stated that he had three males and one female of fragilis. In Wesmael's collection
above this name there now stand eight specimens labelled 'Perilitus fragilis mihi' in Wesmael's
handwriting. Of these eight specimens only two males agree with Wesmael's description of fragilis
and I have chosen the better-preserved of these as lectotype. None of the other specimens can
possibly belong to the syntype-series, disagreeing as they do with the description in most
particulars; four of them are consimilis and two abdominator.
REVISION OF THE WESTERN PALAEARCTIC METEOR US 27
There are 12 specimens standing above the name luridus in Ruthe's collection; I have chosen as
lectotype the specimen which best fits Ruthe's description of the species and which bears a label in
Ruthe's handwriting 'P. luridus m.'. The lectotype and nine other specimens are conspecific with
colon, the two other specimens are discussed under heliophilus.
9- Antennae 30- to 34-segmented; long, thin, all flagellar segments at least twice as long as broad, the basal
two about 4 times as long as broad. Head contracted behind eyes; temple only slightly shorter than eye in
dorsal view; head between ocelli and occipital carina flattened, polished; frons with a blunt tubercle in front
of the fore ocellus so that, in lateral view, the frons appears angled. Ocelli large but not conspicuously so,
OO = 2 times OD. Eyes convergent but not strongly so. Malar space short, slightly less than basal breadth of
mandible. Face not strongly protuberant, generally with a transverse striate-rugose element medially.
Clypeus strongly protuberant; narrower than face and with widely spaced, large punctures; apical border
reflexed. Mandible small, delicate, strongly twisted. Sternaulus foveolate, rest of mesopleurae polished,
punctate. Propodeum finely rugose, occasionally reticulate-rugose, carinate but the carinae often weak and
indistinct. Tergite one long, slender, without dorsal pits, the ventral borders approaching at the mid point of
the segment but generally narrowly separated; ovipositor 1 -5 times length of tergite one. Legs very long and
slender; hind coxa with only a little rugosity on the dorsal surface; tarsal claws small but with a distinct basal
lobe. Recurrent vein always distinctly postfurcal, the second submarginal cell rather elongate (Fig. 45).
Colour black; face, genae inner orbit, propleurae, pronotum posteriorly, mesopleuron ventrally and legs
testaceous, sometimes also tergite two is this colour and occasionally the rest of the tergites, the scutellum
and the propodeum. Completely yellow specimens occur infrequently.
cf. Same as 9 except that antenna 31- to 35-segmented.
MATERIAL EXAMINED
123 9' 1^6 :?• France, Germany, Great Britain, Ireland, Italy, Netherlands, Sweden, Switzerland,
Yugoslavia.
HOSTS. Ladoga Camilla (L.) (Lepidoptera: Nymphalidae); Anticollix sparsata (Treitschke)
(Lepidoptera: Geometridae); Nola cuculatella (L.) (Lepidoptera: Nolidae); Bena prasinana (L.)
(Lepidoptera: Noctuidae).
REMARKS. M. colon is a conspicuously slender species with very long, slender antennae and legs.
The morphology, anatomy and biology of the species (under the namefragilis) has been excellently
monographed by Madel (1963).
Meteorus consimilis (Nees)
(Figs 54, 55)
Perilitus consimilis Nees von Esenbeck, 1834: 42. Syntype(s) tf, GERMANY (destroyed).
Perilitus brevipes Wesmael, 1835: 33. LECTOTYPE tf, BELGIUM: Brussels, coll. Wesmael (IRSNB,
Brussels), here designated [examined]. [Synonymized with albicornis Ruthe by Marshall, 1887: 110.]
Meteorus albicornis Ruthe, 1862: 34. LECTOTYPE 9, GERMANY: Ruthe collection (BMNH), here
designated [examined]. [Synonymized with consimilis by Thomson, 1895: 2160.]
There are four specimens in Wesmael's collection labelled 'Perilitus brevipes mihi' in Wesmael's
handwriting. Three of these specimens are males, one of them extensively damaged; the two others
are conspecific and agree with the description of brevipes, both coming well within the limits of
variation of consimilis. Ruthe (1862: 35) indicated that he had two specimens of albicornis; these
are now present in the BMHH collection. I have chosen as lectotype the specimen bearing Ruthe's
handwritten labels 'P. albicornis m.' and '6.10.55' which has previously been labelled as type of the
species (3C 753).
9- Antennae 32- to 33-segmented; thick, from about the fifteenth the apical segments of the flagellum are
quadrate or even a little broader than long. Head strongly contracted behind eyes; temples about equal in
length to eyes in dorsal view; frons depressed so that antennae are set upon a shelf, the antennal sockets
surrounded by concentric rugae which sometimes extend down onto the face. Ocelli large, OO= 1-5 times
OD. Eyes small, not convergent (Fig. 54). Malar space long, almost twice basal breadth of mandible. Face
protuberant, punctate, sometimes reticulate-punctate laterally. Clypeus strongly protuberant, polished,
punctate, narrower than face. Tentorial pits deep. Mandibles slender; strongly twisted with, at base of teeth, a
28 T. HUDDLESTON
small but distinct tubercle-like swelling which terminates the ventral longitudinal carina of the mandible.
Notaulices deep, narrow, foveolate. Sternaulus deep, broad, reticulate-rugose or sometimes foveolate with
scattered rugae and foveae beneath. Propodeum strongly reticulate-rugose with no distinct longitudinal
carinae but generally with a basal transverse carina and sometimes with a medial transverse carina which
divides the dorsal and posterior faces of the propodeum, the latter generally concave. Tergite one short, wide,
with distinct dorsal pits, dorsal surface strongly longitudinally striate. Ovipositor short, at most equal in
length to tergite one; thick at base, tapering to apex, down-curved. Legs thick, hind tibia swollen, only slightly
less broad than femur; hind coxa strongly sculptured, usually reticulate-rugose; tarsal claws strongly curved,
without a basal lobe but strongly thickened at base. Wings infumate, short, the second abscissa of the radius
short so that the second submarginal cell is strongly narrowed anteriorly even occasionally triangular.
Colour: antennae marked with a band of pale yellow which usually covers the basal half of the flagellum
but is sometimes restricted to the middle segments; head, thorax and tergite one piceous, remaining tergites
and legs testaceous.
$. Same as female except that the antennae are longer, 32- to 36-segmented, all segments of the flagellum
are distinctly longer than broad; there is never a pale band on the flagellum though the basal segments may
be slightly lighter in colour than the rest; colour slightly darker; hind tibia less strongly swollen; punctures on
face smaller so that the face appears smoother and shinier.
MATERIAL EXAMINED
170 (f, 9 9- France, Germany, Great Britain, Hungary, Ireland, Netherlands, Sardinia, Sweden.
HOST. I have examined no reared material of this species. It has frequently been reported as a
parasite of Scolytus multistriatus (Marsham) (Coleoptera: Scolytidae).
REMARKS. The darker parts of this species are dense pitchy black in freshly caught specimens; in
older specimens this colour seems lighter and more reddish. Specimens from Ruthe's collection
have all faded to red while a few of the specimens collected in Sweden in 1938 by Perkins are
distinctly reddish and most show some evidence of such a change. The pale-banded antennae and
the extremely short ovipositor at once distinguish the female of consimilis from all other European
species of Meteorus. The shape of the head is also most characteristic in both sexes.
Meteorus corax Marshall
(Figs 10, 11, 14)
Meteorus corax Marshall, 1898: 220. LECTOTYPE 9, SWITZERLAND: 'Val Somvix/25.vii. 189 I/Marshall
del.' (NMB, Berne), here designated [examined].
Meteorus monochami Fischer, 1957c: 17. Holotype 9- AUSTRIA: Lunz (Haberfelner), ex Monochamus
saltuarius Gebl. (ZSBS, Munich) [not examined]. [Synonymized by Fischer, 1966: 395.]
Marshall stated that he had five specimens before him when describing corax, three from Lunz,
Austria (Konow collection) and two from Val Somvix, Switzerland. The Konow collection is now
housed in Eberswalde and there is in that collection a series of six specimens standing above the
name Meteorus corax\ each of these specimens bears a label 'Col. Konow' and five of them are
labelled 'Lunz'. It is not now possible to distinguish Marshall's three syntypes from amongst these
six specimens nor is it possible therefore to select a lectotype or paralectotypes from them.
Marshall gave the depository for the Swiss specimens of corax as the 'Musee National de Berne'
but there has never been an institution so entitled (Volkart, in lift.). In the collection of the NMB,
Berne, however, there are three specimens standing above the name Meteorus corax, all with
locality labels which bear the data published by Marshall and two of them also labelled 'Marshall
det'. I believe the two latter specimens to belong to the syntype-series of corax and that Marshall
misquoted the name of the depository. The six Konow specimens are conspecific with those in
Berne and all agree closely with the description of corax except that they all have large distinct
dorsal pits whereas Marshall states 'Premier segment de 1'abdomen sans rainures tracheales'. I
believe that the evidence for the identity of corax given above is sufficient to allow this statement to
be discounted as being erroneous. I have examined a paratype of monochami and it is clearly
conspecific with corax, as pointed out by Fischer.
9- Antennae 39- to 4 1 -segmented, long, all segments at least 1-5 times as long as broad. Head expanded
behind the eyes, rather square in face view. Ocelli small, OO about 3 times OD. Eyes small, converging only
REVISION OF THE WESTERN PALAEARCTIC METEOR US 29
very slightly. Face about twice as long as broad. Clypeus distinctly narrower than face; protuberant; dorsal
surface smooth and punctate; ventral surface strongly carunculate. Tentorial pits wide and deep. Mandibles
very stout, twisted, with striate-punctate sculpture and long hairs. Pronotum strongly rugose, becoming
reticulate-foveolate posteriorly. Mesonotum punctate; notaulices slender but distinctly impressed, foveolate.
Sternaulus deep, foveolate with, anteriorly, a reticulate-foveolate patch beneath, rest of mesopleurae smooth,
punctate except for a reticulate-rugose patch dorsally. Propodeum strongly reticulate-rugose with basal and
medial transverse carinae; medial carina extremely prominent, dividing the propodeum into dorsal and
posterior planes; there is also usually a rather indistinct central longitudinal carina. Tergite one with
longitudinal strigose sculpture laterally, reticulate-rugose centrally; spiracles distinctly before the middle;
large dorsal pits present, borders of tergite one not meeting beneath. Tergite two longitudinally striate,
becoming reticulate-rugose basally; thyridia present at about mid-point of tergite; laterotergites large. The
ovipositor about 4 times length of tergite one. Outer surface of hind coxa and femur densely reticulate-
punctate; hind tibia strongly swollen though not as wide as femur; tarsal claws large, strongly curved, not
lobed but strongly swollen at base. Second submarginal cell as high exteriorly as wide anteriorly (Fig. 14).
Colour black, foreleg usually and midleg sometimes testaceous, hind leg always black except for an ivory
band around base of tibia.
(3*. No material examined.
MATERIAL EXAMINED
27 9 Austria: 1 9» Lunz (Haberfelner) 'aus Monocham. saltuarius' (ZSBS Munich) (paratype of Meteorus
monochami Fischer); 5 9» Lunz 'Konow coll.' (IP, Eberswalde); 1 9, 'Konow coll.' (IP, Eberswalde).
Germany: 'Parasit von Mon. sartor F.', no other data (EH, Munich), Sweden: 18 9> DIr. Rattvik, Mun.
Samh. t. (Tfeder) (5 in BMNH, rest in NR, Stockholm). Switzerland: 1 9, Val Somvix, 25.vii.1891, 'Marshall
det.' (NMB, Berne) (paralectotype of Meteorus corax Marshall); 1 9, Val Somvix, 25.vii.1891 (NMB,
Berne).
HOSTS. M. corax has been recorded from two species of Monochamus Guerin-Meneville
(Coleoptera: Cerambycidae). The possession of a long ovipositor is often correlated with the use of
wood-boring larvae as hosts.
REMARKS. This is the largest species of Meteorus, at least in the Palaearctic region, averaging
about 9-0 mm in head and body length. It is exceptionally dark in colour, the legs generally largely
black.
Meteorus eadyi sp. n.
(Fig. 3)
9- Antennae 26- to 27-segmented, all segments at least twice as long as broad. Head rounded behind eyes.
Ocelli small, OO = 2-5-3-0 times OD. Eyes small, protuberant, not strongly convergent (Fig. 3). Face about
1-5 times as broad as high; punctate, usually with transverse rugae beneath antennal sockets. Clypeus not
strongly protuberant, smooth with sparse punctures; the apical border produced laterally into a small flange
(Fig. 3). Mandible not twisted, polished, the upper tooth rather long. Pronotum expanded forwards so that it
projects beyond the mesonotum; smooth laterally with coarse rugae dorsally. Notaulices thin, foveolate.
Sternaulus foveolate, rest of mesopleurae polished, punctate. Propodeum with three longitudinal and two
transverse carinae with scattered rugae between. Tergite one long, slender, about twice as long as apically
broad; without dorsal pits; spiracles distinctly behind the middle; borders of tergite one conjoined beneath
from just behind the base of the segment to just in front of the spiracles; dorsally the tergite rugose or
reticulate-rugose with longitudinal striae laterally, particularly in apical half of segment. Ovipositor short,
about 1 • 5 times as long as tergite one, down-curved. Legs slender, hind coxae rugose, at least on the outer
surface at the base; tarsal claws without a basal lobe.
Colour black; clypeus, pronotum ventrally and legs testaceous, basal segments of the antennae usually
slightly lighter than the rest.
$. Same as 9 except that the eyes are smaller and less convergent.
MATERIAL EXAMINED
35 9, 2 (51. Holotype 9, Great Britain: Northamptonshire, Spratton, x.1975 (Gauld) (BMNH).
Paratypes. Bulgaria: 1 9, Rhodopi, Markovo, 13.viii.1977 (Zaykov) (ZC, Plovdiv). Great Britain: 17 9, 1
c?, England, Northamptonshire, Spratton, v.-x.!975 (Gauld) 1 9, E., Oxfordshire, Waterferry
Common, 5.viii.l969 (Brock} (BMNH); 1 9, E., Greater London, Kew, 22.vii.1979 (Eastop) (BMNH).
Germany (West) : 1 9, Bonn, Waldhauhohenweg, 18.xi.1960 (Schmidt) (CNC, Ottawa). Netherlands: 6 9,
30 T. HUDDLESTON
Wijster (opp. biol. station); 3, 13-26.viii.1973; 1, 7-14.vii.1974; 1, 10-17.ix.1976; 1, 16-23.vi.1978 (van
Achterberg ) (RNH, Leiden); 7 9, Waarder (ZH), Oosteinde; 1, l-15.ix.1973; 1, 17-22.vi.1974; 1, l-5.viii.
1974; 2, l-6.ix.1974; 1, 15-22.vi.1975; 1, 21-31.X.1975 (van Achterberg) (RNH, Leiden); 1 9, Ede,
IS.ix. 1970 (Zwakhals) (RNH, Leiden). Yugoslavia: 1 9, Plitvice, 4-10. vii. 1955 (Coe) (BMNH).
Non-paratypic material. France: 1 9> Chartrettes, 26.viii.1949 (Granger) (MNHN, Paris). Great Britain:
1 c?, Wales, MM, Govilon, Marshall coll. (BMNH).
HOSTS. No reared material examined.
REMARKS. The structure of the thorax, of the propodeum and of tergite one show this species to be
very closely related to filator with which it has often been confused in collections. M. eadyi can be
distinguished, however, by its smaller, less convergent eyes, by the form of its clypeus and of its
antennae and by its shorter ovipositor.
Haliday (1835: 33) noted the similarity between his species filator and delator. The close
resemblance to filator of the species here described at first led me to suppose that it might be
delator, which has never been adequately interpreted. In the description of delator, however, it is
clearly stated that the ovipositor is as long as the abdomen [gaster] which is certainly never the
case in eadyi.
Meteorus filator (Haliday)
(Figs 2, 7)
Perilitus (Meteorus) filator Haliday, 1835: 32. Holotype 9, IRELAND: 'British Haliday 20.2.82' (NMI,
Dublin) [examined].
Perilitus laticeps Wesmael, 1835: 47. LECTOTYPE 9, BELGIUM: Brussels, coll. Wesmael (IRSNB,
Brussels), here designated [examined]. [Considered valid species by Fischer, 1970a.] Syn. n.
Meteorus hodisensis Fischer, 19706: 285. Holotype 9, AUSTRIA: 'Bgld., Markt. hodis, Rechnitz, 7.viii.l961'
(Fischer) (NHM, Vienna) [examined]. Syn. n.
The only specimen of this species in the Haliday collection is that labelled as '9 type' by Stelfox in
1948. 1 therefore regard it as the holotype. The specimen is glued onto a rectangular piece of card
and is in an excellent state of preservation but is rather dirty.
As pointed out by Fischer (1970a) there are two broken specimens in Wesmael's collection
which are labelled 'Perilitus laticeps mihi' in Wesmael's handwriting. Neither of the specimens has a
head but glued to the label of one of them is the head which was illustrated by Fischer (1970a: 53)
in his redescription of laticeps. Largely on the basis of this head Fischer lifted laticeps from
synonymy with filator, where it had been placed by Marshall (1887: 121). The head on the label
does not agree, however, with what Wesmael (1835: 47) said about the head of laticeps and I
consider it to have been mistakenly associated with the specimen on the pin. The parts of the
specimen which remain on the pin are without doubt conspecific -with filator, the head glued to the
label is that of a male consimilis. I consider Marshall's synonymy of the two species further
evidence in support of my view; Marshall had examined Wesmael's collection and he also knew
filator and so gross a difference as that presented by the misassociated head is unlikely to have
escaped his notice. Fischer also stated that one of the specimens possessed a gaster; this is un-
fortunately no longer present. The holotype of hodisensis falls within the range of variation of
filator.
9- Antennae 21- to 25-segmented, short; first two flagellar segments about 3 times as long as broad, five
preapical segments only slightly longer than broad, submoniliform. Head broad, temples rounded. Ocelli
small, OO = 2- 5-3-0 times OD. Eyes large, strongly convergent (Fig. 2). Face about as wide as high, not
strongly protuberant, rugose, at least centrally, sometimes punctate laterally. Clypeus wide, polished,
sparsely punctate, not strongly protuberant. Malar space short, distinctly less than half the basal breadth of
mandible. Mandible large, not twisted. Pronotum projects forwards beyond the mesonotum and is slightly
expanded sideways, strongly rugose. Mesonotum short, smooth; notaulices deep thin foveolate furrows.
Sternaulus wide, foveolate, often with scattered rugae and foveae above and beneath; sculpture occasionally
obsolescent. Propodeum rather depressed with two transverse and three longitudinal carinae, rugose between.
Tergite one long, almost equal in length to rest of gaster, slender, 2-0—2-5 times as long as apically broad,
ventral edges conjoined from near base of segment to its midpoint (Fig. 7), rugose dorsally with laterally in
the apical half of the segment a longitudinal striate element which occasionally replaces the medial rugosity;
REVISION OF THE WESTERN PALAEARCTIC METEOR US 3 1
spiracles distinctly shortly behind mid-segment. Ovipositor long, 2-5 times length of tergite one. Legs long,
slender; hind coxa reticulate-rugose, at least dorso-laterally; tarsal claws simple, long, slightly swollen at
base. Wings large with a faint brown tinge; recurrent antefurcal, rarely interstitial.
Colour black except base of antennae, clypeus, mandibles, palpi, anterior margin of pronotum and legs
yellow; hind coxa usually darker, at least in part, and rest of hind leg may be infuscated. Antennae and face
occasionally completely yellow; in these lighter coloured specimens the pronotum is more extensively yellow
though never completely so. I have seen one specimen in which the normally black parts of the body are deep
reddish brown; this is probably an artefact such as is not uncommon in black insects which have been
exposed overlong to cyanide in a killing bottle.
<$. Like the 9 except eyes are smaller and less convergent so that the face is about twice as broad as high
and the malar space longer; antennae longer, 27- to 30-segmented, all segments of flagellum at least twice as
long as broad, darker in colour, rarely distinctly yellow in basal half of flagellum.
MATERIAL EXAMINED
130 9, 40 tf. Austria, Bulgaria, France, Germany, Great Britain, Ireland, Italy, Netherlands, Norway,
Sweden.
HOSTS. No reared material examined.
REMARKS. The characteristic first segment of the gaster, the shape and size of the eyes and of the
antennal segments at once distinguish filator from all other species.
Meteorus graciliventris Muesebeck
(Figs 46, 47)
Meteorus graciliventris Muesebeck, 1954: 62. Holotype 9, JAPAN: Chosen (USNM, Washington, 62029)
[not examined].
Through the courtesy of Dr P. M. Marsh I have been able to examine seven female paratypes of this
species.
9- Antennae 31- to 33-segmented, the penultimate six or so segments not much longer than broad. Head
strongly narrowed behind the eyes, the temples short, the eyes at least four times the length of the temple.
Ocelli large, OO 1-25-1-50 times OD. Eyes very large, protuberant, strongly convergent so that the malar
space is almost obliterated. Frons and vertex densely, regularly punctured. Face distinctly higher than wide,
reticulate-punctate with some rugosity beneath the antennal sockets. Clypeus not strongly protuberant,
smooth, with scattered large punctures. Tentorial pits small, very close to eye margin. Mandibles small,
moderately twisted. Pronotum laterally coarsely rugose except for a polished punctate area dorsally.
Sternaulus deeply impressed, foveolate, broadened anteriorly by a patch of reticulate-foveolate sculpture on
the mesopleurae beneath the sternaulus, rest of mesopleurae shining punctate except for a subalar patch of
rugosity. Propodeum broad, coarsely rugose, the carinae not distinctly differentiated. Tergite one long,
slender, with dorsal pits, coarsely striate laterally and with a median longitudinal rugulose band. Ovipositor 3
times length of tergite one. Hind coxae strongly sculptured, usually reticulate-rugose; tarsal claws not
strongly curved and with a small, inconspicuous basal lobe. Colour black; antennae, mandibles, palps, tergite
two and legs testaceous except for the hind coxa, femur and tarsus distally which are infuscated.
$. No material examined.
MATERIAL EXAMINED
Japan: 7 9, Yokohama, vii.1939, paratypes of Meteorus graciliventris Muesebeck (USNM, Washington).
HOST. Pectinophora gossypiella Saunders (Lepidoptera: Gelechiidae).
REMARKS. The singular thing about graciliventris is that the paratypes are the progeny of the
holotype, having been bred in the laboratory in Ephestia cautella (Walker) (Lepidoptera:
Pyralidae).
Meteorus gyrator (Thunberg)
(Fig. 56)
Ichneumon gyrator Thunberg, 1822: 261. Holotype <$, SWEDEN (UDE, Uppsala) [examined].
Perilitus scutellator Nees von Esenbeck, 1834: 38. Syntypes $ 9, GERMANY (lost). [Synonymized by
Roman, 1912:289.]
32 T. HUDDLESTON
[Meteorus melanostictus Capron in Marshall, 1 887: 115. Males only from type-series.]
Meteorus parvulus Thomson, 1895: 2156. Holotype 9> SWEDEN: Oland (ZI, Lund) [examined]. Syn. n.
There is a single specimen of gyrator in Thunberg's collection and, although it is in poor condition,
it is readily identifiable. I regard this specimen as the holotype of the species. There is a single
specimen in Thomson's collection standing above the name parvulus which agrees with Thomson's
description of that species and which is labelled '0' (= Oland, the type-locality). I regard this
specimen as the holotype of parvulus', it is rather smaller than the average specimen of gyrator but
has all the structural features of that species and I have no doubt that the two are conspecific.
9. Antennae long, 30- to 33-segmented, at most the two penultimate segments of the flagellum less than twice
as long as broad. Head always contracted behind eyes, occasionally strongly rounded; temple always at least
a little shorter than eye in dorsal view, eye never more than twice as long as temple. Frons with a short,
shallow depression in front of fore ocellus. Ocelli large, OO= 1-0-1-5 times OD (exceptionally OO = 2 times
OD). Eyes large, protuberant, generally but little convergent. Malar space a little less than basal breadth of
mandible. Face generally a little broader than high, moderately protuberant, densely punctate, even
reticulate-punctate, sometimes with some rugosity medially. Clypeus strongly protuberant, densely punctate,
divided from the face by a deep fold. Mandibles long, strongly twisted. Pronotum projecting a little in front of
the mesonotum, laterally rugose though this is occasionally largely obsolescent. Notaulices anteriorly broad,
reticulate-rugose, coalescing posteriorly into a broad reticulate-rugose area which is rather depressed below
the level of the lateral lobes of the mesonotum; rest of mesonotum polished, densley punctate, sometimes
with a weak rugulose element, particularly on the lateral lobes. Sternaulus narrow, foveolate, occasionally
broadened anteriorly by reticulate-foveolate patch beneath and, occasionlly, with some rugae; rest of
mesopleurae smooth with minute punctures except for a rugose area beneath the subalar prominence.
Propodeum broad, generally rather flat, densely but finely rugose, sometimes reticulate-rugose with a central
longitudinal carina which is sometimes weak and occasionally broken but always present, at least in part; the
rugosity is occasionally coarse but is then sparse and not reticulate. Tergite one with distinct dorsal pits;
dorsal surface strongly striate; lateral surface rugose at base, striate apically; glymmae shallow. Ovipositor
short, 1-5-2-0 times length of tergite one, thick, strongly swollen at base. Legs long, thin; hind coxa smooth,
punctate, often with a trace of weak rugosity laterally, occasionally with stronger rugosity dorsally, but never
strongly reticulate-rugose. Tarsal claws always with a basal lobe but this is often difficult to see because the
claws are rather small and often partly buried in the hair of the apical tarsal segment. Wings never infumate,
even in the darkest-bodied specimens; the pterostigma always uniformly pale testaceous.
Colour black; face, clypeus, genae anteriorly, vertex laterally, thorax ventrally, scutellum and tergites 2
and 3 orange-testaceous, legs yellow; few darker specimens occur and I have seen none in which the
scutellum is not light in colour; completely testaceous specimens are not uncommon and all intermediate
patterns occur.
cf. Same as 9 except eyes smaller and malar space longer; propodeum often more coarsely sculptured;
colour often darker, the pterostigma generally dark, often with a pale border.
MATERIAL EXAMINED
220 9, 201 $. Austria, Bulgaria, Cyprus, Finland, France, Germany, Great Britain, Hungary, Ireland,
Netherlands, Sweden.
HOSTS. Lycophotia porphyrea (Denis & Schiffermiiller), Diarsia brunnea (Denis & Schiffermiiller),
Ipimorpha retusa (L.), Xestia xanthographa (Denis & Schiffermuller), Mythimna sp., Lacanobia
oleracea (L.), Agrochola lota (Clerck), Euplexia lucipara (L.), Cosmia trapezina (L.), Thalpophila
matura (Hufnagel), Cleoceris viminalis (Fabricius) (Lepidoptera: Noctuidae).
REMARKS. M. gyrator is superficially similar to pulchricornis, being similar in stature and in
colour. The clypeus of gyrator, however, is shining with scattered punctures and hairs, quite unlike
that of pulchricornis; gyrator also has a deeply impressed, narrow, foveolate sternaulus (whereas
that of pulchricornis is shallow rugose), a more depressed and generally more finely rugose
propodeum and a uniformly pale stigma, at least in the female (the stigma in pulchricornis is dark
and pale-bordered).
Meteorus heliophilm Fischer
(Figs 32, 33)
Meteorus heliophilus Fischer, 19706: 284. Holotype 9 AUSTRIA: Burgenland, Rechnitz, 2.viii.l958 (Fischer)
(NM, Vienna) [examined].
REVISION OF THE WESTERN PALAEARCTIC METEOR US 33
Among Ruthe's material of luridus two specimens stand which are quite distinct from the other
specimens of that species. One of these specimens bears a label 'pallidus' in Ruthe's handwriting
and I take this to indicate that Ruthe considered this specimen to be his var. pallida of luridus.
These two specimens of Ruthe's are clearly conspecific with heliophilus which is the species
misidentified as luridus Ruthe by Marshall (1887), by Lyle (1914) and by Tobias (1976).
9- Antennae long 30- to 32-segmented, all segments of flagellum clearly much longer than broad. Head
strongly contracted behind eyes. Eye in dorsal view 2-5-3-0 times as long as temple. Ocelli large,
OO= 1-0-1-5 times OD. Eyes large, protuberant, not strongly convergent. Malar space short, less than half
basal breadth of mandible. Face about as broad as high, not strongly protuberant, slightly raised medially
where it is transversely rugose, laterally punctate. Clypeus protuberant though not strongly so, smooth with
scattered coarse punctures. Tentorial pits small, indistinct. Mandibles strongly twisted, the upper tooth not
conspicuously long. Pronotum laterally smooth with a little fine rugose sculpture medially. Mesonotum
punctate, sometimes the central lobe reticulate-punctate, at least in part; notaulices weakly impressed.
Sternaulus weakly rugose, sometimes obsolescent posteriorly. Propodeum shining with weak carinae and
weak, largely obsolescent rugae dorsally. Tergite one long, slender, ventral borders meeting beneath at the
mid-part of the segment, not joined at the base of the segment; dorsal surface weakly longitudinally striate;
no glymmae or dorsal pits. Ovipositor about twice length of tergite one, thick, strongly expanded at the base
and the apical sixth much narrowed. Legs long, slender; hind coxa mainly smooth but sometimes with a trace
of rugosity dorsally. Tarsal claws with a distinct basal lobe.
Colour testaceous, the legs generally a paler shade of yellow.
$. Same as 9 except antennae 31- to 33-segmented.
MATERIAL EXAMINED
34 9, 21 tf. Bulgaria: 1 9, Rhodopi, Velingrad, 20.x. 1977 (Zaykov) (ZC, Plovdiv). Germany: 4 9, Harz,
27.iv.1957 (Hinz), ex Agrotis triangulum (BMNH); 2 9, Ruthe coll. (BMNH). Great Britain: 27 9, 19 tf,
England, Hampshire, New Forest (Lyle) (BMNH).
HOSTS. Polia nebulosa (Hufnagel), Lithophane ornitopus (Hufnagel), Noctua fimbriata (Schreber),
Xestia triangulum (Hufnagel), Orthosia stabilis (Denis & SchirTermiiller) (Lepidoptera: Noctuidae).
Both Lyle (1914) and Tobias (1976) recorded this species as being gregarious, several parasite
larvae developing in one host caterpillar.
REMARKS. M. heliophilus is structurally very close to lionotus but the differences in head shape
between the two species together with the characteristics given in the key to species amply
distinguish them. The two species also have a different host-spectrum and heliophilus is gregarious
and lionotus solitary. M. heliophilus is also close to rubens and shares the latter's gregarious habit
and use of noctuids as hosts. M. rubens, however, has shorter antennae, smaller eyes and ocelli and
less strongly contracted temples than heliophilus. The three species are clearly closely related.
Meteorus hirsutipes sp. n.
(Figs 25, 28, 29, 35)
9- Antennae 28- to 30-segmented, thin, all flagellar segments longer than broad, the three basal ones at least
3 times as long as broad. Head strongly narrowed behind eyes, temples distinctly shorter than eyes in dorsal
view; occiput concave. Ocelli small, OO = 3 times OD. Eyes protuberant, slightly convergent. Face about
twice as broad as high, not strongly protuberant. Clypeus not strongly protuberant, as broad as face,
punctate. Mandible very large, stout and not twisted. Prothorax projecting before the mesonotum with strong
vertical rugae laterally. Mesonotum polished with fine punctures; notaulices deep, narrow, foveolate.
Sternaulus deep, foveolate, angled sharply upwards in its anterior half; rest of mesopleurae polished,
punctured. Propodeum with strong longitudinal and transverse carinae with a few weak rugae between,
posteriorly with strong, mainly vertical rugae. Tergite one with coarse longitudinal striae, the innermost striae
turn inwards posteriorly and join to form a transverse rugose area; medial part of tergite anteriorly reticulate-
rugose. Ovipositor 2-5 times length of tergite one, straight. Hind coxa punctate or reticulate-punctate; tarsi
with long hairs the longest of which at the apices of the tarsal segments are at least twice the maximum breadth
of the segment. Tarsal claws simple, very long, sickle-shaped.
Colour black, prothorax and legs testaceous.
. No material examined.
34 T. HUDDLESTON
MATERIAL EXAMINED
3 9. Holotype 9, Finland: Valamo, Ladoga, NE. of monastery Hainwald 24.viii.1935 (Kerrich) (BMNH).
Paratypes. Germany (West): 1 9, Bavaria, Schliersee, 28.vii.1958 (Townes) (CNC, Ottawa). Ireland: 1 9,
Co. Wicklow, Deputy's Pass, 7.viii.l932 (Stelfox) (USNM, Washington).
HOSTS. No reared material examined.
REMARKS. This species is extremely close to brevicauda. The main differences between them are
those cited in the key to species; the hairs of the hind tarsi also differ, those of brevicauda being
much shorter, the longest hairs at most equal in length to the maximum breadth of the tarsi.
Meteorus ictericus (Nees von Esenbeck)
(Figs 52, 53)
Bracon ictericus Nees von Esenbeck, 181 1: 22. Syntype(s) 9> GERMANY (destroyed).
Ichneumon minutor Thunberg, 1822: 266. Holotype 9 (UDE, Uppsala) [examined]. [Synonymized by
Roman, 1912: 267.]
Zele ephippium Curtis, 1832: folio 415. Syntypes, GREAT BRITAIN: 'Coomb Wood' (NMV, Melbourne) [not
examined]. [Synonymized by Curtis, 1837.]
Perilitus xanthomelas Wesmael, 1835: 29. LECTOTYPE tf, BELGIUM: Brussels (IRSNB, Brussels), here
designated [examined]. [Synonymized by Fischer, 19706: 263.]
Meteorus confinis Ruthe, 1862: 18. LECTOTYPE 9, GERMANY: Ruthe coll. (BMNH), here designated
[examined]. Syn. n.
Meteorus fallax Ruthe, 1862: 18. LECTOTYPE 9, GERMANY: Ruthe coll. (BMNH), here designated
[examined]. Syn. n.
Meteorus pleuralis Ruthe, 1862: 19. LECTOTYPE 9, GERMANY: Ruthe coll. (BMNH), here designated
[examined]. Syn. n.
Meteorus liquis Ruthe, 1862: 20. LECTOTYPE 9, GERMANY: Ruthe coll. (BMNH), here designated
[examined], Syn. n.
Meteorus consors Ruthe, 1862: 44. LECTOTYPE 9, GERMANY: Ruthe coll. (BMNH), here designated
[examined]. [Synonymized by Fischer, 19706: 263.]
Meteorus crassicrus Thomson, 1895: 2154. LECTOTYPE 9, SWEDEN: Skane, Arrie (ZI, Lund), here
designated [examined]. Syn. n.
Meteorus lophyriphagus Fahringer in Schonwiese, 1934: 495. Holotype 9' AUSTRIA (IFF, Vienna)
[examined]. Syn. n.
Meteorus adoxophyesi Minamikawa, 1954: 41. Holotype 9> JAPAN: Kanaya Shizuoka-ken, 10.iv.1948
(Sonan) (NIAS, Yatabe) [examined]. Syn. n.
There is one specimen standing above the label 'minutor' in Thunberg's collection. It is in a
reasonably good state of preservation and is clearly within the limits of variation of ictericus. The
specimen is labelled 'Meteorus minutor Thbg.' in Roman's handwriting.
I have based my interpretation of ephippium Curtis on a specimen labelled 'Met. (Zele)
ephippium C. com. w. type' by G. E. J. Nixon who examined the Curtis collection in 1948. Nixon
chose a 'type-specimen' for ephippium but never validated his selection by publication.
Wesmael stated that he had two males of xanthomelas. One male and one female presently stand
in his collection above this name, both of them labelled 'Perilitus xanthomelas mini' in Wesmael's
handwriting. The female clearly cannot be part of the syntype-series and I have therefore chosen the
male as lectotype.
Ruthe stated that he possessed two females of confinis; these are present in the collection of the
BMNH. I have selected as lectotype the specimen bearing Ruthe's label 'P. confinis m.'.
Ruthe stated also that he possessed two females of fallax, these are present in the collection of
the BMNH and I have selected as lectotype the specimen bearing a label 'P. fallax m.' in Ruthe's
handwriting.
Ruthe did not state what material he had of pleuralis or of liquis. There is one specimen of each
of these species in the BMNH, each bearing a label in Ruthe's handwriting; I have designated these
specimens as lectotypes of their respective species.
There are four conspecific specimens standing above the name 'consors' in Ruthe's collection. I
have selected as lectotype the specimen which bears a label 'P. consors m.' in Ruthe's handwriting.
REVISION OF THE WESTERN PALAEARCTIC METEOR US 35
Four conspecific female specimens stand above the name 'crassicrus' in Thomson's collection.
Two of them bear a label 'ar' [=Arrie i Skane, one of the published localities]; one of these
specimens also bears a label 'crassicrus' in Thomson's handwriting. I have chosen the latter
specimen as lectotype of crassicrus and the second specimen from Arrie, labelled paratype by
Fischer, as paralectotype.
Fahringer stated that he possessed one female of lophyriphagus. This specimen, a photograph of
which appears in Schonwiese (1934: 496), stands in the collection of the Universitat fur
Bodenkultur, Vienna. It bears a label 'Meteorus lophyriphagus n. sp. Type' in Fahringer's
handwriting and a second label 'Rinkenberg K., 25.7.32, ex Loph. sif.'. It is well within the limits of
variation of ictericus.
Minamikawa, in his description of adoxophyesi, stated that tergite one was about three times as
long as apically broad and that the tracheal grooves [dorsal pits] were obsolete. These points in the
description certainly do not agree with what is to be observed on the holotype in which tergite one
is twice as long as apically broad and dorsal pits are present. Thus adoxophyesi comes within the
limits of variation of ictericus. Minamikawa formerly used the name Sonan under which name he
cited himself as the author of Meteorus adoxophyesi (1954: 43). Since the paper in which this
species was described appeared under the authorship of Minamikawa I have used this name in
citing authorship of adoxophyesi, as did Shenefelt (1969: 50).
9- Antennae 26- to 3 3 -segmented, long, all segments at least slightly longer than broad. Head strongly
contracted behind the eyes (Fig. 53), generally not strongly rounded; length of eyes 2-5-3-0 times length of
temple in dorsal view. Ocelli large, OO= 1-0-1-5 times OD, protuberant, the ocellar area distinctly raised.
Eyes large, protuberant inner margins converging (Fig. 52) but never strongly so. Malar space short, at most
equal to half basal breadth of mandible. Face not strongly protuberant but often slightly raised medially,
polished, finely punctate. Clypeus narrower than face, protuberant, polished, sparsely punctate, the ventral
border often broadly and shallowly emarginate. Tentorial pits small, indistinct. Mandibles moderately
twisted, short but fairly stout. Pronotum laterally smooth, punctate. Notaulices deeply impressed, foveolate.
Sternaulices foveolate, narrow, often at least partly obsolescent; rest of mesopleuron polished, finely
punctate. Propodeal carinae weakly developed except for the medial transverse carina which is strongly
raised, at least centrally; propodeum dorsally polished with a few obsolescent rugae between the carinae, the
rugose sculpture sometimes rather stronger and occasionally reticulate-rugose though never strongly so.
Tergite one long, slender with distinct dorsal pits and glymmae; dorsal surface striate. Ovipositor long,
2-5—3-0 times length of tergite one, stout, usually slightly down-curved at the apex. Legs long, slender; hind
coxa polished and with fine puncturation, often reticulate-punctate ventrolaterally but never with distinct
rugosity; hind tibia swollen, particularly in its apical half but not usually as broad as the femur; tarsal claws
strongly bent and with a large basal lobe. Recurrent always antefurcal, usually markedly so but occasionally
almost interstitial.
Colour black except head, prothorax, legs and pterostigma which are yellow, tergite two is often also
yellow or at least lighter in colour than rest of gaster; head always yellow except sometimes for a dark patch
on vertex around ocelli and occasionally extending back to the occipital carina, legs also generally yellow
except that the apex of hind tibia and of the tarsal segments are infuscated. I have seen only one specimen of
this species in which the head was almost completely dark and in this specimen the legs were also rather dark.
Lighter-coloured specimens are not infrequent in occurrence; the thorax and gaster either in whole or in part
may be reddish testaceous rather than black. Rarely specimens are found which are wholly reddish
testaceous except for the ovipositor sheaths which are black.
$. Same as 9 except antennae longer.
MATERIAL EXAMINED
655 9' 14 <$. Austria, Czechoslovakia, Finland, France, Germany, Great Britain, Hungary, Ireland,
Japan, Netherlands, Sweden, Turkey.
HOSTS. Archips podana (Scopoli), Epiphyas postvittana (Walker), Adoxophyes orana (Fischer von
Roslerstamm), Adoxophyes privitana (Walker), Tortrix viridana (L.), Croesia bergmanniana (L.),
Acleris hastiana (L.), Epinotia sordidana (Hiibner) (Lepidoptera: Tortricidae). Neodiprion sertifer
(Geoffrey in Fourcroy) (Hymenoptera: Diprionidae).
REMARKS. Meteorus ictericus is a rather slender, conspicuously smooth species; it is rather similar
in appearance to obsoletus from which it is easily distinguished by its possession of dorsal pits on
tergite one.
36 T. HUDDLESTON
I am by no means confident that I have adequately defined the specific limits of ictericus and of
pallipes. The typical forms of the two species are fairly easily differentiated but small specimens
quite frequently occur which are difficult to place. It may be that ictericus and pallipes as I have
defined them comprise a complex of species which I am at present unable to separate, or there may
only be a single large protean species. I am retaining them here as two distinct but closely related
species. I have given ictericus rather broader limits than pallipes which I have restricted to the
typical form. Included in the synonymy of ictericus are confinis, fallax, pleuralis, liquis and
crassicrus although they are all smaller and lighter in colour than typical ictericus. Ruthe gave his
four nominal taxa specific rank on the basis of their differences in colour from typical ictericus and
on their shorter antennae. Colour differences are not a reliable basis for specific distinction and the
differences in the antennae are only such as are to be expected in small specimens. Thomson
differentiated crassicrus principally by its possession of a swollen hind tibia. In Thomson's original
material, however, the hind tibia is not more swollen than in other specimens of ictericus. Although
crassicrus is rather smaller and paler than typical specimens of ictericus it is otherwise within the
limits of variation of that species.
Meteorus jaculator (Haliday)
(Fig. 15)
Perilitus (Meteorus) jaculator Haliday, 1835: 34. Syntypes 9, IRELAND (lost).
Meteorus obscurellus Ruthe, 1862: 29. Holotype 9> GERMANY: Ruthe coll. (BMNH) [examined].
[Synonymized by Marshall, 1887: 108.]
Meteorus tenuicornis Thomson, 1895: 2164. LECTOTYPE 9, SWEDEN: Skane, Palsjo (ZI, Lund), here
designated [examined]. Syn. n.
There is no specimen of this species present in Haliday's collection; I have therefore based my in-
terpretation of jaculator on specimens from Stelfox's comprehensive collection of Irish insects
which Stelfox himself had named as jaculator. These specimens fit Haliday's description and I
believe Stelfox's interpretation to be correct. There is one specimen of obscurellus in the BMNH
collection; this specimen bears a label in Ruthe's handwriting 'P. obscurellus M.' and corresponds
exactly with the description of that species. It is clearly conspecific with the insects I have
interpreted as jaculator. There is one female specimen standing above the name tenuicornis in
Thomson's collection, it is from the type-locality and agrees with Thomson's description of the
species. This specimen has been labelled 'type' by Fischer; there is a second (male) specimen taken at
Molle. .
9- Antennae short, 18- to 26-segmented, slender, all segments of flagellum at least as long as broad. Head
swollen behind eyes; temples longer than eyes (Fig. 15). Ocelli small, OO = 3-4 times OD. Eyes small, height
of eye less than breadth efface; slightly convergent. Malar space at least slightly shorter than basal breadth
of mandible. Face about twice as broad as high, protuberant, reticulate-punctate with some rugosity
medially, occasionally entirely smooth and shining with minute punctures. Clypeus smooth, shining, with
scattered large punctures, not strongly protuberant. Tentorial pits small but distinct. Mandibles stout, not
strongly twisted. Pronotum projecting in front of mesonotum; laterally smooth and weakly rugose.
Notaulices thin, foveolate. Sternaulus narrow, reticulate-foveolate with a few scattered rugae; rest of
mesopleurae smooth, minutely punctured except for a large reticulate-foveolate area beneath the base of the
forewing. Propodeum weakly rugose, the carinae weak but generally distinct, occasionally the rugosity is
strong enough to obscure the carinae. Tergite one generally predominantly irregular reticulate-rugose with
only a few striae laterally, occasionally the striae are predominant but usually then they are obsolescent; the
medial longitudinal area of the tergite is strongly raised. Ovipositor 3—4 times length of tergite one. Hind coxa
generally strongly rugose, occasionally weakly so. Tarsal claws short with no basal lobe.
Colour black; legs testaceous but sometimes the coxae darker; clypeus and mandibles often yellow.
3. Same as 9 except that the antennae much longer, up to 29-segmented, all flagellar segments con-
spicuously longer than broad.
MATERIAL EXAMINED
42 9, 1 (j*. Bulgaria: 1 9, Rhodopi, 23.viii.1977 (Zaykov) (ZC, Plovdiv). Germany (West): 1 9, Hiedeck,
Heidelb. (Haeselbarth) (EH, Munich). Great Britain: 12 9, England, Northamptonshire, Spratton, viii.-
ix.1975 (Gauld) (BMNH); 1 9, E., Hertfordshire, Tring, 8.vi.l936 (Benson) (BMNH); 1 9, E., H., Bricket
REVISION OF THE WESTERN PALAEARCTIC METEOR US 37
Wood, 28.V.1937 (Nixon) (BMNH); 1 9, E., Cambridgeshire, Woodwalton, 21-23.V.1949 (Benson)
(BMNH); 1 9, E., Dorset, Studland Heath, 6.vi.l938 (Benson) (BMNH); 1 9, E., Kent, Eynsford, 6.vi.l939
(Nixon) (BMNH); 1 9, E., Hereford and Worcester, Fownhope, Capler Wood, 31.V.1936 (Perkins)
(BMNH); 1 9, E., H. and W., Fownhope, Pagets Wood, l.vi.1936 (Perkins) (BMNH); 1 9, E., Devon,
Torquay, 19.vi.1960 (Vockeroth) (CNC, Ottawa). Ireland: 1 tf, Co. Dublin, B'Scorney, 15.vii.1945 (Stelfox)
(USNM, Washington); 2 9, Co. Down, Tollymore Park, 27.viii.1961 and 17.ix.1961 (Stelfox) (USNM,
Washington); 2 9, Co. Do., Dramahilly, 20. and 27.vii.1964 (Stelfox) (USNM, Washington); 1 9, Co. Do.,
Donnaird Lo., 17.X.1965 (Stelfox) (USNM, Washington); 5 9, Co. Sligo, Trawallua, 24-29.vi.1933 (Nixon)
(BMNH). Netherlands: 2 9, Waarder, Oosteinde, vii. and ix.1974 (van Achterberg) (RNH, Leiden); 2 9,
Wijster(Dr.)vi.-vii.l973 and vii. 1977 (van Achterberg) (RNH, Leiden); 1 9,Oostvoorne, 15-20.V. 1973 (Paw
Achterberg) (RNH, Leiden); 1 9 ,Meyendal near the Hague, Bierlap outer dunes, 18-25. vii. \914(vanderZon)
(RNH, Leiden); 1 9, Ede,22-28.ix. (van Rossem) (RNH, Leiden); 1 9,Nunspeet, l.x.!975(Zwacfca/.s)(RNH,
Leiden); 1 9, Wassenaar, 18.vii.1971 (Pronk) (RNH, Leiden). Sweden: 1 9, Upl., Vallentuna, 17.ix.1941
(Hedqvist)(HC, Stockholm).
HOSTS. No reared material examined. Van Burgst (1919: 105) recorded jaculator as a parasite of
Trichophaga tapetzella L. (Lepidoptera: Tineidae). I have examined these specimens and found them
to be M. cespitator,
Meteorus lionotus Thomson
(Figs 30, 31,38,40)
Meteorus lionotus Thomson, 1895: 2160. LECTOTYPE tf, SWEDEN: Norland (ZI, Lund), here designated
[examined].
Meteorus ruficoloratus Fischer, 19576: 4. Holotype tf, GERMANY (WEST): Miinchen, 15.vii.1884
(Kriechbaumer) (ZSBS, Munich) [examined]. Syn. n.
There are two specimens standing above the name lionotus in Thomson's collection. I have selected
as lectotype a specimen labelled 'Norl.' (the type locality) in Thomson's handwriting; this specimen
agrees precisely with the description of lionotus. The second specimen has evidently been misplaced
and has no connection with lionotus.
I consider Fischer's placement of ruficoloratus near ictericus to be erroneous despite their
common possession of short, strongly contracted temples. The holotype of ruficoloratus has only
spurious pits on the dorsal surface of tergite one and the ventral borders of this tergite conjoined
beneath at about the mid-point of the segment, and comes well within the limits of variation of
lionotus.
9- Antennae 31- to 33-segmented, long, most segments at least twice as long as broad. Head strongly
contracted behind the eyes and the temples short, about one-third length of eye (Fig. 31). Ocelli very large,
OO = 0-5 times OD, protuberant. Eyes a little broader than high, not strongly protuberant. Clypeus strongly
protuberant. Mandibles strongly twisted, small, rather delicate. Thorax large, a little higher than long.
Mesonotum smooth, weakly rugose-punctate; notaulices weak, indistinct anteriorly. Sternaulus shallow,
rugose, the sculpture often obsolete anteriorly and posteriorly; rest of mesopleurae polished, punctate.
Propodeum generally depressed, usually with distinct carinae laterally and occasionally with a weak medial
transverse carina; usually smooth, polished anteriorly but in some specimens this area is weakly rugose.
Tergite one long with no dorsal pits; ventral borders conjoined at about mid point of segment; dorsal surface
smooth at base and longitudinally striate apically though this is sometimes obsolescent. Ovipositor about 1-5
times length of tergite one. Legs long, slender; tarsal claws with strong basal lobe. Wings large, first abscissa
of CM, longer than cua.
Colour testaceous, usually vertex around ocelli and occipital region black; sometimes propodeum, first
tergite and apex of gaster dark, occasionally the whole body dark except for the legs which are always light in
colour.
J1- Same as 9 except that ocelli and eyes slightly smaller.
MATERIAL EXAMINED
30 9, 22 '. Great Britain, Greece, Sweden.
HOSTS. Them variata (Denis & Schiffermu'ller), Them obeliscata (Hiibner) (Lepidoptera:
Geometridae) on Pinus and Them junipemta (L.) on Juniperus. In Great Britain the parasite has
two generations in April/May and in September.
38 T. HUDDLESTON
REMARKS. The extremely large ocelli and the short strongly contracted temples immediately
distinguish lionotus from any closely related species. The shape of the thorax and the weakness of
the sculpture are also characteristic of lionotus.
Meteor us longicaudis (Ratzeburg)
(Fig. 18)
Perilitus longicaudis Ratzeburg, 1848: 55. Syntypes 9, GERMANY (destroyed).
I have followed Thomson's (1895) interpretation of this species which was based on specimens
reared from the same host species as Ratzeburg's original material. The description of longicaudis,
although short, is sufficient for the species to be identified with tolerable certainty.
9. Antennae 29- to 32-segmented, flagellar segments all clearly longer than broad, flagellum gradually
tapering to apex. Head broad, strongly rounded behind eyes. Ocelli small, OO = 3 times OD. Eyes rather
small, barely convergent. Malar space about equal to basal breadth of mandible. Face not strongly
protuberant, about twice as broad as high, strongly punctate and, at least centrally, reticulate-punctate;
vertex also punctate but frons centrally polished, impunctate. Clypeus moderately convex with scattered deep
punctures and a broad polished reflexed border. Tentorial pits indistinct. Mandibles stout, slightly twisted.
Pronotum rugose in its ventral half, polished dorsolaterally. Notaulices narrow but deeply impressed
foveolate furrows. Sternaulus rugose, always broadened anteriorly into a reticulate-rugose area; rest of
mesopleurae densely punctate and, dorsally, with a rugose area beneath the wing. Propodeum strongly
rugose, the carinae strong but not easily seen. Tergite one very broad, length about 1-5 times apical breadth;
dorsal pits large, distinct; dorsal surface finely striate laterally finely rugose medially. Ovipositor very long,
4-5 times length of tergite one. Legs long, hind coxa weakly rugose dorsally; tarsal claws simple but
somewhat thickened at base, strongly curved.
Colour black; antennae, head, prothorax and legs orange-testaceous; the basal half of hind tibia lighter
yellow, almost ivory; the head is sometimes infuscated, at least in part, and tergite two is often lighter in
colour than rest of body.
$. Same as 9 except antennae longer, up to 36-segmented, all segments at least twice as long as broad;
propodeum smaller; tergite one narrower.
MATERIAL EXAMINED
27 9, 4 tf. Netherlands: 4 9, Well, 1978 (Lefeber) (RNH, Leiden); 1 9, Brisen Dantz (RNH, Leiden).
Sweden: 5 9, Thomson coll. (ZI, Lund); 16 9, 4 <$, SM. Hornso, 28.v.-21.vi.l954 (Lundblad) (5 9 in
BMNH, rest in NR, Stockholm); 1 9, Sk. His vadero, 29.iv.1973, ex Eledenoprius armatus Panzer
(Huggert) (HC, Stockholm).
HOSTS. Type-material reared from Orchesia micans (Panzer) (Coleoptera: Melandryidae).
Thomson's specimens of longicaudis which I have examined were reared from the same host-
species. I have also examined a specimen reared from Eledenoprius armatus Panzer (Coleoptera:
Tenebrionidae). The larvae of both these species of beetle live in bracket fungi on trees.
REMARKS. M. longicaudis most closely resembles obfuscatus, which also parasitizes Orchesia
micans; longicaudis is easily distinguished from obfuscatus by its much longer ovipositor, rugose
Sternaulus and longer antennae. The males of the two species are more difficult to distinguish since
the antennae of both are long and of course they lack ovipositors; the sculpture of the Sternaulus is
the principal means of separating them. M. longicaudis males generally also have slightly smaller
eyes and ocelli and rugose hind coxae, the latter are smooth, punctate in obfuscatus. M. longicaudis
also resembles salicorniae from which it may be distinguished by its longer ovipositor, by the shape
of its head and by its strong facial puncturation.
Meteorus melanostictus C apron
(Fig. 57)
Meteorus melanostictus Capron in Marshall 1887: 115. Syntypes 9, GREAT BRITAIN (lost).
Meteorus niger Lyle, 1913: 244. LECTOTYPE 9, GREAT BRITAIN: England, Hampshire, New Forest,
1 3.v. 19 1 1 (Lyle), H. syringaria (BMNH), here designated [examined]. Syn. n.
REVISION OF THE WESTERN PALAEARCTIC METEORUS 39
I have been unable to locate any of Capron's specimens of melanostictus, I have therefore based my
interpretation of this species on a female standing above the name melanostictus in Marshall's
collection. This specimen agrees precisely with Capron's description. There are two males also from
Marshall's collection placed with the female, one of them bearing a label 'melanostictus C apron' in
Marshall's handwriting. These males are not conspecific with the female and I consider them to be
melanic specimens of M. gyrator. These specimens do agree with Marshall's statements on what he
evidently mistakenly considered to be the male of melanostictus. Lyle had completely the wrong
concept ofmelanostictus, the specimens from his collection named as that species are lionotus and his
description and host records for melanostictus also apply to lionotus. Lyle (1913) then described
specimens of the true melanostictus as a new species, niger. There are four specimens (2 <$, 2 9) of
niger in Lyle's collection labelled 'type' by Lyle; I have selected the best-preserved female as
lectotype.
9- Antennae 25- to 26-segmented, all flagellar segments at least twice as long as broad. Head strongly
contracted behind eyes, rather flat between ocelli and occipital carina; eyes in dorsal view about twice as long
as temples; frons with a blunt tubercle medially, in front of the fore ocellus. Ocelli large, OO= 1-0-1-5 times
OD. Eyes large, protuberant, moderately convergent. Malar space short, distinctly less than half the basal
breadth of mandible. Face about as wide as high, not strongly protuberant, smooth, punctate, with some
weak rugosity. Clypeus strongly protuberant with scattered large punctures. Tentorial pits deep. Mandibles
strongly twisted. Pronotum projecting but little in front of the mesonotum, laterally smooth with a little
rugosity medially. Notaulices thin, foveolate, coalescing posteriorly into a rugose area. Sternaulus thin,
foveolate, often obsolete posteriorly; rest of mesopleurae smooth, punctate; subalar prominence strongly
raised, carinate. Propodeum short, without distinct carinae except for a strongly raised medial transverse
carina which divides the propodeum into dorsal and posterior faces, the sculpture rugose, generally weak and
never reticulate. Tergite one rather stout with distinct dorsal pits and large glymmae; ventral borders of
tergite widely separated; dorsal surface coarsely striate. Ovipositor short, about 1-5 times length of tergite
one. Legs short; hind coxa smooth, punctate; tarsal claws with a distinct basal lobe.
Colour black; face, clypeus, orbits, genae, prothorax, tegulae and legs yellow.
$. Same as female except for differences in terminalia.
MATERIAL EXAMINED
21 9, 6 $. Great Britain: 20 9, 6 tf, England, Hampshire, New Forest (Lyle), ex H. syringaria (BMNH); 1
9, Marshall coll. (BMNH).
HOSTS. Apeira syringaria (L.),Ennomos quercinaria (Hufnagel) (Lepidoptera: Geometridae).
Meteorus micropterus (Haliday)
(Fig. 1)
Perilitus (Meteorus) micropterus Haliday, 1835: 27. LECTOTYPE 9, IRELAND: 'British, Haliday, 20.2.82'
(NMI, Dublin), here designated [examined].
I have chosen and labelled as lectotype the specimen selected as '9 type' by Stelfox in 1948
(unpublished Ms). The specimen is glued on a rectangular piece of card and it is by far the best-
preserved specimen of Haliday's material of this species. It agrees precisely with Haliday's
description though it is rather faded in colour.
9- Antennae 23- to 26-segmented, short, thick; segments obconical and except for the first four flagellar
segments only as long as broad; only first segment of flagellum as much as twice as long as broad. Head with
temples strongly rounded, rather square in face view. Ocelli small, OO = 3 times OD. Eyes small, inner orbits
parallel; length of malar space at least equal to basal breadth of mandible. Face protuberant, punctate,
projecting in front of frons which is depressed. Clypeus broad, smooth, punctured, distinctly divided from the
face. Tentorial pits large, deep. Mandibles moderately twisted, large, stout. Pronotum slightly expanded,
projecting strongly in front of and to the sides of the mesonotum, forming a 'collar'. Mesonotum rather
narrow, closely punctate and very hairy. Mesopleurae mostly smooth with scattered punctures. Sternaulus
foveolate. Propodeum reticulate-rugose, without distinct carinae. Tergite one smooth, shining, with
obsolescent longitudinal striate sculpture posterolaterally; ventral borders fused beneath for at least the
proximal half of the segment. Ovipositor twice length of tergite one, straight. Legs stout; hind coxae rugose
dorsally; tarsal claws without a lobe but strongly swollen at base. Wings short, narrow, usually infumate.
40 T. HUDDLESTON
Colour brownish black; legs testaceous, most usually with the hind coxa dark and frequently also the hind
femur, often tergite two, sometimes side of pronotum and occasionally base of flagellum lighter in colour.
<$. Same as the 9 except that eyes smaller; antennae 24- to 28-segmented, the segments longer, the
flagellum distinctly thickened in its proximal half and gradually tapering to apex and the colour darker; mid
and hind coxae usually darker, sometimes black, this colour change may also affect the rest of the mid and
hind legs so that occasionally the mid and hind legs are almost completely black, only the tarsi retaining a
lighter tinge.
MATERIAL EXAMINED
38 9' 46 (5*. Germany, Great Britain, Ireland.
HOSTS. Hepialus humuli (L.), Hepialus fusconebulosa (De Geer) (Lepidoptera: Hepialidae). This
species evidently uses as hosts larvae living at the base of dense herbage such as grass tussocks.
This correlates with the sturdy build and shortened appendages of the parasite which will enable it
the better to push its way into suitable host-habitats.
REMARKS. M. micropterus bears a superficial resemblance to abdominator and to consimilis. They
all are dark, heavily sculptured insects with short dark wings and short thick legs and antennae.
Both abdominator and consimilis, however, have dorsal pits in tergite one, the ventral borders of
which are widely separated, in addition consimilis has larger ocelli, strongly twisted mandibles and
a long malar space, and abdominator a down-curved ovipositor and a broad reticulate-rugose
sternaulus.
Meteor us nixoni sp. n.
(Figs 8, 12)
9- Antennae 43 -segmented, long, all segments at least longer than broad. Head broad, contracted behind eyes
but not strongly so; temples shorter than eyes. Ocelli small, OO = 2-5 times OD. Eyes small, protuberant,
very slightly convergent. Malar space slightly shorter than basal breadth of mandible. Face about twice as
broad as high, not strongly protuberant, centrally reticulate-rugose, laterally reticulate-punctate. Clypeus
broad, only slightly narrower than face, not strongly protuberant but deeply impressed at the junction with
face and with a strongly reflexed apical margin, shining, sparsely punctured; clypeal hair conspicuously
longer than facial hair. Tentorial pits deep. Mandibles large, not twisted, with conspicuous reticulate-
foveolate sculpture on basal half. Pronotum strongly rugose. Mesonotum rather narrow, densely almost
reticulately punctate, especially on the central lobe; notaulices thin, deeply impressed. Sternaulus reticulate-
rugose; rest of mesopleurae polished, punctate except for a subalar patch of rugosity. Propodeum with basal
and medial transverse carinae and a central longitudinal carina, with rugose sculpture between. Tergite one
rather broad at base, apically only 2-5 times as broad as at narrowest point, the spiracles distinctly before
mid-point; dorsal pits large, deep, behind dorsal pits strongly longitudinally strigose. Tergite two with distinct
thyridia and a trace of longitudinal strigose sculpture. Ovipositor long, about 4 times length of tergite one.
Legs long; hind coxa shining, punctured but not densely so; hind femur densely reticulate-punctate; hind tibia
swollen, though not as broad as femur, and narrow at base; tarsal claws thick, strongly curved and strongly
swollen at base, almost with a basal lobe. Wings large infumate.
Colour piceous; mandibles, palps, fore and mid legs testaceous, hind coxa and femur brown, tibia black
except at the base which is light yellow, hind tarsi black except at base and apex which are light yellow; malar
space and clypeus slightly lighter in colour than rest of head.
(5*. Unknown.
MATERIAL EXAMINED
Holotype 9, Austria: Tyrol, Oberau, vii.1938 (Nixon) (BMNH).
HOSTS. No reared material examined.
REMARKS. The characteristics of tergite one proclaim the close relationship of nixoni with corax
and sulcatus. Although the sculpture on tergite two, which is strongly developed in the two latter
species, is largely obsolete in nixoni, traces of it are discernible and thyridia are well developed in all
three species. The characters given in the key to species serve amply to distinguish the three species.
I name this species after Dr G. E. J. Nixon in appreciation of his contribution to the under-
standing of Hymenoptera.
REVISION OF THE WESTERN PALAEARCTIC METEOR US 4 1
Meteorus obfuscatus (Nees von Esenbeck)
(Fig. 22)
Bracon obfuscatus Nees von Esenbeck, 18 1 1 : 22. Syntypes, GERMANY (destroyed).
Zele thoracicus Curtis, 1832: folio 415. Syntypes, GREAT BRITAIN: England, Regent's Park (NMV,
Melbourne) [not examined]. [Synonymized by Haliday, 1835: 31.]
Perilitus formosus Wesmael, 1835: 36. LECTOTYPE 9, BELGIUM: Liege, coll. Wesmael (IRSNB, Brussels),
here designated [examined], [Synonymized by Marshall, 1887: 106.]
Meteorus fodori Papp, 1973: 3. Holotype 9, YUGOSLAVIA: Montenegro, Bjela Gora, Grahovo, 1929 (Fodor)
(HNHM, Budapest) [examined]. Syn. n.
Wesmael possessed four specimens of formosus. Two specimens presently stand in his collection
above this name; both are labelled 'Perilitus formosus mihi' in Wesmael's handwriting. The two
specimens are female, are conspecific and agree with Wesmael's description of formosus; I have
selected the better preserved of them as lectotype.
The holotype of M. fodori is rather smaller than the average specimens of obfuscatus but is
within the limits of variation of that species.
9- Antennae short, 23- to 29-segmented, the basal segments of the flagellum not more than twice as long as
broad, often shorter; the segments in the apical half of the flagellum only as long as broad. Head behind eyes
contracted, rounded; temples narrower than breadth of head across eyes, slightly shorter than eyes. Ocelli
small, OO = 2- 5 times OD. Eyes large, protuberant, slightly convergent (Fig. 22). Malar space short, not
more than half basal breadth of mandible. Face about 1-5 times as broad as high, not strongly protuberant,
reticulate-punctate, often weakly rugose beneath the antennal sockets, rarely almost completely rugose.
Clypeus protuberant, distinctly divided from face, slightly narrower than face, polished and with scattered
large punctures, sometimes with a slight medial impression in the apical border. Tentorial pits small.
Mandibles stout, moderately twisted. Pronotum projecting slightly in front of mesoscutum, laterally smooth
or at most weakly rugose. Notaulices narrow, foveolate; rest of mesonotum finely punctate. Sternaulus
deeply impressed, foveolate, occasionally with a few rugae or foveae immediately below; rest of mesopleurae
smooth, punctured except for a rugose-foveolate area beneath the base of the forewing. Propodeum with
carinae distinct, occasionally smooth between but usually with more or less well-developed rugosity. Tergite
one stout with distinct dorsal pits; generally somewhat raised medially with a medial longitudinal reticulate-
rugose band and striae laterally; occasionally tergite one is completely striate dorsally. Ovipositor twice
length of tergite one, slender, usually straight but in some specimens down-curved. Legs long, slender; hind
coxa smooth punctate; tarsal claws long, without basal lobe but slightly swollen at base.
Colour brownish black, legs always yellow except occasionally the hind tibia infuscate apically; prothorax
usually orange-testaceous and sometimes the antennae, head, mesothorax and gaster except for tergite one
also this colour, at least partly.
$. Same as 9 except that antennae longer, 29- to 32-segmented, all flagellar segments much longer than
broad; temple as long as eye, eyes smaller.
MATERIAL EXAMINED
41 9> 27 $. France, Germany, Great Britain, Hungary, Sweden, Yugoslavia.
HOSTS. The original material was reared from Orchesia micans (Panzer) (Coleoptera:
Melandryidae); I have examined specimens reared from this same host in Polyporus hispidus and
others recorded merely as 'ex Polyporus'. I have also examined two specimens 'ex cocoons made in
cells of Halomerid beetles' and one specimen recorded as a parasite of IScolytus triarmatus Eggers
(Coleoptera: Scolytidae).
REMARKS. M. obfuscatus is similar to profligator and to punctifrons but the conspicuously short
antennae, the larger eyes and ocelli and the shape of the head of obfuscatus at once distinguish that
species.
Meteorus obsoletus (Wesmael)
(Fig. 42)
Perilitus obsoletus Wesmael, 1835: 49. LECTOTYPE 9, BELGIUM: Brussels, coll. Wesmael (IRSNB,
Brussels), here designated [examined].
42 T. HUDDLESTON
Meteorus viridanae Johansson, 1964: 251. Holotype 9> SWEDEN: Borgholm, 30.vi.1954, ex larva of Tortrix
viridana (L.), coll. 13-16.vi. 1954 (Johansson) (ZI, Lund) [examined]. Syn. n.
Wesmael (1835: 50) stated that he possessed two males and a female of this species; three such
specimens still stand in his collection over the label 'obsoletus' and each is labelled 'Perilitus
obsoletus mihi' in his handwriting. The three specimens are conspecific, are in good condition and
agree precisely with Wesmael's description. The female specimen has been labelled lectotype by
Fischer though he failed to mention this selection in his redescription of the species (1970a).
9- Antennae 27- to 30-segmented, long, all segments at least 1-5 times as long as broad. Head rounded
behind eyes. Ocelli large, OO = OD (Fig. 42). Eyes not protuberant, moderately convergent. Face moderately
convex, punctate or reticulate-punctate, sometimes with a little transverse rugosity close beneath the antennal
sockets. Clypeus not strongly protuberant. Tentorial pits small, shallow. Mandibles small, strongly twisted.
Notaulices shallow rugose grooves. Sternaulus reticulate-rugose, sometimes obsolete anteriorly; rest of
mesopleurae smooth, punctate except for a subalar patch of rugosity. Propodeum evenly rounded with only a
weak medial impression posteriorly, irregularly reticulate-rugose. Tergite one slender, finely longitudinally
striate dorsally, laterally rather smooth at least at the base, without dorsal pits, ventral borders of tergite
joined from base of segment to about mid-point; spiracles slightly behind middle of segment. Ovipositor
2-0-2-5 times length of tergite one, straight, slender with no strong preapical constriction. Hind coxa
smooth, punctate without strong rugose sculpture, tarsal claws with strong basal lobe.
Colour brownish black; head, prothorax and legs testaceous, mesothorax and gaster also sometimes
testaceous; tergite one always dark except occasionally at the base.
$. Same as 9 except eyes and ocelli smaller, eyes slightly less convergent and antennal segments slightly
longer.
MATERIAL EXAMINED
30 9, 21 ^. Austria: 3 9, Partschins, Sudtirol, 1050 m, 31.viii. 1967 (Haeselbarth) (EH, Munich). Belgium:
2(5", Wesmael coll. (paralectotypes of Perilitus obsoletus Wesmael) (IRSNB, Brussels). France: 1 $, Aries sur
Tech, 14.vi.1961 (Clark) (BMNH); 3 9, 3 tf, Vaucluse, Mont Ventoux, 1978, ex Tortix viridana (L.) (du
Merle) (BMNH). Germany (West): 8 9, 7 <^, Geierlambach, Heidelbeere, vii.-viii.1970 (Haeselbarth)
(EH, Munich); 1 tf, 'Oberbiberg bei Miinchen, an Heidelbeere', 15.vii.1970 (Haeselbarth) (EH, Munich).
Great Britain: 1 9, 1 <^, England, Gloucestershire, 1932, ex Tortrix viridana (L.) (Brown) (BMNH); 1 9, E.,
Hertfordshire, Whetstone, 9.vii.l961 (Ward) (BMNH); 3 9, 4 tf, E., Berkshire, Wytham (Varley) (UM,
Oxford); 1 9, E., Hampshire, New Forest, 21.vi.1906, ex Tortrix viridana (L.) (Lyle) (BMNH). Ireland: 1 tf,
Co. Sligo, Trawallua, 24-27.vii.1933 (Nixon) (BMNH); 27.vii.1933 (Stelfox) (USNM, Washington).
Netherlands: 1 tf, Waarder (ZH) Oosteinde, 10-1 l.viii.1971 (van Achterberg) (RNH, Leiden). Sweden: 3 9,
Borgholm, 30. vi. 1954, ex Tortrix viridana (L.) (Johansson) (paratypes of Meteorus viridanae Johansson)
(ZI, Lund); 2 9, Rostanga. 4-6.vii.1938 (Perkins) (BMNH); 2 9, Hoor district, ll-13.vii.1938 (Perkins)
(BMNH). Turkey: 1 9, 1 tf, Bulancek, vi.1974, ex Gypsonoma dealbana (Frolich) (BMNH).
HOSTS. Tortrix viridana (L.), Gypsonoma dealbana (Frolich) (Lepidoptera: Tortricidae), larvae
feeding on oak between spun leaves or shoots.
REMARKS. This species is superficially similar to cinctellus but the closure of tergite one, the larger
ocelli and the moderate convergence of the eyes at once distinguish obsoletus.
Meteorus oculatus Ruthe
(Figs 5, 6)
Meteorus oculatus Ruthe, 1862: 23. LECTOTYPE 9, GERMANY: Ruthe coll. (BMNH), here designated
[examined].
Meteorus pachypus Schmiedeknecht, 1897: 207. LECTOTYPE 9, GERMANY: coll. Schmiedeknecht
(MNHU, Berlin), here designated [examined]. Syn. n.
I have examined three conspecific specimens labelled 'Meteorus pachypus Schmied' in
Schmiedeknecht's handwriting (9(J MNHU, Berlin', 9 AU, Wageningen). The two females agree
precisely with the description of pachypus and I regard them as syntypes of that species. The male
sex is not mentioned in the original description and the male specimen is very possibly a subsequent
capture and is therefore excluded from consideration as a syntype. The specimen in Wageningen is
REVISION OF THE WESTERN PALAEARCTIC METEORUS 43
part of the collection of van Burgst who is known to have bought material from Schmiedeknecht,
probably around 1910-1915 (Zwart, in litt.).
9. Antennae 30- to 33-segmented, rather slender, in apical half of flagellum segments only slightly longer
than broad. Head broad, strongly contracted behind eyes; eye about 3 times length of temple in dorsal view.
Ocelli small, OO about 2-5 times OD. Eyes very large, protuberant, strongly convergent. Malar space very
short, much less than half the basal breadth of a mandible. Face about as broad as high, weakly convex,
densely punctate with some rugosity medially. Clypeus not strongly protuberant, smooth with scattered
punctures. Tentorial pits large. Mandibles slightly twisted. Prothorax stout, expanded forwards and sideways,
rather smooth laterally but weakly rugose in part. Mesothorax rather small, its breadth across the tegulae
distinctly less than breadth of head; notaulices deep, foveolate. Sternaulus deep, reticulate-rugose; rest of
mesopleurae smooth, punctate except dorsally rugose-foveolate. Propodeum rather narrow, strongly rugose
with indistinct carinae, the medial transverse carina, at the junction of the dorsal and posterior faces of the
propodeum, the most distinct. Tergite one long, slender; dorsal pits small but distinct, dorsal surface of tergite
longitudinally striate with a scalariform element medially. Ovipositor long, about 3 times length of tergite
one, straight. Outer face of hind coxa reticulate-punctate; hind tibia strongly swollen, about as thick as
femur, constricted at base. Tarsal claws simple, short, thick and strongly expanded at base.
Colour black, legs testaceous; antennae, clypeus, mandibles, pronotum ventrally and tergite two also
sometimes lighter in colour; wings infumate.
$. Same as 9 except that eyes are less strongly convergent and hind tibia not strongly swollen.
MATERIAL EXAMINED
7 9, 1 <$. Germany (West): 1 9, Ruthe coll. (BMNH); 1 9, 'Oberberg b, Munchen, an Heidelbeere',
2.vii.l969 (Haeselbarth) (EH, Munich); 1 9, Geierlambach, Heidelbeere (Haeselbarth) (EH, Munich); 1 9,
coll. Schmiedeknecht (AU, Wageningen) (paralectotype of Meteoms pachypus Schmiedeknecht); 1 <$, coll.
Schmiedeknecht (MNHU, Berlin). Hungary: 1 9, Budapest, 1926 (Biro) (HNHM, Budapest). Sweden: 1 9,
SM. Alem, Stromsrum, 1958 (Sundholm) (ZI, Lund); 1 9, So., Dalaro Malmen, viii.1976 (Quinlan &
Huddleston) (BMNH).
HOSTS. No reared material examined.
REMARKS. M. oculatus is superficially similar to graciliventris, both having very large, convergent
eyes, short temples and long ovipositors. M. graciliventris, however, has much larger ocelli, small
but distinct lobes on the tarsal claws, and temples which are shorter and more strongly contracted
than those of oculatus; in addition oculatus has a strongly swollen hind tibia which is lacking in
graciliventris.
Meteorus pallipes (Wesmael)
(Figs 50, 51)
Perilitus pallipes Wesmael, 1835: 29. LECTOTYPE 9, BELGIUM: coll. Wesmael (IRSNB, Brussels), here
designated [examined].
Meteorus nigritarsis Ruthe, 1862; 21. Holotype 9, GERMANY: Ruthe coll. (BMNH) [examined]. Syn. n.
Wesmael (1835: 30) stated that he had four males and three females of pallipes. There are
presently eight specimens in Wesmael's collection labelled 'Perilitus pallipes mini' in Wesmael's
handwriting. Four of these specimens are males of M. filator and there is a badly damaged male
specimen which is probably also filator, these cannot have been part of the syntype-series. There
remains a female and two males which agree well with Wesmael's description of pallipes and I have
designated these lectotype and paralectotypes respectively.
Ruthe possessed only a single specimen of nigritarsis. Ruthe himself noted its similarity to
pallipes and the holotype bears a label 'P. nigritarsis m. pallip. Wsm. var?' in Ruthe's handwriting.
Thomson (1895: 2152) placed nigritarsis in synonymy with pallipes, in my opinion correctly.
Schmiedeknecht (1897: 184), however, keyed the two species separately, remarking that they
appear to be related and referring to Thomson's action in suppressing nigritarsis.
9. Antennae 26- to 31-segmented, all flagellar segments at least slightly longer than broad. Head strongly
rounded behind eyes (Fig. 51). Ocelli large, OO = 2 times OD. Eyes strongly convergent. Malar space short,
about half basal breadth of mandible. Face not strongly protuberant but generally with a slightly raised
44 T. HUDDLESTON
medial longitudinal area; smooth, punctate. Tentorial pits deep. Mandibles stout, moderately twisted.
Pronotum laterally smooth, punctate, with but few rugae. Notaulices thin, foveolate; rest of mesonotum
smooth, minutely punctured. Sternaulus foveolate, occasionally anteriorly with a small reticulate-foveolate
area beneath, often obsolescent posteriorly; rest of mesopleuron smooth with minute punctures. Propodeum
rugose, the carinae often indistinct except for the medial transverse carina which is always strongly
developed. Tergite one stout with distinct dorsal pits and glymmae; dorsally generally strongly longitudinally
striate, occasionally reticulate medially. Ovipositor 2-5-3-0 times length of tergite one. Hind coxa smooth,
punctate; tarsal claws with distinct basal lobe.
Colour black, legs always yellow; head often yellow, at least in part, and gaster occasionally marked with
yellow.
tf. Same as 9 except antennae longer, eyes not strongly convergent.
MATERIAL EXAMINED
38 9- Great Britain, Ireland, Netherlands, Sweden.
HOSTS. No reared material examined.
REMARKS. Morphologically pallipes and ictericus are closely similar; the main points of distinction
between them are to be found in the shape of the head and in the convergence of the eyes. Indeed, it
is doubtful whether headless specimens of the two species could be distinguished. M. pallipes is a
conspicuously smooth species; the face, the pronotum, the mesopleuron and the hind coxa all lack
strong sculpture; the sculpture of the sternaulus is sometimes obsolescent and occasionally
obsolete.
Meteorm prqfligator (Haliday)
(Fig. 24)
Perilitus (Meteorus) prqfligator Haliday, 1835: 33. Syntypes 9, IRELAND (lost).
My interpretation of this species is based on material from A. W. Stelfox's comprehensive
collection of Irish insects. The specimens named as prqfligator by Stelfox agree precisely with the
description given by Haliday and I have no doubt that Stelfox's interpretation of the species is the
correct one. Marshall (1887) appears to have had a different interpretation of prqfligator but the
only specimen I have seen so named by Marshall agrees neither with Haliday 's description nor with
Marshall's statements on the species.
9- Antennae 20- to 22-segmented, all segments of flagellum at least slightly longer than broad. Head rounded
behind eyes, temple about equal in length to eye viewed dorsally. Ocelli small, OO = 3 times OD. Eyes small,
not strongly protuberant, slightly convergent. Malar space slightly shorter than basal breadth of mandible.
Face 1-5-2-0 times as broad as high, smooth, punctate, not strongly protuberant. Clypeus narrower than
face, almost flat, its apical border weakly notched medially and usually with a weak vertical impression
behind the notch. Tentorial pits distinct. Mandibles short, stout and moderately twisted. Prothorax not
strongly projecting in front of the mesonotum, laterally rather smooth with only scattered weak rugae and a
few large punctures. Mesothorax short, notaulices thin, foveolate. Sternaulus foveolate, sometimes rather
broadened in its anterior half, becoming reticulate-foveolate. Mesopleuron reticulate-rugose dorsally,
otherwise smooth, punctate. Propodeum with weak carinae between which there is weak rugose sculpture.
Tergite one with large distinct dorsal pits, dorsal surface generally reticulate-rugose with a few longitudinal
striae laterally. Ovipositor about twice length of tergite one, slender. Legs slender; hind coxa never strongly
rugose but often with a trace of obsolescent rugosity at the base and on the outer surface. Tarsal claws not
lobed but somewhat swollen basally.
Colour piceous; antennae in basal half, clypeus, mandibles and usually tergite two reddish testaceous;
tegulae and legs yellow.
$. No material examined.
MATERIAL EXAMINED
30 9. Austria: 1 9, Sudtirol, Partschins, 750 m, l.ix.1967 (HaeselbartK) (EH, Munich). Great Britain: 3
9, England, Northamptonshire, Spratton, ix.1975 (Gauld) (BMNH); 1 9, E., Kent, Bexley, 16.V.1937 (Ford)
(BMNH); 1 9, E., Devon, Torquay, viii.1929 (Nixon) (BMNH). Ireland: 1 9, Co. Wicklow, Cloughlaague,
2.ix.l953 (Stelfox) (USNM, Washington); 19, Co. Wi., Deputy's Pass, 4.ix.l932 (Stelfox) (USNM,
Washington); 1 9, Co. Kildare, Royal Canal, 10.ix.1944 (Stelfox) (USNM, Washington); 1 9, Co. Dublin,
REVISION OF THE WESTERN PALAEARCTIC METEOR US 45
Glendhu, viii.1933, em. 24.iii.1934, ex Cis boleti (Scopoli) (Stelfox) (USNM, Washington). Netherlands: 19
9, ZH, Waarder, Oosteinde (6 9, viii.1973; 1 9, vi.1973; 6 9, viii.1974; 5 9, viii.1975; 1 9, ix.1975) (van
Achterberg) (RNH, Leiden); 1 9, Wijster (Dr.), 27.vii.-9.viii. 1973 (van Achterberg) (RNH, Leiden).
HOST. Cis boleti (Scopoli) (Coleoptera: Cisidae); this species of Meteorus is evidently parasitic
upon the larvae of fungivorous Coleoptera.
Meteorus pulchricornis (Wesmael)
(Fig. 36)
Perilitus pulchricornis Wesmael, 1835: 42. Lectotype 9, BELGIUM: Brussels, coll. Wesmael (IRSNB,
Brussels), designated by Marsh (1979) [examined].
Meteorus striatus Thomson, 1895: 2157. LECTOTYPE 9, SWEDEN: Skane, Palsjo (ZI, Lund), here
designated [examined]. Syn. n.
Meteorus thomsoni Marshall, 1899: 301. [Unnecessary replacement nanie for pulchricornis Wesmael sensu
Thomson.]
Meteorus japonicus Ashmead, 1906: 190. Lectotype 9> JAPAN: Gifu, viii.1902 (USNM, Washington),
designated and synonymized by Marsh (1979) [not examined].
Meteorus nipponensis Viereck, 1912: 624. Holotype 9> JAPAN (USNM, Washington) [not examined].
[Synonymized with japonicus by Watanabe, 1939.]
Meteorus macedonicus Fischer, 1957a: 104. Holotype 9, YUGOSLAVIA: Macedonia, Treskaslucht (NHM,
Vienna) [examined]. Syn. n.
Meteorus graejfei Fischer, 1957a: 107. Holotype tf, ITALY: 'Triest' (NHM, Vienna) [not examined].
[Synonymized with macedonicus by Fischer, 19706: 287.]
Meteorus tuberculifer Fischer, 1957a: 108. Holotype 9, ITALY: Trieste Kiistenland, coll. Graeffe (NHM,
Vienna) [examined]. Syn. n.
Wesmael (1835: 43) stated that he possessed one female and three males of pulchricornis; four
such specimens presently stand in Wesmael's collection above this name. All are conspecific and
agree well with Wesmael's description of pulchricornis and each bears a label 'Perilitus
pulchricornis mini' in Wesmael's handwriting. Marsh (1979) has selected the female specimen as
lectotype. In Thomson's collection six conspecific specimens stand above the name striatus; they
agree well with Thomson's description of that species. Unfortunately the lectotype, the only
specimen from the type-locality, is headless. Marshall (1899: 301) evidently mistook Thomson's
concept of pulchricornis and proposed for it a replacement name. In Thomson's collection,
however, there are seven conspecific specimens standing above the name pulchricornis which agree
with Thomson's description of the species and which are certainly conspecific with Wesmael's
series of pulchricornis. The aciculate sculpture on tergite two in macedonicus led Fischer to
postulate a relationship between this species, corax and sulcatus. M. macedonicus, however, shares
none of the other characteristics which distinguish both corax and sulcatus and I do not believe
macedonicus to be at all closely related to these species. Furthermore I have seen a few specimens
ofgyrator and versicolor which have aciculate sculpture on tergite two to a greater or lesser extent.
I therefore regard this as a fortuitous sculptural aberration of no taxonomic significance. Mason
(1974: 240) reached a similar conclusion with regard to Eubazus Nees von Esenbeck. When the
aciculation of tergite two of macedonicus is disregarded the species falls well within the limits of
variation of pulchricornis. M. tuberculifer is distinguished chiefly by the possession of large
spiracular tubercles upon tergite one. Such tubercles, however, occur occasionally in several species
of Meteorus and are developed to a lesser extent in other specimens of those species. The
possession of prominent spiracular tubercles therefore appears to me to be fortuitous and of no
value as a taxonomic character. Apart from the tubercles, tuberculifer is structurally indis-
tinguishable from typical specimens of pulchricornis.
M. pulchricornis was synonymized with gyrator by Fischer (1970ft: 263) but the two species are
abundantly distinct and neither Tobias (1976) nor Marsh (1979) has accepted this synonymy.
9- Antennae 29- to 33-segmented, long, slender; all flagellar segments distinctly longer than broad. Head
contracted behind eyes (Fig. 36), length of eye about twice length of temple in dorsal view. Ocelli large,
OO= 1-5 times OD. Eyes large, protuberant, moderately convergent. Malar space slightly shorter than
46 T. HUDDLESTON
basal breadth of mandible. Face not strongly protuberant but slightly raised medially, the raised area
finely transversely rugose, rest of face smooth, punctate. Clypeus strongly protuberant, evenly convex,
finely and densely rugose-punctate with a dense pile of erect hairs. Mandibles small, delicate and strongly
twisted. Pronotum laterally rugose. Notaulices foveolate, broadened and rugose anteriorly, coalescing
posteriorly into a reticulate-rugose area. Sternaulus rugose, often becoming reticulate-rugose anteriorly but
never broadened. Rest of mesopleuron polished, punctate except dorsally where it is reticulate-rugose.
Propodeum strongly reticulate-rugose, without distinct carinae. Tergite one longitudinally striate, usually
with no dorsal pits but with indications of pits in the sculpture of the tergite; in these specimens the ventral
borders of the tergite are in contact for a short distance in the mid part of the segment. Tergite one smooth
laterally, sometimes with a few rugae but never with glymmae. Ovipositor 1-5-2-0 times length of tergite
one. Legs long, slender; hind coxa generally completely rugose, this sculpture always fine, never reticulate
though sometimes transverse. Tarsal claws with a strong basal lobe.
Colour yellow except for the notaulices, the sternauli, the subalar area of the mesopleurae, the propodeum
and tergite one which are reddish brown; the tip of the gaster is often darkened and sometimes the preceding
tergites. Completely pale specimens occasionally occur.
<5\ Same as 9 except antennae slightly longer; face occasionally strongly raised medially; propodeum
smaller, more depressed and with greater variation in sculpture.
MATERIAL EXAMINED
96 9, 85 J1. Cyprus, France, Germany, Great Britain, Hungary, Ireland, Japan, Netherlands, Poland,
Portugal, Spain, Sweden, Switzerland, Turkey.
HOSTS. Lycophotia porphyrea (Denis & Schiffermuller), Eupsilia transversa (Hufnagel)
(Lepidoptera: Noctuidae); Lymantria dispar (L.) (Lepidoptera: Lymantriidae); Operophtera
brumata (L.), Agriopis leucophaearia (Denis & Schiffermuller), Agriopis aurantiaria (Hiibner),
Eupithecia nanata (Hiibner) (Lepidoptera: Geometridae); Thecla betulae (L.) (Lepidoptera:
Lycaenidae); Nola cuculatella (L.) (Lepidoptera: Nolidae); Poecilocampa populi (L.) (Lepidoptera:
Lasiocampidae); Charaxes jasius jasius L. (Lepidoptera: Nymphalidae).
Many other hosts of M. pulchricornis have been recorded in the literature, some of them of
considerable economic importance (see Marsh, 1979). The species is evidently catholic in its choice
of hosts, attacking caterpillars of species from diverse groups of Lepidoptera.
REMARKS. The densely punctate clypeus with its dense erect vestiture of hairs make pulchricornis
a distinctive species which is likely to be confused only with abscissus. The differentiation of the
two species is discussed under abscissus.
Meteorus puncttfrons Thomson
(Fig. 23)
Meteorus punctifrons Thomson, 1895: 2166. Holotype 9, SWEDEN: Areskutan i Jemtland (ZI, Lund)
[examined].
There is one specimen under this name in Thomson's collection; it bears a locality label "Are." (=
Areskutan, the type-locality) in Thomson's handwriting. Thomson stated that he had one specimen
and the specimen labelled punctifrons agrees well with the description of that species and may
therefore be regarded as the holotype. The specimen is in rather poor condition, the abdomen and
forewings have become detached, the abdomen has been glued onto the locality label and one
forewing on a small piece of celluloid attached to the pin, the second forewing is missing.
9- Antennae 26- to 27-segmented. Head subcubic. Ocelli very small OO = 4 times OD. Eyes small, only
slightly convergent. Face about twice as broad as high; frons with a short groove in front of fore ocellus and
between ocelli and eye with a patch of large punctures separated at most by a distance equal to their diameter.
Mandible stout, slightly twisted. Pronotum very finely reticulate-rugose ventrally. Mesonotum with scattered
large punctures, notaulices strongly impressed, foveolate; sternaulus foveolate. Propodeum short, depressed,
weakly rugose dorsally, becoming reticulate-rugose on the sides with distinct carinae, excavate apically.
Tergite one short, wide, with small but distinct dorsal pits and strong dorsolateral carinae, tergite laterally
longitudinally striate with a reticulate-rugose element medially. Ovipositor short, about 1 • 5 times length of
tergite one, down-curved. Legs very long, slender; hind coxa only slightly rugose at the base; tarsal claws
long, not lobed but slightly swollen at base.
REVISION OF THE WESTERN PALAEARCTIC METEORUS 47
Colour brownish black; base of antennae, legs, apex of tergite one, tergite two and base of tergite three
testaceous though in some specimens these parts are dark brown and the hind coxae black.
(5*. Same as 9-
MATERIAL EXAMINED
6 9, 3 tf, France: 2 tf, 3 9, Lorris Forest (Loiret), viii.-ix.1978 (Herard) (2 9, 1 tf, BMNH; 1 9, 1 tf
USDA, Sevres). Sweden: 1 9, Sk. Ahus 18.vii.1958 (Hedqvist); 1 9, Bl. Sjoarp, 6.viii.l957 (Ehnstrom); 1 tf,
Og Andebol Mosstorp 2.V.1956 (Hedqvisf) (HC, Stockholm); 1 9, Vb. Edefors, Harads, ix.1935 (BMNH).
HOST. Corticeus longulus (Gyllenhal) (Coleoptera: Tenebrionidae). The larvae of this species of
beetle are scavengers in the burrows of Scolytids in conifers. It is probable that they are sometimes
also facultative predators on the Scolytid larvae when these are present.
The French specimens examined came from logs of Pinus sylvestris which were infested with
Scolytidae (Coleoptera).
REMARKS. This is a small, slender species which is rare, at least in collections; it is easily
recognized by the characteristic frontal puncturation, the subcubic head and the minute ocelli.
Meteorus rubens (Nees von Esenbeck)
(Fig. 37)
Bracon rubens Nees von Esenbeck, 181 1: 22. Syntypes 9, GERMANY (lost).
Perilitus leviventris Wesmael, 1835: 46. LECTOTYPE 9, BELGIUM: Brussels, coll. Wesmael (IRSNB,
Brussels), here designated [examined]. [Synonymized by Fischer, 19706: 260.]
Meteorus islandicus Ruthe, 1859: 317. Syntypes, ICELAND: Staudinger coll. (NM, Vienna) [examined].
[Synonymized by Roman, 1917: 4.]
Meteorus medianus Ruthe, 1862: 53. LECTOTYPE 9, GERMANY: Ruthe coll. (BMNH), here designated
[examined]. [Synonymized with leviventris by Marshall, 1887: 126.]
Meteorus scutatus Costa, 1884: 172. Holotype 9, ITALY: Oristano (MZ, Naples) [examined]. Syn. n.
Meteorus heteroneurus Thomson, 1895: 2158. Holotype 9, SWEDEN: Vestergothland (ZI, Lund) [examined].
[Synonymized by Fischer, 19706: 260.]
Meteorus szechuanensis Fahringer, 1935: 11. LECTOTYPE 9, CHINA: No Szechuan (NR, Stockholm),
here designated [examined]. Syn. n.
Meteorus mesopotamicus Fischer, 1957a: 105. Holotype 9, IRAQ: Mosul (NM, Vienna) [examined]. Syn. n.
Wesmael (1835: 47) stated that he possessed two specimens of leviventris; in his collection there
now stands a single specimen above the name. This specimen, a female, is labelled 'Perilitus
leviventris mini' in Wesmael's handwriting and it agrees perfectly with the description of leviventris,
I have therefore chosen it as lectotype.
Ruthe (1859: 317) stated that he had before him 6 tf and 8 9 of islandicus collected in Iceland
by Staudinger. Six specimens (5 tf, 1 9) presently stand above the name islandicus in Staudinger's
collection. These specimens are conspecific, they agree with Ruthe's description of islandicus and
they are within the limits of variation of rubens.
There is one specimen standing above the label 'heteroneurus' in Thomson's collection; it agrees
precisely with Thomson's description of the species and bears a label ' VG' (= Vestergothland, the
published locality). I therefore regard this specimen as the holotype of heteroneurus. It comes well
within the limits of variation of rubens.
There are two specimens labelled 'szechuanensis n. sp. Type' in Fahringer's handwriting in the
collection of the Riksmuseet, Stockholm, each also bearing the published locality data. Both
specimens are conspecific, agree with Fahringer's description of the species and are within the
limits of variation of rubens. I have chosen as lectotype the specimen bearing a small square ticket
marked 27'5, the second specimen, labelled 25'5, as paralectotype.
9- Antennae short, 24- to 28-segmented; the segments in the distal half of the flagellum often only as long as
broad but sometimes longer than this. Head behind the eyes straight for a short distance, then strongly
contracted. Ocelli large, OO= 1-0-1-5 times OD. Eyes only slightly convergent. Malar space slightly less
than basal breadth of mandible. Face a little less than twice as wide as high, protuberant with a medial
longitudinal raised area which is finely transversely rugose, in more heavily sculptured specimens the rugosity
48 T. HUDDLESTON
extends onto the lateral areas of the face which are normally smooth, punctate. Clypeus protuberant,
transverse, medially reticulate-punctate, laterally finely transversely rugose. Tentorial pits large, distinct.
Mandibles long, moderately twisted, the upper tooth generally rather long. Pronotum laterally rugose.
Mesonotum densely punctate, the punctures large and often forming a reticulate pattern, especially on the
central lobe of the mesonotum. Notaulices short, foveolate, broadened anteriorly and posteriorly, coalescing
into a reticulate-rugose area. Sternaulus a shallow rugose furrow; rest of mesopleuron smooth, punctate.
Propodeum rugose and generally with only a medial longitudinal carina distinct though sometimes, in less
heavily sculptured specimens, the basal and medial transverse carinae can be differentiated from the rugae.
Tergite one generally smooth at the base and with obsolescent striae distally, occasionally almost completely
smooth. Ventral borders of tergite one conjoined in the midpart of the segment; there is some variation in the
extent of this conjunction but it never extends to the base of the segment. Ovipositor twice length of tergite
one, thick, strongly swollen at the base and strongly narrowed in the apical sixth. Hind coxa smooth,
punctate, sometimes with weak rugosity dorsally at the base. Tarsal claws never strongly bent and usually
only swollen at the base but sometimes the swollen base is demarcated as a lobe and occasionally this lobe is
dentate, particularly on the claws of. the forelegs.
Colour varies from completely black to completely yellow with all intermediate stages represented, none of
which is completely characteristic of the species.
$. Same as 9 except antennae longer, 26- to 30-segmented, all flagellar segments distinctly longer than
broad; the propodeum is more depressed, often less strongly rugose and with the carinae more distinct.
MATERIAL EXAMINED
354 9, 144 g. Algeria, Bulgaria, Cyprus, Denmark, Egypt, France, Germany, Great Britain, Hungary,
Iceland, Ireland, Israel, Japan, Mongolia, 'Palestine', Sweden, Turkey.
HOSTS. Agrotis ipsilon (Hufnagel), Agrotis exclamationis (L.), Mamestra brassicae (L.),
Spodoptera exigua (Hiibner), Noctua pronuba (L.) (Lepidoptera: Noctuidae); Cynthia cardui (L.)
(Lepidoptera: Nymphalidae); Idaea muricata (Hufnagel) (Lepidoptera: Geometridae).
Muesebeck (1923: 32) pointed out that rubens [as vulgaris (Cresson)] is 'an important parasite
of the cutworm type of Noctuid larva'.
REMARKS. M. rubens is a common and widespread species. It is known to be at least Holarctic in
distribution and it probably also occurs in other zoogeographical regions. M. rubens belongs to a
group of species (heliophilus, lionotus, versicolor, pulchricornis, gyrator and unicolor) character-
ized by the possession of relatively short, thick ovipositors which are generally distinctly narrowed
shortly before the apex. M. rubens is distinguished from the closely related species chiefly by the
characteristics of head-shape, claw-shape and length of antennae. The first tergite of rubens is
generally rather smooth and sometimes completely so, and this is a useful ancillary character for
the separation of the species.
Meteorus salicorniae Schmiedeknecht
(Fig. 20)
Meteorus salicorniae Schmiedeknecht, 1897: 189. LECTOTYPE 9, GERMANY (EAST): Artern (MNHU,
Berlin), here designated [examined].
Meteorus ocellatus Watanabe, 1951: 45. Holotype 9, JAPAN: Kyushu Hikosan, 17.V.1948 (Yasumatsu) (El,
Sapporo) [not examined]. Syn. n.
I have examined a specimen from the collection of the MNHU, Berlin which bears two labels in
Schmiedeknecht's handwriting: 'Artern [the published type-locality] Meteorus n. sp./Meteorus
salicorniae Schmied. 9'. This specimen agrees well with the description of salicorniae and I have no
doubt that it is a syntype of that species. I have not been able to examine type-material of ocellatus.
The description and figures of ocellatus clearly show that is is conspecific with salicorniae. The
slightly longer malar space and the smaller ocellar diameter given for ocellatus I consider to be
infraspecific variation or artefacts resulting from differing measuring techniques.
9. Antennae 33- to 35-segmented; flagellum thick at base, strongly tapering to apex, the three basal segments
at most twice as long as broad; antenna set in shallow depression in the frons which bears concentric rugae
around the antennal base. Head slightly broader than thorax; temples strongly rounded, slightly longer than
eye in dorsal view; occiput concave (Fig. 20). Ocelli small, OO = 2-5-3-5 times OD. Eyes small, not
REVISION OF THE WESTERN PALAEARCTIC METEOR US 49
convergent, not strongly protuberant. Malar space slightly longer than basal breadth of mandible. Face
strongly protuberant, transverse, aboui twice as broad as high, smooth, minutely punctured. Clypeus
protuberant, not as broad as face. Mandibles large, not twisted. Prothorax projecting slightly in front of
mesonotum, strongly swollen laterally, largely smooth, punctate. Sternaulus very deep, foveolate, rest of
mesopleuron smooth, punctate except for a strongly rugose area beneath base of wing. Propodeum high,
rounded, with indistinct carinae, strongly reticulate-rugose. Tergite one strongly expanded apically, with
small but distinct dorsal pits, dorsally with a medial longitudinal reticulate-foveolate area and fine
longitudinal striae laterally, lateral surface of tergite strongly rugose basally. Ovipositor about 2-5 times
length of tergite one, sometimes slightly more; sheaths with erect hairs which are twice as long as a sheath is
broad.
Colour. Head, prothorax, gaster and legs reddish testaceous, gaster lightefr in shade than the rest, thorax
piceous, the mesonotum, scutellum and base of propodeum also sometimes reddish testaceous; antennae
black except for the basal two or three segments.
$. No material examined.
MATERIAL EXAMINED
Bulgaria: 1 9, Rhodopi, Sh. poljana 24.vi.1975 (Zaykov) (ZC, Plovdiv). Austria: 1 9, Sudtirol,
Partschins, 1000 m, FC, 20.vii.1966 (Haeselbarth) (EH, Munich). Hungary: 1 9, Csobanka, Oszaly,
24.vii.1973 (Papp) (HNHM, Budapest). ?Yugoslavia: 1 9, Graeffe coll. (BMNH).
HOSTS. No reared material examined.
REMARKS. The shape of the head and of the antennae of this species are most distinctive.
Meteorus sulcatus Szepligeti
(Figs 9, 13)
Meteorus sulcatus Szepligeti, 1896: 3 10. Holotype 9, YUGOSLAVIA: Croatia, Grehovica, 10. v. 1885
[published as 1882] (Biro) (HNHM, Budapest) [examined].
Meteorus insignis Muesebeck, 1939: 83. Holotype 9, GREAT BRITAIN: England, Hertfordshire, Brickett
Wood, 17.vi.1936 (Benson) (BMNH) [examined]. [Synonymized by Fischer, 1966: 395.]
Meteorus molorchi Fischer, 1966: 391. Holotype 9, AUSTRIA: 'O'sterreich, Wien XIII, ob. Tirolergarten, ex
Molorchus umbellatarum, vi. 1964 (Holzschuh)' (NHM, Vienna) [examined]. Syn. n.
The types of both sulcatus and insignis are in an excellent state of preservation and I have no doubt
that Fischer's synonymy of the two species is correct. Fischer (1966: 394) differentiated molorchi
by its lighter coloration, the slighter, longer antennal segments and the position of entry of the
recurrent vein into the submarginal cells. I consider these characteristics to have only infraspecific
value and I have been unable to find other structural characteristics by which to distinguish
molorchi.
Muesebeck's (1939) description of insignis leaves little to be added.
9- Antennae 31- to 36-segmented. Head not expanded behind eyes. Clypeus smooth, punctate. Mandibles
short, stout, strongly twisted. Propodeum strongly depressed, without a strong transverse medial carina.
Tergite two longitudinally striate, with distinct thyridia. Ovipositor 3-0-3-5 times length of tergite one.
Tarsal claws without a distinct basal lobe but swollen at the base. Second submarginal cell conspicuously
higher distally than wide (Fig. 13).
(5*. No material examined.
MATERIAL EXAMINED
9 9. Austria: 1 9, Rohrwald, Ober-Rohrbach, 14.iii.1968 (Legorsky) (EH, Munich). Great Britain: 3 9,
England, Hertfordshire, Brickett Wood, 17.vi.1936 (Benson) (paratypes of Meteorus insignis Muesebeck) (1
BMNH; 2 USNM, Washington); 1 9, same data except 8.V.1943 (BMNH). France: Aiguines, Var,
12.vi.1953 (Veyref) (MNHN, Paris). ?Germany: 2 9, no data except 'Parasit von Semanotus undatus' (EH,
Munich). Netherlands: 1 9, Wijster (Dr), l-14.vii.1972 (van Achterberg) (RNH, Leiden).
HOSTS. Semanotus undatus (L.), Molorchus umbellatarum (von Schreber) (Coleoptera:
Cerambycidae).
REMARKS. The characteristics of tergite one show sulcatus to be closely related to corax which is
also parasitic on Cerambycidae. M. sulcatus is generally a much smaller species than corax, though
it exhibits considerable variation in size, a typical feature of parasites of wood-boring beetles.
50 T. HUDDLESTON
Meteorus tabidus (Wesmael)
(Fig. 48)
Perilitus tabidus Wesmael, 1835: 32. LECTOTYPE tf, BELGIUM: Brussels, coll. Wesmael (IRSNB,
Brussels), here designated [examined].
Meteorus fascialis Ruthe, 1862: 22. Holotype 9, GERMANY. Ruthe collection (BMNH) [examined].
[Synonymized by Schmiedeknecht, 1897: 187.]
Meteorus dubius Ruthe, 1862: 27. LECTOTYPE 9, GERMANY: Ruthe collection (BMNH), here designated
[examined]. Syn. n.
Meteorus pentheri Fischer, 19706: 262 [diagnosis in key]. Holotype tf, YUGOSLAVIA: Stolac (Penther)
(NHM, Vienna) [examined]* Syn. n.
Wesmael (1835: 33) stated that he possessed one female and three males of this species. In the
Wesmael collection there are one female and two males labelled 'Perilitus tabidus mini' in
Wesmael's handwriting; they are conspecific and all agree with Wesmael's description of tabidus.
The female specimen is in poor condition, having lost its head and three legs, and I have therefore
chosen as lectotype the best preserved of the male specimens. Ruthe did not indicate what material
he had of fascialis. There is in the collection of the BMNH one damaged specimen from Ruthe's
collection labelled 'P. fascialis m.' in Ruthe's handwriting; this specimen has been labelled as type
(3C 757). It agrees with the description and I am accepting this specimen as holotype of Meteorus
fascialis. There is a female syntype from Ruthe's collection in the BMNH labelled 'P. dubius m.' in
Ruthe's handwriting; this specimen, which agrees with Ruthe's description of dubius and is clearly
within the limits of variation of tabidus, is designated as lectotype. Fischer (1970ft: 258) suppressed
dubius as a synonym of Zemiotes [Zele] caligatus Haliday. Van Achterberg (in litt.) states that in
the MNHU, Berlin there is a female caligatus identified as dubius. It seems probable that Fischer's
misinterpretation of dubius was based on this wrongly identified specimen.
9- Antennae 28- to 3 1 -segmented, thick, the penultimate five or so segments not longer than broad. Head
contracted behind eyes, not distinctly rounded, the occiput not distinctly concave. OO = 2 times OD. Eyes
large, strongly convergent. Malar space short, about half basal breadth of mandible. Face not strongly
convex, rugulose-punctate. Clypeus strongly protuberant, abruptly divided from face, polished, sparsely
punctate. Tentorial pits wide, deep, very close to orbits. Mandibles at most moderately twisted. Pronotum
heavily rugose except for a smooth band along its dorsal border. Notaulices deeply impressed foveolate
grooves which coalesce posteriorly into a reticulate-rugose area. Sternaulus broad, reticulate-rugose, rest of
mesopleurae smooth except for a dorsal rugose area. Propodeum strongly rugose, sometimes reticulate-
rugose, almost always with a distinct basal transverse carina and often with at least an indication of central
longitudinal and medial transverse carinae though neither of these is easily distinguished from the rugae.
Tergite one long, narrow, with distinct dorsal pits; generally strongly longitudinally striate and often with
some scaly-reticulate sculpture medially. Ovipositor long, 2-5-3-0 times length of tergite one, thick. Legs
long; hind coxa strongly rugose; tarsal claws with a distinct basal lobe.
Colour black; tergites 2 and 3 and legs testaceous. The face and base of the antennae sometimes lighter.
There is always a lighter-coloured spot on each orbit at the level of the ocelli (though in darker specimens this
sometimes almost disappears) and sometimes a lighter patch on the genae just behind the eye. The hind legs are
sometimes darker; the hind coxae sometimes black. Wings generally slightly embrowned.
(5s. Same as 9 except that antennae longer, 32- to 35-segmented, all segments of flagellum at least twice as
long as broad; temples longer, about equal to eyes in dorsal view; the ocelli slightly larger; the eyes not
strongly convergent; propodeum depressed, often narrower and much less strongly rugose so that the carinae
are more distinct; tergite one narrower, not so expanded apically sometimes almost linear and with a few
coarse longitudinal striae dorsally; the colour generally darker (in the darker specimens the light orbital mark
on the vertex is often reduced but it is generally still present if only as a trace).
MATERIAL EXAMINED
28 9' 40 • ?. Austria, Bulgaria, France, Germany, Great Britain, Ireland, Netherlands, Sweden,
Switzerland.
HOSTS. No reared material examined.
REMARKS. M. tabidus is very similar to qffinis. In addition to the characters given for their
separation in the key to species, tabidus has a more protuberant clypeus, the area of rugosity at the
junction of the notaulices smaller and the rugosity coarser, and it is generally a slightly more robust
REVISION OF THE WESTERN PALAEARCTIC METEOR US 5 1
insect than qffinis. The male of tabidus is exceptionally distinctive; it most resembles consimilis in
shape of head, sculpture and colouring but is distinguished by the shorter malar space, the frons not
depressed and by a generally more slender habitus.
Meteorus unicolor (Wesmael)
Perilitus unicolor Wesmael, 1835: 41. LECTOTYPE 9, BELGIUM: Brussels, coll. Wesmael (IRSNB,
Brussels), here designated [examined].
Saprotichus chinensis Holmgren, 1868: 430. Holotype 9, CHINA (NR, Stockholm) [examined]. Syn. n.
Wesmael (1835: 41) stated that he had two females and a male of this species. Two specimens
presently stand in his collection above the label 'unicolor'; they are conspecific and each bears a
label 'Perilitus unicolor mini' in Wesmael's handwriting and both agree precisely with Wesmael's
description of the species.
9. Antennae long, up to 36-segmented; all segments of flagellum distinctly longer than broad, most at least
twice as long as broad. Head contracted behind eyes; temples slightly shorter than eyes in dorsal view. Ocelli
large, OO=2 times OD. protuberant. Eyes protuberant, slightly convergent. Malar space about equal in
length to basal breadth of mandible. Face 1-5-2-0 times as broad as high, not strongly protuberant but
distinctly raised medially and there transversely rugose, laterally densely punctate. Clypeus strongly
protuberant, densely rugulose-punctate with scattered long hairs. Mandibles long, slender, strongly twisted.
Pronotum projecting but little in front of the mesonotum; coarsely rugose laterally. Notaulices foveolate,
broadened and reticulate-rugose anteriorly, coalescing posteriorly in a broad densely reticulate-rugose area.
Sternaulus broadened, strongly reticulate-rugose; rest of mesopleurae smooth, punctate except for a large
rugose area around the subalar prominence; prepectal carina strongly raised. Propodeum broad, shallowly
excavate posteriorly; coarsely reticulate-rugose and with no distinct carinae dorsally though there is
sometimes a trace of median longitudinal and transverse medial carinae. Tergite one stout with large dorsal
pits and distinct glymmae; ventral borders of tergite only narrowly separated at mid point of segment; dorsal
surface longitudinally striate. Ovipositor twice length of tergite one, thick, strongly swollen at base, straight.
Hind coxa foveolate laterally, often with a few strong rugae dorsally. Tarsal claws with a large basal lobe.
Colour testaceous, base of tergite one paler yellow.
$. Same as 9 except for differences in genitalia.
MATERIAL EXAMINED
Belgium: 1 tf, Brussels, coll. Wesmael (IRSNB, Brussels) (paralectotype of Perilitus unicolor Wesmael).
Germany (West): 1 9, Bavaria, Schliersee, 28.vii.1958 (Townes) (CNC, Ottawa). Great Britain: 1 9, Wales,
Dyfed 'Pembrokeshire', 2.vii.l973 (Ford) (MSC, Reading).
HOST. Zygaenafilipendulae (L.) (Lepidoptera: Zygaenidae).
REMARKS. I have seen only two specimens of this species apart from the type-material. The only
characters I can find for distinguishing unicolor and abscissus are those given in the key to species
and I am by no means confident that these characters would hold good in a longer series of
specimens. When more material is available for study it should be possible more certainly to show
whether unicolor and abscissus are but variants of a single species. M. unicolor has been
traditionally regarded as a testaceous form of gyrator and the two species do resemble each other;
unicolor, however, is a more robust and heavily sculptured insect but whether these differences will
prove in the long run to be valid at the specific level is open to question.
Meteorus versteolor (Wesmael)
(Figs 4 1,43)
Perilitus versicolor Wesmael, 1835: 43. Lectotype 9» BELGIUM: Charleroy (IRSNB, Brussels), designated by
Marsh (1979) [examined].
Perilitus bimaculatus Wesmael, 1835: 45. Lectotype 9, BELGIUM: Charleroy (IRSNB, Brussels), designated
by Marsh (1979) [examined]. [Synonymized by Muesebeck, 1923: 36.]
Perilitus unicolor Hartig, 1838: 254. LECTOTYPE 9. GERMANY (ZSBS, Munich), here designated
[examined]. [Junior primary homonym of unicolor Wesmael 1835: see hartigi below.]
52 T. HUDDLESTON
Perilitus brevicornis Ratzeburg, 1844: 77. Holotype 9, GERMANY: Ratzeburg collection (IP, Eberswalde)
[examined]. [Synonymized by Konigsmann, 1964: 654.]
Perilitus rugator Ratzeburg 1852: 59. LECTOTYPE 9, GERMANY: *741/ neust Br./ Peril rugator ant 30
art', Ratzeburg collection (IP, Eberswalde), here designated [examined]. Syn. n.
Meteorus decoloratus Ruthe, 1862: 48. LECTOTYPE 9, GERMANY: Ruthe coll. (BMNH), here designated
[examined]. [Synonymized by Muesebeck, 1923: 36.]
Meteorus camptolomae Watanabe, 1939: 25. Holotype 9» JAPAN: 'Okikuba, Tokyo' (El, Sapporo) [not
examined]. Syn. n.
Meteorus ikonomovi Fischer, 1959: 5. Holotype 9> YUGOSLAVIA: Dalmatien, Gravosa (NHM, Vienna)
[examined], Syn. n.
Meteorus hartigi Shenefelt 1969: 69. [Replacement name for unicolor Hartig, 1838.] Syn. n.
Wesmael (1845: 43) stated only that he had many specimens of versicolor; three specimens now
stand in Wesmael's collection above that name, all are conspecific and labelled 'Perilitus versicolor
mini' in Wesmael's handwriting. They agree well with the description of versicolor. There are two
specimens in the collection above the name bimaculatus\ Marsh has chosen as lectotype the
specimen which best fits the description of the species.
Two conspecific female specimens presently stand above the name unicolor in Hartig's
collection. One of these specimens bears a label 'unicolor n' in Hartig's handwriting and I have
chosen this specimen as the lectotype of the species. It comes within the limits of variation of
versicolor.
The principal character used by Ratzeburg to distinguish rugator was the position of entry of the
recurrent vein into the cubital (submarginal) cells. I have found this structure to be variable in most
species and consequently it is of little use as a taxonomic character. The specimens discussed by
Konigsmann (1964: 652) agree with the description of rugator and are within the limits of variation
of versicolor.
I have been able to examine only two male paratypes of camptolomae (USNM, Washington) but
these are clearly within the limits of variation of versicolor.
9- Antennae 29- to 33-segmented; flagellum thick basally, tapering to apex, all segments of flagellum
distinctly longer than broad, most more than twice as long as broad. Head broad, more or less strongly
contracted behind eyes, eyes in dorsal view 2-5-3-0 times length of temple. Ocelli large, OO = OD,
protuberant. Eyes large, protuberant, only slightly convergent. Malar space short, always less than the basal
breadth of the mandible. Face about 1-5 times as broad as high, not strongly protuberant but somewhat
raised medially; generally with transverse rugose sculpture, at least on the raised medial part, occasionally
smooth, punctate. Clypeus protuberant. Mandibles small, delicate, strongly twisted. Pronotum not projecting
before the mesonotum, laterally rugose, sometimes reticulate-rugose. Propleurae rugose-punctate. Notaulices
shallow, anteriorly broadened and reticulate-rugose, posteriorly coalescing into a broad rugose area; rest of
mesonotum punctate, often reticulate-punctate medially. Sternaulus a broad rather shallow furrow weakly
rugose and foveolate, generally broader and sometimes reticulate-foveolate medially; often obsolescent
anteriorly and posteriorly; rest of mesopleurae polished and with minute punctures; much of the sculpture of
the sternaulus is so weak that it is invisible except under the most oblique light. Propodeum short, broad,
strongly rugose, often reticulate-rugose with no distinct carinae but with a distinct medial impression
posteriorly. Tergite one long, rather slender at the base with no dorsal pits, its ventral borders conjoined from
shortly before the midpoint of the segment to its base, dorsal surface finely striate, lateral and ventral
conjoined parts smooth. Ovipositor short, 1-5-2-0 times length of tergite one, thick, strongly tapered shortly
before apex. Legs stout, hind coxa smooth, punctate, sometimes reticulate-punctate, often with weak
transverse rugae dorsally; tarsal claws strongly curved with a pronounced basal lobe.
Colour varies from almost completely testaceous to almost completely brown; specimens occur fairly
commonly in which the propodeum, tergite one except at base, hind coxa, thorax in part and head in part are
black, the rest of the body and legs testaceous; sometimes the normally testaceous parts of the body are
much paler yellow, almost ivory. The base of tergite one generally pale yellow.
$. Same as 9 except that eyes generally smaller and less protuberant. I have examined one male specimen
which is completely brownish black.
MATERIAL EXAMINED
152 9, 108 cf . Austria, Bulgaria, France, Germany, Great Britain, Hungary, Ireland, Japan, Mongolia,
Netherlands, 'Palestine,' Poland, Sweden.
REVISION OF THE WESTERN PALAEARCTIC METEOR US 53
HOSTS. Lasiocampa quercus (L.), Malacosoma neustria (L.), Dendrolimus pint (L.), Macrothylacia
rubi (L.) (Lepidoptera: Lasiocampidae); Leucoma salicis (L.), Euproctis chrysorrhoea (L.),
Dasychira pudibunda (L.) (Lepidoptera: Lymantriidae); Anarta myrtilli (L.), Lycophotia
porphyrea (Denis & Schiffermiiller) (Lepidoptera: Noctuidae); Eulithis testata (L.) (Lepidoptera:
Geometridae); Maniola jurtina (L.) (Lepidoptera: Satyridae); Thaumetopoea processioned (L.)
(Lepidoptera: Thaumetopoeidae).
REMARKS. Traditionally the principal character used for the separation of versicolor from other
species of Meteorus has been the presence of a pale yellow area at the base of the petiole (tergite
one), which strongly contrasts with the rest of the tergite which is darker. Indeed the presence of a
pale yellow band at the base of tergite one is an easy and sure way to recognize versicolor. In about
a quarter of the specimens of versicolor which I have examined, however, the yellow colour at the
base of tergite one is either absent or is so faint as to be indistinguishable from the condition of
tergite one in several other species in which this area is sometimes slightly lighter in colour (e.g.
unicolor, obsoletus}. M. versicolor is structurally very close to obsoletus', in addition to the
characters given in the key to species the pale base of tergite one is a useful means of differentiating
versicolor, also obsoletus is generally a much smaller and less robust insect than versicolor. M.
versicolor and, in particular, the testaceous form of this species bears a superficial resemblance to
several other species of Meteorus (e.g. lionotus, heliophilus, gyrator), being similar in stature, length
and shape of ovipositor, colour and sculpture. The ventral closure of tergite one, however, amply
distinguishes versicolor.
Meteorus vexator (Haliday)
(Fig. 4)
Perilitus (Meteorus) vexator Haliday, 1835: 33. Neotype 9, IRELAND: Co. Kd., R. Canal, (2), ll.vii.1943
(Stelfox) (USNM, Washington), designated by Fischer (1959: 12) [examined].
Fischer chose as neotype a specimen from Stelfox's collection which had been named as vexator by
Stelfox. This specimen agrees with Haliday's exiguous description except that it has 23 antennal
segments whereas Haliday gave 19-20 as the range; 20 is the least number of antennal segments I
have found in the material I have examined of this species. Some workers have been misled as to
the significance of the bracketed figure which appears on some of Stelfox's data labels ((2) in the
type data for vexator). Graham (pers. comm.) states that this figure refers to the number of the
collection made on the date with which it appears. Thus the neotype of vexator was captured in the
second collection which Stelfox made on 1 l.vii.1943.
9- Antennae 20- to 24-segmented, slender, all flagellar segments distinctly longer than broad. Head large,
much broader than thorax, rounded behind eyes and also more or less strongly contracted; eyes in dorsal
view at most twice length of temple. Ocelli small, OO = 2-5— 3-0 times OD. Eyes large, protuberant, very
strongly convergent. Malar space short, much less than basal breadth of mandible. Face small, about as
broad as high, not protuberant, smooth, punctate. Clypeus moderately protuberant, smooth, punctate.
Tentorial pits distinct and very close to margin of eye. Mandibles short, stout, moderately twisted. Pronotum
laterally generally with weak rugose sculpture. Notaulices narrow but distinct, foveolate. Sternaulus narrow,
foveolate, sometimes with a small punctate patch beneath; rest of mesopleurae polished except for a small
reticulate area beneath the base of the forewing. Propodeum small, distinctly carinate, generally rather
smooth between the carinae but sometimes weakly rugulose. Tergite one with distinct dorsal pits at about its
midpoint or slightly before; dorsal surface strongly rugose with at most a few weak striae laterally, the lateral
faces of the tergite strongly rugose. Ovipositor 2-5-3-0 times length of tergite one. Legs long, slender, hind
tibia not unusually swollen, narrower than femur; hind coxa generally at least slightly rugose ventro-laterally
though sometimes reticulate-punctate; tarsal claws without a basal lobe.
Colour brownish black, legs testaceous; face, clypeus, mandibles, antennae at the base, pronotum ventrally
and tergite two sometimes lighter in colour; hind coxa sometimes infuscate in part, occasionally entirely dark.
$. Same as 9 except antennae longer, up to 27-segmented; eyes less convergent and sculpture of
sternaulus and of hind coxa sometimes obsolete.
MATERIAL EXAMINED
105 9- Austria, Great Britain, Ireland, Netherlands, Sweden.
54 T. HUDDLESTON
HOST. Biphyllus lunatus (Fabricius) (Coleoptera: Biphyllidae).
REMARKS. I have examined a male specimen belonging to the series on which Morley (1912) based
his description of vexator. This specimen is certainly conspecific with vexator the interpretation of
which has been fixed by the designation of a neotype (Fischer, 1959: 12). Morley's association of
vexator with Biphyllus lunatus (Fabricius) is as yet the only indication of a probable host of the
species though I consider the evidence upon which Morley based his record to be less conclusive
than he stated.
Nomina dubia
Meteorus dejeanus (Rondani)
Perilitus dejeanus Rondani, 1874: 131. Type-material not found.
Meteorus delator (Haliday)
Perilitus (Meteorus) delator Haliday, 1835: 33. Type-material not found.
Meteorus dilutus (Ratzeburg)
Perilitus dilutus Ratzeburg 1844: 77. Type-material lost.
Meteorus effeminates Ruthe
Meteorus effeminatus Ruthe, 1862: 32.
Ruthe stated that he had two male specimens which he referred to as effeminatus. I have examined
these specimens but I have not been able to reach a conclusion about their placement.
Meteorus flaviceps (Ratzeburg)
Perilitus flaviceps Ratzeburg, 1844. 75. Type-material lost.
Meteorus gracilis (Ratzeburg)
Perilitus gracilis Ratzeburg, 1852: 58. Type-material lost.
Meteorus longicornis (Ratzeburg)
Perilitus longicornis Ratzeburg, 1844: 76. Type-material lost.
Meteorus pallidus (Nees von Esenbeck)
Bracon pallidus Nees von Esenbeck, 1812: 22. Type-material lost.
Fischer (1970Z?: 258) placed pallidus in Zemiotes which then had the rank of a subgenus in
Meteorus. Van Achterberg (1979: 387) has pointed out that Nees stated there were no dorsal pits
on tergite one of pallidus and that this precludes Fischer's placement of that species. Van
Achterberg therefore concludes that pallidus belongs in Meteorus. I have, however, been unable to
identify the species.
Meteorus pendulator (Latreille)
Ichneumon pendulator Latreille, 1799: 138. Type-material lost.
I have unfortunately been unable to interpret pendulator. Marshall (1887: 100) followed Haliday
(1835: 28) in placing pendulator and ictericus in synonymy. The series standing above the name
pendulator in Haliday's collection includes specimens of both ictericus and gyrator. The
description and figure of pendulator show it to have a shorter ovipositor and to be lighter in colour
than ictericus. M. pendulator could quite well be conspecific with gyrator but the description is not
sufficiently precise to preclude other species.
Meteorus rubriceps (Ratzeburg)
Perilitus rubriceps Ratzeburg, 1844: 75. Type-material lost.
Meteorus ruficeps (Nees von Esenbeck)
Perilitus ruficeps Nees von Esenbeck, 1834: 39. Type-material lost.
REVISION OF THE WESTERN PALAEARCTIC METEOR US 55
Meteorus rufus (DeGeer)
Ichneumon rufus DeGeer, 1778: 597. Type-material lost.
There is a label in the DeGeer collection for 'Ichneumon rufus' but there is no specimen present.
There is no pinhole in the cork above the label which indicates that no specimen was present in the
collection when it was transferred to its present cabinet in 1 845.
Meteorus stenostigma Thomson
Meteorus stenostigma Thomson, 1895: 2153. Type-material lost.
Two specimens stand in Thomson's collection above the name stenostigma. Neither of them agrees
precisely with Thomson's description and neither is from the type-locality.
Meteorus wesmaeli (Boie)
Perilitus wesmaeli Boie, 1850: 214. Type-material not found.
Acknowledgements
I wish to thank Professor B.-O. Landin and Mr R. Danielsson (Lund); Professor E. Sylven
(Stockholm) and Dr L. Hedstrom (Uppsala) for the provision of study facilities and cordial
hospitality in their respective departments during August 1976, and Mr G. Nordlander for his
hospitality on that occasion. I wish also to thank the following for the loan of material and for
much other valuable assistance: Dr C. van Achterberg (Leiden); Dr A. A. Allen (Reigate); Mr R.
Danielsson (Lund); Dr E. Diller (Munich); Dr P. Dessart (Brussels); Dr M. Fischer (Vienna); Dr
M. W. R. de V. Graham (Oxford); Dr E. Haeselbarth (Munich); Dr K.-J. Hedqvist (Stockholm);
Dr L. Hedstrom (Uppsala); Dr E. Konigsmann (Berlin); Dr P. M. Marsh (Washington); Dr W. R.
M. Mason (Ottawa); Dr J. P. O'Connor (Dublin); Professor G. Morge (Berlin); Dr J. Papp
(Budapest); Dr M. Shaw (Reading); Dr B. Sigwalt (Paris); Dr H. D. Volkart (Berne); Dr A.
Zaykov (Plovdiv); Drs K. Zwart (Wageningen) and many colleagues in the British Museum
(Natural History).
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Ruthe, J. F. 1859. Verzeichniss der von Dr Staudinger im Jahre 1856 auf Island gesammelten
Hymenopteren. Stettin, ent. Ztg 20: 305-322.
- 1862. Deutsche Braconiden. Drittes Stuck. Berl. ent. Z. 6: 1-58.
Schmiedeknecht, H. L. O. 1897. Die Braconiden-Gattung Meteorus Hal. Illte Wschr. Ent. 2: 150-154,
173-175, 184-190, 204-207, 221-223, 298-302.
Schonwiese, F. 1934. Beobachtungen und Versuche anlasslich einer Ubervermehrung von Lophyrus sertifer
Geoffr. (rufus Panz.) in Sudkarnten in den Jahren 1931' 1932. Z. angew. Ent. 21: 463-500.
Shenefelt, R. D. 1969. Hymenopterorum catalogus (nov. ed.) part 4 Braconidae 1: 1—176, Junk, 'S-
Gravenhage.
Spinola, M. 1808. Insectorum liguriae species novae an rariores, quas in agro Ligustico nuper detexit,
descripsit, et iconibus illustvit (Hymenoptera). 2: 262 pp. Genuae.
Szepligeti, G. V. 1896. Beitrage zur Kenntnis der Ungarischen Braconiden. Termeszetr. Fuz. 19: 359-386
Thomson, C. G. 1895. Bidrag till Braconidernas kannedom. Opusc. Ent. 20: 2141-2339.
Thunberg, C. P. 1822. Ichneumonidea, insecta Hymenoptera, illustrata. Mem. Akad. St. Petersburg 8:
249-281.
Tobias, V. I. 1966. Genera groupings and evolution of the subfamily Euphorinae (Hymenoptera:
Braconidae) part 2. [In Russian.] Ent. Obozr. 45: 612-633.
- 1976. Braconidae of the Caucasus. [In Russian.] Opred. Faune SSSR. 1 10: 1-286.
Viereck, H. L. 1912. Contributions to our knowledge of bees and ichneumon-flies, including the descriptions
of twenty-one new genera and fifty-seven new species of ichneumon-flies. Proc. U.S. natn. Mus. 42:
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1914. Type-species of the genera of ichneumon-flies. Bull. U.S. natn. Mus.no. 83: 186pp.
Watanabe, C. 1939a. Meteorus japonicus Ashmead, a parasite of the Gypsy Moth Lymantria dispar Linne
(Hymenoptera: Braconidae). Insecta matsum. 13: 63-65.
19396. A new species of Meteorus bred from Camptoloma interiorata Walker (Hymenoptera:
Braconidae). Insecta matsum. 14: 25-26.
1951. A new species of the genus Meteorus Haliday from Japan (Hymenoptera: Braconidae). Mushi
22 (6): 45-46.
Wesmael, C. 1835. Monographic des Braconides de Belgique. Nouv. Mem. Acad. R. sci. Bruxelles 9: 1-252.
1838. Monographic des Braconides de Belgique IV. Nouv. Mem. Acad. R. sci. Bruxelles 11: 1-166.
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London.
Index
Invalid names are in italics.
abdominator 20 brevipesalis 20
abscissus 21 brunnipes 20
adoxophyesi 34
affinis 22 camptolomae 52
albicornis 27 cespitator 24
ambiguus 24 chinensis 51
atrator 24 cinctellus 25
colon 26
bimaculatus 51 confinis 34
brevicauda 23 consimilis 27
brevicornis 52 consors 34
brevipes 20, 27 corax 28
58
crassicrus 34
decoloratus 52
dejeanus 54
delator 54
dilutus 54
dub ins 50
eadyi 29
ejfeminatus 54
ephippium 34
T. HUDDLESTON
34
fascialis 50
fasciatus 26
filator 30
flaviceps 54
fodori 41
formosus 41
fragilis 26
fuscipes 25
gracilis (Ratzeburg) 54
gracilis Ruthe 22
graciliventris 31
graeffei 45
gyrator 31
52
heliophilus 32
heteroneurus 47
hirsutipes 33
hodisensis 30
ictericus 34
ikonomovi 52
insignis 49
islandicus 47
jaculator 36
japonicus 45
laticeps 30
leviventris 47
lionotus 37
//(?uu 34
longicaudis 38
longicornis 54
lophyriphagus 34
luridus 26
macedonicus 45
medianus 47
melanostictus 38
mesopotamicus 47
micropterus 39
minutor 34
molorchi 49
mongolicus 23
monochami 28
w'ger 38
nigritarsis 43
nipponensis 45
nixoni 40
obfuscatus 41
obscurellus 36
obsoletus 41
ocellatus 48
oculatus 42
pachypus 42
pallidus 54
pallipes 43
parvulus 32
pendulator 54
pentheri 50
pleuralis 34
profligator 44
pulchricornis 45
punctifrons 46
punctiventris 22
rubens 47
rubriceps 54
ruficeps 54
ruficoloratus 37
rw/i/s 55
rugator 52
ruthei 22
salicorniae 48
scutatus 47
scutellator 31
similator 24
stenostigma 55
striatus 45
sulcatus 49
szechuanensis 47
tabidus 50
tenellus 25
tenuicornis 36
thomsoni 45
thoracicus 41
thuringiacus 23
tuberculifer 45
unicolor Hartig 5 1
unicolor (Wesmael) 51
versicolor 51
vexator 53
viridanae 42
voloscensis 22
wesmaeli 55
xanthomelas 34
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A revision of the Old World Polymorphanisini (Trichoptera : Hydropsychidae)
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Bulletin of the
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'*•<; m«<°.
A revision of the Old World
Polymorphanisini (Trichoptera:
Hydropsychidae)
P. C. Barnard
Entomology series
Vol 4 1 No 2 25 September 1 980
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World List abbreviation : Bull. Br. Mus. nat. Hist. (Ent.)
Trustees of the British Museum (Natural History), 1980
ISSN 0524-6431 Entomology series
Vol41No2pp59-106
British Museum (Natural History)
Cromwell Road
London SW7 5BD Issued 25 September 1 980
A revision of the Old World Polymorphanisini
(Trichoptera : Hy dropsychidae)
P. C. Barnard
Department of Entomology, British Museum (Natural History), Cromwell Road, London SW7
5BD
Contents
Synopsis 59
Introduction .............. 59
Classification of the Macronematinae 60
Abbreviations 60
Acknowledgements 61
Taxonomic section . . . . . . . . . . . . .61
Tribe Polymorphanisini Lestage 61
Geographical distribution 63
Biology 63
Check-list of Old World Polymorphanisini 63
Key to world genera of Polymorphanisini 64
Oestropsyche Brauer 64
Aethaloptera Brauer 66
Key to species of Aethaloptera .......... 68
Polymorphanisus Walker ............ 78
Key to species of Polymorphanisus ......... 78
The nigricornis-group ......... r .. 79
The ocw/ara-group 98
References 103
Index . .105
Synopsis
The Old World species of the tribe Polymorphanisini, with its constituent genera Oestropsyche Brauer,
Aethaloptera Brauer and Polymorphanisus Walker, are revised. One new generic and eight new specific
synonyms are established; four new species are described. Seven lectotypes and one neotype are designated.
Keys are given to the twenty-five species currently recognized.
Introduction
The Polymorphanisini are one of the most discrete tribes within the Hydropsychidae; although
they are superficially very similar to the other species of the subfamily Macronematinae, they
have an obvious diagnostic character in the absence of mouthparts. This paper deals only with
the Old World species ; the endemic Neotropical genus Synoestropsis is distinguished in the key
to genera, but most of its constituent species are dealt with in recent publications by O. S. Flint,
Jr (e.g. Flint, 1978).
The subfamily Macronematinae was revised by Ulmer (1907) and this remains the basis for
any work on this group, Ulmer's monograph being complete for all the literature up to that date.
Although many of the species of the tribe Macronematini, especially of Macronema itself, can be
readily identified by their wing pattern, the Polymorphanisini are relatively uniform in general
Bull. Br. Mus. nat. Hist. (Ent.) 41 (2): 59- 106 Issued 25 September 1980
50 P- C. BARNARD
appearance and have always been considered a difficult group to identify correctly. For example,
in his paper on the Trichoptera of Ghana, Gibbs (1973) remarks 'Because of a lack of modern
figures of male genitalia, African species of Polymorphanisus are usually in some doubt.' I hope
the present paper will remove this difficulty, but in fact a study of the genitalia has not yielded
many useful characters. The keys are based on 'external' characters, namely features of the wing
venation, coloration, thoracic markings and so on, and these, together with the known
distribution of the species, are sufficient to separate all the species in the tribe. The genitalia of
both sexes (where known) of each species are described and figured, however, because they are
often useful in confirming the identity of superficially similar species. Of the 25 species included
in this paper, the males of ten are unknown, which is another reason for the limited use of
genitalic characters.
All the drawings in this paper were made using a camera lucida attachment on a stereo-
microscope. Male genitalia preparations were examined and drawn in glycerine to avoid any
distortion due to flattening. In the females the eighth sternite, which was the only feature found
to exhibit consistent specific characters, was drawn from permanent preparations in Euparal,
deliberately flattened to facilitate comparability of outline of this curved sclerite.
Classification of the Macronematinae
The first division of the Hydropsychidae was made by Brauer (1868) who erected the 'subfamily'
Oestropsidae, for Oestropsis and Polymorphanisus, on the basis of the absence of the palps :
Macronema was retained in the Hydropsychidae sensu stricto. Later, Brauer (1875) described the
genus Phanostoma which he also placed in the Oestropsidae along with Aethaloptera, although
Phanostoma (now synonymized with Amphipsyche) has normal mouthparts.
McLachlan (1878: 350, 353) divided the Hydropsychidae into five sections: his section I
corresponds with Brauer's Oestropsidae, containing Polymorphanisus, Oestropsis, Phanostoma,
Aethaloptera and Amphipsyche. Ulmer (1907) united McLachlan's sections I and II (which
contained Macronema and Blepharopus) to form the Macronematinae, stating that the name
Oestropsidae could not be used, not even in the form Oestropsinae, because Oestropsis had been
synonymized with Polymorphanisus. This change of name has been generally accepted by authors
and is therefore retained under Art. 40 (a) of the International Code of Zoological Nomenclature.
Denning (1943) tried to revive the name Oestropsinae in preference to Ulmer's Macronematinae
but other authors have not accepted this change. Two other minor changes to this group name
have been proposed : Banks (1913) uses the Macronematidae as a family name (apparently in the
same sense as Ulmer's subfamily), and Navas (1926) reduced the group to a tribe, but neither
change has been accepted by subsequent authors.
The first attempt at splitting the Macronematinae sensu Ulmer was by Lestage (1936) who
divided the subfamily into two tribes, the Macronematini and the Polymorphanisini, the latter
distinguished by the loss of the palps. This division has been accepted by most authors and is the
one adopted in the present paper. Surprisingly, Ulmer (1951) continued to use his subfamily
Macronematinae with no further subdivision. Banks's (1939) introduction of the name
Oestropsychinae is difficult to interpret as the group apparently includes Polymorphanisus,
Oestropsyche and Amphipsyche. If it is intended to replace the Polymorphanisini then
Amphipsyche is erroneously included, but in any case it is an unnecessary replacement name.
Marlier (1962) accepts Lestage's division of the subfamily but renames the Polymorphanisini as
the Oestropsychini, which again is an unnecessary replacement name. The only other change
recently proposed is the standardization of the tribe name to Polymorphanisini by Flint (1974;
1978) and Scott (1975).
Abbreviations
AM Albany Museum, Grahamstown
ANIC Australian National Insect Collection, C.S.I.R.O., Canberra
BMNH British Museum (Natural History), London
IM Indian Museum, Calcutta
REVISION OF OLD WORLD POLYMORPHANISINI £j
IRSNB Institut Royal des Sciences Naturelles de Belgique, Brussels
IZPAN Instytut Zoologiczny, Polska Akademia Nauk, Warsaw
MCZ Museum of Comparative Zoology, Harvard
MNHN Museum National d'Histoire Naturelle, Paris
MNHU Museum fur Naturkunde der Humboldt-Universitat, Berlin
MRAC Musee Royal de 1'Afrique Centrale, Tervuren
NM Naturhistorisches Museum, Vienna
NMV National Museum of Victoria, Melbourne
RNH Rijksmuseum van Natuurlijke Historie, Leiden
USNM U.S. National Museum, Smithsonian Institution, Washington
ZI Zoological Institute, Leningrad
ZM Zoologisches Museum, Hamburg
Acknowledgements
I am very grateful to the following for the loan or donation of specimens: Ms J. Cardale,
C.S.I. R.O., Canberra; Dr J. Decelle, MRAC, Tervuren; Dr O. S. Flint, Jr, USNM, Washington;
Dr D. C. Geijskes, RNH, Leiden; Dr H. J. Hanneman, MNHU, Berlin; Dr A. Kaltenbach, NM,
Vienna; Dr J. Legrand, MNHN, Paris; Dr A. Liana, IZPAN, Warsaw; Dr G. Marlier, IRSNB,
Brussels; Dr K. M. F. Scott, AM, Grahamstown; Prof. Dr H. Striimpel, ZM, Hamburg; Ms M.
K. Thayer, MCZ, Harvard. Thanks are also due to Dr A. Neboiss, NMV, Melbourne, for
bringing the Australian material to my attention. I am grateful to my colleague Dr P. E. S.
Whalley for commenting on the manuscript.
Taxonomic section
Tribe POLYMORPHANISINI Lestage
Polymorphanisi Lestage, 1936: 176. Type-genus: Polymorphanisus Walker.
Oestropsychini Marlier, 1962: 118. Type-genus: Oestropsyche Brauer.
Polymorphanisini; Flint, 1974: 115.
Medium to large species, wing length 8-28 mm, yellow or brownish in coloration, probably always
greenish in life. Antennae usually longer than wings, occasionally up to four times fore wing length. Scape
large and bulbous, pedicel narrow, ring-like. Flagellar segments numerous, up to 95 in some species, always
long and narrow. Head with one or two pairs of setigerous warts on vertex ; hind pair, if present, often
reduced, especially in $. Eyes usually small, but greatly enlarged and almost meeting ventrally in J1 of the
Polymorphanisus ocularis-group. Mouthparts always absent, traces of palps occasionally visible. Thorax
with large, shield-shaped mesoscutellum, often with black markings in the P. nigricornis-group. Legs often
with dark tips to tibiae and femora (but not constant within species). Spurs 0.2.2, 0.3.2, 1.2.2, 1.3.2 or 1.3.3.
Mesothoracic legs often greatly broadened in ?, only slightly so (if at all) in J1. Wing-coupling mechanism
consists of one or two rows of curved macrotrichia near costa of hind wing, engaging on anal fold of fore
wing. Discal and median cell usually present in fore wing; in Oestropsyche discal cell absent, in Aethaloptera
'false' discal cell produced by secondary fusion of R4 + 5. Both cells absent in hind wing; 'false' discal cell
occasionally present (though not consistent) in Aethaloptera. In fore wing Sc and Rt separate, but fused for
most of length in Oestropsyche. Cu2 ends on wing margin in Polymorphanisus, but on 2 A in other genera. In
hind wing Sc and /?j either fused (Oestropsyche, Aethaloptera) or Rl ends on Sc (P. ocularis-group) or /?t
free (P. nigricornis-group); R2 and R3 usually separate (fused in Polymorphanisus).
J1 genitalia with well-sclerotized, ring-like ninth segment and one- or two-segmented, elongate gonopods.
Aedeagus greatly enlarged distally, with complex internal structure. 9 genitalia with elongate tubular ninth
segment, membranous ventrally with various tubercles. Eighth sternite partially divided into two plates.
REMARKS. The loss of the mouthparts is the most obvious distinguishing feature of this group;
the separation of this tribe from the Macronematini is also supported by larval characters (Scott,
1975). Betten (1934) described the distinctive wing-coupling mechanism in Macronema, which
has a single row of curved, hook-like macrotrichia along the costal margin of the hind wing. This
seems to be the general pattern throughout the Macronematini, and this type is also found in
Aethaloptera. A more advanced form is seen in Oestropsyche (Fig. 1) where a second row of
62
P. C. BARNARD
1mm
,0.1mm
,0.1mm
Figs 1-7 1-3, Oestropsyche vitrina ?, wing coupling mechanism, (1) costal area of
hind wing; (2) costal hair; (3) radial hair. 4, Aethaloptera dispar $\ costal hair from
hind wing. 5-7, Polymorphanisus semper i $, (5) costal area of hind wing; (6) costal hair; (7)
radial hair.
REVISION OF OLD WORLD POLYMORPHANISINI 63
hooked macrotrichia is present on the stem of Rs. In Polymorphanisus (Fig. 5) and Synoestropsis
this row of macrotrichia has migrated forwards to lie anteriorly to Rs. Both species groups of
Polymorphanisus are similar in this respect. The detailed structure of these macrotrichia also
varies within the group. In Aethaloptera (Fig. 4) the hooks are toothed on the inner side, just as
described in Macronema by Betten (1934). In Oestropsyche and Polymorphanisus the radial
hooks are not toothed (Figs 3, 7) whereas the costal hairs are only slightly curved but have an
enlarged, roughened tip (Figs 2, 6). In all cases the length of the macrotrichia varies according to
their position on the wing. In the fore wing 2 A is greatly thickened and prominent, and the wings
tend to fold along this vein, providing an attachment point for the hooks on the hind wing.
Geographical distribution
Most species of Polymorphanisini are restricted to tropical or subtropical areas. Apart from the
genus Synoestropsis, which is endemic to South America, the tribe is represented in the
Afrotropical region (sensu Crosskey & White, 1977) and Madagascar, and throughout the
Indian, South East Asian, Malaysian and Indonesian regions, including the Philippines. The far
eastern species Aethaloptera sexpunctata just reaches northern Australia. Although several
species are found in southern and central China, the main exception to the broad pattern of
distribution is A. evanescens, which occurs in the Amur Region of the U.S.S.R. and southern
Siberia. A few species have very wide distributions, such as Oestropsyche vitrina and
Aethaloptera sexpunctata (Fig. 46).
Biology
There is very little published information on the habitats of the Polymorphanisini, but one can
infer from the existing data that larvae of many of the species inhabit fairly large rivers, not
necessarily fast flowing.
Very few larvae in the group have been described. Ulmer (1912) and Barnard (1934) both
described what were believed to be larvae of Aethaloptera spp., but these were later referred to
Macronema (Ulmer, 1957). The larva of A. dispar was subsequently described by Gibbs (1973).
Marlier's (1943) 'larve C de Macronematinae' was shown to be a species of Polymorphanisus by
Ulmer (1957) and later Marlier (1958; 1961) redescribed what he believed to be the same species.
Since the adults collected at the same time are P. elisabethae, the larvae are probably the same
species, and not bipunctatus as Marlier believed. Scott (1975) has summarized the larval
characters of these two African genera and has shown that the distinction between the
Polymorphanisini and the Macronematini is supported by the larval features. Outside Africa
only two other species have definitely associated larvae. These are Aethaloptera evanescens in the
U.S.S.R., described (as rossicd) by Lepneva (1970), and Oestropsyche vitrina, described by Ulmer
(1957) from material from Sumatra and the Philippines.
Larvae of the neotropical genus Synoestropsis have been described, but not associated with
any known species (Flint, 1978).
Check-list of Old World Polymorphanisini
OESTROPSYCHE Brauer
vitrina (Hagen)
palingenia Brauer
hageni Banks syn. n.
AETHALOPTERA Brauer
Chloropsyche McLachlan evanescens (McLachlan)
Primerenca Navas rossica Martynov syn. n.
Paraethaloptera Martynov syn. n. gracilis (Martynov) comb. n.
maxima Ulmer
dispar Brauer sexpunctata (Kolenati)
maesi Navas dyakana Banks
maerina Navas punctata Banks syn. n.
64 P. C. BARNARD
POL YMORPHANISUS Walker
Oestropsis Brauer
nigricornis-group semperi (Brauer)
astictm Navas taoninus Navas
hainanensis Martynov syn. n. tumidus Banks
flavipes Banks syn. n. umbripes sp. n.
bipunctatus Brauer unipunctus Banks
elisabethae Navas
,. .„ XT , ocularis-group
bipunctatus pupillatus Navas syn. n. . . T T1
fuscus Ulmer angustipenms Ulmer
hargreavesi^n. *••»«•« Navas
mar/imsp.n oc«/am Ulmer
muluensis sp. n. . !**« Banks syn' n'
nigricornis Walker siimfa Ulmer
quadripunctatus Ulmer &****ie Navas syn. n.
scutellatus Banks
Key to world genera of Polymorphanisini
1 Rs and M in fore wing connected by cross-vein (Fig. 57). (Neotropical species) .
SYNOESTROPSIS
Rs and M not connected by cross-vein. (Old World species) 2
2(1) 'False' discal cell present in fore wing (enclosing corneous spot) formed by secondary fusion
of R4 + 5 (Fig. 19) . AETHALOPTERA (p. 66)
No 'false' discal cell (Figs 8, 47) 3
3 (2) Discal cell absent in fore wing; Sc and R^ fused for most of length (Fig. 8) ...
OESTROPSYCHE (p. 64)
Discal cell present in fore wing; Sc and Rl separate (Fig. 47) POLYMORPHANISUS (p. 78)
OESTROPSYCHE Brauer
Oestropsyche Brauer, 1868: 265. Type-species: Oestropsyche palingenia Brauer [ = Oestropsyche vitrina
(Hagen)], by monotypy.
Oestropsyche Brauer; Ulmer, 1907: 29 [redescription].
Head with slightly raised areas anteriorly on vertex, no tubercles posteriorly (Fig. 10). Frons slightly
inflated. Antennae at least twice as long as fore wing in $, much shorter than fore wing in $. Eyes small, not
meeting ventrally. J1 mesothoracic legs with tibia and tarsal segments slightly dilated (Fig. 9), markedly
dilated in $ (Fig. 15). Spurs 1.2.2 (1.3.2 in J1 from Philippines), not 2.3.3 as stated by most authors.
Venation as in Figs 8, 14; Sc and Rl fused for most of their length, discal cell absent. Median cell long and
broad in J, smaller in $.
REMARKS. Fischer (1963) places this monotypic genus in the Macro nematini rather than the
Polymorphanisini but the reason for this is not stated. The morphology of both the adults (with
the loss of the mouthparts) and of the larvae (described by Ulmer, 1951) clearly show its affinities
with the other genera in the Polymorphanisini.
Oestropsyche vitrina (Hagen)
(Figs 8-18)
Macronema vitrinum Hagen, 1859: 209. Holotype J, SRI LANKA: Rambodde (Nietner) (MCZ, Harvard)
[not examined].
Polymorphanisus vitrinus (Hagen) Hagen, 1864: 875.
Oestropsyche palingenia Brauer, 1868: 266. LECTOTYPE J1, PHILIPPINES: Antipolu (Semper) (IRSNB,
Brussels), here designated [examined]. [Synonymized by Ulmer, 1907: 29; considered valid species by
Banks, 1939: 56; synonymized by Ulmer, 1951 : 188.]
Oestropsyche vitrina (Hagen) Ulmer, 1907: 29.
REVISION OF OLD WORLD POLYMORPHANISINI
65
5mm
, 2mm ,
10
, 1mm
12
1mm
Figs 8-13 Oestropsyche vitrina $. 8, wing venation; 9, mid-leg; 10, head, dorsal view; 11,
genitalia, ventral view; 12, genitalia, dorsal view; 13, genitalia, lateral view.
66 P- C. BARNARD
Oestropsyche hageni Banks, 1939 : 56. Holotype J, INDIA : Mysore, Shimoga, River Tunga, 1865' [560 m], at
light, 21.vii.[? year] (Nathan) (type no. 23469, MCZ, Harvard) [examined]. Syn. n.
Macronema vitrinum Hagen; Ross, 1952: 35. [Holotype depository.]
3. Wing length 1 1-17 mm. Antenna 40-46 mm with about 75 segments. Head pale brown, whitish dorsally,
prothorax and tegulae whitish, rest of thorax brown. Eyes black. Abdomen blackish brown dorsally,
yellowish brown ventrally. Legs yellowish. Antennae yellowish brown with darker annulations. Wings
yellowish.
5. Wing length 11-16 mm. Antennae 5-10 mm with about 35 segments; segments shorter and narrower
than those of j1. Coloration similar to _J.
GENITALIA j (Figs 1 1-13). Ninth tergite with projecting median lobe, gonopods short and stout, apparently
unsegmented but with slight constriction distally. Well-developed lateral tubercles on tenth segment.
GENITALIA 9 (Figs 16-18). Ninth tergite slightly bifid, eighth sternite not completely divided, each half
pointed anteriorly.
REMARKS. Hagen's (1859) description of this species is based quite explicitly on a single male;
Ross (1952) states that this holotype is in the MCZ, Harvard (although both Banks (1939) and
Ulmer (1951) mention two males in Hagen's collection). However, Ulmer (1907; 1951) examined
a 'type' in the Selys-Longchamps collection (now in the IRSNB, Brussels) and there is a further
'type' in the BMNH! I have examined these two specimens: both are labelled 'Ceylon, Nietner'
and were almost certainly collected at the same time as the MCZ specimen, although presumably
not seen by Hagen when he described the species. In the absence of further evidence we must
accept the MCZ specimen as being the holotype; the specimens in Brussels and London therefore
have no type-status, but are nevertheless important 'topotypic' material.
Banks (1939) distinguished hageni from vitrina on size and minor venational differences, but
these are not constant within any one region as several authors, e.g. Ulmer (1951), have noted.
Banks also described a difference in the shape of the tip of the aedeagus of his species hageni, but
since he observed this from a posterior view, the shape depends entirely on the angle of view, as
Ulmer (1951) remarked.
The larva of vitrina was described by Ulmer (1957 : 371); its habitat in Sri Lanka was noted by
Schmid (1958: 107) as 'les rivieres assez agitees, en altitude moyenne, ou elle est souvent
commune'.
DISTRIBUTION. India (Martynov, 1935), Sri Lanka, Philippines, Sumatra, Java, Borneo, Sulawesi
(Ulmer, 1951), New Guinea (Papua New Guinea), China (Navas, 1923a; Ulmer, 1933).
MATERIAL EXAMINED
Sri Lanka: 3 3 [Rambodde] (Nietner) (2 in BMNH, 1 in IRSNB, Brussels); 2 ^, 3 ?, N[awala]piti[y]a
(BMNH); 1 .3 (Green) (BMNH); 1 ?, Pundaluoya, x.1897 (BMNH); \ 3, 1 ?, Lindula, 3.iii.l954 (Schmid)
(USNM, Washington); 1 3, Kitulgala, 2.iii.l954 (Schmid) (USNM, Washington); 1 ?, Uggalkaltota, 500'
[150m], 10-14.X.1970 (Flint) (USNM, Washington). Sumatra: 1 3, Sandaran Agong, Korinchi, 2450'
[740m], v-vi.1914 (Robinson & Kloss) (BMNH). Philippines: 1 3, Antipolu (Semper) (IRSNB, Brussels)
(paralectotype of Oestropsyche palingenia Brauer); 1 j1, 1 $, Luzon, Laur, Nueva Ecija, 1 1-12. ii. 1958
(Thompson) (USNM, Washington); 1 $, Negros, Victorias, at light, 17.vii.1928 (USNM, Washington); 1 J,
2 ?, Luzon, Mt Makiling (Baker) (USNM, Washington). New Guinea: 1 J1, 48 $, Papua New Guinea,
Kokoda, 1200' [360m], vii-ix.1933 (Cheesman) (BMNH).
AETHALOPTERA Brauer
Aethaloptera Brauer, 1875: 71. Type-species: Aethaloptera dispar Brauer, by monotypy.
Chloropsyche McLachlan, 1880: 69. Type-species: Chloropsyche evanescens McLachlan, by monotypy.
[Synonymized by Kimmins, 1962: 96.]
Primer enca Navas, 1915: 181. Type-species: Primer enca maesi Navas, by original designation and
monotypy. [Synonymized by Lestage, 1919: 293.]
Paraethaloptera Martynov, 1935: 193. Type-species: Paraethaloptera gracilis Martynov, by original
designation and monotypy. Syn. n.
Head with two pairs of setigerous warts on vertex in 3 (Fig. 20), posterior pair smaller and weakly
developed in $ (Fig. 25). Antennae up to four times wing length in J1, only slightly longer than wing in 9;
16
REVISION OF OLD WORLD POLYMORPHANISINI
Sc
67
Cu
18
15
5mm
Figs 14-18 Oestropsyche vitrina ?. 14, wing venation; 15, mid-leg; 16, genitalia, lateral
view; 17, genitalia, dorsal view; 18, genitalia, ventral view.
68 P- C. BARNARD
antennal segments shorter and more slender in 9, Eyes small, not meeting ventrally, slightly larger in ? (Figs
20, 25). J* mesothoracic legs with tibia and tarsal segments slightly dilated, greatly dilated in ? (Fig. 26).
Spurs 0.2.2 or 0.3.2. Venation as in Figs 19, 24; a 'false' discal cell, enclosing the corneous spot, formed by
R4 and R5 rejoining almost immediately after separating. J wings narrow, excised on hind margin (Fig. 19),
9 wings broader, triangular, hind margin almost straight (Fig. 24).
Most museum specimens of this group are yellowish brown in colour, with black eyes, but a specimen of
A. sexpunctata in the BMNH bears the description, made at the time of capture by H. T. Pagden, 'Head,
thorax, abdomen, legs and wing veins very pale green . . . eyes piceous.'
REMARKS. Kimmins (1962) provisionally synonymized Paraethaloptera with Aethaloptera, and
despite the differences in venation I have here formally synonymized it. The female genitalia of
Paraethaloptera gracilis (the males being unknown) are almost indistinguishable from those of
Aethaloptera maxima.
The genus is distributed throughout the Afrotropical region, India, Sri Lanka, South East
Asia and Indonesia as far as northern Australia, with one species in the Amur Region of the
U.S.S.R. and southern Siberia.
Key to species of Aethaloptera
1 Fork R4 in hind wing stalked (Figs 29, 33) maxima (p. 72)
Fork R4 in hind wing sessile (Figs 19, 24) 2
2 (1) Fork R2 in fore wing slightly shorter than its stem; /?! fuses apically with Sc (Fig. 39)
gracilis (p. 75)
Fork R2 in fore wing longer than its stem; Rl runs free to wing margin (Fig. 24) .. 3
3 (2) Tip of J1 aedeagus only slightly swollen in ventral view (Fig. 23); sclerites of $ eighth sternite
with outer posterior corners deeply excised (Fig. 27). (African species) . . dispar (p. 68)
Tip of J1 aedeagus strongly swollen in ventral view (Fig. 37) ; sclerites of $ eighth sternite
without deeply excised corners (Figs 38, 45). (Palaearctic and Oriental species) . . 4
4 (3) j* wing length 10-12 mm; posterior edges of? eighth sternite straight or slightly curved (Fig.
45). (Indian, Oriental and N. Australian species) sexpunctata (p. 75)
^ wing length 14-17 mm; posterior edges of 9 eighth sternite with shallow U-shaped excision
(Fig. 38). (Russian and Chinese species) evanescens (p. 72)
Aethaloptera dispar Brauer
(Figs 19-28)
Aethaloptera dispar Brauer, 1875 : 72. LECTOTYPE 3, SENEGAL: near Taoue, xi.1869 (Steindachner) (NM,
Vienna), here designated [examined].
Primerenca maesi Navas, 1915: 182. Holotype $, ZAIRE: Kwamouth (Maes) (MRAC, Tervuren) [not
examined]. [Synonymized by Lestage, 1919: 293.]
Primerenca maerina Navas, 1916: 242. [Unjustified emendation.]
.3. Antennae up to 50 mm, with up to 95 segments. Each segment golden brown, with distinct white annular
band near the base. Head and thorax yellowish brown, occasionally greenish, legs yellow, abdomen
yellowish brown, darker dorsally. Wings pale green, fading to yellowish brown. Brown spots centred on
various cross- veins in fore wing (Fig. 19) but these are variable in number. Wing length 10-15 mm. Spurs
0.3.2. Fork R4 in hind wing sessile, Sc and /?, in fore wing sinuous apically (Fig. 19).
$. Antennae up to 10mm, with 30-35 segments. Coloration as in _?, except that basal white band on
antennal segments is often ill-defined or even absent. Wings broader and shorter than in j1 ; wing length
8-10 mm. Spurs and venations similar to male, except that in fore wing fork M3 is occasionally stalked. Sc
and Rl in fore wing straight apically (Fig. 24).
GENITALIA J1 (Figs 21-23). Ninth tergite slightly produced centrally. Aedeagus with angular projection on
ventral surface, internal armature with broad rounded lobes in lateral view.
GENITALIA $ (Fig. 27). Plates of eighth sternite with outer posterior corners deeply cut away.
REMARKS. This is a very variable species throughout its range; this variability has prompted
several authors to attempt to subdivide it. Lestage (1936) doubted that 'dispar' was the same
REVISION OF OLD WORLD POLYMORPHANISINI
Sc
69
0.5mm
22
23
0.5mm
0.5mm
Figs 19-23 Aethaloptera dispar j. 19, wing venation. 20, head, dorsal view; 21, genitalia,
lateral view; 22, aedeagus, lateral view; 23, aedeagus, ventral view.
species throughout Africa, because of variation in the degree of dilation of the female tibiae, and
so on. Marlier (1943) stated that there were definitely three groups in Zaire, based on differences
in both male and female genitalia, the number of markings on the wings and the colour of the
antennal segments. However, he declined to name these forms, although suggesting that
Primer enca maesi Navas was a valid species. Kimmins (1962) took a rather opposite view,
suggesting that not only are both dispar and maxima variable, but that they may be only local
forms of one widespread species.
Only a detailed study of long series of specimens, coupled with examination of larvae from
different areas, could help to solve this problem. In the present study I have examined a total of
approximately 540 specimens of dispar, and I can find no consistent differences sufficient to split
the species into discrete groups : indeed there can be as much variation between two specimens
caught at the same time at one locality as between examples from the extremes of the
70
P. C. BARNARD
Figs 24-27 Aethaloptera dispar ?. 24, wing venation; 25 head, dorsal view; 26, mid- leg; 27,
eighth sternite, ventral view.
geographical range. A. dispar and maxima have reasonably discrete distributions; both species
occur in Angola but in different river basins (see remarks on A. maxima below). The two species
meet in Zambia at Victoria Falls, but here they can still be separated by the hind wing venation
and the spur formulae; the genitalia in both sexes are very similar. Kimmins (1962) reports that
males from Malawi (Nyasaland) have the typical venation and spur formula of dispar but have
genitalia more like maxima.
The collection of over 200 specimens, mostly females, from Zaire (listed below) is of some
interest, as they were caught at some distance from water. The collector, Dr S. L. Sutton (in litt.,
1978) estimates that the trapping site was about 25 km SSW. of Kindu, and 8 km east of the
River Lualaba, a headwater of the Zaire River. There is another river, the Kasuku, about 3 km
REVISION OF OLD WORLD POLYMORPHANISINI
71
Fig. 28 Distribution map ofAethaloptera dispar and A . maxima.
to the west, but the only nearer water was a slow, shallow stream (at most 3 cm deep) about 1 km
away. Fourteen other species of Trichoptera were caught at the same site.
DISTRIBUTION (Fig. 28). Senegal, Sierra Leone, Ivory Coast (Marlier, 1978), Ghana, Nigeria,
Zaire, Sudan, Uganda, Tanzania, Angola (Marlier, 1965), Zambia, Malawi, Zimbabwe. Also
recorded from Madagascar by Navas (1923&) although this is not confirmed.
MATERIAL EXAMINED
Senegal: 1 J1, 2 ?, near Taoue, xi.1869 (Steindachner) (NM, Vienna) (paralectotypes of dispar). Sudan:
4 1, 1 9, White Nile, L. Loat, 1906 (BMNH); 1 ?, 1914 (Lowe) (BMNH); 2 ?, El Jebelain, 3-4.xii.1961 ; 1 ;,
Renk, 4-7.xii.1961; 3 <?, 6 9, Malakal-Shambe, 8.xii.l961; i'<$, 1 $, Mongalla-Malakal, 24-27.xii.1961
(Cloudsley-Thompsori) (BMNH). Sierra Leone: 1 3, 6 ?, Njala, 2.iv.l926, 2.vi.l926, 19-21. viii. 1930,
17.iii.1932, l.iv.1932, vi.1934 (Hargreaves) (BMNH). Ghana: 2 3, 1 $, Afram R., Mankrong, 13.ix.1950
(Berner) (BMNH); 2 $, Sunyani, 12-13.xii.1965 (Gibbs) (BMNH); 3 ?, Tafo, 9-13.X.1966 (Gibbs) (BMNH);
6 J, Black Volta, 1967 (Petr) (BMNH). Nigeria: 4 ?, Birnin Kebbi, 3.xi.l957 (BMNH); 1 5, Oban District,
1910 (Talbot) (BMNH); 11 ?, Samaru, 18-25.V., l-8.ix.1970 (Ward) (BMNH); 1 ?, Zaria, Samaru,
72 P. C. BARNARD
26.ix.1970 (Deeming) (BMNH); 9 1, Lake Kainji, 1975 (Bidwell) (BMNH). Zaire: 2 ;, 1900 (Andreae)
(BMNH); 1 ;, 4 », Kabalo, v.1926 (Schouteden) (BMNH); 1 ?, Gombe, 27.ii.1949 (Marlier) (MRAC,
Tervuren); 1 j, Leopoldville [Kinshasa], at light, ii.1949 (Marlier) (MRAC, Tervuren); 1 3, Tshuapa,
Lukolela, 13.viii. 1947 (Poll) (MRAC, Tervuren); 10 i, c. 200 ?, Sciere, near Kindu, xi.1974 (Sutton)
(BMNH). Uganda: 1 J, Lake George, xi.1920 (Carpenter) (BMNH); 2 J, Lake Albert, Hi. 1931 (Hopkins)
(BMNH); 1 J, Jinja, iv.1952 (USNM, Washington); 2 J, 49, 2-24.V.1954 (Hickiri) (BMNH); 8 5, 90 v, W.
Nile, near Laropi, 27-28.iv.1956, 18 ;, 75 , , Albert Nile, Pakwach, 26-29.iv.1956 (Corbet) (BMNH); 3 ;,
8 ?, Victoria Nile, L. Kyoga, near Namasale, l.v.1956 (Corbet) (BMNH); 2 ., Kawanda, Kampala,
20-29.viii.1969 (Brown) (BMNH). Tanzania: 4 ,, Old Shinyanga, 4.V.1954, 2-7.iii.1956 (Burn} (BMNH);
1 9, Lake Victoria, Mwanza Pier, ll-13.viii.1956 (Corbet} (BMNH). Zambia: 12 $, 4 ?, Zambezi River,
Katambora, iv.1962 (Pinhey) (BMNH). Malawi: 2 J, Chiromo, 2.xii.l923 (Smee) (BMNH); 5 ^, Ntundu,
7.viii.l952 (Berner) (BMNH). Zimbabwe: 1 j\ Zambezi River, Victoria Falls, 9.vii.l962 (Mason) (AM,
Grahamstown); 2 J, Victoria Falls, xii.1955-i.1956 (USNM, Washington); 2 j1, 2 9, Victoria Falls
National Park, 3-6.iv.1968 (Spangler) (USNM, Washington); 1 2, Gatooma, iv.1956 (USNM,
Washington).
Aethaloptera maxima Ulmer
(Figs 28-34)
Aethaloptera maxima Ulmer, 1906: 62. NEOTYPE J1, SOUTH AFRICA: Waterval River, National Road
between Standerton and Greylingstad, 12.L1959 (ZM, Hamburg), here designated [examined].
Chloropsyche maxima (Ulmer) Ulmer, 1907: 16.
Aethaloptera maxima Ulmer; Weidner, 1964: 66 [J1 holotype destroyed].
[Aethaloptera dispar Brauer, forma A; Marlier, 1965: 40. Misidentification.]
J. Antennae up to 45 mm long, with up to 80 segments. Wing length 14-17 mm. Antennal segments golden
brown, without annulations, otherwise as A. dispar. Spurs 0.2.2. Venation as in j1 dispar except that in
hind wing fork R4 is stalked (Fig. 29).
9- Antennae 15mm long with approximately 45 segments. Wing length 9-15 mm. Posterior warts on
head scarcely developed. Antennal segments golden brown with basal third to half white. Spurs 0.2.2.
Venation as in 9 dispar except that fork R4 in hindwing is stalked (Fig. 33).
GENITALIA j1 (Figs 30-32). Very similar to dispar, differing slightly in the internal armature of the aedeagus.
GENITALIA 9 (Fig. 34). Similar to dispar but with the outer posterior corners of the eighth sternite less deeply
excised. Kimmins (1962) discusses the differences in genitalia between dispar and maxima.
REMARKS. The relationship of this species with A. dispar is discussed under the latter species.
Marlier (1965) recorded maxima from Angola as a form ('forma A') of dispar, but the spur
formula (0.2.2) and his figure of the female genitalia show that the specimens can undoubtedly be
referred to maxima. They were collected in eastern Angola, in the Zambezi basin, and this does
not unduly extend the known distribution (Fig. 28).
DISTRIBUTION (Fig. 28). Angola (Marlier, 1965), Namibia, South Africa, Zimbabwe,
Mozambique, Zambia.
MATERIAL EXAMINED
South Africa: 2 J, data as neotype (BMNH); 1 9, Boshof, 7.J.1921 (BMNH); 2 9, Wilge River,
Frankfort, 9.H.1959 (BMNH); 4 J1, 4 , , Transvaal, Standerton, at light, 23.iii.1960 (AM, Grahamstown);
7 _>, 5 9, Frankfort, at light, 18.iii.1965 (Chutter) (AM, Grahamstown); 4 , , Kruger National Park, Olifants
Camp, 19.ii.1968; 3 9, Pretoriuskop, 20-2 l.ii. 1968; 1 9, Transvaal, Mooketsi, 14-18.ii.1968 (Krombein &
Spangler); 2 9, Transvaal, Potchefstroom, l.iii.1968 (Spangler) (USNM, Washington). Namibia: 1 J, 1 9,
Otjimbumbe, Kunene River, iii. 1923 (Barnard) (BMNH). Zimbabwe: 1 J1, Zambezi River, Victoria Falls,
9.V.1962 (Mason) (AM, Grahamstown). Mozambique: 12 ?, Caia, Zambezi River, 29.viii.1929 (Marshall)
(BMNH); 1 ^, Gare, 13.vii.1929 (Lesne) (BMNH); 4 ;, Nova Choupanga, near Chemba, v.1928, iv.1929
(Lesne) (BMNH). Zambia: 1 j1, Upper Luangwa River, 27.vii.-13.viii.1910; 1 J1, Lower Luangwa River,
4-13.ix.1910 (Neave) (BMNH); 19 $, Chipepo, from tiger fish stomachs, 25.X.1956 (BMNH).
Aethaloptera evanescens (McLachlan)
(Figs 35-38)
Chloropsyche evanescens McLachlan, 1880: 69. Lectotype £, U.S.S.R.: 'Amur Land' (Christoph)
(BMNH), designated by Kimmins (1957: 105) [examined].
REVISION OF OLD WORLD POLYMORPHANISINI
73
0.5mm
Figs 29-34 Aethaloptera maxima. 29, * wing venation; 30, > genitalia, lateral view; 31,
aedeagus, lateral view; 32, aedeagus, ventral view; 33, ? wing venation; 34, ? eighth
sternite, ventral view.
74
P. C. BARNARD
Figs 35-38 Aethaloptera evanescens. 35, _£ wing venation; 36, $ genitalia, lateral view; 37,
aedeagus, ventral view; 38, 9 eighth sternite, ventral view.
Aethaloptera rossica Martynov, 1910: 385. Syntypes J, ?, U.S.S.R. : 'Ussurian Amur Land and S. Siberia to
River Ob" (? ZI Leningrad) [not examined]. Syn. n.
Aethaloptera evanescens (McLachlan) Kimmins, 1962: 96.
3. Antennae up to 40mm long, with approximately 75 segments. Wing length 14-17 mm. Antennal
segments golden brown with basal third white. Head and thorax yellowish brown, legs pale yellow.
Abdomen yellowish brown ventrally, slightly darker dorsally. Usually three brown spots on fore wing (Fig.
35) but these may be reduced in number or absent. Spurs 0.2.2. or 0.3.2. Venation as in A. dispar (Fig. 35).
9. Antennae 10-12 mm long, with up to 40 segments. Wing length 10 mm. Coloration and spurs as j1,
venation as in $ dispar.
GENITALIA 3 (Figs 36, 37). Ninth segment very narrow ventrally, gonopods slender. In ventral view,
aedeagus strongly enlarged apically with square-edged excision.
GENITALIA $ (Fig. 38). Eighth sternite with a shallow U-shaped excision at the outer posterior corners.
REMARKS. Martynov (1934: 305) suspected that his species rossica was a synonym of evanescens,
REVISION OF OLD WORLD POLYMORPHANISINI 75
as it was only separable by the spur formula, which was then considered a generic character to
separate Aethaloptera and Chloropsyche. Kimmins (1962) has synonymized these genera as the
spur formula is unreliable, so the synonymy of the two species is inevitable. I have not examined
the Martynov types, which are presumably in the ZI, Leningrad, but the series in the BMNH
from the Rivers Ob' and Ussuri were determined as rossica by Martynov. An examination of this
series showed that the pre-apical spur of the mesothoracic leg varied from being very long to very
short, and was absent in two specimens, thus emphasizing the variability of this character.
DISTRIBUTION. U.S.S.R. (Amur Region, S. Siberia), S. and E. China.
MATERIAL EXAMINED
China: 1 J1, 1 ?, Fu-chou (Yang); 1 J1, Pai-se, Kwangsi, 29.iii.1939 (Richardson). U.S.S.R.: 3 3, Amur
Region, Raddefka [ = Radde, on Amur River]; 4 j1, River Ob', Tomsk, 6.vii.l925 (Filipjev); 2 3, R. Ussuri,
Bikin, 9.vii.l927 (Martynov). (All specimens in BMNH.)
Aethaloptera gracilis (Martynov) comb. n.
(Figs 39, 40)
Paraethaloptera gracilis Martynov, 1935: 193. 4 9 syntypes, INDIA: Sanjai River, Chakradharpur, Chota
Nagpur, 8-10.ii.1918 (Annandale & Gravely) (IM, Calcutta) [not examined].
cJ. Unknown
9. Antennae up to 10mm long, with about 35 segments. Wing length 8-10 mm. Antennal segments
golden brown, proximal half yellowish white. Head, thorax and abdomen yellowish brown, legs yellow.
Wings yellowish brown with up to four brown spots (Fig. 39) which may be reduced or absent. In fore wing,
fork R2 is shorter than its stalk; in both wings R{ fuses apically with Sc (Fig. 39). Spurs 0.2.2.
GENITALIA $ (Fig. 40). Eighth sternite with a moderately deep excision on the outer posterior corners.
REMARKS. I was unable to borrow the syntypes of this species, but Martynov's (1935) figures of
the distinctive venation make it instantly recognizable. Because of the similar distribution, it is
possible that gracilis specimens are dimorphic females of sexpunctata, but the peculiar venation
makes this unlikely.
DISTRIBUTION. India, Sri Lanka.
MATERIAL EXAMINED
India: 1 $, Alagar Kovil, Madura District, 17.iii.1936 (BMNH); 1 $, New Delhi, at light, 29.xi.1967
(Gibson) (USNM, Washington). Sri Lanka: 1 9, Uggalkaltota, 350' [105 m], 31.i-8.ii.1970; 2 $, Sigiriya,
800' [240m], 25.ii.1970 (Davis & Rowe) (USNM, Washington).
Aethaloptera sexpunctata (Kolenati)
(Figs 41-46)
Setodes sexpunctata Kolenati, 1859: 266. Holotype ^, INDIA (Huge!) (NM, Vienna) [examined].
Polymorphanisus sexpunctatus (Kolenati) Brauer, 1868: 263.
Aethaloptera sexpunctata (Kolenati) Ulmer, 1907: 19.
Aethaloptera dyakana Banks, 1920: 354. Holotype $, BORNEO: Duson Timoc, v.1882 (Grabowsky) (type no.
10885, MCZ, Harvard) [examined]. [Synonymized by Ulmer, 1951: 194.]
Paraethaloptera punctata Banks, 1938: 232. LECTOTYPE $, WEST MALAYSIA: Negri Sembilan, Port
Dickson, lO.i.1935 (Pendlebury) (BMNH), here designated [examined]. Syn. n.
Aethaloptera punctata (Banks) Kimmins, 1962: 96.
J1. Antennae up to 35 mm long, with about 70 segments. Wing length 10-12 mm. Antennal segments golden
brown with proximal third white. Head, thorax and abdomen yellowish brown, legs pale yellow. Wings
greenish or yellowish, with up to seven brown spots (Fig. 41). Spurs 0.3.2.
?. Antennae 10mm long, with about 40 segments. Wing length 8-11 mm. Coloration as in J1. Spurs
0.3.2.
GENITALIA 3 (Figs 42-44). Aedeagus greatly expanded in ventral view to form two rounded lobes with a
flatter lobe between.
76
P. C. BARNARD
Figs 39-45 39, 40, Aethaloptera gracilis ?, (39) wing venation; (40) eighth sternite, ventral
view. 41-45, A. sexpunctata, (41) J1 wing venation; (42) J genitalia, lateral view; (43)
aedeagus, lateral view; (44) aedeagus, ventral view; (45) $ eighth sternite, ventral view.
REVISION OF OLD WORLD POLYMORPHANISINI
77
70 80 90 100 110 120 130 140 150
Fig. 46 Distribution map of Aethaloptera sexpunctata.
GENITALIA ? (Fig. 45). Posterior margin of each half of eighth sternite almost straight, slightly serrated
towards mid-line.
REMARKS. In his original description, Kolenati (1859) described the presence of brown spots on
the posterior cubitus, arculus and thyridium, making six spots on the two fore wings combined,
hence the specific name sexpunctata. As described above, most specimens have more spots, often
up to seven, on each wing. The male holotype does in fact have four spots on each wing;
Kolenati apparently overlooked the anal spot.
Kimmins's (1962) 'type' ofpunctata Banks is so labelled by Banks but is not distinguished in his
description. This specimen is therefore designated the lectotype. Banks (1938) also refers to
specimens from Selangor, Kuala Lumpur, but the location of this material is unknown ; these
would be further paralectotypes (the two females from this locality in the BMNH do not agree
with the dates cited by Banks). Other specimens from Port Dickson in the BMNH were
determined by Banks as sexpunctata, thus supporting the synonymy.
The discovery of A. sexpunctata in N. Queensland is the first record of the tribe
Polymorphanisini in Australia.
DISTRIBUTION (Fig. 46). India, Sri Lanka, Burma, Laos, Cambodia (Ulmer, 1926), West
Malaysia, Sumatra (Ulmer, 1951), Borneo (Ulmer, 1930), New Guinea, Australia (Queensland).
78 P. C. BARNARD
MATERIAL EXAMINED
India: 2 1, NW. India (Home) (BMNH); 14 ^, Bihar, Pusa, various dates and collectors (BMNH). Sri
Lanka: 1 J1, Maha Oya, 12.iii.1954 (Schmid) (USNM, Washington); 2 J1, Hasalaka, 1000' [300 m], at light,
30-3 l.iii. 1973 (Baumann & Cross) (USNM, Washington). Burma: 9 ^, Prome, at light, 17-18. ii.1918
(BMNH). Laos: 1 3, Pakkading, 4.V.1932 (Kerr) (BMNH).
West Malaysia: 2 $, Penang, Tanjong Bungah, at light, ll.iv.1955 (Pagden); 1 ?, Lahat, 1916
(Henderson); 2 ?, Selangor, Kuala Lumpur, at light, 9.ii., 30. ix. 1931 (Pendlebury); 5 9, Negri Sembilan,
Port Dickson, various dates (Pendlebury); 1 ?, data as lectotype (paralectotype of punctata Banks). New
Guinea: 1 $, Papua New Guinea, Port Moresby, 1887 (Kowald). (All specimens in BMNH.)
Australia: 1 ?, Queensland, 20 miles [32 km] W. of Tully, 20. iv.1964 (Common & Upton) (BMNH) [first
record for Australia].
POLYMORPHANISUS Walker
Polymorphanisus Walker, 1852: 78. Type-species: Polymorphanisus nigricornis Walker, by monotypy.
Oestropsis Brauer, 1868: 263. Type-species: Oestropsis semperi Brauer, by monotypy. [Synonymized by
Ulmer, 1907: 19.]
Head with one pair of setigerous warts on vertex with a transverse ridge posteriorly on each side (Fig. 52).
Antennae up to twice wing length, similar in both sexes. Mesothoracic legs of $ with tibia and tarsal
segments slightly broadened, very broad in $. Spurs 1 .3.2, 1 .3.3 or 2.3.3. Venation as in Figs 47, 120; Sc and
/?, in fore wing terminate separately on wing margin. In hind wing of ocw/an's-group /?, ends on Sc; R2 + 3
fused.
REMARKS. It is possible that future work will show that the two species groups of
Polymorphanisus deserve generic status. The ocularis-group particularly has many derived
characters such as the enlarged male eyes, the unsegmented gonopods, and so on. However, there
are also several features which unit the two groups, particularly venational characters and the
unusual wing-coupling mechanism. Further larval descriptions will probably elucidate the
relationship of the two groups as at present the larvae of the ocularis-group are unknown.
The genus is distributed throughout the Afrotropical region, and from India through south
East Asia to Indonesia, including the Philippines.
Key to species of Polymorphanisus
1 In fore wing Ml is closely associated at base with R5 ; M2 is direct continuation of Ml + 2
stem (Fig. 47). J1 eyes well separated ventrally. (nigricornis-group) .... 2
In fore wing Ml is direct continuation of M, + 2 stem; M2 arises from median cell (Fig. 120).
3 eyes almost touching ventrally (Fig. 122). (ocw/am-group) 16
2 (1) African species 3
Asian species 6
3 (2) Thorax unmarked '. . . 4
Thorax with two spots on mesoscutellum (Figs 58-71) 5
4 (3) Antennal flagellum dark brown or black, with dark stripe on scape (Fig. 75) hargreavesi (p. 87)
Antennae yellow marlieri (p. 87)
5 (3) Mesoscutellar spots small and round (Fig. 48). Antennae brown in _>, black in ? .
bipunctatus (p. 81)
Mesoscutellar spots elongate (Figs 58-71). Antennae yellow . . . elisabethae(p.8l)
6 (2) Thorax unmarked
Thorax with one or more dark spots . . 9
7 (6) Fork R2 in fore wing shorter than its stalk (Fig. 110). . . . . taoninm (p. 95)
Fork R2 longer than or equal to its stalk (Fig. Ill) ....... 8
8 (7) Antennae, fore femora and tibiae pale yellowish brown (basal antennal segments sometimes
with a narrow, longitudinal, dark brown stripe) astictus (p. 79)
Antennae, fore femora and tibiae dark brown umbripes(p.95)
9 (6) Mesoscutellum unmarked.
Two large spots anteriorly on mesoscutum and two on metascutum (Fig. 94) .
quadripunctatus (p. 90)
Mesoscutellum with one, two or four spots . ... . . . . . 10
10 (9) Two spots on mesoscutellum (Fig. 105) . . . 11
REVISION OF OLD WORLD POLYMORPHANISINI 79
One or four spots on mesoscutellum (Figs 85, 87, 118) 14
11 (10) Thoracic spots large and oval, meeting sides of mesoscutellum (Fig. 102) . scutellatus (p. 92)
Thoracic spots not touching sides of mesoscutellum (Fig. 105) 12
12(11) Head greatly swollen anteriorly (Fig. 112). Mesoscutellar spots narrow and elongate,
widened posteriorly tumidus (p. 95)
Head not swollen. Mesoscutellar spots not widened posteriorly 13
13(12) Mesoscutellar spots elongate (Fig. 105). j1 fore wings falcate (Fig. 104) . . semperi (p. 92)
Mesoscutellar spots small and round (Fig. 91). $ fore wings not falcate . .nigricornis (p. 90)
14(10) Thorax with single spot in centre of mesoscutellum (Fig. 118). . . . unipunctus (p. 96)
Thorax with more than one spot (Figs 85, 87) 15
15(14) Wings dark brown. Thorax with one large spot on mesoscutellum and two smaller squarish
spots anteriorly on metascutum (Fig. 85) fuscus (p. 84)
Wings pale yellowish brown or greenish. Mesoscutellum with four small round spots (Fig.
87) muluensis(p.%T)
16 (1) Fore wing with no brown spots " . angustipennis (p. 98)
Fore wing with one or two brown spots (Figs 131, 133) 17
17(16) Fore wing with only one spot (Fig. 133) guttatus (p. 98)
Fore wing with two spots (Fig. 131) 18
18 (17) J1 gonopods abruptly narrowed half-way (Fig. 129). Outer posterior corners of $ eighth
sternite sloping evenly towards outer edge (Fig. 132). (African species) . . similis(p. 103)
j gonopods slightly widened in centre (Fig. 136). Outer posterior corners of ? eighth
sternite produced in rounded lobes (Fig. 138). (Indian and Oriental species) . ocularis(p. 100)
The nigricornis-group
Dark green or yellowish brown species, often with black markings on thorax. Wings long and narrow,
without brown markings (except P. fuscus). Antennae about twice forewing length. Eyes small, well
separated ventrally in male. Spurs usually 1 .3.3, in some species 1 .3.2. In fore wing M2 is direct continuation
of A/, + 2 stem ; in hind wing /?, terminates on wing margin, j gonopods long and slender, terminal segment
well defined.
Polymorphanisus astictus Navas
(Figs 79-83)
Polymorphanisus astictus Navas, 1923a: 47. LECTOTYPE 3, CHINA: Kweichow, P'ing-Fa, 1908
(Cavalerie) (MNHN, Paris), here designated [examined].
Polymorphanisus hainanensis Martynov, 1930: 82. Holotype J, CHINA: Hainan Tao I., Mt Wuchih Shan,
20.V.1903 (BMNH) [examined]. Syn. n.
Polymorphanisus flavipes Banks, 1939: 53. Holotype $, INDIA: Mysore, Shimoga, River Tunga, 1865'
[560 m], at light, 10.vi.[? year] (Nathan) (type no. 23467, MCZ, Harvard) [examined]. Syn. n.
J. Antennae up to 45 mm, with about 65 segments. Antennal segments yellow, basally brown, distal
segments yellowish brown. Body yellowish brown, wings pale greenish yellow or yellowish brown. No
markings on thorax. Wing length 19-23 mm, venation as in Fig. 79. Spurs 1.3.3.
$. Antennae up to 50 mm, with up to 80 segments. Scape, pedicel and first flagellar segment sometimes
with thin black stripe externally, but all segments may be yellow as in j1. General coloration as $. Wing
length 18-28 mm. Spurs 1.3.3.
GENITALIA J1 (Figs 80-82). Gonopods long and narrow. Ninth segment produced into a spatulate lobe,
viewed dorsally. Aedeagus with swollen round tip, with small, oval cavity at tip.
GENITALIA $ (Fig. 83). Outer posterior corners of eighth sternite slightly produced as rounded lobes; outer
margins of sternite slightly sinuate.
REMARKS. Banks (1939) seems to have confused this species with some of its allies. He
erroneously states that it belongs in the indicus-ocularis group; Ulmer (1951 : 178) has pointed
out that this is incorrect. In the same paper Banks described flavipes, comparing it with what he
believed to be Walker's species nigricornis, but he was misled by Walker's failure to notice the
mesoscutellar markings (see umbripes, p. 95). Banks also noted two specimens of flavipes with
yellowish wings and black spots each side of the mesonotum. I have examined all of Banks's
material of this species; the two specimens with yellow wings are males of flavipes (i.e. astictus)
80
. 1mm .
^ V-
Figs 47-53 Polymorphanisus bipunctatus. 47, ? wing venation; 48, $ thorax; 49, j
genitalia, lateral view; 50, aedeagus, lateral view; 51,9 wing venation; 52, 9 head, dorsal
view; 53, 9 eighth sternite, ventral view.
REVISION OF OLD WORLD POLYMORPHANISINI gj
but the two with 'black' markings are females with the normal yellowish brown wing colour. The
markings are simply abnormally dark attachment points of the flight muscles showing through
the mesonotum.
DISTRIBUTION. India (Mysore), China, Thailand, West Malaysia.
MATERIAL EXAMINED
India: 3 }, 34 ?, data as holotype offlavipes, various dates; 12 :, Mysore, Bhadravati (Nathan). (All
specimens in MCZ, Harvard: all apparently paratypes offlavipes but not labelled as such.) China: 1 $,
Hainan Tao I., Mt Wuchih Shan, 12. v. 1903 (paratype of hainanensis). Thailand: 1 j1, Upper Pran R.,
14.iv.1926 (Ladell). West Malaysia: 1 J, Pahang, Kuala, Tahan, at light, 300' [90m], 23.xi.1921
(Pendlebury); 1 9, Kedah, nr Jitra, 5.iv.l928 (Pendlebury); 1 9, Selangor, Ulu Langat, at light, 31.viii.1934
(Pendlebury). (All specimens in BMNH.)
Polymorphanisus bipunctatus (Brauer)
(Figs 47-53, 72)
Oestropsis bipunctatus Brauer, 1875: 73. LECTOTYPE 9, ETHIOPIA: Beni Sciangul, Blue Nile, 1871
(Marno) (NM, Vienna), here designated [examined].
Polymorphanisus bipunctatus (Brauer) Ulmer, 1907: 20.
3. Antennae up to 40mm, with up to about 70 segments. Scape and pedicel yellowish brown, flagellar
segments brown with dark annulation at each joint. Body colour yellowish brown. Thorax with one pair of
round black spots on mesoscutellum (Fig. 48). Wings pale green, fading to yellowish brown. Wing length
22-26 mm. Venation as in Fig. 47. Spurs 1.3.3.
$. As 3, except as follows. Scape and pedicel of antennae with black longitudinal stripe on outer side,
flagellum black (Fig. 52). Wing length 20-28 mm. Venation as in Fig. 51. Spurs 1.3.3, occasionally 2.3.3.
GENITALIA J1 (Figs 49, 50). Terminal segment of gonopod short and wide. Ratio of lengths of basal segment
of gonopod to terminal segment 2.6-2.9:1. Ninth segment relatively broad ventrally, rounded dorsally.
GENITALIA 9 (Fig. 53). Inner posterior corners of eighth sternite strongly rounded; thickened edges
extending almost half-way along inner side. Outer corners almost right-angled.
REMARKS. Betten & Mosely (1940) regarded this species as almost certainly a synonym of
nigricornis. In view of the widely differing distribution it would seem best to regard them as
separate, at least until their larvae have been described (the same would apply to ocularis and
similis in the ocular is-group). P. bipunctatus can be separated from nigricornis by the relative
lengths of the segments of the gonopods.
The Angolan specimen listed below is presumably from the south or east of that country, in
the Zambezi basin (cf. Aethaloptera maxima, p. 72).
DISTRIBUTION (Fig. 72). Sudan, Ethiopia, Kenya, Angola, Zimbabwe, South Africa.
MATERIAL EXAMINED
Sudan: 1 5, Senga [Singa], Blue Nile, 30.xii.1914 (Lowe}; 1 ;, Zeidab, 15.xi.1918 (Bedford); 1 : , Shendi,
3 1.x. 1928 (Cowland); 2 3, nr mouth of Dinder River, Blue Nile, 26.vii.1909 (Flower); 1 J, White Nile. (All
specimens in BMNH.) Ethiopia: 1 v paralectotype, data as lectotype (NM, Vienna). Kenya: 2 9, Nzoia
River, Lwamba Ferry, 19-20.iv.1956 (Corbet) (BMNH). Angola: 1 ? (BMNH). Zimbabwe: 1 J1, Victoria
Falls, i.1956 (USNM, Washington). South Africa: 1 ?, Transvaal, Elandshoek, at light, 16.xi.1946
(Capener) (BMNH); 3 $, Natal, Ndumu Reserve, l-10.xii.1963; 1 3, Natal, Tugela River, 9.x. 1953; 4 $,
Natal, Mooi River, 20.xii.1940 (Crass) (AM, Grahamstown).
Polymorphanisus elisabethae Navas
(Figs 54-56, 58-72)
Polymorphanisus elisabethae Navas, 193 la: 140. Holotype 9 [not ^, as stated by Navas], ZAIRE: Malela,
viii.1928 (Queen Elisabeth) (MRAC, Tervuren) [examined].
Polymorphanisus bipunctatus pupillatus Navas, 1931a: 139. Holotype J1, ZAIRE: Bokote, 22.xii.1925
(Hulstaert) (MRAC, Tervuren) [examined]. Syn. n.
[Polymorphanisus bipunctatus (Brauer) Marlier, 1961 : 208. Misidentification.]
Polymorphanisus pupillatus Navas; Marlier, 1965: 77.
[Polymorphanisus bipunctatus (Brauer); Marlier, 1965: 40. Misidentification.]
82
1mm
2A Cu1
Figs 54-57 54-56, Polymorphanisus elisabethae , (54) j1 wing venation; (55) $ genitalia,
lateral view; (56) •: eighth sternite, ventral view. 57, Synoestropsis sp. * fore wing.
REVISION OF OLD WORLD POLYMORPHANISINI
83
60
2mm
Figs 58-71 Variation in thoracic markings of Polymorphanisus elisabethae. 58, V, Zaire,
Kivu; 59, 3, Albertville; 60, 3, Sierra Leone; 61,9 holotype o{ elisabethae; 62, 3 holotype
of pupillatus; 63, 9, Zaire, Popokabaka; 64^66, 9, Ibembo; 67, 9, Kivu; 68, 9, Lulonga;
69, 70, 9, 3, Bilomba; 71, 9, Zambia.
g4 P. C. BARNARD
j . Antennae up to 70 mm, with up to 75 segments. Scape and pedicel yellowish brown, flagellar segments
golden brown, with a dark line at each joint. Body yellowish brown, abdomen darker brown dorsally.
Thorax with a pair of elongate spots, varying greatly (Figs 58-71; _J, 9) but never round. Wing length
15-26 mm. Wings slightly falcate (Fig. 54), pale green to yellowish brown, with golden brown streaks
apically along the branches of R and M; extreme wing tip often dark brown. Spurs 1.3.3 or 2.3.3.
9. Antennae up to 45mm, otherwise as J1. Wings less obviously falcate than in J, being shorter and
broader. Wing length 16-20 mm. Spurs 1.3.3 or 2.3.3.
GENITALIA J1 (Fig. 55). Similar to bipunctatus, armature of aedeagus with enlarged external opening.
GENITALIA 9 (Fig. 56). Outer corners of eighth sternite rounded; inner thickened edges extending less than
half-way.
REMARKS. This species has caused great confusion to previous authors on account of its great
variability, as well as being confused with bipunctatus. Navas (193 la: 139) described the variety
pupillatus as a form of bipunctatus, not realizing that it was a form of his own species elisabethae
described in the same paper! Lestage (1936) recognized that all variations in the form of the
thoracic spots could occur, including their disappearance, and Barnard (1934) made similar
comments about bipunctatus (in the present paper forms lacking thoracic markings are
recognized as valid species). Marlier (1965) took an opposite view, grouping specimens with
similar markings into 'forms' of bipunctatus. These are in fact forms of elisabethae, and Figs
58-7 1 show the enormous variability of these thoracic markings, even in specimens from the
same locality. I do not think that there is any possibility of subdividing this species in any
meaningful way, and even Navas's pupillatus is here considered to fall within the range of
variation, despite Marlier's (1965) elevation of it to a full species.
DISTRIBUTION (Fig. 72). Sierra Leone, Ghana, Nigeria, Cameroun, Congo (Marlier, 1965), Zaire,
Uganda, Zambia, Zimbabwe.
MATERIAL EXAMINED
Sierra Leone: 5 J1, 11 $, Njala, various dates (Hargreaves) (BMNH). Ghana: 1 9, Wassaw District, 45
miles [72 km] inland from Sekondi (BMNH). Nigeria: 1 9, Lagos, 6 miles [9.6 km] NW. of Agege, at light,
24.ii.1973 (Riley) (BMNH); 1 9, Kagoro Forest, 15-17.X.1971 (Deeming) (BMNH); 2 9, Benin, 8.iv.l973
(Medler) (BMNH). Cameroun: 1 $ (Rosevear) (BMNH). Zaire: 1 9, 150-200 miles [240-320 km] W. of
Kambove, 3500-4000' [1050-1200 m], 28.ix.1907 (Neave) (BMNH); 1 9, Tshuapa, Bamania, x.1951
(Hulstaert); 3 9, Uele, Ibembo, 30.vi.1950, x-xi.1951, ii.1952 (Hutsebaut); 1 3, Albertville [Kalemie],
3.1.1919 (Mayne); 1 9, Mayumbe forest, 29.iii.1973 (Allaer); 1 ?, Kivu, Lubero 950 m, xii.1956 (Celts); 1 9,
Mayumbe, Vaku, 22.V.1970 (Elsen); 1 9, Kimwenza, ix.1962 (Deheegher); 1 9, Kwango, Popokabaka,
iii.1952 (Pierquin); 1 9, Lulonga, 22.ii.1949 (Marlier); 1 9, Irangi, River Luhoho, 8-10.L1957 (Leleup), 2 9,
24.xii.1957 (Marlier); 1 $, 1 <J, Bilomba, Kamituga, 25.viii.1950 (Marlier); 2 9, Bolobo-Lukolela,
10.vi.1951 (Marlier); 1 ^, Mombongo, 29.vi.1951 (Marlier); 2 9, Kisangani, l.vii.1951 (Marlier); 1 3,
Meko, Bolobo, nr Tshumbiri, 9.vi.l951 (Marlier); 5 3, 1 9, Lukanga, at light, 7.vi.l955 (Marlier); 1 3,
Ikela, ix.1959 (Leleup); 1 9, Shabunda, Kiamiseke, at light, 27.x. 1954 (Leleup) (all in MRAC, Tervuren).
Uganda: 3 J, 1 9, Karuma Falls, Victoria Nile (Corbet) (BMNH). Zambia: 2 9, Katombora, Zambezi
River, iv.1962 (Pinhey) (BMNH). Zimbabwe: 1 ^, 1 9, Victoria Falls, i.1956 (USNM, Washington).
Polymorphanisus fuscm (Ulmer)
(Figs 84-86)
Oestropsis fusca Ulmer, 1905a: 42. Holotype 9, SUMATRA: Soekaranda [? Soekaradja] (Dohrn) (type no.
1753, IZPAN, Warsaw) [examined].
Polymorphanisus fuscus (Ulmer) Ulmer, 1 905/7 : 31.
Oestropsis fusca Ulmer; Tomaszewski, 1961 : 4. [Holotype depository.]
J1. Unknown.
9. Antennal length unknown (all specimens damaged). Antennal segments golden brown, with dark
joints. Body pale yellowish brown. Thorax with two oblong, black markings on mesonotum;
mesoscutellum almost covered by large oval spot (Fig. 85). Wings dark smoky brown with blackish brown
spot near base of costa on fore wing (Fig. 84). Wing length 24-26 mm. Spurs 1.3.3.
REVISION OF OLD WORLD POLYMORPHANISINI
85
o-
g
S
a,'
c
.1
T3
a
a
o
c
•S-
•C)
I
B-
s
I
2
a
KJ
a
c
o
86
P. C. BARNARD
76
, 5 mm ,
t 1 mm ,
Figs 74-78 74-76, Polymorphanisus hargreavesi ?, (74) wing venation; (75) head, dorsal
view; (76) eighth sternite, ventral view. 77, 78 P. marlieri$, (77) wing venation; (78) eighth
sternite, ventral view.
REVISION OF OLD WORLD POLYMORPHANISINI 87
GENITALIA 9 (Fig. 86). Sides of eighth sternite tapering posteriorly, outer corners broadly rounded.
REMARKS. Even though the male is unknown, this species is one of the most distinctive of the
genus, with its characteristic dark brown wings and the markings on the thorax and wing base.
DISTRIBUTION. Sumatra, Borneo (Sarawak).
MATERIAL EXAMINED
Borneo: 4 ?, Sarawak, Gunong Mulu National Park, at light, ii-iii.1978 (Holloway et al.) (BMNH).
Polymorphanisus hargreavesi sp. n.
(Figs 73-76)
c£. Unknown.
9- Antennae up to 45 mm, with about 65 segments. Scape yellowish brown, with longitudinal black stripe
(Fig. 75), pedicel and flagellum dark brown or black. Body yellowish brown, abdomen dark brown dorsally,
wings pale yellowish brown. No markings on thorax. Wing length 22-28 mm. Venation as in Fig. 74. Spurs
1.3.3.
GENITALIA 9 (Fig. 76). Outer posterior corners of eighth sternite produced into sharp points.
REMARKS. This species is readily distinguished from bipunctatus and elisabethae by the lack of
thoracic markings, and from marlieri by the dark antennae. The specimens examined show a
discontinuous distribution, with a long series from Sierra Leone, and scattered individuals from
Zaire, Zambia and Zimbabwe. Although I am reasonably certain of the identity of these latter
specimens, I am restricting the type-series to the Sierra Leone material.
DISTRIBUTION (Fig. 73). Sierra Leone, Zaire, Zambia, Zimbabwe.
MATERIAL EXAMINED
Holotype 9, Sierra Leone: Njala, 22.x. 1930 (Hargreaves) (BMNH).
Paratypes. 1 1 9, data as holotype, various dates (BMNH).
Material excluded from paratype series. Zaire: 1 9, 1926 (Jackson) (BMNH); 2 9, Lulonga, 22.ii.1949
(Marlier}; 1 9, Ubangi, ll.viii.1947 (Poll); 3 9, Kwango, Popokabaka, iii.1952 (Pierquin); 1 9, Bolobo,
27.vii.1930 (Beheyu); 1 9, Zaire River, Is. Kui, Stanleyville [Kisangani], l.vii.1951 (Marlier); 1 9, Meko,
Bolobo, 9.vi.l951 (Marlier); all in MRAC, Tervuren. Zambia: 1 9, Katombora, iv.1962 (USNM,
Washington). Zimbabwe: 1 9, Victoria Falls, xii.1938 (USNM, Washington).
Polymorphanisus marlieri sp. n.
(Figs 73, 77, 78)
cJ. Unknown.
9. Antennae 45 mm long, with approximately 75 segments. Antennal segments pale yellow, with brown
annulations at joints. Body and wings yellowish brown, no markings on thorax. Wing length 22, 26 mm
(two specimens only). Venation as in Fig. 77, fork R2 in fore wing shorter than, or just equal in length to,
stem. Spurs 1.3.3.
GENITALIA 9 (Fig. 78). Outer corners of eighth sternite rounded, outer sides slightly sinuous.
REMARKS. This species is named after Dr G. Marlier in recognition of his valuable work on the
African Trichopteran fauna. The lack of thoracic markings and the pale antennae distinguish it
from other African species, its nearest relative probably being hargreavesi sp. n., with dark
antennae.
DISTRIBUTION. (Fig. 73). Zaire.
MATERIAL EXAMINED
Holotype 9, Zaire: Feshi, Kwango, R. Kwenge, iv.1959 (Leleup) (MRAC, Tervuren).
Paratype. Zaire: 1 9, Feshi, R. Kwenge, 850m, at light, ii.1959 (Leleup) (BMNH).
Polymorphanisus muluensis sp. n.
(Figs 87-90)
J. Antennae 40 mm long, with up to 70 segments. Antennal segments pale golden yellow, with dark brown
joints. Body yellowish brown, thorax with two pairs of black markings on the mesoscutellum, the anterior
Figs 79-83 Polymorphanisus astictus. 79, j1 wing venation; 80, _> genitalia, lateral view; 81,
j genitalia, dorsal view; 82, aedeagus, ventral view; 83, ? eighth sternite, ventral view.
pair subtriangular, the posterior subquadrangular (Fig. 87). Estimated wing length 16mm, venation
probably typical of nigricornis-group (specimen damaged). Spurs 1.3.2.
9. General appearance as J1, antennae and wings damaged. Estimated wing length 17mm. Spurs 1.3.3.
GENITALIA 3 (Figs 88, 89). Tenth segment divided medially, each half with a double lobe, viewed dorsally.
Terminal segment of gonopod with clubbed tip, viewed dorso-ventrally.
GENITALIA £ (Fig. 90). Thickened inner edges of eighth sternite extend over half-way down segment. Outer
edges of sternite with definite obtuse angle.
REMARKS. This species is probably most closely related to scutellatus. In the female paratype the
mesoscutellar markings are slightly more elongate medially, and rather resemble the form of
scutellatus from Borneo figured by Ulmer (1951 : pi. 10, fig. 224). I think that the differences in
female genitalia (the males of scutellatus being unknown) are sufficient to warrant describing it as
distinct.
REVISION OF OLD WORLD POLYMORPHANISINI
Sc
89
89
, 0.5mm ,
Figs 84-90 84-86, Polymorphanisus fuscus ?, (84) wing venation; (85) thorax, dorsal view;
(86) eighth sternite, ventral view. 87-90, P. muluensis, (87) 3 thorax, dorsal view; (88) $
genitalia, lateral view; (89) 3 genitalia, dorsal view; (90) $ eighth sternite, ventral view.
90 P- C. BARNARD
DISTRIBUTION. Borneo (Sarawak).
MATERIAL EXAMINED
Holotype J1, Borneo: Sarawak, Gunong Mulu National Park, i.1978 (Holloway el al.) (BMNH).
Paratype. Borneo: 1 9, Sarawak, Gunong Mulu National Park, at light, ii.1978 (Holloway et al.)
(BMNH).
Polymorphanisus nigricornis Walker
(Figs 91-93)
Polymorphanisus nigricornis Walker, 1852: 79. Holotype J, INDIA: north (Stevens) (BMNH) [examined].
Polymorphanisus nigricornis Walker; Betten & Mosely, 1940: 212. [Redescription of holotype.]
cJ. Antennae up to 50 mm long, with about 80 segments. Scape and pedicel yellowish brown, flagellum dark
brown. Body and wings yellowish brown or greenish in fresh specimens. Thorax with two small, rounded,
black spots on mesoscutellum (Fig. 91). Wing length 22-23 mm. Spurs 1.3.3.
9. Antennae up to 45 mm long, with about 65 segments. Scape and pedicel with longitudinal black stripe
externally, flagellum dark brown or black (as in bipunctatus, Fig. 52). Rest of coloration as J1. Wing length
20-29 mm. Spurs 1.3.3.
GENITALIA j1 (Fig. 92). Similar to bipunctatus and elisabethae; terminal segment of gonopod long and
narrow. Ratio of lengths of basal and terminal segments of gonopod 1.8-2.3:1.
GENITALIA 9 (Fig. 93). Eighth sternite tapering strongly posteriorly; outer posterior corners produced
sharply.
REMARKS. Much of the confusion over this species has resulted from Walker's (1852) inadequate,
and even misleading, description. Walker failed to notice the thoracic markings, which are partly
obscured by the pin through the holotype, and he also described the male antennae as black,
rather than brown. Betten & Mosely (1940) corrected these errors, but not before several authors
such as Ulmer (1907) and Banks (1939) had misidentified the species. Fischer (1972: 165) has
suggested that some of the specimens from Java and Sumatra listed by Ulmer (1951) are in fact
scutellatus ; Banks's concept of nigricornis is discussed under ast ictus (p. 79) and umbripes (p. 96).
With most of these misidentifications rectified, it now appears that the true nigricornis is much
more rare and restricted than was previously thought.
Betten & Mosely (1940) suggested that bipunctatus was almost certainly a synonym of
nigricornis, but there are differences in the male and female genitalia which, together with the
widely separated distributions, suggest that the two should be kept distinct.
DISTRIBUTION. India, Vietnam, Sumatra, Java.
MATERIAL EXAMINED
India: 1 j1, no further data; 1 9 (Sounders); 2 9, Manipur, Imphal (Ram). Vietnam: 1 9, Cha Pa, viii.1936
(Masseyeff). Sumatra: 1 9, 1892 (Forbes). Java: 1 9 (Horsfield). (All in BMNH.)
Polymorphanisus quadripunctatus Ulmer
(Figs 94-100)
Polymorphanisus quadripunctatus Ulmer, 1951 : 186. Holotype 9, BORNEO: Nanga Raven [? Nangaraun], vii.
1907 (Buttikofer) (ZM, Hamburg) [examined].
Polymorphanisus quadripunctatus Ulmer; Weidner, 1964: 91. [Holotype depository.]
cJ. Antennae 35 mm long, with about 70 segments. Antennal segments pale golden yellow, with dark brown
joints. Body yellowish brown, thorax with four black markings, two on mesoscutum and two on
metascutum (Fig. 94). Wings greenish yellow, wing length estimated 1 8 mm (single specimen damaged).
Spurs 1.3.3, spur on fore leg very long and curved.
9. Similar to J1, wing length 17-23 mm. Venation as in Fig. 98. Spurs 1 .3.3, spur on fore leg curved as in 5
(Fig. 99).
GENITALIA J (Figs 95-97). Distal segment of gonopod long and narrow. Aedeagus with prominent disc-like
enlargement at apex, not gradually thickened.
91
Figs 91-100 91-93, Polymorphanisus nigricornis, (91) $ thorax, dorsal view; (92) $
genitalia, lateral view; (93) ? eighth sternite, ventral view. 94-100, P. quadripunctatus, (94)
J genitalia, dorsal view; (95) 3 genitalia, lateral view; (96) 3 genitalia, dorsal view; (97)
aedeagus, lateral view; (98) y wing venation; (99) 2 fore leg; (100) v eighth sternite,
ventral view.
92 P- C. BARNARD
GENITALIA $ (Fig. 100). Sides of eighth sternite strongly rounded, with prominent setigerous projection on
inner posterior corners.
REMARKS. This is the first time that the male of this species has been described ; the shape of the
aedeagus of the single male specimen is strikingly different from all other species in the genus.
DISTRIBUTION. Borneo (Sarawak), Philippines (Palawan).
MATERIAL EXAMINED
Borneo : 2 9, Sarawak, Bidi (Brooks) ; 2 9, Sarawak, foot of Mt Dulit, junction of rivers Tinjar and Lejok,
l-15.ix.1932 (Hobby & Moore); I J, 14 9, Sarawak, Gunong Mulu National Park, at light, ii-iv.1978
(Holloway et al.). (All specimens in BMNH.) Philippines: 1 9, Palawan, Chromite Mine, 28 km W. Puerto
Princessa, l-7.xii.1965 (Davis) (USNM, Washington).
Polymorphanism scutellatus Banks
(Figs 101-103)
Polymorphanisus scutellatus Banks, 1939: 55. Holotype 9, BORNEO: Sarawak, Baram River district, 1912
(Smith) (type no. 23472, MCZ, Harvard) [examined].
Polymorphanisus scutellaris Banks; Kimmins, 1955: 399. [Incorrect subsequent spelling of scutellatus
Banks.]
J1. Unknown.
9. Antennae up to 50 mm long, with about 95 segments. Antennal segments yellowish brown, slightly
darker distally. Body colour yellowish brown, thorax with two very large oval black markings on
mesoscutellum, which meet sides of sclerite (Fig. 102). Wings pale green or yellowish, wing length
16-24 mm. Venation as in Fig. 101, fork R2 in fore wing shorter than, or just equal to, its stem. Spurs 1.3.2.
GENITALIA 9 (Fig. 103). Plates of eighth sternite long and narrow, with broadly rounded excision in outer
sides.
REMARKS. All the specimens of scutellatus that I have examined have remarkably constant
thoracic markings, yet two variations were figured by Ulmer (1951 : pi. 10, figs 222, 224). The
latter form may be referable to muluensis (see p. 88) and the former may conceivably be
unipunctus ; only the collection of a great deal more material from the Indochina peninsula could
help to resolve these problems.
DISTRIBUTION. Sumatra, Java (Ulmer, 1951), Borneo (Sarawak), Sulawesi.
MATERIAL EXAMINED
Sumatra: 2 9, Medan, 4.ix.l921, Pagar Marbau, 24.vii.1921 (Corporaal) (IRSNB, Brussels). Borneo: 3 9,
Sarawak, foot of Mt Dulit, junction of rivers Tinjar and Lejok, at light, 28.viii-l.ix.1932 (Hobby & Moore);
1 9, Sarawak, Long Lama, at light, 2. i. 1967 (Kueh); 1 9, Sarawak, Gunong Mulu National Park, Batu base
camp, at light, 27. v. 1978 (Hammond & Marshall); 5 9, Sarawak, Gunong Mulu National Park, at light,
i-iv.1978 (Holloway et al.). Sulawesi: 2 9, Maros, ix.1923 (Brooks); 2 9, Macassar. (All specimens in
BMNH.)
Polymorphanisus semperi (Brauer)
(Figs 104-109)
Oestropsis semperi Brauer, 1868: 264. LECTOTYPE J, PHILIPPINES: Mindanao, vii-x.1864 (Semper)
(IRSNB, Brussels), here designated [examined].
Polymorphanisus semperi (Brauer) Ulmer, 1907: 23.
J. Antennae 50 mm long, with about 90 segments. Scape and pedicel with faint longitudinal brown stripe
externally, flagellar segments dark golden brown. Body yellowish brown, thorax with a pair of elongate
dark brown or black markings on mesoscutellum, not touching sides of sclerite (Fig. 105). Wings falcate,
greenish yellow, with golden yellow stripes along the apical veins. Wing length 19-22 mm, venation as in
Fig. 104. Spurs 1.3.3.
9. Antennae 40 mm long, with up to 85 segments. General coloration as 5 , but scape and pedicel usually
lacking brown stripe. Thoracic markings similar to J1, but often fainter. Wings not falcate, and without
apical stripes; venation as in Fig. 108. Wing length 21-27 mm. Spurs 1.3.3.
REVISION OF OLD WORLD POLYMORPHANISINI
93
101
Figs 101-103 Polymorphanisus scutellatus . 101, wing venation; 102, thorax, dorsal view;
103, eighth sternite, ventral view.
GENITALIA J1 (Figs 106, 107). Terminal segment of gonopod stout. Tenth segment bilobed, viewed dorsally.
GENITALIA 9 (Fig. 109). Plates of eighth sternite straight-sided, subquadrangular, with only slightly rounded
posterior corners.
REMARKS. I have been unable to identify the type-localities cited by Brauer (1868) as 'Dugang'
[? = Duguan] on Mindanao, and 'Baubo'. None of the four syntypes examined was labelled with
a locality. I am including the two specimens in the BMNH in the syntype series because, as well
as having been labelled as types by McLachlan (from whose collection they originated), one
bears a small label with the number '546'. One of the syntypes from the IRSNB, Brussels, bears
the number '545' in the same hand, suggesting that both were in the same collected series,
although the significance of the numbers is unknown. Brauer did not indicate how many
specimens constituted his type-series.
DISTRIBUTION. Philippines.
Figs 104-109 Polymorphanisus semperi. 104, J1 wing venation; 105, J1 thorax, dorsal view;
106, J1 genitalia, lateral view; 107, J genitalia, dorsal view; 108, $ wing venation; 109, $
eighth sternite, ventral view.
REVISION OF OLD WORLD POLYMORPHANISINI 95
MATERIAL EXAMINED
Philippines: 2 J, 1 9, Mindanao, vii-x.1864 (Semper) (1 $ in IRSNB, Brussels; 1 j1, 1 9 in BMNH)
(paralectotypes) ; 8 J1, no further data (BMNH); 2 J1, 3 ?, Luzon, Mt Makiling; 2 ^, Mindanao, Surigao
(Baker) (USNM, Washington).
Polymorphanisus taoninus Navas
(Fig. 110)
Polymorphanisus taoninus Navas, 1936: 128. Holotype 9, CHINA: Kwangsi-Chuang, Wuchow, 9. v. 1934
(Tao) (depository unknown) [not examined].
J1. Unknown.
9 (translated from Navas, 1936). 'Head yellow-fulvous; eyes black; antennae black, first two segments
same colour as head, apical segments testaceous, bases of segments dark; longer than 25mm. Thorax
whitish green below, greenish fulvous above. Abdomen yellowish green? (badly preserved). Legs fulvous,
second tarsi fulvous-green. Wings unmarked; membrane lightly coloured fulvous-green; venation fulvous-
yellow. Anterior wing [Fig. 1 10]; apical fork 1 [fork R2] shorter than its stalk; discal cell short and broad,
wider than long, outer margin perpendicular to radial sector ; median cell more than three times as long, a
little wider . . . [etc.]'
'Wing length 23 mm.'
REMARKS. Navas (1936) describes this species as being similar to ast ictus, but the dark antennae
and short fork R2 are very similar to unipunctus (described on p. 96). It is possible that Navas
may have overlooked a central thoracic spot, which could easily be obscured by the pin, in which
case Navas's name would take priority over unipunctus.
DISTRIBUTION. China (Kwangsi-Chuang).
Polymorphanisus tumidus Banks
(Figs 111-113)
Polymorphanisus tumidus Banks, 1939: 54. Holotype 9, INDIA: Mysore, Shimoga, R. Tunga, 1865' [560 m],
at light, 17.iv.[? year] (Nathan) (type no. 23468, MCZ, Harvard) [examined].
J. Unknown.
?. Antennal length unknown (specimen damaged). Scape with rounded black spot anteriorly (Fig. 1 12),
other segments greenish yellow. Head yellow, with greatly swollen, green frons (Fig. 112). Rest of body
greenish yellow, thorax with two elongate black spots, each expanded postero-laterally (Fig. 112). Wing
length 20mm (holotype only), fore wing venation as in Fig. Ill : fork R2 very long. Spurs 1.3.3.
GENITALIA 9 (Fig. 1 13). Plates of eighth sternite subquadrangular, with moderately rounded outer corners.
REMARKS. The curiously inflated frons of the unique holotype of this species appears
teratological, yet the thoracic and antennal markings distinguish it from all other species of the
genus.
DISTRIBUTION. India (Mysore).
MATERIAL EXAMINED
Holotype only.
Polymorphanisus umbripes sp. n.
(Figs 114-116)
[Polymorphanisus nigricornis Walker; Banks, 1939: 53. Misidentification.]
c£. Unknown.
9. Antennae up to 40 mm long, with about 75 segments. Scape and pedicel greenish yellow, scape with
small dark brown spot distally (occasionally absent), pedicel with lateral brown stripe ; flagellar segments
dark brown (Fig. 1 15). Body yellowish brown, abdomen darker dorsally; thorax unmarked. Forelegs with
most of femur and whole of tibia dark brown; tarsi yellowish brown. Spurs 1.3.3. Wing length 21-24 mm,
venation as in Fig. 114.
GENITALIA 9 (Fig. 116). Eighth sternite tapering posteriorly, outer posterior corners rounded in a gentle
curve.
96
P. C. BARNARD
R.
111 5mm
<J
r T
113
1 mm
112
, 1mm
Figs 110-113 1 10, Polymorphanisus taoninus ?, part of fore wing (after Navas). 1 1 1-113, P.
tumidus ?, (Ill) fore wing venation; (112) head and thorax, dorsal view; (113) eighth
sternite, ventral view.
REMARKS. This species is based on the series of specimens which Banks (1939) assumed to be
nigricornis Walker (see remarks under nigricornis, p. 90). The two Indian localities of this species
are the same for Banks' sflavipes ( = ast ictus) and this would suggest that umbripes may be merely
a colour variant of the latter with dark antennae and forelegs, but the differences in female
genitalia are also consistent in the specimens examined.
DISTRIBUTION. India (Mysore).
MATERIAL EXAMINED
Holotype $, India: Mysore, Shimoga, R. Tunga, 1865' [560 m], at light, IS.vi. [? year] (Nathan) (type no.
32396, MCZ, Harvard).
Paratypes. India: 29 $, data as holotype, various dates (26 in MCZ, Harvard; 2 in USNM, Washington;
1 in BMNH); 4 $, Mysore, Bhadravati (Nathan) (MCZ, Harvard).
Polymorphanisus unipunctus Banks
(Figs 117-119)
Polymorphanisus unipunctus Banks, 1939: 53. Holotype 9, CHINA: Szechwan, Suifu [=Ipin], viii.1928
(Graham) (type no. 53164, USNM, Washington) [examined].
Polymorphanisus unipunctus Banks; Banks, 1940: 206. [Redescription as new species.]
REVISION OF OLD WORLD POLYMORPHANISINI
Sc
97
Figs 114-119 114-116, Polymorphanisus umbripes ?, (114) wing venation; (115) head,
dorsal view; (116) eighth sternite, ventral view. 117-119, P. unipunctus $, (117) wing
venation; (118) thorax, dorsal view; (119) eighth sternite, ventral view.
98 P. C. BARNARD
c£. Unknown.
9. Antennal length unknown (specimen damaged). Scape and pedicel yellow, flagellar segments dark
brown. Body yellowish brown, thorax with a single, partially divided, black spot in anterior half of
mesoscutellum (Fig. 118). Wing length 25 mm, venation as in Fig. 1 17; fork R2 shorter than its stem. Spurs
1.3.2.
GENITALIA 9 (Fig. 119). Setigerous posterior margin of each half of eighth sternite produced in a rounded
lobe.
REMARKS. The possible similarity of this species with Navas's description oftaoninus is discussed
under the latter species. Banks's (1939) first description of this species was obviously intended to
be published later than his second paper (1940). In the Trichopterorum Catalogus Fischer (1963 :
209) lists the first reference as 'nomenclatorially invalid', presumably as a nomen nudum, but in
fact this first (1939) paper contains sufficient detail to constitute a valid description.
DISTRIBUTION. China (Szechwan).
MATERIAL EXAMINED
Holotype only.
The ocularis-group
Pale greenish white species, never with markings on thorax. Wings broad, with brown markings (absent in
angustipennis). Antennae up to one and a half times fore wing length. Male eyes large, almost meeting
ventrally. Spurs 1.3.2. In fore wing M, is direct continuation of Ml+2 stem, M2 arises from median cell. In
hind wing Rl ends on Sc. Basal segment of male gonopods expanded, terminal segment not differentiated.
Polymorphanisus angustipennis Ulmer
(Figs 120-128)
Polymorphanisus angustipennis Ulmer, 1912: 97. Holotype j1, CAMEROUN: Ma[y]o Godi, 9-13. vi. 1909
(Riggenbach) (MNHU, Berlin) [examined].
Polymorphanisus sp.; Marlier & Botosaneanu, 1968: 11. [Compared with P. similis Ulmer.]
cJ. Antennae up to 20 mm long, with about 50 segments. Antennae and rest of body pale yellowish brown,
legs greenish, wings greenish white, with no dark markings. Eyes very large, almost meeting ventrally (Fig.
122). Wing length 12-14 mm, venation as in Fig. 120. Spurs 1.3.2.
?. Antennae and general coloration as in £. Eyes small, well separated ventrally (Fig. 127). Wing length
15-17 mm, venation as in Fig. 125. Spurs 1.3.2.
GENITALIA J (Figs 123, 124). Ninth segment bifurcate dorsally. Gonopods only moderately expanded
basally.
GENITALIA 9 (Fig. 128). Eighth sternite long and narrow, straight-sided, with only slightly roundec
posterior corners.
REMARKS. This species is easily distinguished from similis, with which it shares a simila
distribution, by the absence of wing markings.
DISTRIBUTION. Ghana, Nigeria, Cameroun, Sudan, Uganda.
MATERIAL EXAMINED
Ghana: 1 ^, Volta R., Yeji, 14.X.1950 (Berner). Nigeria: 1 9, Kaduna, 16.X.1957; 16 9, Samaru, at light
7-14.vii.1970 (Ward); 38J, 8 9, Lake Kainji, 1975 (Bidwell). Sudan: 2 9, Malakal-Shambe, 8-14.xii.196
(Cloudsley-Thompsori). Uganda: 2 9, Ugweano, 22.viii.1949 (Lowe); 2 9, Lake Victoria, Bukakata
13.xii.1950 (Lowe), 6.viii.l959 (Corbet); 2 J1, Mengo, Entebbe, 28-30.iii.1956 (Corbet). (All specimens in
BMNH.)
Polymorphanisus guttatus Navas
(Figs 133, 134)
Polymorphanisus guttatus Navas, 1935: 71. Holotype 9, MADAGASCAR: Ambodirefia, 70km NW. o
Tamatave, ii.1934 (MNHN, Paris) [examined].
3*. Unknown.
122 . 1mm
REVISION OF OLD WORLD POLYMORPHANISINI
Sc
99
. 0.5mm
Figs 120-124 Polymorphanisus angustipennis j1. 120, wing venation; 121, head, dorsal
view; 122, head, ventral view; 123, genitalia, lateral view; 124, genitalia, dorsal view.
?. Antennal length at least 25 mm (specimens damaged). Scape and pedicel yellow, first flagellar segment
dark brown distally, rest of flagellum dark brown. Body yellowish brown. Wing length 16-20 mm,
membrane greenish, with one brown spot on median cell of fore wing, at base of M2 and M3 (Fig. 133).
Spurs 1.3.2.
GENITALIA ? (Fig. 134). Eighth sternite broad centrally, posterior corners obtusely rounded. Setigerous lobe
on posterior margin strongly produced.
REMARKS. This species, apparently endemic to Madagascar, is easily recognized by the dark
antennae and single spot on the fore wing.
DISTRIBUTION. Madagascar.
MATERIAL EXAMINED
Madagascar: 1 <?, Station Perinet, 149km E. of Tananarive, 20.x-10.xi.1930 (d'Olsoufieff) (BMNH).
100
P. C. BARNARD
Figs 125-128 Polymorphanisus angustipennis $. 125, wing venation; 126, head, dorsal view;
127, head, ventral view; 128, eighth sternite, ventral view.
Polymorphanisus ocularis Ulmer
(Figs 135-138)
Polymorphanisus ocularis Ulmer, 1906: 60. LECTOTYPE ?, JAVA (Piepers) (RNH, Leiden), here
designated [examined].
Polymorphanisus indicus Banks, 1911 : 105. Holotype ?, INDIA: Bengal, Pusa, at light, 4.viii.l909 (type no.
11787, MCZ, Harvard) [examined]. Syn. n.
Polymorphanisus ocularis Ulmer, unnamed var. ; Martynov, 1935: 195.
$. Antennae up to 23 mm, with about 50 segments. Antennal segments pale golden yellow. Body pale
yellowish white, abdomen yellowish brown dorsally. Wings white, occasionally greenish, with two brown
REVISION OF OLD WORLD POLYMORPHANISINI
101
0.5mm .
130
0.5mm
131 . 5mm .
132
5 mm
1 mm
Figs 129-134 129-132, Polymorphanisus similis, (129) ? genitalia, lateral view; (130) $
genitalia, dorsal view; (131) 9 wing venation; (132) $ eighth sternite, ventral view. 133,
134, P. guttatus $, (133) wing venation; (134) eighth sternite, ventral view.
102
P. C. BARNARD
135 • 2mm . R,
Cu
137
. 0.5mm .
138
1 mm
Figs 135-138 Polymorphanisus ocularis. 135, j1, wing venation; 136, j1 genitalia,
lateral view; 137, j1 genitalia, dorsal view; 138, ? eighth sternite, ventral view.
spots on fore wing, one on discal cell at base of R2 + 3, the other on median cell at base of M2 and M3 (Fig.
135). Wing length 14-16 mm. Spurs 1.3.2.
9. Antennae up to 22 mm, with about 50 segments. Coloration as in j1. Wing length 14-18 mm. Spurs
1.3.2.
GENITALIA $ (Figs, 136, 137). Ninth segment produced in a squarish lobe dorsally. Gonopods relatively
narrow at base, thickened in centre.
GENITALIA ? (Fig. 138). Outer posterior corners of eighth sternite produced into rounded lobes, almost as
long as setigerous lobes on inner corners.
REVISION OF OLD WORLD POLYMORPHANISINI JQ3
REMARKS. Banks's (1911) species indicus was described without reference to ocularis, although he
later (1939) listed both in the same paper. Martynov (1935) remarked that he could not separate
the two, and Ulmer (1951) noted their great similarity; in fact they cannot be separated and
indicus is here synonymized.
Ulmer described ocularis from one male and one female, but the male is now apparently
missing (Geijskes, in litt.}. I have therefore designated the female as lectotype.
DISTRIBUTION. India, Sri Lanka, Burma (Martynov, 1935), China (Fuchow), West Malaysia,
Borneo (Sarawak), Java, Sumatra (Ulmer, 1951).
MATERIAL EXAMINED
1 .?, 1 9, no data (Chapman) (BMNH). Sri Lanka: 1 j1 Colombo, iii.1927 (BMNH); 2 9, Marai Villu,
19-21. iii. 1933 (BMNH); 1 9, Marichchukkaddi, 28.iii.1933 (BMNH); 1 9, Puttalam (BMNH); 1 9,
Boyagama, at light, 20-2 l.viii. 1973 (Ekis) (USNM, Washington); 2 9, Wilpattu Park, Talawila, at light,
9-10.iv.1973 (Baumann & Cross) (USNM, Washington), 1 9, Hunuwilagama, Wilpattu, 10-19.iii.1970
(Davis & Rowe) (USNM, Washington); 2 9, Polonnaruwa, 16-21. iii. 1954, 1 9, Mi Oya, 25.iii.1954 (Schmid)
(USNM, Washington). China: 1 9, Fuchow (Yang) (BMNH). West Malaysia: 1 9, Kuala Pilah (BMNH);
3 9, Kuala Lumpur, 28.vi.1929, 23.iv.1931, 6.vi.l932 (Pendlebury) (BMNH); 2 9, Kota Tinggi, Johore,
viii.1917 (BMNH). Borneo: 1 3, no further data (BMNH); 2 9, Sarawak, Gunong Mulu National Park,
i-iv.1978 (Holloway et al.) (BMNH).
Polymorphanisus similis Ulmer
(Figs 129-132)
Polymorphanisus similis Ulmer, 1912 : 96. Holotype 3, CAMEROUN : Lolodorf, 16.V.1896 (Conradt) (MNHU,
Berlin) [examined].
Polymorphanisus bisignatus Navas, 19316: 276. Holotype 9, ZAIRE: Katanga, Kafakumba, 16.V.1925
(Overlaet) (MRAC, Tervuren) [examined]. Syn. n.
c?. Antennal length 23 mm (holotype), antennal segments pale yellow. Body pale yellow, abdomen darker
dorsally. Wings white, occasionally greenish, with two brown spots on fore wing, one on discal cell at base
of R2 + 3 and one on median cell at base of M2 and M3. Wing length 13-16 mm. Spurs 1.3.2.
9. Antennal length up to 30 mm, with up to 60 segments. Coloration as j\ wing venation as in Fig. 131.
Wing length 16-23 mm. Spurs 1.3.2.
GENITALIA <$ (Figs 129, 130). Ninth segment produced into a broad rounded lobe dorsally. Gonopods very
broad at base, narrowing suddenly in centre.
GENITALIA 9 (Fig. 132). Eighth sternite gradually tapering posteriorly, posterior edge sloping evenly to
setigerous lobes.
REMARKS. Although Ulmer's descriptions were usually accurate, he mistakenly described the
spur formula of this species as being 2.3.3 (Ulmer, 1912). The male holotype is clearly seen to
have the spur formula 1.3.2, typical of this group. This inaccuracy has confused authors such as
Marlier (1961) who draws attention to the apparent discrepancy between his female specimen
and the description of the male holotype. However, Marlier & Botosaneanu's (1968)
'Polymorphanisus cf. similis'' is angustipennis, as shown by the lack of wing markings.
DISTRIBUTION. Sierra Leone, Nigeria, Cameroun, Zaire, Uganda.
MATERIAL EXAMINED
Sierra Leone: 1 9, Bo, ix.1968 (Revell); 2 9, Njala, 1 l.viii. 1929, ix.1934 (Hargreaves). Nigeria: 1 J, Ilesha
(Humfrey); 1 9, Zungeru, xi.1910 (Simpson); 1 9, Abuja, at light, 21.xi.1970 (Deeming); 1 9, Kagoro Forest,
15-17.X.1971 (Deeming). (All specimens in BMNH.) Uganda: 1 9, Ankole, 25 m [40 km] S. of Kichwamba,
Kalinzu Forest, 28.iv.1968 (Spangler) (USNM, Washington).
References
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104 P- C. BARNARD
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— 1957. Lectotypes of Trichoptera from the McLachlan collection now in the British Museum (Natural
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1962. New African caddis-flies (Order Trichoptera). Bull. Br. Mus. nat. Hist. (Ent.) 12: 81-121.
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Nat. Mosc. 11: 141-296.
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[English translation.]
Lestage, J. A. 1919. Les Trichopteres d'Afrique. Catalogue synonymique et systematique des especes
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- 1936. Notes trichopterologiques. XIV. Les composantes de la faune sud-africaine et la dispersion
transafricaine de quelques especes. Bull. Annls Soc. r. ent. Belg. 76: 165-192.
Marlier, G. 1943. Trichopteres du Congo beige. Revue Zool. Bot. afr. 37: 64-88.
- 1958. Recherches hydrobiologiques au Lac Tumba. Hydrobiologia 10: 352-385.
- 1961. Hydropsychidae du Kivu (Trichoptera). Revue Zool. Bot. afr. 63: 158-212.
- 1962. Genera des Trichopteres de 1'Afrique. Annls Mus. r. Afr. cent. Ser. 8°. Sciences Zoologiques.
109: 1-263.
- 1965. Les Trichopteres du musee de Dundo. Publcoes cult. Co. Diam. Angola 72: 13-80.
1978. Sur une collection de Trichopteres de 1'Afrique occidentale. Revue zool. afr. 92: 283-302.
Marlier, G. & Botosaneanu, L. 1968. Trichopteres du Ghana et de la Cote d'lvoire. Bull. Inst. r. Sci. nat.
Belg. 44 (16): 1-16.
Martynov, A. V. 1910. The Trichoptera of Siberia and adjacent regions. [In Russian.] Ezheg. zool. Mus. 15:
351-429.
- 1930. On the Trichopterous fauna of China and eastern Tibet. Proc. zool. Soc. Land. 1930: 65-1 12.
- 1934. Trichoptera Annulipalpia of the U.S.S.R. [In Russian, English summary.] Opred. Faune SSSR
13: 1-343.
1935. On a collection of Trichoptera from the Indian Museum. Rec. Indian Mus. 37: 93-209.
McLachlan, R. 1878. A monographic revision and synopsis of the Trichoptera of the European fauna. Part VII.
pp. 349-428. London.
— 1880. A monographic revision and synopsis of the Trichoptera of the European fauna. Supplement. Part
II. pp. 13-84. London.
Navas, L. 1915. Notes sur quelques Nevropteres du Congo beige. Revue zool. afr. 4: 172-182.
- 1916. Neuropteros nuevos o poco conocidas. Mems R. Acad. Cienc. Artes Barcelona 12: 219-243.
- 1923a. Algunos insectos del museo de Paris. Revta Acad. Cienc. exact, fis. quim. nat. Zaragoza 7
(1922): 15-51.
REVISION OF OLD WORLD POLYMORPHANISINI 105
- 19236. Insecta nova. Memorie Accad. pent. Nuovi Lincei (2) 6: 19-27.
- 1926. Nevropteres d'Egypte et de Palestine. Bull. Soc. ent. Egypte 19: 192-216.
- 193 la. Insectes du Congo beige. Revue zool. afr. 21: 123-144.
- 19316. Insectes du Congo beige. Revue zool. afr. 20: 257-279.
- 1935. Insectos de Madagascar. Revta Acad. Cienc. exact, fis. quim. nat. Zaragoza 18 (1934): 42-74.
- 1936. Nevropteres et insectes voisins. Notes Ent. chin. 3: 117-132.
Ross, H. H. 1952. Lectotypes of Hagen species belonging to certain families of Trichoptera. Psyche, Camb.
59: 31-36.
Schmid, F. 1958. Trichopteres de Ceylan. Arch. Hydrobiol. 54: 1-173.
Scott, K. M. F. 1975. The value of larval stages in systematic studies of the Trichoptera, with particular
reference to the Hydropsychidae from Africa south of the Sahara. Proc. 1st Congr. ent. Soc. S. Afr.
1975: 41-52.
Tomaszewski, C. 1961. List of type specimens in the collection of the Institute of Zoology of the Polish
Academy of Sciences in Warszawa. IV. Caddis flies (Trichoptera). Annls zool. Warsz. 20: 1-6.
Ulmer, G. 19050. Zur Kenntniss aussereuropaischer Trichopteren. Stettin, ent. Ztg 66: 3-119.
- 19056. Uber die geographische Verbreitung der Trichopteren. Z. wiss. InsektBiol. 1: 16-32.
- 1906. Neuer Beitrag zur Kenntnis aussereuropaischer Trichopteren. Notes Leyden Mus. 28: 1-116.
- 1907. Trichopteren. Monographic der Macronematinae. Collections Zoologiques du Baron Edm. de
Selys Longchamps 6 (2): 1-121. Brussels.
- 1912. Trichopteren von Aquatorial-Afrika. Ergebn. Zw. dt. zent. Afr. Exped. 4: 81-125.
- 1926. Beitrage zur Fauna sinica. III. Trichopteren und Ephemeropteren. Arch. Naturgesch. 91 (A) 5
(1925): 19-110.
- 1930. Trichopteren von den Philippinen und von den Sunda-Inseln. Treubia 11: 373^498.
— 1933. Aquatic insects of China. IV. Neue chinesische Trichopteren. Peking nat. Hist. Bull. 7: 135-157.
— 1951. Kocherfliegen (Trichopteren) von den Sunda-Inseln (Teil I). Arch. Hydrobiol. Suppl. 19: 1-528.
1957. Kocherfliegen (Trichopteren) von den Sunda-Inseln (Teil III). Arch. Hydrobiol. Suppl. 23:
109-470.
Walker, F. 1852. Catalogue of the specimens of Neuropterous insects in the collection of the British Museum.
Part I. 192 pp. London.
Weidner, H. 1964. Die entomologischen Sammlungen des Zoologischen Staatsinstituts und Zoologischen
Museums Hamburg. Mitt. hamb. zool. Mus. Inst. 62: 55-100.
Index
Synonyms are in italics; main references are in bold. References to the Check-list (p. 63) are not included.
Aethaloptera 60, 61, 63, 64, 66, 68, 75 gracilis 66, 68, 75
Amphipsyche 60 guttatus 79, 98
angustipennis 79, 98, 103
astictus 78, 79, 90, 95, 96 hageni 66
hainanensis 79, 81
bipunctatus 63, 78, 81, 84, 87, 90 hargreavesi 78, 87
bisignatus 103
Blepharopus 60 .
mdicus 100, 103
Chloropsyche 66, 75
Macronema 59, 60, 61, 63
dispar 63, 66, 68, 69, 70, 72, 74 Macronematidae 60
dyakana 75 Macronematinae 59, 60
Macronematini 59, 60, 61, 63, 64
elisabethae 63, 78, 81, 84, 87, 90 maerina 68
evanescens 63, 66, 68, 72, 74 maesi 66, 68, 69
marlieri 78, 87
flavipes 79, 81, 96 maxima 68, 69, 70, 72, 81
fuscus 79, 84 muluensis 79, 87
106
nigricornis 78, 79, 81, 90, 95, 96
ocularis 79, 81, 100, 103
Oestropsidae 60
Oestropsinae 60
Oestropsis 60, 78
Oestropsyche 60, 61, 63, 64
Oestropsychinae 60
Oestropsychini 60, 61
palingenia 64
Paraethaloptera 66, 68
Phanostoma 60
Polymorphanisus 60, 61, 63, 64, 78
Primerenca 66, 68
punctata 75, 77, 78
pupillatus 81, 84
P. C. BARNARD
quadripunctatus 78, 90
rossica 63, 74, 75
scutellaris 92
scutellatus 79, 88, 92
semperi 78, 79, 92
sexpunctata 63, 68, 75, 77
similis 79, 81, 98, 103
Synoestropsis 59, 63, 64
taoninus 78, 95, 98
tumidus 79, 95
umbripes 78, 79, 90, 95, 96
unipunctus 79, 92, 95, 96
vitrina 63, 64
British Museum (Natural History)
Monographs on Trichoptera
A revision of the genus Leptonema (Trichoptera). M. E. Mosely
1933, 69 pp, 200 text figures, frontispiece, 8vo, £2.20
The Francis Walker types of Trichoptera in the British Museum
(Natural History). (Re-described and figured). C. Bretten and M. E.
Mosely
1940, ix + 248 pp, frontispiece, 122 text figures, 4to £7.75
The Trichoptera (Caddis-Flies) of Australia and New Zealand. M. E.
Mosely and D. E. Kimmins
1953, 550 pp, 364 text figures, £10.50
Titles to be published in Volume 41
A revision of the western Palaearctic species of the genus Meteorus
(Hymenoptera: Braconidae). By T. Huddleston.
A revision of the Old World Polymorphanisini (Trichoptera : Hydropsychidae).
By P. C. Barnard.
A review of the genera of Neotropical Encyrtidae (Hymenoptera:
Chalcidoidea). By John S. Noyes.
A revision of the British species of Dendrocerus Ratzeburg (Hymenoptera:
Ceraphronoidea) with a review of their biology as aphid hyperparasites.
By N. D. M. Fergusson.
A revision of the Pacific species of Conocepkalus Thunberg (Orthoptera :
Tettigoniidae). By Linda M. Pitkin.
Printed by The Whitefriars Press Ltd, London and Tonbridge
^•'
CD
Bulletin of the
British Museum (Natural History
A review of the genera of Neotropical
Encyrtidae (Hymenoptera : Chalcidoidea)
John S. Noyes
Entomology series
Vol 41 No 3 30 October 1980
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World List abbreviation: Bull. Br. Mus. nat. Hist. (Ent.)
Trustees of the British Museum (Natural History), 1980
ISSN 0524-6431 Entomology series
Vol 41 No 3 pp 107-253
British Museum (Natural History)
Cromwell Road
London SW7 5BD Issued 30 October 1980
A review of the genera of Neotropical Encyrtidae
(Hymenoptera : Chalcidoidea) W LIB-,
John S. Noyes
Department of Entomology, British Museum (Natural History), Cromwell Road, London SW7
5BD
Contents
Synopsis 107
Introduction 107
Notes on terms and measurements 108
Abbreviations Ill
Key to genera (females) Ill
Key to genera (males). . . . . . . . . . . . .150
Notes on genera 169
Host index 236
Proposed new synonymies 238
Proposed new combinations 238
Lectotype designations ............ 239
Acknowledgements 239
References 239
Index 246
Synopsis
Keys are presented to both sexes of the 148 genera* of Encyrtidae known from the Neotropical region.
Notes on each genus are also included; these give information on known world distribution, number of
species known, distribution of the genus within the Neotropics, a list of species known from the region,
notes on biology, systematic placement and references to original descriptions, redescriptions, revisions or
other useful papers. Lectotypes are designated for six species, 19 new genera and 18 new species are
described, 16 new generic synonymies and 1 new specific synonymy are proposed.
Introduction
The Neotropical Encyrtidae, probably more diverse than those of any other part of the world
(except perhaps Australasia), have been little studied beyond the work of De Santis (1964) and
remain poorly known. So far 148 genera* and 380 species are recognized as occurring in the
region covered by the present study, i.e. South and Central America, including associated islands
and the whole of Mexico, the Antilles from Trinidad to Cuba, and Bermuda; it is very likely that
this is only a small proportion of the number that actually occur there. Over 20 000 specimens
comprising well over 500 species have been examined in the course of this study; many of these
species are undescribed and belong to unknown genera. Nineteen genera are here described as
new and as many as twice this number are ignored because of the condition of the material or
because they are represented by single specimens. It is possible that a number of the genera
considered here as valid may eventually fall in synonymy when more is known about the family
as a whole. Some may have already been described by Girault from Australia, since there is a
close similarity between the eastern Australian and Neotropical encyrtid faunas, but this
,
Bull. Br. Mus. nat. Hist. (Em.) 41 (3): 107-253 Issued 30 October 1980
* Note added while this paper was in press. De Santis (1979) also includes the following genera not dealt with in this
review: Asitus Erdos, Boucekiella Hoffer, Cerapterocerus Westwood, Platencyrtus Ferriere and Tachinaephagus
Ashmead.
108 JOHN S. NOYES
possibility has had to be ignored because of the general unavailability of the Girault type-
material. In this review a species has been placed in an already described genus whenever
possible, even though it may be considered somewhat out of place there.
An examination of collections received after the first draft of the keys was completed has
shown that about a third of the material belonged to genera not included (described or
undescribed), or to aberrant species that did not key out as expected (e.g. a Zeteticontus sp. with
punctate sculpture on the scutellum or a Metaphycus sp. with a long exserted ovipositor). It
would therefore seem unlikely that anyone using the keys could expect better than a 60 per cent
success rate, even if the keys are used correctly. The keys are almost entirely 'artificial' and
identifications should be confirmed by reference to original or subsequent generic descriptions.
Some of the characters used to separate groups of genera, e.g. relative width of scape or length of
funicle segments, may prove to be rather weak, but in such cases the genera concerned have
usually been keyed out via both alternatives. More 'natural' keys would be very difficult to use
and might be no more reliable than the present ones, since the number of genera and species
remaining to be discovered in the Neotropics must be very high, probably at least twenty times
greater than the number known at present. The keys are best regarded as an introductory guide
to the Neotropical Encyrtidae and will be in constant need of revision as more information and
material become available.
The main difficulty in producing a more 'natural' key to encyrtid genera is that, unlike most
other chalcidoid families, the Encyrtidae do not easily break down into distinct subgroups. Most
past authors have divided the family into three subfamilies: Arrhenophaginae, Antheminae and
Encyrtinae, the last-mentioned one containing almost all the known genera. Trjapitzin (1973a;
19736) divides the family into two large subfamilies only: Tetracneminae and Encyrtinae,
separated mainly on the presence or absence of paratergites. This character may not be a
reflection of natural grouping within the Encyrtidae since the genera with paratergites are almost
all parasites of Pseudococcidae and these structures may have been evolved more than once as an
adaptation to this form of parasitism; however, some genera of Encyrtinae, lacking paratergites,
are also parasites of Pseudococcidae. Trijapitzin's study is nevertheless the most detailed to date
and I have therefore attempted in this review to place all the included Neotropical genera
according to his classification. This is often difficult or impossible because many of the tribes and
subtribes are not clearly defined; some are even defined entirely by their biology, so that it is
virtually impossible to assign to them genera of which the biology is unknown.
Included in the notes for each genus is the number of known world species and a list of those
known from the Neotropics. References are given to original descriptions or redescriptions for
most of the species listed, or to revisions or other works where these can be found. Also given are
notes on the distribution and biology of each genus. Additional information on the hosts of
related genera is given by Tachikawa (1978).
A key to the genera of Encyrtidae supposedly found in the Caribbean, around Cuba, is
provided by Alayo & Hernandez (1978: 21-27), but this is probably based mainly on
descriptions. Some of the genera included (e.g. Calocerinus Howard, Habrolepopteryx Ashmead)
are known from Florida but not from the Neotropics, although it is quite possible that these
genera may eventually be found on the islands around Cuba.
Keys to the genera of Encyrtidae found in other zoogeographical regions have been published
by the following authors: Trjapitzin (19716) for the Palaearctic; Trjapitzin & Gordh (19780;
19786) for the Nearctic; and Prinsloo & Annecke (1979) for the Ethiopian Region.
The keys in this review are based on the examination of material in the BMNH, London and
on loan from the MLP, Argentina; USNM, Washington; UC, Riverside; CNC, Ottawa; CIBC,
Trinidad and PPRI, Pretoria. For an explanation of these abbreviations see p. 111.
Notes on terms and measurements
Head (Figs 2, 5)
Antennal funicle. This does not include the anellus (or false ring-joint of Timberlake, 1922: 168,
172), which may be present or absent but is almost always hidden by the pedicel and
NEOTROPICAL ENCYRTIDAE
109
mesoscutum
ocelli
ubmarginal
vein
Figs 1-5 1, Homalotylus flaminius (Dalman) $, thorax in dorsal view; 2, generalized encyrtid head in
frontal aspect; 3, generalized encyrtid left forewing in dorsal view; 4, generalized encyrtid $ gaster,
viewed from left side; 5, generalized encyrtid $ antenna, left, outer aspect.
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.
110 JOHN S. NO YES
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 frontovertex is narrowest, as stated in text.
Head width. The maximum width of the head.
Malar space. The minimum distance between the eye and the mouth margin. The measurement is
taken as for the eye.
Malar sulcus. The sulcus joining the lower margin of the eye and mouth margin (see Fig. 2),
sometimes absent but usually indicated by a slight change of sculpture.
Mandibles. The dentition can vary as follows: without teeth, with one long curved tooth
(Fig. 54), one tooth and a truncation (Fig. 91), two teeth (Figs 40, 55, 56), two teeth and a
truncation (Figs 47, 72, 73, 100, 103, 104, 105, although those in Figs 100 and 104 may also
be considered tridentate), two teeth and a rudimentary third tooth (Fig. 102), three teeth
(Fig. 97) or four teeth (Fig. 46).
OOL. The minimum distance between the eye margin and the adjacent posterior ocellus.
POL. The minimum distance between the posterior ocelli.
Thorax (Figs 1, 3)
Forewing.
Marginal vein : measured from where the submarginal vein reaches the anterior margin of the
wing (as shown in Fig. 3) or from where the anterior edge of the venation at the junction of
the submarginal vein and marginal vein is abruptly angled and not from the subapical
hyaline break of the submarginal vein.
Postmarginal vein : measured as shown in Fig. 3, its apex usually indicated by a single,
relatively long seta.
Speculum (or hairless line) : sometimes interrupted in the posterior half or closed near the
posterior margin of the wing by from one to several lines of setae on the dorsal surface of the
wing.
Stigmal vein (or radial vein of some authors) : measured as shown in Fig. 3.
Notaular lines (or parapsidal lines of some authors). 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. 4)
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 to the apex.
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 measured in
dry-mounted material.
Ovipositor. The relative length of the exserted part measured in dry-mounted material. If
material has been in alcohol the gaster may be distorted and the ovipositor appear to be
relatively more exserted; in this case it would be better to use relative lengths of the exserted
parts of the gonostyli.
Ovipositor sheath. The gonostylus as seen in dry-mounted material.
NEOTROPICAL ENCYRTIDAE 1 \ \
Abbreviations
AMNH, New York American Museum of Natural History, New York, U.S.A.
BMNH, London British Museum (Natural History), London, U.K.
CIBC, Trinidad Commonwealth Institute of Biological Control, Trinidad, West Indies.
CNC, Ottawa Biosystematics Research Institute, Ottawa, Canada.
MLP, Argentina Facultad de Ciencas Naturales y Museo, La Plata, Argentina.
PPRI, Pretoria Plant Protection Research Institute, Pretoria, South Africa.
UC, Riverside University of California, Riverside, California, U.S.A.
USNM, Washington National Museum of Natural History, Washington, D.C., U.S.A.
ZI, Leningrad Zoological Institute, Academy of Sciences, Leningrad, U.S.S.R.
Key to genera (females)
The following genera are not included because the females are unknown, the unique type is
missing or their presence in the Neotropics is doubtful: Atelaphycus, Baeoencyrtus,
Echthroplexiella, Parencyrtus, Prionomitus.
1 Antenna with funicle composed of two to four anelliform segments, clava long and
unsegmented (Fig. 6). Tarsi four-segmented . . . ARRHENOPHAGUS (p. 178)
Antenna not as above. Tarsi five-segmented 2
2 (1) Funicle four-segmented ............ 3
Funicle fiv- or six-segmented 7
3 (2) Body dark. Propodeum relatively short centrally, at most one-sixth as long as scutellum 4
Body orange, or, if dark, then propodeum smooth and relatively long centrally, at least
one-third as long as scutellum.
Funicle five-segmented,, but first funicle segment anelliform and almost hidden,
giving funicle four-segmented appearance (Fig. 7) 7
4 (3) All funicle segments longer than broad, usually at least twice as long as broad .
HOMALOPODA (p. 203)
Funicle segments at most quadrate, usually transverse 5
5 (4) Anterior margin of frontovertex above antennal scrobes sharply margined.
Frontovertex and mesoscutum very smooth and shiny. Forewing at least slightly
infumate towards base but without distinct hyaline spots. Marginal vein more or less
quadrate (Fig. 9).
Stigmal vein sessile BRACHYPLATYCERUS (p. 183)
Anterior margin of frontovertex above antennal scrobes rounded. Frontovertex with
sculpture distinct and deeper than that on mesoscutum. Forewing hyaline, or, if
darkened then with several hyaline spots. Marginal vein at least twice as long as
broad ............... 6
6 (5) Forewing infuscate with several hyaline spots . . PSEUDHOMALOPODA (p. 225)
Forewing hyaline ... PLAG1OMERUS (p. 222)
7 (2, 3) Funicle five-segmented 8
Funicle six-segmented 28
8 (7) Forewing shortened, hardly reaching base of gaster 9
Forewing of normal length, at least reaching apex of gaster 10
9 (8) Clava three-segmented. Notaular lines present on mesoscutum. Scutellum of normal
shape with rounded apex which extends well beyond posterior margins of axillae
(Fig. 8). Cerci situated in basal half of gaster . . . SOLENAPHYCUS (p. 230)
Clava two-segmented. Notaular lines absent. Scutellum almost rectangular with
posterior margin hardly extending past posterior margins of axillae (Fig. 11).
Cerci situated in apical half of gaster. . . . STEMMA TOSTERES (p. 230)
112 JOHN S. NOYES
10 (8) Funicle segments distinctly flattened (the first longer than broad), broadening apically
so that sixth is transverse and about one and a half times as broad as and three-
quarters as long as first and about one and a half times as broad as scape (Fig. 10)
ANARHOPUS(p. 173)
Funicle segments cylindrical, first segment often anelliform 11
11 (10) First funicle segment anelliform, at most one-seventh as long as and about half as wide
as second which is subquadrate or longer than broad (Fig. 7) ... . . . 12
First funicle segment not anelliform, or, if anelliform then second is at most only slightly
longer and broader and distinctly transverse . . . . . . 13
12 (11) Exserted part of ovipositor equal to about one-third length of gaster. Propodeum
relatively short centrally, at most one-eighth as long as scutellum ....
APHYCOMASTIX (p. 175)
Ovipositor not or hardly protruding. Propodeum relatively long and shiny centrally, at
least one-third as long as scutellum .... MEROMYZOB1A (p. 212)
13 (11) Exserted part of ovipositor equal to at least one-sixth length of gaster, or, if ovipositor
not as far exserted then body completely yellow except for one or two dark brown
bands on gaster . . 14
Ovipositor not or hardly protruding, gaster never yellow with dark bands . . .20
14 (13) Clava one- or two-segmented TIMBERLAKIA (p. 233)
Clava three-segmented 15
15 (14) First and second funicle segments anelliform, contrasting greatly with third to fifth
segments which are subquadrate or longer than broad (Fig. 14)
AMEROMYZOBIA (p. 172)
All funicle segments transverse (occasionally apical segments subquadrate) and
gradually lengthening towards apex of antenna, never with first and second segments
contrasting as greatly with third as above ,. 16
16 (15) Head and thorax generally shining dark brown, mesopleuron dark brown . . . 17
Head and thorax yellow or orange, or, if darker then not shiny and with venter and sides
of thorax, including mesopleuron, white or yellow or occasionally pale brown . 19
17 (16) Interantennal prominence clothed in dense white setae. Sculpture of scutellum deep and
contrasting with the much shallower sculpture of mesoscutum. Mid tibia yellow
with two brown bands ARCHINUS (p. 176)
Interantennal prominence clothed in sparse dark setae. Sculpture of scutellum more or
less the same as mesoscutum. Mid tibia without brown bands 18
18 (17) First to fourth funicle segments transverse, fifth quadrate. Legs completely yellow
BOTHRIOCRAERA (p. 182)
All funicle segments longer than broad. Legs at least partially darkened . ;!i :v
ARHOPOIDIELLA (p. 177)
19 (16) Antenna yellow or orange, unicolorous ACEROPHAGUS (p. 169)
Clava with at least the two apical segments white contrasting sharply with the dark
brown segments of the funicle ..... PSEUDAPHYCUS (p. 225)
20 (13) Scutellum long with apical tongue-like extension which overhangs propodeum and curls
downwards so that its apex touches posterior margin of propodeum (as in Fig. 12).
Frontovertex with a few, very long, conspicuous setae. Clava very large and as wide
as long, much wider than funicle. Submarginal vein of forewing with very well-
marked triangular expansion near its apex (as in Fig. 13)
BLANCHARDISCUS (p. 182)
Scutellum of normal form, not overhanging propodeum. Frontovertex with short setae.
Clava longer than broad. Submarginal vein occasionally with triangular expansion. 21
21 (20) Malar space very long, about one and a half times length of eye (Fig. 21) .
ARRHENOPHAGOIDEA (p. 178)
Malar space much less than length of eye . . , ... ; . . .* . 22
NEOTROPICAL ENCYRTIDAE 1 1 3
22 (21) Forewing hyaline, or, if slightly infumate then mesoscutum has more or less complete
notaular lines .............. 23
Forewing more or less uniformly infumate. Mesoscutum entire.
Funicle six-segmented, but first funicle segment very short and sometimes hidden
by pedicel in dry mounted specimens, giving funicle five-segmented appearance . 28
23 (22) Clava four-segmented (Fig. 16) TETARTICLAVA (p. 232)
Clava at most three-segmented 24
24 (23) Hypopygium not reaching apex of gaster, at most reaching about two-thirds along
gaster. Marginal vein broader than long, or, if slightly longer than broad then colour
of head and thorax orange 25
Hypopygium reaching apex of gaster. Marginal vein longer than broad. Head and
thorax dark brown or black ........... 26
25 (24) Mesoscutum with complete notaular lines (Fig. 8). Marginal vein punctiform or slightly
longer than broad. Clava apically rounded. Head and thorax orange
SOLENAPHYCUS (p. 230)
Mesoscutum entire. Marginal vein distinctly broader than long (as in Fig. 126). Clava
obliquely truncate (Fig. 15). Head and thorax dark brown
PENTALITOMASTIX (p. 222)
26 (24) Notaular lines complete (difficult to see in dry mounted material). Scutellum matt with
much deeper sculpture than mesoscutum .... COCCIDAPHYCUS (p. 187)
Notaular lines absent or indicated anteriorly only. Scutellum shiny with sculpture not
noticeably deeper than mesoscutum 27
27 (26) Funicle segments transverse. Forewing with speculum narrow, about same width as
length of marginal vein and not interrupted (Fig. 20), but closed on ventral surface of
wing by two lines of setae near dorsum. Mandibles tridentate
ACEROPHAGOIDES (p. 169)
Funicle segments subquadrate or longer than broad. Forewing with speculum broad,
much broader than length of marginal vein (Fig. 24) and interrupted by two lines of
setae on dorsal surface of wing near dorsum. Mandibles bidentate ....
TETRACNEMOIDEA (p. 233)
28 (7, 22) Wings shortened, not reaching apex of gaster 29
Wings of normal length, reaching or exceeding apex of gaster ..... 40
29 (28) Scutellum concave, or with a pit or indentation basally 30
Scutellum flat or convex, without pit or indentation basally 33
30 (29) Posterior margin of scutellum with line of longer, slightly flattened, upright setae. Scape
less than twice as long as broad COELASPIDIA (p. 188)
Posterior margin of scutellum with at most two upright setae. Scape at least two and a
half times as long as broad 31
31 (30) Outer margins of antennal scrobes rounded. Occipital margin rounded. Scape widest at
extreme apex and here with internal and external flanges that completely enclose
pedicel. Eyes small, only slightly longer than minimum width of fronto vertex .
PHEIDOLOXENUS (p. 222)
Antennal scrobes sharply margined above and at sides. Occipital margin acute. Scape
widest in middle or towards base, pedicel not enclosed. Eyes larger, more than twice
as long as minimum width of frontovertex 32
32 (31) Scape foliaceously flattened, widest in middle and about three times as long as broad.
Mesoscutum uniformly sculptured and convex. Scutellum more or less triangular
in shape with apices rounded AEPTENCYRTUS (p. 171)
Scape not foliaceously flattened, widest in basal third and about five times as long as
greatest width. Mesoscutum distinctly sculptured and convex centrally, completely
smooth, shiny and concave laterally. Scutellum as seen from above with sides
concave in outline and almost T-shaped . . . PELMATENCYRTUS (p. 221)
33 (29) Apex of scutellum with distinct tuft of setae 34
Apex of scutellum without distinct tuft of setae 35
114 JOHN S. NO YES
34 (33) Dorsal margin of scape almost straight or uniformly curved. Clava three-segmented.
Ovipositor longer than mid tibia CHEILONEURUS (p. 184)
Dorsal margin of scape abruptly bent at middle (Fig. 17). Clava two-segmented.
Ovipositor shorter than mid tibia J " -1 ;^;:.;" . . . ACROASPIDIA (p. 170)
35 (33) First funicle segment longer than pedicel, scape about eight times as long as broad.
Rudiments of forewing entirely infuscate . . •; . . ECTROMATOPSIS (p. 194)
First funicle segment shorter than pedicel, scape at most six times as long as broad.
Rudiments of forewing hyaline, suffused with yellow or infuscate at base and apex 36
36 (35) Frontovertex broad, at narrowest point at least half as wide as head . . •: . . 37
Frontovertex narrower, at narrowest point at most three- fifths as wide as head . . 38
37 (36) Mesoscutum very transverse, about four times as broad as long. Scutellum with apex
more or less square. Axillae separated by almost their own width (Fig. 18). Cerci very
near base of gaster. Antenna with clava white contrasting with dark funicle
segments. Body fairly robust NEODUSMETIA (p. 216)
Mesoscutum not as transverse as above, only slightly broader than long. Scutellum with
apex well rounded. Axillae more or less touching (as in Fig. 19). Cerci nearly halfway
along gaster. Antenna with clava and funicle usually unicolorous or with clava darker
than funicle, never with clava paler than funicle. Body dorso-ventrally flattened
RHOPUS (p. 226)
38 (36) Rudiment of forewing infuscate at base and apex. Mandibles bidentate
ECTROMA (p. 194)
Rudiment of forewing hyaline or suffused pale yellow. Mandibles not bidentate. . 39
39 (38) Head and thorax completely orange. Forewing rudiment transversely truncate.
Mandibles with three acute teeth MUCRENCYRTUS (p. 213)
At least thorax dark and metallic. Forewing rudiment more or less rounded, not
transversely truncate. Mandibles with one or two teeth and a truncation
OOENCYRTUS (p 217)
40 (28) Scape not more than three times as long as its greatest width 41
Scape more than three times as long as its greatest width 90
41 (40) Forewing darkened (excluding those species with very small darkened area below
marginal vein which hardly extends past apex of stigmal vein, or those where
infumation is very weak and almost imperceptible) 42
Forewing hyaline or with almost imperceptible infumation (also including those species
with a very small darkened area below marginal vein which hardly extends past apex
of stigmal) 66
42 (41 ) Funicle segments and clava flattened, at least segments four to six of funicle more or less
as wide as clava and almost parallel sided (Figs 22, 23). Funicle segments often
articulated near dorsal edge and with long curved ventral process .... 43
Funicle segments and clava not flattened, more or less cylindrical or oval in cross-
section, or, if appearing slightly flattened then flagellum distinctly club-shaped, the
clava much wider than majority of funicle segments which gradually widen towards
apex of antenna. Funicle segments articulated in middle and never with ventral
process 50
43 (42) Stigmal vein short, at most one-third as long as marginal (Fig. 25). Exserted part of
ovipositor at least half as long as gaster TETRACNEMUS (p. 233)
Stigmal vein longer, at least half as long as marginal. Exserted part of ovipositor at most
one-third as long as gaster in all known South American species . . "^ . . 44
44 (43) Forewing with two fuscous fasciae, one arising from marginal vein (incomplete) and
other about half way along submarginal, sometimes expanded into a triangular
blotch, areas either side of the outermost fascia paler than wing apex. Head and
thorax orange or reddish with rather coarse punctate sculpture which gives a silky
appearance . . . ANAGYRUS (p. 173)
Infuscate areas of forewing not represented by fasciae, usually more extensive. Head
and thorax dark, or, if orange then mesoscutum with shallow reticulate sculpture
which is quite shiny . . . . . . . . , . . . . 45
NEOTROPICAL ENCYRTIDAE 1 \ 5
45 (44) Marginal vein very short, almost punctiform, stigmal and postmarginal veins both at
least four times as long as marginal, stigmal curved towards anterior margin of wing
(Figs 26, 27) 46
Marginal vein relatively longer, at least half as long as stigmal 47
46 (45) Forewing : postmarginal vein distinctly longer than stigmal, costal cell with two or three
lines of setae ventrally, upper surface of wing below apex of submarginal vein with at
least four lines of normal setae nearly reaching the dorsum (often basal cell
completely pilose), basal cell infumate (Fig. 27). Mesoscutum and scutellum entirely
of similar sculpture, scutellum at most slightly convex and without distinct tuft or line
of setae at apex ZAPLATYCERUS (p. 235)
Forewing: postmarginal vein at most as long as stigmal, costal cell with no more than
one complete line of setae ventrally, upper surface of forewing below apex of
submarginal vein with at most one line of setae nearly reaching dorsum (usually not
reaching half way across wing), some of these flattened and scale-like, basal cell
almost completely hyaline (Fig. 26). At least part of mesoscutum or scutellum smooth
and shiny, contrasting with sculptured areas, scutellum strongly convex and with tuft
or line of setae at apex . . . CHRYSOPLA TYCERUS (p. 185)
47 (45)* Body dorso-ventrally flattened. Forewing with one or two longitudinal fuscous streaks.
Frontovertex usually yellow with median dark stripe . .COMPERIELLA (p. 189)
Body robust. Forewing more or less uniformly infuscate from speculum to near apex.
Frontovertex unicolorous, occasionally darker between ocelli ..... 48
48 (47) Infuscation of forewing enclosed subapically by darker band, or at least with setae
appearing to be more dense in this band (Fig. 30) . . . .ANICETUS (p. 174)
Infuscation of forewing not enclosed subapically by darker band .... 49
49 (48) Funicle segments broadening towards apex of antenna so that clava is distinctly broader
than first three (Fig. 23). Frontofacial ridge prominent and continuous across face .
HOMOSEMION (p. 203)
Funicle segments slightly but distinctly narrowing towards apex of antenna so that clava
is narrower than funicle (Fig. 22). Interantennal prominence confluent with
frontovertex so that frontofacial ridge is interrupted in middle ....
AMMONOENCYRTUS (p. 172)
50 (42) Clava entire 51
Clava three-segmented 52
51 (50) Pedicel apically with several conspicuous very long setae on dorsal surface (Fig. 31).
Marginal and postmarginal veins more or less equal in length and stigmal about one
and a half times as long as either (Fig. 29). Body colour orange to dark
brown. Mandibles bidentate HAMBLETONIA (p. 200)
Pedicel apically without conspicuous setae on dorsal surface. Marginal vein short,
almost punctiform and several times shorter than the long postmarginal or stigmal,
the latter being strongly curved towards anterior margin of wing (Fig. 28). Body
colour shining green or blue. Mandibles tridentate . . . ZARHOPALUS (p. 235)
52 (50) First funicle segment about twice as long as pedicel. Forewing with two fuscous bands,
one arising at marginal vein and other about half way along submarginal (latter may
extend almost to base of wing), apex and base of wing infumate but paler.
Mandibles bidentate (as in Fig. 40) ANAGYRUS (p. 173)
First funicle segment at most one and half times as long as, usually shorter than, pedicel.
Forewing not infuscate as above. Mandibles rarely bidentate, usually tridentate or
quadridentate 53
53 (52) Apex of scutellum and occipital margin between eyes each with two lamelliform setae
(Fig. 32) . HABROLEPIS (p. 199)
Apex of scutellum and occipital margin without lamelliform setae .... 54
* Note added while this paper was in press: Cerapterocerus Westwood (see footnote, p. 107) runs here, but differs from
the other included genera in the body not being dorso-ventrally flattened, the forewing having several sharply delimited
wedge-shaped hyaline areas and the submarginal vein having a distinct triangular expansion.
116 JOHN S. NO YES
54 (53) Apex of scutellum with long tuft of setae, or, if tuft not distinct then exserted part of
ovipositor is about half as long as gaster. Forewing more or less uniformly infuscate
from bend of submarginal vein to at least near apex (occasionally a broad hyaline
band at apex) •< 55
Apex of scutellum without long tuft of setae. Ovipositor hidden or if slightly protruding
than exserted part is less than one-fifth as long as gaster. Forewing with distinct
hyaline spots or band of paler setae between marginal vein and apex ... 59
55 (54) Mesoscutum usually with median tuft of setae, or, if absent then submarginal vein has
small but distinct triangular expansion in apical third below which is a tuft of bristles
and infuscate area delimited subapically by darker band. Marginal vein not or hardly
longer than stigmal 56
Mesoscutum without central tuft of setae. Submarginal vein without small triangular
expansion. Marginal vein at least slightly longer than stigmal, usually more than
twice as long . - * . 57
56 (55) Mesoscutum with median tuft of setae. Infuscate area of forewing not delimited by
darker band subapically , * . DIVERSINERVUS (p. 193)
Mesoscutum without median tuft of setae. Infuscate area of forewing delimited
subapically by darker band (as in Fig. 30) . . . . . . genus A (p. 169)
57 (55) Marginal vein at most about one and a half times as long as stigmal, postmarginal at
least half as long as marginal (Fig. 38), infuscation of forewing very slight.
Hypopygium extending at most half way along gaster. Exserted part of ovipositor
equal to at least half length of gaster ICEROMYIA (p. 204)
Marginal vein at least twice as long as stigmal and four times postmarginal. Infuscation
of forewing strong (Figs 35, 36). If ovipositor is as long as above then hypopygium
extends to apex of gaster or infuscation of forewing is very strong .... 58
58 (57) Epipygium shortened, giving apex of gaster a squared-off appearance (gaster sometimes
collapses upwards and inwards laterally giving it a tapered appearance). Hypopygium
reaching apex of gaster. Apical hyaline area of forewing, at widest, about equal to
half maximum width of wing and with well-delimited strongly curved inner margin,
subparallel to wing apex (Fig. 35). Exserted part of ovipositor always at least half as
long as gaster PROCHILONEURUS (p. 223)
Epipygium long and triangular giving gaster a gradually tapered appearance.
Hypopygium at most reaching about two-thirds along gaster. Forewing without
apical hyaline area, or, if area present then its inner margin not well delimited but
usually wedge-shaped, not subparallel to apical margin (Fig. 36). Ovipositor usually
more or less hidden, occasionally exserted part long, but always shorter than half
length gaster . . . . . CHEILONEURUS (p. 184)
59 (54) Forewing with several hyaline spots on margin. Head triangular in side view with planes
of face and frontovertex forming a strongly acute angle (Fig. 33) ANABROLEPIS (p. 172)
Forewing without hyaline spots, generally uniformly infuscate or gradually becoming
paler towards apex or with one or more transverse, often interrupted, hyaline fasciae.
Head in side view more or less gradually curved (except in Hemaenasius which has
base of forewing infumate and gradually fading towards wing apex) ... 60
60 (59) First funicle segment distinctly transverse, or, if subquadrate then at most half as long as
pedicel 61
First funicle segment longer than broad, or, if subquadrate then at least two-thirds as
long as pedicel 65
61 (60) Forewing more or less uniformly infuscate or gradually becoming paler towards apex,
without hyaline bands or band of paler setae between marginal vein and apex.
Pronotum more or less same colour as mesoscutum .... 62
Forewing with at least one hyaline band or a band of paler setae distal of marginal vein.
Pronotum often paler than mesoscutum : .... 64
62 (61) Occipital margin sharp. Forewing with basal cell totally infumate. Hypopygium
reaching apex of gaster. Eyes reaching occipital margin. Head and thorax shining
black, blue or green . . /: . .-'. 63
NEOTROPICAL ENCYRTIDAE 1 17
Occipital margin not sharp. Forewing with basal cell hyaline. Hypopygium not reaching
apex of gaster. Eyes separated from occipital margin by distance about the same as
diameter of posterior ocellus (Fig. 34). Head and thorax brown ....
HUNTERELLUS (p. 204)
63 (62) Head more or less lenticular, usually with thimble-punctured sculpture. Flagellum club-
shaped and clava as least as long as funicle (Fig. 62). Costal cell with at least two or
three lines of setae dorsally
AENASIUS(p. 170)
Head not lenticular, often abruptly bent at level with top of antennal scrobes,
frontovertex smooth and shiny. Flagellum not club-shaped and clava slightly shorter
than funicle (Fig. 51). Costal cell dorsally with at most a few setae at apex
HEMAENASWS (p. 202)
64 (61) Femora as well as tibia with contrasting light and dark bands or areas. Forewing: basal
cell with setae on dorsal surface extremely dense, as dense as in centre of wing, distal
side of infuscate area with a distinct band of paler setae
[Dicarnosis] blanchardi De Santis (p. 193)
At most only mid and hind tibiae with pronounced dark bands. Forewing: basal cell
usually with setae on dorsal surface markedly less dense than, rarely as dense as, in
centre of wing, distal side of infuscate area without band of paler setae .
METAPHYCUS (p. 212)
65 (60) Dorsum of thorax, frontovertex and mesopleura with dense punctiform sculpture.
Upper surface of basal cell with setae as dense as in centre of wing. Eyes distinctly
hairy. Mandibles with two teeth and a truncation
COMPERIA (p. 189)
Dorsum of thorax almost smooth, with shallower less dense sculpture. Upper surface of
basal cell with setae much less dense than centre of wing and usually with bare area.
Eyes not distinctly hairy. Mandibles usually with three acute teeth ....
MICROTERYS (p. 213)
66 (41) All funicle segments longer than broad 67
Not all funicle segments longer than broad 71
67 (66) First funicle segment at most one and a half times as long as broad. Mandibles
tridentate or with one tooth and a broad truncation or without teeth ... 68
First funicle segment at least twice as long as broad. Mandibles bidentate (Fig. 40) . 69
68 (67) Scutellum rather convex and with similar sculpture to mesoscutum. Pedicel subequal in
length to first funicle segment, clava at most as long as the combined lengths of
funicle segments four to six (Fig. 37) GAHANIELLA (p. 197)
Scutellum flat with deeper sculpture than mesoscutum. Pedicel longer than first funicle
segment, clava nearly as long or longer than the funicle (as in Fig. 39) .
COCCIDENCYRTUS (p. 188)
69 (67) Sculpture of frontovertex shallow and finer so that it is distinctly shiny, never orange or
yellow, usually black APOANAGYRUS (p. 176)
Sculpture on frontovertex coarsely punctate giving it a matt or silky appearance, never
shiny and often yellow or orange .......... 70
70 (69) Postmarginal vein shorter than stigmal (Figs 70, 71). Face usually with conspicuous,
moderately dense white setae. Sculpture on frontovertex irregular and silky in
appearance ANAGYRUS (p. 173)
Postmarginal vein at least about twice as long as stigmal. Pale setae present on head, but
never conspicuous. Sculpture on frontovertex more regular and polygonal, not silky
in appearance GYRANUSOIDEA (p. 199)
71 (66) Apex of scutellum with a tuft of setae, or, if this is inconspicuous then normally at least
posterior half of mesoscutum with silvery white setae which contrast with the darker
colour of the mesoscutum, scutellum with dark setae. Usually at least anterior one-
third of mesoscutum with fine striate sculpture. Frontovertex shiny, thorax and head
dark ZAOMMA (p. 235)
118 JOHN S. NO YES
Apex of scutellum without tuft of setae (in majority of species only two upright setae).
Mesoscutum without silvery setae, or, if these are present and contrast with dark
colour of mesoscutum then scutellum also has silvery setae, frontovertex is matt and
thorax and head usually have paler areas . . .72
72 (71)* Body distinctly dorso-ventrally flattened. Cerci situated about half way along gaster.
Ovipositor not protruding. Mandibles long and narrow with teeth of equal length .
RHOPUS (p. 226)
Body robust, or if flattened, then cerci are situated near base of gaster (mandibles may
be bidentate, but here upper tooth is very long and curved and lower tooth short) or
mandibles are clearly tridentate and the exserted part of the ovipositor is equal to at
least one-quarter length of gaster . . . . . . ... . 73
73 (72) Clava two-segmented . . . 74
Clava three-segmented ',........ . , 75
74 (73) Frontovertex with distinct sculpture. Antennal scrobes moderately impressed and horse-
shoe shaped. Exserted part of ovipositor equal to about one-quarter length of gaster
and more or less straight, not curved downwards . . TANYENCYRTUS (p. 231)
Frontovertex highly polished. Antennal scrobes absent or represented by an extremely
shallow groove much shorter than length of torulus (as in Fig. 41). Exserted part of
ovipositor equal to at least about half length of gaster and distinctly downcurved .
SHENAHETIA (p. 227)
75 (73) Body foliaceously dorso-ventrally flattened (Fig. 42). Exserted part of ovipositor at least
one-third length of gaster and with sheaths bilaterally flattened. . MARIOLA (p. 210)
Body robust. Ovipositor not exserted as much as above, or if so, then sheaths are
cylindrical 76
76 (75) Head at least with some orange or yellow areas, or, if head completely darkened then
matt in appearance and pronotum paler than mesoscutum 77
Head dark and shiny, or, if head not distinctly shiny then pronotum the same colour as
mesoscutum 79
77 (76) Clava apically rounded METAPHYCUS (p. 212)
Clava obliquely truncate (Fig. 43, cf. Fig. 44) ........ 78
78 (77) Exserted part of ovipositor as long as at least half length of gaster. At least mesoscutum
dark and metallic. Marginal vein at least twice as long as broad (as in Fig. 38)
ICEROMYIA (p. 204)
Ovipositor not or hardly visible. Thorax completely orange; marginal vein punctiform
(Fig. 150) .... BENNETTISCA (p. 180)
79 (76) Pedicel and all funicle segments subequal in length and breadth and subquadrate or
longer than broad (as in Fig. 37), frontovertex at narrowest point at least three-
eighths head width.
Mandibles without teeth or with one tooth and a broad truncation . . f! t' o
GAHANIELLA (p. 197)
Pedicel distinctly longer than first funicle segment, or, if not then funicle distinctly
broadens apically or the frontovertex at its narrowest point is less than one-third head
width and the funicle segments are distinctly not quadrate 80
80 (79) Clava shorter than funicle, at most as long as the combined lengths of funicle segments
three to six 81
Clava almost as long or longer than funicle, at least as long as the combined lengths of
funicle segments two to six . . . . , . . , . .... 86
* Note added while this paper was in press. Asitus Erdos, Boucekiella Hoffer and Platencyrtus Ferriere (see footnote on
p. 107) all run here but may be distinguished by having an extremely flattened leaf-like body. Asitus has bidentate
mandibles, a longitudinally divided pronotum and the forewing with a very long marginal fringe, at least one-fifth
maximum width of wing; Boucekiella has tridentate mandibles, as does Platencyrtus, and has the submarginal vein with a
subapical triangular expansion (absent in Platencyrtus) and the ovipositor exserted for a length equal to one-third of
gaster (the ovipositor is not exserted in Platencyrtus).
NEOTROPICAL ENCYRTIDAE 1 19
81 (80) Head with large, shiny bottomed thimble-punctures. Scutellum convex with rough
sculpture. Hypopygium not reaching apex of gaster . . HELMECEPHALA (p. 200)
Head without large thimble-punctures, or if present (Zeteticontus), then scutellum is
quite flat, smooth and shiny, and the hypopygium reaches apex of gaster . . 82
82 (81) Eyes over-reaching occipital margin and with very short indistinct hairs. Frontovertex
with inconspicuous hairs. Ocelli usually forming a distinctly acute angle. Scutellum
convex, usually with deep sculpture, but occasionally smooth.
Mandibles with one or two teeth and truncation . . . OOENCYRTUS (p. 217)
Eyes not over-reaching occipital margin, or, if so then with conspicuous hairs.
Frontovertex with long, conspicuous hairs. Ocelli usually forming a right or obtuse
angle, or, occasionally a slightly acute angle. Scutellum more or less flat, usually
smooth and shiny, without distinct sculpture 83
83 (82) Eyes separated from occipital margin, which is more or less rounded, by a distance
about same as diameter of posterior ocellus (Fig. 34) . HVNTERELLUS (p. 204)
Eyes separated from occipital margin, which is sharp, by much less than half diameter of
posterior ocellus ............. 84
84 (83) Mandibles with three acute teeth. Hypopygium reaching apex of gaster
ZETETICONTUS (p. 236)
Either the hypopygium does not reach apex of gaster, or mandibles are bidentate with
upper tooth long and curved and lower tooth short (Figs 55, 56) . . . . 85
85 (84) Mandibles tridentate. Propodeum at most one-eighth as long as scutellum and without
distinct sculpture centrally EXORISTOBIA (p. 196)
Mandibles bidentate (Figs 55, 56). Propodeum at least one-sixth as long as scutellum
and with distinct sculpture centrally (as in Fig. 121) . . RHYTIDOTHORAX (p. 226)
86 (80) Mesoscutum with distinct longitudinally striate sculpture . . . DESOBIUS (p. 192)
Mesoscutum not with striate sculpture 87
87 (86) Scutellum with coarser sculpture than mesoscutum, giving it a matt (or less shiny)
appearance in contrast with the shiny appearance of the mesoscutum ... 88
Scutellum with similar sculpture to mesoscutum, both shiny . . . . 89
88 (87) Mandibles quadridentate (Fig. 46). Mid tibia completely yellow without dark bands .
ADELENCYRTVS (p. 170)
Mandibles with one or two teeth and a truncation, mid tibia often brown, or, yellow
with dark bands .... . COCCIDENCYRTUS (p. 188)
89 (87) Funicle and clava unicolorous, brown . . . .COELOPENCYRTUS (p. 189)
Funicle yellow, contrasting with the brown clava
[ICoccidencyrtus maculicornis (Blanchard)] (p. 188)
90 (40) Hind tibia distinctly flattened (Fig. 45). Forewing with central oval fuscous spot or
fascia. Mesoscutum with notaular lines (one specimen from Panama which may
belong here has mesoscutum entire and clava obliquely truncate) . MOORELLA (p. 213)
Hind tibia not flattened, or, if flattened then mesoscutum entire or forewing hyaline . 91
9 1 (90) Eyes nearly reaching base of mandible, malar space at most about one-fifth length of eye
(usually not more than one-tenth), apex of clypeus slightly produced outwards
(Fig. 48).
Notaular lines completely absent. Head and thorax completely metallic green or
blue. Clava obliquely truncate .... PARABLASTOTHRIX (p. 218)
Malar space much longer than above and clypeus normal ... 92
92 (91) Forewing darkened (excluding those species with small fuscous spot below marginal
vein which does not or hardly extends past apex of stigmal vein, or, with forewing
suffused pale yellow) .......
Forewing hyaline (including those species with small fuscous spot below marginal vein
which does not or hardly extends past apex of stigmal vein, or with forewing suffused
pale yellow) 145
120 JOHN S. NO YES
93 (92) Apex of scutellum with tuft of long setae at apex ....... 94
Apex of scutellum without tuft of setae (usually two erect setae in most species) . 96
94 (93) Marginal vein short, postmarginal and stigmal veins long, at least about twice as long as
marginal (Fig. 49). Mandibles without teeth .... ENCYRTUS (p. 195)
Marginal vein at least twice as long as stigmal (Figs 35, 36). Mandibles with at least two
acute teeth -•- 95
95 (94) Epipygium shortened giving gaster a truncate appearance (gaster sometimes collapses
upwards and inwards laterally giving it a tapered appearance). Hypopygium reaching
apex of gaster. Apical hyaline area of forewing, at widest, equal to about half
maximum width of wing and with a well-delimited, strongly curved inner margin
subparallel to wing apex (Fig. 35). Exserted part of ovipositor always equal to at least
half length of gaster . PROCHILONEURUS (p. 223)
Epipygium long and triangular giving it a gradually tapered appearance. Hypopygium
at most reaching two-thirds along gaster. Forewing without apical hyaline area, or, if
area present then its inner margin not well delimited but usually wedge-shaped, not
subparallel to apical margin (Fig. 36). Ovipositor generally more or less hidden,
occasionally exserted part long, but always shorter than half length of gaster..
CHEILONEURUS (p. 184)
96 (93) Exserted part of ovipositor at least one-fifth as long as gaster . . : :'r . . 97
Exserted part of ovipositor at most one-sixth as long as gaster 108
97 (96) Mesoscutum with complete notaular lines (as in Fig. 1) . . . . . . 98
Mesoscutum entire or with notaular lines indicated in anterior half only ... 99
98 (97) Clava apically rounded. Mid and hind tibiae each yellow with two brown rings .
TYNDARICHOIDES (p. 234)
Clava apically obliquely truncate (as in Fig. 50). Mid and hind tibiae without dark rings 99
99 (97, 98) Infuscation of forewing restricted to median oval spot below stigmal and marginal
veins, occasionally moderately large and extending over whole of central third of
wing, but at least basal and apical thirds of wing hyaline. At least one funicle segment
quadrate or transverse. Notaular lines usually anteriorly indicated. Exserted part of
ovipositor equal to at least half length of gaster. Hypopygium reaching apex of gaster 100
Infuscation of forewing usually more extensive (less extensive in some genera) and often
pale, or, if similar to above then all funicle segments longer than broad. Notaular
lines completely absent. Exserted part of ovipositor occasionally as long or longer
than above. Hypopygium sometimes reaching apex of gaster 102
100 (99) Notaular lines more or less complete, at least nearly reaching posterior margin of
mesoscutum (as in Fig. 1). Clava solid and obliquely truncate (as in Fig. 50) .
HOMALOTYLUS (p. 203)
Either notaular lines are indicated at anterior margin of mesoscutum only, or, clava is
segmented and apically rounded and notaular lines reach at most two-thirds across
mesoscutum 101
101 (100) Ovipositor sheaths, in side view, distinctly tapering towards apex and relatively deep at
apex of gaster, at least one and a half times as deep as maximum breadth of scape
AZTECENCYRTUS (p. 180)
Ovipositor sheaths completely cylindrical and not tapering, at apex of gaster about half
as deep as maximum breadth of scape .... CIRRHENCYRTUS (p. 187)
102 (99) Forewing with a large, hyaline, triangular spot on anterior margin confluent with apices
of postmarginal and stigmal veins and joined by a slightly curved hyaline band to a
similar spot on the opposite side of the wing (Fig. 52). Hind tibia yellow with two
dark bands NEOCOCCWENCYRTUS (p. 216)
Forewing without hyaline areas as above. Hind tibia at most with one brown band . 103
103 (102) Occipital margin rounded . . . . . . ^ .. . . . . 104
Occipital margin sharp , - ; , . . 105
NEOTROPICAL ENCYRTIDAE 121
104 (103) Epipygium broadly rounded apically giving gaster truncate apearance. Exserted part of
ovipositor slightly shorter than gaster. Body more or less orange ....
CALLIENCYRTUS (p. 183)
Epipygium elongate and triangular, overhanging base of exserted part of ovipositor.
Exserted part of ovipositor longer than gaster. Body dark metallic green
DELOENCYRTUS (p. 191)
105 (103) Ovipositor sheaths distinctly flattened, together more than twice as deep as broad in
dorsal view HEMENCYRTUS (p. 202)
Ovipositor sheaths cylindrical or oval in cross section 106
106 (105) Stigmal vein longer than marginal. Infuscation of forewing weak, not extending past
stigmal vein towards wing apex. Hypopygium extending about halfway along gaster.
Mesoscutum orange with contrasting dark brown setae . APHYCOMORPHA (p. 175)
Stigmal vein short, less than one-third as long as marginal (as in Fig. 35). Infuscation of
forewing stronger and extending past stigmal vein almost to wing apex. Hypopygium
reaching apex of gaster. Mesoscutum with pale (usually white) setae . . .107
107 (106) All funicle segments longer than broad. Gaster long and tapered, longer than thorax.
TINEOPHOCTONVS (p. 234)
At least one funicle segment quadrate or transverse. Gaster apically truncate and
shorter than thorax . PROCHILONEURUS (p. 223)
108 (96) All funicle segments longer than broad 109
Not all funcle segments longer than broad 121
109 (108) Mesoscutum with more or less complete notaular lines (Fig. 1). Clava obliquely truncate
(Fig. 50) HOMALOTYLUS (p. 203)
Mesoscutum without notaular lines, clava apically rounded 110
110 (109) Forewing with one or two distinct, dark sometimes incomplete fasciae (occasionally
dark spot beyond second fascia and confluent with it). First funicle segment as long or
longer than pedicel Ill
Either forewing without distinct dark fasciae, or, first funicle segment distinctly shorter
than pedicel 115
111 (110) Cerci in apical third of gaster 112
Cerci in basal third of gaster 113
112 (111) Axillae very convex and very prominent, raised about level of scutellum, scutellum
posteriorly raised above level of mesoscutum. Scape reaching well above level of
vertex. First funicle segment about three times as long as pedicel. Eyes separated from
occipital margin by distance greater than diameter of posterior ocellus. Occipital
margin strongly concave in dorsal view. Forewing with two dark fasciae, basal cell
and costal cell almost completely naked HEX ACL ADI A (p. 203)
Axillae almost flat and on a level with scutellum which is not posteriorly raised. Scape
not reaching further than anterior ocellus. First funicle segment less than twice as
long as pedicel. Eyes separated from occipital margin by distance less than diameter
of posterior ocellus. Occipital margin only slightly concave in dorsal view. Forewing
with only one dark fascia, basal and costal cells hairy . PRIONOMASTIX (p. 223)
113(111) Postmarginal vein longer than stigmal LEPTOMASTIDEA (p. 206)
Postmarginal vein shorter than stigmal (as in Figs 70, 71) . . . .114
114 (113) Malar space relatively short, less than one-quarter length of eye. Scape cylindrical,
about eight times as long as broad. Forewing with only one fascia, this situated about
half way along submarginal vein. Body black .... ANATHRIX (p. 174)
Malar space longer, nearly half as long as eye. Scape distinctly broadened and flattened,
just greater than three times as long as broad. Forewings with two dark fasciae, one
two-thirds along submarginal vein and the other at marginal vein. Head and thorax
orange ANAGYRUS (p. 173)
115 (110) Infuscation of forewing restricted to single wedge-shaped streak (Fig. 57) or triangular
mark arising at or near apex (Fig. 53). Thorax green 116
122 JOHN S. NOYES
Infuscation not triangular or wedge-shaped, or, if similar to above then thorax
completely orange .;;•„•. 117
116 (115) Forewing with marginal vein about seven times as long as broad, stigmal and
postmarginal veins both longer than marginal (Fig. 53). Pedicel less than half as long
as first funicle segment. Occipital margin acute. Mandibles bidentate .
ERICYDNUS (p. 195)
Forewing with marginal vein at most five times as long as broad, stigmal and
postmarginal veins shorter than marginal (Fig. 57). Pedicel less than half as long as
first funicle segment. Occipital margin rounded. Mandibles tridentate
PARECHTHRODRYINUS (p. 221)
117 (116) Marginal form punctiform, stigmal vein at least twice as long as marginal, postmarginal
vein short or absent (as in Fig. 90). Infuscation restricted to large, more or less oval
spot extending completely across wing below marginal vein MUCRENCYRTUS (p. 213)
Marginal vein at least twice as long as broad, or nearly as long as stigmal, if stigmal
longer than marginal then postmarginal long and distinct. Infuscation usually more
extensive 118
118 (i!7) Occipital margin more or less rounded. Head triangular in side view (as in Fig. 33).
Forewing with distinct hyaline fascia distal to marginal vein, postmarginal vein very
short or absent (Fig. 52) NEOCOCCIDENCYRTUS (p. 216)
Occipital margin acute. Head rounded in side view. Forewing never with hyaline fascia,
postmarginal vein long 119
119 (118) Eyes not distinctly hairy. Forewing with speculum interrupted in lower half (as in Figs
70, 71). First funicle segment at least twice as long as pedicel. Head and thorax orange
LEPTOMASTIX (p. 206)
Eyes distinctly hairy, speculum not interrupted. First funicle segment not or hardly
longer than pedicel. Head and thorax dark, from green to black . . . .120
120 (119) Ovipositor usually visible with sheaths bilaterally flattened. Mandibles broad and
tridentate, lower teeth subequal in length, upper tooth often short and indistinct.
Scutellum either flat with a distinct translucent apical flange (Fig. 60) or very convex
with flange indistinct or absent. Hypopygium reaching apex of gaster
HEMENCYRTUS (p. 202)
Ovipositor usually hidden, but if visible then sheaths are cylindrical. Mandibles
unidentate (Fig. 54) or bidentate with upper tooth long and curved, distinctly longer
than the short lower tooth (Figs 55, 56). Scutellum always fairly flat and without
apical translucent flange (although occasionally a flange is present but this is not
translucent and is distinctly sculptured). Hypopygium not reaching apex of gaster,
often quite long laterally and nearly reaching apex, but deeply incised centrally
RHYTIDOTHORAX (p. 226)
121 (108) Funicle segments flattened, distinctly broadest in middle of funicle and gradually
tapering towards base and apex (Fig. 66). Forewing with patch of golden setae
situated in centre just below marginal vein and bordered along inner and outer
margins by dark setae, basal cell completely naked. Marginal vein long and subequal
in length to the long postmarginal, stigmal vein distinctly shorter than either .
ANUSIOPTERA (p. 175)
Funicle segments subequal in width or broadening towards clava, never with segments
widest in middle, forewing not as above . 122
122 (121) Forewing with one or two hyaline fasciae, or a pair of opposite triangular hyaline spots
(sometimes indistinct) distal to stigmal vein, apex of forewing infuscate. Mesoscutum
entire 123
Forewing without hyaline fasciae, or triangular spots distal to stigmal vein, or, if fascia of
paler setae present then notaular lines complete or at least extending half way across
mesoscutum with apex of forewing hyaline . ••. 127
123 (122) Occipital margin rounded. Forewing with infuscation on proximal side of hyaline spots,
which often appear joined, sometimes very weak. Scutellum flat with sculpture
distinctly deeper than that on mesoscutum . .,, . . ANABROLEPIS (p. 172)
NEOTROPICAL ENCYRTIDAE
123
Figs 6-14 6, Arrhenophagus chionaspidis Aurivillius ?, right antenna, outer aspect; 7
annulata De Santis ?, left antenna, outer aspect; 8, Solenaphycus vianai De Santis
and scutellum in dorsal view; 9, Brachyplatycerus sp. $, venation of left forewing;
sidneyensis Timberlake 2, left antenna, outer aspect; 11, Stemmatosteres apterus
mesoscutum and scutellum in dorsal view; 12, Blanchardiscus sp. $, scutellum and
dorsal view; 13, Blanchardiscus sp. j1, outline of left forewing and venation; 14,
aphelinoides Girault 9, left antenna, outer aspect.
, Aphycomastix
$, mesoscutum
10, Anarhopus
Timberlake $,
propodeum in
Ameromyzobia
124 JOHN S. NOYES
Occipital margin acute. Forewing with infuscation on proximal side of hyaline areas
strong. If sculpture on scutellum distinctly deeper than that of mesoscutum then
scutellum convex 124
124 (122) Mid tibial spur of unusual form, very flattened and apically rounded, spatulate (Fig. 61)
first two segments of funicle very short, at most half as long as the remainder which
are subequal in length and breadth DESANTISELLA (p. 191)
Mid tibial spur of normal form, gradually tapering with pointed apex. Antennae not as
above - . . .125
125 (124) Hind tibia distinctly flattened, not or hardly more than four times as long as broad (Fig.
64) CARABUNIA (p. 184)
Hind tibia not or hardly flattened, at least six times as long as broad. . . . 126
126 (125) Forewing with two opposite, more or less triangular hyaline spots HOPLOPSIS (p. 204)
Forewing with one or two hyaline fasciae, if two present then outer one often
interrupted MICROTERYS (p. 213)
127 (122) Forewing with longitudinal wedge-shaped streak, broadest at apex of wing, submarginal
vein with triangular expansion in apical third (sometimes indistinct) (Fig. 57).
Occipital margin more or less rounded . . . PARECHTHRODRY1NUS (p. 221)
Forewing without longitudinal wedge-shaped streak, submarginal vein without
triangular expansion. Occipital margin sharp 128
128 (127) Scutellum, mesoscutum and fronto vertex with deep thimble-punctures . . .129
At most either head or dorsum of thorax with thimble-punctures . . . .130
129 (128) At least first funicle segment longer than broad, funicle segments shortening towards
clava, clava at most as long as combined lengths of funicle segments three to six .
BOTHRIOTHORAX (p. 182)
All funicle segments distinctly transverse and lengthening towards clava, clava as long
or longer than funicle 130
130 (128, 129) First funicle segment distinctly longer than broad. Clava apically rounded . . 131
Either first funicle segment transverse or clava obliquely truncate . . . .134
131 (130) Forewing with distinct dark fascia situated about half way along submarginal vein,
remainder of forewing hyaline ANATHRIX (p. 174)
Forewing without distinct dark fascia, more generally darkened or with fascia arising at
marginal vein and gradually becoming paler towards wing apex . . . .132
132 (131) Cerci situated in apical third of gaster. Mandibles with one tooth and a broad
truncation PRIONOMASTIX (p. 223)
Cerci situated in basal half of gaster. Mandibles unidentate, bidentate or tridentate . 133
133 (132)* Ovipositor usually visible with sheaths bilaterally flattened. Mandibles broad and
tridentate, lower teeth subequal in length, upper tooth often short and indistinct.
Scutellum either flat with a distinct translucent apical flange (Fig. 60) or very convex
with flange indistinct or absent. Hypopygium reaching apex of gaster
HEMENCYRTUS (p. 202)
Ovipositor usually hidden, but if visible then sheaths are cylindrical. Mandibles
unidentate (Fig. 54) or bidentate with upper tooth long and curved, distinctly longer
than the short lower tooth (Figs 55, 56). Scutellum always fairly flat and without
apical translucent flange (although occasionally flange is present but distinctly
sculptured and not translucent). Hypopygium not reaching apex of gaster, often quite
long laterally and nearly reaching apex, but deeply incised medially
RHYTIDOTHORAX (p. 226)
134 (130) Apex of clava rounded. Hypopygium rarely extending further than half way along
gaster and never to near apex . ..... •)<••. '.'- .) . . . 135
* Note added while this paper was in press. Tachinaephagus Ashmead may run here but differs from Rhytido thorax in
having strongly tridentate mandibles and the hypopygium reaching the apex of the gaster, and from Hemencyrtus in
having a smooth convex scutellum with the apical flange very small, almost absent, and the eyes separated from the
occipital margin by about half the diameter of an ocellus (very nearly touching occipital margin in Hemencyrtus).
NEOTROPICAL ENCYRTIDAE
Figs 15-25 15, Pentalitomastix plethoricus Caltagirone 9, left antenna, outer aspect; 16, Tetartidava
yoshimotoi sp. n. 9, left antenna, outer aspect; 17, Acroaspidia myrmicoides Compere & Zinna 9, left
antenna, outer aspect; 18, Neodusmetia sangwani (Subba Rao) 9, mesoscutum and scutellum in
dorsal view; 19, Rhopus desantisellus Ghesquiere 9, scutellum in dorsal view; 20, Acerophagoides sp.
9, base of left forewing, upper surface; 21, Arrhenophagoidea sp. 9, head (slightly collapsed) viewed
from left side; 22, Ammonoencyrtus bonariensis (Brethes) 9, left antenna, outer aspect; 23,
Homosemion bennetti Annecke 9, left antenna, outer aspect; 24, Tetracnemoidea pretiosa
(Timberlake) 9, base of left forewing, upper surface; 25, Tetracnemus maculipennis (De Santis) 9,
venation of left forewing, upper surface.
126 JOHN S. NO YES
Apex of clava obliquely truncate (Figs 50, 59, 63, 101, cf. Figs 58, 62). Hypopygium
reaching or very nearly reaching apex of gaster (except in Epiencyrtus where clava is
very distinctly obliquely truncate).
In some genera clava may appear apically rounded in dry mounted material
depending on the way it has collapsed or the angle from which it is viewed, but in
these genera the hypopygium very distinctly reaches apex of gaster . . . .137
135 (134) Thorax orange with contrasting dark setae dorsally. Eyes with long hairs, each longer
than diameter of facet. Gaster very short, at most as long as scutellum (care!; in
material that has been in alcohol the gaster may have become distended) ,t— -• -»— .
APHYCOMORPHA (p. 175)
Thorax usually dark with contrasting pale setae, or, if orange then setae are pale. Eyes
with numerous short inconspicuous hairs, each no longer than diameter of facet.
Gaster much longer than scutellum . . 136
136 (135) Clava and segments three to six of funicle white contrasting with the brown scape,
pedicel and first two funicle segments .... TYNDARICHOIDES (p. 234)
Clava dark or yellow, never white METAPHYCUS (p. 212)
137 (134) Hypopygium reaching just over half way along gaster. Forewing almost hyaline, very
slightly and evenly infumate EPIENCYRTUS (p. 195)
Hypopygium reaching apex of gaster. Infuscation of forewing generally strong, or, if
weak then not evenly distributed 138
138 (137) Notaular lines at least anteriorly indicated. Infuscation of forewing limited to central
spot below marginal vein with occasionally middle third of wing infuscate, base and
apex hyaline. Clava usually distinctly shorter than funicle 139
Notaular lines completely absent. Infuscation of forewing usually more extensive with
either base or apex darkened. Clava often longer than funicle 141
139 (138) Notaular lines complete or meeting in middle (Fig. 1). Funicle segments and clava more
or less subequal in width so that clava is not, or hardly, wider than first funicle
segment (Fig. 50) HOMALOTYLUS (p. 203)
Notaular lines incomplete and never meeting in middle, at most extending two-thirds
across mesoscutum. Funicle segments distinctly broadening apically so that clava is at
least one and a half times as wide as first funicle segment (Figs 58, 63) . . ./. 140
140 (139) First funicle segment very short, less than half as long as second (Fig. 63). Notaular
lines anteriorly indicated only. Mid tibial spur very long and slightly longer than mid
basitarsus. Mesoscutum and scutellum completely orange . BRETHESIELLA (p. 183)
First funicle segment at least three-quarters as long as second (as in Fig. 58). Notaular
lines reaching half way across mesoscutum. Mid tibial spur at most as long as
basitarsus. Mesoscutum and scutellum usually with dark areas ISODROMUS (p. 206)
141 (138) Mesoscutum relatively short, at most slightly more than half as long as scutellum.
Frontovertex at narrowest point about two-thirds of head width or more
CHALCASPIS (p. 184)
Mesoscutum longer, at least about two-thirds as long as scutellum. Frontovertex
narrower, at narrowest point at most two-fifths head width 142
142 (141) Frontovertex narrower, at narrowest point at most one-sixth head width .
EURYRHOPALUS (p. 196)
Frontovertex broader, at narrowest point more than one-fifth head width . . .143
143 (142) Postmarginal vein distinctly longer, at least one and a half times as long as stigmal,
forewing most strongly infumate towards apex . ; . . . BLEPYRUS (p. 182)
Postmarginal vein shorter, not longer than stigmal, forewing most strongly infumate
towards base . . . . . . . . .'";-. ... 144
144 (143) Clava as long or longer than funicle and usually much wider than funicle, antennae
club-shaped (as in Fig. 62), all funicle segments transverse. Forewing : costal cell with
two or three lines of setae dorsally AENASWS (p. 170)
NEOTROPICAL ENCYRTIDAE
127
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L^:&£±^^-^
VJU
Figs 26-34 26, Chrysoplatycerus splendens (Howard) 2, left forewing, upper surface; 27,
Zaplatycercus fullawayi Timberlake 2, base of left forewing, upper surface; 28, Zarphopalus
putophilus Bennett 9, venation of left forewing, upper surface; 29, Hambletonia pseudococcina
Compere 2, venation of left forewing, upper surface; 30, Anicetus annulatus Timberlake 2, left
forewing, upper surface ; 3 1 , Hambletonia pseudococcina Compere 2, right antenna, outer aspect ; 32,
Habrolepis dalmanni (Westwood) 2, scutellum in dorsal view, showing apical lamelliform setae; 33,
Anabrolepis zetterstedtii (Westwood) $, head viewed from left side; 34, Hunterellus hookeri Howard
2, head in dorsal view.
128 JOHN S. NOYES
Clava about half as long as funicle and not much wider than funicle, antenna not club-
shaped, at least some funicle segments longer than broad or quadrate (Fig. 59).
Forewing : costal cell dorsally with at most a few setae at its apex ....
HEMAENASIUS (p. 202)
145 (92) All funicle segments longer than broad 146
Not all funicle segments longer than broad 177
146 (145) Head more or less purplish brown with area below level of dorsal part of antennal
scrobes shiny, metallic green with shallow shiny bottomed thimble-punctures.
Frontovertex with five pale yellow areas : two circular ones about midway between
anterior ocellus and antennal toruli on either side of frontovertex and touching eye
margins, and three elongate spots on same level just above toruli and interantennal
prominence all more or less joined to form a continuous transverse line (Fig. 67) .
PLATYLYCA (p. 222)
Head not as above "'.•'. 1 47
147(146) Exserted part of ovipositor at least as long as one-quarter length of gaster . . . 148
Exserted part of ovipositor at most as long as one-fifth length of gaster . . .156
148 (147) Forewing narrow, more than three times as long as broad. Costal cell about as wide as
diameter of submarginal vein (Fig. 70). Mandibles long, narrow and bidentate (as in
Fig. 40) ANAGYRUS(p. 173)
Forewing less than three times as long as broad, costal cell at least twice as broad as
diameter of submarginal vein. Mandibles usually tridentate, or, if bidentate then
relatively broad 149
149 (148) Postmarginal vein much longer than stigmal, speculum interrupted in lower half by two
or three lines of setae on dorsal surface of wing (Fig. 68) 150
Postmarginal vein not longer than stigmal, or, if slightly longer then speculum not
interrupted ; . 151
150 (149) Exserted part of ovipositor much longer than gaster and with sheaths cylindrical. Eyes
with moderately long dark setae DELOENCYRTUS (p. 191)
Exserted part of ovipositor shorter than gaster and with sheaths slightly flattened. Eyes
more or less naked NEAPSILOPHRYS (p. 215)
151 (149) Ovipositor sheaths bilaterally flattened, together at least twice as deep (at deepest point
of exserted part) as broad (at apex of gaster) "... 152
Ovipositor sheaths cylindrical, or, if slightly flattened then together less than twice as
deep (at deepest point of exserted part) as broad (at apex of gaster) . . 1 54
152(151) Mandibles bidentate ASEIRBA (p. 179)
Mandibles tridentate . , 153
153 (152) Hypopygium reaching apex of gaster. Scutellum flat with an apical translucent flange
(Fig. 60) or very convex. Occipital margin sharp . . HEMENCYRTUS (p. 202)
Hypopygium extending about half way along gaster. Scutellum quite flat but without
apical translucent flange. Occipital margin not sharp . . . CERCHYSIUS (p. 184)
154 (151) Occipital margin not sharp. Hypopygium reaching about halfway along gaster .
CERCHYSIUS (p. 184)
Occipital margin sharp. Hypopygium longer, reaching more or less to apex of gaster . 155
155 (154) Marginal vein punctiform (Fig. 69). Sculpture of scutellum often deep but never in
distinct whorls. Setae on mesoscutum dark. Axillae usually hardly raised above level
of scutellum and with posterior margin distinctly sculptured, very rarely as below .
COPIDOSOMA (p. 189)
Marginal vein at least about twice as long as broad (Fig. 65). Sculpture of scutellum
usually longitudinally striate-reticulate, more or less forming distinct whorls.
Mesoscutum usually with conspicuous white setae. Axillae step-like and distinctly
raised above level of scutellum with posterior margin almost vertical and very
polished . . I . APSILOPHRYS (p. 176)
NEOTROPICAL ENCYRTIDAE
129
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Figs 35-42 35, Prochiloneurus dactylopii (Howard) ?, left forewing, upper surface ; 36, Cheiloneurus sp.
?, left forewing, upper surface; 37, Gahaniella tertia Kerrich $, left antenna, outer aspect; 38,
Iceromyia flavifrons sp. n. 9, base of left forewing, upper surface; 39, Coccidencyrtus sp. ?, left
antenna, outer aspect; 40, Anagyrus insolitus (Howard) 9, left mandible; 41, Shenahetia masneri sp.
n. 9, head in frontal aspect; 42, Mariolaflava sp. n. 9, whole insect viewed from left side.
130 JOHN S. NOYES
156 (147) Forewing: speculum interrupted by 2 or 3 lines of setae in lower half, or completely
closed at this point by at least 6 or 7 lines of setae, basal cell completely pilose dorsally
with setae as dense or nearly as dense as in centre of wing (as in Figs 70, 71) . . 157
Forewing: speculum not interrupted or interrupted only by one line of setae, or
sometimes closed near hind margin of wing by two or three lines of setae, or, if
speculum interrupted then at least proximal one-third of basal cell naked dorsally and
setae are noticeably less dense than in centre of wing 160
157 (156) Postmarginal vein distinctly longer than stigmal, dorsum of thorax usually orange
(rarely black) with conspicuous dark setae on mesoscutum. Pedicel and flagellum
often concolorous, black ... . . LEPTOMASTIX (p. 206)
Postmarginal vein not longer than stigmal (Figs 70, 71). Dorsum of thorax dark or
occasionally orange, but always with pale setae on mesoscutum. Pedicel and flagellum
not unicolorous, at least apex of pedicel white, or clava yellowish . . . ^158
158 (157) Postmarginal vein very short or absent (as in Figs 70, 71). Sculpture of scutellum more
or less same as that on mesoscutum ANAGYRUS (p. 173)
Postmarginal vein as long, or nearly as long, as stigmal. Sculpture of scutellum deeper
and coarser than that on mesoscutum . . .159
159 (158) Scape distinctly broadest at middle. Scutellum with deep, dense punctiform sculpture.
Head in side view with posterior margin of eye adjacent to occipital margin for over
half its length. Funicle and whole of pedicel black, clava white . ANATHR1X (p. 174)
Scape broadest just above middle or parallel-sided. Scutellum with slightly elongate
sculpture. Head in side view with posterior margin of eye diverging from occipital
margin at much less than half its length. At least one funicle segment or apex of
pedicel paler than rest of flagellum APOANAGYRUS (p. 176)
160 (156) Apex of scutellum with a tuft of longer setae, or, if tuft indistinct then at least posterior
third of mesoscutum clothed in dense white setae which contrast strongly with dark
colour of mesoscutum. Scutellum with deep sculpture and not shiny. Submarginal
vein usually slightly expanded in apical third ..... ZAOMMA (p. 235)
Apex of scutellum without tuft of setae, at most with a few upright setae. Mesoscutum
not with contrasting pale setae, or, if with some pale setae then posterior third of
scutellum shiny. Submarginal vein usually not expanded in apical third . . .161
161 (160) Apical third of Submarginal vein with triangular expansion (Fig. 57) (sometimes
indistinct but usually indicated by a single long seta)
PARECHTHRODRYINUS (p. 221)
Submarginal vein without triangular expansion . . . . . . . .162
1 62 (161) Scutellum very convex and with numerous distinct short ridges running backwards from
axillae (Fig. 74) LIRENCYRTUS (p. 207)
Scutellum without numerous short ridges at base 163
163 (162) Eyes distinctly hairy, occasionally hairs pale and inconspicuous but always as long as
the diameter of facet. Propodeum medially at least one-fifth as long as scutellum and
usually with sculpture centrally which contrasts with the smoother lateral areas
(Fig. 60, cf. Fig. 121). Marginal vein at least about three times as long as broad;
occipital margin sharp .164
Eyes not hairy, or, if slightly hairy the following characters are all present: hairs on eyes
pale and inconspicuous and not longer than a facet, and occipital margin not sharp,
and marginal vein less than twice as long as broad. Propodeum medially usually less
than one-sixth as long as scutellum and without sculpture centrally (except in
Helegonatopus and Mucrencyrtus where the eyes are completely naked) . . . 165
164 (163) Ovipositor usually visible with sheaths bilaterally flattened. Mandibles broad and
tridentate, lower teeth subequal in length, upper tooth often short and indistinct.
Scutellum either flat with distinct translucent apical flange (Fig. 60) or very convex
with flange indistinct or absent. Hypopygium reaching apex of gaster
HEMENCYRTUS (p. 202)
Ovipositor usually hidden, but if visible then sheaths are cylindrical. Mandibles
unidentate (Fig. 54) or bidentate with upper tooth long and curved (Figs 55, 56),
NEOTROPICAL ENCYRTIDAE
131
^ii^.'/^iO-:-.;. • .-^ •-. \\ K^- V
53 56
Figs 43-56 43, Bennettisca flavigena sp. n. 9, left antenna, outer aspect ; 44, Iceromyiaflavifrons sp. n.
v, left antenna, outer aspect; 45, Moorella fulviceps Cameron 9, left hind tibia and tarsus, outer
aspect; 46, Adelencyrtus moderatus (Howard) 9, left mandible; 47, Coccidencyrtus sp. $, left
mandible; 48, Parablastothrix sp. 9, head viewed from left side; 49, Encyrtus lecaniorum (Mayr) $,
left forewing venation, upper surface; 50, Homalotylus terminalis (Say) $, left antenna, outer aspect;
51, Hemaenasius sp. 9, left antenna, outer aspect; 52, Neococcidencyrtus sp. $, left forewing, upper
surface; 53, Ericydnus lamasi (Domenichini) 9, left forewing, upper surface; 54-56, Rhytidothorax
spp., left mandibles.
132 JOHN S. NOYES
distinctly longer than lower tooth. Scutellum always fairly flat and without apical
translucent flange (although occasionally a flange is present, but distinctly sculptured
and not translucent). Hypopygium not reaching apex of gaster, often quite long
laterally and nearly reaching apex, but deeply incised medially ....
RHYTWOTHORAX (p. 226)
165 (163) Flagellum bicolorous: funicle brown, clava white or yellow. Mandibles with two teeth
and a truncation ' \ LOHIELLA (p. 209)
Flagellum unicolorous, or if as above, then mandibles are bidentate . . . 1 66
166 (165) Scape relatively short, not longer than malar space. Antennal toruli high on head, more
than twice the length of each torulus from mouth margin. All funicle sements about
as long and as wide as pedicel (as in Fig. 37).
Mandibles without teeth or with one tooth and a truncation GAHANIELLA (p. 197)
Scape distinctly longer than malar space or antennal torulus less than twice its own
length from mouth margin. First funicle segment distinctly shorter than pedicel, at
most three-quarters as long and distinctly less in diameter . .r* . . .167
167 (166) Clava at least three-quarters as long as funicle. Legs completely yellowish white.
[Coccidencyrtus malloi Blanchard] (p. 188)
Clava at most two-thirds as long as funicle. Legs orange or at least partially darkened 168
168 (167) Mandibles with one or two teeth and a truncation (Figs 72, 73). Hypopygium at most
reaching just over halfway along gaster. Clava apically round or with a short oblique
truncation .,, ; ~ .. ..169
Mandibles with three acute teeth (Fig. 97), or, if third (upper) tooth developed but
truncate, then hypopygium reaches or nearly reaches apex of gaster and apex of clava
is transversely truncate 172
169 (168) Either marginal vein more r less punctiform (at most about one and a half times as long
as broad) or postmarginal absent or nearly absent (Figs 75, 76, 81). Mesoscutum
often bright metallic green 170
Marginal vein at least twice as long as broad, postmarginal distinct, usually as long or
nearly as long as radial (Fig. 79). Mesoscutum rarely green 172
170 (169) Head lenticular, in side view more than twice as long as deep (Fig. 82) and in dorsal view
over two and a half times as wide as deep (Fig. 83). Antennal scrobes very shallow
and indistinct , SIMMONDSIELLA (p. 229)
Head of more normal proportions, in side view less than twice as long as deep and in
dorsal view less than twice as wide as deep. Antennal scrobes moderately to deeply
impressed 171
171 (170) Posterior margin of mesoscutum gently curved or with either side straight and centrally
slightly angled so that axillae appear to meet centrally and not greatly separated by
posterior margin of mesoscutum (Fig. 78). Setae on antenna very short, the longest
less than half the diameter of the first funicle segment. Stigmal vein distinctly longer
than marginal, postmarginal distinct and only slightly shorter than marginal (Fig. 75).
Frontovertex at narrowest point usually at least one-third head width. Parasites of
psyllid nymphs. . PSYLLAEPHAGUS (p. 226)
Posterior margin of mesoscutum produced backwards in middle third so that axillae
appear to be widely separated (Fig. 77). Setae of antennae longer, the longest about as
long or longer than the diameter of the first funicle segment. Stigmal vein rarely
longer than marginal, postmarginal usually not distinct or at most not as long as
marginal (Fig. 76). Frontovertex generally less than one-fifth head width, rarely a
great deal wider OOENCYRTUS (p. 217)
172 (168, 169) Marginal vein punctiform and at most half as long as stigmal (Figs 85, 90, 109, also
as in 69). Hypopygium extending to at least five-sixths along gaster . . .173
Marginal vein distinctly longer than broad, at least slightly more than half as long as
stigmal (Fig. 79). Hypopygium extending to at most three-quarters along gaster . 175
173 (172) Hypopygium with apex rounded, not produced as narrow projecting tongue. Head and
thorax metallic COPIDOSOMA (p. 189)
NEOTROPICAL ENCYRTIDAE
Figs 57-68 57, Parechthrodryinus nitidus (Howard) $, left forewing, upper surface; 58, Isodromus
iceryae Howard ?, left antenna, outer aspect; 59, Hemaenasius confusus Ashmead ?, left antenna,
outer aspect; 60, Hemencyrtus sp. ?, scutellum and propodeum, dorsal view; 61, Desantisella
trifasciata De Santis $, mid tibial spur; 62, Aenasius phenacocci Bennett $, left antenna, outer aspect;
63, Brethesiella sp. ?, left antenna, outer aspect; 64, Carabunia myersi Waterston $, left hind tibia
and tarsus, outer aspect; 65, Apsilophrys sp. 9, left forewing venation, upper surface; 66, Anusioptera
aureocincta Brues $, left antenna, outer aspect (clava missing); 67, Platylyca quadraticeps De Santis
$, head in frontal aspect; 68, Neapsilophrys flavipes sp. n. ?, left forewing base, upper surface.
134 JOHN S. NOYES
Hypopygium apically produced into narrow projecting tongue which may be almost as
long as half length of gaster (Fig. 84). Head and thorax orange . . . .174
174 (173) Forewing: setae on dorsal surface of costal cell limited to line from about halfway along
to apex of cell near wing margin, basal cell distinctly less pilose than centre of wing
(Fig. 85). Antennal scrobes not deep or well defined and with interantennal
prominence dorsally more or less rounded . . . . PARAMUCRONA (p. 219)
Forewing : setae on dorsal surface of costal cell rather dense, particularly in distal half,
not limited to a single line near wing margin, basal cell about as densely pilose as
centre of wing (Fig. 90). Antennal scrobes quite deep and well defined and with
interantennal prominence dorsally produced as a long, very acute ridge (Fig. 80) .
MUCRENCYRTUS (p. 213)
175(172) Scutellum longitudinally striate with silky lustre . . MERCETENCYRTUS (p. 211)
Scutellum not longitudinally striate and without silky lustre . . . . .176
176 (175) Basal two-thirds or three-quarters of Scutellum with raised very rough sculpture which
contrasts very strongly with the highly polished apex. Centre of propodeum with
distinct sculpture. Venation of forewing yellowish or testaceous. Apex of clava
rounded HELEGONATOPUS (p. 200)
Scutellum more or less evenly sculptured, never with sculpture at base contrasting so
strongly with that at apex. Centre of propodeum smooth. Venation of forewing
usually dark brown. Clava apically with at least a short oblique truncation .
SYRPHOPHAGUS (p. 230)
177 (145) First funicle segment longer than broad . .178
First funicle segment not longer than broad 207
178 (177) Exserted part of ovipositor with sheaths distinctly flattened, together more than twice as
deep as broad at apex of gaster. Mesoscutum not with distinct thimble-punctured
sculpture and scutellum never smooth and shiny . . . . . . .179
Ovipositor not exserted, or, if exserted then sheaths more or less cylindrical, or, if
slightly flattened (Zeteticontus) then mesoscutum has deep thimble-punctured
sculpture or scutellum is smooth and shiny . " . 181
179 (178) Eyes relatively small, frontovertex at narrowest point about half head width, malar
space about two-thirds length of eye. Posterior ocellus separated from eye margin by
at least its own diameter. Postmarginal vein very short, about one-third length of
marginal which is shorter than stigmal. Exserted part of ovipositor longer than gaster.
Mandible with two strong teeth GONZALEZIA (p. 197)
Eyes larger, frontovertex at narrowest less than half head width, malar space at most
one-third as long as an eye. Posterior ocellus separated from eye margin by less than
its own diameter. Postmarginal vein nearly as long or longer than stigmal. Exserted
part of ovipositor much shorter than gaster, often only just visible. Mandible with at
least a very short third (upper) tooth • . . .180
180(179) Hypopygium reaching apex of gaster ..... HEMENCYRTUS (p. 202)
Hypopygium reaching to about halfway along gaster. Encyrtus conformis Howard (p. 195)
181 (179) Frontovertex or mesoscutum with conspicuous deep thimble-punctures usually
separated by at most their own diameters 182
Sculpture not as above, generally reticulate and shallow, occasionally with distinct
piliferous punctures but these are small and separated by much more than their own
diameters 186
182 (181) Scutellum very smooth and shiny, devoid of sculpture except occasionally in basal half,
at least apical third very polished (occasionally basal two-thirds has thimble-
punctured sculpture, but spaces between punctures are polished) ....
ZETETICONTUS (p. 236)
All of scutellum distinctly sculptured . . . . . . . . . .183
183 (182) Marginal vein punctiform or absent (Figs 86, 87) 184
Marginal vein at least twice as long as broad 185
135
NEOTROPICAL ENCYRTIDAE
u'/A v^v^*
vlMvV W,\>
\ \ H r \ \^\v ,\ v \
Kx^VlMiXx\\\^
\,\ XN I \ \ X\N\V\\.\
74
78
Figs 69-78 69, Copidosoma virescens De Santis 9, left forewing venation, upper surface; 70, Anagyrus
bellator (De Santis) 9, right forewing base, upper surface; 71, Anagyrus sp. 9, left forewing base,
upper surface; 72, Psyllaephagus trellesi (Blanchard) 9, left mandible; 73, Psyllaephagus
rotundiformis (Howard) ?, left mandible; 74, Lirencyrtus primus sp. n. ^, scutellum, dorsal aspect;
75, Psyllaephagus rotundiformis (Howard) ?, left forewing venation, upper surface ; 76, 77, Ooencyrtus
trinidadensis Crawford 9, (76) left forewing venation, upper aspect, (77) scutellum, dorsal aspect; 78,
Psyllaephagus rotundiformis (Howard) 9, scutellum, dorsal aspect.
136 JOHN S. NOYES
184 (183) Marginal vein more or less absent, stigmal vein usually arising before submarginal
reaches wing margin (Fig. 86). Frontovertex with shiny bottomed piliferous
punctures. Mandibles with two teeth and a truncation LOCHITOENCYRTUS (p. 209)
Marginal vein present, punctiform (as in Fig. 87). Piliferous punctures on frontovertex
not shiny bottomed. Mandibles with three acute teeth . BOTHRIOTHORAX (p. 182)
185 (183)* Clava apically rounded. Punctures on frontovertex not shiny bottomed. Mandibles with
three acute teeth .; . .«-. . . HEMENCYRTUS (p. 202)
Clava obliquely truncate (Fig. 88). Punctures on frontovertex shiny bottomed.
Mandibles with two teeth and a truncation or with a rudimentary third (upper) tooth
HELMECEPHALA (p. 200)
186 (181) Hypopygium clearly extending past apex of epipygium (Fig. 89). Postmarginal vein
longer than stigmal. Exserted part of ovipositor at least half as long as gaster
COCCIDOCTONUS (p. 188)
Hypopygium clearly not extending past apex of epipygium. Postmarginal vein not
longer than stigmal, or, if so then ovipositor is more or less hidden or only slightly
exserted .... 187
187 (186) All funicle segments longer than broad, except the second which is anelliform (Fig. 92)
CYDERIUS (p. 190)
Funicle not as above :„, , . 188
188 (187) Propodeum medially at least one-sixth as long as scutellum and with some sculpture
centrally (Fig. 60) 189
Propodeum medially at most one-seventh as long as scutellum and more or less smooth
centrally 194
189 (188) Eyes and frontovertex with long conspicuous setae. Hypopygium reaching apex of
gaster. Mandibles tridentate. Apex of scutellum with translucent flange (Fig. 60)
(occasionally if scutellum is very convex flange may be indistinct or absent). Body
dark and shiny HEMENCYRTUS (p. 202)
Eyes and frontovertex without conspicuous long setae, or, if present then mandibles
unidentate or bidentate and hypopygium does not reach apex of gaster. Apex of
scutellum never with a translucent flange, or, if flange present then body completely
orange and not shiny 190
190 (189) Scutellum very convex with very deep, rough, elongate sculpture, apex with a small,
thin, more or less triangular, foliate extension (Fig. 94). Clava obliquely truncate
(Fig. 106) . . . .... AMAURILYMA (p. 171)
Scutellum rather flat, occasionally with deep elongate sculpture, but never rough and
without apical extension. Clava apically rounded or transversely truncate . . 191
191 (190) Mandibles unidentate (Fig. 52) or bidentate (Figs 55, 56). Eyes with conspicuous long
hairs. Apical margin of scutellum usually with an inconspicuous carina (as in Fig.
121) . RHYTIDOTHORAX (p. 226)
Mandibles tridentate. Eyes without conspicuous hairs, at most with a few short pale
ones. Apical margin of scutellum without carina, often quite smooth . . . 192
192 (191) Head and thorax mostly metallic green or greenish blue. Gaster longer than thorax .
PARECHTHRODRYINUS (p. 221)
Head or thorax largely orange. Gaster shorter than thorax. .. ' . . . .. 193
193 (192) Head black. Clava apically truncate (Fig. 200) . . PROTYNDARICHOIDES (p. 224)
Head orange. Clava apically rounded (Fig. 189) . . . MUCRENCYRTUS (p. 213)
194 (188) Thorax and head mostly orange and yellow, never metallic. ...
Metaphycus ceroplastae (Dozier) (p. 212)
At least thorax, excluding legs, dark brown or green and metallic or shiny. . . 195
* Note added while this paper was in press. Tachinaephagus Ashmead may run here, but differs from Hemencyrtus in
having a smooth convex scutellum with the apical flange very small, almost absent, and the eyes separated from the
occipital margin by about half the diameter of an ocellus (very nearly reaching the occipital margin in Hemencyrtus).
NEOTROPICAL ENCYRTIDAE
137
86
Figs 79-86 79, Syrphophagus sp. $, left forewing venation, upper surface; 80, Mucrencyrtus insulanus
sp. n. 2, head in frontal aspect; 81-83, Simmondsiella flaviptera sp. n., $, (81) left forewing base,
upper surface, (82) head viewed from left side, (83) head, dorsal view; 84, Mucrencyrtus insulanus sp.
n. $, gaster viewed from right side; 85, Paramucrona brasiliensissp. n. 1, left forewing base, upper
dorsal view; 90, Mucrencyrtus insulanus sp. n. j1, left forewing base, upper surface; 91,
138 JOHN S. NOYES
195 (194) Head not unicolorous, mostly dark brown with white or pale yellow areas on lower part
of face, particularly around antennal scrobes and on interantennal prominence (Fig.
93) . TACHARDIOBIUS (p. 231)
Head unicolorous without yellow or white markings 196
196 (195) Scape short, not longer than malar space, all funicle segments and pedicel subquadrate
and subequal in length and breadth, not distinctly widening towards apex of antenna,
clava about as wide as first funicle of segment (as in Fig. 37)
Mandibles without teeth or with one tooth and a broad truncation
GAHANIELLA (p. 197)
Scape longer, longer than malar space, funicle segments and pedicel not subequal in
length and breadth, with either the first funicle segment distinctly shorter than
pedicel, or funicle segments broadening distinctly so that clava is wider than first
funicle segment 197
197 (196) Mandibles with one or two teeth and a truncation (Figs 72, 73, 91). Clava apically
rounded or obliquely truncate. Hypopygium, at most, reaching only just over half
way along gaster 198
Mandibles with three acute teeth (Fig. 97), or, if the upper tooth is truncate then
hypopygium more or less reaches apex of gaster and apex of clava is transversely
truncate .200
198 (197) Posterior margin of mesoscutum gently curved or with either side straight and centrally
slightly angled so that axillae appear to meet centrally or are not greatly separated by
posterior margin of mesoscutum (Fig. 78). Marginal vein punctiform, postmarginal
at least half as long as stigmal (Fig. 75). Antenna with short setae, the longest less
than half as long as the diameter of the first funicle segment. Mesoscutum bright
metallic green or blue-green.
Parasites of psyllid nymphs ... . PSYLLAEPHAGUS (p. 226)
Posterior margin of mesoscutum usually produced backwards in middle third so that
axillae appear to be widely separated (as in Fig. 77). Marginal vein longer than broad,
or, if punctiform then stigmal vein is very short and rarely longer than marginal and
postmarginal usually indistinct. Antenna with longer setae, the longest about as long,
or longer than diameter of first funicle segment. Mesoscutum usually not bright
metallic green or blue-green 199
199 (198) Stigmal vein very short, not longer than the very short more or less punctiform
marginal, post-marginal indistinct or absent (as in Fig. 76) . OOENCYRTUS (p. 217)
Marginal vein always longer than broad, stigmal relatively long, postmarginal distinct
and nearly as long as stigmal . . . 200
200 (197, 199) Clava entire and obliquely truncate (as in Fig. 96) . . LITOMAST1X (p. 208)
Either clava three-segmented or apically rounded . . . . . . . 201
201 (200) Hypopygium reaching to at least five-sixths along gaster. Antennal torulus very close to
mouth margin, separated by less than half its length. Funicle segments more or less
subequal in width. Marginal vein punctiform, postmarginal vein sometimes very
short or almost absent. Mandibles always with three acute teeth or rarely with upper
tooth blunt or truncate . COPIDOSOMA (p. 189)
Hypopygium reaching to at most two-thirds along gaster. Antennal torulus separated
from mouth margin by more than half its length. Funicle segments distinctly
widening towards apex of antenna. Marginal vein longer than broad, postmarginal
vein always at least half as long as stigmal. Mandibles usually not with three acute
teeth .202
202 (201) Exserted part of ovipositor at least as long as one-third length of gaster. Scutellum
dorsally very flat with deep punctate to slightly elongate sculpture GR1SSELL1A (p. 197)
Ovipositor not, or hardly, protruding. Either scutellum convex, occasionally with
punctate sculpture, or flat and quite smooth . . . . . . . 203
203 (202) Scutellum longitudinally striate with silky lustre ....' . MERCETENCYRTUS (p. 211)
Scutellum not longitudinally striate and without silky lustre 204
NEOTROPICAL ENCYRTIDAE
97
•> /
Figs 87-97 87, Bothriothorax sp. 9, left forewing venation, upper surface; 88, Helmecephala
albisetosa sp. n. 9, left antenna, outer aspect; 89, Coccidoctonus trinidadensis Crawford 9, gaster in
dorsal view; 90, Mucrencyrtus insulanus sp. n. <$, left forewing base, upper surface; 91,
Echthrodryinus sp. 9, left mandible, 9; 92, Cyderius urbicola sp. n., left antenna, outer aspect; 93,
Tachardiobius nigr icons De Santis 9, head in frontal aspect (reconstructed from collapsed head of
holotype); 94, Amaurilyma sp. 9, scutellum in dorsal view; 95, Szelenyiola sp. 9, head in frontal
aspect ; 96, Litomastix truncatella (Dalman) 9, left antenna, outer aspect ; 97, Copidosoma koehleri
Blanchard 9, left mandible.
140 JOHN S. NO YES
204 (203) Eyes and frontovertex with conspicuous long dark setae. Scutellum moderately flat,
quite smooth and shiny EXORISTOB1A (p. 196)
Eyes and frontovertex without conspicuous long dark setae. Scutellum very convex and
with distinct sculpture . . .. .. .. • " . . . . . . 205
205 (204) Scutellum (without axillae) relatively long and narrow, a little longer than broad at base
and not evenly rounded. Forewing venation usually dark brown .......
SYRPHOPHAGUS (p. 230)
Scutellum (without axillae) relatively broader, little wider than long and rather evenly
rounded. Forewing venation usually yellowish or testaceous 206
206 (205) Apex of scutellum very shiny, contrasting with anterior two-thirds or three-quarters
which has distinct (sometimes deep) sculpture. Marginal vein not more than twice as
long as broad. Head, in side view, less than twice as long as deep
ECHTHRODRYINUS (p. 193)
Apex of scutellum with distinct sculpture. Marginal vein at least twice as long as broad.
Head, in side view, at least twice as long as deep . . TR1CHOMASTHUS (p. 234)
207 (177) Dorsum of thorax, including axillae, orange or yellow . . . .. ,. ' .. 208
At least axillae or mesoscutum dark •'«*: ,-' tf./ - . .-- rr-k 212
208 (207) Body dorso-ventrally flattened. Mesoscutum entire. Mandibles bidentate with teeth of
equal length . . . RHOPUS (p. 226)
Body not flattened. Mandibles not bidentate, or, if bidentate then either upper tooth is
much longer than lower (Rhytidothorax) or mesoscutum has notaular lines anteriorly
indicated (Cirrhencyrtus) . . -. -. 209
Propodeum medially at least one-sixth as long as scutellum and with rough sculpture
centrally (as in Fig. 121). Frontovertex and eyes with distinct long setae. Scutellum
only slightly convex. Forewing with setae in basal cell longer than those in centre of
wing RHYTIDOTHORAX (p. 226)
Propodeum medially at most one-eighth as long as scutellum and without sculpture
centrally. Frontovertex and eyes not usually with conspicuous long setae. Forewing
with setae in basal cell usually not, or hardly, longer than those in centre of wing.
Scutellum usually distinctly convex . ,„.,.».. 210
Mesoscutum and scutellum clothed in conspicuous dark setae, notaular lines completely
absent. Gaster very short, at most as long as scutellum (care! if material has been in
alcohol gaster may be distended) .... APHYCOMORPHA (p. 175)
Mesoscutum and scutellum usually with pale setae, at most scutellum with conspicuous
dark setae, notaular lines usually anteriorly indicated or complete. Gaster much
longer than scutellum . . . . . . 211
211 (210) Mandibles bidentate, or with third (upper) tooth rudimentary. Marginal vein about
three times as long as broad. Setae on scutellum often dark and conspicuous.
Exserted part of ovipositor over half as long as gaster. Hypopygium reaching apex of
gaster *•'**. CIRRHENCYRTUS (p. 187)
Mandibles with three acute teeth. Marginal vein more or less punctiform. Setae on
mesoscutum and scutellum pale. Exserted part of ovipositor rarely as long as half
length of gaster. Hypopygium occasionally reaching apex of gaster ....
METAPHYCUS (p. 212)
212 (207) Head with at least three pale membranous lines, two of which run near inner eye
margins from near anterior ocellus to near antennal toruli and a third transverse line
which connects these near the top of the antennal scrobes (Fig. 95 also as in 128)
Body slightly dorso-ventrally flattened, mandibles tridentate . ~ -^ , . 213
Head without membranous lines. .... 214
213(212) Clava entire , ,-,,.. SZELENYIOLA (p. 231)
Clava three-segmented , , AVETIANELLA (p. 179)
214 (212) Body dorso-ventrally flattened. Cerci situated about halfway along gaster. Ovipositor
not or hardly visible. Mandibles bidentate . . RHOPUS (p. 226)
NEOTROPICAL ENCYRTIDAE
141
m--^*
Figs 98-112 98, Forcipestricis sp. 2, scutellum showing tubercles; 99, Forcipestricis gazeaui Burks
scutellum showing tubercles; 100, Coelopencyrtus gargaris (Walker) 2, left mandible; 101,
Epiencyrtus sp. 2, left antenna, outer aspect; 102, Papaka confusor sp. n. 2, left mandible; 103,
Neococcidencyrtus crouzelae De Santis 2, left mandible; 104, Grissellia terebrata sp. n. 2, left
mandible; 105, Copidosoma sp. 2 near silvestrii (Costa Lima), left mandible; 106, Amaurilyma sp. 2,
left antenna, outer aspect; 107, Copidosoma sp. 2 near silvestrii (Costa Lima), right antenna, outer
aspect; 108, Cicoencyrtus angustifrons sp. n. 2, head, frontal aspect; 109, Copidosoma sp. 2 near
silvestrii (Costa Lima), left forewing venation, upper surface; 110, Grissellia terebrata sp. n. 2, left
forewing base, upper surface; 111, Anabrolepis zetterstedtii (Westwood) J1, left antenna, outer
aspect; 112, Anarhopus sidneyensis Timberlake J, left antenna, outer aspect.
142 JOHN S. NO YES
Body not dorso-ventrally flattened, or, if slightly flattened then cerci are nearer to base
of gaster than to middle, or mandibles are not bidentate, or the exserted part of the
ovipositor is at least as long as about one-quarter length of gaster . . . .215
215 (212) Clava one- or two-segmented . . . 216
Clava three-segmented .... . 223
216 (215) Clava one-segmented ' V . 217
Clava two-segmented . . 220
217 (216) Notaular lines present in anterior half of mesoscutum. Head and large areas of thorax
yellow or orange ISODROMUS (p. 206)
Mesoscutum entire. Head and thorax dark . 218
218 (217) Apex of clava obliquely truncate (Fig. 96). Postmarginal vein shorter than stigmal
LITOMASTIX (p. 208)
Apex of clava apically rounded or only very slightly truncate. Postmarginal vein
distinctly longer than stigmal (as in Fig. 130) . . . . . . . . 219
219 (218) Mesoscutum shiny with shallow, transverse sculpture contrasting with the deep,
longitudinally semi-striate sculpture of scutellum which is silky in appearance. First
two funicle segments not much shorter than third . . PARALEUROCERUS (p. 219)
Both mesoscutum and scutellum with longitudinal, semi-striate sculpture and both more
or less silky in appearance. Each of first two funicle segments about half as long as
third . AGENIASPIS (p. 171)
220 (216) Antennal torulus separated from mouth margin by much less than own length. Body at
least slightly dorso-ventrally flattened. Mesoscutum entire. Cerci situated about half
way along gaster. Mandibles with three acute teeth. . . . . . .221
Antennal torulus separated from mouth margin by about own length. Body
occasionally slightly dorso-ventrally flattened. Mesoscutum sometimes with notaular
lines anteriorly indicated. Cerci situated near apex of gaster. Mandibles with two
teeth and a truncation, or quadridentate • . . 222
221 (220) Antennal scrobes moderately impressed and horseshoe-shaped. Protruding part of
ovipositor equal to about one-quarter length of gaster and more or less straight, not
curved downwards towards apex. Frontovertex with distinct sculpture .
TANYENCYRTUS (p. 231)
Antennal scrobes absent or represented by extremely short, shallow grooves, much
shorter than the length of an antennal torulus (Fig. 41). Protruding part of ovipositor
equal to at least about half length of gaster and curved downwards towards apex.
Frontovertex highly polished SHENAHETIA (p. 227)
222 (220) Mandibles quadridentate. Notaular lines anteriorly indicated . . ,..'".
ANDINOENCYRTUS (p. 174)
Mandibles bidentate with truncation. Mesoscutum entire ......
[Coccidencyrtus] obesus De Santis (p. 188)
223 (215) Exserted part of ovipositor at least as long as half length of gaster. Hypopygium
extending well past apex of epipygium so that it is clearly visible in dorsal view
(Fig. 89). Postmarginal vein longer than stigmal . . COCCIDOCTONUS (p. 188)
Ovipositor not or hardly protruding, or, if as long or longer than half length of gaster
then hypopygium does not extend past apex of epipygium and postmarginal vein is
not longer than stigmal . . . . ' . . •','»'• • . . . 224
224 (223) Scutellum smooth and shiny, not sculptured except occasionally at extreme base . 225
At least anterior half of scutellum with distinct sculpture (occasionally quite shallow) 228
225 (224) Hypopygium clearly reaching apex of gaster. Frontovertex and mesoscutum often with
thimble-punctured sculpture . . . .; . . . . ZETETICONTUS (p. 236)
Hypopygium reaching at most two-thirds along gaster, usually no more than half way.
Frontovertex and mesoscutum never with thimble-punctured sculpture . . . 226
226 (225) Propodeum medially at least one-sixth as long as scutellum and with rough central area
which contrasts with smooth lateral areas (as in Fig. 121). Frontovertex and eyes with
NEOTROPICAL ENCYRTIDAE
143
Figs 113-125 113, Aenasius advena (Compere) J, left antenna, outer aspect; 114, Helegonatopus
pseudophanes Perkins J, left antenna, outer aspect; 115, B lane har discus sp. j1, right antenna, outer
aspect; 116, Ham bletonia pseudococcina Compere ^, left antenna, outer aspect; 117, Ameromyzobia
aphelinoides Girault J, left antenna, outer aspect; 118, Hambletonia pseudococcina Compere 3,
scutellum in dorsal view; 119, Desobius sp. J1, left antenna, outer aspect; 120, Coelopencyrtus
gargaris (Walker) J, right antenna, outer aspect; 121, Rhytidothorax sp. J, scutellum and
propodeum in dorsal view; 122, Litomastix truncatella (Dalman) J, left antenna, outer aspect; 123,
Homalotylus flaminius (Dalman) j\ left antenna, outer aspect; 124, Isodromus iceryae Howard J1,
right forewing venation, upper surface; 125, Homalotylus flaminius (Dalman) J, right forewing
venation, upper surface.
144 JOHN S. NO YES
very distinct long setae. Scutellum without tubercles. Mandibles unidentate or
bidentate . . . . . . . . . RHYTWOTHORAX (p. 226)
Propodeum medially at most one-eighth as long as scutellum and without distinct
central rough area. Frontovertex and eyes without long setae, or, if present, then
either scutellum has tubercles (Fig. 98, also cf. Fig. 99) (best seen in cleared slide
mounts) or mandibles are tridentate .. - . • • - ~ 227
227 (226) Scutellum distinctly convex in side view with tubercles or pits (Fig. 98, also cf. Fig. 99)
(best seen in cleared slide mounts). Frontovertex smooth and shiny, or rarely
sculptured, always with conspicuous long setae. Eyes hairy. Mandibles bidentate or
tridentate with middle tooth longest .... FORC1PESTRICIS (p. 196)
Scutellum almost flat and without tubercles. Frontovertex with distinct sculpture and
without any very conspicuous long setae. Eyes not conspicuously hairy. Mandibles
with three teeth of about equal length .... HABROLEPOIDEA (p. 199)
228 (224) Apex of scutellum with tuft of setae, or, if tuft indistinct then usually at least posterior
one-third of mesoscutum clothed in white setae which contrast strongly with dark
colour of mesoscutum. Scutellum not shiny. Submarginal vein slightly expanded in its
apical three-quarters ZAOMMA (p. 235)
Apex of scutellum without tuft of setae, at most with few longer upright setae.
Mesoscutum without contrasting pale setae, or, if pale setae present then submarginal
vein is not expanded apically 229
229 (228) Both mesoscutum and scutellum with fine, longitudinal, more or less striate sculpture
giving both a silky appearance. Clava as long or longer than funicle (Fig. 160) ^ -
DESOBIUS(p 192)
Thorax sculpture different : at most either only scutellum or mesoscutum with striate
sculpture, never both. Clava occasionally as long or longer than funicle, but usually
shorter ; ... 230
230(229) Notaular lines present in anterior half of mesoscutum, or complete . . . .231
Notaular lines absent or indicated at extreme anterior margin of mesoscutum only . 233
23 1 (230) Setae on dorsal surface of forewing, proximal to the speculum, in four lines, remainder
of basal cell naked. Scutellum with deeper more elongate sculpture than mesoscutum.
Notaular lines complete. Gaster completely yellow ventrally *---
CIBDELOENCYR TVS (p. 185)
Basal cell more or less completely pilose, although occasionally sparsely pilose towards
base. Sculpture of scutellum more or less the same as mesoscutum. Notaular lines not
complete, extending about half way across mesoscutum. Venter of gaster not yellow,
except perhaps in some Metaphycus 232
232 (231) Marginal vein at least one and a half times as long as broad. Mid tibia without dark
rings SOLENOENCYRTUS (p. 230)
Marginal vein punctiform, or hardly longer than broad. Mid tibia yellow, usually with
dark rings METAPHYCUS (p. 212)
233 (230) Mesoscutum dark brown or black, not shiny or metallic, posterior margin of pronotum
usually white or distinctly paler than mesoscutum. Hypopygium never extending to
apex of gaster. Mandibles with three acute teeth . . . METAPHYCUS (p. 212)
Mesoscutum at least slightly shiny, posterior margin of pronotum the same colour as
mesoscutum, or darker and not distinctly paler. Hypopygium often reaching apex of
gaster - 234
234 (233) Mandibles unidentate or bidentate 235
Mandibles tridentate (third (upper) tooth sometimes rudimentary) or with one or two
teeth and a truncation . . . , , „ ' , " , , ... 238
235 (234) Hypopygium convex, more or less boat-shaped and reaching apex of gaster . . 236
Hypopygium hardly convex, reaching at most to three-quarters along gaster . , 237
236 (235) Postmarginal vein very short, much shorter than stigmal, submarginal vein with a
triangular expansion in its apical third (as in Fig. 20). Frontovertex with reticulate
sculpture. Body not robust ,' , . PAURIDIA (p. 221)
NEOTROPICAL ENCYRTIDAE
145
136
figs 126-136 126, Pentalitomastix plethoricus Caltagirone 3, left forewing venation, upper surface;
127, Baeoencyrtus platys De Santis J\ scutellum in dorsal view; 128, Avetianella sp. J1, head in
frontal aspect; 129, Metaphycus sp. J1, left forewing base, upper surface; 130, Ageniaspis fuscicollis
(Dalman) J1, left forewing venation, upper surface; 131, Anagyrus sp. J1, apical segments of right
antenna to show perpendicular sensillae; 132, Hunterellus hookeri Howard 3, left hind tibia and
tarsus, outer aspect ; 1 33, Microterys ! flaws (Howard) J1, left forewing venation, upper surface ; 1 34,
Anagyrus sp. J1, gaster in dorsal view; 135, Prionomastix sp. J1, left forewing venation, upper surface;
136, Encyrtus infelix (Embleton) J, left forewing venation, upper surface.
146 JOHN S. NOYES
Postmarginal vein much longer than stigmal, submarginal vein without triangular
expansion. Frontovertex with thimble-punctured sculpture. Body very robust .
BLEPYRUS(p. 182)
237 (235) Scutellum with tubercles or pits (as in Figs 98, 99) (best seen in cleared slide mounts).
Frontovertex almost always smooth and shiny. Propodeum short medially, at most
one-sixth as long as scutellum and smooth. Proximal funicle segments often
anelliform, contrasting with longer distal segments. Mandibles bidentate.
FORCIPESTRICIS (p. 196)
Scutellum without tubercles or pits. Frontovertex with distinct sculpture and not very
shiny. Propodeum medially usually longer than one-sixth scutellum, sculptured
centrally and contrasting with the smoother lateral areas (as in Fig. 121). Funicle
segments usually becoming shorter distally. Mandibles unidentate (Fig. 54) or
bidentate (Figs 55, 56) RHYTIDOTHORAX (p. 226)
238 (234) Propodeum shiny and with distinct median carina and medially at least one-fifth as
long as the flat scutellum which has deeper more elongate sculpture than mesoscutum.
Submarginal vein with triangular expansion in its apical one-third (Fig. 57) (often
indistinct). Marginal vein at least twice as long as broad. Occipital margin more or
less rounded PARECHTHRODRYINUS (p. 221)
Propodeum normally without distinct median carina and medially at most one-sixth as
long as scutellum, or, if longer and with median carina, then scutellum distinctly
convex, submarginal vein usually without a triangular expansion and occipital
margin acute. Marginal vein often punctiform and scutellum occasionally with
elongate sculpture 239
239 (238) Scutellum with distinct tubercles or pits, either scattered or in a distinct group on each
side (as in Figs 98, 99) (best seen in cleared slide mounts). Frontovertex with
conspicuous long setae and almost always smooth and very shiny. Eyes very hairy.
Hypopygium not reaching apex of gaster.
Mandibles with middle tooth longest. Antennae often with at least one or two
proximal funicular segments anelliform, contrasting with distal segments which are
subquadrate FORCIPESTRICIS (p. 196)
Scutellum without tubercles or pits, or, if some present then hypopygium reaches apex
of gaster (Amaurilymd). Frontovertex with distinct sculpture and usually without
long conspicuous setae. Eyes rarely very hairy 240
240 (239) Exserted part of ovipositor well over half length of gaster. Frontovertex very narrow, at
narrowest point at most about one-fifth head width and sometimes only slightly wider
than anterior ocellus (Fig. 108) CICOENCYRTUS (p. 186)
Ovipositor not or hardly protruding, never with protruding part as long as half length of
gaster. Frontovertex never as narrow as above, or, if as narrow (Euryrhopalus) then
ovipositor more or less hidden . . . . . . . . ' . .241
241 (240) Clava large, about twice as wide as sixth funicle segment and longer than funicle and
with a very strong oblique truncation which is longer than length of ventral surface of
clava, all funicle segments transverse and unicolorous (Fig. 101). Marginal vein about
three times as long as broad. Hypopygium not reaching apex of gaster .
EPIENCYRTUS (p. 195)
Clava usually relatively smaller (although sometimes much longer than funicle) and with
rounded apex, or, if truncate then truncation is transverse and marginal vein is
punctiform, or, if truncation is oblique then it is very short, much shorter than length
of ventral side of clava and the clava is shorter than funicle or the hypopygium
reaches the apex of gaster 242
242 (241) Scutellum at most, only slightly convex, more or less flat, not shiny, relatively shallow in
side view and with deep punctate or elongate sculpture which is distinctly deeper than
that on shiny mesoscutum (except in Papaka where mesoscutum may also have
punctate sculpture), sculpture never shallower than that on mesoscutum. Head often
triangular in side view with frontovertex flat in profile 243
Scutellum distinctly convex, or, if dorsally flat then sides are long and convex and it is
relatively deep in side view, or it is rather shiny with very much shallower sculpture
NEOTROPICAL ENCYRTIDAE
Figs 137-144 137, Shenahetia masneri sp. n. j1, left forewing venation, upper surface; 138,
Parechthrodryinus nitidus (Howard) J, left forewing venation, upper surface; 139, Tanyencyrtus
divisus De Santis 3, left forewing venation, upper surface; 140, Helegonatopus sp. J, scutellum and
propodeum in dorsal view; 141, 142, Arrhenophagoidea sp., $, (141) left hind tibia and tarsus, outer
aspect, (142) left forewing, upper surface; 143, 144, Arhopoidiella carinata sp. n., 3, (143) left
antenna, outer aspect, (144) genitalia.
148 JOHN S. NOYES
than mesoscutum. Scutellum only occasionally with deeper sculpture than
mesoscutum. Head rounded in profile 248
243 (242) Mandibles with four acute teeth (Fig. 46). Mid tibia completely yellow without dark
bands . .,..-. . . . •«.-.," . ADELENCYRTUS (p. 170)
Mandibles with one tooth or two teeth and a truncation (the latter may have the
truncate third (upper) tooth rudimentary (Fig. 102)). Mid tibia often brown or yellow
with dark bands 244
244 (243) Frontovertex, at narrowest point, at least one-third head width. Clava yellow or
testaceous, funicle usually concolorous or slightly paler, rarely darker. Clava long,
almost as long as, or longer than, funicle, usually much longer . , -.
COCCIDENCYRTUS (p. 188)
Frontovertex, at narrowest point, at most one-quarter head width. Clava and first four
funicle segments brown, often segments five and six yellowish or testaceous. Clava at
most as long as funicle . . -. •. -. . . . 245
245 (244) Sides and apex of scutellum steep, flat and polished. Clava about as long as funicle and
segments five and six of funicle usually yellow . NEOCOCCIDENCYRTUS (p. 216)
Sides of scutellum short and convex, not flat or polished. Clava at most about two-
thirds as long as funicle, or, if longer then funicle segments five and six are
concolorous with rest of funicle .......... 246
246 (245) Mandibles with upper tooth rudimentary, almost absent (Fig. 102). Sixth funicle
segment often yellow and contrasting with rest of funicle and clava. Setae on funicle
short, the longest much shorter than diameter of first funicle segment PAPAKA (p. 217)
Mandibles with third (upper) tooth broad and truncate (Fig. 104). Sixth funicle segment
concolorous with rest of flagellum. Setae on funicle long, the longest at least as long
as diameter of first funicle segment 247
247 (246) Postmarginal vein distinct, nearly as long as stigmal (as in Fig. 110) GRISSELLIA (p. 197)
Postmarginal vein more or less absent or extremely short (as in Fig. 76) .
OOENCYRTUS (p. 217)
248 (242) Antenna with funicle not unicolorous, with contrasting dark and light segments.
PARALITOMASTIX (p. 219)
Antenna with funicle unicolorous without contrasting dark and light segments . . 249
249 (248) Postmarginal vein much longer than stigmal y^. . 250
Postmarginal vein not longer than stigmal , , / 251
250 (249) Frontovertex, at narrowest point, at least one-sixth head width . BLEPYRUS (p. 182)
Frontovertex, at narrowest point, at most one-eighth head width >
EURYRHOPALUS (p. 196)
251 (249) Hypopygium reaching apex of gaster, or, if reaching to only four-fifths of gaster then
marginal vein punctiform (Fig. 109) and mandibles have three acute teeth (Fig. 97) (if
third (upper) tooth blunt (Fig. 105) then clava transversely truncate (Fig. 107)) . 252
Hypopygium, at most, reaching just over halfway along gaster. Marginal vein distinctly
longer than broad, or, if punctiform then mandibles have at most two acute teeth (if
with third (upper) tooth developed and blunt then clava apically rounded or distinctly
obliquely truncate) 254
252 (251) Antenna very short, about as long as width of head, mandibles with third (upper) tooth
apically rounded or truncate (Fig. 100) .... COELOPENCYRTUS (p. 189)
Antenna longer, at least one and a half times as long as width of head. Mandibles with
upper tooth acute , 253
253 (252) Antenna with clava obliquely truncate (Fig. 106). Scutellum with very deep, rough,
longitudinal sculpture and short, thin, apical extension (Fig. 94) ....
AMAURILYMA (p. 171)
Antenna with clava apically rounded or with transverse truncation (Fig. 107) not
obliquely truncate. Scutellum not with rough longitudinal sculpture and without
apical extension . , . COPIDOSOMA (p. 189)
NEOTROPICAL ENCYRTIDAE
149
Figs 145-152 145-148, Arhopoidiella carinata sp. n. ?, (145) left antenna, outer aspect, (146) head
in frontal aspect; (147) left forewing base, upper surface, (148) genitalia; 149, 150, Bennettisca
flavigena sp. n. ?, (149) genitalia, (150) left forewing base, upper surface; 151, 152, Cicoencyrtus
angustifrons sp. n. 31, (151) left antenna, outer aspect, (152) genitalia.
150 JOHN S. NO YES
254 (251) Posterior margin of mesoscutum gently curved or slightly angled so that axillae appear
to be more or less touching apically (Fig. 78). Forewing: marginal vein punctiform or
hardly longer than broad, stigmal vein long and postmarginal distinct, longer than
marginal (Fig. 75). Antenna with setae relatively short, the longest much shorter than
half diameter of the first funicle segment. Mesoscutum bright metallic green or blue-
green.
Parasites of psyllid nymphs PSYLLAEPHAGUS (p. 226)
Posterior margin of mesoscutum usually rather convex in median third so that axillae
appear to be widely separated (as in Fig. 77). Marginal vein at least about twice as
long as broad, or, if punctiform (as in Fig. 76) then stigmal is short, not longer than
marginal and postmarginal very short or indistinct, not longer than marginal and
axillae distinctly separated by posterior margin of mesoscutum. Antenna with setae
relatively longer, the longest at least half as long as diameter of the first funicle
segment; mesoscutum occasionally green or blue-green but never very bright . . 255
255 (254) Marginal vein punctiform, rarely longer than broad and often relatively thick, stigmal
vein short, almost sessile, postmarginal vein very short or absent (as in Fig. 76).
Egg parasites OOENCYRTUS (p. 217)
Marginal vein at least one and a half times, and usually at least twice, as long as broad,
stigmal vein always relatively long and postmarginal always distinct and as long or
nearly as long as stigmal (as in Fig. 74) (occasionally marginal vein punctiform, but
postmarginal vein nearly as long as stigmal vein) 256
256 (255) Scutellum quite flat and shiny with very shallow inconspicuous sculpture. Eyes and
frontovertex with very distinct, long, dark setae. . . . EXORISTOBIA (p. 196)
Scutellum never flat and shiny, usually distinctly convex or with deep sculpture. Eyes
and frontovertex without conspicuous setae, or, if these are present then scutellum
very convex or with deep, conspicuous sculpture 257
257 (256) Scutellum (without axillae) at most as long as broad, usually broader than long and
almost uniformly convex, sculpture (of scutellum) often deeply punctate, particularly
in basal half, apical half or third often smooth and shiny. Eyes usually rather hairy.
Marginal vein sometimes punctiform .... ECHTHRODRYINUS (p. 193)
Scutellum (without axillae) longer than broad and normally abruptly convex at sides
and apex, sculpture uniform and never deeply punctate, occasionally striate (in latter
case the extreme apex is quite shiny). Eyes more or less naked. Marginal vein always
distinctly longer than broad 258
258(257) Scutellum longitudinally striate and with silky lustre . MERCETENCYRTUS (p. 211)
Scutellum longitudinally striate and without silky lustre 259
259 (258) Setae on flagellum finer and shorter, usually with the longest not longer than diameter of
first funicle segment. Generally smaller species: — length less than 1 mm. Parasites of
Aphididae (or perhaps psyllid nymphs) .... APHIDENCYRTUS (p. 175)
Setae on flagellum usually coarser and longer, the longest about as long or longer than
the diameter of the first funicle segment, generally larger species: — length greater
than 1-5 mm. Parasites of syrphid larvae (Diptera) . . SYRPHOPHAGUS (p. 230)
Key to genera (males)
The following genera are not included because the males are unknown or their presence in the
Neotropics is doubtful: Acroaspidia, Anathrix, Andinoencyrtus , Anusioptera, Aphycomastix,
Arrhenophagoidea, Atelaphycus, Aztecencyrtus, Bennettisca, Calliencyrtus, Cerchysius,
Cibdeloencyrtus, Cyderius, Deloencyrtus , Dicarnosis, Ectroma, Epiencyrtus, Exoristobia,
Habrolepoidea, Homosemion, Hoplopsis, Lohiella, Mariola, Moorella, Neapsilophrys,
Paramucrona, Parencyrtus, Pelmatencyrtus, Pheidoloxenus, Platylyca, Prionomitus,
Protyndarichoides, Simmondsiella, Solenoencyrtus, Tetarticlava, Tineophoctonus, Tyndarichoides.
1 Funicle with long rami (Fig. 112) . > . . . . . . 2
Funicle simple, without rami 6
NEOTROPICAL ENCYRTIDAE
151
Figs 153-160 153-155, Cicoencyrtus angustifrons sp. n. $, (153) left antenna, outer aspect, (154) left
forewing base, upper surface, J, (155) genitalia; 156-158, Amaurilyma sp., (156) genitalia, 9, (157)
left forewing base, upper surface, 9, (158) left antenna, outer aspect, $ ; 159, Cyderius urbicola sp. n.
9, left forewing base, upper surface, 160, Desobius convexus sp. n. 9, left antenna, outer aspect.
152 JOHN S. NO YES
2 Funicle five-segmented .. ••-'.. - .. .. ' .. .. ., . . .... 3
Funicle six-segmented . ». ....... . . . . 4
3 (2) Fifth funicle segment with short apical ramus (Fig. 112) about one-quarter length of
segment. Scutellum with slightly deeper, coarser sculpture than mesoscutum, greenish
ANARHOPUS(p. 173)
Fifth funicle segment simple. Scutellum with slightly shallower sculpture than
mesoscutum, purple, contrasting with the green mesoscutum
TETRACNEMOIDEA (p. 233)
4 (2) First funicle segment simple and transverse . , -..'• TETRACNEMUS (p. 233)
First funicle segment with ramus . . . . . .. . 5
5 (4) Scape not reaching anterior ocellus. Forewings hyaline. Body metallic green or blue-
green .... PARABLASTOTHRIX (p. 218)
Scape reaching well above level of posterior ocelli. Forewings usually with some
infuscate areas. Body orange to dark brown, never metallic . HEXACLADIA (p. 203)
6 (1) Funicle composed of two to five anelliform segments, clava extremely long and sausage-
shaped (Figs 111, 113), more than twice (usually more than three times) as long as
scape . .. 7<
If all funicle segments are anelliform then clava at most one and a half times as long as
scape and not sausage-shaped 8
7 (6) Mandibles with four teeth or with two acute teeth and a truncation. Funicle two-
segmented, clava with base rounded and articulated to funicle in centre (Fig. 111).
Marginal vein as long or longer than short stigmal, postmarginal shorter than radial
ADELENCYRTUS (p. 170), PLAG1OMERUS (p. 222), ANABROLEPIS
(p. 172), HABROLEPIS (p. 199), WOMALOPODA (p. 203),
1PSEUDHOMALOPODA (p. 225)
Mandibles with two or three teeth. Funicle two- to five-segmented (sometimes in dorsal
view only two segments visible, the others being hidden by base of clava, this being
most easily observed in slide-mounted material), clava with base not rounded,
dorsally produced towards pedicel thus partially obscuring funicle and articulated on
ventral side (Fig. 1 13). Marginal vein more or less punctiform, postmarginal as long
or longer than stigmal which is several times longer than marginal .
CHALCASPIS (p. 184), A ENASWS (p. 170)
8 (6) Funicle of fewer than six segments . . . .. 9
Funicle of six or more segments .. .. 25>
9 (8) Funicle three- or four-segmented, all segments more or less anelliform
BLEPYRUS(p 182)
Funicle five-segmented.
First segment may be anelliform or partially obscured by pedicel or second funicle
segment giving funicle a four-segmented appearance .. .. .. - ., , . .. 10)
10 (9) Mesoscutum with notaular lines complete (as in Fig. 8) or extending at least about
quarter of way towards posterior margin of mesoscutum ! •..- . ; -^ .. ,. Ill
Notaular lines completely absent, or indicated only at extreme anterior margin of
mesoscutum .... . . . .. "'.. .. • w .. / .. . '..-. 14
llll (10) Notaular lies only visible in anterior half of mesoscutum . .. . .. . .. 1C£
Notaular lines complete or very nearly reaching posterior margin of mesoscutum . 1CP
12 (1:1) First funicle segment anelliform, the remaining four much longer than broad and with
very short setae, shorter than one-third diameter of segments. Propodeum at least
one-third length of scutellum and shiny. Forewing with a large infumate area below
marginal vein extending to posterior margin of wing . . MEROMYZOBIA (p. 212)
First funicle segment at least about as long as broad, not shorter than remainder. Setae
on funicle well over half diameter of segments. Propodeum at most one-fourth as long
as scutellum and not very shiny. Forewing very slightly infumate in anterior half .
ACEROPHAGOIDES (p. 169)
NEOTROPICAL ENCYRTIDAE
153
Figs 161-169 161, Desobius sp. 31, genitalia; 162, 163, Desobius convexus sp. n. ?, (162) genitalia,
(163) left forewing base, upper surface; 164, Grissellia terebrata sp. n. $, left antenna, outer aspect,
9; 165, Grissellia sp. J1, left antenna, outer aspect; 166, Grissellia terebrata 9, genitalia; 167,
Exoristobia sp. 9, left antenna, outer aspect; 168, Exoristobia sp. 9, left forewing base, upper
surface; 169, Exoristobia sp. 9, genitalia.
154 JOHN S: NOYES
13 (11) Thorax yellow or orange, scutellum with apex well rounded and with sculpture not
deeper than that on mesoscutum SOLENAPHYCUS (p. 230)
Thorax dark and metallic. Scutellum with apex more or less pointed and with sculpture
distinctly deeper and coarser than that on mesoscutum . COCCIDAPHYCUS (p. 187)
14 (10) Wings absent STEMMA TOSTERES (p. 230)
Wings fully developed or extending past apex of gaster . . . . . . 15
15 (14) Antennae about as long as body, all funicle segments at least about twice as long as
broad and with setae at least as long as diameter of segments, scape triangular and
broadest at base (Fig. 1 14). Body metallic green . . HELEGONATOPUS (p. 200)
Antennae much shorter than body, generally not more than half as long, usually some
funicle segments transverse or subquadrate, rarely all longer than broad, none as
much as twice as long as broad, setae on funicle snorter than diameter of segments,
scape usually widest at about middle. Body often orange, rarely green . . . 16
16 (15) Clava two-segmented, funicle with a small, very transverse first segment, the remainder
much longer and each about twice as broad as long (Fig. 115). Scutellum produced
apically in a tongue-like extension which curls downwards towards the posterior
margin of propodeum (Fig. 12) BLANCH A RDISCUS (p. 182)
Clava entire, funicle not as above, scutellum not produced apically . . . . 17
17 (16) Postmarginal vein longer than stigmal .... EURYRHOPALUS (p. 196)
Postmarginal vein not longer than stigmal 18
18 (17) Scutellum with central ridge which arises between the apices of the axillae and extends
posteriorly for about half its length (Fig. 118). Scape triangular in side view, about
twice as long as broad and widest in the middle (Fig. 1 16). Mandibles bidentate
HAMBLETONIA (p. 200)
Scutellum without central ridge. Scape not triangular, if broadened then ventral edge
gradually curved. Mandibles tridentate 19
19 (18) Head and thorax dark orange-brown to black and at least moderately shiny. Pronotum
not paler than mesoscutum. Neither mesoscutum nor scutellum with contrasting pale
setae 20
Head and thorax usually orange or yellow, or, if dark then matt and without metallic
reflections and with contrasting pale setae and pronotum distinctly paler than
mesoscutum <. 22
20 (19) Eyes covered with numerous short hairs. Interantennal prominence not covered in dense
white setae 21
Eyes naked. Interantennal prominence clothed in dense white setae ARCHINUS (p. 176)
21 (20) All funicle segments transverse, clava about as long as funicle. Top of antennal scrobes
without carina. Legs completely yellow .... BOTHRIOCRAERA (p. 182)
All funicle segments longer than broad, clava much shorter than funicle (Fig. 143), top
of antennal scrobes often with a distinct carina which runs transversely from eye to
eye (as in Fig. 146). Legs darkened ARHOPOIDIELLA (p. 177)
22 (19) First two funicle segments anelliform, their combined lengths less than length of third
which is more or less subquadrate as are remaining segments (Fig. 117).
AMEROMYZOBIA (p. 172)
First two funicle segments not contrasting as greatly with other segments as above, each
usually slightly smaller than the third 23
23 (22) Maxillary palpi two-segmented, labial palpi one-segmented. . TIMBERLAKIA (p. 233)
Maxillary palpi three- or four-segmented, labial palpi two-segmented .... 24
24 (23) Antenna more or less unicolorous, yellow or yellowish orange. Pronotum and
mesoscutum usually concolorous . . '.'- . . ACEROPHAGUS (p. 169)
Antenna with clava white or yellowish white and contrasting with dark brown funicle
segments, or, if these are white then with dark brown scape. Pronotum often
distinctly paler than mesoscutum PSEUDAPHYCUS (p. 225)
NEOTROPICAL ENCYRTIDAE
155
Figs 170-179 170, 171, Helmecephala sp. J\ (170) left antenna, outer aspect, (171) genitalia; 172, 173,
Helmecephala albisetosa sp. n. ?, (172) base of left forewing, upper surface, (173) genitalia; 174, 175,
Iceromyia flavifrons sp. n. 3, (174) left antenna, outer aspect, (175) left forewing venation; 176,
Iceromyia sp. $, genitalia; 177-179, Lirencyrtus primus sp. n., (177) left antenna, outer aspect, ?,
(178) left forewing base, upper surface, $, (179) left antenna, outer aspect, j1.
156 JOHN S. NOYES
25 (8) Frontovertex with very conspicuous shiny-bottomed thimble-punctures. Forewing with
stigmal vein arising before submarginal reaches wing margin (as in Fig. 86). Occipital
margin sharp. Funicle segments at least as wide as pedicel, the first five longer than
broad and with extremely short hairs. . . LOCHITOENCYRTUS (p. 209)
Combination of characters not as above . . . . . .., . . . 26
26 (25) Both mesoscutum and scutellum convex and with longitudinally strigose sculpture. Post-
marginal vein shorter than stigmal (as in Fig. 163). Flagellum variable. Funicle seven-
segmented with clava one- or two-segmented (Fig. 1 19) or funicle six-segmented with
clava two- or three-segmented DESOBIUS (p. 192)
If both mesoscutum and scutellum have longitudinally strigose sculpture then scutellum
flat or postmarginal vein longer than stigmal (Fig. 130). Flagellum with funicle always
six-segmented, clava usually entire* . . 27
27 (26) Funicle not unicolorate, with basal segments dark and apical segments white.
Frontovertex narrow, not more than twice as broad as anterior ocellus. Forewing
darkened with a transverse hyaline band in apical third . . COMPERIA (p. 189)
Combination of characters not as above 28
28 (27) Not all funicle segments longer than broad 29
All funicle segments longer than broad 56
29 (28) Funicle with at least one segment with setae at least one and a half times as long as its
diameter 30
Setae on funicle segments at most as long as diameter of funicle segments . . . 31
30 (29) Tarsi four-segmented. Scutellum without tubercles or pits . ARRHENOPHAGUS (p. 178)
Tarsi five-segmented. Scutellum with tubercles or pits (easy to see on cleared, slide
mounted material) (as in Figs 98, 99) . . . . FORC1PESTRICIS (p. 196)
31 (29) Mid-tibial spur expanded at apex and sack-like (as in Fig. 61).
Funicle consisting of two anelliform segments and four segments which are longer
than broad DESANTISELLA (p. 191)
Mid-tibial spur normal, long and pointed at apex 32
32 (31) Wings shortened, not reaching apex of gaster. Body yellow. Notaular lines more or less
complete ECHTHROPLEX1ELLA (p. 194)
Wings fully developed, at least reaching apex of gaster 33
33 (32) Postmarginal vein longer than stigmal (as in Fig. 130) 34
Postmarginal vein not longer than stigmal .40
34 (33) First funicle segment at least slightly narrower than pedicel, at least some funicle
segments as long as, or longer than broad . . . PARALEUROCERVS (p. 219)
First funicle segment distinctly broader than pedicel, or, if not then all funicle segments
transverse 35
35 (34) Frontovertex, above antennal scrobes, with coarser sculpture than below this point
which is smooth and shiny, the area below the top of the antennal scrobes contrasting
greatly in colour with the area above and also more shiny. Antennal torulus separated
from mouth margin by distance equal to about its own length. Hairs on eyes generally
short and inconspicuous 36
Frontovertex and area below top of antennal scrobes of similar sculpture and colour.
Antennal torulus often separated from mouth margin by much less than its own
length. Setae on eyes usually long and very conspicuous . . . - •* • -. . 37
36 (35) Sculpture of scutellum slightly coarser and more elongate (particularly in centre) than
that on mesoscutum. Postero-lateral margins of mesoscutum not slightly raised and
without smoother sculpture than rest of mesoscutum . ZAPLATYCERUS (p. 235)
* Note added while this paper was in press. Tachinaephagus Ashmead has a seven-segmented funicle and scutellum which
is smooth and shiny.
NEOTROPICAL ENCYRTIDAE
157
186
Figs 180-189 180, 181, Lirencyrtus primus sp. n., (180) genitalia, ?, (181) genitalia, ^; 182, 183,
Lohiella flaviclava (Howard), ?, (182) left antenna, outer aspect, (183) base of left forewing, upper
surface; 184, 185, Mariolaflava sp. n., ?, (184) left forewing base, upper surface, (185) left antenna,
outer aspect; 186-189, Mucrencyrtus insulanus sp. n., (186) genitalia, J1, (187) genitalia, ?, (188) left
antenna, outer aspect, J, (189) left antenna, outer aspect, ?.
158 JOHN S. NOYES
Sculpture of scutellum more or less same as, or shallower than, on mesoscutum.
Postero-lateral margins of mesoscutum slightly turned upwards and with slightly
smoother sculpture (particularly on inner face) than rest of mesoscutum .
CHR YSOPLA TYCERUS (p. 185)
37 (35) Cerci in apical half of gaster . . ,. ' . . .' AEPTENCYR TVS (p. 171)
Cerci in basal half of gaster > . . . . 38
38 (37) Posterior ocellus nearly touching margin of eye, separated by at most the diameter of a
facet, also separated from occipital margin by distance slightly greater than from eye.
Frontovertex usually with distinct thimble-punctured sculpture . AENASWS (p. 170)
Posterior ocellus separated from eye margin by distance equal to diameter of at least two
facets and often much closer to occipital margin than to eye. Frontovertex never with
thimble-punctured sculpture . . 39
39 (38) Posterior ocellus separated from occipital margin by about diameter of a facet or nearer
occipital margin than eye margin. POL: OOL about 8:1 . ZARHOPALUS (p. 235)
Posterior ocellus separated from occipital margin by about diameter of two facets and
about equidistant from occipital and eye margins. POL : OOL about 6:1.
COELASPIDIA (p. 188)
40 (33) Base of scape with distinct spine (Fig. 120). Vertex at anterior ocellus abruptly raised
much above dorsal level of eyes COELOPENCYRTUS (p. 189)
Base of scape without spine. Vertex at anterior ocellus hardly raised above dorsal level
of eyes . . 41
41 (40) Apical two-thirds of submarginal vein with a triangular expansion (as in Fig. 20)
(mandibles bidentate, notaular lines indicated at anterior margin of mesoscutum) .
PAURIDIA (p. 221)
Submarginal vein without triangular expansion, or, if it is slightly thickened apically
then notaular lines more or less complete or mandibles tridentate .... 42
42 (41)* Clava two-segmented, scutellum more or less flat, smooth and shiny . . . .
ZETETICONTUS (p. 236)
Clava entire, scutellum with clear sculpture and usually distinctly convex ... 43
43 (42) Mandibles unidentate or bidentate 44
Mandibles tridentate or with one or two teeth and a truncation "46
44 (43) Notaular lines reaching one-third across mesoscutum. Frontovertex at narrowest point
at most one-third length of head width. Head and thorax mostly orange, clava and
apical funicle segment white C1RRHENCYRTVS (p. 187)
Notaular lines completely absent. Frontovertex at narrowest point at least about half
length of head width. Head and thorax dark and metallic. Clava and funicle brown 45
45 (44) Frontovertex and eyes with conspicuous long setae, frontovertex at narrowest point
about half length of head width. Propodeum medially at least one-sixth as long as
scutellum (Fig. 121) RHYTIDOTHORAX (p. 226)
Eyes naked and frontovertex without conspicuous setae and very wide, at narrowest
point well over three-fifths as wide as head. Propodeum medially at most one-eighth
as long as scutellum NEODUSMETIA (p. 216)
46 (43) Clava obliquely truncate (Figs 122, 123) ;' 47
Clava apically rounded i . . . . .'"" 50
47 (46) Notaular lines present and extending at least half way across mesoscutum ... 48
Notaular lines completely absent or indicated at anterior margin of mesoscutum only. 49
48 (47) Marginal vein slightly longer than broad, postmarginal distinct (Fig. 125). Mesoscutum
with notaular lines more or less reaching posterior margin (as in Fig. 1).
HOMALOTYLUS (p. 203)
* Note added while this paper was in press. Asitus Erdos runs here but differs from other included genera in having an
extremely flattened leaf-like body, a longitudinally divided pronotum and the forewing with the marginal fringe at least
one-fifth as long as maximum width of wing.
NEOTROPICAL ENCYRTIDAE
159
Figs 190-200 1 90, 1 9 1 , Neapsilophrysflavipes sp. n. $, ( 1 90) genitalia, ( 1 9 1 ) left antenna, outer aspect ;
192-196, Papaka confusor sp. n., (192) left antenna, outer aspect, ?, (193) genitalia, ?, (194) left
forewing base, upper surface, ?, (195) genitalia, J1, (196) left antenna, outer aspect, J1; 197, 198,
Paramucrona brasiliensis sp. n. $, (197) left antenna, outer aspect, (198) genitalia; 199, 200,
Protyndarichoides nigriceps sp. n. 9, (199) left forewing base, upper surface, (200) left antenna, outer
aspect.
160 JOHN S. NO YES
Marginal vein punctiform or absent, postmarginal very short or absent (Fig. 124).
Notaular lines at most only reaching half way across mesoscutum ....
ISODROMUS (p. 206)
49 (47) Head and thorax dark, metallic.
Scutellum with purple reflections contrasting with the green mesoscutum . . '
LITOMASTIX (p. 208)
Head and thorax yellow or orange yellow, not metallic . . BRETHESIELLA (p. 183)
50 (46) Mesoscutum metallic green, scutellum convex and metallic green with strong purple
reflections . . 51
Mesoscutum and scutellum not as above, or, if metallic then scutellum flat . . 52
51 (50) Marginal vein punctiform, postmarginal absent (Fig. 126) PENT A LITOMASTIX (p. 222)
Marginal vein at least slightly longer than broad, postmarginal distinct, about half as
long as stigmal LITOMASTIX (p. 208)
52 (50) Scutellum triangular in dorsal view, the sides more or less straight and with apex acute
(Fig. 127), sculpture elongate and contrasting with shallower more polygonal
sculpture of mesoscutum. Clava (in slide mounted material) with indications of being
three-segmented. Mandibles with three acute teeth . . BAEOENCYRTUS (p. 180)
Scutellum shield-shaped in dorsal view with sides and apex more or less rounded and
occasionally with deep elongate sculpture. Clava without indications of
segmentation. If scutellum is almost triangular in dorsal view then mandibles have
upper tooth rudimentary. ........... 53
53 (52) Thorax orange. Scutellum and posterior margin of pronotum with a few contrasting
dark setae. Mandibles with third (upper) tooth rudimentary
CIRRHENCYRTUS (p. 187)
Thorax dark brown, or, if orange then without contrasting dark setae. Legs often with
dark areas. Mandibles with three acute teeth or sometimes with upper tooth
rudimentary (Papaka, as in Fig. 102) 54
54 (53) Frontovertex narrow, at narrowest point not more than about one-seventh of head
width. Speculum not interrupted (Fig. 154) . . . CICOENCYRTUS (p. 186)
Frontovertex wider, at narrowest point at least about one-fifth of head width. Speculum
sometimes interrupted by one to three lines of setae on dorsal surface of wing (Fig.
129) 55
55 (54) Mandibles with three acute teeth METAPHYCUS (p. 212)
Mandibles with upper tooth rudimentary and not acute (as in Fig. 102) PAPAKA (p. 217)
56 (28) Head dark with several pale markings around antennal scrobes, eyes margined white,
interantennal prominence with an inverted dark 'V originating at the antennal toruli
(as in Fig. 93) (setae on funicle about as long as the diameter of the funicle segments)
TACHARDIOBWS (p. 231)
Not as above 57
57 (56) Head with at least three membranous lines, two longitudinal ones from about level with
anterior ocellus along inner margins of the eyes and outer margins of scrobes to near
antennal toruli, these being joined by a third transverse line along the top of the
scrobes (Fig. 128, also cf. Fig. 95).
Body flattened, mandibles tridentate 58
Head without such membranous lines ~l_'i_l ._ ..•". .... 59
58 (57) Setae on funicle not longer than diameter of funicle segments. Labial palpi one-
segmented SZELENYIOLA (p. 231)
Setae on funicle at least about twice as long as diameter of funicle segments. Labial palpi
three-segmented : . . . . AVETIANELLA (p. 179)
59 (57) Funicle segments with longest setae at most equal to diameter of corresponding segment 60
Funicle segments with at least setae longer (usually at least one and a quarter times as
long) than diameter of corresponding segment . 85
NEOTROPICAL ENCYRTIDAE
161
Figs 201-209 201, Protyndarichoides nigriceps sp. n. 9, genitalia; 202-205, Shenahetia masneri sp. n.,
(202) left antenna, outer aspect, 2, (203) left forewing base, upper surface, 9, (204) left antenna, outer
aspect, j, (205) genitalia, j1; 206, Shenahetia sp. 2, genitalia; 207, Simmondsiella ftaviptera sp. n. ?,
left antenna, outer aspect; 208, 209, Tetarticlava yoshimotoi sp. n. i, (208) left forewing base, upper
surface, (209) genitalia.
162 JOHN S. NO YES
60 (59) Hind tibia distinctly bilaterally flattened, at most four times as long as broad (as in
Fig. 64) . CARABUNIA (p. 184)
Hind tibia not flattened, or, if slightly flattened then more than five times as long as
broad . . ... . . . , ' ' . . ... . . 61
61 (60) Postmarginal vein at least slightly longer than stigmal (Figs 133, 135, 136). . . 62
Postmarginal vein not longer than stigmal 68
62 (61) Scutellum with a distinct apical tuft of setae. Mandibles without teeth . . .
EI\CYRTUS(p. 195)
Scutellum without apical tuft of setae (occasionally a few longer setae at apex).
Mandibles with at least one distinct tooth . . . ... . . 63
63 (62) At least anterior three-quarters of mesoscutum with dark setae and longitudinally
striate sculpture, posterior one-half to one-third of mesoscutum clothed in pale setae 64
Mesoscutum not with elongate sculpture, or if sculpture elongate then setae are entirely
dark throughout : « . 65
64 (63) Frontovertex, particularly between ocelli, with coarse, regular, punctate sculpture.
Sculpture in centre of mesonotum not, or hardly, longitudinally striate in contrast to
that at sides which is very elongate DIVERSINERVUS (p. 193)
Frontovertex with irregular, shallow, reticulate sculpture. Sculpture in centre of
mesoscutum longitudinally striate, although not as strongly as that at sides .
genus A (p. 169)
65 (63) Stigmal vein very short, not as long as marginal (Fig. 130) . . AGEN1ASPIS (p. 171)
Stigmal vein very long and curved towards anterior margin of forewing or longer than
marginal (Figs 133, 135, also cf. Fig. 53) 66
66 (65) Legs completely yellow. Forewings hyaline. Mandibles tridentate . MICROTERYS (p. 213)
Legs not completely yellow. Forewings often partially infuscate. Mandibles not
tridentate 67
67 (66) Marginal vein rather short, hardly twice as long as broad, costal cell dorsally with only a
few setae at apex (Fig. 135). Posterior ocellus separated from eye margin by at least
about twice its own diameter. Mandibles with one tooth and a broad truncation .
PRIONOMASTIX (p. 223)
Marginal vein at least about six times as long as broad, costal cell dorsally with a
complete line of setae extending from base to apex along wing margin (as in Fig. 53).
Posterior ocellus nearly touching eye margin. Mandibles bidentate .
ERICYDNUS(p. 195)
68 (61) Speculum interrupted in lower half by at least one line of setae on dorsal surface of wing
(Fig. 129). Notaular lines indicated anteriorly or complete, clava never apically
truncate.
Marginal vein punctiform. Thorax never shiny, if dark then matt. Mandibles
tridentate METAPHYCUS (p. 212)
Speculum not interrupted. Notaular lines absent except in Homalotylus where they are
more or less complete (as in Fig. 1) and the clava is apically obliquely truncate (as in
Fig. 123) . . 69
69 (68) Notaular lines more or less complete (as in Fig. 1). Clava apically obliquely truncate (as
in Fig. 123) HOMALOTYLUS (p. 203)
Notaular lines absent. Clava apically rounded 70
70 (69) Clava two-segmented 71
Clava solid . . . ... 73
71 (70) Mandibles with one tooth and a broad truncation (as in Fig. 91). Marginal vein
punctiform. Frontovertex broad, much broader than length of eye (eyes small).
Length of hind tarsus rather longer than corresponding hind tibia (Fig. 132) .
HUNTERELLUS (p. 204)
Mandibles tridentate or with two teeth and a truncation. Marginal vein longer than
broad. Frontovertex narrower than the length of an eye. Length of hind tarsus
shorter than length of corresponding hind tibia 72
NEOTROPICAL ENCYRTIDAE 163
72 (71) Scutellum smooth and shiny ZETETICONTUS (p. 236)
Scutellum with distinct sculpture.
Clava entire but appearing two-segmented in dry mounted specimens ... 73
73 (70, 72) Mandibles without teeth. Apex of scutellum usually with a tuft of long setae .
ENCYRTUS(p. 195)
Mandibles with three acute teeth or with one or two teeth and a truncation. Apex of
scutellum without tuft of setae 74
74 (73)* Mandibles with three acute teeth 75
Mandibles with one or two teeth and a truncation 81
75 (74) Marginal vein punctiform 76
Marginal vein at least twice as long as broad 78
76 (75) Thorax and head orange MUCRENCYRTUS (p. 213)
Thorax and head dark and metallic 77
77 (76) Dorsum of scutellum with deep longitudinally striate sculpture, contrasting with the
much more shallow, polygonal sculpture of mesoscutum. Scutellum not very shiny
(except at sides) and more or less dark purplish brown contrasting with the shiny
green mesoscutum. Spur of mid tibia shorter than corresponding basitarsus .
PARALITOMASTIX (p. 219)
Dorsum of scutellum not with longitudinally striate sculpture and often quite shiny,
occasionally purplish. Spur of mid tibia usually about as long or longer than
corresponding basitarsus COPIDOSOMA (p. 189)
78 (75) Scutellum very convex with deep, rough striate sculpture. Apex of scutellum with very
short, smooth, thin, leaf-like extension (as in Fig. 94) . . AMAURILYMA (p. 171)
If scutellum convex with deep striate sculpture then apex is without leaf-like extension 79
79 (78) Both mesoscutum and scutellum with very fine longitudinally striate-reticulate
sculpture, anterior two-thirds of mesoscutum with dark setae, posterior one-third
with pale setae ........... genus A (p. 169)
Never both mesoscutum and scutellum with sculpture as above, mesoscutum
occasionally with pale setae but these always evenly distributed .... 80
80 (79) Axillae distinctly raised above level of scutellum with posterior margin step-like, almost
vertical and mirror shiny, mesoscutum often with conspicuous white setae
APSILOPHRYS (p. 176)
Axillae hardly raised above level of scutellum and with posterior margin distinctly
sculptured, mirror never shiny. Setae on mesoscutum dark . . . . . 81
8 1 (74, 80) Marginal vein punctiform or only slightly longer than broad, usually as long or longer
than stigmal, postmarginal generally not distinct and therefore shorter than marginal
(as in Fig. 76). Posterior margin of mesoscutum produced backwards in middle third
so that axillae appear to be widely separated (as in Fig. 77). Parasites of eggs of
Lepidoptera, Hemiptera, etc OOENCYRTUS (p. 217)
Marginal vein usually at least twice as long as broad, stigmal and postmarginal veins
relatively long, or, if marginal punctiform then stigmal vein distinctly longer than
marginal. Posterior margin of mesoscutum gently curved or with either side straight
and centrally slightly angled so that axillae appear to meet or nearly meet centrally
and are not greatly separated by posterior margin of mesoscutum, or, if as above then
marginal vein is distinctly at least twice as long as broad 82
82 (81) Marginal vein punctiform and shorter than stigmal (as in Fig. 75). Funicle segments
often flattened and broadened, much wider than pedicel. Parasites of Psyllid nymphs
PSYLLAEPHAGUS (p. 226)
Marginal vein at least twice as long as broad and as long or longer than stigmal. Funicle
segments cylindrical 83
* Note added while this paper was in press. Boucekiella Hoffer and Platencyrtus Ferriere run here but differ from other
genera in having an extremely flattened leaf-like body. Boucekiella has the submarginal vein with a triangular expansion
which is absent in Platencyrtus.
164 JOHN S. NO YES
83 (82) Tegulae with at least base yellow or distinctly paler than apex (rarely completely dark).
Scutellum never with sculpture conspicuously deeper than that on mesoscutum
MICROTERYS(p 213)
Tegulae completely black or dark brown. Scutellum often with sculpture much deeper
than that on mesoscutum '""'.'• . . . . . ' . . . '. . 84
84 (83) At least outer face of scape distinctly paler (usually yellow) than flagellum (usually
brown). Parasites of Coccoidea . .... TRICHOMASTHUS (p. 234)
Scape always as dark or darker than flagellum. Parasites of Syrphidae
SYRPHOPHAGUS (p. 230)
85 (59) Wings shortened, not reaching apex of gaster 86
Wings reaching apex of gaster . . 87
86 (85) Body flattened. Mandibles bidentate. Wing rudiments hyaline . . RHOPUS (p. 226)
Body not flattened. Mandibles tridentate, the third (upper) tooth often truncate. Wing
rudiments sometimes darkened CHEILONEURUS (p. 184)
87 (85) Mandibles without teeth. Apex of scutellum usually with a tuft of long setae. Clava
entire. Stigmal vein longer than marginal, postmarginal as long or longer than stigmal
ENCYRTUS (p. 195)
Mandibles with from one to three acute teeth. Apex of scutellum usually without a tuft
of setae, if tuft present then stigmal vein not longer than marginal and postmarginal
shorter than stigmal (Gahaniella may have mandibles without teeth but clava two-
segmented) . 88
88 (87) Mandibles unidentate or bidentate 89
Mandibles tridentate, with one or two teeth and a truncation, or without teeth . . 100
89 (88) Epipygium covering less than half dorsal surface of gaster. Neither sixth funicle segment
nor clava with perpendicular sensilla. Speculum never interrupted .... 90
Epipygium covering at least about two-thirds dorsal surface of gaster (Fig. 1 34), sixth
funicle segment or clava usually with perpendicular sensilla (Fig. 131). Speculum
usually interrupted in lower half by at least one line of setae on dorsal surface of wing
(as in Figs 70, 71) (Rhopus may have shortened wings) . . . . . . 94
90 (89) Cerci in apical half of gaster GONZALEZIA (p. 197)
Cerci in basal half of gaster . 91
91 (90) Scutellum with tubercles or pits (as in Figs 98, 99) (these are best seen in cleared slide
mounted material) . . ... FORCIPESTRICIS (p. 196)
Scutellum without tubercles or pits 92
92 (91) Marginal vein punctiform or only very slightly longer than broad and postmarginal
absent. Area between stigmal vein and anterior margin of wing completely naked.
Propodeum short, medially at most one-eighth as long as scutellum
BRACHYPLATYCERUS (p. 183), HEMAENASWS (p. 202)
Marginal vein distinctly longer than broad, postmarginal and stigmal veins relatively
long. Area between stigmal vein and wing margin with several setae. Propodeum
medially at least one-sixth as long as scutellum 93
93 (92) Centre of propodeum completely smooth or with only very light sculpture and apex of
scutellum without distinct carina along its edge. Eyes with rather short inconspicuous
setae. Mandibles rather broad with two teeth of almost equal length. ASEIRBA (p. 179)
Centre of propodeum with distinct sculpture and apex of scutellum with carina along its
margin (Fig. 121) (often indistinct or absent in those species with a smooth
scutellum). Eyes usually with conspicuous setae. Mandibles narrow with teeth of
unequal length, the upper being long and the lower very short or absent
RHYTWOTHORAX (p. 226)
94 (89) Body dorso-ventrally flattened (wings sometimes shortened) . . . RHOPUS (p. 226)
Body not flattened (wings always fully developed) 95
95 (94) Frontovertex and mesoscutum with sculpture punctate, giving it a silky or granulate
appearance . ... . . 96
NEOTROPICAL ENCYRTIDAE 1 65
Frontovertex and mesoscutum with shallow reticulate sculpture, often quite shiny . 97
96 (95) Postmarginal vein longer than stigmal. Frontovertex with regular, raised, polygonal
sculpture GYRANUSOIDEA (p. 199)
Postmarginal vein short, much shorter than stigmal. Frontovertex with rather fine
raised, irregular, sculpture ANAGYRUS (p. 173)
97 (95) Postmarginal vein at least one and a half times as long as stigmal. Longest cilia on hind
wings at least half as long as maximum width of wing. Forewing usually with at least
indications of two or three dark fasciae .... LEPTOMASTIDEA (p. 206)
Postmarginal vein at most one and one third times as long as radial. Longest cilia on
hind wings usually much less than half as long as maximum width of wing. Forewing
completely hyaline 96
98 (97) Sculpture of scutellum deeper and coarser than on mesoscutum so that scutellum is
noticeably less shiny than mesoscutum .... APO ANAGYRUS (p. 176)
Sculpture on scutellum not distinctly deeper and coarser than on mesoscutum so that
scutellum is about as shiny as mesoscutum 99
99 (98) Postmarginal vein slightly shorter than stigmal . . . EXTROMATOPSIS (p. 194)
Postmarginal vein longer than stigmal LEPTOMAST1X (p. 206)
100 (88) Postmarginal vein longer than stigmal (Figs 130, 133, 175) 101
Postmarginal vein not longer than stigmal 106
101 (100) Head with punctate sculpture and dorsum of thorax with very fine semi-striate sculpture
which gives it a silky appearance. Speculum interrupted in its posterior half by a line
of setae on dorsal surface of wing ; first funicle segment hardly longer than pedicel,
funicle with setae not much longer than diameter of corresponding segment. Head
usually blue, thorax dorsally dark purplish brown . . . AGENIASPIS (p. 171)
Sculpture and coloration of head and thorax not as above. Speculum not interrupted.
First funicle segment usually at least about twice as long as pedicel, funicle with much
longer setae than above (except Microterys) 102
102 (101) Sculpture on scutellum elongate and arranged in a whorl-like pattern, distinctly deeper
than that on mesoscutum, apex of scutellum sometimes with a tuft of setae. Tegulae
brown 103
Sculpture on scutellum not elongate or arranged in whorls, if much deeper than that on
mesoscutum, then tegulae are yellow, apex of scutellum never with a tuft of setae . 104
103 (102) Frontovertex, particularly between ocelli and below anterior ocellus, with regular
punctate sculpture. Posterior one-third of mesoscutum very slightly concave and
often with pale setae in contrast to the anterior two-thirds which is convex and
usually with dark setae DIVERSINERVUS (p. 193)
Frontovertex with irregular, slightly transversely elongate sculpture. Mesoscutum
regularly convex and without pale setae ICEROMYIA (p. 204)
104 (102) Propodeum long, centrally at least one-sixth as long as scutellum and with sculpture
centrally (as in Fig. 60). Apex of scutellum often produced as translucent flange (if
scutellum convex then flange is absent or very indistinct). Eyes often very hairy.
Frontovertex and dorsum of thorax usually with thimble-punctured sculpture (often
sparse). Setae on dorsal side of funicle usually about twice as long as those ventrally.
Clava apparently two-segmented HEMENCYRTUS (p. 202)
Propodeum centrally at most one-seventh as long as scutellum and smooth. Apex of
scutellum never produced as a flange. Eyes naked or with short setae. Frontovertex
and thorax never with thimble-punctured sculpture. Setae on funicle the same length
dorsally as ventrally. Clava entire 105
105 (104) Antennal torulus separated from mouth margin by much more than its own length.
Head and thorax usually metallic green. Mandibles often with third upper tooth
truncate. Tegulae often yellow or orange MICROTERYS (p. 213)
Antennal torulus separated from mouth margin by less than its own length. Head and
thorax brown with some green, blue and purple reflections. Mandibles with three
acute teeth. Tegulae dark brown COCCIDOCTONUS (p. 188)
166 JOHN S. NO YES
106(100) Clava two-segmented . p$|< ....:» 107
Clava entire 109
107 (106) Scutellum smooth and shiny, except occasionally the basal two-thirds which may have
thimble-punctured sculpture but apical one-third always smooth and shiny
ZETETICONTUS (p. 236)
Scutellum with distinct sculpture, apical one-third never smooth or shiny . . .108
108 (107) Propodeum long, centrally at least one-sixth as long as scutellum and with distinct
sculpture centrally (as in Fig. 60). Apex of scutellum often produced as a translucent
flange. Eyes often very hairy. Frontovertex and dorsum of thorax usually with
thimble-punctured sculpture (often sparse). Mandibles tridentate ....
HEMENCYRTUS (p. 202)
Propodeum centrally at most one-seventh as long as scutellum and without sculpture.
Apex of scutellum without flange. Eyes naked or with very short setae. Frontovertex
and thorax usually with fine punctate sculpture, never thimble-punctured. Mandibles
without teeth or with one tooth and a truncation . . . GAHANIELLA (p. 197)
109 (106) Scutellum with tubercles or pits (as in Figs 98, 99) (best seen in cleared slide mounts).
Frontovertex usually smooth and shiny and with setae. Eyes hairy •;.!. f .
FORCIPESTRICIS (p. 196)
Scutellum without tubercles or pits 110
110 (109) At least head and usually dorsum of thorax with thimble-punctured sculpture. If
punctures indistinct then scutellum is flat and apically produced as a translucent
flange Ill
Head and dorsum of thorax without thimble-punctured sculpture and apex of scutellum
not produced as a flange. . . . . . . . . . . .113
111 (110) Marginal vein punctiform (as in Fig. 87). Thimble punctures on head and thorax deep
and distinct. Scutellum convex without any indication of apical translucent flange .
BOTHRIOTHORAX (p. 182)
Marginal vein much longer than broad. Thimble-punctures on head and thorax often
indistinct. Scutellum flat with distinct apical translucent flange, or convex with flange
very narrow and indistinct or absent 112
112 (111) Frontovertex with shiny-bottomed piliferous punctures. Mandibles with two acute teeth
and a truncation. Apex of scutellum without translucent flange
HELMECEPHALA (p. 200)
Punctures on frontovertex not shiny- bottomed. Mandibles with three acute teeth. Apex
of scutellum often produced as translucent flange (as in Fig. 60) ....
HEMENCYRTUS (p. 202)
113 (110) Funicle without whorls of long setae. Setae less than twice as along the diameter of
corresponding segment ' . .114
Funicle segments with whorls of long setae or setae which are at least twice as long as
diameter of corresponding segment . . 124
114(113) Marginal vein punctiform hardly longer than broad, postmarginal very short
or absent (as in Fig. 76) . . . . ,. . . OOENCYRTUS (p. 217)
Marginal vein at least about twice as long as broad, postmarginal distinct, occasionally
nearly as long as stigmal . .115
115 (1 14) Forewing darkened. Apex of scutellum with tuft of longer setae (sometimes indistinct or
lost) CHEILONEURUS (p 184)
Forewing hyaline, or, if darkened, then with a barely discernible longitudinal wedge-
shaped mark arising at apex (similar to Fig. 57). Apex of scutellum without tuft of
setae •-...-«•- . ' • . .116
116 (115) Submarginal vein of forewing with a triangular expansion in its apical one-third (Fig.
138) (sometimes indistinct but always indicated by a conspicuous, long, erect seta).
Forewing often with an indistinct longitudinal wedge-shaped fuscous streak arising at
apex (similar to Fig. 57) PARECHTHRODRYINUS (p. 221)
Submarginal vein of forewing without a triangular expansion, forewing without dark
streak 1 17
NEOTROPICAL ENCYRTIDAE 167
117 (116) Tegulae with at least base yellow or paler than apex (rarely completely dark). Scutellum
never with sculpture conspicuously deeper than mesoscutum . MICROTERYS (p. 213)
Tegulae completely black or dark brown. Scutellum often with conspicuously deeper
sculpture than mesoscutum 118
118 (117) Axillae with posterior face long and shiny. Propodeum centrally about one-sixth length
of scutellum and with distinct sculpture medially (Fig. 140). Mesoscutum metallic
green with purplish reflections anteriorly and covered with translucent hairs.
Scutellum at base with deep, rough, irregular sculpture and with apical one-third
highly polished (Fig. 140) . . . HELEGONATOPUS (p. 200)
Axillae more or less level with scutellum, not with posterior face long and shiny.
Propodeum shorter and smooth centrally. Mesoscutum not as above. Scutellum
occasionally rough at base and shiny at apex (some Trichomasthus and
Echthrodryinus) but usually with similar sculpture throughout 119
119 (118) Outer face of scape usually much paler (normally yellow) than flagellum (normally
brown). Parasites of Coccoidea TRICHOMASTHUS (p. 234)
Scape always about as dark or darker than flagellum. Not parasites of Coccoidea . 120
120 (1 19) Axillae not touching apically, always separated by posterior margin of mesoscutum . 121
Axillae more or less touching apically 123
121 (120) Scutellum abruptly convex at sides and at apex, dorsally almost flat, never with punctate
sculpture, usually with uniform sculpture. Scutellum (without axillae) at least slightly
longer than broad SYRPHOPHAGUS (p. 230)
Scutellum uniformly convex, or flat dorsally, often with deep punctate sculpture with
apical third of half smooth and shiny. Scutellum (without axillae) at most as long as
broad 122
122(121) Postmarginal vein long and distinct, half to nearly as long as stigmal .
ECHTHRODRYINUS (p. 193)
Postmarginal vein very short or absent (as in Fig. 76) . . . OOENCYRTUS (p. 217)
123 (120) Body usually not longer than 1 mm. Parasites of Aphidoidea APHIDENCYRTUS (p. 175)
Body usually longer than 1 mm. Parasites of Syrphidae . SYRPHOPHAGUS (p. 230)
124 (113) Scutellum very convex and, at base, with several distinct longitudinal ridges running
backwards from axillae (as in Fig. 74). Setae on dorsal side of funicle several times
longer than those ventrally (Fig. 179) LIRENCYRTUS (p. 207)
Scutellum occasionally very convex but never with longitudinal ridges at base. Setae on
funicle usually about the same length dorsally as ventrally 125
125 (124)* Propodeum centrally at least one-fifth as long as scutellum and with at least one
longitudinal carina. Submarginal vein with triangular expansion in its apical one-
third (Fig. 138) (often indistinct), forewing occasionally with fuscous wedge-shaped
streak from apex to near centre (similar to Fig. 57). PARECHTHRODRY1NUS (p. 221)
Propodeum shorter than one-sixth length of scutellum or, if longer then there are no
distinct carinae. Submarginal vein usually without a triangular expansion, forewing
never with fuscous wedge-shaped streak 126
126 (125) Mandibles with three acute teeth 127
Mandibles with one or two teeth and a truncation 137
127 (126) First funicle segment at most as long as pedicel 128
First funicle segment at least one and a half times as long as pedicel . . . .131
128(127) Scutellum flat and polished. Body slightly dorso-ventrally flattened .... 129
Either scutellum distinctly convex or flat with distinct sculpture. Body robust . . 130
129 (128) Stigmal vein as long as marginal and with stigma hardly wider than vein (Fig. 139).
Head with strong sculpture which is distinct on genae . . TANYENCYRTUS (p. 231)
* Note added while this paper was in press. Cerapterocerus Westwood runs here but the Submarginal vein of the forewing
has a distinct subapical triangular expansion; it differs from Parechthrodryinus in having hyaline forewings and the
scutellum with a thin apical flange which overhangs the propodeum centrally for over half its median length
(Parechthrodryinus has no distinct apical flange on the scutellum).
1 68 JOHN S. NO YES
Stigmal vein shorter than marginal and with stigma enlarged, at least two and a half
times as wide as stigmal vein at base (Fig. 137). Head with very weak sculpture so that
genae are very smooth and shiny SHENAHET1A (p. 227)
130 (128) Scape at least three and a half times as long as broad . MERCETENCYRTUS (p. 21 1)
Scape at most three times as long as broad . * -.•..;,„,'< [COCCIDENCYRTUS] (p. 188)
131(127) Forewing darkened . .... . . . CHEILONEURUS (p. 184)
Forewing hyaline 132
132 (131) Head triangular in side view, the junction of the frontovertex and face forming an acute
angle. Head orange, contrasting with dark brown thorax . . ANICETUS (p. 174)
Head not triangular in side view, more or less gradually rounded. Head not contrasting
so strongly with thorax ' 133
133(132) Scutellum longitudinally striate with silky lustre . . MERCETENCYRTUS (p. 211)
Scutellum not longitudinally striate with silky lustre . " '., "_ 134
134(133) Thorax and head dark and metallic. Mesoscutum with dark setae . . . . 135
If thorax and head are dark and metallic then setae on mesoscutum are mostly white 136
135 (134) Scutellum, without axillae, slightly longer than wide or about as long as wide. Hind
femur and tibia yellow, mid-tibial spur not shorter than corresponding basitarsus .
ICEROMYIA (p. 204)
Scutellum, without axillae, slightly wider than long. Hind femur and tibia mostly brown,
mid tibial spur shorter than corresponding basitarsus AMMONOENCYRTUS (p. 172)
136 (134) Thorax and head dark and metallic. Mesoscutum with contrasting pale setae
CHEILONEURUS (p. 184), PROCHILONEURUS (p. 223)
Thorax and head orange, with darker areas not dark and metallic . ...
CHEILONEURUS (p. 184)
137 (126) Body dorso-ventrally flattened. Scutellum flat, with punctate sculpture centrally and
with a shiny, rounded border about as wide as one-quarter length of scutellum.
Speculum often interrupted on dorsal surface of wing . . COMPERIELLA (p. 189)
Body not dorso-ventrally flattened. Scutellum occasionally flat with punctate sculpture
but border not smooth and shiny or, if so then sides are flat and not rounded.
Speculum not interrupted . . " f . „ 138
138 (137) Gaster only slightly longer than scutellum. First funicle segment at most only slightly
longer than pedicel. Thorax not metallic, mesoscutum with dark setae. Mandibles
with one tooth and a broad truncation .... APHYCOMORPHA (p. 175)
Gaster at least one and a half times as long as scutellum. If first funicle segment is not at
least twice as long as pedicel then mesoscutum is dark with at least a few white setae
or the mandibles have two acute teeth with the third (upper) tooth truncate . . 139
139 (138) First funicle segment at most as long as pedicel ._- 140
First funicle segment longer than pedicel . . , . . . . . .141
140 (139) Scape at least three and a half times as long as broad . . SYRPHOPHAGUS (p. 230)
Scape' at most three times as long as broad . . . [COCCIDENCYRTUS] (p. 188)
141 (139) Postmarginal vein absent or short, at most half as long as stigmal (as in Fig. 110).
Mesoscutum dark with dark setae (may be orange in species not examined).
Scutellum flat , . ' . . . .142
Postmarginal vein nearly as long or as long as stigmal. If mesoscutum dark then with at
least a few contrasting white setae or scutellum convex . . :.. . . . . 144
142 (141) Scutellum with steep, very flat, polished sides . . NEOCOCCIDENCYRTUS (p. 216)
Sides of scutellum short and curved, occasionally shiny 143
143 (142) Scutellum with apex not shiny, never metallic green, legs usually yellow with one or two
dark bands COCCIDENCYRTUS (p. 188)
Scutellum with extreme apex metallic green, legs extensively darkened, never yellow
with dark bands . GRISSELLIA (p. 197)
NEOTROPICAL ENCYRTIDAE 169
144 (Hi) Forewing darkened or, if hyaline then thorax mostly orange . CHE1LONEURUS (p. 184)
Forewing hyaline and thorax completely dark 145
145 (144) Setae on funicle not arranged in distinct whorls. Scutellum very convex with base
sculptured and apical third shiny. Mesoscutum with numerous dark setae
ECHTHRODRYINUS (p. 193)
Setae on funicle normally arranged in two or three distinct whorls per segment.
Scutellum usually fairly flat, never very convex, and with uniform sculpture
throughout or with sides and apex shiny. Mesoscutum usually with at least some
appressed white setae 146
146 (145) Submarginal vein distinctly expanded in its apical two-thirds . . ZAOMMA (p. 235)
Submarginal vein not expanded in its apical two-thirds. . CHEILONEURUS (p. 184)
Notes on genera
Genus A
(Key couplets: ? 56; J1 64, 79)
This genus is being formally described in Acta Zoologica Lilloana by P. Fidalgo from San Miguel
de Tucuman, Argentina. It is close to Cheiloneurus, Diversinervus and Anicetus, from which it can
be separated using the characters given in the key.
ACEROPHAGOIDES Blanchard
(Key couplets: I1 27; j1 12. Fig. 20)
Acerophagoides Blanchard, 1940: 106. Type- species : Acerophagoides triangularis Blanchard, by monotypy.
INCLUDED SPECIES. One: triangularis Blanchard (De Santis, 1964: 104); also one unidentified
species from Trinidad (BMNH).
DISTRIBUTION. Argentina, Trinidad.
BIOLOGY. Reared from Eriococcidae (Homoptera).
COMMENTS. Compere & Annecke (1960: 381) correctly placed the genus near Pauridia
Timberlake (tribe Pauridiini), from which it differs in the number of funicle segments and
dentition of mandibles (the third tooth of Acerophagoides being nearer the apex of the mandible
than in Pauridia). Also examined is a single specimen from Trinidad (BMNH) with only four
funicle segments.
ACEROPHAGUS Smith
(Key couplets : ? 19; ^24)
Acerophagus Smith, 1880: 83. Type-species: Acerophagus coccois Smith, by monotypy.
Rhopoideus Howard, 18986: 235. Type-species: Rhopoideus citrinus Howard, by monotypy.
INCLUDED SPECIES. Fourteen, eight from the Neotropics : coccois Smith, debilis Timberlake, flavus
Rosen, luteolus Rosen, notativentris (Girault), nubilipennis Dozier, pallidus Timberlake and
texanus (Howard).
REFERENCE. World revision: Rosen (1969).
DISTRIBUTION. Mexico, Trinidad, Jamaica, Puerto Rico. World: New World.
BIOLOGY. Parasites of Pseudococcidae (Homoptera).
COMMENTS. Very close to Pseudaphycus and Timberlakia (tribe Aphycini, subtribe Aphycina)
from which it can be separated using the characters given in the key.
170 JOHN S. NOYES
ACROASPIDIA Compere & Zinna
(Key couplet: \ 34. Fig. 17)
Acroaspidia Compere & Zinna, 1955: 96. Type-species: Acroaspidia myrmicoides Compere & Zinna, by
monotypy.
INCLUDED SPECIES. One: myrmicoides Compere & Zinna (1955: 98).
DISTRIBUTION. Trinidad.
BIOLOGY. Parasite of Pseudococcidae (Homoptera).
COMMENTS. Trjapitzin (1973a) places the genus in the tribe Acroaspidiini which includes
Aeptencyrtus and Pelmatencyrtus. Kerrich's (1978) interpretation is broader and places it in the
subtribe Dinocarsiina which also includes Zarhopalus, Chrysoplatycerus, Hambletonia and
Zaplatycerus. It can be easily separated from these genera and also from Cheiloneurus, which it
superficially resembles, using the characters given in the key.
ADELENCYRTUS Ashmead
(Key couplets: ? 88, 243; 3 1. Fig. 46)
Adelencyrtus Ashmead, 1900: 401. Type-species: Encyrtus chionaspidis Howard, by original designation.
Epiencyrtoides Girault, \9l5d: 108. Type-species: Epiencyrtoides quadridentatus Girault, by original
designation.
Rotrencyrtus Risbec, 1958: 39. Types-species: Rotrencyrtus depressus Risbec, by monotypy.
INCLUDED SPECIES. Twenty-five, three from the Neotropics: aulacaspidis (Brethes), moderatus
(Howard) and odonaspidis Fullaway; also one unidentified species from Trinidad (BMNH).
REFERENCES. World key: Compere & Annecke (1961), see also Noyes (1979).
DISTRIBUTION. Throughout Neotropics. World: cosmopolitan.
BIOLOGY. Parasites of Diaspididae (Homoptera).
COMMENTS. Placed in the tribe Habrolepidini, subtribe Habrolepidina by Trjapitzin (1973ft)
which includes other genera which have quadridentate mandibles and, in the female, the head
triangular in side view. The female can be separated from the included genera using the
characters given in key. The males of this subtribe all have a two-segmented funicle and a long
unsegmented clava and are extremely difficult to separate. A key to genera included in this
subtribe is also given by Tachikawa & Valentine (1969).
AENASWS Walker
(Key couplets: 9 63, 144; 3 7, 38. Figs 62, 113)
Aenasius Walker, 1846: 181. Type-species: Encyrtus hyettus Walker, by monotypy.
INCLUDED SPECIES. Twenty-seven, 26 being found in the Neotropics : acuminatus Kerrich, advena
Compere, bolowi Mercet, brasiliensis (Mercet), brethesi De Santis, caeruleus Brues, cariocus
Compere, chapadae Ashmead, connectens Kerrich, flandersi Kerrich, frontalis Compere, hyettus
Walker, insularis Compere, longiscapus Compere, maplei Compere, masii Domenichini, pacificus
Compere, paulistus Compere, personatus Kerrich, phenacocci Bennett, punctatus Compere,
regularis Kerrich, similis Kerrich, tachigaliae (Brues), vadosus Kerrich and vexans Kerrich ; also a
further seven or eight undescribed species (BMNH).
REFERENCE. Revision: Kerrich (1967).
BIOLOGY. Parasites of Pseudococcidae (Homoptera).
DISTRIBUTION. Throughout the Neotropics. World: Nearctic region, Africa, India.
NEOTROPICAL ENCYRTIDAE 171
COMMENTS. Kerrich (1967) did not deal with Aenasius bolowi Mercet since at that time the type
had not been located. Since then the type has been found in Mercet's collection and is being dealt
with in a paper on Mercet's encyrtid types by the present author. The species is very close to
regular is and similis, but differs in the scape bulging outwards at the apex, the top of the antennal
scrobes lacking a keel or carina and the outer margin of the forewing being slightly concave.
Placed in the tribe Rhinoencyrtini, subtribe Aenasiina which also includes Chalcaspis, from
which it can be separated using the characters given in the key.
AEPTENCYRTUS De Santis
(Key couplets: ? 32; j1 37)
Aeptencyrtus De Santis, 1964: 119. Type-species: Pheidoloxenus bruchi De Santis, by monotypy.
INCLUDED SPECIES. One: bruchi (De Santis, 1964: 119).
DISTRIBUTION. Argentina, Trinidad, St Vincent.
BIOLOGY. Parasites of Pseudococcidae (Homoptera).
COMMENTS. See under Acroaspidia.
AGENIASPIS Dahlbom
(Key couplets: ? 219; j1 65, 101. Fig. 130)
Ageniaspis Dahlbom, 1857: 293. Type-species: Encyrtus fuscicollis Dalman, by designation of Ashmead
(1904: 303).
INCLUDED SPECIES. Six, one from the Neotropics : fuscicollis (Dalman, 1820: 359; De Santis,
1964: 312).
DISTRIBUTION. Argentina. World : cosmopolitan.
BIOLOGY. Polyembryonic parasites of larvae of Yponomeutidae (Lepidoptera).
COMMENTS. Placed in the tribe Copidosomatini, subtribe Ageniaspidiina which also includes
Paraleurocerus and Holcothorax Mayr. It can be separated from Holcothorax (not known from
South America) in having a six-segmented funicle (Holcothorax has five) and from
Paraleurocerus by the characters given in the key. It is very close to Paraleurocerus and may
eventually be considered synonymous.
AMAURILYMA Graham
(Key couplets: ? 190, 253; $ 78. Figs 94, 106, 156-158)
Amaurilyma Graham, 1958: 153. Type-species: Encyrtus brevicornis Dalman, by monotypy.
INCLUDED SPECIES. Three, none of which is found in the Neotropics, but at least one
undetermined species from South America (BMNH; CNC, Ottawa; USNM, Washington).
DISTRIBUTION. Trinidad, Jamaica, Brazil. World: Palaearctic and Ethiopian regions.
BIOLOGY. Unknown.
COMMENTS. The material from each of the three countries mentioned above may belong to three
distinct species since there are differences in the proportions of the antennal segments, sculpture
of the propodeum and colour of the legs. I have also examined one specimen, which obviously
belongs to this genus, from Zimbabwe (Rhodesia).
I am unable to place this genus according to Trjapitzin's (\913b) classification of the
Encyrtidae. However, the genus does seem to be closely related to Desobius, but can be separated
using the characters given in the key.
172 JOHN s- NOYES
AMEROMYZOBIA Girault
(Key couplets: j> 15; ; 22. Figs 14, 117)
Ameromyzobia Girault, 19166: 232. Type-species: Ameromyzobia aphelinoides Girault, by monotypy.
INCLUDED SPECIES. Two: aphelinoides Girault, 19166: 233 and bulyginskayae Trjapitzin, 1971c:
290; also one, probably undescribed, species from Brazil (BMNH).
DISTRIBUTION. St Vincent, Trinidad, Cuba. World: U.S.A., Florida (one undetermined species,
BMNH).
BIOLOGY. Reared from Diaspis sp. (Homoptera: Diaspididae) on grass in Trinidad (BMNH).
COMMENTS. Both of the described species and that found in Florida are very close and in fact may
be a single species. The genus appears to be related to those included in the tribe Aphycini,
subtribe Paraphycina, e.g. Metaphycus.
AMMONOENCYRTUS De Santis
(Key couplets: ? 49; $ 135. Fig. 22)
Ammonoencyrtus De Santis, 1964: 376. Type-species: Cerapterocerus bonariensis Brethes, by monotypy.
INCLUDED SPECIES. Two, one from the Neotropics: bonariensis (Brethes; De Santis, 1964: 378);
also one undescribed species from Trinidad (BMNH) which appears to differ from both
described species in the form of the antennae.
REFERENCE. Revision: Annecke (1967: 159-160).
DISTRIBUTION. Argentina, Brazil, Chile, Trinidad. World: Nearctic region.
BIOLOGY. Parasites of Coccidae (Homoptera).
COMMENTS. Placed in the tribe Cerapterocerini which includes Anicetus and Homosemion; it can
be separated from these genera using the characters given in the key. Annecke (1967) includes a
key to separate all other genera included in this tribe as well as a few superficially similar genera.
ANABROLEPIS Timberlake
(Key couplets: ? 59, 123; ^ 7. Figs 33, 111)
Anabrolepis Timberlake, 19206: 431. Type-species: Anabrolepis extranea Timberlake, by original
designation.
INCLUDED SPECIES. Five, one species (? zetterstedti (Westwood)) apparently recorded from
Bermuda (F. D. Bennett, in ////.)•
REFERENCE. World review: Tachikawa (1955).
DISTRIBUTION. Bermuda. World: Nearctic region, Palaearctic region, Pacific Islands, North
Africa.
BIOLOGY. Parasites of Diaspididae (Homoptera).
COMMENTS. I have not seen any material of this genus from South America and it may be that the
record from Bermuda is actually of an Adelencyrtus. It is possible that Anabrolepis does occur in
the Neotropical region and for that reason I have left it in the key.
The genus is included in the tribe Habrolepidini, subtribe Habrolepidina and can be separated
from other included genera by characters given in the key and also in a key given by Tachikawa
& Valentine (1969).
NEOTROPICAL ENCYRTIDAE 173
ANAGYRUS Howard
(Key couplets: 9 44, 52, 70, 114, 148, 158; 3 96. Figs 40, 70, 71, 131, 134)
Anagyrus Howard in Howard & Ashmead, 1896: 638. Type-species: Anagyrus greeni Howard, by
monotypy.
Heterarthrellus Howard, 18986: 239. Type-species: Heterarthrellus australiensis Howard, by monotypy.
Epidinocarsis Girault, 19136: 83. Type-species: Epidinocarsis tricolor Girault, by monotypy.
Paranusia Brethes, 1913: 102. Type-species: Paranusia bifasciata Brethes, by monotypy. Syn. n.
Philoponectroma Brethes, 1913: 104. Type-species: Philoponectroma pectinatum Brethes, by monotypy.
Syn. n.
Gyranusia Brethes, 1920: 137. Type-species: Gyranusia porteri Brethes, by monotypy. Syn. n.
Gyranusa Mercet, 1921 : 123. Type-species: Gyranusa matritensis Mercet, by original designation.
Protanagyrus Blanchard, 1940: 115. Type-species: Protanagyrus aciculatus Blanchard, by monotypy.
Xiphomastix De Santis, 19726: 45. Type-species: Xiphomastix bellator De Santis, by original designation.
INCLUDED SPECIES. One hundred and thirty-two, 19 from the Neotropics: aciculatus (Blanchard),
ananaitis Gahan, antoninae Timberlake, 1920: 409, bellator (De Santis, 19726: 46), bifasciatus
(Brethes) (comb, n.), brevistigma De Santis, chilensis Brethes, 19160: 8, coccidivorus Dozier,
1932: 8, diver sicornis Mercet, 1921: 134, graminicolens Dozier, 1937: 123, insolitus (Howard;
Noyes, 1979: \45),jucundus De Santis, nigriceps (De Santis, 19726: 46), porteri (Brethes) (comb,
n.), pseudococci (Girault, 19156: 185), pulchricornis (Howard; Noyes, 1979: 146), saccharicola
Timberlake, 1932: 159, tanystis De Santis, terebratus (Howard; Noyes, 1979: 87); also at least a
further 10 undescribed species from Trinidad, Brazil, Panama and Colombia (BMNH; CNC,
Ottawa).
REFERENCES. South America: De Santis (1964). World: Peck (1963: 335).
DISTRIBUTION. Throughout the Neotropics. World: cosmopolitan.
BIOLOGY. Parasites of Pseudococcidae (Homoptera). Some species in Australia are parasites on
coccinellid larvae (Coleoptera) which produce a waxy secretion and are predatory on mealybugs,
e.g. Anagyrus australiensis (Howard).
COMMENTS. I have examined specimens (syntypes?) of Paranusia bifasciata Brethes and these
belong to the group of species previously placed under Gyranusa (which is considered by some
authors to be a good genus). I have not examined the types of Philoponectroma pectinatum
Brethes or Gyranusia porteri Brethes but since De Santis (1950: 54, 60) synonymized the genera
with Paranusia, they are here treated as synonyms of Anagyrus.
Two species, nigriceps and terebratus, and a further species from Panama (CNC, Ottawa)
(which runs in the 9 key to Apoanagyrus, couplet 1 59), belong to a group with a smooth, shiny,
frontovertex. According to present understanding of the group of genera to which Anagyrus
belongs (subtribe Anagyrina), these three species could be considered as belonging to an
undescribed genus. I am not convinced of this and therefore a new genus is not described here to
accommodate them. Indeed, the generic limits of the genera belonging to this group, e.g.
Anagyrus, Anathrix, Apoanagyrus, Gyranusoidea, Leptomastix, Pseudleptomastix, etc., are in
some cases very difficult to define and it may be possible that further study and new material may
show that some ought to be considered synonymous.
ANARHOPUS Timberlake
(Key couplets: 9 10; J 3. Figs 10, 112)
Anarhopus Timberlake, 1929: 15. Type-species: Anarhopus sydneyensis Timberlake, by monotypy.
INCLUDED SPECIES. One: sydneyensis Timberlake (1929: 18).
DISTRIBUTION. Trinidad. World: Nearctic region, Australia.
BIOLOGY. Parasites of Pseudococcidae (Homoptera).
174 JOHN S. NO YES
COMMENTS. I have not seen any South American specimens of this genus but, according to De
Santis (in ////.), it has been found (?introduced) in Trinidad.
Placed in the tribe Tetracnemini, subtribe Arhopoideina, which also includes Tetracnemoidea
from which it can be separated using the characters in the key.
ANATHR1X Burks
(Key couplets:? 114, 131, 159)
Anathrix Burks, 1952: 179. Type-species: Anathrix argyrus Burks, by monotypy.
Aglyptoideus De Santis, 1964: 99. Ty pe- species : Aglyptoideus rusticus De Santis, by monotypy. Syn. n.
INCLUDED SPECIES. Four, one from the Neotropics: rustica (De Santis, 1964: 101) (comb. n.).
REFERENCE. Review: Myartseva (\911a).
DISTRIBUTION. Argentina. World: Nearctic and Palaearctic regions.
BIOLOGY. Parasites of Pseudococcidae (Homoptera).
COMMENTS. Placed in the tribe Anagyrini, subtribe Anagyrina. See comments under Anagyrus.
ANDINOENCYRTUS Blanchard
(Key couplet: ? 222)
Andinoencyrtus Blanchard, 1940: 106. Type-species: Andinoencyrtus ocellatus Blanchard, by monotypy.
INCLUDED SPECIES. One: ocellatus Blanchard (De Santis, 1964: 276).
DISTRIBUTION. Argentina.
BIOLOGY. Parasites of Eriococcidae (Homoptera).
COMMENTS. I have examined the syntypes of this species. They are mounted under a single
coverslip on a slide and are very badly crushed and distorted, but it is apparent that De Santis'
(1964) redescription of the genus is slightly incorrect in that the marginal vein is actually slightly
longer than broad and not as transverse as in his figure 135. It is probable that the cheeks are not
subparallel but of more normal form, even though the cheeks of one syntype appear to be more
or less parallel ; this is very probably due to the head being distorted when it was squashed by the
coverslip.
Andinoencyrtus is the sole genus included in the tribe Andinoencyrtini.
ANICETUS Howard
(Key couplets: 9 46; 3 132. Fig. 30)
Anicetus Howard in Howard & Ashmead, 1896: 639. Type-species: Anicetus ceylonensis Howard, by
monotypy.
Asteropaeus Howard, 1 898£ : 23 1 . Type-species : Asteropaeus primus Howard, by monotypy.
Habrolepopterygis Girault, \9\5d: 86. Type-species: Habrolepopterygis felix Girault, by monotypy.
Krishnieriella Mani, 1935: 421. Type-species: Krishnieriella ceroplastodis Mani, by monotypy.
INCLUDED SPECIES. Twenty-six, three from the Neotropics: annulatus Timberlake, primus
(Howard) and quintanai De Santis.
REFERENCE. World revision: Annecke (1967: 105-130).
DISTRIBUTION. Throughout the Neotropics. World : cosmopolitan, except more northern latitudes
(40° + ).
BIOLOGY. Parasites of Coccidae (Homoptera).
COMMENTS. Placed in the tribe Cerapterocerini. See comments under Ammonoencyrtus.
NEOTROPICAL ENCYRTIDAE 175
ANUSIOPTERA Brues
(Key couplet: $ 121. Fig. 66)
Anusioptera Brues, 1910: 83. Type-species: Anusioptera aureocincta Brues, by monotypy.
INCLUDED SPECIES. One, aureocincta Brues, 1910: 83.
DISTRIBUTION. Mexico.
BIOLOGY. Probably parasitic on Pseudococcidae (Homoptera).
COMMENTS. The holotype female of aureocincta is in good condition except that it is lacking the
left antenna and right clava.
The genus very probably belongs to the tribe Anagyrini, subtribe Leptomastideina.
APHIDENCYRTUS Ashmead
(Key couplets: 9 259; $ 123)
Aphidencyrtus Ashmead, 1900: 398. Type-species: Encyrtus aphidiphagus Ashmead, by original
designation.
INCLUDED SPECIES. Twenty-eight, two from the Neotropics: aphidivorus (Mayr, 1876: 712, 713 &
724) and brasiliensis Ashmead (1904a: 497); also one unidentified species from St Vincent
(BMNH).
REFERENCE. Key to Palaearctic species: Hoffer (1970: 78-79).
DISTRIBUTION. Argentina, Brazil, Chile, Peru, Puerto Rico, St Vincent. World: cosmopolitan.
BIOLOGY. Parasites of Aphididae (secondary or primary) and in other parts of the world of
Psyllidae (Homoptera).
COMMENTS. I have not seen the type of brasiliensis but from the description it is very probably
incorrectly placed in this genus. Psyllaephagus trellesi may be an Aphidencyrtus, but the wing
venation and colour of the mesoscutum point to it being a Psyllaephagus. The types of this
species are badly mounted on slides.
Placed in the tribe Microteryini, subtribe Syrphophagina, which also includes Syrphophagus
Ashmead. The two genera are extremely close and sometimes impossible to separate unless the
biology is known. In general specimens of Aphidencyrtus are smaller than Syrphophagus, but this
is only a reflection of the size of the host. Possibly the two genera ought to be synonymized.
APHYCOMASTIX De Sands
(Key couplet: ? 12)
Aphycomastix De Santis, 19726: 49. Type-species: Aphycomastix annulata De Santis, by monotypy.
INCLUDED SPECIES. One: annulata De Santis, 19726: 49.
DISTRIBUTION. Argentina, Brazil.
BIOLOGY. Unknown.
COMMENTS. I am unable to place this genus according to Trjapitzin's classification except to say
that it belongs to the Encyrtinae, possibly the tribe Aphycini, and may be close to Cyderius.
APHYCOMORPHA Timberlake
(Key couplets: ? 106, 135, 210; £ 138)
Aphycomorpha Timberlake, 19196: 225. Type-species: Aphycomorpha araucariae Timberlake, by
monotypy.
176 JOHN S. NOYES
INCLUDED SPECIES. Two, one from the Neotropics: araucariae Timberlake, 19196: 227.
DISTRIBUTION. Mexico. World: Hawaii, New Zealand.
BIOLOGY. Parasites of Diaspididae and Eriococcidae (Homoptera).
COMMENTS. Placed in the tribe Aphycini by Trjapitzin (19736).
APOANAGYRVS Compere
(Key couplets: 9 69, 159; ^ 98)
Apoanagyrus Compere, 1947: 18. Type-species: Apoanagyrus californicus Compere, by monotypy.
INCLUDED SPECIES. Seven, five of which occur in the Neotropics : diversicornis (Howard ; Noyes,
1979: 148), lopezi De Santis, 1964: 74, malenotus (De Santis, 19726: 44) (comb. n. from
Leptomastix), montivagus (De Santis, 1964: 86) (comb. n. from Leptomastix) and trinidadensis
Kerrich, 1953: 792, also two further species being described by Dr G. J. Kerrich in a paper in
preparation.
DISTRIBUTION. Throughout the Neotropics. World: Nearctic and Palaearctic regions.
BIOLOGY. Parasites of Pseudococcidae (Homoptera).
COMMENTS. It is possible that both malenotus and montivagus are merely colour forms of
diversicornis.
Placed in the subtribe Anagyrina. See comments under Anagyrus.
APSILOPHRYS De Santis
(Key couplets: ? 155; 3 80. Fig. 65)
Apsilophrys De Santis, 1964: 296. Type-species: Copidosoma oeceticola De Santis, by monotypy.
Copidencyrtus De Santis, 1964: 304. Type-species: Copidencyrtus gracilis De Santis, by monotypy. Syn. n.
INCLUDED SPECIES. Three, all from the Neotropics: capsicum (Burks, 1967: 54) (comb. n. from
Copidosoma), gracilis (De Santis, 1964: 305) (comb, n.) and oeceticola De Santis (1964: 296); in
addition two further unidentified species from Peru and Brazil (BMNH).
DISTRIBUTION. Argentina, Brazil, Peru, Trinidad, Puerto Rico.
BIOLOGY. Polyembryonic parasites of larvae of Gelechiidae (Lepidoptera).
COMMENTS. I have examined the holotype of Copidencyrtus gracilis; it is a male and belongs to
Apsilophrys. I have not seen the holotype of Copidosoma capsicum but specimens which fit the
description of the species and are reared from the same host and locality as the holotype
undoubtedly belong to Apsilophrys.
Apsilophrys is very close to Copidosoma (Copidosomatini) but can be separated using the
characters given in the key.
ARCHINUS Howard
(Key couplets : ? 17; <$ 20)
Archinus Howard, 1897: 154. Type-species: Archinus occupatus Howard, by monotypy.
INCLUDED SPECIES. One: occupatus Howard (Noyes, 1979: 149).
DISTRIBUTION. Grenada, Trinidad.
BIOLOGY. Unknown.
COMMENTS. See Noyes (1979: 148-149).
NEOTROPICAL ENCYRTIDAE 177
ARHOPOIDIELLA gen. n.
(Key couplets: ? 18; 3 21. Figs 143-148)
Type-species: Arhopoidiella carinata sp. n.
?. Head. In side view, moderately deep and most rounded at level of top of antennal scrobes, sometimes
abruptly angled at this point, frontovertex slightly convex. Eyes slightly longer than broad, with
conspicuous pubescence, overreaching occipital margin which is more or less sharp. Malar space about half
length of eye with malar sulcus present. Frontovertex about one-third head width; posterior ocellus
separated from occipital margin by slightly more than, to twice as much as its diameter and from eye
margin by about half its diameter, ocelli forming an acute angle. Antennal scrobes deep, meeting dorsally
and reaching about two-thirds from antennal toruli to anterior ocellus, top of scrobes sometimes with a
transverse carina which almost reaches inner eye margins ; antennal torulus separated from mouth margin
by about half its length and from other torulus by about one and a half times its length, top of torulus
below level of lower eye margin ; scape subcylindrical and stout, pedicel conical and longer than first funicle
segment, funicle five-segmented, all segments longer than broad and cylindrical, clava three-segmented,
pointed apically and about as wide as funicle, longitudinal sensilla on all flagellar segments. Frontovertex
with coriaceous or punctate sculpture, the latter giving it a granulate appearance, below top of antennal
scrobes more shallow and elongate and almost alutaceous on genae. Frontovertex with a few short setae,
those on interantennal prominence longer. Mandibles with three acute teeth, the middle tooth slightly
longest, maxillary palpi three-segmented, labial palpi three-segmented.
Thorax. In side view, moderately deep and dorsally quite flat. Mesoscutum without notaular lines, about
a half broader than long, axillae slightly separated by posterior margin of mesoscutum, scutellum about as
long as broad and with apex pointed, propodeum medially short, less than one-eighth as long as scutellum.
Pronotum and mesoscutum with shallow alutaceous sculpture, which is slightly deeper on scutellum where
it is centrally of smaller mesh, mesopleura with similar sculpture, but more elongate anteriorly and tending
towards punctate posteriorly, propodeum with irregular very shallow sculpture. Forewing hyaline, slightly
more than three times as long as broad, basal cell sparsely hairy and completely naked in proximal one-
third, speculum not interrupted and open ; submarginal vein slightly expanded towards apex and with an
apical hyaline break, marginal vein about twice as long as broad, stigmal vein about twice as long as
marginal or post-marginal veins. Hindwing about three and a half times as long as broad. Mid tibial spur
slightly shorter than corresponding basal tarsal segment.
Caster. Slightly shorter than thorax, cerci in basal one-third, apex of hypopygium reaching to about two-
thirds along gaster or to apex and slightly incised, paratergites absent. Ovipositor protruding, exserted part
equal to about one-quarter to one-third length of gaster or from one-quarter to one-half as long as mid
tibia; ovipositor about twice as long as mid tibia, gonostyli slightly longer than half length of mid tibia.
3. Similar to female except clava entire. Aedeagus about two-thirds as long as mid tibia and each digitus
with one thick spine apically.
Arhopoidiella carinata sp. n.
(Figs 143-148)
?. Length (excluding ovipositor) 1-00-1-27 mm (holotype 1-06 mm).
Colour. Body generally black with weak metallic green reflections, frontovertex slightly paler; antennae
testaceous to dark brown except scape and pedicel which may be darker, occasionally terminal funicle
segments and clava slightly paler; legs dark brown except tarsi and apices of tibiae which are testaceous or
yellow, occasionally fore tibia and tarsus completely brown; ovipositor sheaths brown.
Head. In profile sharply angled at top of antennal scrobes which have a transverse carina extending
across width of frontovertex at this point (Fig. 146); posterior ocellus separated from occipital margin by
slightly more than its own diameter, ocelli forming an angle of about 60°. Sculpture of frontovertex
punctate so that it has a granulate appearance. Relative measurements of head of holotype: maximum head
width 62, width of frontovertex across median ocellus 16-5, malar space 20, eye length 39, eye width 34,
POL 7, OOL 2, length of scape 32, other proportions of antenna as in Figure 145.
Thorax. Relative measurements of forewing of holotype: length 144, maximum width 46, length of
submarginal vein 60, of marginal vein 4, of stigmal vein 7, of postmarginal vein 4; of hindwing: length 93,
maximum width 26. Base of forewing as in Fig. 147.
178 JOHN s- NOYES
Caster. Apex of hypopygium reaching to about two-thirds along gaster. Relative measurements of
paratype: length of ovipositor 95, of gonostylus 33 [mid tibia 54]. Ovipositor as in Fig. 148.
cJ. Length 0-92- 1-13 mm.
Similar to female except antennae (Fig. 143) and genitalia (Fig. 144). Relative measurements of head and
forewings similar to those given for female except that scape is proportionately about one-eighth shorter.
DISTRIBUTION. Trinidad, Brazil.
BIOLOGY. Unknown.
MATERIAL EXAMINED
Holotype $, Trinidad: St George, Guanapo Valley, 29.vii.1976 (J. S. Noyes) (BMNH).
Paratypes. Brazil: 2 $ 1 J, Santa Catarina, Nova Teutonia, 29.iv.1941 ; 2 $, Nova Teutonia, 9. v. 1941 ; 10
•$, 10 3, Nova Teutonia, iii.1950; 1 $, Parana, Rondon, 29.x. 1952 (F. Plaumann) (BMNH; MLP, La Plata;
CIBC, Trinidad; USNM, Washington; UC, Riverside; PPRI, Pretoria; ZI, Leningrad).
COMMENTS. Also a further two undescribed species from Trinidad (BMNH) and Brazil (CNC,
Ottawa). Both differ from the type-species in lacking the transverse carina at the top of the
antennal scrobes.
Superficially the females of this genus resemble Tetracnemoidea, but can be easily separated
using the characters given in the key. The males differ in having the antennae simple, without
rami.
The genus is best placed in the subfamily Encyrtinae (as defined by Trjapitzin, 19736) but
placement to tribe is difficult. It may be related to Ooencyrtus and Echthrodryinus but differs
from these in the mandibles having three acute teeth, the hypopygium reaching or very nearly
reaching the apex of the gaster, and the male having very similar antennae to the female as in
some genera included in the subtribe Aphycina.
ARRHENOPHAGOIDEA Girault
(Key couplet 9 21. Figs 21, 141, 142)
Arrhenophagoidea Girault, \9\5d: 73. Type-species: Arrhenophagoidea coloripes Girault, by monotypy.
INCLUDED SPECIES. Four, none of which are known to occur in the Neotropics, but one
undescribed species from Trinidad (BMNH).
REFERENCE. Revision: Annecke & Prinsloo (1974).
DISTRIBUTION. Trinidad. World: South Africa, Australia.
BIOLOGY. Parasites of Diaspididae (Homoptera).
COMMENTS. Fig. 21 is drawn from the collapsed head of the only specimen of this genus taken
from Trinidad. The head of dry mounted material of this genus generally collapses in this way,
giving it a rather narrow, elongate appearance, but in live specimens or in slide mounted material
the head is of more normal encyrtid shape.
The South American species differs from the other known species in having clearly tridentate
mandibles, rather flattened hind tibia and enlarged fifth tarsal segment (Fig. 141), and a long
distinct stigmal vein (Fig. 142). I had originally considered describing a new genus to
accommodate this species, but now feel that it is best to include it in the present genus.
Placed in the tribe Psyllechthrini.
ARRHENOPHAGUS Aurivillius
(Key couplets : 9 1 ; 3 30. Fig. 6)
Arrhenophagus Aurivillius, 1888: 144. Type-species: Arrhenophagus chionaspidis Aurivillius, by monotypy.
Mymariella Risbec, 1951 : 402. Type-species: Mymariella parlatoriae Risbec, by monotypy.
NEOTROPICAL ENCYRTIDAE 179
INCLUDED SPECIES. Three, one of which occurs in the Neotropics : chionaspidis Aurivillius.
REFERENCE. Annecke & Prinsloo (1974).
DISTRIBUTION. Argentina, Peru, Guyana, Puerto Rico, Jamaica and Barbados. World: Nearctic
and Palaearctic regions, India, Africa.
BIOLOGY. Parasites of Diaspididae (Homoptera).
COMMENTS. The only genus included in the tribe Arrhenophagini.
ASEIRBA Cameron
(Key couplets : ? 152; J 93)
Aseirba Cameron, 1884: 127. Type-species: Aseirba caudata Cameron, by monotypy.
INCLUDED SPECIES. One : caudata Cameron ( = cultrata Kerrich) and a further four undescribed
species from Brazil (BMNH; CNC, Ottawa).
DISTRIBUTION. Brazil, Guatemala.
BIOLOGY. Unknown.
COMMENTS. One of the two female syntypes of caudata Cameron (labelled: 'Torola, 1000ft,
champion, B.C.A. Hymen. I Aseirba caudata, Cam., "Aseirba caudata Cam. type. BCA Hy.
1.128'") in the BMNH is here designated LECTOTYPE.
Kerrich (1954: 371) treated Aseirba as a junior synonym ofCerchysius and proposed the name
cultratus as a replacement name for caudatus Cameron nee Forster, 1841. Since Aseirba is here
considered distinct from Cerchysius the original species name is reverted to since cultratus has
not been used since it was originally proposed.
The four undescribed species differ from caudata in having the hypopygium not extending to
the apex of the gaster whereas in caudata it reaches the apex. The mandibles are bidentate and
not tridentate as stated by Kerrich. I hesitate to describe a new genus to accommodate the four
undescribed species on the differences exhibited by the hypopygium until the taxonomic
significance of this character can be fully ascertained. The genus may possibly belong to the tribe
Bothriothoracini, and is very probably near Hemencyrtus.
ATELAPHYCUS Blanchard
Atelaphycus Blanchard, 1940: 107. Type-species, Atelaphycus eriococci Blanchard, by monotypy.
INCLUDED SPECIES. One: eriococci Blanchard (1940: 107).
DISTRIBUTION. Argentina.
BIOLOGY. Parasites of Eriococcidae (Homoptera).
COMMENTS. The type of eriococci Blanchard is apparently lost (De Santis, 1964: 119). It is
impossible to place the genus from the short description, but it may be close to and possibly
synonymous with Metaphycus. Species of Metaphycus with exserted ovipositors are known from
South America.
A VET1ANELLA Trjapitzin
(Key couplets: 9 213; ^ 58. Fig. 128)
Avetianella Trjapitzin, 1968: 97. Type-species: Avetianella capnodiobia Trjapitzin, by monotypy.
INCLUDED SPECIES. Two, neither known from the Neotropics, but two undescribed species from
Trinidad (BMNH; CNC, Ottawa) and Brazil (BMNH).
180 JOHN S. NOYES
REFERENCE. Trjapitzin (197 Id).
DISTRIBUTION. Trinidad, Brazil. World: Holarctic.
BIOLOGY. Parasites of eggs of Cerambycidae and Scolytidae (Coleoptera).
COMMENTS. Placed in the tribe Microteryini, subtribe Oobiina.
AZTECENCYRTUS Timberlake
(Key couplet: 9 101)
Aztecencyrtus Timberlake, 1926: 16. Type-species: Aztecencyrtus flavus Timberlake, by monotypy.
INCLUDED SPECIES. Two : flavus Timberlake (1926: 18) and iceryae (Howard, 1882 : 379) (comb. n.
from Cerchysius West wood).
DISTRIBUTION. Mexico, Jamaica.
BIOLOGY. Parasites of Margarodidae (Homoptera).
COMMENTS. Apparently close to Cirrencyrtus from which it can be separated using the characters
given in the key. Possibly best placed in the tribe Aphycini, subtribe Aphycina, although
Trjapitzin (19736) includes it in the tribe Homalotylini, which may be correct. The only known
specimens of the two included species are the holotypes and both lack the clavae which appear to
be the only certain means of separating the Aphycina from the Homalotylini.
BAEOENCYRTUS De Santis
(Key couplet: ^ 52. Fig. 127)
Baeoencyrtus De Santis, 1964: 204. Type-species: Baeoencyrtus platys De Santis, by monotypy.
INCLUDED SPECIES. One: platys De Santis (1964: 205).
DISTRIBUTION. Argentina.
BIOLOGY. Unknown.
COMMENTS. The holotype of platys is on a slide and is therefore difficult to place systematically. It
may be near Metaphycus (tribe Aphycini, subtribe Paraphycina).
BENNETT1SCA gen. n.
(Key couplet: 9 78. Figs 43, 149, 150)
Type-species: Bennettisca flavigena sp. n.
$. Head. In side view, moderately deep and most strongly curved at a level with top of antennal scrobes,
frontovertex only slightly convex. Eye about one-third longer than broad, naked, very nearly reaching
occipital margin which is sharp. Malar space nearly three-quarters as long as eye and with malar sulcus
absent. Frontovertex a little more than one-third head width; posterior ocellus separated from eye margin
and occipital margin by less than its own diameter, ocelli more or less forming a right angle. Antennal
scrobes moderately deep, meeting dorsally and reaching about half way between antennal toruli and
anterior ocellus; antennal torulus separated from mouth margin by slightly less than its own length and
from other torulus by nearly twice its own length, top of torulus slightly below level of lower eye margin;
scape slightly broadened and flattened, shorter than malar space, pedicel conical and longer than half length
of funicle which is six-segmented, all funicle segments transverse and cylindrical, distinctly broadening
distally, clava three-segmented, much wider and slightly longer than funicle and with a broad oblique apical
truncation, longitudinal sensilla on sixth funicle segment and clava. Frontovertex with numerous
colourless setae and with fine, closely meshed, coriaceous sculpture which gives it an almost granulate
appearance, sculpture becoming more elongate on lower parts of face, almost transversely strigose between
eyes and scrobes, more normal on cheeks and interantennal prominence. Mandibles with two short, acute
teeth and a short truncation, maxillary palpi four-segmented, labial palpi three-segmented.
NEOTROPICAL ENCYRTIDAE 181
Thorax. In side view, moderately deep, scutellum flat. Mesoscutum without notaular lines and about two
and half times as broad as long, axillae meeting apically, scutellum slightly longer than broad with apex
acute and with a very broad translucent flange along sides and at apex, flange about one-eighth as long as
scutellum, completely covering the propodeum medially and slightly downcurved apically as in
Blanchar discus (Fig. 12), propodeum medially about one-eighth as long as scutellum (without flange).
Mesoscutum, axillae, scutellum and mesopleura with fine, closely meshed coriaceous sculpture and covered
with numerous short, pale setae (except mesopleura), propodeum with slightly elongate, reticulate sculpture
extending to spiracles, metanotum with similar sculpture. Forewing hyaline, slightly more than twice as
long as broad, densely hairy, basal cell and costal cell dorsally with setae as dense as in centre of wing;
submarginal vein with an apical hyaline break which extends to anterior wing margin and is more or less
confluent with the speculum ; marginal vein very transverse, almost absent and not reaching anterior margin
of wing, postmarginal vein absent, stigmal vein about one-quarter as long as the submarginal ; marginal
cilia absent at wing apex and along anterior margin of wing. Hind wing about three times as long as broad.
Mid tibial spur about as long as mid basal tarsal segment.
Caster. About as long as thorax and sides with strigose sculpture and very hairy. Cerci about one-third
along gaster, paratergites absent, hypopygium more or less reaching apex of gaster and produced as in
Mucrencyrtus (Fig. 84) but with projection relatively shorter. Ovipositor about two-thirds as long as mid
tibia and hidden, gonostylus about one-fifth as long as mid tibial spur.
cJ. Unknown.
Named in honour of Dr F. D. Bennett.
Bennettisca flavigena sp. n.
(Figs 43, 149, 150)
?. Length 0-85-0-90 mm (holotype 0-85 mm).
Colour. Body generally yellowish orange, slightly paler ventrally. Upper and lower margins of scape,
metanotum, anterior margin of propodeum and dorsum of gaster brown ; cheeks, temples and clypeal
margin very slightly infuscate ; mid and hind tibia with some faintly darker areas.
Head. Relative measurements of head of holotype: maximum head width 66, frontovertex width at
median ocellus 23, malar space 25, eye length 37, eye width 30, POL 15, OOL 2-5, scape length 18, other
proportions of antenna as in Fig. 43.
Thorax. Relative measurements of forewing of holotype: length 122, maximum width 53, length of
submarginal vein 58, marginal vein 2, stigmal vein 13; of hindwing: length 87, maximum width 28. Base of
forewing as in Fig. 150.
Gaster. Relative measurements of paratype: ovipositor length 38, gonostylus length 3 [mid tibia 48, mid
tibial spur 16]. Ovipositor as in Fig. 149.
J. Unknown.
DISTRIBUTION. Trinidad.
BIOLOGY. Unknown, but probably parasites of Aclerdidae (Homoptera).
MATERIAL EXAMINED
Holotype $, Trinidad; Curepe, Santa Margarita Circular Road, 9-23.vi.1974, Moericke trap (F. D.
Bennett} (CNC, Ottawa).
Paratype. 1 ?, same data as holotype (BMNH).
COMMENTS. I have examined a further female from Trinidad (CNC, Ottawa) which was
provisionally placed in this genus, but it has a much longer marginal vein (about four times as
long as broad), more dense setae on the thorax and gaster, and the setae on the submarginal vein,
marginal vein and along the distal margin of the basal cell are very much thicker. I feel it best to
exclude this species from Bennettisca for the present, but future work may show that it can be
satisfactorily accommodated here.
Bennettisca belongs to the same group of genera as Allencyrtus Annecke & Mynhardt and
Mucrencyrtus (subtribe Microteryina) which both include parasites of Aclerdidae. It differs in
182 JOHN S. NOYES
the form of the antennae, sharp occipital margin, broad translucent flange on scutellum and
forewing venation. Both Allencyrtus and Mucrencyrtus have funicle segments all longer than
broad, rounded occipital margin, very short or non-existent flange on scutellum and longer
marginal vein. Also included in this group of genera is Paramucrona which has setae in basal and
costal cell of forewing rather sparse and all the funicle segments longer than broad.
BLANCHARDISCVS De Santis
(Key couplets: ? 20; ^ 16. Figs 12, 13, 115)
Blanchar discus De Santis, 1964: 268. Type-species: Blanchar discus scutellaris De Santis, by monotypy.
INCLUDED SPECIES. One: scutellaris De Santis, 1964: 268.
DISTRIBUTION. Argentina, Brazil.
BIOLOGY. Unknown.
COMMENTS. Placed in the tribe Blanchardiscini.
BLEPYRUS Howard
(Key couplets: 9 143, 236, 250; <£ 9)
Blepyrus Howard, 1 8986 : 234. Type-species : Blepyrus mexicanus Howard, by designation of Ashmead,
(1900: 373).
Coccophoctonus Ashmead, 1900: 375. Type-species: Coccophoctonus dactylopii Ashmead, by monotypy.
INCLUDED SPECIES. Two : clavicornis (Compere) and insularis (Cameron) ; also a further two or
three undescribed species from Trinidad (BMNH).
REFERENCE. Revision: Kerrich (1967: 225-228).
DISTRIBUTION. Brazil, Guyana, Colombia, Trinidad, Mexico. World: Nearctic region, Pacific
Islands, South East Asia, India, Africa.
BIOLOGY. Parasites of Pseudococcidae (Homoptera).
COMMENTS. Placed in the tribe Neodiscini.
BO THRIOCRA ERA Timberlake
(Key couplets: $ 18; J 21)
Bothriocraera Timberlake, 1916: 567. Type-species: Bothriocraera flavipes Timberlake, by monotypy.
INCLUDED SPECIES. Two, one from the Neotropics: bicolor Compere & Zinna, 1955: 101.
DISTRIBUTION. Trinidad. World: Nearctic.
BIOLOGY. Parasites of Pseudococcidae (Homoptera).
COMMENTS. Placed in the tribe Aphycini, subtribe Aphycina.
BOTHRIOTHORAX Ratzeburg
(Key couplets: 9 129, 184; J 111. Fig. 87)
Bothriothorax Ratzeburg, 1844: 209. Type-species: Bothriothorax altensteinii Ratzeburg, by monotypy.
Trimorphocerus Dahlbom, 1857 : 292. Type-species : Bothriothorax altensteinii Ratzeburg, by designation of
Gahan & Pagan (1923: 149).
INCLUDED SPECIES. Twenty-seven, none of which occurs in the Neotropics, but one undetermined
species from Trinidad (BMNH).
NEOTROPICAL ENCYRTIDAE 183
REFERENCE. Peck (1963: 375).
DISTRIBUTION. Trinidad. World: Holarctic.
BIOLOGY. Parasites of larvae of Syrphidae (Diptera).
COMMENTS. Placed in the tribe Bothriothoracini, subtribe Bothriothoracina.
BRACHYPLATYCERUS De Santis
(Key couplets: ? 5; ^ 92. Fig. 9)
Brachyplatycerus De Santis, 19726: 49. Type-species: Brachyplatycerus minutus De Santis, by monotypy.
INCLUDED SPECIES. One: minutus De Santis, 19726: 50; also two further undescribed species
from Trinidad and one from Brazil (BMNH).
DISTRIBUTION. Argentina, Brazil, Trinidad.
BIOLOGY. Unknown.
COMMENTS. Appears to be very close to Hemaenasius, differing only in the number of funicle
segments. Hemaenasius has been placed in the tribe Discodini, subtribe Hemaenasiina by
Trjapitzin & Gordh (19786).
BRETHESIELLA Porter
(Key couplets: 5 140; 3 49. Fig. 63)
Brethesia Timberlake, 1919a: 190. Type-species: Brethesia latifrons Timberlake, by original designation.
[Homonym of Brethesia Schrottky, 1909.]
Brethesiella Porter, 1920: 16. [Replacement name for Brethesia Timberlake.]
Brethesiella Timberlake, 1920a: 96. [Replacement name for Brethesia Timberlake.]
Perissodromus Blanchard, 1940: 107. Type-species: Perissodromus longipes Blanchard, by monotypy.
Neocopidosoma Blanchard, 1940: 126. Type-species: Neocopidosoma coccidophaga Blanchard, by
monotypy. [Homonym of Neocopidosoma Ishii, 1923.] Syn. n.
Noblanchardia Ghesquiere, 1946: 369. [Replacement name for Neocopidosoma Blanchard.] Syn. n.
INCLUDED SPECIES. Four: abnormicornis (Girault, 19170: 118), coccidophaga (Blanchard; De
Santis, 1964: 176) (comb, n.), latifrons (Timberlake; De Santis, 1964: 171) and longipes
(Blanchard; De Santis, 1964: 172).
DISTRIBUTION. Argentina, Brazil, Trinidad.
BIOLOGY. Parasites of Margarodidae (Homoptera).
COMMENTS. I have examined the holotype of Neocopidosoma coccidophaga. It is a male and
belongs to Brethesiella. All four species are extremely close and may just be forms of a single
species.
Both Timberlake (\9\9a: 193) and Tachikawa (1978: 49) state that they believe the host of
Brethesiella is actually chrysopid or hemerobiid larvae (Neuroptera) predatory on Icerya
Signoret (Homoptera: Coccoidea, Margarodidae). However, both Dr F. D. Bennett and I
(CIBC, Trinidad) have reared specimens of this genus in very large numbers from Icerya
montserratensis Riley & Howard.
Placed in the tribe Homalotylini, subtribe Brethesiella.
CALLIENCYRTUS De Santis
(Key couplet: 9 104)
Calliencyrtus De Santis, 1960: 61. Type-species: Calliencyrtus bucculentus De Santis, by monotypy.
INCLUDED SPECIES. One: bucculentus De Santis, 1960: 63.
184 JOHN s- NOYES
DISTRIBUTION. Brazil.
BIOLOGY. Unknown.
COMMENTS. I am unable to place the genus according to Trjapitzin (19736) classification of the
Encyrtidae.
CARABUNIA Waterston
(Key couplets: ? 125; ^ 60. Fig. 64)
Carabunia Waterston, 1928: 249. Type-species: Carabunia myersi Waterston, by monotypy.
INCLUDED SPECIES. Five, two from the Neotropics : myersi Waterston and waterstoni Subba Rao,
and one undetermined species from Colombia (AMNH, New York).
REFERENCE. Revision of world species: Subba Rao (1973).
DISTRIBUTION. Colombia, Cuba, Puerto Rico, Jamaica. World: Africa and Asia.
BIOLOGY. Parasites of Cercopidae (Homoptera).
COMMENTS. Placed in the tribe Neocladini.
CERCHYSIUS Westwood
(Key couplets: $ 153, 154)
Cerchysius Westwood, 1832: 128. Type-species: Encyrtus urocerus Dalman, by designation of Westwood
(1840: 73).
INCLUDED SPECIES. Nineteen, none of which occurs in the Neotropics, but two undetermined
species from Ecuador (CNC, Ottawa) and Peru (BMNH) and perhaps also Prionomitus
fuscipalpis Kieffer in Kieffer & Jorgensen, 1910: 413.
REFERENCE. Partial world revision: Kerrich (1954).
DISTRIBUTION. Ecuador, Peru. World: cosmopolitan.
BIOLOGY. Parasites of Chamaemyiidae (Diptera).
COMMENTS. Placed in the tribe Microteryini, subtribe Pseudencyrtina.
CHALCASPIS Howard
(Key couplets : $ 141; J 7)
Chalcaspis Howard, 1895: 606. Type-species: Chalcaspis pergandei Howard, by monotypy.
INCLUDED SPECIES. Four, one from the Neotropics : lucida Kerrich.
REFERENCE. Revision: Kerrich (1967: 191-195).
DISTRIBUTION. Cuba. World: Nearctic.
BIOLOGY. Parasites of Pseudococcidae (Homoptera).
COMMENTS. Placed in the tribe Rhinoencyrtini, subtribe Aenasiina.
CHEILONEURUS Westwood
(Key couplets: $ 34, 58, 95; 3 86, 115, 131, 136, 144, 146. Fig. 36)
Cheiloneurus Westwood, 1833: 343. Type-species: Encyrtus elegans Dalman, by monotypy.
Chrysopophagus Ashmead, 1894: 246. Type-species: Chrysopophagus compressicornis Ashmead, by
monotypy.
NEOTROPICAL ENCYRTIDAE 185
Blatticida Ashmead, 1904: 305. Type-species: Blatticida pulchra Ashmead, by monotypy.
Cristatothorax Girault, 1911 : 169. Type-species: Cristatothorax pulcher Girault, by monotypy.
Metacheiloneurus Hoffer, 1957 : 336. Type-species: Metacheiloneurus moestus Hoffer, by monotypy. Syn. n.
INCLUDED SPECIES. Ninety-four, 1 1 from the Neotropics : albinotatus De Santis, angulatus De
santis, cristatus De Santis, cupreicollis Ashmead, elegans (Dalman), gahani (Dozier, 1927: 270)
(comb. n. from Achrysopophagus), inimicus Compere, 1925: 297, loretanus De Santis, 1972: 60,
nigrescens Howard (= longisetaceus De Santis), praenitens Waterston, 1922: 49 and pulvinariae
Dozier, 1927: 363; also a further ten or more undetermined species (BMNH; USNM,
Washington).
REFERENCE. Revision of Argentinian species: De Santis (1964: 332-353).
DISTRIBUTION. Throughout the Neotropics. World: cosmopolitan.
BIOLOGY. Hyperparasites of other Chalcidoidea, mainly Encyrtidae and Aphelinidae
(Hymenoptera).
COMMENTS. Placed in the tribe Cheiloneurini.
CHR YSOPLA TYCERUS Ashmead
(Key couplets: 9 46; J1 36. Fig. 26)
Rileya Howard, 1888: 80. Type-species: Rileya splendens Howard, by monotypy. [Homonym of Rileya
Ashmead, 1888.]
Chrysoplatycerus Ashmead, 1889: 38. [Replacement name for Rileya Howard.]
Encyrtolophus De Santis, 19726: 49. Ty pe- species : Encyrtolophus flavicollis De Santis, by monotypy.
Paraplatycerus Hall, 1974: 19. Type-species: Paraplatycerus citriculus Hall, by monotypy.
Metaplatycerus Gordh & Trjapitzin in Trjapitzin & Gordh, 1978a: 384. Type-species: Chrysoplatycerus
ferrisi Timberlake, by monotypy. Syn. n.
INCLUDED SPECIES. Four, three of which occur in the Neotropics : colombiensis Kerrich, flavicollis
(De Santis) and splendens (Howard).
REFERENCE. Revision: Kerrich (1978: 136-142).
DISTRIBUTION. Throughout the Neotropics. World: Nearctic region, South Africa.
BIOLOGY. Parasites of Pseudoccidae (Homoptera).
COMMENTS. I disagree with Trjapitzin & Gordh's (1978a) view that ferrisi requires generic status.
C. flavicollis exhibits characters intermediate between splendens and ferrisi, notably in the
structure of the mesoscutum and scutellum, although it is nearest to ferrisi. If Metaplatycerus
and Chrysoplatycerus were to be maintained as separate genera then Metaplatycerus should be
considered a junior synonym of Encyrtolophus.
Placed in the tribe Chrysoplatycerini, subtribe Chrysoplatycerina.
CIBDELOENCYRTUS De Santis
(Key couplet: 9 231)
Cibdeloencyrtus De Santis, 1964: 209. Type-species: Cibdeloencyrtus aphelinoides De Santis, by monotypy.
INCLUDED SPECIES. One: aphelinoides De Santis, 1964: 210.
DISTRIBUTION. Argentina.
BIOLOGY. Unknown.
COMMENTS. Difficult to place according to Trjapitzin's (19736) classification, but it is possibly
related to Echthroplexiella (tribe Miraini, subtribe Echthroplexiellina).
186 JOHN s- NOYES
CICOENCYRTUS gen. n.
(Key couplets: 9 240; 3 54. Figs 108, 151-155)
Type-species : Cicoencyrtus angustifrons sp. n.
9. Head. In side view moderately deep and slightly angled at a level with dorsal margin of antennal scrobes,
frontovertex slightly convex. Eye about one-half longer than broad, clothed with short, pale setae and
overreaching occipital margin which is rounded. Malar space less than half as long as eye and with malar
sulcus absent. Frontovertex narrow, less than one-fifth head width; posterior ocellus separated from
occipital margin by slightly more than its own diameter and almost touching eye margin, ocelli forming a
strongly acute angle of about 25-45°. Antennal scrobes moderately impressed, long and narrow, meeting
dorsally, reaching slightly more than half way between antennal toruli and anterior ocellus and sharply
separated from frontovertex; antennal torulus separated from mouth margin by much less than half its own
length and from other torulus by slightly less than twice its own length, top of torulus slightly below level of
lower eye margin ; scape subcylindrical and stout, pedicel conical and about as long as the first four funicle
segments together, funicle six-segmented, segments transverse, cylindrical and broadening distally, distal
segments occasionally quadrate or slightly longer than broad, clava three-segmented and rounded apically,
more than half as long as funicle, longitudinal sensilla on apical two or three funicle segments and clava.
Frontovertex with shallow coriaceous sculpture except between and behind ocelli where it is more coarse
and sometimes granulate in appearance, more elongate on lower parts of face, sparse brown setae on
frontovertex and cheeks, interantennal prominence with at least a few conspicuous white setae. Mandibles
with three acute teeth, maxillary palpi four-segmented, labial palpi three-segmented.
Thorax. In side view moderately deep and dorsally more or less flat. Mesoscutum without notaular lines,
slightly broader than long, axillae touching, scutellum about as long as broad and with apex more or less
rounded, propodeum very short, medially less than one-tenth as long as scutellum. Pronotum, mesoscutum
and axillae slightly shiny and with coriaceous sculpture, scutellum matt and with punctiform sculpture
giving it an almost granulate appearance, mesopleura anteriorly with shallow strigose sculpture becoming
more regular and reticulate posteriorly, propodeum centrally smooth, but with some irregular, shallow
sculpture laterally. Forewing hyaline, nearly two and half times as long as broad, basal cell sparsely hairy
and naked proximally, speculum not interrupted but more or less closed ; submarginal vein with an apical
hyaline break, marginal vein about as long as broad to nearly twice as long as broad, stigmal at least about
twice as long as marginal, postmarginal about as long as marginal or slightly shorter. Hind wing about
three and a half times as long as broad. Mid tibial spur much shorter than mid basal tarsal segment.
Caster. Slightly shorter than thorax. Cerci slightly more than one-third along gaster, paratergites absent,
apex of hypopygium from about one half to three-quarters along gaster and incised apically, exserted part
of ovipositor equal to about two-thirds length of gaster or mid tibia. Ovipositor about one and half times as
long as gaster or slightly more than twice as long as mid tibia, gonostylus slightly longer than mid tibia.
cJ. Similar to female except for genitalia and clava which is entire.
Cicoencyrtus angustifrons sp. n.
(Figs 108, 151-155)
?. Length (excluding ovipositor): 1-33-1-49 mm (holotype 1-33 mm).
Colour. Frontovertex reddish brown, black between ocelli and on occipital margin between eyes; lower
parts of face and antennal scrobes black, slightly tinged green, particularly on genae ; eyes reddish ; scape
and basal half of pedicel brown, apical half of pedicel and flagellum yellowish organge, slightly darker
proximally; thorax black, tinged green with some slightly paler areas which are dark brownish orange; legs
brown, except apical half of all tibiae and arsi which are yellowish testaceous ; metanotum and centre of
propodeum brownish orange; gaster dark brown with cereal plates ringed orange-brown, ovipositor
sheaths dark brown.
Head. In frontal aspect as in Fig. 108. Ocelli forming an angle of about 25-30°. Relative measurements of
head of holotype : maximum head width about 67, minimum width of frontovertex 8, malar space 25, eye
length about 55, eye width about 36, POL 5, OOL 1, scape length 38, other proportions of antenna as in
Fig. 154.
NEOTROPICAL ENCYRTIDAE 1 g?
Thorax. Relative measurements of forewing of holotype: length 172, maximum width 78, length of
submarginal vein 72, narginal vein 5-5, stigmal vein 14, postmarginal vein 5-5; of hindwing: length 112,
maximum width 30.
Caster. Apex of hypopygium about halfway along gaster. Relative measurements of gaster of paratype:
length of exserted part of ovipositor 52, length of ovipositor 147, length of gonostylus 74 [mid tibia 66].
Ovipositor as in Fig. 155.
cJ. Length 1-23-1 -29 mm.
As for female. Antenna as in Fig. 151, base of forewing 154, genitalia 152. Aedeagus just over half as long
as mid tibia, digiti each with two spines at apex.
DISTRIBUTION. Brazil.
BIOLOGY. Unknown.
MATERIAL EXAMINED
Holotype ?, Brazil: Santa Catarina, Nova Teutonia, 2.xi.l944 (F. Plaumanri) (BMNH).
Paratypes. Brazil: 1 ?, 2 J, Santa Catarina, Nova Teutonia, 20.ix.1944, 14.x. 1944, 13.x. 1944 respectively
(F. Plaumann) (BMNH).
COMMENTS. Also one further undescribed species from Brazil (BMNH) and Trinidad (CNC,
Ottawa), the female of which differs from the type-species in having a wider frontovertex and a
more or less anelliform fourth funicle segment.
The genus appears to be related to Archinus but can be easily separated by having six funicle
segments (Archinus has five). Possibly best placed in the tribe Aphycini, subtribe Paraphycina.
CIRRHENCYRTUS Timberlake
(Key couplets: $ 101, 211; J1 44, 53)
Cirrhencyrtus Timberlake, 1918: 360. Type-species: Cirrhencyrtus ehrhorni Timberlake, by monotypy.
Neoanisotylus De Santis, 1939: 330. Type-species: Neoanisotylus bimaculatus De Santis, by monotypy.
INCLUDED SPECIES. Four, two of which are found in the Neotropics : bimaculatus (De Santis,
1939: 331) and diversicolor (Compere, 1939: 62) (comb. n. from Waterstonia}.
DISTRIBUTION. Brazil, Argentina, Trinidad. World: Holarctic region.
BIOLOGY. Parasites of Pseudococcidae (Homoptera).
COMMENTS. I have not seen any types of Waterstonia diversicolor, but it is clear from the
description that it belongs to Cirrhencyrtus and is very close to bimaculatus.
Trjapitzin & Gordh (19786) place the genus in the tribe Homalotylini, subtribe Homalotylina.
In my view this genus would be better placed in the tribe Aphycini, subtribe Aphycina. I have
examined specimens determined by Timberlake as Cirrhencyrtus ehrhorni and also the holotype
of Neoanisotylus bimaculatus and both show remarkable similarities to the species of Aphycus
Mayr, particularly the latter. They differ principally in having the well-marked notaular lines
(absent in Aphycus) and relatively longer ovipositor. Perhaps this genus shows a closer
relationship between the two subtribes than Trjapitzin's (19736) classification would infer.
COCCID APHYCUS Blanchard
(Key couplets:? 26; J 13)
Coccidaphycus Blanchard, 1940: 110. Type-species: Coccidaphycus nigricans Blanchard, by monotypy.
INCLUDED SPECIES. One: nigricans Blanchard (De Santis, 1964: 244); also one undescribed
species from Trinidad (BMNH).
DISTRIBUTION. Argentina, Trinidad.
BIOLOGY. Parasites of Coccidae (Homoptera).
188 JOHN S. NOYES
COMMENTS. The genus appears to be very close to Trechnites Thomson (tribe Trechnitini,
subtribe Trechnitina) but differs in the longer marginal vein, very slight triangular expansion at
apex of submarginal vein and host (Trechnites spp. are psyllid parasites).
COCCIDENCYRTUS Ashmead
(Key couplets: $ 68, 88, 89, 167, 222, 244; J1 130, 140, 143. Figs 39, 47)
Coccidencyrtus Ashmead, 1900: 383. Type-species: Encyrtus ensifer Howard, by monotypy.
Coccidencyrtoides Blanchard, 1940: 107. Type-species: Coccidencyrtoides annulipes Blanchard, by
designation of De Santis (19676: 161).
INCLUDED SPECIES. Twenty-four, seven from the Neotropics : annulipes (Blanchard), blanchardi
(De Santis), denieri Blanchard, grioti Blanchard, maculicornis (Blanchard), malloi Blanchard and
obesus De Santis; also a further four or five undetermined species from Brazil and Trinidad
(BMNH;CNC, Ottawa).
REFERENCE. Revision of Argentinian species: De Santis (1964: 218-231).
DISTRIBUTION. Argentina, Brazil, Trinidad. World: cosmopolitan.
BIOLOGY. Parasites of Diaspididae (Homoptera).
COMMENTS. The types of denieri and grioti are apparently lost and the names should be
considered nomina dubia, but in any case denieri cannot belong in Coccidencyrtus because the
frontovertex is too narrow. I have not see the types of either maculicornis or malloi but, from De
Sands' (1964) redescriptions of these species, they may be correctly placed. The only described
species from South America which are with certainty correctly placed are blanchardi and
annulipes; obesus does not belong in Coccidencyrtus but at present I cannot place it generically.
Placed in the tribe Habrolepidini (Trjapitzin & Gordh, 19786).
COCCIDOCTONUS Crawford
(Key couplets: $ 186, 223; $ 105. Fig. 89)
Coccidoctonus Crawford, 1912: 167. Type-species: Coccidoctonus trinidadensis Crawford, by monotypy.
Quay lea Timberlake, 19196 : 214. Type-species : Cerchysius whittieri Girault, by original designation. Syn. n.
INCLUDED SPECIES. Two, one of which is found in the Neotropics: trinidadensis Crawford, 1912:
168.
DISTRIBUTION. Trinidad, Barbados, Grenada, St Vincent. World: Nearctic region, South East
Asia, Australia.
BIOLOGY. Hyperparasite of other Encyrtidae and of Pteromalidae (Hymenoptera) parasitizing
Coccidae and Pseudococcidae (Homoptera).
I have examined specimens determined by Timberlake as Quay lea whittieri and these belong to
Coccidoctonus (comb, n.), although they are a distinct species from trinidadensis.
I am unable to place the genus according to Trjapitzin's (19736) classification of the
Encyrtidae.
COELASPIDIA Timberlake
(Key couplets : 9 30; 3 39)
Coelaspidia Timberlake, 1923: 326. Type-species: Coelaspidia osborni Timberlake, by monotypy.
INCLUDED SPECIES. One: osborni Timberlake, 1923: 330.
DISTRIBUTION. Mexico.
BIOLOGY. Parasites of Pseudococcidae (Homoptera).
COMMENTS. Placed in the tribe Acroaspidiini.
NEOTROPICAL ENCYRTIDAE 1 89
COELOPENCYRTUS Timberlake
(Key couplets: ? 89, 252; J 40. Figs 100, 120)
Coelopencyrtus Timberlake, 19196: 218. Type-species: Coelopencyrtus odyneri Timberlake, by original
designation.
Nesencyrtus Timberlake, 19196: 223. Ty pe- species : Adelencyrtus kaalae Ashmead, by monotypy.
Batrachencyrtus Jansson, 1957: 71. Type-species: Batrachencyrtus callidii Jansson, by monotypy.
INCLUDED SPECIES. Nineteen, one of which is found in the Neotropics: gargaris (Walker)
(= Giraultella lopesi Costa Lima & Ferreira, 1963: 431).
REFERENCE. World revision: Burks (1958: 22-26); revision African species: Annecke (1968:
250-258).
DISTRIBUTION. Brazil, Trinidad, St Vincent. World: cosmopolitan.
BIOLOGY. Polyembryonic parasites of larvae of Xylocopidae (Hymenoptera) (and also Hylaeidae
in other parts of the world).
COMMENTS. Placed in the tribe Copidosomatini, subtribe Coelopencyrtina.
COMPERIA Gomes
(Key couplets: ? 65; J1 27)
Comperia Gomes, 1942: 41. Type-species: Dicarnosis merceti Compere, by monotypy.
INCLUDED SPECIES. Seven, one of which is found in the Neotropics: merceti (Compere, 1938:
317).
REFERENCE. Key to world species: Prinsloo & Annecke (1978: 315-316).
DISTRIBUTION. Throughout the Neotropics. World: Nearctic region, African continent,
Australia.
BIOLOGY. Parasites of cockroach oothecae (Orthoptera : Blattodea).
COMMENTS. Placed in the tribe Comperiini.
COMPERIELLA Howard
(Key couplets : $ 47; 3 137)
Comperiella Howard, 1906: 121. Type-species: Comperiella bifasciata Howard, by monotypy.
Pseudanusia Girault, \9\5d: 155. Type-species: Pseudanusia pia Girault, by monotypy.
Habrolepistia Mercet, 1921 : 668. Type-species: Habrolepistia cerapterocera Mercet, by monotypy.
INCLUDED SPECIES. Nine, one of which is from the Neotropics: bifasciata Howard
(= Habrolepistia cerapterocera Mercet, 1921 : 669).
DISTRIBUTION. Argentina, Chile, Mexico, Bermuda. World: cosmopolitan.
BIOLOGY. Parasites of Diaspididae (Homoptera).
COMMENTS. Placed in the tribe Habrolepidini, subtribe Comperiellina.
COP1DOSOMA Ratzeburg
(Key couplets: 9 155, 173, 201, 253; 3 77. Figs 69, 97, 105, 107, 109)
Copidosoma Ratzeburg, 1844: 157. Type-species: Copidosoma boucheanum Ratzeburg, by monotypy.
Verdunia Mercet, 19176: 203. Type-species: Verdunia gloriosa Mercet, by monotypy.
Neocopidosoma Ishii, 1923: 101. Type-species: Neocopidosoma komabae Ishii, by monotypy.
Arrenoclavus Doutt, 1948: 145. Type-species: Copidosoma koehleri Blanchard, by monotypy.
190 JOHN S. NOYES
INCLUDED SPECIES. Eighty-one, five of which are from the Neotropics : desantisi Annecke &
Mynhardt, 1974: 32, hyalinistigma De Santis, 1964: 281, koehleri Blanchard (Annecke &
Mynhardt, 1974: 32), silvestrii (Costa Lima, 1956: 31) (comb. n. from Paralitomastix) and
virescens De Santis, 19726: 57, also a further six undetermined species from Brazil and West
Indies (BMNH ; CNC, Ottawa).
DISTRIBUTION. Throughout the Neotropics. World: cosmopolitan.
BIOLOGY. Polyembryonic parasites of larvae of Lepidoptera.
COMMENTS. Placed in the tribe Copidosomatini, subtribe Copidosomatina. All the genera in this
subtribe are extremely close. Litomastix and Pentalitomastix both usually have a strongly,
obliquely truncate clava (sometimes more rounded on ventral surface than obliquely truncate)
whereas all the other genera have the apex of the clava transversely, or very slightly obliquely,
truncate or rounded. The latter can be separated using the characters given in the key.
CYDERWS gen. n.
(Key couplets: ? 187. Fis 92, 159)
Type-species : Cyderius urbicola sp. n.
?. Head. In profile, moderately deep and most rounded at level with top of antennal scrobes, frontovertex
slightly convex. Eye slightly longer than broad, with sparse, short pale hairs and very nearly reaching
occipital margin which is acute. Malar space about half length of eye with malar sulcus weakly indicated.
Frontovertex about one-third head width; posterior ocellus separated from eye and occipital margin by
about half its diameter, ocelli forming an acute angle of about 75°. Antennal scrobes moderately deep,
meeting dorsally and reaching slightly further than half way between antennal toruli and anterior ocellus ;
antennal torulus separated from mouth margin by slightly less than its own length and from other torulus
by nearly one and a half times its own length, dorsal margin of torulus slightly below ventral level of eyes ;
scape subcylindrical and stout, pedicel conical and about as long as first funicle segment, funicle six-
segmented, all segments distinctly longer than broad, except the second which is anelliform, clava three-
segmented, about half as long as funicle and with apex more or less rounded ; longitudinal sensilla very
conspicuous and present on all flagellar segments except second. Mouth opening narrow, slightly narrower
than minimum width of frontovertex. Frontovertex with shallow, reticulate sculpture, more elongate and
coriaceous on lower parts of face. Mandibles with three acute teeth.
Thorax. In side view moderately deep, with mesoscutum and scutellum more or less flat. Mesoscutum
without notaular lines, nearly twice as broad as long, axillae meeting, scutellum about as broad as long with
apex pointed and produced as a short thin flange, propodeum medially about one-eighth as long as
scutellum. Pronotum, mesoscutum and axillae with shallow coriaceous sculpture, scutellum with deeper,
reticulate sculpture which becomes shallower, more irregular and elongate towards sides and apex,
mesopleura almost smooth with very shallow, elongate, reticulate sculpture. Forewing hyaline, about two
and a half times as long as broad; basal cell sparsely pilose throughout except for a small, naked, triangular
area proximally, speculum not interrupted but more or less closed ; submarginal vein with an apical hyaline
break, marginal vein slightly longer than broad, about twice as long as postmarginal and about half as long
as stigmal. Hindwing about four and a half times as long as broad. Mid tibial spur nearly as long as
corresponding basal tarsal segment.
Caster. About as long as thorax, with exserted part of ovipositor about one-fifth length of gaster or one-
quarter mid tibia, cerci about three-eighths along gaster, apex of hypopygium not reaching half way along
gaster.
cJ. Unknown.
Cyderius urbicola sp. n.
(Figs 92, 159)
$. Length (excluding ovipositor): 0-73 mm.
Colour. Head and thorax dark brown and shiny; scape testaceous, pedicel and flagellum brown; some
NEOTROPICAL ENCYRTIDAE 191
orange areas along top of antennal scrobes just above interantennal prominence to outer margins of toruli
and from there to lower margin of eye (these may represent lines of weakness in head capsule); legs
completely yellow; gaster dark brown, basally and medially orange-brown, ventrally orange; ovipositor
sheaths orange-brown.
Head. Relative measurements of head : maximum width 58, frontovertex width at median ocellus 22-5,
malar space about 20, eye length 35, eye width 29, POL 9, OOL 2-5, scape length 28, other proportions of
antenna as in Fig. 92.
Thorax. Relative measurements of forewing (Fig. 159): length 164, maximum width 63, length of
submarginal vein 66, marginal vein 6, stigmal vein 13, postmarginal 3; of hindwing: length 108, maximum
width 23.
j. Unknown.
DISTRIBUTION. Trinidad.
BIOLOGY. Unknown.
MATERIAL EXAMINED
Holotype ?, Trinidad: St George, St Augustine, 15.vii.-13.viii.1976, Malaise trap (J. S. Noyes) (BMNH).
COMMENTS. I am unable to place this genus according to Trjapitzin's (19736) classification of the
Encyrtinae, although the wings and the structure of the thorax suggest that it may be related to
Archinus and Cicoencyrtus which probably belong in the tribe Aphycini, subtribe Paraphycina. It
can easily be separated on the unusual form of the antenna i.e. all segments longer than broad,
except the second which is anelliform, and also the presence of an apical flange on the scutellum.
DELOENCYRTUS De Santis
(Key couplets : $ 104, 150)
Deloencyrtus De Santis, \961a: 68. Type-species: Hemencyrtus kuscheli De Santis, by monotypy.
INCLUDED SPECIES. One: kuscheli (De Santis, 1955: 193).
DISTRIBUTION. Juan Fernandez Island, Chile.
BIOLOGY. Unknown.
COMMENTS. Very probably belongs to the tribe Microteryini, and possibly subtribe
Pseudencyrtina.
DESANTISELLA Subba Rao
(Key couplets: $ 124; ^ 31. Fig. 61)
Desantisella Subba Rao, 1971: 215. Type-species: Desantisella brasiliensis Subba Rao, by original
designation.
INCLUDED SPECIES. Three: brasiliensis Subba Rao, 1971 : 216, plaumanni Subba Rao, 1971 : 218
and trifasciata De Santis, 19720: 22.
DISTRIBUTION. Brazil.
BIOI OGY. Hyperparasites of Chamaemyiidae (Diptera) on Aclerdidae (Homoptera).
COMMENTS. Extremely close to Meromyzobia (tribe Miraini, subtribe Mayridina) and differs only
in the form of the mid tibial spur (Meromyzobia has the mid tibial spur of normal encyrtid form).
I think it very likely that the two genera will eventually be considered synonymous, but I hesitate
to formally synonymize them here because material of Ericydnus maculipennis (type-species of
the genus) has not been seen although I have examined some material from Florida which very
probably belongs to this genus.
192 JOHN S. NO YES
DESOBWS gen. n.
(Key couplets: ? 86, 229; j1 26. Figs 119, 160-163)
Type-species : Desobius convexus sp. n.
$. Head. In profile, moderately deep and more or less evenly curved. Eye about a half longer than broad
with conspicuous long setae, and reaching occipital margin which is sharp. Malar space about half eye
length and with malar sulcus present. Frontovertex about one-quarter to one-half head width and with
numerous long dark setae; posterior ocellus separated from occipital margin by less than to slightly more
than its own diameter and from eye margin by much less than its own diameter, ocelli forming an acute
angle of about 60°. Antennal scrobes moderately impressed, meeting dorsally and reaching about halfway
from antennal toruli to anterior ocellus ; antennal torulus separated from mouth margin by about its own
length and from other torulus by nearly twice its own length and with dorsal margin below lowest margin of
eye; scape subcylindrical or very slightly broadened and flattened with several strong, dark bristles along
ventral margin, pedicel conical and slightly less than half as long as funicle, funicle six-segmented, all
segments transverse (mostly anelliform) except perhaps the last which may be subquadrate, clava very large,
longer and broader than funicle, three-segmented, apically rounded or with a short oblique truncation;
longitudinal sensilla on sixth funicle segment and clava only. Frontovertex usually appearing fairly smooth
and shiny, but with shallow coriaceous sculpture which becomes more elongate and of larger mesh on
cheeks and lower parts of face, occasionally sculpture deeper and therefore frontovertex less shiny.
Mandibles with three acute teeth, the middle the longest, maxillary palpi four-segmented, labial palpi three-
segmented.
Thorax. Deep in side view and dorsally convex. Mesoscutum convex and without notaular lines, about a
half broader than long, axillae not meeting centrally, scutellum very convex, longer than broad, propodeum
medially from about one-sixth to one-tenth as long as scutellum. Scutellum with very inconspicuous
tubercles or pits like those found in Forcipestricis. Mesoscutum and scutellum with deep, longitudinally
semi-striate or strigose sculpture, matt, axillae with deep reticulate-coriaceous sculpture, mesopleura with
irregular shallow sculpture, propodeum smooth medially. Forewing hyaline with some slight infumation
basally and beneath marginal vein, about two and a half times as long as broad; basal cell with sparse setae
throughout except a large naked area proximally, speculum not interrupted and open; submarginal vein
with an apical hyaline break, marginal vein about three or four times as long as broad, slightly shorter than
stigmal and about as long as postmarginal. Hindwing about four and a half times as long as broad. Mid
tibial spur slightly shorter than corresponding basal tarsal segment.
Caster. Slightly shorter than thorax. Cerci in basal third, paratergites absent, apex of hypopygium nearly
reaching apex of gaster. Ovipositor hidden or very slightly protruding, ovipositor short, only slightly longer
than half of gaster or about three-fifths to three-quarters length of mid tibia, gonostylus about as long as
mid tibial spur.
J1. Essentially similar to female except the frontovertex is less smooth and therefore less shiny, antennae
(Fig. 1 19) and genitalia (Fig. 161), digiti each with an apical spine and aedeagus slightly less than two-fifths
as long as mid tibia. Antennae with flagellum, in some cases having seven-segmented funicle with a one- or
two-segmented clava, or in others the funicle is six-segmented with a two- or three-segmented clava. In dry
specimens the clava always appears two- or three-segmented, but in cleared slide mounted material the
division of the apical segment into two is incomplete or only indicated by a slight thinning of cuticle and
distribution of sensillae. Longitudinal sensilla very conspicuous (even on dry mounted material) and
present on all flagellar segments.
Desobius convexus sp. n.
(Figs 160, 162, 163)
?. Length: 0-78-0-86 mm (holotype 0-78 mm).
Colour. Head, dorsum of thorax, propodeum and gaster black, frontovertex and cheeks shiny with faint
blue or purple sheen, dorsum of thorax matt, gaster dorsally with some brassy, blue and purple reflections;
antenna with scape testaceous, pedicel and clava brown, funicle brownish testaceous ; mesopleura and legs
pale orange, fore and hind legs, except coxae, mixed brown, particularly fore femora and tibiae and hind
femora.
Head. Apex of clava with an indistinct oblique truncation which is much shorter than ventral margin of
NEOTROPICAL ENCYRTIDAE 193
clava (Fig. 160); posterior ocellus separated from occipital margin by slightly more than half its major
diameter; fronto vertex fairly smooth with shallow coriaceous sculpture. Relative measurements of head of
holotype: maximum head width 48, width of frontovertex at median ocellus 12-5, malar space 19, eye length
34, eye width 26, POL 6, OOL 2, scape length 20, other proportions of antenna as in Fig. 160.
Thorax. Relative measurements of forewing of holotype: length 134, maximum width 55, length of
submarginal vein 51, of marginal vein 6, of stigmal vein 8, of postmarginal vein 6. Base of forewing as in
Fig. 163.
Caster. Exserted part of ovipositor about half length of mid tibial spur. Relative measurements of gaster
of paratype: length of ovipositor 33, length of gonostylus 13 [mid tibia 51, mid tibial spur 13].
$. Unknown.
DISTRIBUTION. Trinidad.
BIOLOGY. Unknown.
MATERIAL EXAMINED
Holotype ?, Trinidad: St George, El Tucuche (S. slope), 25.vii.1976 (J. S. Noyes) (BMNH).
Paratype. 1 ?, same data as holotype (BMNH).
COMMENTS. The genus also includes Desobius sylvicola (De Santis, \961a: 71) (comb. n. from
Tyndarichus) from Argentina. I have also examined a possible ten further undescribed species
from Trinidad, St Vincent (BMNH), Ecuador, Brazil and Panama (CNC, Ottawa), differing in
the sculpture of the frontovertex, relative positions of ocelli, proportions of antennal segments,
length of exserted part of ovipositor and coloration. The genus also occurs in India and possibly
in Europe (Coccidencyrtus pinicola Mercet and Parageniaspis montana Mercet may belong here)
and Australia. See comments under Amaurilyma.
DICARNOSIS Mercet
(Key couplet : 9 64)
Dicarnosis Mercet, 1921 : 141. Type-species: Dicarnosis superbus Mercet, by monotypy.
Neither blanchardi De Santis, 19726: 48 nor ogloblini De Santis, 19726: 48 are correctly placed
here, although originally described in Dicarnosis: D. ogloblini belongs to Metaphycus and
blanchardi very probably belongs to an undescribed genus near Metaphycus.
DIVERSINERVUS Silvestri
(Key couplets: $ 56; J 64, 103)
Diversinervus Silvestri, \9l5a: 301. Type-species: Diver sinervus elegans Silvestri, by monotypy.
Cheiloneuroides Girault, \9l5d: 96. Type-species: Cheiloneuroides bicristatus Girault, by monotypy.
INCLUDED SPECIES. Ten, one of which is found in the Neotropics : elegans Silvestri.
REFERENCE. Key to world species: Hayat, Alam & Agarwal (1975: 39-41).
DISTRIBUTION. Brazil, Peru. World: cosmopolitan.
BIOLOGY. Parasites of Coccidae (Homoptera).
COMMENTS. Placed in the tribe Cheiloneurini.
ECHTHRODRYINUS Perkins
(Key couplets: $ 206, 257; J 122, 145. Fig. 91)
Echthrodryinus Perkins, 1906: 252. Type-species: Echthrodryinus destructor Perkins, by monotypy.
INCLUDED SPECIES. Six, one of which is from the Neotropics: saccharalis Gordh & Trjapitzin;
also a further four or five undetermined species from Trinidad, Brazil and Guyana (BMNH;
CNC, Ottawa).
194 JOHN S. NO YES
REFERENCE. Revision: Gordh & Trjapitzin (1978).
DISTRIBUTION. Venezuela, Brazil, Trinidad, Guyana. World: cosmopolitan.
BIOLOGY. Hyperparasites via other parasitic Hymenoptera (mainly Dryinidae and Braconidae).
COMMENTS. Two of the undetermined species that I have included in this genus are perhaps
atypical; one is very common in South America and has been reared solely from syrphid
(Diptera) puparia, the other has the thorax exceptionally deep and the body very squat. The
former is best placed in the present genus even though the stigmal vein forms a slightly greater
angle with the postmarginal vein than is usual for this genus.
Placed in the tribe Microteryini, subtribe Syrphophagina. Some species of this genus strongly
resemble some species of Ooencyrtus and Syrphophagus. It can best be separated from the latter
by the relative length of the scutellum (see key), and from the former by the much longer
postmarginal vein and more strongly clavate antennae.
ECHTHROPLEX1ELLA Mercet
(Key couplet : J 32)
Echthroplexiella Mercet, 1921: 183. Type-species: Echthroplexiella submetalica Mercet, by original
designation.
INCLUDED SPECIES. Twenty, one of which is found in the Neotropics : josephi De Santis, 1965: 4.
DISTRIBUTION. Argentina. World : Palaearctic region.
BIOLOGY. Parasites or hyperparasites (via other Encyrtidae) on Eriococcidae (Homoptera).
COMMENTS. I am very doubtful about the generic placement of josephi. The unique holotype male
is dissected on a slide and may be a brachypterous male Metaphycus.
ECTROMA Westwood
(Key couplet : ? 38)
Ectroma Westwood, 1833: 344. Type-species: Ectroma fulvescens Westwood, by monotypy.
Metallon Walker, 1848: 219. Type-species: Metallon acacallis Walker, by monotypy.
Pezobius Forster, 1860: 129. Type-species: Pezobius polychromus Forster, by monotypy.
INCLUDED SPECIES. Eleven, one of which is found in the Neotropics: semifacta De Santis, 19726:
52.
DISTRIBUTION. Argentina. World: Palaearctic region.
BIOLOGY. Unknown.
COMMENTS. I have examined the holotype of semifacta and it appears to be correctly assigned to
Ectroma.
Placed in the tribe Miraini, subtribe Mayridiina.
ECTROMATOPSIS Compere
(Key couplets: 9 35; 3 99)
Ectromatopsis Compere, 1947: 13. Type-species: Ectroma americanum Howard, by monotypy.
INCLUDED SPECIES. One: americana (Howard, 18986: 248).
DISTRIBUTION. Mexico. World: Nearctic region.
BIOLOGY. Parasites of Pseudococcidae (Homoptera).
COMMENTS. Very probably best placed in the tribe Anagyrini.
NEOTROPICAL ENCYRTIDAE 1 95
ENCYRTUS Latreille
(Key couplets: 9 94; J 62, 73, 87. Figs 49, 136)
Encyrtus Latreille, 1809: 31. Type-species: Chrysis infida Rossi, by designation of Latreille (1810: 436).
Eucomys Forster, 1856: 32. Type-species: Encyrtus swederi Dalman, by original designation.
Comys Forster, 1856: 144. [Replacement name for Eucomys Forster.]
Howardia Dalla Torre, 1897: 86. Type-species: Bothriothorax peckhami Ashmead, by monotypy.
[Homonym of Howardia Berlese, 1896.]
Howardiella Dalla Torre, 1898: 228. [Replacement name for Howardia Dalla Torre.]
Allorhopoideus Brethes, 19166: 425. Type-species: Allorhopoideus mirabilis Brethes, by monotypy.
Prorhopoideus Brethes, 19216: 80. Type-species: Prorhopoideus baezi Brethes, by monotypy.
INCLUDED SPECIES. Ninety-five, most of which are probably incorrectly placed, eight from the
Neotropics: baezi (Brethes), bicolor (Howard), hay war di De Santis, infelix (Embleton, 1902:
223), lecaniorum (Mayr), littoralis (Blanchard), mexicanus (Girault, 19176: 21), vianaiDe Santis;
also a further two undetermined species from Venezuela and Brazil (BMNH).
REFERENCE. Revision of species from Argentina: De Santis (1964: 314-332).
DISTRIBUTION. Throughout the Neotropics. World: cosmopolitan.
BIOLOGY. Parasites of Coccidae (Homoptera).
COMMENTS. Placed in the tribe Encyrtini.
Encyrtus conformis Howard
(Key couplet: 9 180)
Encyrtus conformis Howard, 1897: 152.
I am unable to place this species, although it may be close to Exoristobia, but differs in the eyes
being naked, the scutellum much more convex and the propodeum relatively longer. For
redescription of species see Noyes (1979: 152-154).
Encyrtus quadricolor Howard
Encyrtus quadricolor Howard in Riley, Ashmead & Howard, 1894: 93.
Type lost. The description may refer to a male Rhytidothorax which has lost the conspicuous
setae on the frontovertex or perhaps a male Habrolepoidea. The species may be recognizable if
further material is collected in St Vincent, but until that time the name should be considered a
nomen dubium.
EPIENCYRTUS Ashmead
(Key couplets: 9 137, 241. Fig. 101)
Epiencyrtus Ashmead, 1900: 396. Type-species: Encyrtus thyreodontis Ashmead, by original designation.
INCLUDED SPECIES. Two, one an undetermined species from Panama (CNC, Ottawa).
DISTRIBUTION. Panama. World: Nearctic region.
BIOLOGY. Hyperparasite of other Encyrtidae parasitic in larvae of Lepidoptera.
COMMENTS. Placed in the tribe Cheiloneurini, subtribe Epiencyrtina. For key to related genera
see Trjapitzin & Gordh (19786).
ERICYDNUS Walker
(Key couplets: ? 116; J 67. Fig. 53)
Ericydnus Walker, 1837: 363. Ty pe- species : Ericydnus paludatus Walker, by designation of Westwood
(1840: 72).
196 JOHN S. NOYES
Grandoriella Domenichini, 1951: 171. Type-species: Grandoriella lamasi Domenichini, by monotypy.
Syn. n.
INCLUDED SPECIES. Thirteen, one of which is found in the Neotropics : lamasi (Domenichini,
1951: 173) (comb. n.).
REFERENCE. World revision: Kerrich (1967: 166-180).
DISTRIBUTION. Peru, Panama. World : cosmopolitan.
BIOLOGY. Parasites of Pseudococcidae (Homoptera).
COMMENTS. I do not consider the differences between Grandoriella and Ericydnus warrant
separate generic status. The Australian species belonging to this genus are intermediate between
the two genera.
Placed in the tribe Ericydnini (Grandoriella is placed in a separate tribe of its own by Trjapitzin
(19730), i.e. Grandoriellini).
EURYRHOPALUS Howard
(Key couplets: $ 142, 250; 3 17)
Euryrhopalus Howard, 18986: 237. Type-species: Euryrhopalus schwarzi Howard, by monotypy.
Synaspidia Timberlake, 1924: 397. Type-species: Synaspidia pretiosa Timberlake, by monotypy.
INCLUDED SPECIES. Nine, four of which are found in the Neotropics: kirkpatricki (Kerrich),
pretiosus (Timberlake), propinquus Kerrich, pulchrior Kerrich.
REFERENCE. World revision: Kerrich (1967: 235-246).
DISTRIBUTION. Mexico, Guatemala, Jamaica, Trinidad, Guyana, Panama, Colombia. World:
Nearctic region and Hawaii.
BIOLOGY. Parasites of Pseudococcidae (Homoptera).
COMMENTS. Placed in the tribe Neodiscodini.
EXORISTOBIA Ashmead
(Key couplets: 9 85, 204, 256. Figs 167-169)
Exoristobia Ashmead, 19046: 15. Type-species: Exoristobia philippinensis Ashmead, by monotypy.
INCLUDED SPECIES. Three, none of which is found in the Neotropics, but three undetermined
species from Trinidad and St Vincent (BMNH).
REFERENCE. Revision: Subba Rao (1970).
DISTRIBUTION. Trinidad, St Vincent. World: Oriental, Australian and Ethiopian regions.
BIOLOGY. Parasites of Syrphidae and Tachinidae (Diptera).
COMMENTS. Encyrtus conformis Howard may also belong to Exoristobia, but differs in having a
rather more convex scutellum and relatively longer propodeum. The species from the Neotropics
differ from the other known species in having relatively longer funicle segments.
I am unable to place the genus according to Trjapitzin's (1973&) classification of the
Encyrtidae.
FORCIPESTRIC1S Burks
(Key couplets: 9 227, 237, 239; 3 30, 91, 109. Figs 98, 99)
Fordpestricis Burks, 1968: 237. Type-species: Forcipestricis gaseaui Burks, by monotypy.
INCLUDED SPECIES. Three, two of which are found in the Neotropics: portoricensis Gordh (1975:
239), sordidus (Howard, 1897: 153); also at least a further 25 undetermined species from the
West Indies, Brazil and Ecuador (BMNH; CNC, Ottawa).
NEOTROPICAL ENCYRTIDAE 197
DISTRIBUTION. Puerto Rico, Brazil, Ecuador, West Indies. World: Nearctic region.
BIOLOGY. Parasites of Ceratopogonidae (Diptera).
COMMENTS. Apparently a large and taxonomically difficult genus. The presence of the tubercles
or pits on the scutellum (Figs 98, 99) seems fairly unique in the Encyrtidae, although they are
present in the probably unrelated genera Amaurilyma and Desobius. The pits in these two genera
are very difficult to see, even on slide mounted material, whereas those of Forcipestricis can be
seen, with practice, on dry mounted material. Useful characters for separating the species into
groups are : number of teeth in mandibles (bi- or tridentate), scutellum flat or convex, scutellum
with sculpture or smooth and distribution of pits i.e. grouped or scattered.
The genus cannot be placed according to Trjapitzin's (19736) classification of the Encyrtidae.
GAHANIELLA Timberlake
(Key couplets: 9 68, 79, 166, 196; j1 108. Fig. 37)
Gahaniella Timberlake, 1926: 23. Type-species: Gahaniella calif arnica Timberlake, by original designation.
INCLUDED SPECIES. Three, all from the Neotropics : californica Timberlake, saissetiae Timberlake
and tertia Kerrich.
REFERENCE. Kerrich (1953: 800-802).
DISTRIBUTION. Argentina, Brazil, Uruguay, Cuba, Trinidad, Virgin Islands. World: Nearctic
region.
BIOLOGY. Hyperparasites via other Encyrtidae in Coccidae and Pseudoccidae, possibly also
primary parasites of Coccidae (Homoptera).
COMMENTS. Placed in the tribe Microteryini by Trjapitzin & Gordh (19786).
GONZALEZIA De Santis
(Key couplets : 9 179, J1 90)
Gonzalezia De Santis, 1964: 292. Type-species: Gonzalezia gloriosa De Santis, by monotypy.
INCLUDED SPECIES. One: gloriosa De Santis, 1964: 294.
DISTRIBUTION. Argentina.
BIOLOGY. Unknown.
COMMENTS. Very similar to Cerchysius and Aseirba from which it can easily be separated using
the characters given in the key, notably the relatively small eyes.
I am unable to place the genus according to Trjapitzin's (19736) classification of the
Encyrtidae.
GRISSELLIA gen. n.
(Key couplets: 9 202, 247; 3 143. Figs 104, 110, 164-166)
Type-species: Grissellia terebrata sp. n.
?. Head. In side view moderately deep, most rounded level with top of antennal scrobes, fro nto vertex
slightly convex. Eye slightly longer than broad with short inconspicuous pale hairs and overreaching
occipital margin which is rounded. Malar space slightly more than half to three-fifths length of eye with
malar sulcus weakly indicated. Frontovertex, at narrowest point, about one-quarter of head width, ocelli
forming an acute angle of about 60 °, posterior ocellus separated from occipital margin by about one and a
half times to twice its own diameter and from eye margin by less than half its own diameter. Centre of
clypeus sometimes slightly produced downwards and outwards. Antennal scrobes moderately deep, fairly
long and joined dorsally, their uppermost point slightly more than half way between antennal toruli and
anterior ocellus, interantennal prominence dorsally rounded and about one and a half times as long as a
198 JOHN S. NOYES
torulus, antennal torulus separated from mouth margin by about half its own length and from other torulus
by about one and a half times its own length, dorsal margin slightly lower than lowest eye margin; scape
subcylindrical or slightly expanded and flattened in middle, pedicel conical, longer than first two funicle
segments together, funicle six-segmented with segments cylindrical and distinctly transverse to longer than
broad, gradually broadening apically, clava three-segmented with apex rounded, slightly broader than and
more than half as long as funicle; longitudinal sensilla on fifth and sixth funicle segments and clava.
Frontovertex with punctiform-reticulate sculpture between ocelli gradually becoming shallower and more
irregular towards top of antennal scrobes, cheeks with longitudinally elongate coriaceous sculpture but
quite smooth immediately below eyes, antennal scrobes smooth and polished. Mandibles with one or two
acute teeth and a truncation, maxillary palpi four-segmented, labial palpi three-segmented.
Thorax. Moderately deep in side view with mesoscutum and scutellum flat. Mesoscutum without
notaular lines, about one half broader than long, axillae separated by posterior margin of mesoscutum,
scutellum dorsally flat and slightly broader than long, propodeum centrally less than one-eighth as long as
scutellum. Pronotum, mesoscutum and axillae with coriaceous sculpture, scutellum with deep punctate-
reticulate sculpture, sides and extreme apex smooth, mesopleuron with shallow, irregularly elongate
sculpture, propodeum smooth. Forewing hyaline, slightly more than twice as long as broad; basal cell
sparsely hairy in distal half, naked proximally, speculum open and not interrupted ; submarginal vein with
an apical hyaline break, marginal vein about twice as long as broad, slightly longer than or about as long as
postmarginal and about half as long as stigmal. Hindwing about four times as long as broad. Mid tibial
spur nearly as long as mid basal tarsal segment.
Caster. About as long as thorax, cerci at about one-third, apex of hypopygium reaching to about one-half
to two-thirds along gaster, paratergites absent, epipygium occasionally with apex very acute and
overhanging base of ovipositor as in Pseudencyrtus, exserted part of ovipositor from one-seventh to one-
third as long as gaster. Ovipositor about one and a half to three times as long as mid tibia, gonostylus about
two-fifths to as long as mid tibia.
c?. Differs from female as follows: antennal scrobes nearly reaching anterior ocellus, antennae as in Fig.
165, frontovertex at narrowest point slightly less than one half of head width, scutellum slightly convex and
gaster shorter than thorax. The genitalia are unknown since the only available male is dry mounted.
Named in honour of Dr E. E. Grissell.
Grissellia terebrata sp. n.
(Figs 104, 110, 164, 166)
9- Length (excluding ovipositor): 1-24-1-43 mm (holotype 1-37 mm).
Colour. Head, thorax (except scutellum) and gaster generally dark brown with slight blue, green and
brassy reflections, antennal scrobes and cheeks with stronger purple or blue reflections. Antennae with
scape dark brown, paler apically and with some metallic green reflections, pedicel basally dark brown, apex
of pedicel and flagellum pale brown ; scutellum dark green, sculptured area matt and with some brassy
reflections, smoother areas shiny and tinged blue; legs with coxae, femora and tibiae brown with apices of
tibiae and tarsi yellow; basal tergite of gaster metallic blue-green, ovipositor sheaths dark brown.
Head. Centre of clypeus very slightly produced outwards and downwards; mandibles with two teeth and
a truncation (Fig. 104). Relative measurements of head of holotype: maximum head width 67, frontovertex
width at median ocellus 18, malar space 25, eye length 41, eye width 35, POL 8, OOL 2, scape length 30,
other proportions of antennae as in Fig. 164.
Thorax. Relative measurements of forewing of holotype: length 172, maximum width 76, length of
submarginal vein 70, of marginal vein 5, of stigmal vein 10, of postmarginal vein 5; of hindwing: length 1 17,
maximum width 29. Base of forewing as in Fig. 1 10.
Gaster. Slightly longer than thorax, epipygium very elongate and very acute, apex of hypopygium
reaching to about two-thirds along gaster, exserted part of ovipositor about one-third as long as gaster.
Relative measurement of paratype: ovipositor length 158, gonostylus length 62 [mid tibia length 59].
Ovipositor as in Fig. 166.
cJ. Unknown.
NEOTROPICAL ENCYRTIDAE 199
DISTRIBUTION. Brazil.
BIOLOGY. Unknown.
MATERIAL EXAMINED
Holotype 9. Brazil: Santa Catarina, Nova Teutonia, 15.x. 1949 (F. Plaumann) (BMNH).
Paratypes. Brazil: 2 ?, Nova Teutonia, 17.x. 1949, 20.x. 1949 (F. Plaumann) (BMNH).
COMMENTS. Two additional undescribed species from Brazil and St Vincent (BMNH), also an
undescribed species from India (BMNH).
The genus is very close to Echthrodryinus and Ooencyrtus (tribe Microteryini) but differs from
both in having a flat scutellum with punctate sculpture and exserted ovipositor, and from
Ooencyrtus in having a long postmarginal vein. In both Echthrodryinus and Ooencyrtus the
scutellum is rather evenly convex (except perhaps in Ooencyrtus chrysopae where the scutellum
appears to be flat, but this may be due to the scutellum collapsing as a result of the small size of
this species), and rarely with punctate sculpture, the ovipositor is usually hidden, although
sometimes slightly exserted, and in Ooencyrtus the postmarginal vein is short or absent.
GYRANUSOIDEA Compere
(Key couplets : 9 70; J 96)
Gyranusoidea Compere, 1947: 17. Type-species: Gyranusa citrina Compere, by monotypy.
INCLUDED SPECIES. Ten, none from the Neotropics, but two undetermined species from Brazil
and Trinidad (BMNH).
DISTRIBUTION. Brazil, Trinidad. World: Hawaii, Africa, India, Australia.
BIOLOGY. Parasites of Pseudococcidae (Homoptera).
COMMENTS. Placed in the tribe Anagyrini, subtribe Anagyrina. Close to Anagyrus and probably
should be considered synonymous. See 'Comments' under Anagyrus.
HABROLEPIS Forster
(Key couplets: 9 53; J 7. Fig. 32)
Habrolepis Forster, 1856: 34. Type-species: Encyrtus nubilipennis Walker, by original designation.
Gymnoneura Risbec, 1951 : 157. Type-species: Gymnoneura bambeyi Risbec, by monotypy.
INCLUDED SPECIES. Twenty-one, two of which are found in the Neotropics : dalmanii (Westwood),
rouxi Compere.
REFERENCE. Annecke & Mynhardt (1970: 128-146).
DISTRIBUTION. Chile, Bermuda. World: cosmopolitan.
BIOLOGY. Parasites of Diaspididae and Asterolecaniidae (Homoptera).
COMMENTS. Placed in the tribe Habrolepidini, subtribe Habrolepidina.
HABROLEPOIDEA Howard
(Key couplet: 9227)
Habrolepoidea Howard in Riley, Ashmead & Howard, 1894: 89. Type-species: Habrolepoidea glauca
Howard, by monotypy.
INCLUDED SPECIES. Six, two of which are found in the Neotropics: glauca Howard (Noyes, 1979:
157), celia Girault, 19176:22.
DISTRIBUTION. St Vincent.
BIOLOGY. Unknown.
200 JOHN S. NO YES
COMMENTS. The species described under Habrolepoidea by Girault from North America and
Australia are very probably incorrectly placed. Girault (1920: 187) synonymized Zarhopaloides
Girault with Habrolepoidea but from the description of the type-species this is evidently
incorrect. I have not seen the type of celia.
I am unable to place the genus satisfactorily according to Trjapitzin's (19736) classification of
the Encyrtidae, but it may belong to the Microteryini.
HAMBLETONIA Compere
(Key couplets: 9 51; J1 18. Figs 29, 31, 116, 118)
Hambletonia Compere, 1936: 172. Type-species: Hambletonia pseudococcina Compere, by monotypy.
INCLUDED SPECIES. One: pseudococcina Compere, 1936: 173.
DISTRIBUTION. Brazil, Argentina, Colombia, Puerto Rico, West Indies. World : Nearctic region,
Hawaii.
BIOLOGY. Parasites of Pseudococcidae (Homoptera).
COMMENTS. Placed in the tribe Chrysoplatycerini, subtribe Chrysoplatycerina (Trjapitzin &
Gordh, 19786: 648).
HELEGONATOPUS Perkins
(Key couplets: $ 176; $ 15, 118. Figs 114, 140)
Helegonatopus Perkins, 1906: 257. Type-species: Helegonatopus pseudophanes Perkins, by monotypy.
Schedioides Mercet, 1919a: 96. Type-species: Schedioides formosus Mercet, by monotypy.
Hazmburkia Hoffer, 1954: 172. Type-species: Hazmburkia dimorpha Hoffer, by monotypy.
Masencyrtus Hoffer, 1960: 98. Type-species: Masencyrtus concupiens Hoffer, by monotypy.
Paludencyrtus Hoffer, 1965: 16. Type-species: Paludencyrtus nikolskajae Hoffer, by monotypy.
INCLUDED SPECIES. Twelve, one of which is found in the Neotropics : pseudophanes Perkins, 1906 :
257, also one undescribed species from Trinidad (BMNH).
REFERENCE. Szelenyi (1972: 349-352).
DISTRIBUTION. Argentina, Trinidad, St Vincent, Jamaica. World: cosmopolitan.
BIOLOGY. Hyperparasites of Auchenorrhyncha (Homoptera) via Dryinidae (Hymenoptera).
COMMENTS. Placed in the tribe Chalcerinyini.
HELMECEPHALA gen. n.
(Key couplets: 9 81, 185; $ 112. Figs 88, 170-173)
Type-species: Helmecephala albisetosa sp. n.
9. Head. In profile moderately deep and with frontovertex and face evenly rounded, except for lower parts
of face (below ventral eye margin) which is flat. Eye slightly longer to one-third longer than broad, naked
and dorsally reaching occipital margin which is sharply marginal to base of mandibles. Malar space about
two-thirds length of eye and with malar sulcus absent. Frontovertex at narrowest point slightly more than
one-third head width, ocelli forming an acute angle of about 50-85 °, posterior ocellus separated from
occipital margin by about its own diameter and almost touching inner eye margin. Antennal scrobes
shallow and V-shaped, meeting dorsally, not reaching half way from antennal toruli to anterior ocellus and
shiny, antennal torulus separated from mouth margin and from other torulus by half its own length, its
dorsal margin below lowest level of eyes by about its own length; scape subcylindrical or slightly flattened
and expanded in middle, pedicel conical, longer than first funicle segment which is longer than broad,
funicle six-segmented with segments cylindrical, becoming shorter and broader distally so that distal
segments are distinctly transverse, clava three-segmented, longer than half length of funicle and with a long
oblique truncation; longitudinal sensilla on all flagellar segments. Frontovertex, face and cheeks with deep
thimble-punctured sculpture, punctures mostly separated by less than their own diameters and shiny
NEOTROPICAL ENCYRTIDAE 201
bottomed, punctures extending down interantennal prominence to clypeal margin, between punctures
rather irregular, raised shallow coriaceous to reticulate sculpture which is usually not very shiny and
contrasts rather strongly, in colour and shine, with punctures. Mandibles with two long, narrow, curved,
acute teeth and a truncation, occasionally upper tooth shorter and broader, maxillary palpi four-
segmented, the apical segment as long or longer than the first three together, labial palpi three-segmented.
Thorax. In side view deep with mesoscutum slightly convex and scutellum moderately convex. Pronotum
with strongly concave posterior margin, mesoscutum without notaular lines, less than a half broader than
long, axillae slightly separated by posterior margin of mesoscutum, scutellum slightly broader than long
and with apex rounded, propodeum short and with spiracles surrounded by dense, appressed, white setae.
Pronotum, mesoscutum and axillae with coriaceous sculpture, scutellum with fairly deep, irregular,
reticulate sculpture (distinctly deeper than that on mesoscutum) which becomes shallower and of larger
mesh towards sides and apex, mesopleura with raised longitudinally strigose sculpture, propodeum more or
less smooth but with irregular, shallow sculpture. Forewing hyaline, about two and a half times as long as
broad; basal cell completely pilose but with setae in middle very pale and inconspicuous, those towards
base and apex rather darker and stronger, costal cell with setae scattered over whole of dorsal surface,
speculum closed but not interrupted, extreme apex of wing with cilia absent, marginal cilia along anterior
margin very short; submarginal vein without an apical hyaline break, marginal vein about two to three
times as long as broad, about as long as stigmal vein and about twice as long as postmarginal. Hindwing
about three times as long as broad. Mid tibial spur about as long as mid basal tarsal segment.
Caster. Slightly shorter than thorax, cerci in basal one-third, paratergites absent, apex of hypopygium
reaching about halfway along gaster, ovipositor slightly protruding to exserted part, being about one-third
as long as gaster, and with sheaths slightly flattened. Ovipositor nearly one and a half times as long as mid
tibia, gonostylus about one-third as long as mid tibia.
J. Essentially same as female except antennal toruli are higher on head, i.e. separated from mouth margin
by about one and a half times to twice their own lengths and nearly level with lower eye margins ; antennae
as in Fig. 170 and genitalia as in Fig. 171 ; aedeagus slightly longer than mid tibial spur, each digitus with
one very strong spine.
Helmecephala albisetosa sp. n.
(Figs 88, 172, 173)
?. Length (excluding ovipositor): 1-70-1-90 mm (holotype 1-70 mm).
Colour. Head dark brownish purple and slightly shiny, thimble-punctures very shiny green, each with a
single, slightly flattened white hair arising from it, scape yellowish, pedicel dark brown, yellowish at apex
and ventrally, funicle testaceous yellow, dorsally and distally darker, clava and sixth funicle segment
brown ; pronotum and mesoscutum dark brown with a strong purple shine, mesoscutum covered (except
anteriorly) with numerous appressed, white setae, scutellum dark green, purplish in centre, with some bluish
reflections, mesopleuron mixed shining blue and purple, propodeum shiny, metallic blue medially ; coxae,
femora and hind tibia black, knees testaceous, fore and mid tibiae proximally dark brown, gradually
becoming testaceous distally, tarsi yellowish white gradually darkening to testaceous distally, mid tibial
spur white, brown at apex ; gaster black, basal tergite shining greenish blue mixed with some purple.
Head. Ocelli forming an angle at about 75-80°. Relative measurements of head of holotype: head width
99, frontovertex width at median ocellus 36, malar space 45, eye length 61, eye width 51, POL 26, OOL 2,
scape length 48, other proportions of antennae as in Fig. 88. There is some variation in the relative length of
the scape, from slightly less than to slightly longer than malar space and therefore scape may vary from
about four and a half to five times as long as broad.
Thorax. Relative measurements of forewing of holotype: length 204, maximum width 84, length of
submarginal vein 85, of marginal vein 10, of stigmal vein 10, of postmarginal vein 5. Base of forewing as in
Fig. 172.
Gaster. Exserted part of ovipositor slightly less than one-fifth of gaster or half of mid tibial spur. Relative
measurements of gaster of paratype: length of ovipositor 136, of gonostylus 30 [mid tibia length 106].
Genitalia as in Fig. 173.
3. Unknown.
202 JOHN s- NOYES
DISTRIBUTION. Trinidad.
BIOLOGY. Unknown.
MATERIAL EXAMINED
Holotype ,, Trinidad: Nariva, Ecclesville, 29.vii.1976 (J. S. Noyes) (BMNH).
Paratypes. Trinidad: 1 9, St Andrew, Oropuche, 28.vi.1976; 1 ?, St George, Aripo Valley, 4.viii.l976; 1 9,
St Patrick, Bonasse, 12.viii.1976; 2 9, St Patrick, Coora, 14.viii.1976; 1 ?, St George, Tumpuna Reserve,
9.viii.l976 (J. S. Noyes); 1 9, Curepe, i.1978 (F. D. Bennett). (BMNH; CNC, Ottawa; MLP, Argentina.)
COMMENTS. Three further undescribed species from Trinidad (BMNH ; CNC, Ottawa) differ in
sculpture of frontovertex, coloration of setae on mesoscutum, proportions of antennal segments
and relative length of exserted part of ovipositor.
The shape of the head suggests that the nearest relatives to Helmecephala are the genera placed
in the tribe Prionomasticini (Trjapitzin & Gordh, 19786). This is very probably true, but
Helmecephala appears to be much less primitive than any of the genera included in this tribe. The
cerci are well advanced on the gaster and the dentition of the mandibles is more developed. The
venation is also very different, the marginal vein being relatively long and the postmarginal and
stigmal veins relatively short. The antennae of the male are also considerably different from
those of the males of this tribe.
HEMAENASWS Ashmead
(Key couplets: ? 63, 144, S 92. Figs 51, 59)
Hemaenasius Ashmead, 1900: 374. Type-species: Hemaenasius confusus Ashmead, by monotypy.
Epaenasomyia Girault, 1917c: 3. Type-species: Epaenasomyia varicornis Girault, by monotypy.
INCLUDED SPECIES. Two, one of which is found in the Neotropics : confusus Ashmead, 1900: 374;
also one undetermined species from Panama (CNC, Ottawa).
DISTRIBUTION. Brazil, Panama. World: Nearctic region.
BIOLOGY. Unknown.
COMMENTS. Placed in the tribe Discodini, subtribe Hemaenasiina by Trjapitzin & Gordh (19786).
HEMENCYRTUS Ashmead
(Key couplets: 9 105, 120, 133, 153, 164, 180, 185, 189; S 104, 108, 112. Fig. 60)
Hemencyrtus Ashmead, 1900: 374. Type-species: Hemencyrtus Herbert ii Ashmead, by monotypy.
INCLUDED SPECIES. Three : brasiliensis (Ashmead) (comb. n. from Bothriothorax), casali De Santis,
19670 : 69 and Herbert ii Ashmead, 1900 : 375 ; also a further five or six undetermined species from
Trinidad, Brazil, Panama and Ecuador (BMNH; CNC, Ottawa).
DISTRIBUTION. Argentina, Brazil, Ecuador, Panama, Trinidad.
BIOLOGY. Unknown.
COMMENTS. I have examined three syntype females of Hemencyrtus herbertii. The one which fits
the description best (labelled: 'Aug., Chapada, H. H. Smith coll, Type No. 41381 U.S.N.M.',
here designated LECTOTYPE) is lacking wings ; of the other two, one is in good condition but
fits the description poorly and the other is lacking antennae and one pair of wings. All are
congeneric but may not be conspecific.
Bothriothorax brasiliensis (holotype j examined) is best placed in Hemencyrtus although it has
some characteristics in common with Zeteticontus.
The species included in this genus appear to fall into two distinct groups: those where the
scutellum is very flat and with a distinct translucent apical flange (includes herbertii and casali)
and those where the scutellum is convex and the flange is distinct or absent (includes brasiliensis).
The genus belongs to the tribe Bothriothoracini.
NEOTROPICAL ENCYRTIDAE 203
HEX ACL ADI A Ashmead
(Key couplets : 9 112; ^ 5)
Hexacladia Ashmead, 1891 : 456. Type-species: Hexacladia smithii Ashmead, by monotypy.
Thysanomastix Perkins, 1907: 55. Type-species: Thysanomastix koebelei Perkins, by monotypy.
Sophencyrtus Crawford, 1911 : 275. Type-species: Sophencyrtus townsendi Crawford, by monotypy.
INCLUDED SPECIES. Seven, four of which are found in the Neotropics : blanchardi De Santis,
mexicana Girault, smithii Ashmead, townsendi Crawford; also a further five undescribed species
(BMNH; CNC, Ottawa).
REFERENCE. Revision: Burks (1972).
DISTRIBUTION. Argentina, Brazil, Ecuador, Trinidad. World: Nearctic region.
BIOLOGY. Parasites of adult Scutelleridae and Pentatomidae (Heteroptera).
COMMENTS. Placed in the tribe Prionomasticini, subtribe Hexacladiina.
HOMALOPODA Howard
(Key couplets: 9 4; j* ?7)
Homalopoda Howard in Riley, Ashmead & Howard, 1894: 91. Type-species: Homalopoda cristata Howard,
by monotypy.
INCLUDED SPECIES. One: cristata Howard (Noyes, 1979: 157); also a further four undetermined
species (BMNH; CNC, Ottawa; USNM, Washington).
DISTRIBUTION. Trinidad, St Vincent, Panama.
BIOLOGY. Unknown, but probably parasites of Diaspididae (Homoptera).
COMMENTS. Placed in the tribe Habrolepidini, subtribe Habrolepidina.
HOMALOTYLUS Mayr
Homalotylus Mayr, 1876: 752. Type-species: Encyrtus flaminius Dalman, by designation of Ashmead
(1900:377).
Nobrimus Thomson, 1876: 116. Type-species: Encyrtus flaminius Dalman, by designation of Timberlake
(1919a: 134).
Mendozaniella Brethes, 1913: 97. Type-species: Mendozaniella mirabilis Brethes, by monotypy.
Hemaenasoidea Girault, 191 60: 307. Type-species: Hemaenasoidea oculata Girault, by monotypy.
Lepidaphycus Blanchard, 1936: 13. Type-species: Lepidaphycus bosqi Blanchard, by monotypy.
Neoaenasioidea Agarwal, 1966: 71. Type-species: Neoaenasioidea indica Agarwal, by monotypy.
INCLUDED SPECIES. Twenty-eight, seven of which are found in the Neotropics: brevicauda
Timberlake, cockerelli Timberlake, flaminius (Dalman), latipes Girault, mexicanus Timberlake,
mirabilis (Brethes), terminalis (Say) ; also a further two undetermined species from Panama and
Surinam (BMNH; CNC, Ottawa; CIBC, Trinidad).
REFERENCE. Revision: Timberlake (1919a: 133-170).
DISTRIBUTION. Throughout the Neotropics. World : cosmopolitan.
BIOLOGY. Parasites of coccinellid larvae (Coleoptera).
COMMENTS. Placed in the tribe Homalotylini, subtribe Homalotylina.
HOMOSEMION Annecke
(Key couplet: ? 49. Fig. 23)
Homosemion Annecke, 1967: 119. Type-species: Homosemion bennetti Annecke, by monotypy.
INCLUDED SPECIES. One: bennetti Annecke, 1967: 157.
9
204 JOHN S. NO YES
DISTRIBUTION. Trinidad.
BIOLOGY. Parasites of Margarodidae (Homoptera).
COMMENTS. Placed in the tribe Cerapterocerini.
HOPLOPSIS Destefani
(Key couplet: 9 126)
Hoplopsis Destefani, 1889: 140. Type-species: Hoplopsis mayri Destefani, by monotypy.
INCLUDED SPECIES. Two, one of which is found in the Neotropics : cristulata De Santis, 1 9726 : 56.
DISTRIBUTION. Argentina, Brazil. World: Europe.
BIOLOGY. Unknown.
COMMENTS. Placed in the tribe Microteryini, subtribe Microteryina.
HUNTERELLUS Howard
(Key couplets: $ 62, 83; ^ 71. Figs 34, 132)
Hunterellus Howard, 1908: 241. Type-species: Hunterellus hookeri Howard, by monotypy.
INCLUDED SPECIES. Three, one of which is found in the Neotropics : hookeri Howard, 1908 : 241 ;
also one undetermined species from Trinidad (BMNH).
DISTRIBUTION. Brazil, Trinidad, Puerto Rico, Cuba, Mexico. World: cosmopolitan.
BIOLOGY. Parasites of nymphs of Ixodidae (Acarina).
COMMENTS. Placed in the tribe Ixodiphagini.
ICEROMYIA gen. n.
(Key couplets: $ 57, 78; <? 103, 135. Figs 38, 44, 174-176)
Type-species: Iceromyia flavifrons sp. n.
$. Head. In profile, moderately deep, most rounded level with top of antennal scrobes, fronto vertex slightly
convex. Eye nearly as broad as long, naked and reaching occipital margin which is sharp. Malar space
slightly more than half length of eye, malar sulcus absent. Frontovertex, at narrowest, about one-eighth
head width or twice diameter of anterior ocellus, ocelli forming an acute angle, posterior ocellus separated
from occipital margin by about its own diameter and more or less touching eye margin. Antennal scrobes
fairly deeply impressed, meeting dorsally and reaching about half way between toruli and anterior ocellus,
antennal torulus separated from mouth margin by less than its own length and from other torulus by about
one and a half times its length, its dorsal margin slightly below lowest margin of eye. Scape broadened and
flattened, pedicel conical and about half length of funicle, funicle six-segmented, all segments transverse,
lengthening and broadening distally, clava very large, three-segmented, longer than funicle and with a
strong, apical oblique truncation; longitudinal sensilla on the apical two or three funicle segments and
clava. Frontovertex with shallow coriaceous sculpture. Mandibles with two acute teeth and a truncation,
maxillary palpi four-segmented, labial palpi three-segmented.
Thorax. In side view, moderately deep and with mesoscutum and scutellum slightly convex. Mesoscutum
without notaular lines, about a half broader than long, axillae more or less touching, scutellum about as
long as broad and without apical tuft of setae but with two to six conspicuous longer setae, propodeum
medially at most one-seventh as long as scutellum. Pronotum, mesoscutum and axillae with shallow
coriaceous sculpture, scutellum with quite deep, longitudinally strigose sculpture tending towards reticulate
in centre, mesopleuron with shallow, raised slightly elongate or strigose sculpture, propodeum smooth
medially. Forewing hyaline or very slightly infumate in a small triangular area about half way along
submarginal vein and across wing from marginal vein and extending to wing apex, becoming slightly paler
distally, with a hyaline streak from apex of postmarginal vein to apex of stigmal vein, forewing about two
and a half times as long as broad ; basal cell more or less naked, but with a few transverse lines of setae
NEOTROPICAL ENCYRTIDAE 205
distally, speculum not interrupted and open; submarginal vein with an indistinct apical hyaline break,
marginal vein slightly more than three times as long as broad, slightly longer than stigmal and about twice
as long as postmarginal. Hindwing about four and a half times as long as broad. Mid tibial spur about as
long as mid basal tarsal segment.
Caster. Slightly shorter than thorax. Cerci at about one-third, paratergites absent, apex of hypopygium
reaching about half way along gaster, exserted part of ovipositor about half as long as gaster. Ovipositor
(Fig. 176) more than one and a half times as long as gaster or about twice as long as mid tibia, gonostylus
about as long as mid tibia.
J1. Differs from female as follows: head with antennal scrobes shallower and longer, reaching further
than half way between toruli and anterior ocellus, antennal torulus higher on head, separated from mouth
margin by more than its own length, its upper margin slightly below lowest level of eye ; malar sulcus
present; posterior ocellus separated from occipital margin by less than half its diameter, ocelli forming an
angle of about 60°, fronto vertex nearly one-third head width; antennae as in Fig. 174; sculpture of
frontovertex rugose and partly reticulate between ocelli, gradually becoming shallower and coriaceous on
lower parts of face and longitudinally strigose on cheeks ; forewing completely hyaline with postmarginal
vein about as long as stigmal; aedeagus about half as long as mid tibia.
Iceromyia flavifrons sp. n.
fFi.gs 38, 44, 174, 175)
$. Length (excluding ovipositor): 1-85 mm.
Colour. Head yellow except for vertex between and behind ocelli and a line across clypeus and lower parts
of genae which are dark brown, cheeks with conspicuous, long, dark setae ; antenna, except for clava,
yellowish orange, clava dark brown; pronotum, axillae and mesoscutum brown with purple, blue, green
and brassy reflections, the reflected colour on the mesoscutum depending on angle viewed from ; scutellum
yellowish orange, more orange apically, propodeum dark orange medially, dark brown laterally,
mesopleuron dark brown with purple and blue reflections, mesoscutum with sparse pale brown setae ; legs
yellow; forewings (Fig. 38) with basal area hyaline except for a small, triangular infuscate spot about half
way along and below submarginal vein, wing slightly infumate from level with apex of submarginal vein to
apex, becoming paler distally ; gaster dark brown, basal tergite with strong metallic green and purple sheen,
ovipositor sheaths yellowish orange.
Head. Antennal scrobes more or less semicircular. Relative measurements of head: head width 104,
frontovertex width at median ocellus 15-5, malar space 42, eye length 70, eye width 66, POL 8-5, OOL 0-5,
scape length 51, other proportions of antenna as in Fig. 44.
Thorax. Mesopleuron with shallow reticulate sculpture, anterior half strigose. Relative measurements of
forewing (Fig. 38): length 248, maximum width 97, other proportions as in Fig. 48.
Gaster. Relative measurements : length of exserted part of ovipositor 63, length of gaster 125, length from
base of gaster to apex of hypopygium 67.
c£. Length: 1-56 mm.
Colour. Head and thorax black with weak metallic purple, green and brassy reflections; lower parts of
face strongly metallic green ; antennae yellow except clava which is pale brown ; mesopleuron with distinct
purple coloration ; coxae dark brown, remainder of legs yellow ; gaster dark brown. Relative measurements ;
head : width 92, frontovertex width at median ocellus 32, malar space 32, eye length 59, eye width 45, POL
13, OOL 2, scape length 38, other proportions of antennae as in Fig. 174; forewing: length 212, maximum
width 99, length of submarginal vein 85, of marginal vein 10, other proportions as in Fig. 175.
DISTRIBUTION. Trinidad.
BIOLOGY. Reared from Margarodidae (Homoptera) (probably hyperparasitic on other
Encyrtidae).
MATERIAL EXAMINED
Holotype 2, Trinidad: iv.1963, ex Icerya montserratensis Riley & Howard (F. D. Bennett) (BMNH).
Paratype. 1 3, same data as holotype (BMNH).
206 JOHN s- NOYES
COMMENTS. Also a further undescribed species from Guyana reared from the same host as the
type-species (BMNH).
Closely related to Cheiloneurus (tribe Cheiloneurini) but can be separated using the characters
given in the key.
ISODROMUS Howard
(Key couplets: $ 140, 217; j1 48. Figs 58, 124)
Isodromus Howard, 1887: 488. Type-species: Isodromus iceryae Howard, by monotypy.
Parataneostigma Girault, 1915a: 275. Type-species: Parataneostigma nigriaxillae Girault, by monotypy.
INCLUDED SPECIES. Fourteen, one of which is found in the Neotropics : iceryae Howard ; also a
further undetermined species from Peru (BMNH).
REFERENCE. Revision: Timberlake (\9\9a: 176-190).
DISTRIBUTION. Argentina, Brazil, Chile, Peru, Mexico, Cuba. World: cosmopolitan.
BIOLOGY. Parasites of Chrysopidae and Hemerobiidae (Neuroptera).
COMMENTS. Placed in the tribe Homalotylini, subtribe Homalotylina.
LEPTOMASTIDEA Mercet
(Key couplets:? 113; 3 98)
Leptomastidea Mercet, 1916: 112. Type-species: Leptomastidea aurantiaca Mercet, by monotypy.
Tanaomastix Timberlake, 1918: 362. Type-species: Paraleptomastix abnormis Girault, by original
designation.
Leptanusia De Santis, 1964: 80. Type-species: Leptomastidea pseudococci Brethes, by monotypy. Syn. n.
INCLUDED SPECIES. Seventeen, five of which are found in the Neotropics: abnormis Girault,
antillicola Dozier, 1937: 121, bahiensis (Compere) (comb. n. from Leptomastix), dispar (Kerrich)
(comb. n. from Leptomastix) and pseudococci Brethes (De Santis, 1964: 82).
REFERENCES. Mercet (1924: 252-258), Kerrich (1953: 790-791).
DISTRIBUTION. Throughout the Neotropics. World: cosmopolitan.
BIOLOGY. Parasites of Pseudococcidae (Homoptera).
COMMENTS. I have examined the holotype of Leptomastidea brethesi Blanchard which was
synonymized with Leptomastidea pseudococci Brethes, the type-species of Leptanusia, by De
Santis (1964: 82). It is a Leptomastidea belonging to the same group as bahiensis and dispar. It
differs from all other species of Leptomastidea in having relatively shorter marginal cilia on the
hind wing, but I do not think that this difference and the differences given by Trjapitzin (1971c:
86) warrant separate generic status. Kerrich (pers. comm.) suggested that bahiensis belongs to
Leptomastidea.
The holotype male of antillicola Dozier has not been seen, but the species may be correctly
placed in Leptomastidea.
Placed in the tribe Anagyrina, subtribe Leptomastideina.
LEPTOMASTIX Forster
(Key couplets: 2 119, 157; ^ 99)
Leptomastix Forster, 1856: 34. Type-species: Leptomastix histrio Mayr, by monotypy through reference of
Mayr(1876: 730).
Stenoterys Thomson, 1876: 115. Type-species: Stenoterys orbitalis Thomson, by monotypy.
Pseudleptomastix Girault, 1915a: 272. Type-species: Pseudleptomastix squammulatus Girault, by
monotypy.
NEOTROPICAL ENCYRTIDAE 207
INCLUDED SPECIES. Thirty-seven, one of which is found in the Neotropics : dactylopii Howard
(Dozier, 1927: 267), also one species from Mexico, near squammulata Girault, and a further
species from Brazil (BMNH).
DISTRIBUTION. Throughout the Neotropics. World: cosmopolitan.
BIOLOGY. Parasites of Pseudococcidae (Homoptera).
COMMENTS. The undetermined species from Brazil apparently has a seven-segmented funicle and
two-segmented clava as in Alamella Agarwal, but since the venation is more typical of
Leptomastix I include it in the latter.
Placed in the tribe Anagyrini, subtribe Anagyrina. See comments under Anagyrus.
L1RENCYRTUS gen. n.
(Key couplets: 9 162; 3 124. Figs 74, 177-181)
Type-species : Lirencyrtus primus sp. n.
$. Head. In side view moderately deep with frontovertex and face more or less uniformly curved but mostly
so level with top of antennal scrobes. Eye about one-third longer than broad, with numerous short hairs
and reaching occipital margin which is sharp. Malar space from much less than to slightly more than half
length of eye with malar sulcus absent or present. Frontovertex at narrowest point about one-third head
width and with fairly conspicuous dark setae; ocelli more or less forming a right angle, posterior ocellus
separated from occipital margin by less than its own diameter and from eye margin by about half its own
diameter. Antennal scrobes moderately impressed and semicircular, meeting dorsally but quite short, not
quite reaching half way from toruli to anterior ocellus ; antennal torulus separated from mouth margin by
slightly more than its own length and from other torulus by about one and a half times its own length, its
dorsal margin about level with lowest margin of eye. Antenna with scape subcylindrical, pedicel conical and
about as long as first funicle segment, all six funicle segments cylindrical and longer than broad, clava three-
segmented, about half as long as and slightly broader than the funicle and with apex rounded, pointed or
with a very short oblique truncation; longitudinal sensilla on all flagellar segments. Sculpture on-
frontovertex shallow coriaceous, sometimes very smooth or occasionally rougher behind ocelli, on lower
parts of face becoming more elongate and shallower, cheeks almost smooth. Mandibles with three acute
teeth, maxillary palpi four-segmented, labial palpi three-segmented.
Thorax. In side view moderately deep and dorsally convex, scutellum very convex. Dorsum of thorax
with strong dark setae. Mesoscutum without notaular lines, about one-half broader than long, axillae
touching, scutellum about as broad as long, propodeum medially relatively long, more than one-sixth as
long as scutellum. Pronotum with fairly deep coriaceous sculpture, mesoscutum with coriaceous reticulate,
reticulate-strigose, coriaceous-strigose or strigose sculpture, axillae dorsally with alutaceous sculpture,
posterior margin steep and polished, scutellum anteriorly with numerous longitudinal ridges which run
backwards from axillae, these sometimes short, but often about half length of scutellum, usually some
sculpture between ridges which may extend nearly to apex but at least apex and usually apical one-third to
half of scutellum very smooth and polished ; mesopleuron more or less smooth but with some shallow,
irregular sculpture anteriorly ; propodeum usually with some sculpture in centre and along anterior margin
from either side of middle third nearly to spiracles. Forewing hyaline, about two and a half times as long as
broad ; basal cell completely naked except for a few long setae in its apical half and a line below submarginal
vein, speculum not interrupted and closed ; submarginal vein with an apical hyaline break, marginal vein
about three to four times as long as broad, about a half longer than stigmal which is slightly longer than
postmarginal. Hindwing about four times as long as broad. Mid tibial spur about as long as mid basal
tarsal segment.
Caster. Slightly shorter than thorax, cerci at about one-third, paratergites absent, ovipositor not visible in
dry mounted material, hypopygium reaching apex of gaster and medially incised at apex. Ovipositor short,
less than half as long as mid tibia, gonostylus about one-quarter as long as mid tibial spur.
31. Differs from female as follows: antennae with pedicel distinctly shorter than first funicle segment, all
funicle segments longer than broad, clava entire, funicle dorsally with two or three groups of very long setae
about three to four times as long as diameter of corresponding segment, ventrally setae not much longer
than diameter of segment ; aedeagus about two-fifths as long as mid tibia and digitus with one apical spine.
208 JOHN s- NO YES
Lirencyrtus primus sp. n.
(Figs 74, 177-181)
$. Length: 1-22-1-44 mm (holotype 1-35 mm).
Colour. Frontovertex dark metallic green, purple along eye margins and below level of top of antennal
toruli, scape testaceous, pedicel and flagellum brown; pronotum purplish brown, mesoscutum metallic
green usually with purple reflections along posterior and anterior margins, sculptured area of scutellum
purple, polished area green with some purple reflections, propodeum brown ; foreleg, mid tarsus, mid tibial
spur and hind coxae yellow to yellowish orange, mid coxae brown, mid femora in basal half, mid tibia in
apical two-thirds, hind femora in basal third and hind tibia in apical half yellow to yellowish orange,
remainder dark brown, but occasionally hind femora almost completely brown or mid femora, tibiae and
hind tarsi almost completely yellow ; gaster dark purplish brown with basal tergite metallic green with some
purple reflections.
Head. Upper margin of antennal torulus about level with lowest margin of eye, vertex between posterior
ocelli and occipital margin quite smooth and polished, malar sulcus absent, clava with apex more or less
pointed. Relative measurements of head of holotype: head width 70, frontovertex width at median ocellus
25, malar space 14, eye length 43, eye width 34, POL 12, OOL 2-5, scape length 35, other proportions of
antenna as in Fig. 177.
Thorax. Mesoscutum with coriaceous sculpture, of smaller mesh centrally, more transversely elongate
laterally, scutellum with slightly more than apical half smooth and polished and with about seven to eleven
strong ridges running backwards from axillae (Fig. 74) for a distance up to about half medial length of
scutellum, propodeum with some very shallow sculpture in middle and along anterior margin either side of
middle third almost to spiracles, anterior margin with a transverse shallow trough which extends almost to
spiracle, the trough traversed by several carinae. Forewing (Fig. 178) with marginal vein variable in
thickness, from three to four times as long as broad. Relative measurements of forewing of holotype : length
207, maximum width 83, length of submarginal vein 78, of marginal vein 13-5, of stigmal vein 10, of
postmarginal vein 8-5; of hindwing: length 143, maximum width 37.
Gaster. Relative measurements of paratype : length of ovipositor 34, of gonostylus 5 [mid tibia 78, mid
tibial spur 20]. Ovipositor as in Fig. 180.
cJ. Length: 1-11-1-40 mm.
Similar to female except the frontovertex, which is very slightly broader, antennae (Fig. 179) and
genitalia (Fig. 181).
DISTRIBUTION. Brazil.
BIOLOGY. Unknown.
MATERIAL EXAMINED
Holotype ?, Brazil: Santa Catarina, Nova Teutonia, 9.vii.l943 (F. Plaumanri) (BMNH).
Paratypes. Brazil: 1 ?, Nova Teutonia, 8.viii.l935; 2 ?, 2 <£, Nova Teutonia, 18.V.1943; 3 $, 1 cJ, Nova
Teutonia, vi.1943; 3 $, Nova Teutonia, vii.1943; 1 ?, 1 <$, Nova Teutonia, ix.1943; 1 9, Nova Teutonia,
lO.x.1943; 1 cJ, Nova Teutonia, xi.1949; 1 ?, Nova Teutonia, vi.1971 (F. Plaumann). (BMNH; CNC,
Ottawa; MLP, Argentina; CIBC, Trinidad; UC, Riverside; PPRI, Pretoria; ZI, Leningrad).
COMMENTS. I have examined a further nine undescribed species from Brazil, Trinidad, Jamaica,
Dominica and Panama (BMNH; CNC, Ottawa; USNM, Washington).
The genus appears to be close to Helegonatopus (tribe Chalcerinyini), but the female mainly
differs in having a short ovipositor, very short gonostyli, anterior ridges on the scutellum and the
hypopygium reaching the apex of the gaster. Helegonatopus has the ovipositor much longer than
the mid tibia, the gonostyli about twice as long as the mid tibial spur, no anterior ridges on the
scutellum, and the hypopygium reaching at most about two-thirds along the gaster.
LITOMASTIX Thomson
(Key couplets: ? 200, 218; ^ 49, 51. Figs 96, 122)
Litomastix Thomson, 1876: 171. Type-species: Encyrtus chalconotus Dalman, by designation of Ashmead
(1900: 363).
NEOTROPICAL ENCYRTIDAE 209
Pentacnemus Howard, 1892: 366. Type-species: Pentacnemus bucculatricis Howard, by monotypy.
Berecyntus Howard, 18986: 237. Type-species: Berecyntus bakeri Howard, by monotypy.
Paracopidosomopsis Girault, \9\6d: 49. Type-species: Berecyntus floridanus Ashmead, by monotypy.
Litomastiellus Mercet, 1921: 443. Type-species: Litomastix claviger Mercet, by designation of Peck in
Muesebeck et al. (1951 : 481).
Limastotix Mercet, 1921 : 443. Type-species: Litomastix hispanicus Mercet, by monotypy.
Berecyntiscus Ghesquiere, 1946: 368. [ Unnecessary replacement name for Berecyntus Howard.]
INCLUDED SPECIES. Thirty-six, three of which are found in the Neotropics: calypso (Crawford,
1914: 88) (comb. n. from Holcencyrtus), floridana Ashmead, 1900: 365, truncatella
(Dalman) ( = aestivalis Mercet, 1921 : 447); also a further two undescribed species from Ecuador
and Trinidad (BMNH; CNC, Ottawa).
DISTRIBUTION. Throughout the Neotropics. World : cosmopolitan.
BIOLOGY. Polyembryonic parasites of larvae of Lepidoptera.
COMMENTS. It is very likely that calypso and floridana are the same species as truncatella. The two
undetermined species from Trinidad and Ecuador are very distinct from this species and belong
to the thebe-group.
Placed in the tribe Copidosomatini, subtribe Copidosomatina. See comments under
Copidosoma.
LOCHITOENCYRTUS De Santis
(Key couplets: ? 184; J* 25. Fig. 86)
Lochitoencyrtus De Santis, 1964: 264. Type-species: Lochitoencyrtus gahani De Santis, by monotypy.
INCLUDED SPECIES. One: gahani De Santis, 1964: 265.
DISTRIBUTION. Argentina, Brazil, Trinidad.
BIOLOGY. Unknown.
COMMENTS. Placed in the tribe Trechnitini by Trjapitzin (19736); this is very probably wrong.
The genus is similar in appearance to Aminellus Masi (tribe Bothriothoracini, subtribe
Aminellina) but differs in the head being relatively more deep (i.e. less lenticular), the stigmal
vein straight, the punctures on the head shiny bottomed and the mesoscutum and scutellum
lacking deep piliferous punctures. Even though it superficially resembles Aminellus I do not think
that the genus belongs in the tribe Bothriothoracini. I am unable to satisfactorily place it
according to Trjapitzin's classification of the Encyrtidae.
LOH1ELLA gen. n.
(Key couplet: 9 165. Figs 182, 183)
Type-species: Encyrtus flaviclavus Howard in Riley, Ashmead & Howard, 1894: 96.
?. Head. Moderately deep inside view and more or less abruptly bent at level of top of antennal scrobes. Eye
about one-third longer than broad, with numerous short hairs and reaching occipital margin which is sharp.
Malar space about half as long as eye and with malar sulcus absent. Frontovertex at narrowest point
slightly less than one-quarter head width; ocelli forming a slightly obtuse angle, posterior ocellus separated
from occipital margin by slightly less than its own diameter and from eye margin by less than half its own
diameter. Antennal scrobes shallow but meeting dorsally, reaching about halfway from antennal toruli to
anterior ocellus, antennal torulus separated from mouth margin by about one and a half times its own
length and from other torulus by about its own length, torulus high on head, its ventral margin only very
slightly below ventral margin of eye. Antennal scape subcylindrical, pedicel conical and subequal in length
to all six funicle segments which are all slightly longer than broad and cylindrical, clava three-segmented,
about as wide as funicle and with apex rounded (Fig. 182); longitudinal sensilla on all flagellar segments.
Sculpture on frontovertex shallow, shagreened, more elongate between eyes and antennal scrobes, and on
genae. Mandibles with two acute teeth and a truncation, maxillary palpi four-segmented, labial palpi three-
segmented.
210 JOHN S. NOYES
Thorax. In side view moderately deep, mesoscutum convex and scutellum almost flat. Mesoscutum
without notaular lines and about a half broader than long, axillae slightly separated by posterior margin of
mesoscutum, scutellum about as broad as long, propodeum medially about one-sixth as long as scutellum.
Pronotum, mesoscutum and axillae with similar sculpture to frontovertex but slightly deeper, laterally and
on axillae more transversely elongate, scutellum with deeper, raised, elongate sculpture more or less
arranged in whorls which is similar in appearance to a fingerprint, apex and sides of scutellum entirely
smooth, mid pleuron almost smooth but with raised elongate sculpture, propodeum medially with some
shallow sculpture. Forewing (Fig. 183) hyaline and about two and a half times as long as broad; basal cell
naked in its proximal half, speculum not interrupted and open ; submarginal vein with apical hyaline break,
marginal vein distinctly longer than broad, longer than radial which in turn is longer than postmarginal.
Hindwing about three and a half times as long as broad. Mid tibial spur slightly longer than mid basal tarsal
segment.
Caster. Slightly shorter than thorax with cerci at about one-third, apex of hypopygium slightly more than
half way along gaster, ovipositor only slightly exserted.
J. Unknown.
INCLUDED SPECIES. One: flavic lava (Howard) (Noyes, 1979: 154).
DISTRIBUTION. St Vincent, Trinidad.
BIOLOGY. Unknown.
COMMENTS. The species previously undetermined by Noyes (1979: 154) is merely a form of
flaviclava; the mandibles have two acute teeth and a truncation, not three acute teeth as stated.
This genus bears a superficial resemblance to some species of Rhytidothorax, but undoubtedly
belongs to the tribe Microteryini and very probably the subtribe Microteryina.
MARIO LA gen. n.
(Key couplet: ? 75. Figs 42, 184, 185)
Type-species: Mariolaflava sp. n.
?. Body generally dorso-ventrally flattened (Fig. 42).
Head. Occipital foramen high on head at about four-fifths head length. Eye about two and a half to three
times as long as broad, naked and very nearly reaching occipital margin which is sharp. Malar space about
one-fifth length of eye, with malar sulcus absent. Frontovertex at narrowest point slightly less than half
head width, inner eye margins slightly concave but almost parallel, ocelli more or less forming a right angle,
posterior ocellus separated from occipital margin by slightly more than its own diameter and from eye
margin by slightly more than twice its own diameter. Antennal scrobes absent, torulus very nearly touching
mouth margin and separated from other torulus by about three times its own length. Antennal scape
slightly expanded and flattened, pedicel conical, about half as long as funicle, funicle six-segmented, all
segments transverse, the distal ones much longer and broader, clava three-segmented, broader than and
about as long as funicle and with apex rounded. Frontovertex with shallow coriaceous sculpture. Mandibles
narrow, with three acute teeth, the middle one slightly the longest.
Thorax. Pronotum not longitudinally divided in centre, mesoscutum without notaular lines, about a half
broader than long, axillae meeting, scutellum only very slightly broader than long, with sides (in dorsal
view) rounded and apex pointed, propodeum medially about one-seventh as long as scutellum. Pronotum,
mesoscutum and axillae with shallow coriaceous sculpture, similar to that on frontovertex, scutellum more
or less smooth and polished. Forewing more or less hyaline but slightly and irregularly infumate from about
half way along submarginal vein to apex of postmarginal ; wing about three times as long as broad ; basal
cell sparsely but almost entirely with setae, costal cell narrow, about as wide as submarginal vein, speculum
interrupted by a few setae and closed; submarginal vein with an apical hyaline break, marginal vein slightly
more than twice as long as broad, slightly shorter than stigmal and about twice as long as postmarginal.
Hindwing slightly more than five times as long as broad. Mid tibial spur shorter than mid basal tarsal
segment.
Gaster. About one-quarter longer than thorax, cerci about half way along but just in apical half,
hypopygium nearly reaching apex of gaster, exserted part of ovipositor slightly longer than one-third of
NEOTROPICAL ENCYRTIDAE 2 1 1
gaster and with sheaths distinctly flattened, deepest at about middle of exserted part and gradually tapering
towards apex.
J. Unknown.
Named in honour of my wife.
Mariola fluva sp. n.
(Figs 42, 184, 185)
9. Length (excluding ovipositor) 0-99 mm.
Colour. Body completely yellowish orange except antenna which has scape and pedicel testaceous and
ftagellum brown, propodeum slightly infuscate, gaster with a small area just anterior of each cercus and
sides near apex brown.
Head. Relative measurements of head : maximum width 45, frontovertex width at median ocellus 1 8,
malar space 6, eye length 28, eye width approximately 10, POL 7, OOL 4-5, scape length 18, other
proportions of antenna as in Fig. 185.
Thorax. Relative measurements of forewing (Fig. 184): length 115, maximum width 34, length of
submarginal vein 50, marginal vein 5, stigmal vein 7, postmarginal vein 2.
Gaster. About one-quarter longer than thorax, cerci about half way along but just in apical half,
3. Unknown.
DISTRIBUTION. St Vincent.
BIOLOGY. Unknown.
MATERIAL EXAMINED
Holotype ?, St Vincent: St George, Belmont, 6.vii.l976 (J. S. Noyes) (BMNH).
COMMENTS. Belongs to the tribe Microteryini, subtribe Platencyrtina and differs from the only
included genus, Platencyrtus Ferriere, in having the ovipositor well exserted with sheaths
flattened, relatively longer scutellum, narrower frontovertex and cerci more advanced on the
gaster. Platencyrtus has the ovipositor hardly exserted with the sheaths cylindrical, scutellum
triangular in shape and about one and three-quarter times as broad as long, frontovertex about
two-thirds head width and cerci nearer apex of gaster at about four-fifths.
MERCETENCYRTUS Trjapitzin
(Key couplets: $ 175, 203, 258; 3 130, 133)
Mercetencyrtus Trjapitzin, 1963: 884. Type-species: Encyrtus ambiguus Nees, by monotypy.
INCLUDED SPECIES. Two, one of which is found in the Neotropics : gracilicornis (De Santis, 1 964 :
191) (comb. n. from Syrphophagus); also a further one (or two) undescribed species from
Trinidad and Panama (BMNH; CNC, Ottawa).
DISTRIBUTION. Argentina, Brazil, Trinidad, Panama. World: cosmopolitan.
BIOLOGY. Unknown.
COMMENTS. The South American species are very close to ambiguus Nees but differ in the relative
lengths of the funicle segments and coloration of the legs. The mandibles are also different in
lacking, or apparently lacking the fourth (second lowest) tooth. However, one slide-mounted
specimen from Trinidad appears to have a four-toothed right mandible and a three-toothed left
mandible.
The species of Mercetencyrtus have a pit, or some similar structure, laterally on each side of
the scutellum which is clearly visible in slide-mounted material. In ambiguus it is fairly small and
situated close to the axillae whereas in gracilicornis it is relatively larger and nearly half way
along the scutellum. It is possible that there are many other species present in other parts of the
212 JOHN S. NO YES
world since those I have seen from Africa and Australia appear to represent unknown species.
The genus cannot be placed according to Trjapitzin's classification of the Encyrtidae.
MEROMYZOBIA Ashmead
(Key couplets: ? 12; 3 12)
Meromyzobia Ashmead, 1900: 349. Type-species: Ericydnus maculipennis Ashmead, by original
designation.
INCLUDED SPECIES. Eight, two of which are found in the Neotropics \flavipes De Santis, 19726 : 52
and gripha De Santis, 1968: 152.
DISTRIBUTION. Argentina. World: Nearctic region.
BIOLOGY. Parasites of eggs of Orthoptera.
COMMENTS. Both of the South American species differ from other species of Meromyzobia in
having only five funicle segments (one anelliform) instead of six (two anelliform). However, I
think these species are well placed here.
Placed in the tribe Miraini, subtribe Mayridiina by Trjapitzin & Gordh (19786).
METAPHYCUS Mercet
(Key couplets: 9 64, 77, 136, 194, 211, 232, 233; S 55, 68. Fig. 129)
Metaphycus Mercet, 1917a: 138. Type-species: Aphycus zebratus Mercet, by monotypy.
Euaphycus Mercet, 1921 : 197. Type-species: Encyrtus hederaceus sensu Mercet nee West wood (see Graham
1969: 224-225), designated by Mercet (1925: 23).
Mercetiella Dozier, 1926: 98. Type-species: Mercetiella reticulata Dozier, by monotypy.
Melanphycus Compere, 1947: 5. Type-species: Pseudococcobius fumipennis Timberlake, by original
designation.
Notoencyrtus De Santis, 1964: 211. Type-species: Notoencyrtus guttofasciatus De Santis, by monotypy.
Syn. n.
INCLUDED SPECIES. One hundred and twelve, 22 of which are found in the Neotropics:
alboclavatus Compere, 1939: 63, amoenus (Howard; Noyes, 1979: 160), angustifrons Compere,
1957: 227, brasiliensis Compere & Annecke, 1961 : 36, ceroplastae (Dozier, 1936: 185) (comb. n.
from Coccidoctonus), discolor (De Santis, 1970: 13) (comb. n. from Aphycus), eruptor (Howard;
Timberlake, 1916: 628),flavus (Howard; Mercet, 1921: 209), guttofasciatus (De Santis, 1964:
212) (comb. n. from Notoencyrtus), helvolus (Compere, 1926: 25), lounsburyi (Howard, 18986:
244), luteolus (Timberlake, 1916: 636), maculipes (Howard; Timberlake, 1916: 621) (comb. n.
from Aphycus), mexicanus (Howard; Timberlake, 1916: 627), monticolens Dozier, 1937: 124,
niger (Brethes; De Santis, 1964: 143) (comb. n. from Aphycus), oaxacae (Howard, 18986: 246)
(comb. n. from Aphycus), ogloblini (De Santis, 19726: 48) (comb. n. from Dicarnosis),
portoricensis (Dozier, 1926: 100) (comb. n. from Euaphycus}, reticulatus (Dozier, 1926: 98), rusti
(Timberlake, 1916: 617) (comb. n. from Aphycus), Stanley i Compere, 1940: 20; also many
undetermined species from Brazil, Colombia, Panama and Trinidad (BMNH; CNC, Ottawa).
REFERENCES. Timberlake (1916: 587-639); Annecke & Mynhardt (1971; 1972).
DISTRIBUTION. Throughout the Neotropics. World : cosmopolitan.
BIOLOGY. Parasites of Coccidae, Diaspididae, Lacciferidae and Eriococcidae (Homoptera).
COMMENTS. I have examined a paratype of Notoencyrtus guttofasciatus. It is a male Metaphycus
(not female as stated by De Santis).
The species described by Dozier as Coccidoctonus ceroplastae is very probably misplaced in
Metaphycus and may even belong to the tribe Micro teryini. However, I feel that it is best placec
here until its taxonomic position can be better determined.
Two of the South American species of Metaphycus are unusual (including discolor) in that the
ovipositor is well exserted, the exserted part being about as long as the gaster.
Placed in the tribe Aphycini, subtribe Paraphycina.
NEOTROPICAL ENCYRTIDAE 213
MICRO TERYS Thomson
(Key couplets: ? 65, 126; 3 66, 83, 105, 117. Fig. 133)
Microterys Thomson, 1876 : 155. Type-species : Encyrtus sylvius Dalman, by designation of Ashmead (1900 :
390).
Sceptrophorus Forster, 1856: 34. Type-species: Encyrtus sceptriger Forster, by monotypy. [Suppressed in
favour of Microterys Thomson: Opinion 1110, 1978, Opin. Decl. int. Commn zool. Nom. 35: 99-100.]
Apentelicus Fullaway, 1913: 26. Type-species: Apentelicus kotinskyi Fullaway, by monotypy.
INCLUDED SPECIES. One hundred and twenty-five, four of which are found in the Neotropics :
elegans Blanchard (De Santis, 1964: 183), flavus (Howard) ( = frontatus Mercet, 1921: 413),
kotinskyi (Fullaway, 1913: 26), tricoloricornis (Destefani) ( = consobrinus Mercet, 1921: 417,
709) ; also one undetermined species from Brazil (BMNH).
REFERENCE. Rosen (1976).
DISTRIBUTION. Throughout the Neotropics. World : cosmopolitan.
BIOLOGY. Parasites of Coccidae, Aclerdidae, Eriococcidae, Kermesidae, Kerridae,
Lecaniodiaspididae and Pseudococcidae (Homoptera).
COMMENTS. I have examined the syntypes of elegans; they are very probably the same as flavus,
differing only in the extent of the infuscation of the forewing.
Placed in the tribe Microteryini, subtribe Micro teryina.
MOORELLA Cameron
(Key couplet: 9 90. Fig. 45)
Moorella Cameron, 1913: 124. Type-species: Moorella fulviceps Cameron, by monotypy.
Chrysopophilus Timberlake, 1925: 178. Type-species: Chrysopophilus compressiventris Timberlake, by
monotypy. Syn. n.
INCLUDED SPECIES. Two: compressiventris (Timberlake, 1925: 179) (comb, n.) and fulviceps
Cameron, 1913: 125; also probably a further undetermined species from Panama.
DISTRIBUTION. Guyana, Panama.
BIOLOGY. Parasites of larvae of Chrysopidae (Neuroptera).
COMMENTS. I have examined a paratype of Chrysopophilus compressiventris ; it is congeneric with
M. fulviceps but is a distinct species.
The genus belongs to the tribe Homalotylini, subtribe Homalotylina.
MUCRENCYRTUS gen. n.
(Key couplets: ? 39, 117, 174, 193; j1 76. Figs 80, 84, 90, 186-189)
Type-species : Mucrencyrtus insulanus sp. n.
?. Head. In profile, deep and most rounded level with top of antennal scrobes. Eye about one-quarter longer
than broad, naked and reaching occipital margin which is rounded. Malar space less than half length of eye
with malar sulcus weakly indicated. Frontovertex at narrowest point about two-fifths head width, ocelli
small and more or less forming a right angle, posterior ocellus about two to four times its own diameter
from occipital margin and less than its own diameter from eye margin. Antennal scrobes deep with outer
margins, at extreme uppermost point, sharply margined, interantennal prominence, in outline like an
inverted Y with edge sharp above antennal toruli and produced as a long, narrow, sharp ridge which nearly
reaches frontovertex ; antennal torulus separated from mouth margin by slightly less than its own length
and from other torulus by slightly more than its own length, its dorsal margin slightly above level of lowest
eye margin. Antennal scape subcylindrical or very slightly flattened, pedicel conical, longer than first funicle
segment, funicle six-segmented, cylindrical and widening very slightly distally, clava three-segmented, with
apex rounded and about half as long as funicle; longitudinal sensilla on all flagellar segments.
Frontovertex with shallow reticulate to punctiform reticulate sculpture which becomes slightly elongate and
sometimes shallow coriaceous on lower parts of face and genae. Mandibles with three teeth, upper tooth
slightly rounded, maxillary palpi four-segmented, labial palpi three-segmented.
214 JOHN S. NOYES
Thorax. In side view moderately deep, mesoscutum and scutellum flat. Mesoscutum about two to two
and a half times as broad as long, without notaular lines or with notaular lines vaguely indicated anteriorly,
axillae meeting, scutellum about as broad as long, very flat with apex pointed and produced as a very
narrow translucent flange which overhangs the propodeum, propodeum medially longer than one-sixth
length of scutellum. Pronotum, mesoscutum and axillae with shallow coriaceous sculpture, sometimes
almost smooth ; scutellum with distinctly deeper coriaceous, almost reticulate, sculpture of smaller mesh ;
metanotum and propodeum with reticulate sculpture, area either side of median area of propodeum
transversely strigose almost to spiracles, propodeum sometimes smooth medially; mesopleuron with
shallow, slightly elongate, reticulate sculpture. Forewing hyaline or with a central fuscous spot, basal cell
and costal cell (sometimes less so) densely hairy, speculum occasionally interrupted but always closed,
submarginal vein without apical hyaline break, marginal vein not reaching anterior margin of wing, at most
as long as broad, apical cilia absent or very short. Macropterous forms: forewing about two and a half
times as long as broad, marginal vein about one-quarter to half as long as stigmal, postmarginal very short,
almost absent; hindwing about two and a half times as long as broad. Micropterous forms: forewing
slightly more than twice as long as broad with apex truncate, marginal vein only slightly shorter than
stigmal; hindwing about two and a half times as long as broad with apex obliquely truncate. Mid tibial spur
very slightly shorter than corresponding basitarsus.
Caster. About as long as thorax, dorsally flat, ventrally very convex, cerci at about one-third, paratergites
absent, hypopygium with a long, thin apical extension which reaches apex of gaster (as in Fig. 84), not
notched apically. Ovipositor about as long as mid tibia, gonostyli apparently absent.
j1. Essentially similar to female except no micropterous forms known; antennae (as in Fig. 188), setae in
basal cell of forewing less dense, last gastral sternite of normal shape.
Mucrencyrtus insulanus sp. n.
(Figs 80, 84, 90, 186-189)
?. Length: 1-27-1-43 mm (holotype 1-35 mm).
Colour. Body completely orange, metanotum, propodeum and extreme of gaster slightly mixed fuscous,
forewing hyaline.
Head. Posterior ocellus about two and a half times its own diameter from occipital margin; frontovertex
with shallow, punctiform-reticulate sculpture which becomes more irregular on lower parts of face and
cheeks. Relative measurements of head of holotype; head width 91, frontovertex width at median ocellus
36, malar space 23, eye length 55, eye width 43, POL 26, OOL 3, scape length 34, other proportions of
antenna as in Fig. 189. Head in frontal aspect as in Fig. 80.
Thorax. Mesoscutum and scutellum with numerous translucent setae which appear white in some light.
Propodeum with dense reticulate sculpture, anteriorly from either side of middle nearly to spiracles almost
transversely striate. Macropterous form: forewing at extreme apex and along anterior margin without
marginal cilia, basal cell with setae nearly as dense as, but markedly stronger than, those in centre of wing,
costal cell with setae as dense as basal cell, speculum not interrupted ; relative measurements of forewing of
holotype: length 172, maximum width 68, length of submarginal vein 74, of marginal vein 5, of stigmal vein
11, of postmarginal vein 0-5; of hindwing: length 112, maximum width 31. Micropterous form: setae of
forewing similar to macropterous form, wing margin without setae, apex of wing very slightly obliquely
truncate; relative measurements of forewing of paratype: length 75, maximum width 31, length of
submarginal vein 52, of marginal vein 3, of stigmal vein 5-5, of postmarginal vein 0-5; of hindwing: length
40, maximum width 12, apex distinctly obliquely truncate.
Gaster. In side view as in Fig. 84. Relative measurements of paratype: length of ovipositor 59 [mid tibia
65]. Ovipositor as in Fig. 187.
cJ. Length: 1-08-1 -20 mm.
Essentially similar to female except the following: lower parts of face whitish, occiput, pronotum,
propodeum and metanotum marked with dark brown, centre of thorax yellowish white, dorsum of gaster
brownish ; ocelli forming a distinctly obtuse angle ; forewing (Fig. 90) with setae in basal and costal cells not
quite as dense. Male genitalia (Fig. 186): each digitus with two apical spines. Relative measurements of
paratype: head width 75, frontovertex width at median ocellus 35, malar space 20, eye length 40, POL 24,
OOL 3, scape length, proportions of antenna as in Fig. 188.
NEOTROPICAL ENCYRTIDAE 215
DISTRIBUTION. Trinidad, St Vincent.
BIOLOGY. Reared from Aclerda sp. (Homoptera : Aclerdidae).
MATERIAL EXAMINED
Holotype 9, Trinidad: St George, St Augustine, x.1976 (F. D. Bennett} (BMNH).
Paratypes. Trinidad: 2 y, Orange Grove, i.1973, ex Aclerda sp. on sugarcane (F. D. Bennett); 1 $, Curepe,
Santa Margarita Circular Road, 15-28.U974 (F. D. Bennett}; 2 $, 2 ^, St Augustine, 21. vi. 1976 (J. S.
Noyes); 1 3, St Vincent, St George, Argyle, 8.vii.l976 (/. S. Noyes). (BMNH; CIBC, Trinidad; CNC,
Ottawa.)
COMMENTS. Two further species examined: Mucrencyrtus aclerdae (De Santis) (comb. n. from
Aenasioided) and an undetermined species from Ecuador (CNC, Ottawa). M. aclerdae has a large
infuscate spot in the centre of the forewings and the setae in the costal cell less dense than in the
type-species.
Very close to Allencyrtus Annecke & Mynhardt (tribe Microteryini, subtribe Micro teryina)
which differs in having a metallic body (greenish blue), hypopygium barely extended apically and
deeply notched, propodeum rather smooth, mandibles with upper tooth rudimentary and
forewings more or less generally suffused greyish.
NEAPSILOPHRYS gen. n.
(Key couplet: ? 150. Figs 68, 190-191)
Type-species: Neapsilophrys flavipes sp. n.
9. Head. In side view, moderately deep and most rounded level with top of antennal scrobes, frontovertex
slightly convex. Eye about one-third longer than broad, naked and reaching occipital margin. Malar space
slightly longer than half length of eye with malar sulcus absent. Frontovertex at narrowest point, slightly
less than half head width, ocelli forming a slightly obtuse angle, posterior ocellus separated from occipital
and eye margins by less than its own diameter. Antennal scrobes shallow, separated dorsally by
continuation of interantennal prominence, and reaching about half way from antennal toruli to anterior
ocellus; antennal torulus separated from mouth margin and other torulus by about its own length. Scape
subcylindrical, pedicel conical and about same length as first funicle segment, all six funicle segments longer
than broad and cylindrical, clava three-segmented, apically rounded and about as wide as funicle;
longitudinal sensilla on all flagellar segments. Frontovertex with irregular reticulate sculpture which is
much shallower on lower parts of face so that genae and areas below eyes are almost completely smooth.
Mandibles tridentate with upper tooth rounded or blunt, maxillary palpi four-segmented, labial palpi three-
segmented.
Thorax. In side view moderately deep and dorsally convex. Mesoscutum without notaular lines, about a
half broader than long, axillae almost meeting, scutellum distinctly longer than broad and with apex
rounded and slightly produced as a flange, propodeum medially about one-seventh as long as scutellum.
Pronotum with coriaceous sculpture, mesoscutum with raised reticulate sculpture medially but laterally
with strigose sculpture, scutellum with fine strigose sculpture except extreme apex which is polished,
mesopleuron with strigose sculpture which posteriorly becomes more reticulate, propodeum smooth.
Forewing hyaline, about two and a half times as long as broad, basal cell hairy except for proximal one-
third which is hyaline, speculum open but interrupted by two or three lines of setae; submarginal vein with
an apical hyaline break, marginal vein more than twice as long as broad and about as long as stigmal,
postmarginal vein nearly twice as long as stigmal. Mid tibial spur slightly shorter than mid basal tarsal
segment.
Caster. Slightly longer than thorax and apically compressed from side to side. Cerci at about one-third,
paratergites absent, hypopygium very nearly reaching apex of gaster, exserted part of ovipositor slightly
longer than half length of gaster. Ovipositor one and a half times as long as gaster or about two and a half
times as long as mid tibia, gonostylus about as long as mid tibia.
3- Unknown.
216 JOHN S. NOYES
Neapsilophrys flavipes sp. n.
(Figs 68, 190, 191)
?. Length (excluding ovipositor): 1-75-1-96 mm (holotype 1-81 mm).
Colour. Frontovertex and dorsum of thorax metallic green, pronotum and scutellum strongly tinged
purple, antenna with scape and pedicel yellow to yellowish brown, flagellum brown, lower parts of face,
genae, temples and mesopleura purple, legs yellow except mid and hind coxae which are brown, base of
gaster yellow, propodeum and remainder of gaster, including ovipositor sheaths, brown.
Head. Relative measurements of head of holotype : head width 76-5, frontovertex width at median ocellus
34, malar space 28, eye length 45, eye width 34, POL 17, OOL 3, length of scape 49, proportions of antenna
as in Fig. 191.
Thorax. Relative measurements of forewing of holotype: length 228, maximum width 89, length of
submarginal vein 87, of marginal vein 15, of stigmal vein 14-5, of postmarginal vein 27; of hindwing: length
148, maximum width 42. Base of forewing Fig. 68.
Gaster. Relative length of paratype: length of ovipositor 238, of gonostylus 104 [mid tibia 96]. Genitalia
as in Fig. 190.
DISTRIBUTION. Brazil.
BIOLOGY. Unknown.
MATERIAL EXAMINED
Holotype 9, Brazil: Santa Catarina, Nova Teutonia, 14.xi.1944 (F. Plaumanri) (BMNH).
Paratypes. Brazil: 1 ?, Nova Teutonia, 14.xi.1944; 4 $, Nova Teutonia 16.xi.1944; 8 $, Nova Teutonia,
30.iii.1945; 2 ?, Nova Teutonia, 27.iv.1945 (F. Plaumann). (BMNH; USNM, Washington; CIBC,
Trinidad; CNC, Ottawa; MLP, Argentina; PPRI, Pretoria; ZI, Leningrad.)
COMMENTS. The genus belongs to the tribe Copidosomatini, subtribe Ageniaspidina. It can be
separated from other members of the subtribe found in South America using the characters given
in the key. In many ways the genus is intermediate between this subtribe and the
Copidosomatina, the venation and arrangement of the setae at the base of the forewing being
typical of the former but the body is generally typical of the latter.
Also superficially similar to Deloencyrtus but differs in having the gaster apically bilaterally
flattened, the eyes naked, the scutellum convex and the hypopygium without a strong apical
median incision, whereas Deloencyrtus has the gaster dorso-ventrally flattened, the eyes distinctly
hairy, the scutellum flat and the apex of the hypopygium with a strong apical median incision.
NEOCOCCIDENCYRTUS Compere
(Key couplets: ? 102, 118, 245; 3 142. Figs 52, 103)
Neococcidencyrtus Compere, 1928: 209. Type-species: Neococcidencyrtus alula Compere, by monotypy.
Xenocomys Blanchard, 1940: 106. Type-species: Xenocomys chrysomphali Blanchard, by monotypy.
INCLUDED SPECIES. Five, two of which are found in the Neotropics: chrysomphali (Blanchard ; De
Santis, 1964: 188), crouzelae De Santis, 1964: 233; also two undetermined species from Brazil
and Trinidad (BMNH).
REFERENCE. Key to world species: Myartseva (19776: 175).
DISTRIBUTION. Argentina, Brazil, Trinidad. World : Nearctic, Palaearctic and Ethiopian regions.
BIOLOGY. Parasites of Diaspididae (Homoptera).
COMMENTS. Very probably a member of the tribe Habrolepidini.
NEODUSMET1A Kerrich
(Key couplets: 9 37; ^ 45. Fig. 18)
Neodusmetia Kerrich, 1964: 76. Type-species: Dusmetia sangwani Subba Rao, by monotypy.
NEOTROPICAL ENCYRTIDAE 217
INCLUDED SPECIES. One: sangwani (Subba Rao, 1957: 385).
DISTRIBUTION. Mexico, Brazil. World: Nearctic, Ethiopian and Oriental regions.
BIOLOGY. Parasites of Pseudococcidae (Homoptera).
COMMENTS. Placed in the tribe Dinocarsini by Gordh & Trjapitzin (19786).
OOENCYRTUS Ashmead
(Key couplets: 9 39, 82, 171, 199, 247, 255; J1 81, 114, 122. Figs 76, 77)
Ooencyrtus Ashmead, 1900: 381. Type-species: Encyrtus clisiocampe Ashmead, by original designation.
Schedius Howard, 1910: 2. Type-species: Schedius kuwanae Howard, by monotypy.
Pseudlitomastix Risbec, 1954: 1068. Type-species: Litomastix creona Risbec, by monotypy.
INCLUDED SPECIES. Seventy-eight, six of which are found in the Neotropics : chrysopae Crawford,
19136: 341,johnsoni (Howard, 18980: 18), latiscapus Gahan, 1927: 15, prenidis Gahan, 1943:
137, submetallicus (Howard; Noyes, 1979: 160) and trinidadensis Crawford, 19136: 347; also five
or six undetermined species from Trinidad and Brazil (BMNH ; CNC, Ottawa).
DISTRIBUTION. Throughout the Neotropics. World : cosmopolitan.
BIOLOGY. Parasites of eggs of various insects, notably Lepidoptera and Heteroptera.
COMMENTS. Ooencyrtus trinidadensis venatorius De Santis & Sarmiento (in De Santis et al., 1976 :
7-9) may be a distinct species (De Santis, pers. comm.).
I have selected and labelled LECTOTYPE females from the syntype-series of each of O.
chrysopae Crawford (labelled: 'Verdantvale, Arima, Trinidad, eggs Chrysopa 51-1 sp., F. W.
Urich Collector, U.S.N.M. Type No. 16040') and O. trinidadensis Crawford (labelled 'reared
from eggs of Pentatomid, Port of Spain, Trinidad, BWI, Feb. 1912, F. W. Urich Collector, Type
No. 16039 U.S.N.M.') (USNM, Washington).
Placed in the tribe Microteryini, subtribe Ooencyrtina. See Comments under Echthrodryinus.
PAPAKA gen. n.
(Key couplets: 9 246; 3 55. Figs 102, 192-196)
Type-species: Papaka confusor sp. n.
9- Head. In side view moderately deep and quite triangular, frontovertex moderately convex. Eye about
one-third longer than broad, naked and over-reaching occipital margin which is rounded. Clypeal margin
broadly emarginate. Malar space nearly two-thirds eye length and with malar sulcus absent. Frontovertex
about one-quarter head width; ocelli forming an acute angle of about 60°, posterior ocellus separated from
occipital margin by about twice its diameter and nearly touching eye margin. Antennal scrobes shallow,
meeting dorsally and reaching slightly further than half way from antennal toruli to anterior ocellus;
antennal torulus separated from mouth margin by less than its own length and from other torulus by about
one and a half times its own length, its dorsal margin well below ventral margin of eye. Antennal scape
slightly expanded and flattened in its basal half, pedicel conical and longer than the first two funicle
segments together, funicle six-segmented and cylindrical, the first two segments transverse or quadrate, the
remainder longer than broad and hardly broadening distally, clava hardly wider than funicle, three-
segmented with apex pointed and about half length of funicle; longitudinal sensilla on all flagellar segments
except proximal three. Frontovertex with irregular coriaceous sculpture, more elongate on interantennal
prominence and cheeks. Mandibles with two acute teeth and a rudimentary third (upper) tooth (Fig. 102),
maxillary palpi four-segmented, labial palpi three-segmented.
Thorax. In side view, moderately deep with mesoscutum and scutellum flat. Mesoscutum without
notaular lines, about one-third broader than long, axillae more or less meeting, scutellum slightly broader
than long, very flat and with apex acute, propodeum medially about one-fifteenth as long as scutellum.
Pronotum and mesoscutum slightly shiny with fairly regular shallow reticulate sculpture, more elongate
towards side of mesoscutum, axillae with deep coriaceous sculpture, scutellum matt, with more or less
regular punctate sculpture, propodeum almost smooth, mesopleuron almost smooth but with some very
218 JOHN S. NO YES
shallow, reticulate sculpture. Forewing hyaline, about two and a half times as long as broad; basal cell
completely hairy, speculum interrupted by two or three lines of setae and closed ; submarginal vein slightly
expanded apically and with an apical hyaline break, marginal vein about twice as long as broad, less than
half as long as stigmal and about twice as long as postmarginal. Hindwing about four and a half times as
long as broad. Mid tibial spur slightly shorter than mid basal tarsal segment.
Caster. Slightly shorter than thorax, cerci at about one-third, paratergites absent, apex of hypopygium
reaching about three-quarters along gaster, ovipositor slightly exserted. Ovipositor about as long as gaster
or one and a half times length of mid tibia, gonostylus about as long as mid tibial spur.
J1. Essentially similar to female except fronto vertex about one-third head width, antennae with setae
longer and clava entire; genitalia : digitus with two apical spines and aedeagus about two-thirds as long as
mid tibia.
Papaka confusor sp. n.
(Figs 102, 192-196)
9. Length: 0-86-1-17 mm (holotype 1-01 mm).
Colour. Body generally black or very dark brown, frontovertex and mesoscutum with some obscure green
or brassy reflections ; antenna with scape and pedicel dark brown, both yellowish white apically, first five
funicle segments and clava brown, sixth funicle segment yellow; legs with coxae brown, femora brown but
basally and apically yellow, tibiae and tarsi yellow, the fore tibia with a broad dark brown median band,
mid and hind tibiae each with two dark brown bands, ovipositor sheaths yellow.
Head. Relative measurements of holotype : head width 55, frontovertex width at median ocellus 14, malar
space 23, eye length 37, eye width 30, POL 5-5, OOL 1 -5, scape length 26, proportions of antenna as in Fig.
192.
Thorax. Relative measurements of forewing of holotype: length 152, maximum width 64, length of
submarginal vein 60, of marginal vein 5-5, of stigmal vein 12, of postmarginal vein 3; of hindwing: length
100, maximum width 23. Base of forewing as in Fig. 194.
Gaster. Exserted part of ovipositor equal to about one-eighth length of gaster or one-sixth mid tibia.
Relative measurements of paratype: length of ovipositor 73, of gonostylus 14 [mid tibia 45, mid tibial spur
13]. Ovipositor as in Fig. 193.
3. Length: 0-68-0-73 mm.
Antenna as in Fig. 196, genitalia as in Fig. 195. Otherwise similar to female.
DISTRIBUTION. Brazil.
BIOLOGY. Unknown.
MATERIAL EXAMINED
Holotype ?, Brazil: Santa Catarina, Nova Teutonia, 14.x. 1949 (F. Plaumann) (BMNH).
Paratypes. Brazil: 2 $, 2 J, Nova Teutonia, ix.1943; 1 $, Nova Teutonia, 31.viii.1949; 6 ?, 1 J1, Nova
Teutonia, x.1949 (F. Plaumann). (BMNH; MLP, Argentina; UC, Riverside; ZI, Leningrad; PPRI,
Pretoria.)
COMMENTS. One further undescribed species from Brazil (BMNH) which differs from the type-
species in the relative lengths of the funicle segments, the sixth funicle segment concolorous with
rest of the funicle and the exserted part of the ovipositor relatively longer.
The genus is superficially very similar to some species of Neococcidencyrtus and
Coccidencyrtus but the forewing venation suggests that it is very probably more closely related to
Cicoencyrtus and Archinus (Tribe Aphycini). It differs from both in having the upper tooth of the
mandibles rudimentary, and in the sculpture of the mesoscutum, the relatively longer funicle
segments and more densely hairy basal cell.
PARABLASTOTHRIX Mercet
(Key couplets: 9 91 ; ^ 5. Fig 48)
Parablastothrix Mercet, 1917c: 538. Type-species: Parablastothrix vespertina Mercet, by monotypy.
NEOTROPICAL ENCYRTIDAE 2 1 9
INCLUDED SPECIES. Seven, one of which is found in the Neotropics: chilensis (Brethes, 1919: 165)
(comb. n. from Archinus), also three undetermined species from Brazil, Trinidad and St Vincent
(BMNH).
REFERENCE. Key to world species: Trjapitzin (1971 : 36).
DISTRIBUTION. Chile, Brazil, Trinidad, St Vincent. World : cosmopolitan.
BIOLOGY. Parasites of larvae of Lyonetiidae and Nepticulidae (Lepidoptera).
COMMENTS. I have not seen the type of Archinus chilensis but, from the description, it is very
probably a species of Parablastothrix.
Placed in the tribe Copidosomatini, subtribe Pentacnemina. The genus is very close to
Calometopia Mercet, but differs in the female only in having a slightly narrower clava with a
shorter truncation, and in the male in having one segment less with a ramus. It is quite possible
that the two genera will eventually be synonymized.
PARALEUROCERUS Girault
(Key couplets : $ 219; 3 34)
Paraleurocerus Girault, 191 5c: 172. Type-species: Paraleurocerus bicoloripes Girault, by monotypy.
Gibberella Miller, 1961 : 496. Type-species: Gibberella scutellata Miller, by monotypy.
INCLUDED SPECIES. Three, one of which is found in the Neotropics: reticulatus (De Santis, 1964:
310) (comb. n. from Ageniaspis}.
DISTRIBUTION. Argentina, Brazil. World : Nearctic region.
BIOLOGY. Polyembryonic parasites of larvae of Gracillariidae (Lepidoptera).
COMMENTS. I have examined a paratype of Ageniaspis reticulatus; it is clearly best placed in
Paraleurocerus, although it is very probable that with further study the two genera will be
considered synonymous.
Placed in the tribe Copidosomatini, subtribe Ageniaspidina.
PARALITOMAST1X Mercet
(Key couplets: $ 248; 3 77)
Paralitomastix Mercet, 1921 : 438. Type-species: Encyrtus varicornis Nees, by monotypy.
INCLUDED SPECIES. Fifteen species, one of which is found in the Neotropics : teciae Blanchard (De
Santis 1964: 290), and one undetermined species near varicornis (Nees) from Brazil (BMNH).
DISTRIBUTION. Chile, Argentina, Brazil. World: cosmopolitan.
BIOLOGY. Polyembryonic parasites of larvae of Pyralidae and Gelechiidae (Lepidoptera).
COMMENTS. Placed in the tribe Copidosomatini, subtribe Copidosomatina.
PARAMUCRONA gen. n.
(Key couplet: $ 174. Figs 85, 197, 198)
Type-species : Paramucrona brasiliensis sp. n.
9. Head. In profile, moderately deep and most rounded level with top of antennal scrobes, frontovertex
slightly convex. Eye distinctly bulging, slightly longer than broad, naked and not quite reaching occipital
margin which is rounded except near middle where it is more or less sharp. Malar space slightly longer than
half length of eye and with malar sulcus present, but weak. Frontovertex at narrowest point nearly half
head width, ocelli forming an obtuse angle, posterior ocellus separated from occipital margin and eye
margin by about its own diameter. Antennal scrobes quite shallow and not well delimited, reaching slightly
more than half way between toruli and anterior ocellus ; interantennal prominence relatively protuberant
and rounded, reaching slightly further than dorsal margins of toruli and with a single, slightly curved line of
220 JOHN s- NO YES
about seven very conspicuous long, erect setae about halfway between toruli and mouth margin; antennal
torulus separated from mouth margin by about its own length and from other torulus by slightly more than
its length. Antennal scape slightly to distinctly flattened and expanded in middle, pedicel shorter than first
funicle segment, funicle six-segmented, all funicle segments longer than broad and cylindrical, clava three-
segmented, about as long as fifth and sixth funicle segments together and with apex more or less rounded ;
longitudinal sensilla on all flagellar segments but relatively short, mostly only two to three times as long as
broad, sometimes hardly longer than broad. Frontovertex with scattered inconspicuous pale setae,
sculpture very shallow coriaceous becoming more longitudinally elongate on lower parts of face, except on
interantennal prominence. Mandibles with three teeth, the lowest acute and the upper two slightly rounded,
maxillary palpi four-segmented, labial palpi three-segmented.
Thorax. In profile moderately deep, mesoscutum and scutellum fairly flat. Mesoscutum with notaular
lines weakly indicated anteriorly, otherwise absent, axillae meeting, scutellum slightly longer than broad,
slightly shorter than mesoscutum, with apex pointed and produced as a short flange, propodeum medially
less than one-seventh as long as scutellum. Dorsum of thorax with very shallow coriaceous sculpture,
propodeum medially with one or two shallow incomplete carinae, mesopleuron smooth. Forewing hyaline,
about two and a half times as long as broad, with short cilia along apical and costal margins ; basal cell
sparsely hairy with a bare triangle in proximal half, costal cell with one or two lines of setae dorsally which
extend for about three-quarters its length to apex, speculum not interrupted and open ; submarginal vein
without apical hyaline break, marginal vein absent, stigmal vein arising directly from submarginal,
postmarginal vein present but very short. Hindwing almost three and a half times as long as broad. Mid
tibial spur slightly shorter than mid basal tarsal segment.
Caster. Much shorter than, to nearly as long as, thorax, cerci at about one-third, ovipositor hidden or
very slightly exserted, hypopygium with short apical triangular mucro (as in Fig. 84), which is at most
slightly longer than mid tibial spur, but reaches apex of gaster. Ovipositor slightly shorter than mid tibia,
gonostylus slightly less than half as long as mid tibial spur.
c?. Unknown.
Paramucrona brasiliensis sp. n.
(Figs 85, 197, 198)
9. Length: 1-89-2-00 mm (holotype 1-92 mm).
Colour. Body orange except posterior three-quarters of mesopleuron which is brown, and metanotum
and dorsum of gaster which are slightly infuscate.
Head. Ocelli forming an angle of about 105-110°, occipital margin not sharp, only slightly acute
medially. Relative measurements of holotype : head width 99, frontovertex width at median ocellus 45,
malar space 29, eye length 52, eye width 44, POL 18, OOL 8, scape length 45, proportions of antennae as in
Fig. 197. Longitudinal sensilla on flagellum at least twice as long as broad, some on clava five times as long
as broad.
Thorax. Relative measurements of forewing of holotype: length 300, maximum width 121, length of
submarginal vein 120, of stigmal vein 20, of postmarginal vein 4; of hindwing: length 176, maximum width
50. Base of forewing as in Fig. 85.
Gaster. Only slightly shorter than thorax. Relative measurements of paratype: length of ovipositor 90, of
gonostylus 13 [mid tibia 1 12, mid tibial spur 28] ; apical extension of hypopygium about as long as mid tibial
spur.
J. Unknown.
DISTRIBUTION. Brazil.
BIOLOGY. Unknown.
MATERIAL EXAMINED
Holotype ?, Brazil: Santa Catarina, Nova Teutonia, 3 1.x. 1949 (F. Plaumann) (BMNH).
Paratypes. Brazil: 2 ?, Nova Teutonia, 30.x. 1949 (F. Plaumann) (BMNH).
COMMENTS. Two further undescribed species from Brazil (BMNH; CNC, Ottawa) differ in
relative lengths of ovipositor, gonostylus, apical extension of hypopygium, coloration,
proportions of antennal segments and sharpness of occipital margin.
NEOTROPICAL ENCYRTIDAE 22 1
Belongs to the same group as Bennettisca, Mucrencyrtus and Allencyrtus Annecke &
Mynhardt (tribe Microteryini, subtribe Microteryina), but differs from the first two by
characters given in the key and from Allencyrtus in lacking the marginal vein, and in the pilosity
of the base of the forewing, colour of body, structure of antennae, presence of costal cilia and
lack of carinate extension of interantennal prominence. Allencyrtus has a dark, metallic body,
densely pilose basal and costal cells, marginal vein present (although not reaching anterior
margin of forewing), relatively short clava, costal cilia lacking and carinate extension of
interantennal prominence (as in Mucrencyrtus).
PARECHTHRODRYINUS Girault
(Key couplets: ? 116, 127, 161, 192, 238; J 116, 125. Figs 57, 138)
Parechthrodryinus Girault, 1916c: 480. Type-species: Parechthrodryinus convexus Girault, by monotypy.
Tyndarichoides Mercet, 1921: 649. Type-species: Tyndarichoides metallicus Mercet, by monotypy.
[Homonym of Tyndarichoides Girault, 1920.]
Protyndarichus Mercet, 1922: 479. [Replacement name for Tyndarichoides Mercet.]
INCLUDED SPECIES. Fifteen, one of which is found in the Neotropics: nitidus (Howard; Noyes,
1979: 163) (comb. n. from Protyndarichus).
DISTRIBUTION. Argentina, Trinidad, Grenada, Tobago, St Vincent. World: cosmopolitan.
BIOLOGY. Parasites of Lacciferidae and Pseudococcidae (Homoptera).
COMMENTS. Placed in the tribe Cheiloneurini.
PARENCYRTUS Ashmead
Parencyrtus Ashmead, 1900: 368. Type-species: Parencyrtus brasiliensis Ashmead, by monotypy.
INCLUDED SPECIES. One: brasiliensis Ashmead, 1900: 368.
DISTRIBUTION. Brazil.
BIOLOGY. Unknown.
COMMENTS. The holotype is lost, see Kerrich (1967: 152) and Gordh & Trjapitzin (1978: 712).
This genus is very possibly the same as Rhytidothorax or Hemencyrtus.
PAURIDIA Timberlake
(Key couplets : 9 236, ^41)
Pauridia Timberlake, 19196: 206. Type-species: Pauridia peregrina Timberlake, by monotypy.
INCLUDED SPECIES. One: peregrina Timberlake, 19196: 208.
DISTRIBUTION. Chile, Peru, Bermuda. World : Nearctic and Ethiopian regions.
BIOLOGY. Parasites of Pseudococcoidea (Homoptera).
COMMENTS. Placed in the tribe Pauridiini.
PELMATENCYRTUS De Santis
(Key couplet: ? 32)
Pelmatencyrtus De Santis, 1964: 125. Type-species: Pelmatencyrtus bonariensis De Santis, by monotypy.
INCLUDED SPECIES. One: bonariensis De Santis (Kerrich, 1978: 156).
DISTRIBUTION. Argentina.
BIOLOGY. Unknown.
COMMENTS. Placed in the tribe Acroaspidiini by Trjapitzin (1973a), which may be correct.
222 JOHN s- NOYES
PENTALITOMASTIX Eady
(Key couplets: ? 25; ^ 51. Figs 15, 126)
Pseudolitomastix Eady, 1960a: 667. Type-species: Pseudolitomastix nacoleiae Eady, by monotypy.
[Homonym of Pseudolitomastix Risbec, 1954.]
Pentalitomastix Eady, 19606: 173. [Replacement name for Pseudolitomastix Eady.]
INCLUDED SPECIES. Five, one of which is found in the Neotropics: plethorica Caltagirone, 1966:
145.
DISTRIBUTION. Mexico. World: Palaearctic, South East Asia.
BIOLOGY. Polyembryonic parasites of Pyralidae and Tortricidae (Lepidoptera).
COMMENTS. Placed in the tribe Copidosomatini, subtribe Copidosomatina.
PHEIDOLOXENUS Ashmead
(Key couplet: 9 31)
Pheidoloxenus Ashmead, 1904: 328. Type-species: Pheidoloxenus wheeleri Ashmead, by monotypy.
Pheidoloxenus Girault, 1915a: 273. Type-species: Pheidoloxenus wheeleri Girault, by monotypy.
[Homonym of Pheidoloxenus Ashmead, 1904.]
Pheidoloxeniscus Ghesquiere, 1946: 369. [Replacement name for Pheidoloxenus Girault.]
INCLUDED SPECIES. Two : one not from the Neotropics, the other is being described from Mexico
by Dr G. Gordh ( = wheeleri sensu Mann, 1914, nee Ashmead).
DISTRIBUTION. Mexico. World: Nearctic.
BIOLOGY. Reared from ant nests (Formicidae), probably parasitic on mealybugs (Homoptera:
Pseudococcidae) present in nest.
COMMENTS. Placed in the tribe Acroaspidiini.
PLAGIOMERUS Crawford
(Key couplet: $ 6; 3 7)
Plagiomerus Crawford, 1910: 89. Type-species: Plagiomerus diaspidis Crawford, by monotypy.
Parahomalopoda Girault, 191 5c: 170. Type-species: Parahomalopoda peruviensis Girault, by monotypy.
Syn. n.
INCLUDED SPECIES. Six, three of which are found in the Neotropics : cyanea (Ashmead ; Dozier,
1927: 273), diaspidis Crawford, 1910: 90 and peruviensis (Girault, 1915c: 171) (comb. n. from
Parahomalopoda); also one undetermined species from Trinidad (BMNH).
DISTRIBUTION. Mexico, Cuba, Puerto Rico, Trinidad, Peru. World: Nearctic, Palaearctic and
Oriental regions.
BIOLOGY. Parasites of Diaspididae (Homoptera).
COMMENTS. Parahomalopoda peruviensis differs from the true Plagiomerus only in lacking the
four lamelliform setae at the apex of scutellum. The undetermined species from Trinidad has a
line of four normal setae at the apex of the scutellum. I can therefore see no case for keeping the
two genera separate.
Placed in the tribe Habrolepidini, subtribe Habrolepidina.
PLATYLYCA De Santis
(Key couplet: ? 146. Fig. 67)
Platylyca De Santis, 19726: 58. Type-species: Platylyca quadraticeps De Santis, by monotypy.
INCLUDED SPECIES. One: quadraticeps De Santis, 19726: 58.
NEOTROPICAL ENCYRTIDAE 223
DISTRIBUTION. Argentina.
BIOLOGY. Unknown.
COMMENTS. Very probably best placed in the tribe Habrolepidini.
PRIONOMASTIX Mayr
(Key couplets: • 112, 132; ^ 67. Fig. 135)
Prionomastix Mayr, 1876: 725. Type-species: Encyrtus morio Dalman, by monotypy.
Liocarus Thomson, 1876: 115, 121. Type-species: Encyrtus morio Dalman, by monotypy.
Chestomorpha Ashmead, 1900: 370. Type-species: Chestomorpha biformis Ashmead, by monotypy.
Aprionomastix Girault, 1913a: 68. Type-species: Aprionomastix fasciatipennis Girault, by monotypy.
INCLUDED SPECIES. Seven, one of which is found in the Neotropics: fasciatipennis
(Girault) ( = bicarinata De Santis, 1964: 179), also three undetermined species from Trinidad,
Ecuador and Chile (BMNH; CNC, Ottawa).
REFERENCE. Revision of world species: Annecke (1962).
DISTRIBUTION. Argentina, Paraguay, Ecuador, Chile, Trinidad. World: cosmopolitan.
BIOLOGY. Parasites of nymphs of Membracidae (Homoptera).
COMMENTS. Placed in the tribe Prionomasticini, subtribe Prionomasticina.
One undetermined species from Trinidad (CNC, Ottawa) (not included above) falls between
Prionomastix and Hexacladia. The structure of the thorax is close to that of Prionomastix but the
antennae are typical of Hexacladia. On this evidence I believe that the two subtribes
Prionomasticina and Hexacladiina should be considered the same.
PRIONOMITUS Mayr
Prionomitus Mayr, 1876: 701. Type-species: Encyrtus chlorinus Dalman, by monotypy.
INCLUDED SPECIES. Ten, one of which is found in the Neotropics : fuscipalpis Kieffer (in Kieffer &
Jorgensen, 1910: 413).
DISTRIBUTION. Argentina. World: Palaearctic region, India.
BIOLOGY. Parasites of nymphs of Psyllidae (Homoptera) (in Europe).
COMMENTS. The holotype j of fuscipalpis is apparently lost, but from the original description and
biology it is quite unlikely that it is correctly placed in Prionomitus. It may be a Cerchysius.
Howard (in Riley et al., 1894: 97) recorded Prionomitus tiliaris (Dalman) from St Vincent.
There is one specimen of the original Smith collection from St Vincent labelled as this species by
Howard in the BMNH. It is a female Psyllaephagus rotundiformis (Howard).
PROCH1LONEURUS Silvestri
(Key couplets: 9 58, 95, 107; $ 136. Fig. 35)
Prochiloneurus Silvestri, 19156: 350. Type-species: Prochiloneurus pulchellus Silvestri, by monotypy.
Achrysopophagus Girault, 191 5d: 89. Type-species: Achrysopophagus oviductus Girault, by original
designation.
Parachrysopophagus Agarwal, 1965: 65. Type-species: Achrysopophagus insolitus Alam, by original
designation.
Neoprochiloneurus Viggiani, 1966: 95. Type-species: Prochiloneurus bolivari Mercet, by monotypy.
INCLUDED SPECIES. Twenty-nine, four of which are found in the Neotropics : dactylopii (Howard)
(=argentinensis De Santis, 1964: 357. Syn. n.), io (Girault, 1920: 187), rex (Girault, 1920: 188)
and seini (Dozier, 1927: 269); also three undetermined species (BMNH; USNM, Washington).
DISTRIBUTION. Throughout the Neotropics. World: cosmopolitan.
224 JOHN S. NO YES
BIOLOGY. Hyperparasites via other encyrtids parasitizing various families of Coccoidea
(Homoptera), mainly Pseudococcidae and Coccidae, and Coccinellidae (Coleoptera).
COMMENTS. I have labelled one syntype female of Cheiloneurus dactylopii as LECTOTYPE
(USNM, Washington). It is labelled as follows: '117001, Cotype 2641 USNM, "Cheiloneurus
dactylopii How. $ Cotype.'"
Placed in the tribe Cheiloneurini.
PROTYNDARICHOIDES gen. n.
(Key couplet: 9 193. Figs 199-201)
Type-species: Protyndarichoides nigriceps sp. n.
$. Head. In side view moderately deep, with frontovertex and face more or less gradually curved, but
slightly angled just above antennal scrobes. Eye about one-third longer than broad, clothed with a few short
inconspicuous pale setae and reaching occipital margin which is sharp. Malar space about two-thirds length
of eye and with malar sulcus present. Frontovertex about one-third head width, ocelli forming a slightly
acute angle, posterior ocellus separated from occipital margin by less than its own diameter and from eye
margin by one half to slightly more than its own diameter. Antennal scrobes shallow and short, not
reaching half way from antennal toruli to anterior ocellus and separated dorsally by interantennal
prominence which is confluent with frontovertex; antennal torulus separated from mouth margin by
slightly more than its own length and from other torulus by about twice its length, its upper margin well
below lowest margin of eye. Scape subcylindrical, pedicel conical and about as long or longer than the first
two funicle segments together, funicle six-segmented, cylindrical and widening distally, clava three-
segmented, about half as long as, and slightly broader than funicle, with a short, oblique, apical truncation;
longitudinal sensilla on all flagellar segments except first two. Frontovertex with shallow coriaceous
sculpture which becomes shallower and more elongate between eyes and antennal scrobes and on lower
parts of face and genae. Frontovertex with a few scattered brown setae, more dense on interantennal
prominence. Mandibles with three acute teeth, maxillary palpi four-segmented, labial palpi three-
segmented.
Thorax. In side view moderately deep with mesoscutum and scutellum quite flat. Mesoscutum without
notaular lines, about a half broader than long, axillae more or less touching, scutellum slightly longer than
broad, flat and with apex rounded, propodeum medially about one-quarter as long as scutellum. Pronotum,
mesoscutum and axillae with shallow coriaceous sculpture, scutellum with deep, regular, reticulate
sculpture but smooth at extreme apex and sides, sculpture on mesopleuron very shallow, elongate anteriorly
and more or less reticulate posteriorly, propodeum with a few longitudinal carinae medially and spiracles
more or less enclosed by three carinae. Forewing hyaline, slightly less than three times as long as broad ;
basal cell naked in proximal half, speculum not interrupted and open ; submarginal vein slightly expanded
apically and with an apical hyaline break, marginal vein about four to five times as long as broad, about
twice as long as stigmal which is about as long as postmarginal. Hindwing about five times as long as broad.
Mid tibial spur about as long as corresponding basitarsus.
Caster. About two-thirds as long as thorax, with cerci about half way along, paratergites absent,
hypopygium nearly reaching apex of gaster, ovipositor very slightly exserted. Ovipositor short, about half
as long as mid tibia, gonostyli slightly less than half length of mid tibial spur.
<$. Unknown.
Protyndarichoides nigriceps sp. n.
(Figs 199-201)
$. Length: 1-06 mm.
Colour. Frontovertex blackish, face and genae blackish with metallic green and purple reflections ; scape,
pedicel and first five funicle segments yellow, sixth funicle segment and clava black ; thorax largely reddish
orange except pronotum, anterior margin of mesoscutum and scutellum which are dark brown with some
dark green reflections, legs completely yellow except fore and mid coxae which are yellowish white ; gaster
black dorsally, except basal tergite which is orange with two dark spots laterally, basal half of ventral
surface of gaster orange, distal half black.
NEOTROPICAL ENCYRTIDAE 225
Head. Posterior ocellus separated from occipital margin by slightly less than its own diameter and from
eye margin by about half of its own diameter. Relative measurements of holotype: head width 59,
frontovertex width at anterior ocellus 20-5, malar space 25, eye length 37, eye width 30, POL 8, OOL 2,
scape length 35, proportions of antenna as in Fig. 200.
Thorax. Mid tibial spur slightly shorter than corresponding basal tarsal segment. Relative measurements
of forewing of holotype: length 174, maximum width 63, length of submarginal vein 71, of marginal vein
14, of stigmal vein 7, of postmarginal vein 5-5. Base of forewing as in Fig. 199.
Caster. Relative measurements of paratype: length of ovipositor 28, of gonostylus 6 [mid tibia length 52,
mid tibial spur length 12-5]. Ovipositor as in Fig. 201.
c?. Unknown.
DISTRIBUTION. Trinidad, Tobago.
BIOLOGY. Unknown.
MATERIAL EXAMINED
Holotype $, Trinidad: St George, Caura, 2.viii.l976 (/. S. Noyes) (BMNH).
Paratype. Tobago: 1 ?, Mt Irvine Bay, 19.vii.1976 (J. S. Noyes) (BMNH).
COMMENTS. Also examined, one further undescribed species from Brazil (BMNH).
In some respects this genus is similar to Rhytidothorax, i.e. in the flat scutellum, the relatively
long propodeum and the short ovipositor. However, the forewing venation suggests a close
affinity to the genera included in the tribe Cheiloneurini, and it may be closest to
Parechthrodryinus. This is suggested by the structure of the scutellum, propodeum and venation,
although the relatively short gaster and ovipositor distinguish it.
PSEUDAPHYCUS Clausen
(Key couplets : 9 19; J 24)
Pseudaphycus Clausen, 1915: 41. Type-species: Aphycus angelicas Howard, by monotypy.
Psilomirinus Brethes, 19166: 424. Type-species: Psilomirinus flavidulus Brethes, by monotypy.
INCLUDED SPECIES. Twenty-three, eleven of which are found in the Neotropics : abtrusus Gahan,
angelicas (Howard), angustifrons Gahan, dysmicocci Bennett, ferrisianae Bennett, flavidulus
(Brethes), griseus De Santis, mundus Gahan, perdignus Compere & Zinna, 1955: 105, prosopidis
Timberlake and utilis Timberlake; also two undetermined species from Trinidad and Brazil
(BMNH).
REFERENCES. Revision: Gahan (1946); see also Bennett (1955) and De Santis (1964: 150-161).
DISTRIBUTION. Throughout the Neotropics. World: cosmopolitan.
BIOLOGY. Parasites of Pseudococcidae (Homoptera).
COMMENTS. Placed in the tribe Aphycini, subtribe Aphycina. It is very close to Acerophagus and
Timberlakia from which it can be separated using the characters given in the key.
PSEUDHOMALOPODA Girault
(Key couplets : $ 6; 3 ?7)
Pseudhomalopoda Girault, 1915c: 171. type-species: Pseudhomalopoda prima Girault, by monotypy.
INCLUDED SPECIES. Four, one of which is found in the Neotropics: prima Girault, 1915c: 171.
DISTRIBUTION. Mexico, West Indies. World: Australia, Indonesia, Japan, Hawaii.
BIOLOGY. Parasites of Diaspididae (Homoptera).
COMMENTS. Placed in the tribe Habrolepidini, subtribe Habrolepidina.
226 JOHN S. NOYKS
PSYLLAEPHAGUS Ashmead
(Key couplets: 9 171, 198, 254; j1 82. Figs 72, 73, 75, 78)
Psyllaephagus Ashmead, 1900: 382. Type-species: Encyrtus pachypsyllae Howard, by original designation.
Mirocerus Ashmead, 1904: 309. Type-species: Mirocerm pyelae Ashmead, by monotypy.
Calluniphilus Erdos, 1961 : 413. Type-species: Calluniphilus vendicus Erdos, by monotypy.
Ooencyrtoides Hoffer, 1963: 568. Type-species: Ooencyrtus albopilosus Hoffer, by original designation.
Propsyllaephagus Blanchard in De Santis, 1964: 235. Type-species: Propsyllaephagus trellesi Blanchard, by
monotypy.
Mercetia Bakkendorf, 1965: 139. Type-species: Copidosoma lusitanicum Mercet, by monotypy.
Kaszabicyrtus Szelenyi, 1971: 389. Type-species: Kaszabicyrtus acutigastris Szelenyi, by original
designation.
INCLUDED SPECIES. One hundred and four, two of which are found in the Neotropics:
rotundiformis (Howard; Noyes, 1979: 165) and trellesi (Blanchard; De Santis, 1964: 236); also
four undetermined species from Cuba and Brazil (BMNH).
DISTRIBUTION. Argentina, Brazil, Trinidad, Tobago, St Vincent, Cuba. World: cosmopolitan.
BIOLOGY. Parasites of nymphs of Psyllidae (Homoptera).
COMMENTS. The generic placement of trellesi is not too clear since all the types are badly mounted
on slides. Trjapitzin (pers. comm.) believes that it may be an Aphidencyrtus (sometimes parasitic
on psyllid nymphs), but the wing venation indicates that it is best accommodated in
Psyllaephagus. Placed in the tribe Trechnitini, subtribe Metaprionomitina. The two genera most
easily confused with it in the Neotropics are Aphidencyrtus and Ooencyrtus (tribe Microteryini)
which can be distinguished by the characters given in the key.
RHOPUS Forster
(Key couplets: 9 37, 72, 208, 214; ^ 86, 94. Fig. 19)
Rhopus Forster, 1856: 34. Type-species: Encyrtus piso Walker, by monotypy.
Xanthoencyrtus Ashmead, 1902: 302. Type-species: Xanthoencyrtus nigroclavatus Ashmead, by monotypy.
Scelioencyrtus Girault, \9\5d: 161. Type-species: Scelioencyrtus nigriclavus Girault, by original
designation.
Mirastymachus Girault, \9\5d: 166. Type-species: Mirastymachus europaeus Girault, by original
designation.
Pholidoceras Mercet, 1918: 237. Type-species: Pholidoceras brachyptera Mercet, by monotypy.
Pholidocerodes Ferriere, 1956: 358. Type-species: Pholidoceras parvula Mercet, by monotypy.
INCLUDED SPECIES. Forty-one, two of which are found in the Neotropics: desantisellus
Ghesquiere, 1957: 18 and nigroclavatus (Ashmead, 1902: 302).
DISTRIBUTION. Argentina. World: cosmopolitan.
BIOLOGY. Parasites of Pseudococcidae (Homoptera).
COMMENTS. Placed in the tribe Anagyrini, subtribe Rhopina.
RHYTIDOTHORAX Ashmead
(Key couplets: 9 85, 120, 133, 164, 191, 209, 226, 237; 3 45, 93. Figs 54-56, 121)
Rhytidothorax Ashmead, 1900: 377. Type-species: Rhytidothorax marlatti Ashmead, by monotypy.
INCLUDED SPECIES. Six, five of which are found in the Neotropics: bollowi (Mercet, 1928: 10)
(comb. n. from Parencyrtus), flaviclava (De Santis, 1964: 95) (comb. n. from Parastenoterys),
flavicornis (De Santis, 1967: 74) (comb. n. from Parastenoterys), hirtus (Howard; Noyes, 1979:
159) (comb. n. from Encyrtus] and perhispidus (De Santis, 1964: 97) (comb. n. from
Parastenoterys) ; also at least 25 undetermined species (BMNH ; CNC, Ottawa).
DISTRIBUTION. Argentina, Brazil, Panama, Ecuador, West Indies. World: Nearctic region,
?Africa, Australia.
NEOTROPICAL ENCYRTIDAE 227
BIOLOGY. Unknown.
COMMENTS. I have examined the holotype ? of Rhytidothorax marlatti. In my opinion it is
congeneric with the South American species previously incorrectly placed in Parastenoterys
Girault (this genus is distinct from Rhytidothorax) and differs only in the shape of the head
(similar in shape to Simondsiella, Figs 82, 83) and in having stronger setigerous punctures on the
frontovertex. The shape of the head of the South American species varies greatly and in most it it
relatively deep, although some do approach that of marlatti. A few also have well-marked
setigerous punctures on the frontovertex. The genus also occurs in Australia and Africa except
that the African species appear to have a short third (upper) tooth.
The genus appears to be morphologically diverse; in some groups of species the head is very
deep and almost similar in shape to some Hunterellus, and the fore tibia is quite expanded and
flattened with the apical tarsal segment from slightly to greatly enlarged. In this group also, the
funicle segments are relatively short and the frontovertex and scutellum are sometimes highly
polished, although occasionally quite deeply sculptured. This group is perhaps the most self-
contained, although it is difficult to reliably separate it from Rhytidothorax. The group includes
perhispidus.flavicornis and hirtus. The other main group contains species with a shallower head,
more deeply sculptured scutellum and longer funicle segments. At either extreme the species are
very different, but taken as a whole it seems that they would all be best placed in a single genus.
All species have the eyes and frontovertex with long conspicuous setae, the mandibles
unidentate or bidentate (upper tooth relatively long) or perhaps with a very short third (upper)
tooth, the scutellum flat and in larger species with a very fine apical carina with two upright setae
at apex, a relatively long propodeum, medially at least one-sixth as long as the scutellum and
with distinct sculpture, a short ovipositor about half the length of the gaster and a hypopygium
which does not reach the apex of the gaster. In addition the antenna is quite characteristic :
pedicel relatively long, first funicle segment relatively long and following segments progressively
becoming shorter.
Trjapitzin & Gordh (19786) place the genus in the tribe Bothriothoracini, subtribe
Bothriothoracina, which I believe is incorrect. The marginal vein is relatively long, the
hypopygium does not extend to the apex of the gaster and the mandibles are narrower and
usually only with one or two very unequal teeth. The characters are conflicting with those given
by Trjapitzin for this tribe.
SHENAHETIA gen. n.
(Key couplets: ? 74, 221; J 129. Figs 41, 137, 202-206)
Type-species: Shenahetia masneri sp. n.
9. Head. In side view similar to that of Mariola (Fig. 42), very shallow, hardly convex and prognathous, the
occipital foramen situated more than half way up head. Eye about one half times longer than broad, with
short to moderately long hairs and nearly reaching occipital margin which is acute. Malar space about half
length of eye with malar sulcus absent. Frontovertex just greater than half head width; ocelli forming an
obtuse angle, posterior ocellus situated less than its own diameter from occipital margin and about its own
diameter from eye margin. Antennal scrobes very shallow and short (Fig. 41), not as long as an antennal
torulus ; antennal torulus situated almost at mouth margin and separated from other torulus by about two
and a half to three times its own length. Scape subcylindrical or foliaceously flattened, pedicel conical and
about half as long as funicle which is six-segmented, all segments transverse, broadening apically and more
or less cylindrical, clava two-segmented, broader than funicle and rounded apically ; longitudinal sensilla
on fifth and sixth funicle segments and clava. Frontovertex with a few sparse setae, sculpture absent or
extremely shallow and irregular so that it has a polished appearance. Mandibles narrow, with three acute
teeth, the middle the longest, maxillary palpi four-segmented, labial palpi three-segmented.
Thorax. In side view quite shallow and dorsally flattened. Mesoscutum without notaular lines, about one
half broader than long, axillae almost meeting, scutellum slightly broader than long with apex rounded and
produced as a short flange, propodeum not more than one-fifth as long as scutellum. Pronotum and
mesoscutum with very shallow shagreened sculpture, scutellum devoid of sculpture and highly polished,
propodeum smooth except for some very shallow sculpture medially. Forewing hyaline or slightly infumate,
228 JOHN S. NOYES
about two and a half times as long as broad ; basal cell sparsely hairy and naked in its proximal one-third,
speculum not interrupted and open ; submarginal vein slightly expanded apically and with an apical hyaline
break, marginal vein about three times as long as broad, stigmal slightly shorter, postmarginal very short.
Mid tibial spur about as long as mid basal tarsal segment.
Caster. About one-half longer than thorax, cerci at about two-fifths along gaster, paratergites absent,
hypopygium extending past apex of gaster so that it is clearly visible in dorsal view, exserted part of
ovipositor about half as long as gaster and with sheaths distinctly downcurved apically. Ovipositor (Fig.
206) about twice as long as mid tibia with gonostylus slightly longer than mid tibia.
3. Body moderately dorso-ventrally flattened. Antennal scrobes semicircular, slightly longer than in
female, antennal torulus just less than its own length from mouth margin and separated from other torulus
by about its own length ; antenna with pedicel about as long as first two funicle segments together, funicle
segments about as long as broad or slightly longer than broad and clothed with setae which are more than
twice as long as diameter of segments, clava entire, longitudinal sensilla present on sixth funicle segment
and clava. Gaster slightly shorter than thorax; aedeagus about half as long as mid tibia and digiti each with
one apical spine.
Shenahetia masneri sp. n.
(Figs 41, 137, 202-205)
$. Length (excluding ovipositor): 0-92-1-00 mm (holotype 1-00 mm).
Colour. Body dark brown with some purplish and golden reflections on face and thorax, basal tergite of
gaster with bluish green reflections ; scape and pedicel testaceous with dorsum of scape and basal half of
pedicel pale brown, flagellum brown ; legs testaceous yellow, except the following which are brown : fore
and hind femora, distal half of mid femur, extreme base of each tibia and apical segment of hind tarsus ;
forewing hyaline; occasionally ovipositor sheaths paler towards base, but usually pale brown.
Head. Frontovertex with extremely shallow, fine, shagreened sculpture which is slightly deeper and more
irregular between ocelli and more elongate on genae; a few scattered setae near ocelli, along inner eye
margins, on clypeus and genae; ocelli forming an angle of about 115°; eyes clothed in short pale setae; fifth
funicle segment without longitudinal sensilla. Relative measurements of paratype: head width 46,
fronto vertex width at anterior ocellus 25, malar space 13, eye length 28, eye width 18, scape length 21,
proportions of antennae as in Fig. 202.
Thorax. Dorsum of thorax with a few scattered dark setae on pronotum and mesoscutum, scutellum with
fewer setae; mid tibial spur shorter than mid basal tarsal segment. Relative measurements of forewing of
paratype: length 120, maximum width 43, length of submarginal vein 45, of marginal vein 6-5, of stigmal
vein 5, of postmarginal vein 1 ; of hindwing: length 85, maximum width 15. Base of forewing as in Fig. 203.
Gaster. Exserted part of ovipositor about three-quarters as long as mid tibia or nearly half as long as
gaster. Relative lengths of paratype: length of ovipositor 78, of gonostylus 33 (mid tibia 40).
S. Length: 0-75-1 -00 mm.
Similar to female except for antenna (Fig. 204) and genitalia (Fig. 205). Relative measurements of
paratype: head width 47, frontovertex width at anterior ocellus 26-5, scape length 19-5, proportions of
antennae as in Fig. 204, length of hindwing 80, width of hind wing 20.
DISTRIBUTION. Trinidad.
BIOLOGY. Unknown.
MATERIAL EXAMINED
Holotype $, Trinidad: St George, Chaguaramas Bay, 16.vii.1976 (J. S. Noyes) (BMNH).
Paratypes. Trinidad: 4 $, Curepe, Santa Margarita Circular Road, iii-iv.1974, Moericke trap; 2 3,
Curepe, Santa Margarita Circular Road, 24.v.-8.vi.l974 and 15-27.ix.1974, Moericke trap (F. D. Bennett);
1 ^, Caroni, Gran Couva, 27.vi.1976 (/. S. Noyes). (BMNH; CNC, Ottawa.)
COMMENTS. I have also examined one other undescribed species from Trinidad which differs from
the type-species in having the scape broadened and flattened, the frontovertex with conspicuous
setae and the forewings slightly infumate.
The genus is similar in appearance to some species of A vetianella and Szelenyiola but differs in
NEOTROPICAL ENCYRTIDAE 229
lacking the membranous lines on the frontovertex. It is very probably closely related to
Tanyencyrtus from which it can be separated by the characters given in the key.
Probably best placed in the tribe Microteryini near the subtribes Oobina and Platencyrtina.
Named in honour of Dr L. Masner.
S1MMONDS1ELLA gen. n.
(Key couplet: 9 170. Figs 81-83, 207)
Type-species: Simmondsiella flaviptera sp. n.
9- Head. In side view relatively shallow, similar to Trichomasthus but rather more evenly curved. Eye
about one-third longer than broad, with a few very short hairs and overreaching occipital margin which is
sharp. Malar space about half as long as eye, with malar sulcus present but weak. Frontovertex about one-
quarter head width ; ocelli forming an acute angle of about 60°, posterior ocellus separated from occipital
margin by about its own diameter and from eye margin by less than half its diameter. Antennal scrobes
semicircular, shallow and short, meeting dorsally but not reaching half way between antennal toruli and
anterior ocellus, antennal torulus separated from mouth margin and other torulus by about its own length,
its dorsal margin about level with lowest margin of eyes. Antennal scape subcylindrical and with two very
strong, conspicuous setae on outer face near apex, pedicel conical and longer than first funicle segment,
funicle six-segmented, all segments longer than broad, cylindrical and slightly widening distally, clava three-
segmented and with apex rounded. Frontovertex with regular, reticulate, almost punctiform sculpture,
becoming shallower and more irregular on lower parts of face and genae, a few sparse setae present.
Mandibles with two acute teeth and a broad truncation which is about as broad as the two teeth together
across their bases.
Thorax. Moderately deep in side view and dorsally convex. Mesoscutum without notaular lines, about
twice as broad as long, axillae widely separated by posterior margin of mesoscutum, scutellum slightly
wider than long, evenly convex and with apex rounded, propodeum medially about one-eighth as long as
scutellum. Pronotum, mesoscutum and axillae with shallow coriaceous sculpture, scutellum with basal half
similar but sculpture more regular and tending towards punctiform, apical half completely smooth and
polished, mesopleuron with raised elongate sculpture anteriorly becoming more regular and reticulate
posteriorly, propodeum with shallow, irregular sculpture medially. Forewing hyaline, slightly more than
twice as long as broad ; basal cell sparsely pilose and naked in proximal third, speculum not interrupted and
open ; submarginal vein with an apical hyaline break, marginal vein punctiform, stigmal vein about as long
as marginal and postmarginal veins together. Mid tibial spur about as long as mid basitarsus.
Caster. Slightly shorter than thorax and with cerci in basal third, ovipositor only slightly protruding,
apex of hypopygium about two-thirds along gaster.
c£. Unknown.
Named in honour of Dr F. J. Simmonds.
Simmondsiella flaviptera sp. n.
(Figs 81-83, 207)
9. Length: 1-24 mm.
Colour. Frontovertex dark purplish green, paler and more shiny on antennal scrobes, interantennal
prominence and cheeks below eyes tinged coppery ; antenna testaceous with basal half of pedicel and
dorsum of scape brown ; pronotum and mesoscutum with long, dark brown setae, pronotum, mesoscutum,
axillae and anterior half of scutellum dark purplish brown ; apical half of scutellum very shiny deep blue
mixed with purple, mesopleuron brown ; coxae and femora brown, mid and hind femora with apices yellow,
tibiae and tarsi yellow; forewings faintly suffused yellow, particularly towards base; gaster with basal-
tergite dark metallic green, other tergites coppery mixed metallic pale green.
Head. Relative measurements (Figs 82, 83): head width 101, frontovertex width at anterior ocellus 26,
malar space 31, eye length 64, eye width 48, POL 14, OOL 2, scape length 42, proportions of antenna as in
Fig. 207.
Thorax. Relative measurements of forewing (Fig. 81): length 222, maximum width 100, length of
230 JOHN s- NOYES
submarginal vein 88, of marginal vein 6, of stigmal vein 15-5, of postmarginal vein 9-5; of hind wing: length
148, maximum width 43.
cJ. Unknown.
DISTRIBUTION. Brazil.
BIOLOGY . Unknown.
MATERIAL EXAMINED
Holotype ?, Brazil: Santa Catarina, Nova Teutonia, 25.viii.1944 (F. Plaumann) (BMNH).
COMMENTS. Close to Trichomasthus and Hadrencyrtus Annecke & Mynhardt (tribe Microteryini,
subtribe Microteryina). It differs from both in having a punctiform marginal vein, and from
Hadrencyrtus in having subhyaline forewings, eyes overreaching the occipital margin, a shorter
malar space and long setae on the antennal flagellum. Both Trichomasthus and Hadrencyrtus
have relatively long marginal veins (usually at least three times as long as broad) and
Hadrencyrtus has infuscate forewings, eyes only just reaching occipital margin, a longer malar
space (about three-quarters of the length of the eye) and the setae on the antennal flagellum very
short.
SOLENAPHYCUS De Santis
(Key couplets: 9 9, 25; $ 13. Fig. 8)
Solenaphycus De Santis, \912b: 54. Type-species: Solenaphycus vianai De Santis, by monotypy.
INCLUDED SPECIES. One: vianai De Santis, 19726: 54.
DISTRIBUTION. Argentina.
BIOLOGY. Unknown.
COMMENTS. I am unable to place the genus according to Trjapitzin's (19736) classification of the
Encyrtidae.
SOLENOENCYRTUS De Santis
(Key couplet : V 232)
Solenoencyrtus De Santis, 1964: 207. Type-species: Solenoencyrtus platensis De Santis, by monotypy.
INCLUDED SPECIES. One: platensis De Santis, 1964: 207.
DISTRIBUTION. Argentina.
BIOLOGY. Unknown.
COMMENTS. Possibly related to Metaphycus (tribe Aphycini, subtribe Paraphycina).
STEMMATOSTERES Timberlake
(Key couplets: 9 9; J1 14. Fig. 11)
Stemmatosteres Timberlake, 1918: 352. Type-species: Stemmatosteres apterus Timberlake, by monotypy.
INCLUDED SPECIES. Three, one of which is found in the Neotropics: apterus Timberlake, 1918:
354.
DISTRIBUTION. Uruguay. World : Nearctic region, Europe.
BIOLOGY. Parasites of Pseudococcidae (Homoptera).
COMMENTS. Placed in the tribe Aphycini, subtribe Aphycina.
SYRPHOPHAGUS Ashmead
(Key couplets: 9 176, 205, 259; J 84, 121, 123, 140. Fig. 79)
NEOTROPICAL ENCYRTIDAE 231
Syrphophagus Ashmead, 1900 : 397. Type-species : Encyrtus mesograptae Ashmead, by original designation.
Echthrobaccha Perkins, 1906: 253. Type-species: Echthrobaccha injuriosa Perkins, by monotypy.
Syrphidencyrtus Blanchard, 1940: 107. Type-species: Syrphidencyrtus bacchae Blanchard, by monotypy.
Syn. n.
INCLUDED SPECIES. Twenty-nine, six of which are found in the Neotropics : bacchae (Blanchard)
(comb. n. from Syrphidencyrtus), flavitibiae (De Santis) (comb. n. from Syrphidencyrtus),
nigricornis (De Santis) (comb. n. from Syrphidencyrtus), nubeculus De Santis, quadrimaculati
(Ashmead, 1881: 171) and silvai (Brethes, 1921a: 8), also three undetermined species from
Trinidad and Brazil (BMNH).
REFERENCE. De Santis (1964: 128-135, 194).
DISTRIBUTION. Argentina, Chile, Brazil, Trinidad, Puerto Rico. World: cosmopolitan.
BIOLOGY. Parasites of larvae of Syrphidae (Diptera).
COMMENTS. I have examined a specimen determined by De Santis as Syrphidencyrtus bacchae.
Syrphophagus flavitibiae may be out of place in this genus since the scutellum appears to be very
flat and shiny but this may be because the holotype is slide mounted and the specimen slightly
flattened by the pressure of the coverslip.
The genus is close to Aphidencyrtus and they are sometimes almost impossible to separate,
except on relative size, unless the host is known. I believe that the two genera should eventually
be synonymized.
Placed in the tribe Microteryini, subtribe Syrphophagina.
SZELENYIOLA Trjapitzin
(Key couplets: 9 213; 3 58. Fig. 95)
Szelenyiola Trjapitzin, 1977: 160. Type-species: Szelenyiola nearctica Trjapitzin, by monotypy.
INCLUDED SPECIES. One, not found in the Neotropics, but one undescribed species from Brazil
(BMNH).
DISTRIBUTION. Brazil. World: Nearctic.
BIOLOGY. Probably parasitic on eggs of Buprestidae or Scolytidae (Coleoptera).
COMMENTS. Placed in the tribe Microteryini, subtribe Oobiina.
TACHARDIOBIUS Timberlake
(Key couplets: 9 195; <? 56. Fig. 93)
Tachardiobius Timberlake, 1926: 19. Type-species: Tachardiobius nigricans Timberlake, by monotypy.
INCLUDED SPECIES. Two, one of which is found in the Neotropics : silvestrii De Santis, 1956 : 188.
DISTRIBUTION. Argentina. World: Nearctic region.
BIOLOGY. Parasites of Lacciferidae (Homoptera).
COMMENTS. The South American species is very close to the type-species, but differs in having
more extensive pale markings on the face, two lateral white spots on the base of the gaster and
paler legs.
Placed in the tribe Aphycini, subtribe Paraphycina by Trjapitzin & Gordh (19786).
TANYENCYRTUS De Santis
(Key couplets: ? 74, 221; ^ 129. Fig. 139)
Tanyencyrtus De Santis, \912b: 57. Type-species: Tanyencyrtus divisus De Santis, by monotypy.
INCLUDED SPECIES. One: divisus De Santis, 19726: 57.
10
232 JOHN S. NOYES
DISTRIBUTION. Argentina.
BIOLOGY. Parasites of fruit-mining larvae of Lepidoptera.
COMMENTS. See comments under Shenahetia.
TETARTICLAVA gen. n.
(Key couplet: 9 23. Figs 16, 208, 209)
Type-species: Tetarticlava yoshimotoi sp. n.
9. Head. In side view, moderately deep, most rounded level with top of antennal scrobes, frontovertex
slightly convex. Eye about one-third longer than broad, hairy and reaching occipital margin which is acute.
Malar space slightly longer than half length of eye and with malar sulcus present. Frontovertex at
narrowest point about one-third head width ; ocelli forming an acute angle of about 60°, posterior ocellus
separated from occipital and eye margins by less than its own diameter. Antennal scrobes moderately
impressed, reaching much further than half way from antennal toruli to anterior ocellus and joined
dorsally ; antennal torulus slightly more than its own length from mouth margin and nearly twice its own
length from other torulus, its upper margin slightly below lowest margin of eye. Scape subcylindrical and
stout, pedicel longer than first two funicle segments together, funicle five-segmented, all segments transverse
and cylindrical, broadening distally, clava four-segmented, nearly as long as funicle and with apex slightly
pointed; longitudinal sensilla on all flagellar segments except the first two. Frontovertex with fairly
conspicuous dark setae, sculpture shallow coriaceous, more transversely elongate between anterior ocellus
and top of antennal scrobes, becoming more longitudinally elongate on cheeks. Mandibles with two acute
teeth and a rudimentary third (lowest) tooth, the middle tooth much the longest; maxillary palpi four-
segnented, labial palpi three-segmented.
Thorax. In side view, moderately deep with mesoscutum and scutellum fairly convex. Mesoscutum
without notaular lines and about twice as broad as long, axillae meeting, scutellum slightly broader than
long, propodeum medially about one-eighth as long as scutellum. Pronotum, mesoscutum, axillae and
scutellum with shallow coriaceous sculpture, except on scutellum where it is distinctly deeper, mesopleuron
with irregular, shallow, coriaceous sculpture, propodeum medially quite smooth. Forewing hyaline, but
with some slight infumation in basal cell, slightly more than twice as long as broad; basal cell sparsely hairy
with a completely naked triangular area proximally, speculum not interrupted and open ; submarginal vein
with an apical hyaline break, marginal vein about two to three times as long as broad, about as long as
stigmal and slightly longer than postmarginal. Hindwing nearly four times as long as broad. Mid tibial spur
about as long as mid basal tarsal segment.
Caster. Slightly shorter than thorax, cerci in basal third, hypopygium reaching apex of gaster, ovipositor
not or hardly protruding. Ovipositor about as long as gaster or slightly longer than mid tibia, gonostylus
about one-third as long as mid tibia.
$. Unknown.
Tetarticlava yoshimotoi sp. n.
(Figs 16, 208, 209)
?. Length: 0-73-0-89 mm (holotype 0-81 mm).
Colour. Body black with faint green, purple and brassy reflections ; antennae brown, legs brown except
for tarsi and apex of mid tibia which are testaceous.
Head. Relative measurements of holotype: head width 55, frontovertex width at anterior ocellus 21,
malar space 23, eye length 35-5, eye width 26, POL 7-5, OOL 3-5, scape length 26, proportions of antenna as
in Fig. 16.
Thorax. Relative measurements of forewing of holotype: length 126, maximum width 57, length of
submarginal vein 52, of marginal vein 9, of stigmal vein 8-5, of postmarginal vein 6-5; of hindwing: length
88, maximum width 24. Base of forewing as in Fig. 208.
Gaster. Relative measurements of paratype: length of ovipositor 67, of gonostylus 18 [mid tibia 55].
Ovipositor as in Fig. 209.
J. Unknown.
NEOTROPICAL ENCYRTIDAE 233
DISTRIBUTION. Trinidad, St Vincent.
BIOLOGY. Unknown.
MATERIAL EXAMINED
Holotype ?, Trinidad: St George, Lopinot, 10.viii.1976 (/. S. Noyes) (BMNH).
Paratypes. Trinidad: 1 ?, Curepe, Santa Margarita, Circular Road, 25.x.-7.xii.l974 (M. N. Beg). St
Vincent: 1 ?, Richmond, 10.vii.1976 (J. S. Noyes). (BMNH; CNC, Ottawa.)
COMMENTS. The genus bears some resemblance to Forcipestricis and is possibly related. It differs
in the antennal structure and the lack of tubercles on the scutellum. I am unable to place it
according to Trjapitzin's (19736) classification of the Encyrtidae.
Named in honour of Dr C. Yoshimoto.
TETRACNEMOIDEA Howard
(Key couplets: , 27, j 3. Fig. 24)
Tetrac nemo idea Howard, 18986: 232. Type-species: Tetracnemoidea australiensis Howard, by monotypy.
Tetracnemopsis Ashmead, 1900: 358. Ty pe- species : Tetracnemus westwoodi Cockerell, by monotypy.
Arhopoideus Girault, \9\5d: 174. Type-species: Arhopoideus brevicornis Girault, by original designation.
Hungariella Erdos, 1946: 144. Type-species: Hungariella piceae Erdos, by monotypy.
INCLUDED SPECIES. Sixteen, two of which are found in the Neotropics: peregrina (Compere)
(comb. n. from Arhopoideus) and brevicornis (Girault) ( = pretiosus Timberlake) (comb. n. from
Arhopoideus).
REFERENCE. World revision: Kerrich (1967: 154-166).
DISTRIBUTION. Brazil, Argentina, Chile, Trinidad, Bermuda. World: cosmopolitan.
BIOLOGY. Parasites of Pseudococcidae (Homoptera).
COMMENTS. Placed in the tribe Tetracnemini, subtribe Arhopoideina.
TETRACNEMUS Westwood
(Key couplets : 9 43 ; J 4. Fig. 25)
Tetracnemus Westwood, 1837: 258. Type-species: Tetracnemus diversicornis Westwood, by monotypy.
Tetracladia Howard, 1892: 367. Type-species: Tetracladia texana Howard, by designation of Ashmead,
1900: 358.
Tetralophidea Ashmead, 1900: 348. Type-species: Tetralophidea bakeri Ashmead, by monotypy.
Tetralophiellus Ashmead, 1900: 357. Type-species: Tetralophiellus brivicollis Ashmead, by monotypy.
Paracalocerinus Girault, \9\5d: 142. Type-species: Paracalocerinus australiensis Girault, by monotypy.
Masia Mercet, 19196: 470. Type-species: Masia bifasciatella Mercet, by monotypy.
Anusiella Mercet, 19236: 286. Type-species: Anusia heydeni Mayr, by monotypy.
Placoceras Erdos, 1946: 1. Type-species: Placoceras colocense Erdos, by monotypy.
Comperencyrtus De Santis, 1964: 106. Type-species: Comperencyrtus maculipennis De Santis, by monotypy.
INCLUDED SPECIES. Twenty, one of which is found in the Neotropics : maculipennis (De Santis,
1964: 108; 1968: 150).
DISTRIBUTION. Argentina, Brazil. World: cosmopolitan except apparently Ethiopian region.
BIOLOGY. Parasites of Pseudococcidae (Homoptera).
COMMENTS. Placed in the tribe Tetracnemini, subtribe Tetracnemina.
TIMBERLAKIA Mercet
(Key couplets : $ 14; ^ 23)
Timberlakia Mercet, 1925: 9. Type-species: Acerophagus europaeus Mercet, by monotypy.
234 JOHN S. NOYES
INCLUDED SPECIES. Two, one of which is found in the Neotropics: europaea (Mercet, 1921 : 191).
DISTRIBUTION. Mexico, Trinidad. World: cosmopolitan.
BIOLOGY. Parasites of Pseudococcidae (Homoptera).
COMMENTS. Placed in the tribe Aphycini, subtribe Aphycina.
TINEOPHOCTONUS Ashmead
(Key couplet: 9 107)
Tineophoctonus Ashmead, 1900: 351. Type-species: Phaenodiscus armatus Ashmead, by original
designation.
INCLUDED SPECIES. Two, one of which is found in the Neotropics : armatus (Ashmead ; De Santis,
1964: 364).
DISTRIBUTION. Argentina. World: Nearctic region, Europe.
BIOLOGY. Parasites of gall-inhabiting Tineidae (Lepidoptera), Cynipidae (Hymenoptera) and
larvae of Anobiidae and Cerambycidae (Coleoptera).
COMMENTS. Placed in the tribe Cheiloneurini.
TRICHOMASTHUS Thomson
(Key couplets: 9 206; J 84, 119)
Trichomasthus Thomson, 1876: 142. Type-species: Encyrtus cyaneus Dalman, by designation of Gahan &
Pagan, 1923: 148.
Coccidoxenus Crawford, 1913a: 248. Type-species: Coccidoxenus portoricensis Crawford, by monotypy.
Tetracnemella Girault, \9\5d: 170. Type-species: Tetracnemella australiensis Girault, by original
designation.
Stenoteropsis Girault, 191 5d: 176. Type-species: Stenoteropsis abjectus Girault, by monotypy.
INCLUDED SPECIES. Fifty, four of which are found in the Neotropics : tucumanus (De Santis, 1964:
251) (comb. n. from Coccidoxenus), mexicanus (Girault, 19176: 21) (comb. n. from
Coccidoxenus), brasiliensis (Gomes, 1941: 402; 1942: 36) (comb. n. from Coccidoxenus) and
portoricensis (Crawford, 1913a: 248); also two further undetermined species from Brazil and
Trinidad (BMNH).
DISTRIBUTION. Brazil, Mexico, West Indies. World : cosmopolitan.
BIOLOGY. Parasites of Diaspididae, Coccidae, Pseudococcidae and Eriococcidae (Homoptera).
COMMENTS. The undetermined species from Trinidad is unusual in that the marginal vein of the
female is very long (about nine times as long as broad) and it has been reared from Phenococcus
grenadensis Green & Laing (Homoptera: Pseudococcidae). The clava of the male of this species
has the appearance of being two-segmented in dry-mounted material, but in slide-mounted
material it is clearly only one-segmented.
Placed in the tribe Microteryini, subtribe Microteryina.
TYNDARICHOIDES Girault
(Key couplets: 9 98, 136)
Tyndarichoides Girault, 1920: 189. Type-species: Tyndarichoides mexicanus Girault, by monotypy.
INCLUDED SPECIES. One: mexicanus Girault, 1920: 189.
DISTRIBUTION. Mexico.
BIOLOGY. Unknown.
NEOTROPICAL ENCYRTIDAE 235
COMMENTS. Very close to Metaphycus (tribe Aphycini, subtribe Paraphycina) and can only be
reliably separated on the characters given in the key. The two genera should very probably be
considered synonymous.
ZAOMMA Ashmead
(Key couplets: $ 71, 160, 228, J 146)
Zaomma Ashmead, 1900: 401. Type-species: Encyrtus argentipes Howard, by monotypy.
Apterencyrtus Ashmead, 1905: 5. Type-species: Apterencyrtus pulchricornis Ashmead, by monotypy.
Metallonoidea Girault, 1915c: 170. Type-species: Metallonoidea brittanica Girault, by monotypy.
Chiloneurinus Mercet, 1921 : 646. Type-species: Chiloneurus microphagus Mayr, by monotypy.
Richardsius Alam, 1957: 439. Type-species: Apterencyrtus thomsoniscae Alam, by original designation.
Metapterencyrtus Tachikawa, 1963: 213. Type-species: Metapterencyrtus eriococci Tachikawa, by
monotypy.
INCLUDED SPECIES. Fourteen, three of which are found in the Neotropics : argentipes (Howard ;
Noyes, 1979: 166), epytus (Walker, 1839: 69) (comb. n. from Encyrtus) and lambinus (Walker)
(=microphagus Mayr); De Santis, 1964: 361); also six or seven further undetermined species
from Brazil, Argentina and Trinidad (BMNH; CNC, Ottawa).
REFERENCE. Key to species: Gordh & Trjapitzin (1979).
DISTRIBUTION. Peru, Argentina, Brazil, Trinidad and St Vincent. World: Nearctic, Ethiopian
and Palaearctic regions.
BIOLOGY. Hyperparasites of other chalcids, mainly Encyrtidae, parasitizing Diaspididae
(Homoptera).
COMMENTS. The single female of Encyrtus epytus in the collections of the BMNH (labelled
'Encyrtus Epytus Walker, 1302a Bahia') is here designated LECTOTYPE. It differs from the
other South American species in having all the funicle segments subequal in size and
subquadrate.
Placed in the tribe Cheiloneurini.
ZAPLATYCERUS Timberlake
(Key couplets: ? 46; J1 36. Fig. 27)
Zaplatycerus Timberlake, 1925: 173. Type-species: Zaplatycerus fullawayi Timberlake, by monotypy.
INCLUDED SPECIES. Two : fullawayi Timberlake and planiscutellum Kerrich.
REFERENCE. Revision: Kerrich (1978: 152-155).
DISTRIBUTION. Colombia, Panama, Trinidad.
BIOLOGY. Parasites in Pseudococcidae (Homoptera).
COMMENTS. Placed in the tribe Chrysoplatycerini, subtribe Chrysoplatycerina.
ZARHOPALVS Ashmead
(Key couplets: ? 51; 3 39. Fig. 28)
Zarhopalus Ashmead, 1900: 406. Type-species: Zarhopalus sheldoni Ashmead, by original designation.
Anagyrella Girault, 191 5c: 168. Type-species: Anagyrella corvina Girault, by monotypy.
INCLUDED SPECIES. Five, three of which are found in the Neotropics : clavatus Kerrich, inquisitor
(Howard) and putophilus Bennett; also three undetermined species from Brazil, Belize and
Trinidad (BMNH; CNC, Ottawa).
REFERENCE. Revision: Kerrich (1978: 126-131).
DISTRIBUTION. Brazil, Trinidad, Belize and Mexico. World : Nearctic region.
236
JOHN S. NOYES
BIOLOGY. Parasites of Pseudococcidae (Homoptera).
COMMENTS. Placed in the tribe Neodiscodini (Encyrtinae) by Trjapitzin (19736) but transferred
later to the Tetracneminae by Trjapitzin & Gordh (19786).
ZETETICONTUS Silvestri
(Key couplets: ? 84, 182, 225; 3 41, 72, 107)
Aratus Howard, 1897: 155. Type-species: Aratus scutellatus Howard, by monotypy. [Homonym of Aratus
Milne-Edwards, 1853.]
Zeteticontus Silvestri, 19156: 343. Type-species: Zeteticontus abilis Silvestri, by monotypy.
Mirrencyrtus Girault, \9\5d: 115. Type-species: Mirrencyrtus glabriscutellum Girault, by monotypy.
Aratiscus Ghesquiere, 1946: 368. [Replacement name for Aratus Howard.]
INCLUDED SPECIES. Thirteen, three of which are found in the Neotropics : insularis (Howard),
laevigatus (De Santis, 19726 : 55) and scutellatus (Howard), also three or four undescribed species
from Brazil, Ecuador and Trinidad (BMNH; CNC, Ottawa).
REFERENCES. Revision: Subba Rao (1972); key to South American species: Noyes (1979:
167-168).
DISTRIBUTION. Throughout the Neotropics. World : cosmopolitan.
BIOLOGY. Parasites of larvae of Nitidulidae and Silvanidae (Coleoptera).
COMMENTS. Placed in the tribe Bothriothoracini, subtribe Coenocercina.
Host index
ARACHNIDA
ACARINA
Ixodidae
Hunterellus
INSECTA
NEUROPTERA
Chrysopidae
Isodromus, Moorella
Hemerobiidae
Isodromus
ORTHOPTERA
Meromyzobia
BLATTODEA
Comperia
HETEROPTERA
Ooencyrtus
Pentatomidae
Hexacladia, Ooencyrtus
Scutelleridae
Hexacladia
HOMOPTERA
AUCHENORRHYNCHA
Cheiloneurus (via Dryinidae), Helegonatopus
(via Dryinidae)
Cercopidae
Carabunia
Delphacidae
Echthrodryinus (via other hymenopterous
parasites)
Membracidae
Prionomastix
STERNORRHYNCHA
Aphididae
Aphidencyrtus
Psyllidae
Aphidencyrtus, Prionomitus, Psyllaephagus
COCCOIDEA
Aclerdidae
IBennettisca, Cheiloneurus (via other
Encyrtidae), Desantisella (via
Chamaemyiidae), Microterys, Mucrencyrtus,
IParamucrona
Asterolecaniidae
Habrolepis
Coccidae
Ammonoencyrtus, Anicetus, Cheiloneurus (via
other Encyrtidae and Aphelinidae),
Coccidaphycus, Coccidoctonus (via other
Encyrtidae and Pteromalidae),
Diver sinervus, Gahaniella (via other
Encyrtidae), Metaphycus, Microterys,
Prochiloneurus (via other Encyrtidae),
Trichomasthus
Dactylopiidae
A nag yr us
NEOTROPICAL ENCYRTIDAE
237
Diaspididae
Adelencyrtus, Ameromyzobia, Anabrolepis,
Aphycomorpha, Arrhenophagoidea,
Arrhenophagus, Cheiloneurus (via other
Encyrtidae and Aphelinidae),
Coccidencyrtus, Comperiella, Habrolepis,
IHomalopoda, Metaphycus,
Neococcidencyrtus, Plagiomerus,
Pseudhomalopoda, Trichomasthus, Zaomma
(via other Encyrtidae)
Eriococcidae
A cerophago ides, A ndinoencyrtus,
Aphycomorpha, Atelaphycus, Metaphycus,
Microterys, Trichomasthus
Kermesidae
Microterys
Kerriidae
Metaphycus, Microterys, Parechthrodryinus,
Tachardiobius
Lecanodiaspididae
Genus A, Microterys
Margarodidae
Aztecencyrtus, Brethesiella, Cheiloneurus (via
other Encyrtidae), Homosemion
Ortheziidae
Cheiloneurus (via Drosophilidae)
Pseudococcidae
Acerophagus, Acroaspidia, Aenasius,
Aeptencyrtus, Anagyrus, Anarhopus,
Anathrix, lAnusioptera, Apoanagyrus,
Blepyrus, Bothriocraera, Chalcaspis,
Cheiloneurus (via Encyrtidae),
Chrysoplatycerus, Cirrhencyrtus,
Coccidoctonus (via Encyrtidae and
Pteromalidae), Coelaspidia, Ericydnus,
Euryrhopalus, Gahaniella (via Encyrtidae),
Gyranusoidea, Hambletonia, Leptomastix,
Leptomastidea, Microterys, Neodusmetia,
Parechthrodryinus, Pauridia,
IPelmatencyrtus, IPheidoloxenus,
Prochiloneurus (via other Encyrtidae),
Pseudaphycus, Rhopus, Stemmatosteres,
Tetracnemoidea, Tetracnemus, Timberlakia,
Zaplatycerus, Zarhopalus
LEPIDOPTERA
Eggs: Ooencyrtus
Larvae : Copidosoma, Echthrodryinus (via
other parasitic Hymenoptera), Epiencyrtus,
Litomastix, INeapsilophrys, Tanyencyrtus
Gelechiidae
Apsilophrys, Paralitomastix
Gracillariidae
Paraleurocerus
Lyonetiidae
Echthrodryinus (via other parasitic
Hymenoptera), Parablastothrix
Nepticulidae
Parablastothrix
Pterophoridae
Ageniaspis
Pyralidae
Paralitomastix, Pentalitomastix
Tineidae
Ageniaspis, Tineophoctonus
Tortricidae
Pentalitomastix
Yponomeutidae
Ageniaspis, Copidosoma
COLEOPTERA
Anobiidae
Tineophoctonus
Buprestidae
Avetianella, Szelenyiola
Cerambycidae
Avetianella, Tineophoctonus
Coccinellidae
Anagyrus, Coccidoctonus, Homalotylus,
Prochiloneurus (via other Encyrtidae)
Nitidulidae
Zeteticontus
Silvanidae
Zeteticontus
Scolytidae
Avetianella, Szelenyiola
DIPTERA
Cecidomyiidae
Coccidoctonus
Ceratopogonidae
Forcipestricis
Chamaemyiidae
Cerchysius, Cheiloneurus, Desantisella
Drosophilidae
Cheiloneurus
Syrphidae
Bothriothorax, Exoristobia, Syrphophagus
Tachinidae
Exoristobia
HYMENOPTERA
Echthrodryinus
Aphelinidae
Cheiloneurus, Aphidencyrtus
Aphidiidae
Aphidencyrtus
Braconidae
Echthrodryinus
Cynipidae
? Tineophoctonus
Dryinidae
Cheiloneurus, Echthrodryinus, Helegonatopus
Encyrtidae
Cheiloneurus, Coccidoctonus, Epiencyrtus,
Gahaniella, Prochiloneurus, Zaomma
Formicidae (?)
Pheidoloxenus
238
JOHN S. NO YES
Pteromalidae
Coccidoctonus
Xylocopidae
Coelopencyrtus
Proposed new synonymies
(Junior synonyms on right)
Generk
Anagyrus Howard = Gyranusia Brethes syn. n.,
= Paranusia Brethes syn. n.,
= Philoponectroma Brethes syn. n.
Anathrix Burks = Aglyptoideus De Santis syn. n.
Apsilophrys De Santis = Copidencyrtus De Santis
syn. n.
Brethesiella Porter = Neocopidosoma Blanchard
syn. n., = Noblanchardia Ghesquiere syn. n.
Cheiloneurus Westwood = Metacheiloneurus
H offer syn. n.
Chrysoplatycerus Ashmead = Metaplatycerus
Gordh & Trjapitzin syn. n.
Coccidoctonus Craw f or d = Quay lea Timberlake
syn. n.
Ericydnus Wa\ker = Grandoriella Domenichini
syn. n.
Leptomastidea Mercel = Leptanusia De Santis
syn. n.
Metaphycus Mercet — Notoencyrtus De Santis
syn. n.
Moorella Cameron — Chrysopophilus Timberlake
syn. n.
Plagiomerus Crawford = Parahomalopoda Girault
syn. n.
Syrphophagus Ashmead = Syrphidencyrtus
Blanchard syn. n.
Specific
Prochiloneurus dactylopii
(Howard) = Achrysopophagus argentinensis De
Santis syn. n.
Proposed new combinations
(Original genus in brackets)
Anagyrus bifasciatus (Brethes) comb. n.
(Paranusia)
Anagyrus porteri (Brethes) comb. n. (Gyranusia)
Anathrix rustica (De Santis) comb. n.
(Aglyptoideus}
Apoanagyrus malenotus (De Santis) comb. n.
(Leptomastix}
Apoanagyrus montivagus (De Santis) comb. n.
(Leptomastix}
Apsilophrys capsicum (Burks) comb. n.
(Copidosomd)
Apsilophrys gracilis (De Santis) comb. n.
(Copidencyrtus)
Aztecencyrtus iceryae (Howard) comb. n.
(Cerchysius}
Brethesiella coccidophaga (Blanchard) comb. n.
(Neocopidosomd)
Cheiloneurus gahani (Dozier) comb. n.
(Achrysopophagus)
Cirrhencyrtus diversicolor (Compere) comb. n.
(Aphycus)
Coccidoctonus whittieri (Girault) comb. n.
(Cerchysius}
Copidosoma silvestrii (Costa Lima) comb. n.
(Paralitomastix}
Desobius sylvicola (De Santis) comb. n.
(Tyndarichus}
Ericydnus lamasi (Domenichini) comb. n.
(Grandoriella}
Hemencyrtus brasiliensis (Ashmead) comb. n.
(Bothriothorax}
Leptomastidea bahiensis (Compere) comb. n.
(Leptomastix}
Leptomastidea dispar (Kerrich) comb. n.
(Leptomastix)
Litomastix calypso (Crawford) comb. n.
(Holcencyrtus)
Lohiella flaviclava (Howard) comb. n. (Encyrtus}
Mercetencyrtus gracilicornis (De Santis) comb. n.
(Syrphophagus}
Metaphycus ceroplastae (Dozier) comb. n.
(Coccidoctonus)
Metaphycus discolor (De Santis) comb. n.
(Aphycus}
Metaphycus guttofasciatus (De Santis) comb. n.
(Notoencyrtus)
Metaphycus maculipes (Howard) comb. n.
(Aphycus}
Metaphycus niger (Brethes) comb. n. (Aphycus)
Metaphycus oaxacae (Howard) comb. n.
(Aphycus}
Metaphycus ogloblini (De Santis) comb. n.
(Dicarnosis}
Metaphycus portoricensis (Dozier) comb. n.
(Euaphycus)
Metaphycus rusti (Timberlake) comb. n.
(Aphycus}
Moorella compressiventris (Timberlake) comb. n.
(Chrysopophilus}
Mucrencyrtus aclerdae (De Santis) comb. n.
(Aenasioidea}
Parablastothrix chilensis (Brethes) comb. n.
(Archinus}
Paraleurocerus reticulatus (De Santis) comb. n.
(Ageniaspis)
Parechthrodryinus nitidus (Howard) comb. n.
(Encyrtus)
Plagiomerus peruviensis (Girault) comb. n.
(Parahomalopoda}
NEOTROPICAL ENCYRTIDAE 239
Rhytidothorax bollowi (Mercet) comb. n. Trichomasthus brasiliensis (Gomes) comb. n.
(Parencyrtus) (Coccidoxenus)
Rhytidothorax flaviclava (De Santis) comb. n. Trichomasthus mexicanus (Girault) comb. n.
(Parastenoterys) (Coccidoxenus)
Rhytidothorax flavicornis (De Santis) comb. n. Trichomasthus tucumanus (De Santis) comb. n.
(Parastenoterys) (Coccidoxenus)
Rhytidothorax hirtus (Howard) comb. n. Zaomma epytus (Walker) comb. n. (Encyrtus)
(Encyrtus)
Rhytidothorax perhispidus (De Santis) comb. n.
(Parastenoterys)
Syrphophagus bacchae (Blanchard) comb. n. LectotypC designations
(Syrphidencyrtus) (Present genus in brackets)
Syrphophagus flavitibiae (De Santis) comb. n.
(Syrphidencyrtus) Aseirba caudata Cameron (Aseirba)
Syrphophagus nigricornis (De Santis) comb. n. Cheiloneurus dactylopii (Howard)
(Syrphidencyrtus) (Prochiloneurus)
Tetracnemoidea peregrina (Compere) comb. n. Encyrtus epytus Walker (Zaomma)
(Tetracnemus) Hemencyrtus Herbert ii Ashmead (Hemencyrtus)
Tetracnemoidea brevicornis (Girault) comb. n. Ooencyrtus chrysopae Crawford (Ooencyrtus)
(Arhopoideus) Ooencyrtus trinidadensis Crawford (Ooencyrtus)
Acknowledgements
I wish to thank Dr M. Favreau (AMNH, New York), Dr G. Prinsloo (PPRI, Pretoria), Dr G.
Gordh (UC, Riverside), Mr P. Fidalgo (Tucuman, Argentina), and particularly Dr L. Masner
and Dr C. Yoshimoto (CNC, Ottawa) for their generous loans or gifts of material. My special
thanks to Professor L. De Santis (MLP, Argentina) and Dr E. E. Grissell (USNM, Washington)
for arranging loans of important type- and other material. Also to Dr F. D. Bennett (CIBC,
Trinidad) for all his help and generosity, to Dr Z. Boucek for his comments and to my colleagues
at the BMNH for their comments on the keys. Finally I am indebted to Miss V. Dick who typed
the final manuscript.
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242 JOHN s- NOYES
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NEOTROPICAL ENCYRTIDAE 243
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244 JOHN S. NOYES
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NEOTROPICAL ENCYRTIDAE 245
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246 JOHN s- NOYES
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177-185, 270-272.
1848. List of the specimens of Hymenopterous insects in the collection of the British Museum, Part 2,
Chalcidites, additional species, iv, 237 pp. London.
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19: 249-251.
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insects. Phil. Mag. (3) 1: 127-129.
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Index
Invalid names are in italics; principal page references are in bold.
abilis 236
abjectus 234
abnormicornis 183
abnormfs 206
abtrusus 225
acacallis 194
Acerophagoides 113, 152, 169, 237
Acerophagus 112, 154, 169, 225, 237
Achrysopophagus 185, 223
aciculatus 173
aclerdae 215, 238
Acroaspidia 114, 150, 170, 171, 237
acuminatus 170
acutigastris 226
Adelencyrtus 119, 148, 152, 170, 172, 237
advena 170
Aenasioidea 215
Aenasius 117, 126, 152, 158, 170, 237
Aeptencyrtus 113, 158, 170, 171, 237
aestivalis 209
Ageniaspis 142, 162, 165, 171, 219, 237
Aglyptoideus 174, 238
Alamella 207
albinotatus 185
albisetosa 200, 201
alboclavatus 212
albopilosus 226
Allencyrtus 181, 215, 221
Allorhopoideus 195
altensteinii 182
alula 216
Amaurilyma 136, 146, 148, 163, 171, 193, 197
ambiguus 211
americana 194
Ameromyzobia 112, 154, 172, 237
Aminellus 209
Ammonoencyrtus 115, 168, 172, 174, 236
amoenus 212
Anabrolepis 116, 122, 152, 172, 237
Anagyrella 235
Anagyrus 114, 115, 117, 121, 128, 130, 165, 173,
176, 199, 236, 237, 238
ananaitis 173
Anarhopus 112, 152, 173, 237
Anathrix 121, 124, 130, 150, 173, 174, 237, 238
Andinoencyrtus 142, 150, 174, 237
NEOTROPICAL ENCYRTIDAE
247
angelicus 225
angulatus 185
angustifrons (Cicoencyrtus) 186
angustifrons (Metaphycus) 212
angustifrons (Pseudaphycus) 225
Anicetus 115, 168, 169, 172, 174, 236
annulata (Aphycomastix) 175
annulatus (Anicetis) 174
annulipes 188
antillicola 206
antoninae 173
Anusiella 233
Anusioptera 122, 150, 175, 237
Apentelicus 213
aphelinoides (Ameromyzobia) 172
aphelinoides (Cibdeloencyrtus) 185
Aphidencyrtus 150, 167, 175, 226, 231, 236, 237
aphidiphagus 175
aphidivorus 175
Aphycomastix 112, 150, 175
Aphycomorpha 121, 126, 140, 168, 175, 237
Aphycus 187, 212
Apoanagyrus 117, 130, 165, 173, 176, 237
Aprionomastix 223
Apsilophrys 128, 163, 176, 237, 238
Apterencyrtus 235
apterus 230
Aratiscus 236
Araius 236
araucariae 175, 176
Archinus 112, 154, 176, 187, 191, 218, 219
argentinensis 223, 238
argentipes 235
argyrus 174
Arhopoidiella 112, 154, 177
Arhopoideus 233
armatus 234
Arrenoclavus 189
Arrhenophagoidea 112, 150, 178, 237
Arrhenophagus 111, 156, 178, 237
Aseirba 128, 164, 179, 197
Asitus 107, 118, 158
Asteropaeus 174
Atelaphycus 111, 150, 179, 237
aulacaspidis 170
aurantiaca 206
aureocincta 175
australiensis (Anagyrus) 173
australiensis (Tetracnemoidea) 233
australiensis (Tetracnemus) 233
australiensis (Trichomasthus) 234
Avetianella 140, 160, 179, 228, 237
Aztecencyrtus 120, 150, 180, 237
bacchae 231, 239
Baeoencyrtus 111, 160, 180
baezi 195
bahiensis 206, 238
bakeri (Litomastix) 209
bakeri (Tetracnemus) 233
bambeyi 199
Batrachencyrtus 189
bellator 173
bennetti 203
Bennettisca 118, 150, 180, 221, 236
Berecyntiscus 209
Berecyntus 209
bicarinata 223
bicolor (Bothriocraera) 182
bicolor (Encyrtus) 195
bicoloripes 219
bicristatus 193
bifasciata 189
bifasciatus 173, 238
bifasciatella 233
biformis 223
bimaculatus 187
blanchardi (Coccidencyrtus) 188
blanchardi (Dicarnosis) 117, 193
blanchardi (Hexacladia) 203
Blanchardiscus 112, 154, 181, 182
Blattidda 185
Blepyrus 126, 146, 148, 152, 182, 237
bolivari 223
bollowi 226, 239
bolowi 170, 171
bonariensis (Ammonoencyrtus) 172
bonariensis (Pelmatencyrtus) 221
bosqi 203
Bothriocraera 112, 154, 182, 237
Bothriothorax 124, 136, 166, 182, 202, 237
Boucekiella 107, 118, 163
boucheanum 189
Brachyplatycerus 111, 164, 183
brachyptera 226
brasiliensis (Aenasius) 170
brasiliensis (Aphidencyrtus) 175
brasiliensis (Desantisella) 191
brasiliensis (Hemencyrtus) 202, 238
brasiliensis (Metaphycus) 212
brasiliensis (Paramucrona) 219, 220
brasiliensis (Parencyrtus) 221
brasiliensis (Trichomasthus) 234, 239
brethesi (Aenasius) 170
brethesi (Leptomastidea) 206
Brethesia 183
Brethesiella 126, 160, 183, 237, 238
brevicauda 203
brevicornis (Amaurilyma) 171
brevicornis (Tetracnemoidea) 233, 239
brevistigma 173
brittanica 235
brivicollis 233
bruchi 171
bucculatricis 209
bucculentus 183
bulyginskayae 172
248
JOHN S. NOYES
caeruleus 170
californica (Gahaniella) 197
californicus (Apoanagyrus) 176
callidii 189
Calliencyrtus 121, 150, 183
Calluniphilus 226
Calocerinus 108
Calometopia 219
calypso 209, 238
capnodiobia 179
capsicum 176, 238
Carabunia 124, 162, 184, 236
carinata 177
cariocus 170
casali 202
caudata 179, 239
celia 199, 200
cerapterocera 189
Cerapterocerus 107, 115, 167
Cerchysius 128, 150, 179, 180, 184, 197, 223, 237
ceroplastae 136, 212, 238
ceroplastodis 174
ceylonensis 174
Chalcaspis 126, 152, 184, 237
chalconotus 208
chapadae 170
Cheiloneuroides 193
Cheiloneurus 114, 116, 120, 164, 166, 168, 169,
170, 184, 206, 236, 237, 238
Chestomorpha 223
chilensis (Anagyrus) 173
chilensis (Parablastothrix) 219, 238
Chiloneurinus 235
chionaspidis (Adelencyrtus) 170
chionaspidis (Arrhenophagus) 178, 179
chlorinus 223
chrysomphali 216
chrysopae 199, 217, 239
Chrysoplatycerus 115, 158, 170, 185, 237, 238
Chrysopophagus 184
Chrysopophilus 213, 238
Cibdeloencyrtus 144, 150, 185
Cicoencyrtus 146, 160, 186, 191, 218
Cirrhencyrtus 120, 140, 158, 160, 180, 187, 237
citriculus 185
citrina (Gyranusoidea) 199
citrinus (Acerophagus) 169
clavatus 235
clavicornis 182
claviger 209
clisiocampae 217
Coccidaphycus 113, 154, 187, 236
Coccidencyrtoides 188
Coccidencyrtus 117, 119, 132, 148, 168, 188, 218,
237
coccidivorus 173
coccidophaga 183, 238
Coccidoctonus 136, 142, 165, 188, 212, 236, 237,
238
Coccidoxenus 234
coccois 169
Coccophoctonus 182
cockerelli 203
Coelaspidia 113, 158, 188, 237
Coelopencyrtus 119, 148, 158, 189, 238
colocense 233
colombiensis 185
coloripes 178
Comperencyrtus 233
Comperia 117, 156, 189,236
Compendia 115, 168, 189, 237
compressicornis 184
compressiventris 213, 238
Corny s 195
concupiens 200
conformis 134, 195, 196
confusor 217, 218
confusus 202
connectens 170
consobrinus 213
convexus (Desobius) 192
convexus (Parechthrodryinus) 221
Copidencyrtus 176, 238
Copidosoma 128, 132, 138, 148, 163, 176, 189,
209, 237
corvina 235
creona 217
cristata 203
Cristatothorax 185
cristatus (Cheiloneurus) 185
cristulata 204
crouzelae 216
cultrata 179
cupreicollis 185
cyanea (Plagiomerus) 222
cyaneus (Trichomasthus) 234
Cyderius 136, 150, 175, 190
dactylopii (Blepyrus) 182
dactylopii (Leptomastix) 207
dactylopii (Prochiloneurus) 223, 224, 238, 239
dalmanii 199
debilis 169
Deloencyrtus 121, 128, 150, 191, 216
denieri 188
depressus 170
Desantisella 124, 156, 191, 236, 237
desantisellus 226
desantisi 190
Desobius 119, 144, 156, 171, 192, 197
destructor 193
diaspidis 222
Dicarnosis 150, 193, 212
dimorpha 200
discolor 212, 238
dispar 206, 238
diversicolor 187, 238
diversicornis (Anagyrus) 173
NEOTROPICAL ENCYRTIDAE
249
diversicornis (Apoanagyrus) 176
diversicornis (Tetracnemus) 233
Diversinervus 116, 162, 165, 169, 193, 236
divisus 231
dysmicocci 225
Echthrobaccha 231
Echthrodryinus 140, 150, 167, 169, 178, 193, 199,
217, 236, 237
Echthroplexiella 111, 156, 185, 194
Ectroma 114, 150, 194
Ectromatopsis 114, 165, 194
ehrhorni 187
elegans (Cheiloneurus) 184, 185
elegans (Diversineurus) 193
elegans (Microterys) 213
Encyrtolophus 185
Encyrtus 120, 162, 163, 164, 195
ensifer 188
Epaenasomyia 202
Epidinocarsis 173
Epiencyrtoides 170
Epiencyrtus 124, 126, 146, 150, 195, 237
epytus 235, 239
Ericydnus 122, 162, 195, 237, 238
eriococci (Atelaphycus) 179
eriococci (Zaomma) 235
eruptor 212
Euaphycus 212
Eucomys 195
europaea (Timberlakia) 233
europaeus (Rhopus) 226
Euryrhopalus 126, 148, 154, 196, 237
Exoristobia 119, 140, 150, 195, 196, 237
extranea 172
fasciatipennis 223
felix 174
ferrisi 185
ferrisianae 225
flaminius 203
flandersi 170
flava (Mariola) 210, 211
flaviclava (Lohiella) 209, 210, 238
flaviclava (Rhytidothorax) 226, 239
flavicollis 185
flavicornis 226, 227, 239
flavidulus 225
flavifrons 204, 205
flavigena 180, 181
flavipes (Bothriocraera) 182
flavipes (Meromyzobia) 212
flavipes (Neapsilophrys) 215, 216
flaviptera 229
flavitibiae 231, 239
flavus (Acerophagus) 169
flavus (Aztecencyrtus) 180
flavus (Metaphycus) 212
flavus (Microterys) 213
floridana 209
Forcipestricis 144, 146, 156, 164, 166, 196, 233,
237
formosus 200
frontalis 170
frontatus 213
fullawayi 235
fulvescens 194
fulviceps 213
fumipennis 212
fuscicollis 171
fuscipalpis 184, 223
gahani (Cheiloneurus) 185, 238
gahani (Lochitoencyrtus) 209
Gahaniella 117, 118, 132, 138, 164, 166, 197, 236,
237
gargaris 189
gaseaui 196
Gibber ella 219
glabriscutellum 236
glauca 199
gloriosa (Copidosoma) 189
gloriosa (Gonzalezia) 197
Gonzalezia 134, 164, 197
gracilicornis 211, 238
gracilis 176, 238
graminicolens 173
Grandoriella 196, 238
greeni 173
grioti 188
gripha 212
griseus 225
Grissellia 138, 148, 168, 197
guttofasciatus 212, 238
Gymnoneura 199
Gyranusa 173
Gyranusia 173, 238
Gyranusoidea 117, 165, 173, 199, 237
Habrolepis 115, 152, 199, 236, 237
Habrolepistia 189
Habrolepoidea 144, 150, 195, 199
Habrolepopterygis 174
Habrolepopteryx 108
Hadrencyrtus 230
Hambletonia 115, 154, 170, 200, 237
haywardi 195
Hazmburkia 200
hederaceus 212
Helegonatopus 130, 134, 154, 167, 200, 208, 236,
237
Helmecephala 119, 136, 166, 200
helvolus 212
Hemaenasius 116, 117, 128, 164, 183, 202
Hemaenasoidea 203
Hemencyrtus 121, 122, 124, 128, 130, 134, 136,
165, 166, 179, 202, 221
herbertii 202, 239
250
JOHN S. NOYES
Heterarthrellus 173
Hexacladia 121, 152, 203, 223, 236
heydeni 233
hirtus 226, 227, 239
hispanicus 209
histrio 206
Holcencyrtus 209
Holcothorax 171
Homalopoda 111, 152, 203, 237
Homalotylus 120, 121, 126, 158, 162, 203, 237
Homosemion 115, 150, 172, 203, 237
hookeri 204
Hoplopsis 124, 150, 204
Howardia 195
Howardiella 195
Hungariella 233
Hunterellus 117, 119, 162, 204, 227, 236
hyalinistigma 190
hyettus 170
iceryae (Aztecencyrtus) 180, 238
iceryae (Isodromus) 206
Iceromyia 116, 118, 165, 168, 204
indica 203
infelix 195
infida 195
inimicus 185
injuriosa 231
inquisitor 235
insolitus (Anagyrus) 173
insolitus (Prochiloneurus) 233
insulanus 213, 214
insularis (Aenasius) 170
insularis (Blepyrus) 182
insularis (Zeteticontus) 236
io223
Isodromus 126, 142, 158, 206, 236
johnsoni 217
josephi 194
jucundus 173
kaalae 189
Kaszabicyrtus 226
kirkpatricki 196
koebelei 203
koehleri 189, 190
komabae 189
kotynskyi 213
Krishneriella 174
kuscheli 191
kuwanae 217
laevigatus 236
lamasi 196, 238
lambinus 235
latifrons 183
latipes 203
latiscapus 217
lecaniorum 195
Lepidaphycus 203
Leptanusia 206, 238
Leptomastidea 121, 165, 206, 237, 238
Leptomastix 122, 130, 165, 173, 176, 206, 237
Limastotix 209
Liocarus 223
Lirencyrtus 130, 167, 207
Litomastiellus 209
Litomastix 138, 142, 160, 190, 208, 237
littoralis 195
Lochitoencyrtus 136, 156, 209
Lohiella 132, 150, 209
longipes 183
longiscapus 170
longisetaceus 185
lopesi 189
lopezi 176
loretanus 185
lounsburyi 212
lucida 184
lusitanicum 226
luteolus (Acerophagus) 169
luteolus (Metaphycus) 212
maculicornis 119, 188
maculipennis (Meromyzobia) 191, 212
maculipennis (Tetracnemus) 233
maculipes 212, 238
malenotus 176, 238
malloi 132, 188
maplei 170
Mariola 118, 150, 210
marlatti 226
Masencyrtus 200
Masia 233
masii 170
masneri 227, 228
matritensis 173
mayri 204
Melanaphycus 212
Mendozaniella 203
Mercetencyrtus 134, 138, 150, 168, 211
merceti 189
Mercetia 226
Mercetiella 212
Meromyzobia 112, 152, 191, 212, 236
mesograptae 231
Metacheiloneurus 185, 238
metallicus 221
Metallon 194
Metallonoidea 235
Metaphycus 108, 117, 118, 126, 136, 140, 144, 160,
162, 172, 179, 180, 193, 194, 212, 230, 235,
236, 237, 238
Metaplatycerus 185, 238
Metapterencyrtus 235
mexicana (Hexacladia) 203
NEOTROPICAL ENCYRTIDAE
251
mexicanus (Blepyrus) 182
mexicanus (Encyrtus) 195
mexicanus (Homalotylus) 203
mexicanus (Metaphycus) 212
mexicanus (Trichomasthus) 234, 239
mexicanus (Tyndarichoides) 234
microphagus 235
Microterys 117, 124, 162, 164, 165, 167, 213, 236,
237
minutus (Brachyplatycerus) 183
mirabilis (Encyrtus) 195
mirabilis (Homalotylus) 203
Mirastymachus 226
Mirocerus 226
Mirrencyrtus 236
moderatus 170
moestus 185
montana 193
monticolens 212
montivagus 176, 238
Moorella 119, 150, 213, 236, 238
morio 223
Mucrencyrtus 114, 122, 130, 134, 136, 163, 181,
213, 221, 236
mundus 225
myersi 184
Mymariella 178
myrmicoides 170
nacoliae 222
Neapsilophrys 128, 150, 215, 237
nearctica 231
Neoaenasioidea 203
Neoanisotylus 187
Neococcidencyrtus 120, 122, 148, 168, 216, 218,
237
Neocopidosoma Blanchard 183
Neocopidosoma Ishii 189
Neodusmetia 114, 158, 216, 237, 238
Neoprochiloneurus 223
Nesencyrtus 189
niger 212, 238
nigriaxillae 206
nigrescens 185
nigricans (Coccidaphycus) 187
nigricans (Tachardiobius) 231
nigriceps (Anagyrus) 173
nigriceps (Protyndarichoides) 224
nigriclavus 226
nigricornis 231, 239
nigroclavatus 226
nikolskajae 200
nitidus 221, 238
Noblanchardia 183, 238
Nobrimus 203
notativentris 169
Notoencyrtus 212, 238
nubeculus 231
nubilipennis (Acerophagus) 169
nubilipennis (Habrolepis) 199
oaxacae 212, 238
obesus 142, 188
occupatus 176
ocellatus 174
oculata 203
odonaspidis 170
odyneri 189
oeceticola 176
ogloblini 193, 212, 238
Ooencyrtoides 226
Ooencyrtus 114, 119, 132, 138, 148, 150, 163, 166,
167, 178, 194, 199, 217, 226, 236, 237
orbitalis 206
osborni 188
oviductus 223
pachypsyllae 226
pacificus 170
pallidus 169
paludatus 195
Paludencyrtus 200
Papaka 146, 148, 160, 217
Parablastothrix 119, 152, 218, 237
Paracalocerinus 233
Parachrysopophagus 223
Paracopidosomopsis 209
Parahomalopoda 222, 238
Paraleurocerus 142, 156, 171, 219, 237
Paralitomastix 148, 163, 190, 219, 237
Paramucrona 134, 150, 182, 219, 236
Paranusia 173, 238
Paraplatycerus 185
Parastenoterys 226
Parataneostigma 206
Parechthrodryinus 122, 124, 130, 136, 146, 166,
167, 221, 225, 237
Parencyrtus 111, 150, 221, 226
parlatoriae 178
parvula 226
paulistus 170
Pauridia 144, 158, 169, 221, 237
peckhami 195
pectinatum 173
Pelmatencyrtus 113, 150, 170, 221, 237
Pentacnemus 209
Pentalitomastix 113, 160, 190, 222, 237
perdignus 225
peregrina (Pauridia) 221
peregrina (Tetracnemoidea) 233, 239
pergandei 184
perhispidus 226, 227, 239
Perissodromus 183
personatus 170
peruviensis 222, 238
Pezobius 194
Pheidoloxeniscus 222
252
JOHN S. NOYES
Pheidoloxenus Ashmead 113, 150, 222, 237
Pheidoloxenus Girault 222
phenacocci 170
philippinensis 196
Philoponectroma 173, 238
Pholidoceras 226
Pholidocerodes 226
pia 189
piceae 233
pinicola 193
piso 226
Placoceras 233
Plagiomerus 111, 152, 222, 237, 238
planiscutellum 235
Platencyrtus 107, 118, 163, 211
platensis 230
Platylyca 128, 150, 222
platys 180
plaumanni 191
plethorica 222
polychromus 194
porteri 173, 238
portoricensis (Forcipestricis) 196
portoricensis (Metaphycus) 212, 238
portoricensis (Trichomasthus) 234
praenitens 185
prenidis 217
pretiosus (Euryrhopalus) 196
pretiosus (Tetracnemoidea) 233
prima (Pseudhomalopoda) 225
primus (Anicetus) 174
primus (Lirencyrtus) 207, 208
Prionomastix 121, 124, 162, 223, 236
Prionomitus 111, 150, 184, 223, 236
Prochiloneurus 116, 120, 121, 168, 223, 236, 237
propinquus 196
Propsyllaephagus 226
Prorhopoideus 195
prosopidis 225
Protanagyrus 173
Protyndarichoides 136, 150, 224
Protyndarichus 221
Pseudanusia 189
Pseudaphycus 112, 154, 169, 225, 237
Pseudhomalopoda 111, 152, 225, 237
Pseudleptomastix 173, 206
Pseudlitomastix 217
pseud ococci (Anagyrus) 173
pseudococci (Leptomastidea) 206
pseudococcina 200
Pseudolitomastix 222
pseudophanes 200
Psilomirinus 225
Psyllaephagus 132, 138, 150, 163, 175, 226, 236
pulchellus 223
pulcher 185
pulchra 185
pulchricornis (Anagyrus) 173
pulchricornis (Zaomma) 235
pulchrior 196
pulvinariae 185
punctatus 170
putophilus 235
pyelae 226
quadraticeps 222
quadricolor 195
quadridentatus 170
quadrimaculati 231
Quaylea 188, 238
quintanai 174
regularis 170, 171
reticulatus (Metaphycus) 212
reticulatus (Paraleurocerus) 219, 238
rex 223
Rhopoideus 169
Rhopus 114, 118, 140, 164, 226, 237
Rhytidothorax 119, 122, 124, 132, 136, 140, 144,
146, 158, 164, 195, 210, 221, 225, 226
Richardsius 235
Rileya 185
Rotrencyrtus 170
rotundiformis 223, 226
rouxi 199
rusti 212, 238
rustica 174, 238
saccharalis 193
saccharicola 173
saissetiae 197
sangwani 216
Scelioencyrtus 226
sceptriger 213
Sceptrophoms 213
Schedioides 200
Schedius 217
schwarzi 196
scutellaris 182
scutellata (Paraleurocerus) 219
scutellatus (Zeteticontus) 236
seini 223
semifacta 194
sheldoni 235
Shenahetia 118, 142, 168, 227, 232
silvai 231
silvestrii (Copidosoma) 190, 238
silvestrii (Tachardiobius) 231
similis 170, 171
Simmondsiella 132, 150, 227, 229
smithii 203
Solenaphycus 111, 154, 230
Solenoencyrtus 113, 144, 150, 230
Sophencyrtus 203
sordidus 196
splendens 185
squammulata 206, 207
stanleyi 212
NEOTROPICAL ENCYRTIDAE
253
Stemmatosteres 111, 154, 230, 237
Stenoteropsis 234
Stenoterys 206
submetalica 194
submetallicus 217
superbus 193
swederi 195
sydneyensis 173
sylvicola 193, 238
sylvius 213
Synaspidia 196
Syrphidencyrtus 231, 238
Syrphophagus 134, 140, 150, 164, 167, 168, 175,
194,211,230,237, 238
Szelenyiola 140, 160, 228, 231, 237
Tachardiobius 138, 160, 231, 237
tachigaliae 170
Tachinaephagus 107, 124, 136, 156
Tanaomastix 206
Tanyencyrtus 118, 142, 167, 229, 231, 237
tanystis 173
teciae 219
terebrata (Grissellia) 197, 198
terebratus (Anagyrus) 173
terminalis 203
tertia 197
Tetarticlava 113, 150, 232
Tetracladia 233
Tetracnemella 234
Tetracnemoidea 113, 152, 174, 178, 233, 237
Tetracnemopsis 233
Tetracnemus 114, 152, 233, 237
Tetralophidea 233
Tetralophiellus 233
texana (Tetracnemus) 233
texanus (Acerophagus) 169
thebe 209
thomsoniscae 235
thyreodontis 195
Thysanomastix 203
tiliaris 223
Timberlakia 112, 154, 169, 225, 233, 237
Tineophoctonus 121, 150, 234, 237
townsendi 203
Trechnites 188
trellesi 175, 226
triangularis 169
Trichomasthus 140, 164, 167, 229. 230, 234, 236,
237
tricolor 173
tricoloricornis 213
trifasciata 191
Trimorphocerus 182
trinidadensis (Apoanagyrus) 116
trinidadensis (Coccidoctonus) 188
trinidadensis (Ooencyrtus) 217, 239
truncatella 209
tucumanus 234, 239
Tyndarichoides Girault 120, 126, 150, 234
Tyndarichoides Mercet 221
Tyndarichus 193
urbicola 190
urocerus 184
utilis 225
vadosus 170
varicornis (Hemaenasius) 202
varicornis (Paralitomastix) 219
venatorius 217
vendicus 226
Verdunia 189
vespertina 218
vexans 170
vianai (Encyrtus) 195
vianai (Solenaphycus) 230
virescens 190
waterstoni 184
Waterstonia 187
westwoodi 233
wheeleri Ashmead 222
wheeleri Girault 222
whittieri 188, 238
Xanthoencyrtus 226
Xenocomys 216
Xiphomastix 173
yoshimotoi 232
Zaomma 117, 130, 144, 169, 235, 237
Zaplatycerus 115, 156, 170, 235, 237
Zarhopaloides 200
Zarhopalus 115, 158, 170, 235, 237
zebratus 212
Zeteticontus 108, 119, 134, 142, 158, 163, 166,202,
236, 237
zetterstedti 172
Other British Museum (Natural History)
Publications on the Hymenoptera :
African bees of the genera Ceratina, Halictus and Megachile. T. D. A. Cockerell.
1937, xvi + 254 pp, 50 text figures, 8vo, £7-15
A revision of the Leucospidae (Hymenoptera : Chalcidoidea) of the World.
Z. Boucek.
Bull. British Museum (Natural History) Entom. Supp. No 23, 1974, 241 pp, 272 text figures,
4to paper, £14-00
The Pteromatidae of north-western Europe (Hymenoptera : Chalcidoidea).
M. W. R. de V. Graham.
Bull. British Museum (Natural History) Entom. Supp. No 16, 1969, 908 pp, 686 text figures,
4to paper, £33-00
A revisional study of the Masarid wasps (Hymenoptera : Vespoidea).
O. W. Richards.
1962, 294 pp, 241 text figures, 4to, £9-35
The social wasps of the Americas excluding the Vespinae. O. W. Richards.
1978, 571 pp, 159 text figures, 4 plates (colour), 4to cloth, £32-50
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Titles to be published in Volume 41
A revision of the western Palaearctic species of the genus Meteorus
(Hymenoptera: Braconidae). By T. Huddleston.
A revision of the Old World Polymorphanisini (Trichoptera: Hydropsychidae).
By P. C. Barnard.
A review of the genera of Neotropical Encyrtidae (Hymenoptera:
Chalcidoidea). By John S. Noyes.
A revision of the British species of Dendrocerus Ratzeburg (Hymenoptera
Ceraphronoidea) with a review of their biology as aphid hyperparasites.
By N. D. M. Fergusson.
A revision of the Pacific species of Conocephalus Thunberg (Orthoptera :
Tettigoniidae). By Linda M. Pitkin.
Printed by The Whitefriars Press Ltd, London and Tonbridge
Bulletin of the
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A revision of the British species of
Dendrocerus Ratzeburg (Hymenoptera :
Ceraphronoidea) with a review of their
biology as aphid hyperparasites
N. D. M. Fergusson
Entomology series
Vol 41 No 4 18 December 1980
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World List abbreviation : Bull. Br. Mus. nat. Hist. (Ent.)
Trustees of the British Museum (Natural History), 1980
ISSN 0524-6431 Entomology series
Vol 41 No 4 pp 255-314
British Museum (Natural History)
Cromwell Road
London SW7 5BD Issued 18 December 1980
A revision of the British species of Dendrocerus
Ratzeburg (Hymenoptera : Ceraphronoidea)
with a review of their biology as aphid
hyperparasites
N. D. M. Fergusson ^
Department of Entomology, British Museum (Natural History), Cromwell Road, London SW7
5BD
Contents
Synopsis ............ 255
Introduction ............ 255
Terminology and methods .......... 256
Abbreviations ............ 257
Depositories ............ 257
Suprageneric classification 257
Key to families and subfamilies of the Ceraphronoidea . . . . -257
Generic classification in the Megaspilidae . . . . . . -258
Key to the British genera of Megaspilidae ....... 258
Dendrocerus Ratzeburg . . ....... 259
Subgenera
Synonymic list of British species
Key to British species
Species descriptions
Review of the biology of Dendrocerus
261
261
263
266
293
Host aphid list for Dendrocerus species ....... 293
List of the Aphidiidae (Hymenoptera) parasitized by Dendrocerus . . . 295
Other hosts 297
Host information on some non-British species of Dendrocerus. . . . 300
Non-British species . . . . . . . . . . -301
Doubtfully placed species .......... 309
Acknowledgements ........... 309
References ............ 309
Index 313
Synopsis
The species of Dendrocerus occurring in Britain are revised; 72 nominal species are involved, of which 14 are
considered to be valid. Six species are recorded as British for the first time and 1 7 new specific synonyms are
established. All the European and some non-European species are discussed. Keys are given to the families
and subfamilies of Ceraphronoidea, to the genera of Megaspilidae and to the British species of Dendrocerus.
The genus Dendrocerus is redefined and its subgenera discussed, and one new generic synonym established.
The biology of each species is discussed and the biology of the genus as a whole is reviewed.
Introduction
Dendrocerus Ratzeburg is the most economically important genus of the Ceraphronoidea as it
contains most, if not all, of the aphid hyperparasites known from this superfamily. Species of
Dendrocerus are also associated with other Homoptera, and with Neuroptera and Diptera. The
Bull. Br. Mus. not. Hist. (Ent.) 4 1 (4): 255-314 Issued 1 8 December 1 980
256 N. D. M. FERGUSSON
genus has a world wide distribution except that it has not yet been recorded from the Madagascan
region.
The original idea for this study came from Dr V. F. Eastop. As a result of discussions with him
and other aphid-workers, it has been possible to obtain many new host records and to present the
data in a way which should be useful to them as well as to hymenopterists. The principal aims of
this study are to distinguish Dendrocerus from other genera of Ceraphronoidea, especially
Conostigmus, to redefine the limits of the species in Dendrocerus, to establish the extent of the
British fauna, and to assess the basic host-parasite relationships. In addition, the subgenera have
been critically reviewed.
In the course of this work nearly 3000 specimens have been examined, and most of the European
species were found to occur in Britain. Examination of much foreign material, especially a large
Dutch collection given to the British Museum (Natural History) by Dr Evenhuis, has helped to
clarify the taxonomy and add a number of host records.
Terminology and methods
The morphological nomenclature used in this work follows that of Richards (1977) and Snodgrass
(1935). The terms thorax and gaster have been used in preference to the 'mesosoma' and
'metasoma' used by Dessart (1972a). The microsculpture terminology follows Eady (1968). The
term frontal dent refers to a small median depression on the frons, closer to the interantennal carina
than to the median ocellus. This is the 'frontal pit' of Dessart (1972a) but not the same as the
'frontal pit' used by Bin (1977). The preoccipital crescent (Figs 25, 26) is a central area behind the
lateral ocelli which is almost semicircular in shape. It is not as narrow or as well defined as the
analogous postocellar area of some sawflies (Hymenoptera: Symphyta). The gastral collar (Figs
18-20) is a large projection around the petiole arising from the main tergite and sternite of the
gaster. The term notaulices has been used for the pair of long furrows, one on each side of the
median mesonotal furrow (Figs 13—17), which some authors have incorrectly called parapsidal
furrows. Two other pairs of furrows (Fig. 14) are sometimes present as short lines; one pair at the
anterior margin of the mesonotum close to the median furrow, the other pair half way down the
mesonotum exterior to the notaulices. As the terminology of these furrows is confused I have
followed Dessart (1972a) in calling them collectively the secondary furrows.
A standardized arrangement of information has been adopted for each species. The synonymy
contains all nomenclatural changes but it was thought unnecessary to include every reference to
the species. The species description is not based solely on type-material as it includes a considera-
tion of the infraspecific variation. The male has been characterized separately, but only features
differing from the female have been mentioned. The biology section contains all published host
records and as many new records as it has been possible to obtain. The information has been
presented as follows:
x<$, y9, host/primary parasite/host plant, date of collection or emergence (authority for record
if previously published).
A dash has been inserted in the appropriate place if the host, parasite or host plant is unknown. The
records are listed in alphabetical order. The discussion of the biology is based on the classification of
faunistic complexes employed by Stary (1970: 312).
All measurements used in the text are maximum distances unless otherwise stated. In males of
D. halidayi and D. ramicornis the antennal process is excluded from the measurement of segment
breadth. The lateral length of the head is measured from the vertex to the base of the mandible. The
length of the scutellum is measured from the mesonotal border to the apex of the scutellum, and the
breadth is taken as the maximum distance across the raised area. The length and breadth of the
pterostigma are measured respectively parallel and perpendicular to the front edge of the wing.
(The wing edge must not be curled over, making the pterostigma appear long and narrow.)
In this paper the classification and nomenclature of aphids has been taken from Eastop & Hille Ris
Lambers (1976) and Kloet & Hincks (1964: 67); and that of the Aphidiidae from Kloet & Hincks
(1978:61),Mackauer(1968),Stary(1973)andTakada(1973:l).
BRITISH SPECIES OF DENDROCERUS 257
Abbreviations
AI, All, AIII, AIV, etc. - Antennal segments one, two, three, four, etc. The scape is AI, the pedicel
All and the first flagellar segment is AIII.
L/B - Length divided by the breadth of a structure, that is the ratio of the length to the breadth
with the breadth expressed as one.
I.C.Z.N. - International Code of Zoological Nomenclature.
Depositories
The following abbreviations have been used for depositories containing type- or other material
which has been examined.
MP, Amiens Musee de Picardie, Amiens, France
MNHU, Berlin Museum fur Naturkunde der Humboldt-Universitat, Berlin, East Germany
BMNH British Museum (Natural History), London, England
IRSNB, Brussels Institut Royal des Sciences Naturelles de Belgique, Brussels, Belgium
TM, Budapest Termeszettudomanyi Muzeum, Budapest, Hungary
Chambers coll. Collection of Dr V. Chambers, Meppershall, Bedfordshire, England
NMI, Dublin National Museum of Ireland, Dublin, Ireland
IP, Eberswalde Institut fur Pflanzenschutzforschung, Eberswalde, East Germany
MZU, Florence Museo Zoologico della specola Universita degli studi, Florence, Italy
MHN, Geneva Museum d'Histoire Naturelle, Geneva, Switzerland
MCSN, Genoa Museo Civico di Storia Naturale, Genoa, Italy
RES, Harpenden Rothamsted Experimental Station, Harpenden, England
ZMU, Helsinki Zoological Museum of the University, Helsinki, Finland
UM, Kuala Lumpur University of Malaya, Kuala Lumpur, Malaya
MCM, Liverpool Merseyside County Museum, Liverpool, England
ZI, Lund Zoologiska Institution, Lund, Sweden
NHM, Maastricht Natuurhistorisch Museum, Maastricht, Netherlands
MM, Manchester Manchester Museum, Manchester, England
ZMMLSU, Moscow Zoological Museum of the Moscow Lomonosov State University, U.S.S.R.
UM, Oxford University Museum, Oxford, England
IP, Paris Institut Pasteur, Paris, France
MNHN, Paris Museum National d'Histoire Naturelle, Paris, France
NR, Stockholm Naturhistoriska Riksmuseet, Stockholm, Sweden
MCSN, Verona Museo Civico di Storia Naturale, Verona, Italy
NM, Victoria National Museum of Victoria, Melbourne, Victoria, Australia
IPO, Wageningen Instituut voor Plantenziektenkundig Onderzoek, Wageningen, Netherlands
USNM, Washington National Museum of Natural History, Washington D.C., U.S.A.
The specimens listed under 'Material examined' without a named depository are in the BMNH.
Suprageneric classification
The Ceraphronoidea consist of two families: Ceraphronidae and Megaspilidae. The Megaspilidae
are further divided into the subfamilies Megaspilinae and Lagynodinae. The following key has been
designed for ease of use; for this reason characters best seen in slide preparations (e.g. genitalia
characters) have not been used.
Key to the families and subfamilies of the Ceraphronoidea
1 Midleg with one tibial spur; mesonotum without notaulices (except Ceraphron abnormis Perkins,
which has mesepisternum strongly striate), with or without median longitudinal furrow; longer
tibial spur of foreleg not forked apically; antenna eight- to eleven-segmented
CERAPHRONIDAE
Midleg with two tibial spurs; mesonotum, at least anteriorly, with notaulices and nearly always
with median longitudinal furrow, or if notaulices absent then pronotum strongly elongate and
ocelli absent or forewing has only costal vein and gastral collar has three carinae;
mesepisternum with or without striations; longer tibial spur of foreleg forked apically (Fig. 1 1);
antenna eleven-segmented (MEGASPILIDAE) 2
258 N. D. M. FERGUSSON
2 Ocelli usually absent; pronotum markedly elongated, mesonotum reduced (Fig. 12), gastral collar
with three distinct dorsal carinae (Fig. 12). Apterous, females . . LAGYNODINAE (part)
Ocelli present; pronotum and mesonotum normal, or if (rarely) thorax is modified then gastral
collar has many carinae. Mostly macropterous, males and females .... 3
3 Forewing with large, obvious pterostigma (Figs 1, 33—36, 38—46); in brachypterous forms large
pterostigma is conspicuous near wing apex (Fig. 9) (except in Conostigmus dimidiatus
(Thomson), in which median longitudinal furrow of mesonotum is absent posteriorly); gastral
collar not with three carinae, usually with many ..... MEGASPILINAE
Forewing without pterostigma, or pterostigma linear (Fig. 10) and longitudinal furrow complete;
gastral collar with three distinct carinae dorsally (Fig. 12) . . . LAGYNODINAE (part)
Generic classification in the Megaspilidae
The genera of the Megaspilidae are difficult to distinguish and Conostigmus and Dendrocerus have
never been adequately delimited. The following key includes an attempt to produce a genuine
disjunction for the two genera. The couplet separating females of Conostigmus and Dendrocerus is
long and complex but nevertheless will not adequately distinguish D. spissicornis and C.
fasciatipennis, which have therefore been keyed separately. Conostigmus fasciatipennis has many
characters typical of Dendrocerus species; conversely, D.punctipes has several characters normally
found in Conostigmus species.
Key to the British genera of Megaspilidae
1 Disc of forewing sparsely covered with very short pubescence; radial vein short, 0-5-0-8 times as
long as breadth of pterostigma; propodeal spiracle large and conspicuous; hypopygium of
female with distinct tuft of upright hairs; interantennal carina incomplete; wings hyaline;
macropterous TRICHOSTERESIS Forster
(Monotypic: T. glabra (Boheman), a parasite of Diptera: Syrphidae)
- Forewing with normal pubescence (except Dendrocerus punctipes, which has inter-antennal
carina complete and dark area on forewing); radial vein long, at least as long as breadth of
pterostigma; propodeal spiracle not large or conspicuous; hypopygium without distinct tuft of
upright hairs; macropterous or brachypterous ........ 2
2 Propodeum always with distinctive median double-toothed lamellate projection (Figs 2, 3).
Precoxal sulcus strongly developed, foveolate and curved (Fig. 4). Disc of scutellum always
conspicuously bordered laterally and apically by strong foveae (Fig. 2). Head sculpture and
ocular suture strongly foveolate. Antenna long and broad, AIII at least 4 times longer than
broad, AIII of females about twice (1-7-2- 1) as long as AIV, AIV of males as long as scape.
Interantennal carina without sharp median projection. Macropterous ....
(A rarely collected genus) MEGASPILUS Westwood
Propodeum without median double-toothed lamellate projection, often without any projection, or
with median fovea, keel, small spine or double tooth. Precoxal sulcus absent, or if present then
weakly developed and virtually straight. Disc of scutellum only rarely bordered laterally and
apically by strong foveae. Head sculpture usually alutaceous, ocular suture normally without
strong foveae. If antenna! segments are long they are usually thin, only occasionally is AIII 4
times longer than broad, AIII of females normally less than twice as long as AIV, AIV of males
usually shorter than scape. Interantennal carina sometimes with small sharp median projection
(Figs 6, 12). Macropterous or brachypterous ......
3 Entire breadth of gaster covered anteriorly by strigose sculpture radiating from gastral collar and
nearly reaching gastrocoeli (Fig. 19); head rugose; ocellar triangle with broad base; gastral
collar broad and very short . . . DENDROCERUS (part, D. spissicornis) (p. 292)
Area of sculpture at anterior of gaster shaped and sculptured differently; if head rugose then
ocellar triangle has short base or gastral collar is narrower or longer . . 4
4 Gastral collar very short and broad (Fig. 18); hairs on costal vein long, up to 0- 1 mm (Fig. 39)
(these hairs can be lost); ocellar triangle short with a broad base; notaulices complete and
sharply angled outwards anteriorly; basal flagellar segments of male virtually cylindrical; large
brown area under pterostigma in female, present but less distinct in male; frontal dent absent;
disc of forewing with normal pubescence; preoccipital crescent not reaching eyes or ocelli;
interantennal carina complete . . CONOSTIGMUS Dahlbom (part, C.fasdatipennis Kieffer)
BRITISH SPECIES OF DENDROCERUS 259
Gastral collar long (Fig. 20), or if short then hairs on costal vein much shorter, or ocellar triangle
long with short base, or notaulices posteriorly absent (Fig. 15) or slightly smoothly angled, or
basal flagellar segments of male clearly asymmetrical (Figs 30-32, 47-53), or forewing
hyaline, or frontal dent present (Fig. 8), or disc of forewing with reduced pubescence, or
preoccipital crescent reaching eyes or ocelli, or interantennal carina incomplete or absent 5
5 Males (All globular, as in Figs 51-55) ........ 6
Females (All elongate, as in Figs 56-63) ....... 7
6 Basal flagellar segments clearly asymmetrical (Figs 30, 47, 49), often strongly serrate (Figs 48,
50-53) or with long projections (Figs 31,32). If only slightly asymmetrical then mesepisternum
smooth and pterostigma at least twice as long as broad (Fig. 38), or parameres exposed, large
and distinctively angled (Fig. 29). Ocellar triangle always with broad base (e.g. Fig. 7); eye
pubescence short; notaulices sharply angled towards anterior corners of mesonotum (Figs
14, 17) or absent posteriorly (Fig. 15) .... DENDROCERUS (part) (p. 259)
All flagellar segments virtually cylindrical; ocellar triangle usually with short base (Figs 5, 6); eye
pubescence often long and conspicuous; notaulices complete and usually only slightly smoothly
angled towards anterior corners of mesonotum . . . CONOSTIGMUS Dahlbom (part)
7 Ocellar triangle short with broad base (except D. punctipes, which lacks forewing fringe), often
almost linear (Fig. 7). Notaulices sometimes posteriorly absent (Fig. 15), always sharply angled
outwards anteriorly (when viewed from directly above) (Figs 14-17). Eye pubescence short
and inconspicuous, pubescence of upper face short, or if long then ocellar triangle linear.
Interantennal carina without projection, frons behind this carina flat or slightly raised with
depression inwards from each torulus. Frontal dent small except in D. pupparum (Fig. 8). Head
alutaceous, without clear frontal line or sculptured ocular suture. Occipital carina weak.
Scutellum usually with raised central area, or if not raised then ocellar triangle broad. Raised
area of scutellum without margin of sculpture. Mesonotum usually quadrate (Figs 14-17).
Head often clearly transverse (Fig. 7). Fresh specimens are brown to black, or if lighter then
notaulices incomplete, or wing pubescence strongly reduced, or brachypterous with AIII
elongate (Fig. 58). Brachypterous forms are exceptionally rare . DENDROCERUS (part) (p. 259)
Ocellar triangle usually long with short base (Figs 5, 6) and forewing fringe present. Notaulices
complete and usually only slightly, smoothly angled outwards anteriorly (when viewed from
directly above) (Fig. 13). Eye pubescence often long or dense and conspicuous, pubescence of
face frequently long. Interantennal carina often with a small sharp median projection (Fig. 6),
frons behind this carina frequently distinctly concave (Fig. 6). Frontal dent often strongly
developed. Head and ocular suture sometimes strongly sculptured and foveolate frontal line
occasionally present. Occipital carina sometimes projecting and foveolate. Scutellum often flat,
sometimes with margin of foveolate sculpture. Mesonotum often obviously narrowed anteriorly
(Fig. 13). Head frequently globular or rounded (Fig. 6). Body colour often light brown.
Frequently brachypterous CONOSTIGMUS Dahlbom (part)
DENDROCERUS Ratzeburg
Dendrocerus Ratzeburg, 1852: 180—181. Type-species: Dendrocerus lichtensteinii Ratzeburg [— Ceraphron
halidayi Curtis], by monotypy.
Lygocerus Foerster, 1856: 97. Type-species: Ceraphron ramicornis Boheman, by subsequent designation
(Ashmead, 1893: 107).
Macrostigma Rondani, 1877: 184. Type-species: Macrostigma aphidum Rondani, by monotypy.
[Synonymized by Dessart, 19650: 157.]
Atritomus Foerster, 1878: 56. Type-species: Atritomus coccophagus Foerster [= Ceraphron laevis
Ratzeburg], by monotypy. [Homonym of Atritomus Reitter, 1877: 384.]
Prodendrocerus Kieffer, 1907: 11. Type-species: Lygocerus ratzeburgi Ashmead [= Ceraphron ramicornis
Boheman], by subsequent designation (Muesebeck & Walkley, 1956: 389). [Synonymized by Dessart,
1966: 5.]
A tritomellus Kieffer, 1914: 141. [Replacement name for Atritomus Foerster.] [Synonymized by Hellen, 1966:
9.]
Neolygocerus Ishii, 1951: 16. Type-species: Neolygocerus koyamai Ishii, by monotypy. [Synonymized by
Dessart, 1966: 11.]
Basoko Risbec, 1958: 111. Type-species: Basoko africana Risbec, by monotypy. Syn. n.
Antenna eleven-segmented in both sexes (except single fossil female of D. dubitatus Brues, which has ten
segments). In females scape long, often longer than combined length of next three segments. Pedicel
260
N. D. M. FERGUSSON
Fig. 1 Dendrocerus carpenteri (Curtis), male.
elongate (Figs 56-63), flagellar segments cylindrical. In males scape shorter and stout, usually shorter than
combined length of next three segments. Pedicel small and almost globular (Figs 51-55). Basal flagellar
segments of males never completely radially symmetrical, varying from almost cylindrical in D. bifoveatus
(Fig. 55) to serrate in many species (Figs 50-52), strongly serrate in D. serricornis (Fig. 53) and ramose in
D. ramicornis (Fig. 31); asymmetry progressively decreases towards distal segments. Pubescence of flagellum
in males often longer than breadth of segment, pubescence in females shorter. Antennae alutaceous or
papillate, inserted low on face close to clypeus.
Head lightly sculptured (except D. spissicornis, in which it is rugose); pubescence usually very short, eye
pubescence short and inconspicuous; generally flat and transverse; usually black or dark brown. Mandibles
bidentate. Toruli not strongly extended. Interantennal carina absent or distinctly developed, if present often
with slight central hump. (Some species of Conostigmus have short sharp medial projection between toruli
(Fig. 6) but this does not occur in Dendrocerus.) Frons behind interantennal carina flat, or slightly raised with
depression inwards from each torulus, without large concave area. Except in D. pupparum (Fig. 8), frontal
dent absent or present but small. Vertex slightly convex, or biconvex with slight longitudinal median
depression but with no clear frontal line. Ocular suture not strongly sculptured. Ocelli present in both sexes.
Ocellar triangle short with broad base (e.g. Fig. 7). Anterior ocellus situated just in front of postocellar line.
Preoccipital crescent absent, or strongly developed and nearly reaching eyes and ocelli, sharply angled away
from vertex. Occipital carina weak.
Side of pronotum with Y-shaped furrow, posterior arm of furrow sometimes faint. Mesonotum usually
quadrate (Figs 14, 16, 17), not obviously narrowed anteriorly. Median furrow present. Notaulices sometimes
absent posteriorly (Fig. 15) but always present anteriorly, sharply angled outwards (when viewed from
directly above) towards anterior corners of mesonotum. In D. spissicornis notaulices not quite so sharply
angled. Secondary furrows often present (Fig. 14). Scutellum generally convex with raised central area, only
rarely with sculpture on margin of raised area. Precoxal sulcus absent. Metanotum reduced. Propodeum
often with median fovea. Propodeal spiracle normal. Tibiae of all legs with two spines. Large spine of anterior
leg forked at its apex. Claws simple. Brachypterous forms are exceptionally rare. Forewing with large
pterostigma (Figs 33-36, 38-46). Radial vein longer than pterostigma (except in two non-British species).
BRITISH SPECIES OF DENDROCERUS 261
Hindwing with three hamuli, proximal one almost straight, others strongly curved. Anal lobe present on
hindwing in D. ramicornis (Fig. 37). Wings fully pubescent with fringe except in D.punctipes.
Large tergite of gaster with crenulate collar (Fig. 10), clearly with neck, except in D. spissicornis which has
short broad collar (Fig. 19). Rest of gaster alutaceous or smooth. Gastrocoeli often visible. Hypopygium
without distinct tuft of upright hairs. Genitalia with lamina volsellaris fused for greater part along its
boundary with paramere.
FUNCTIONAL MORPHOLOGY OF MALE ANTENNAE. The most notable character of Dendrocerus
males is the lateral extension of the basal flagellar segments. This character reaches its greatest
development in the branches of D. ramicornis (Fig. 3 1) and D. halidayi (Fig. 32). It is likely that this
is a device to increase the olfactory efficiency of the male, probably in its response to sexual
attractants liberated by the female. Kaissling (1971: 351) has shown that the efficiency of an
antenna as an odour filter can be defined by three terms: the outline area, the adsorption quotient
and the effective fraction of adsorption. All these functions are mainly dependent upon the external
geometry of the antenna. The most sensitive olfactory apparatus could be expressed as an odour
filter with an extended outline area and a good subdivision of form. The antennae of male
Saturniidae (Lepidoptera) approach most closely to this ideal.
Subgenera
After clarifying the generic position of Dendrocerus, Dessart (1972a: 26) retained some of the
generic synonyms as subgenera; subgenus Dendrocerus for males with branched antenna,
subgenus Atritomellus for species with the notaulices absent posteriorly, subgenus Neolygocerus
for a Japanese species with a specialized scutellum, and subgenus Macrostigma for the remaining
species.
The use of these subgenera is neither significant nor helpful. The separation of subgenus
Dendrocerus applies only to males. The male of D. (Neolygocerus) koyamai (Ishii) has branched
antennae like those found in subgenus Dendrocerus, and D. (Dendrocerus) halidayi often has the
notaulices absent posteriorly as in the species of subgenus Atritomellus. The two European species
of subgenus Dendrocerus have very little in common in either morphology or biology. Atritomellus
is easily split into two groups on morphology, and the dumping of the bulk of the genus under
Macrostigma has no advantage. For the above reasons these subgenera are listed as generic
synonyms and omitted from further consideration in this paper.
Synonymic list of British species
(* = new to the British list)
aphidum (Rondani, 1877)
rufipes (Thomson, 1858) (junior homonym) syn. n.
aphidorum (Rondani, 1874) nomen nudum
koebelei (Ashmead, 1904) syn. n.
subquadratus (Kieffer, 1907) syn. n.
fusciventris (Kieffer, 1907) syn. n.
frenalis (Kieffer, 1907) syn. n.
breadalbimensis (Kieffer, 1907) syn. n.
bicolor (Kieffer, 1907) syn. n.
fuscipennis (Kieffer, 1907)
neglectus (Kieffer, 1907)
lundensis Dessart, 1966 syn. n.
btfoveatus (Kieffer, 1907)
sordidipes (Kieffer, 1907)
carpenteri (Curtis, 1829)
crispus (Curtis, 1829) nomen nudum
elegans (Curtis, 18 29) nomen nudum
hyalinatus (Thomson, 1858)
262 N. D. M. FERGUSSON
niger (Howard, 1890)
proximus (Kieffer, 1907)
punctatipennis (Kieffer, 1907) syn. n.
rufiventris (Kieffer, 1907) (junior homonym)
campestris (Kieffer, 1907)
aphidivorus (Kieffer, 1907)
testaceimanus (Kieffer, 1907)
aphidum (Kieffer, 1907) (junior homonym)
giraudi (Kieffer, 1907)
cameroni (Kieffer, 1907)
thomsoni (Kieffer, 1907)
inquilinus (Kieffer, 1917)
ambianus (Dessart, 1965)
britannicus Dessart, 1966
tischbeini Dessart, 1966
dubiosus (Kieffer, 1907) sp. rev.
longicornis (Thomson, 1858) (junior homonym)
pallipes (Kirchner, 1867) nomen nudum
flavipes (Kieffer, 1907) (junior homonym) syn. n.
claripennis (Kieffer, 1907) syn. n.
rectangularis (Kieffer, 1907) syn. n.
alpestris (Kieffer, 1907) syn. n.
navaensis Dessart, 1966 syn. n.
flavus (Hellen, 1966) syn. n.
*flavipes Kieffer, 1907
fuscipes Kieffer, 1907
halidayi (Curtis, 1829)
lichtensteini Ratzeburg, 1852
damicornis (Foerster, 1856) nomen nudum
callicerus (Thomson, 1858)
*laevis (Ratzeburg, 1852)
frontalis (Thomson, 1858)
coccophagus (Foerster, 1878)
smirnoffi (Ghesquiere, 1960)
applanatus Dessart, 1972 syn. n.
*terice/«(Hedicke, 1929)
*liebscheri Dessart, 1972
tenuicornis (Thomson, 1858) (junior homonym)
*punctipes (Boheman, 1832)
parvulus (Wollaston, 1858)
pupparum (Boheman, 1832)
ancyloneurus (Ratzeburg, 1844)
syrphidarum (Kieffer, 1907)
ramicornis (Boheman, 1832)
glabriculus (Thomson, 1858)
japonicus (Ashmead, 1904)
ratzeburgi (Ashmead, 1904)
serricornis (Boheman, 1832)
piceus (Ratzeburg, 1852)
lapponicus (Thomson, 1858)
semiramosus (Kieffer, 1907) syn. n.
subramosus (Kieffer, 1907)
zetterstedti (Ghesquiere, 1960)
*spissicornis (Hellen, 1966)
BRITISH SPECIES OF DENDROCERUS 263
Key to British species
The recognition of D. liebscheri and the separation of D. aphidum and D. dubiosus is difficult and should be
based on series of both males and females. The male of D. punctipes is not known; it may well have the
forewing pubescence reduced as in the female. An explanation of AI, L/B etc. is given on p. 257.
1 Males (All globular, as in Figs 5 1-55) .......... 2
Females (All elongate, as in Figs 56-63) ......... 14
2 AIII to AVI each bearing long process always considerably longer than the segment which bears
it(Figs31,32) . 3
Antenna without long processes; any projection present always equal to or shorter than segment
which bears it (Figs 30, 47-55) 4
3 AVII longer than other antennal segments, longer than process it carries (Fig. 32); raised area of
scutellum long and narrow, 1-5-2-0 times longer than broad. Hindwing without anal lobe;
pterostigma semioval (Fig. 45); body colour brown ..... halidayi(p.219)
AVIII longer than other antennal segments; AVII shorter than process it carries (Fig. 31); raised
area of scutellum broad, 1-1-1-3 times longer than broad. Hindwing with small anal lobe (Fig.
37);pterostigmasemicircular(Fig.41);bodycolourblack .... ramicornis (p. 288)
4 Notaulices incomplete, fading out anteriorly just after bend (Fig. 15). . . . . 5
Notaulices complete, reaching posterior margin of mesonotum (Figs 14, 16, 17) ... 7
5 AIII approximately same length as scape (Fig. 30); flagellum densely pubescent; proximal
flagellar segments strongly papillate; flagellar pubescence short, shorter than maximum breadth
of flagellar segments; mesonotum and scutellum relatively elongate (i.e. L/B of mesonotum
0-7-0-8 . flavipes(p.218)
AIII clearly shorter than scape (Figs 47, 48); flagellum sparsely pubescent; proximal flagellar
segments at most with only a few papillae; flagellar pubescence long, at least as long as
maximum breadth of flagellar segments; mesonotum and scutellum usually broad, only
elongate in small specimens (i.e. L/B of mesonotum 0-4— 0-7) .... 6
6 Flagellar segments short (Fig. 48) (e.g. L/B for AIII 1-5-2-5); scape short, about equal to length
of eye; preoccipital crescent normally strongly developed, close to margin of eye (Fig. 25);
scape without distinct light-coloured basal ring, usually completely dark (considerable light
coloration occasionally present but vaguely delimited) ..... laevis(p. 281)
Flagellar segments elongate (Fig. 47) (e.g. L/B for AIII 2-4-3-0); scape often long, always longer
than length of eye; preoccipital crescent weakly developed, not reaching margin of eye (Fig. 26);
scape nearly always with distinct light-coloured basal ring, never completely dark, often with
sharply defined longitudinal light-coloured stripe ventrally .... laticeps(p. 283)
7 Entire breadth of large gastral tergite covered by strigose sculpture, radiating from gastral collar
and nearly reaching to gastrocoeli (Fig. 19); apex of scutellum with transverse carina;
propodeum with large bilobed transverse keel (Fig. 19); sculpture coarse, mostly rugose .
spissicornis (p. 292)
Large gastral tergite at most with short, narrow area of carinae not covering its entire breadth
and not reaching gastrocoeli; apex of scutellum without transverse carina; propodeum without
bilobed transverse keel, usually with median fovea ....... 8
8 Antenna strongly serrate (Figs 51-53) (L/B for AIII 2-0 or less). Interantennal carina absent or
indistinct, if weakly present then L/B for AIV approximately 1-0 (Fig. 53). Forewing always
clear. Pterostigma approximately semicircular and radius deeply angled (Figs 40, 42, 43) . 9
Antenna almost cylindrical (Figs 54, 55) or only moderately serrate (Figs 49, 50) (L/B for AIII
usually greater than 2-0, if less then interantennal carina present and forewing with area of
brown coloration). Interantennal carina often present and forewing often with brown area.
Pterostigma semioval, radius shallowly angled or evenly curved (Figs 33, 38, 44, 46) . 1 1
9 Basal flagellar segments very strongly serrate (Fig. 53), AIV as long as broad (L/B 0-9-1-2).
Preoccipital crescent very strongly developed, reaching eyes. Longest hairs on AVIII at least as
long and usually much longer than maximum length of AIX. Thorax and gaster often long and
narrow (L/B for thorax 1-4—1-8). Anterior corners of mesonotum often strongly curved
downwards and rounded ......... serricornis (p. 290)
Basal flagellar segments less strongly serrate (Figs 51, 52), AIV longer than broad (L/B 1-4 or
more). Preoccipital crescent well developed but not quite reaching eyes. Longest hairs on AVIII
not quite as long as maximum length of AIX. Thorax and gaster always broad (L/B for thorax
1 • 5 or less). Anterior corners of mesonotum never strongly curved downwards . . .10
10 Notaulices strongly convergent posteriorly, meeting or almost meeting median furrow at scutal
suture (Fig. 17). Parameres (viewed laterally in situ) expanded apically and broadly truncate
264 N. D. M. FERGUSSON
(Fig. 27). Pterostigma (Fig. 42) slightly less semicircular, radius (Fig. 42) slightly less deeply
angled than in following species. Gaster often narrow, scutellum narrow and strongly raised. A
rare species hyperparasitic on lachnid aphids on conifers .... liebscheri (p. 285)
Notaulices only slightly convergent, not meeting median furrow (Fig. 14), or if they meet or
almost meet median furrow then they converge in curve (Fig. 16). Parameres (viewed laterally
in situ) narrow and apically rounded (Fig. 28). Pterostigma (Fig. 40) (without leading edge
curled over) approximately semicircular, radius (Fig. 40) deeply angled. Gaster usually broad,
scutellum broad and not so high. An extremely common and widespread species on a wide
range of aphids and plants cwpenteri(p.21G)
11 Pterostigma distinctly long and narrow (Fig. 38), more than 2-1 times longer than broad.
Mesepisternum almost totally smooth and shining. Flagellar segments almost cylindrical, only
very weakly serrate (Fig. 55), flagellar pubescence longer than breadth of segment
btfoveatus (p. 269)
Pterostigma (without leading edge curled over) relatively broad (Figs 33, 44, 46), less than 2-0
times longer than broad. Mesepisternum clearly with alutaceous sculpture, only rarely is
sculpture fine. Basal flagellar segments moderately serrate (Figs 49, 50), or if almost cylindrical
(Fig. 54) then flagellar pubescence shorter than breadth of segment . •- . . . 12
12 Apex of parameres strongly upturned or expanded along dorsal edge (Fig. 29). Flagellar
pubescence shorter than breadth of segment. Frontal dent strongly developed, circular and
slightly linear (Fig. 8). Flagellum from AIV onwards weakly serrate, almost cylindrical (Fig.
54). Radius evenly and shallowly curved (Fig. 44) .... . pupparum(p. 287)
- Apex of parameres simple, not upturned or expanded. Flagellar pubescence at least as long as
breadth of segment. Frontal dent small and shallow, or absent. Flagellum serrate, or if weakly
serrate then radius angled (Figs 33, 46) ......... 13
13 Scape short and often stout, 4-0-5-1 times longer than broad, clearly shorter than lateral length
of head. Scape often completely black, or with a small amount of yellow at base, sometimes
entirely yellow but then scape short and broad, about 4-2 times longer than broad. Legs usually
mostly dark, if yellow then scape short and broad (L/B about 4-2) . . . aphidum (p. 266)
Scape longer and thinner, 5-0 or more times as long as broad, nearly equal to lateral length of
head. Basal third of scape usually yellow, legs not totally dark, often mostly yellow dubiosus (p. 276)
14 Thorax long and narrow, 1-5-1-9 times longer than broad, notaulices only clearly present
anteriorly, fading out just after bend (Fig. 15). Body colour brown 15
Thorax broad and stocky, at most 1-4 times as long as broad, or if long and thin then notaulices
complete and distinct. Notaulices complete (Fig. 14), or if absent posteriorly then body length
1 • 5 mm or less, or body colour black (in fresh specimens) 16
15 Forewing with a distinct brown band under pterostigma and radial vein (Fig. 35); gastral collar
white or light yellow, strongly contrasting with surrounding body colour; antenna dark brown
but often scape lighter basally ........ flavipes (p. 278)
Forewing virtually clear; body colour uniformly brown; antenna entirely brown . halidayi(p. 279)
1 6 Notaulices incomplete, present only anteriorly, fading out just after bend (Fig. 15). . . 17
Notaulices complete, reaching posterior margin of mesonotum (Fig. 14) . . . . 18
1 7 Flagellar segments elongate (e.g. AIII longer than 0- 1 mm, except in teneral specimens) (Fig. 60),
first flagellar segment longer than pedicel; preoccipital crescent weakly developed (Fig. 26),
clearly separated from margin of eye; scape never completely dark, normally with a distinct
light-coloured basal ring (sometimes indistinct in teneral and old specimens); gastral collar long
(mid lateral length normally greater than 0-04 mm); total body length very variable
(1-3-2-0 mm) laticeps (p. 283)
Flagellar segments short (e.g. AIII less than 0-1 mm) (Fig. 61), first flagellar segment at most as
long as pedicel, usually shorter; preoccipital crescent normally well developed (Fig. 25), often
close to margin of eye; scape usually completely dark, without distinct light-coloured basal
ring, occasionally considerable area is light-coloured but vaguely delimited; gastral collar short
(mid lateral length less than 0-04mm); total body length less than 1-5 mm, often much smaller
/aevfc(p.281)
18 Entire breadth of large gastral tergite covered by strigose sculpture radiating from gastral collar
and nearly reaching gastrocoeli (Fig. 19); apex of scutellum with transverse carina; propodeum
with large bilobed transverse keel (Fig. 19); sculpture coarse, mostly rugose . spissicornis (p. 292)
Large gastral tergite at most with short, narrow area of carinae, not covering entire breadth of
tergite and not reaching gastrocoeli; apex of scutellum without transverse carina; pro-
podeum without bilobed transverse keel, usually with median fovea 19
BRITISH SPECIES OF DENDROCERUS 265
19 Pubescence of forewing virtually absent, few fringe hairs present on costal vein and hairs on wing
ultra-short. Scutellum completely flat, interantennal carina complete and prominent. Eyes
large, occupying most of lateral length of head. Forewings with dark brown mark near
pterostigma. Brachypterous specimens are known ...... punctipes (p. 286)
Pubescence of forewing normal, long fringe hairs and hairs on centre of wing conspicuous.
Scutellum mostly convex, or if slightly flattened then interantennal carina absent, interrup-
ted, or only weakly developed ........... 20
20 Mesepisternum almost totally smooth and shining. Pterostigma distinctly long and narrow (Fig.
38), more than 2-1 times longer than broad. Legs mostly dark, interantennal carina weak but
present, forewing evenly light brown, scape dark btfoveatus (p. 269)
- Mesepisternum clearly with alutaceous sculpture, only rarely is sculpture fine. Pterostigma (when
flat and without leading edge curled over) relatively broad (Figs 33, 40-44, 46), less than 2-0
times longer than broad; if pterostigma is almost twice as long as broad then legs mostly
yellow, or interantennal carina absent, or forewing with conspicuous brown area, or base of
scape yellow .............. 2 1
2 1 Frontal dent strongly developed, circular and slightly linear (Fig. 8). Forewing with brown area
under radius, which is very shallowly curved (Fig. 44). Interantennal carina incomplete (Fig. 8),
absent or very indistinct centrally. Scape brown. Body length 1-9-3-2 mm . pupparum (p. 287)
Frontal dent small and shallow, or absent. Forewing clear, radius distinctly angled (Figs 40—43),
if otherwise then interantennal carina complete and prominent, or scape distinctly yellow at base,
or body length less than 1-9 mm .......... 22
22 Forewing with area of brown pigmentation near pterostigma (Fig. 33). Interantennal carina
present, usually conspicuous. Body length less than 1-9 mm, body not long and narrow (L/B
for thorax 1-5 or less). Pterostigma semioval (Figs 33, 45). Notaulices only moderately
convergent, not meeting median furrow. Legs often mostly yellow or yellow-brown . . 23
Forewing uniformly clear. Interantennal carina absent, or if very weakly developed then body
length greater than 2- 1 mm, or head (in dorsal view) rounded, anterior corners of mesonotum
distinctly rounded and body clearly long and narrow )e.g. L/B for thorax 1-4-1-8). Pterostigma
almost semicircular (Figs 40, 41, 43), or if semioval (Fig. 42) then notaulices strongly
convergent posteriorly, meeting median furrow at scutal suture (Fig. 17). Legs often mostly
dark 24
23 AIII clearly much longer than All (Fig. 58). Scape 5-7-7-0 times longer than broad, much longer
than head. Legs entirely yellow, very rarely basal half of hind coxa dark. Scape yellow basally.
(Three brachypterous specimens are known and these have the scape slightly shorter than
normal) dubiosus(p.216)
— AIII about equal to or slightly longer than All (Fig. 59). Scape less than 5-7 times longer than
broad, about equal to or shorter than head. Legs always dark on basal half of hind coxa,
usually dark on outside of hind femur and sometimes dark elsewhere. Scape usually mostly
black but sometimes entirely yellow. (No known brachypterous specimens) . . aphidum (p. 266)
24 Head (in dorsal view) rounded; anterior corners of mesonotum strongly downcurved and
rounded. Body clearly long and narrow (e.g. L/B for thorax 1-4-1-8). Preoccipital crescent
strongly developed and virtually reaching eye margin.
Body length 1-4-1-8 mm, pterostigma as in Fig. 43. Parasite of Diptera: Chamaemyiidae on
Hemiptera: Adelgidae serricornis (p. 290)
- Head clearly transverse, not rounded; anterior corners of mesonotum not obviously downcurved
or rounded. Body not notably long and narrow (e.g. L/B for thorax 1-5 or less). Preoccipital
crescent present but not reaching eye margin, or if it does then body length 2- 1 mm or more 25
25 Body large, 2- 1-2-8 mm long; AIII clearly longer than All (about 1-5 times longer).
Interantennal carina present but not prominent. Hindwing with small pronounced anal lobe
(Fig. 37). Legs often mostly light brown. Pterostigma distinctively shaped, approximately
semicircular (Fig. 41).
Due to large size, head appears to be strongly transverse. Preoccipital crescent usually
reaching close to margin of eye. Hyperparasite of Hemiptera: Lachnidae, often on coniferous
and deciduous trees .......... ramicornis (p. 288)
- Body smaller, only rarely as long as 2- 1 mm; AIII approximately as long as AIL Interantennal
carina absent. Anal lobe indistinct or absent. Legs mostly dark. Pterostigma roughly
semicircular (Fig. 40) or semioval (Fig. 42) ......... 26
26 Scape long (5-7 times longer than broad), longer than lateral length of head. AV to AIX elongate
(Fig. 56) (e.g. L/B for AVI 1-6-2-0). Notaulices strongly convergent posteriorly, meeting or
266 N. D. M. FERGUSSON
almost meeting median furrow at scutal suture (Fig. 17). Pterostigma (Fig. 42) slightly less
semicircular and radius (Fig. 42) slightly less deeply angled than in following species. A rare
species hyperparasitic on lachnid aphids on conifers .... . liebscheri (p. 285)
Scape shorter (at most 5 times as long as broad), shorter than lateral length of head. AV to AIX
not so long (Fig. 57) (e.g. L/B for AVI 1-1-1-6). Notaulices only slightly convergent, not
meeting median furrow (Fig. 14), or if they meet or almost meet median furrow then they
converge in curve (Fig. 16). Pterostigma (Fig. 40) (without leading edge curled over) approx-
imately semicircular, radius (Fig. 40) deeply angled. An extremely common and widespread
species on a wide range of aphids and plants . . . . - . . carpenteri(p.21Q)
Species descriptions
Dendrocerus aphidum (Rondani )
(Figs 3 3, 50, 59)
Ceraphron rufipes Thomson, 1858: 293. Lectotype 9, SWEDEN (NR, Stockholm), designated by Dessart
(1972a: 102) [examined]. [Primary homonym of Ceraphron rufipes Nees, 1834: 277. ] Syn. n.
Macrostigma aphidorum Rondani, 1874: 134. [Nomen nudum.] ['Synonymized' by Dessart, 1965a: 157.]
Macrostigma aphidum Rondani, 1877: 184-185, pi. 1, figs 34, 36. Lectotype tf, ITALY (MZU, Florence),
designated by Dessart (1965a: 158) [examined].
Lygocerus koebelei Ashmead, 1904: 70. Lectotype $, JAPAN (USNM, Washington), designated by Masner
& Muesebeck (1968: 1 12) [examined]. Syn. n.
Lygocerus subquadratus Kieffer, 1907: 42. Lectotype $, GREAT BRITAIN (BMNH), designated by Masner
(1965: 21) [examined]. Syn. n.
Lygocerus fusciventris Kieffer, 1907: 47. Lectotype <$, GREAT BRITAIN (BMNH), designated by Masner
(1965: 20) [examined]. Syn. n.
Lygocerus frenalis Kieffer, 1907: 47-48. Lectotype <$, GREAT BRITAIN (BMNH), designated by Masner
(1965: 20) [examined]. Syn. n.
Lygocerus breadalbimensis Kieffer, 1907: 53-54. Lectotype 9» GREAT BRITAIN (BMNH), designated by
Masner (1965: 20) [examined]. Syn. n.
Lygocerus bicolor Kieffer, 1907: 62. Lectotype 9, GREAT BRITAIN (BMNH), designated by Masner (1965: 20)
[examined]. Syn. n.
Lygocerus fuscipennis Kieffer, 1907: 64. Lectotype 9» ITALY (MCSN, Genoa), designated by Dessart
(1972a: 59) [examined]. [Synonymized by Dessart, 1972a: 52.]
Lygocerus neglectus Kieffer, 1907: 64-65. Syntype(s) 9, FRANCE: Amiens, August (Carpentier) (lost).
[Synonymized by Dessart 1972a: 52.]
Lygocerus aphidum (Rondani) Dessart, 1965a: 157.
Dendrocerus aphidum (Rondani) Dessart, 1966: 12.
Dendrocerus lundensis Dessart, 1966: 12. [Replacement name for Ceraphron rufipes Thomson, 1858.]
Syn. n.
FEMALE. Scape length variable (L/B less than 5-7), but longer than All, AIII and AIV together, about equal
to or less than length of head. Scape usually black with a little yellow at base, sometimes entirely yellow. All
approximately equal to length of AIII, often slightly shorter than AIII. Flagellar segments flat ventrally.
Apex of All slightly lighter than rest of flagellum which is black or brown. Antenna weakly papillate,
pubescence short, quite dense.
Head alutaceous; pubescence short, except on clypeus, black; mandibles brown with red teeth, palps
colourless. Head somewhat rounded but still transverse. Frons rather flat but with central swelling and small
depression behind each torulus. Frontal dent absent, or small and shallow. Depression in front of median
ocellus small, depressions by lateral ocelli shallow. Interantennal carina with central hump. Toruli not
prominent. Ocellar triangle broad and short, Lateral Ocelli linked posteriorly by faint groove. Vertical furrow
absent or very faint. Preoccipital crescent small, not reaching eyes and only rarely near ocelli.
Thorax black or very dark brown, alutaceous, with short pubescence. Notaulices complete, angled
outwards but not as acutely angled as in most Dendrocerus species. Secondary furrows not always visible.
Axillae only downcurved near outer edge. Scutellum distinctly arched but central area occasionally rather flat.
Metanotal furrow strongly foveolate. Pleural sulcus foveolate. Propodeum with distinct median fovea. Leg
colour variable but basal half of hind coxa always dark, usually outside of hind femur and often rest of legs
dark. Tibiae and tarsi often mostly yellow. Forewing with brown area under pterostigma and often dark in
proximal region so that basal veins are indicated as clear lines. Pterostigma semiovoid, 1-6-1-9 times longer
than broad. Radius about 1 • 3 times longer than pterostigma. Wing pubescence normal.
BRITISH SPECIES OF DENDROCERUS 267
Gastral collar long with many carinae, of which three are usually prominent. Gaster mostly smooth.
Gastrocoeli visible as light brown areas. Basal half of gaster sometimes lighter in colour than posterior half.
MALE. As for female except scape short and often stout, at most 5- 1 times longer than broad, clearly shorter
than lateral length of head; longer than All and AIII together. Scape often completely black or with small
area of yellow at base, if entirely yellow, then it is also short and broad, about 4-2 times longer than broad.
AIII longer and more slender than AIV (Fig. 50). Basal flagellar segments serrate (Fig. 50), their pubescence
longer than breadth of segment.
REMARKS. The study of many specimens has shown that D. aphidum is very variable in size and
coloration. I have been unable to find any constant method of separating the usually larger and
lighter specimens (breadalbimensis) from the usually smaller and darker specimens of D. aphidum.
Previously D. aphidum and D. breadalbimensis were distinguished by leg, forewing and scape
coloration; length of certain antenna! segments; degree of development of interantennal carina; and
differences in the male genitalia. Leg coloration varies to such an extent that it is an unreliable
character at the species level. Forewing colour provides some discrimination but several specimens
with quite dark wings (showing basal veins) are otherwise typical examples of D. breadalbimensis.
The differences in length of the scape and flagellar segments reflect size variation. Several series of
very dark and very small specimens of D. aphidum from Iceland may have unduly influenced
previous concepts of this species. The scape coloration does not appear to give a reliable distinction
and there are no interantennal carina differences. The main separating character was the male
genitalia. In D. aphidum the volsella plates appeared to have a distinctive sinuous, angular
termination and a non-apical hair, whereas in D. breadalbimensis (and D. bicolor) they were
rounded with an apical hair. The 'sinuate form' of genitalia is in fact a slightly lateral view of the
'round form' of genitalia. The pubescence changes its position relative to the apex in the same way.
In view of these considerations I am regarding all these specimens as conspecific.
The lectotypes of Lygocerus bicolor, L. fusciventris, L. frenalis and L. subquadratus are all
large specimens of D. aphidum (= L. breadalbimensis). (The paralectotype of L. bicolor and the
lectotype of L. breadalbimensis were collected together.) D. bicolor auct. nee Kieffer is D. dubiosus
(see p. 276), the females of this species being easily recognizable by their long AIII; females of D.
aphidum have All and AIII about equal in length. The paralectotype of Conostigmus dubiosus is a
specimen of D. aphidum and thus not conspecific with the lectotype of C. dubiosus. The lectotype of
Lygocerus koebelei is a specimen of D. aphidum; the paralectotypes are D. carpenteri and D.
serricornis.
It is not certain that this complex consists of only two species. Apart from the large specimens
(breadalbimensis) there is a small group of males that I have tentatively placed in D. aphidum.
These have yellow legs and scape and the latter is very broad and short, but I am not convinced
that they are separable from D. aphidum on such characters; also they occur with typical specimens
of D. aphidum.
The species separation detailed above and in the keys is the most practical arrangement possible
at present, although it might be improved when reared series of both sexes are obtained and the
biology is well known.
After D. carpenteri, D. aphidum (as defined above) is one of the most common species of
Dendrocerus.
BIOLOGY. The following three host records for D. rufipes are probably misidentifications of other
Dendrocerus or Conostigmus species. Henriksen (1918: 140) recorded D. rufipes from fly pupae.
Mani (1941: 27) recorded D. rufipes from a chrysopid (Neuroptera) on sugar-cane and Pruthi &
Mani (1942:421) gave Chrysopa virgestes (Neuroptera) as a host. All other host data indicate that
D. aphidum (= rufipes) is a secondary parasite of aphids. The records for D. aphidum are too
extensive to register in full; the following are alphabetical lists of hosts and parasites.
Hosts (Aphidoidea): Acyrthosophon malvae, A. pisum, A. sp., Aphis evonymi, A. nasturtii, A.
pomi, A. sp., Brachycaudus persicae, Cavariellia aegopodii, Clethrobius comes, Diuraphis
frequens, D. noxius, Eucallipterus tiliae, Hyalopterus pruni, Liosomaphis berberidis,
Macrosiphoniella usquertensis, Metopolophium dirhodum, M. sp., Microlophium carnosum, M.
sp., Myzus persicae, Paraschizaphis scirpi, Rhopalosiphum nymphaeae, Schizaphis graminum,
Sipha maydis, Sitobion avenae, S.fragariae, Uroleucon cirsii, U. sp.
268 N. D. M. FERGUSSON
Primary parasites (Aphidiidae): Aphidius ervi, A. picipes, A. platensis, A. sonchi, A.
uzbekistanicus, A. sp., Binodoxys acalephae, Diaeretiella rapae, Ephedrus persicae, E. plagiator,
E. sp., Praon abjectum, P. necans, P. volucre, P. sp., Trioxys betulae.
The records of hosts and parasites listed above are taken from material examined and from the
following references: Boness (1958: 322); Dessart (1972a: 58; 1975: 267-269); Ivanova-Kazas
(1955: 151); Kieffer (1907 63); Kryger & Schmiedeknecht (1938: 76); Malaquin & Moitie (1914:
804); Meyer (1929: 234-244); Rondani (1877: 184) and Stary (1977: 2).
The host information for D. aphidum includes records that are typical elements of certain
faunistic complexes. The record Aphis sp./Binodoxys acalephae /Caltha palustris along with
several other records indicate that D. aphidum is part of the Eurasian steppe faunistic complex. The
record Clethrobius comesl Trioxys betulae/ Alnus incana represents the Holarctic forest tundra
faunistic complex, and Rhopalosiphum nymphaeae/ Praon necans/ Sagittaria sp. is typical of the
Boreal Europe faunistic complex. There are several records from grass, one from the complex of
parasites on apple, and a few from deciduous forest. The aphids are Chaitophoridae (1 record),
Callaphididae (2 records) and Aphididae; over half of the last family are Macrosiphinae,
Acyrthosiphon (11 records) and Sitobion (8 records) being the most frequently recorded genera.
The Callaphididae are a deciduous forest group. The Aphidiidae listed include most of the
common genera, Aphidius and Praon being the most frequently recorded. The recorded plants are
shrubs, herbs, a few trees, and many crop and grassland plants, and include both Monocotyledons
and Dicotyledons. Several plants are associated with wet habitats, e.g. Sagittaria, Typha and
Angelica. Thus D. aphidum is associated with cooler conditions, wet places, grass and deciduous
trees and is commonly found on crops. It has been collected from March to November, its numbers
reaching a peak in June and July.
MATERIAL EXAMINED
Ceraphron rufipes Thomson, lectotype 9, Sweden: 'Ld. [Lund] 9/57' (NR, Stockholm). Macrostigma
aphidum Rondani, lectotype $, Italy: (locality unknown) (MZU, Florence). Lygocerus koebelei Ashmead,
lectotype <$, Japan: Atami (Koebele) (USNM, Washington). Lygocerus subquadratus Kieffer, lectotype <$,
Great Britain: [Scotland, Central] Clober (Cameron). Lygocerus fusciventris Kieffer, lectotype <$, Great
Britain: [Scotland, Central] Mugdock (Cameron). Lygocerus frenalis Kieffer, lectotype $, Great Britain:
[Scotland, Strathclyde] Lock Libo (Cameron). Lygocerus breadalbimensis Kieffer, lectotype 9> Great Britain:
[Scotland, Tayside] 'Ben Lawers, along with 1' [1 = paralectotype ofbicolor]] (Cameron). Lygocerus bicolor
Kieffer, lectotype 9, Great Britain: [Scotland, Dumfries and Galloway] New Galloway (Cameron). Lygocerus
fuscipennis Kieffer, lectotype 9, Italy: Giglio I., iv. 1902 (Dorid) (MCSN, Genoa).
239 cf, 147 9- Algeria: Esbarres; Oran. Canary Is.: Tenerife. Egypt: Siwa. France: Dordogne,
Mouleydier; Pyrenees-Orientales. Germany (East): Harz Mts, Brocken dist.; Hinnundviedust, Altvater.
Great Britain. Bedfordshire: Dunstable Downs, Eaton Bray, Steppingly; Berkshire: nr Streatly, Lambourn
Down, Silwood Park, Thatcham Moor, nr Theale, Wytham; Buckinghamshire: Burnham Beeches, Farnham;
Cambridgeshire: Wicken Fen; Cheshire; Cumbria: Wray Castle; Devon: Bovey Heathfield, Lustleigh,
Torquay dist., Woolacombe; Dorset: nr Wareham; Dyfed: Llangranog; Essex: Hornchurch; Glamorgan;
Gloucestershire: Gloucester, Staunton; Greater London: Highgate, Norwood; Gwynedd: Llanddona;
Hampshire: Farley, New Forest, Southampton; Hereford and Worcester: Fawnhope; Hertfordshire:
Rothamsted, Royston; Highland; Isles of Scilly; Kent: Barnehurst, Eynsford; Lancashire: Ainsdale; Norfolk:
Norwich, Foulden; Northamptonshire: Spratton; North Yorkshire: Malham Tarn; Oxfordshire: Headington,
Oxford, Mapledurham, Summertown; Salop; Caer Caradoc; Somerset: Winsford; Suffolk: Minnesmere,
Santon Downham, Wangford; Surrey: Ashtead, Coulsdon, East Sheen; Effingham, Esher Common, Horsley,
Kew, Oxshott, Richmond; Tayside: Ben Lawers (paralectotype of Lygocerus bicolor Kieffer); Western Isles:
Lewis, Stornoway Castle; [?] Thornhill (paralectotype of Conostigmus dubiosus Kieffer). Greece: Kifisia; Mt.
Parnis. Greenland: Noret, nr Mesters Vig. Iceland: Akureyri; Dyrholaey; Eyjafjall, Skogasandur; Seljaland;
Vestmannaeyjar. Ireland: Co. Dublin, Bohernabreena; Leitrim; Sligo, Trawallua; Wicklow, Blessington,
Tonelagee Mt. Israel: Bet Dagen. Italy: Cortina, D'Ampezzo. Morocco: Beni Mellal. Lebanon: Beirut.
Netherlands: Helmond; Tiel; Urk; Wageningen. Portugal: Lisbon; Spain: Marbella, Malaga. Sweden: Jmt
Fors; Skane, Lund; Skane, Stensoffa; Sulnasker. Yugoslavia: Istria.
FURTHER DISTRIBUTION. Austria (Hellen, 1966: 13); Belgium (Dessart, 1972a: 59); Czechoslovakia (Stary,
1977: 2); Denmark (Henriksen, 1918: 140); Finland (Hellen, 1966: 13); Norway (Hellen, 1966: 13);
U.S.S.R. (Ivanova-Kazas, 1955: 151). Asia: ? India (Mani, 1941: 27). South America: Chile (Dessart, 1975:
268). Australasia: Australia (Dessart, 1975: 267).
BRITISH SPECIES OF DENDROCERUS 269
Dendrocerus bifoveatus (Kieffer)
(Figs 38, 55)
Lygocerus bifoveatus Kieffer, 1907: 55. Holotype 9> FRANCE: Amiens, ex Aphis on Cirsium arvense
(Carpentier) (lost).
Lygocerus sordidipes Kieffer, 1907: 63. Lectotype 9> GREAT BRITAIN (BMNH), designated by Masner
(1965: 21) [examined]. [Synonymized by Dessart, 1972a: 88.]
Dendrocerus bifoveatus (Kieffer) Dessart, 1972a: 88.
FEMALE. Scape relatively long, usually longer than All, AIII and AIV together, nearly as long as head.
Flagellum thin, AIII 3-4 times as long as broad. Only AI and usually AXI longer than AIII. Segments AV
to AX approximately equal in length, segments AVI to AX about equal in breadth. All clearly broader than
AIII. Pubescence short but slightly longer on the proximal than on distal segments. Ventrally, flagellar
segments flat with single central groove. Antenna weakly papillate; dark brown except distal end of All
which is often lighter in colour. Old specimens can fade to light brown.
Head smooth or weakly alutaceous; pubescence short but dense on gena, long on clypeus; dark brown,
mandibles brown with red teeth, palps clear brown; transverse but not strongly so, broader than thorax. Eyes
variable in size but never large. Frontal dent shallow or absent, depression behind each torulus shallow.
Depression in front of median ocellus distinct and sometimes deep. Depressions near lateral ocelli almost
absent. Interantennal carina weakly developed, usually almost complete over central area. Ocellar triangle
broad, ocelli almost in line. Horizontal groove behind ocelli absent. Preoccipital crescent present but
rounded, not limited by ridge, not reaching eyes or ocelli. Vertical furrow indistinct.
Thorax dark brown or black, weakly alutaceous. Pubescence not long except for region of dense hairs on
mesepimeron and side of propodeum. Mesonotum slightly narrowed anteriorly. Anterior corners very sharp
due to deep impression of notaulices in this region. Notaulices complete, usually sharply angled but can be
more smoothly angled in some specimens, not meeting posteriorly. Lateral secondary furrows short; pair of
anterior central furrows also present and far more evident on each side of median furrow. Median furrow
distinct. Scutellum with raised central area strongly arched. Scutellum up to twice as long as broad.
Propodeum weakly rugose or smooth between carinae. Propodeal fovea present, transverse propodeal carina
strongly developed. Pronotum alutaceous, only bottom arm of Y-shaped furrow obvious. Mesepisternum
almost completely smooth, shiny and only with peripheral pubescence; strong furrow present from near top
of anterior mesepisternal boundary to episternal scrobe. Legs mostly dark brown; joints light brown in some
specimens.
Forewing slightly and evenly light brown, sometimes lighter in distal region. Fringe long. Pterostigma long
and thin (Fig. 38), over twice as long as broad. Radius as long as or little longer than pterostigma and only
moderately curved.
Gaster 1-5-1-8 times longer than broad. Gastral collar well developed (but rather short in lectotype of D.
sordidipes), with tuft of long hairs on each side. When visible gastrocoeli are lighter brown than rest of gaster.
Gaster smooth, densely pubescent ventrally, in distal third.
A small species about 1 • 5 mm long.
MALE. As for female except scape short, over four times longer than broad and longer than All plus AIII.
Flagellar segments weakly serrate, nearly cylindrical (Fig. 55); elongate, AIII three times as long as broad.
Only AI and usually AXI longer than AIII.
REMARKS. The holotype of D. bifoveatus is lost; it is therefore unfortunate that the name
bifoveatus was chosen by the first reviser in preference to sordidipes, the lectotype of which is extant.
The antennae of males are unusually weakly serrated for a species of Dendrocerus but the
ocellar triangle and other characters, and the biology, confirm the placement of bifoveatus in this
genus. The antennae of males, the elongate pterostigma and the smooth mesepisternum distinguish
D. bifoveatus but it is difficult to assess its affinities.
BIOLOGY. Stary (1977: 2) recorded D. bifoveatus from Paraschizaphis scirpi / Aphidius urticae /
Typha angustifolia and P. scirpi / Diaeretiella rapae I Typha angustifolia, from moorland. In the
BMNH collection there are seven specimens with host records: two from Acyrthosiphon malvae on
Achillea millefolium, four from A. auctus and one from Rhopalosiphonius calthae. Kieffer (1907:
55) recorded D. bifoveatus from 'Aphis noir' on Cirsum arvense.
Kieffer's black aphid may have been an Aphis species of thefabae group, viz. Brachycaudus
cardui (adults black) or Uroleucon cirsii (dark red). The A. malvae listed above had wandered from
270 N. D. M. FERGUSSON
its normal host plants (Potentilla crantzii or Geum sp.). A. auctus has a north European coastal dis-
tribution, R. calthae is specific to Caltha palustris, and Typha angustifolia grows in shallow water,
therefore a wet marsh or moor habitat is indicated for D. bifoveatus. This habitat preference
explains the large number of records from Iceland (the climatic and soil conditions result in a pre-
ponderance of wet moorland habitat). D. bifoveatus has been collected mostly in July and August.
MATERIAL EXAMINED
Lygocerus sordidipes, lectotype 9> Great Britain: 'Bonar [= Bonar Bridge, Todcaster, Yorkshire (Binnie)],
483' (BMNH).
9 c?, 11 9. Great Britain. Cheshire: Rostherne; North Yorkshire: Malham Tarn (Hincks) (MM,
Manchester); Western Isles: Lewis, L. na Gainmheich. Iceland: Akureyri; Hornafjordur, Dynjandi (ZI,
Lund); Hornafjordur, Thveit (ZI, Lund); Modruvellir, Eyjabjordur; Vestmannaeyjar, Alfsey (ZI, Lund);
Vestmannaeyjar, Sudurey (ZI, Lund).
FURTHER DISTRIBUTION. France (Kieffer, 1907: 55); Czechoslovakia (Stary, 1977: 2); Sweden (Dessart,
1972a:94).
Dendrocerus carpenteri (Curtis)
(Figs 1, 14, 16, 21-24, 28, 40, 51, 57)
Ceraphron carpenteri Curtis, 1829a: 249. Syntype(s), GREAT BRITAIN: ex aphids (Carpenter) (lost). Neotype
$, BELGIUM (IRSNB, Brussels), designated by Dessart (1972a: 1 19) [examined].
Ceraphron crispus Curtis, 1829a: 249. Nomen nudum.
Ceraphron elegans Curtis, 1829a: 249. Nomen nudum.
Megaspilus carpenteri (Curtis) Stephens, 1829: 401.
Calliceras carpenteri (Curtis) Westwood, 1840: 77.
[Ceraphron fuscipes Nees; Ratzeburg, 1852: 180. Misidentification.]
Ceraphron hyalinatus Thomson, 1858: 291. Lectotype $, SWEDEN (ZI, Lund), designated by Dessart
(1972a: 11 9) [examined]. [Synonymized by Dessart, 1970: 143.]
[Ceraphron stigma Nees; Thomson, 1858: 290. Misidentification.]
Lygocerus carpenteri (Curtis) Marshall, 1868: 158.
Coryne carpenteri (Curtis) Buckton, 1876: pi. 7.
Megaspilus niger Howard, 1890: 247. Lectotype 9, U.S.A.: Michigan, Selkirk, vii. 1889 (USNM,
Washington), designated by Masner & Muesebeck (1968: 113). [Synonymized by Dessart, 19726: 240.]
Lygocerus carpentieri: Marshall, 1896: 569. [Incorrect subsequent spelling of carpenteri Curtis.]
Trichosteresis proximo Kieffer, 1907: 32. Lectotype 9, GREAT BRITAIN: Bugbroak (Marshall) (TM,
Budapest), designated by Dessart (19746: 413). [Synonymized by Dessart, 19746: 413.]
Trichosteresis punctatipennis Kieffer, 1907: 33. Holotype 9, GREAT BRITAIN (BMNH) [examined]. Syn. n.
Lygocerus rufiventris Kieffer, 1907: 45. Lectotype 9, GREAT BRITAIN (BMNH), designated by Masner (1965:
21) [examined]. [Secondary homonym ofAtritomus rufiventris Ashmead, 1887. [Synonymized by Dessart,
1970: 143.]
Lygocerus campestris Kieffer, 1907: 48. Syntype(s), FRANCE: Amiens (Carpentier) (lost). [Synonymized by
Dessart, 1972a: 107.]
Lygocerus aphidivorus Kieffer, 1907: 49-50. Lectotype 9, GREAT BRITAIN (BMNH), designated by Masner
(1965: 19) [examined]. [Synonymized by Dessart, 1970: 143.]
Lygocerus aphidivorus var. inconspicuus Kieffer, 1907: 50. Syntype(s), FRANCE: Amiens (Carpentier) (lost).
[Synonymized by Dessart, 1972a: 107.]
Lygocerus testaceimanus Kieffer, 1907: 51. Syntype(s), FRANCE (? lost). [Synonymized by Dessart, 1970:
143.]
Lygocerus aphidum Kieffer, 1907: 52. Syntype(s), FRANCE: Amiens, ex Aphis on Medicago sativa
(Carpentier) (lost). [Secondary homonym of Macros tigma aphidum Rondani, 1877.] [Synonymized by
Dessart, 1972a: 107.]
Lygocerus giraudi Kieffer, 1907: 52-53. Syntype(s), AUSTRIA, FRANCE (lost). [Synonymized by Dessart,
1970: 143.]
Lygocerus cameroni Kieffer, 1907: 56-57. Lectotype 9, GREAT BRITAIN (BMNH), designated by Masner
(1965: 20) [examined]. [Synonymized by Dessart, 1970: 143.]
BRITISH SPECIES OF DENDROCERUS 271
Lygocerus thomsoni Kieffer, 1907: 57-58. Lectotype 9, SWEDEN (NR, Stockholm), designated by Dessart
(1972a: 119) [examined]. [Replacement name for Ceraphron stigma Nees; Thomson, 1858: 290.]
[Synonymized by Dessart, 1970: 143.]
Lygocerus inquilinus Kieffer, 1917: 349. Holotype $, GERMANY (WEST) (NHM, Maastricht) [examined].
[Synonymized by Dessart, 1970: 143.]
Trichosteresis carpenteri (Curtis) Morley, 1929: 55.
Lygocerus ambianus Dessart, 1965a: 158. [Replacement name for Lygocerus aphidum Kieffer, 1907.]
[Synonymized by Dessart, 1972a: 107.]
Dendrocerus britannicus Dessart, 1966: 12. [Replacement name for Lygocerus rufiventris Kieffer, 1907.]
[Synonymized by Dessart, 1970: 143.]
Dendrocerus tischbeini Dessart, 1966: 12. Holotype <$, GERMANY (EAST) (IP, Eberswalde). [Replacement
name for Ceraphron fuscipes Nees; Ratzeburg, 1852: 180.] [Synonymized by Dessart, 1972a: 107.]
Dendrocerus carpenteri (Curtis) Dessart, 1970: 142.
FEMALE. Scape short but longer than All, AIII and AIV together. AH and AIII approximately equal in
length. Rest of flagellar segments short (L/B about 1-1-1-6), except AXI which is usually twice as long as
broad (Fig. 57). Flagellar segments flattened ventrally. Antenna black but distal end of All sometimes
lighter; weakly papillate, with short dense pubescence.
Head alutaceous, with short pubescence except on or near clypeus; black, or brown in faded specimens,
mandibles brown with red teeth, palps colourless; transverse, frons broad. Depressions behind toruli large,
separated by distinct hump, those near lateral ocelli small, that in front of median ocellus larger, deep and in
some specimens prolonged anteriorly as faint short frontal line. Interantennal carina absent. Ocellar triangle
broad, the ocelli almost in line. Preoccipital crescent well developed, anteriorly limited by distinct ridge
behind ocelli but not reaching eyes. Vertical furrow present.
Thorax alutaceous, black (brown in faded specimens) with short, quite dense pubescence. Mesonotum
broad, almost quadrate. Notaulices complete, strongly angled and often only slightly convergent posteriorly.
Four secondary furrows often visible. Scutellum broad and raised but flattened centrally. Propodeum
medially very short, fovea not strongly developed. Posterior propodeal carina strongly developed, especially
laterally. Y-shaped furrow on pronotum deeply impressed. Legs mostly brown-black but anterior tibiae
usually lighter, as are articulations.
Wings clear, pterostigma nearly semicircular but distal edge almost straight. Radius strongly curved (Fig.
40), often continuing distally without pigmentation.
Gaster black or brown, mostly smooth. Gastral collar with strong carinae. Gastrocoeli not always visible
but when present broad and lighter brown than rest of gaster.
MALE. As for female except scape short, slightly longer than All and AIII together. Basal flagellar segments
ventrally slightly concave; strongly serrate (Fig. 51), becoming less serrate distally, segments X and XI
cylindrical. AIII almost twice as long as broad. Antenna brown. Flagellar pubescence long, at least as long as
breadth of segment. Preoccipital crescent more developed and transverse than in female, eye slightly more
rounded. Parameres rather narrow.
REMARKS. D. carpenteri is often found in large numbers and has been recorded from most parts of
the world. Stary (1977: 6) found that D. carpenteri accounted for 92-6% of the 1865 specimens of
Dendrocerus he studied. D. carpenteri is easily separated from most other species of Dendrocerus
but D. liebscheri is very similar; very small specimens of D. carpenteri look like D. aphidum. The
lectotype of Lygocerus koebelei is a specimen of D. aphidum but three of the four paralectotypes are
faded specimens of D. carpenteri. In the Haliday collection there is a labelled specimen of
Ceraphron crispus (listed by Curtis, 1829a: 249 but not described). It is badly damaged (only the
antenna, legs and wings are left) but identifiable as a specimen of D. carpenteri. There is further
material of C. crispus in the NM, Victoria and, according to Curtis's note book (in NM, Victoria),
these were collected in 1850; one specimen is D. aphidum, the rest are D. carpenteri. In both the
BMNH and the NM, Victoria, there is a specimen of C. elegans (listed by Curtis, 1829a: 249 but
not described); both specimens are conspecific with D. carpenteri. Also in the BMNH is the
holotype of Trichosteresis punctatipennis. The specimen is from Clober, as stated by Cameron
(1907: 161), and bears the species name in Cameron's handwriting. The antennae are brown, not
testaceous, but otherwise the specimen fits the published description. It is conspecific with D.
carpenteri.
In the Curtis collection there is a male and female of D. carpenteri mounted on one card. Written
on the reverse of the card is 'Bred 28.6 off Roses Gl. Wool'. Curtis did not mention the type-
272 N. D. M. FERGUSSON
locality but he did say that the species was collected by T. Carpenter. If Gl. Wool is the collector's
name then the male and female have no type-status; if it is a locality then they could be syntypes.
G. E. J. Nixon designated (unpublished) the male as 'type'; he thought the last word on the label
was Wood. I am convinced that the word in question is Wool and I am unable to ascertain the
type-status of these specimens. There is a pin in the Haliday collection which bears a label
'carpenterC but the specimen is missing from the card and there is no other evidence, apart from the
label, to indicate the type-status of the missing specimen. The microscope slides of larval stages (see
'Biology') of carpenteri prepared by Mrs Brindley (Miss M. D. Haviland) were donated to the
BMNH. There are 36 slides, mostly serial sections of third or fourth instar larvae. There are two
whole mounts labelled Lygocerus cameroni, and I can confirm that they are D. carpenteri
(= cameroni). This is particularly important as Haviland's work forms the basis of our knowledge
of larval development etc. in the genus.
BIOLOGY. As the records for D. carpenteri are too extensive to register in full, the following are
alphabetical lists of the hosts and parasites.
Hosts (Aphidoidea): Acyrthosiphon caraganae, A. cyparissiae, A. malvae, A. pisum, A. sp.,
Amphorophora ampullata, A. idaei, Aphis citricola, A. craccae, A. craccivora, A. fabae, A.
farinosa, A. frangulae, A. gossypii, A. marthae, A. nerii, A. pomi, A. sambuci, A. schneideri, A.
urticata, A. viburni, A. sp., Aulacorthum ibotum, A. magnoliae, A. solani, A. syringae, A. sp.,
Betulaphis quadrituberculata, Brachycaudus cardui, B. helichrysi, B. sp., Brevicoryne brassicae,
Callipterinella calliptera, Capitophorus hippophaes, C. sp., Cavariella aegopodii, C. araliae, C.
salicicola, C. theobaldi, C. sp., Ceruraphis eriophori, Chaetosiphon tetrarhodus, Cinara juniperi,
C. lands, C. piceae, C. pinea, C. pini, C. sp., Coloradoa sp., Corylobium avellanae, Cryptomyzus
sp., Diur aphis frequens, D. noxius, Drepanosiphum platanoidis, D. sp., Dysaphis plantaginea, D.
sorbi, Elatobium abietinum, E. sp., Euceraphis punctipennis, E. sp., Eucallipterus tiliae,
Hamamelistes shirakabae, Hayhurstia atriplicis, Hyadaphis foeniculi, H. sp., Hyalopterus pruni,
H. sp., Hyperomyzus carduellinus, H. lactucae, H. niger, Indomegoura indica, Lipaphis erysimi,
Liosomaphis berberidis, Macrosiphoniella absinthii, M. artemisiae, M. grandicauda, M. linariae,
M, major, M. millefolii, M. oblonga, M. tanacetaria, M. sp., Macrosiphum avenae, M. euphorbiae,
M. gei, M. hellebori, M. rosae, M. rudbeckiarum, M. sorbi, M. sp., Melanaphis bambusae,
Metopeurum fuscoviride, Metopolophium albidum, M. dirhodium, M. festucae, M. poacearum, M.
sp., Microlophium carnosum, M. evansi, M. sp., Mindarus japonicus, Myzaphis rosarum,
Myzocallis coryli, Myzus boehmeriae, M. cerasi, M. ligustri, M. persicae, M. varians, M. sp.,
Nasonovia nigra, Neophyllaphis totarae, Ovatus sp., Parachaitophorus spiraeae, Periphyllus
acericola, P. aceris, P. calif orniensis, P. hirticornis, P. lyropictus, P. sp., Phorodon humuli,
Pleotrichophorus glandulosus, Prociphilus fraxini, P. xylostei, Protolachnus agilis, P. thunbergii,
Pseudobrevicoryne erysimi, Pterocallis sp., Pterocomma salicis, Rhopalomyzus lonicerae,
Rhopalosiphoninus deutzifoliae, R. sp., Rhopalosiphum insertum, R. nymphaeae, R. padi,
Schizaphis agrostis, S. longicaudata, Schizolachnus pineri, S. pineti, S. sp., Sipha maydis, S. sp.,
Sitobion akebiae, S. avenae, S. fragariae, S. ibarae, S. sp., Staticobium limonii, Symydobius
oblongus, Tinocallis platani, Titanosiphon artemisiae, Toxoptera odinae, Toxopterella
drepanosiphoides, Tuberocephalus sp., Tuberculoides sp., Tuberolachnus salignus, Uroleucon
aeneus, U. ambrosiae, U. campanulae, U. cephalonoli, U. chondrillae, U. cichorii, U. cirsii, U.
gobonis, U. hypochoeridis, U.jaceae, U.picridis, U. rudbeckiae, U. sonchi, U. sp.
Primary parasites (Aphidiidae): Aphidius absinthii, A. aquilus, A. areolatus, A. cingulatus, A.
equiseticola, A. ervi, A. funebris, A. gifuensis, A. hieraciorum, A. hortensis, A. longipetiolus, A.
matricariae, A. phalangomyzi, A. picipes, A. polygonaphis, A. rosae, A. salicis, A. schimitscheki, A.
setiger, A. smithi, A. tanacetarius, A. urticae, A. uzbekistanicus, A. sp., Areopraon nipponicum,
Binodoxys acalephae, B. angelicae, B. brevicornis, B. centaurae, Calaphidius watanabei,
Diaeretiella rapae, Diaeretus leucopterus, Dyscritulus planiceps, D. sp., Ephedrus californicus, E.
lacertosus, E. minor, E. nacheri, E. niger, E. persicae, E. plagiator, E. salicicola, E. sp., Lipolexis
gracilis, Lysaphidus erysimi, L. matsuyamensis, L. pleotrichophori, Lysiphlebia japonica,
Lysiphlebus ambiguus, L. arvicola, L. dissolutus, L. fabarum, L. fritzmuelleri, L. testaceipes,
Metaphidius aterrimus, Monoctonus angustivalvus, M. cerasi, M.pseudoplatani, Pauesia abietis, P.
akamatsucola, P. laricis, P. piceaecollis, P. pini, P. salignae, P. unilachni, P. sp., Praon abjectum,
BRITISH SPECIES OF DENDROCERUS 273
P. absinthii, P. barbatum, P. bicolor, P. capitophori, P. dorsale, P.flavinode, P. gallicum, P. necans,
P. pequodorum, P. silvestre, P. valuer -e, P. sp., Trioxys betulae, T. cirsii, T. curvicaudus, T.
euceraphis, T.falcatus, T.pallidus, T. pannonicus, T.parauctus.
The following genera of Cynipoidea and Chalcidoidea have also been recorded as forming part of
the host complex of D. carpenteri: Cynipoidea — Alloxysta, Phaenoglyphis; Chalcidoidea -
Aphelinus, Aphidencyrtus, Asaphes, Coruna, Cyrtogaster, Encyrtus, Pachyneuron.
D. carpenteri has also been associated with the ants Formica rufa and Lasius fuliginosus
(Hymenoptera: Formicidae) (Kieffer, 1907: 349) and with Meromyza saltatrix (Diptera:
Chloropidae) (Meyer, 1929: 241). In the BMNH there is a specimen of D. carpenteri bred from a
Rose Bedeguar gall.
The records of hosts, parasites etc. listed above are taken from material examined, and from the
following references: Bankowska (1975: 312); Buckton (1876: 1 17); Campbell & Mackauer (1975:
419); Chua (1977: 125); Cumber & Eyles (1961: 404); Curtis (1829: 249; 1860: 293); Dean (1974:
414); Dessart (1972a: 1 15-1 19); Dessart (1975: 267); Dill (1937: 233); Dours (1873: 113); Dunn
(1949: 105); Ferriere & Voukassovitch (1928: 28); Fulmek (1968: 900-909); Garbarczyk (1977:
447-448); Gatenby (1919: 405); Gourlay (1930: 339); Hafez (1961: 530); Haviland (1920a: 293;
19206: 104); Howard (1890: 247); Kieffer (1907: 48-53); Lowe (1968: 821); Maneval (1940: 96);
Marshall (1872: 124; 1896: 569-572); Meyer (1929: 234-244); Muesebeck et al. (1951: 669);
Paetzold & Vater (1967: 83); Pass & Parr (1971: 1153); Petherbridge & Mellor (1936: 336);
Ratzeburg (1852: 180); Rothschild (1963: 126); Skripchinskii (1930: 281); Spencer (1926:
142-150); Stary (1977: 2-3); Sullivan & Van Den Bosch (1971: 389); Takada (1973: 9-11;
1976: 237); Thuneberg (1959: 23); Tillyard (1926: 283); Todd (1957: 720); Treherne (1916: 186);
Valentine (1975: 61); Voukassovitch (1925: 655; 1928: 54); Walden(1926: 295-298); Ward (1934:
144); Westwood (1840: 170); Wimshurst (1925: 93); Yasumatsu et al. (1946: 12; 1947: 113).
The aphids listed above are from most of the systematic groups of the Aphidoidea. The only
families not represented are the Adelgidae and Phylloxeridae. The most frequently recorded genera
are Microlophium (261 records, mostly of M. carnosuni), Acyrthosiphon (154 records),
Macrosiphum (100), Aphis (78, many A.pomi records), Metopolophium (70, mostly M. dirhodum\
Sitobion (66), Periphyllus (60), Schizolachnus (56), Uroleucon (44), Macrosiphoniella (34),
Rhopalosiphum (32) and Myzus (28). All the major genera of Aphidiidae are included in the above
records. The most frequently recorded genera are: Ephedrus (281 records), Praon (258), Aphidius
(220) and Pauesia (74). Both forms of pupation (inside and outside the mummy) are represented.
D. carpenteri has been recorded from a wide range of plants, including coniferous and deciduous
trees, herbs, shrubs, cultivated plants and crops, fruit-trees, and plants of wasteland, hedgerows and
damp habitats.
The host information for D. carpenteri includes records that are typical elements of the following
faunistic complexes: Eurasian steppe — Macrosiphoniella spp. / Aphidius absinthii I Artemisia
sp.; Titanosiphon artemisiae / Trioxys pannonicus /-; Microlophium spp. / Aphidius ervi /-;
Uroleucon spp. / Praon dorsale; Acyrthosiphon caraganae / Aphidius ervi: Pseudobrevicoryne
erysimi/ Lysaphidius erysimi /—. European and Far Eastern deciduous forest — Drepanosiphum
platanoidis I Dyscritulus planiceps - Acer pseudoplatanus; Periphyllus californiensis I Aphidius
areolatus /Acer sp. Coniferous forest — Cinaria lands I Pauesia pini I Larix leptolepis. Holarctic
forest tundra — Pterocomma salicis I Aphidius cingulatus I Salix sp.; Symydobius oblongus I
Trioxys betulae. The most frequently represented of the faunistic complexes is the Eurasian steppe.
Several gall-forming aphids are represented which are commonly parasitized by Ephedrus persicae
or E. plagiator. Some wax-producing aphids and their parasites are also listed, e.g. Brevicoryne
brassicae I Diaretiella rapae /-. Stary (1977: 6) records D. carpenteri mostly from parks, fields,
meadows and waste places, and found that it was numerically the most abundant Dendrocerus
species, but in some habitats other species were dominant, e.g. D. bifoveolatus in moor/pond
habitats.
D. carpenteri, which is the only species of Dendrocerus to have had its biology well studied, is a
hyperparasite of mature larvae or young pupae of Aphidiidae (or on secondary parasites); the
Aphidiidae are in turn parasites of Aphidoidea. The aphidiid larva consumes the aphid and pupates
inside the empty aphid skin. As the female of D. carpenteri only oviposits into aphids containing a
274 N. D. M. FERGUSSON
mature parasite larva or pupa, D. carpenteri cannot be a direct parasite of the aphid. According to
Haviland (19206: 103) and Spencer (1926: 142) D. carpenteri can parasitize other larvae of D.
carpenteri, and Haviland could not find any other hymenopteron that parasitized D. carpenteri
larvae. The complexity of parasitism and hyperparasitism can be seen from one example recorded
by Haviland (19206: 103): a Microlophium carnosum (= Macrosiphum urticae) was parasitized by
Aphidius ervi which had been hyperparasitized by an undetermined species of Chalcid which in
turn had been partially devoured by a second hyperparasite which had been hyperparasitized by D.
carpenteri.
The development of D. carpenteri occurs outside the body of the host. The egg is elliptical,
becoming more spherical as development proceeds. It measures 0-25 mm by 0-1 mm, is white,
semi-translucent, and has a minute teat-like protuberance at one end. (Spencer, 1926: 143 records a
protuberance at both ends.) The chorion surface has numerous longitudinal striae. According to
Haviland (19206: 106) and Spencer (1926: 143) the egg hatches in about 24 hours, but Rothschild
(1963: 127) states that at 25 °C the first instar larva hatches after two days.
The first instar larva grows to 0-45 by 0-22 mm. It is white, transparent, with thirteen body
segments (Fig. 21) (there is some variation in the number of body segments given by authors for the
instars). The distinct head has two small tactile papillae and the mouth has two slender mandibles.
When newly hatched only two pairs of spiracles are open, one near the posterior margin of the first
body segment and the other on the anterior part of the fourth body segment, but the spiracles of the
third and fifth segments open shortly afterwards. The midgut is closed, it is large and its contents
are yellow. The first instar lasts 20-24 hours. The larva can progress by a kind of peristaltic
movement but it probably does not move around as the host skins normally have only one
puncture mark. If the host is in the pupal stage the D. carpenteri larva is usually found feeding on
the posterior part of the abdomen where the integument is still soft. As the egg is deposited on the
third or fourth segment of the aphidiid the D. carpenteri larva must move to this position.
The second stage larva (Fig. 22) measures 0-7 by 0-35 mm and differs from the first instar chiefly
by the more developed tracheal system and the greater development of the anterior part of the body
so that the head appears to be divided from the thorax by a constriction. The stigmatic trunks of
segments six, seven and eight are visible but the corresponding spiracles are closed. This stage lasts
24-36 hours. The hyperparasite seems to feed by suction and the skin of the aphidiid (otherwise
uninjured) is gradually emptied of its contents.
The third stage larva (Fig. 23) is globose and about 1-00 mm long by 0-75 mm broad. In this
stage the head is bent round ventrally, the papillae on the head disappear and the spiracles on body
segments six, seven and eight are open. The mandibles work vigorously and are probably used to
macerate the host organs. This stage lasts for about 40 hours.
The fourth stage larva (Fig. 24) measures 1 -0 by 0-83 mm when fully fed (1 -4 mm long according
to Rothschild, 1963: 127). It becomes creamy white and ingests the last of its host. It differs
considerably from the third stage larva. The first four body segments are greatly developed, the
small head is bent round to the ventral side and is almost hidden by the large prothorax. The last
abdominal segment bears dorsally a conical caudal appendage of unknown function. The body
segments bear a row of chitinous papillae or spines. The salivary glands and malpighian tubes have
considerable lumina. The midgut, which fills the greater part of the body cavity, contains a mass of
fluid which is churned to and fro by muscular contractions. At this stage there are seven pairs of
spiracles, the first pair near the posterior margin of segment one, the second on the posterior side of
the third segment and the rest on the following five segments. The eighth spiracle does not become
functional until metamorphosis. Rudimentary stigmatic trunks can sometimes be seen on the ninth
and tenth segments. The fourth instar lasts from one and a half to three days.
D. carpenteri pupates within the cocoon spun by the aphidiid. Just before metamorphosis the
midgut opens to the hindgut and the contents are voided. The pupa is yellow at first but gradually
darkens until it is black. According to Spencer (1926: 145) this stage lasts six to eight days but
Haviland (19206: 121) states that the period of pupation is from 14-16 days and Rothschild
(1963: 127) gives the mean duration of the pupal stage (at 25 °C) as eight days. The imago gnaws a
hole in the aphid mummy and emerges. The hole has irregular margins and thus differs from the
emergence hole of aphidiids.
BRITISH SPECIES OF DENDROCERUS 275
According to Spencer (1926: 145) a generation can occupy 11-16 days, the average being 12
days, while Haviland (19206: 106-121) indicates 22-25 days and Rothschild (1963: 127) 18 days.
These variations in development times are presumably due to different temperatures. Campbell &
Mackauer (1975: 422) studied the influence of temperature on development rate. It took 9-4 days at
25-9°C and 42-69 days at 10-3°C for the parasite to develop from egg to adult. The parasite
developed more slowly than its host, at the same temperature. This may ensure that the parasites
do not appear too early in the season and that a continued minimum host supply is available.
Copulation may last from 20 seconds to 15 minutes and can take place a few hours after
emergence. Rothschild (1963: 126) noted that the males mate repeatedly with the same female but
Haviland (19206: 105) stated that they only paired once.
According to Rothschild (1963: 126) oviposition may last from 90 seconds to 15 minutes. When
a suitable host is found the female of D. carpenteri energetically examines it with her antennae. The
female stands on or behind the aphid so that the ovipositor, when inserted, punctures the aphidiid
larva (lying, bent head to tail in the cocoon) at its centre of curvature. Usually only one egg is laid.
Spencer (1929: 143) observed that two eggs were sometimes laid but only one adult emerged.
Rothschild (1963: 126) recorded one instance where an unmated D. carpenteri female appeared to
oviposit in the normal way but no eggs were laid. Haviland (19206: 106) estimated the total
number of eggs laid by a single female to be less than 25.
Adults live up to 10 days and Takada (1976: 383) gives the possible number of generations in a
year for D. carpenteri as between two and six. D. carpenteri is active from May to November.
Kieffer (1907: 50) records D. carpenteri (as aphidivorus var. inconspicuus) over-wintering in moss,
and Takada (1976: 383) and Dunn (1949: 106) indicate that it over-winters within the aphid
mummy or can sometimes survive the winter as adults.
Haviland (19206: 105) recorded that the sex ratio was 2:3 (males: females). Dunn (1949: 105)
gave the sex ratio as 2:1. The work of Garbarczyk (1977:445-454) showed a preponderance of
females (61.9%). Spencer (1926: 145) induced virgin females to oviposit; the resultant adults were all
males.
Although D. carpenteri is a very common species the numbers of aphids are so great that the
overall rate of parasitism is quite low. Hafez (1961: 471) and Chua (1977: 138) both give a figure
of under 2%, Bankowska (1975: 312) recorded 3.4%, Low (1968: 825) recorded a 5.5% parasitism
rate and Dean (1974: 415) gave 3.2% and 12% for two different hosts. Because it is last in the
chain of parasites D. carpenteri is often the least common member of the aphid parasite complex.
Haviland (19206: 125) recorded it as comprising 32—44% of the parasites of Aphidiidae. Takada
(1976: 252), studying the parasites of aphids on potatoes and cruciferous crops, found that it
emerged in a higher proportion from parasitized aphids on the upper leaves, than from those on the
lower leaves.
MATERIAL EXAMINED
Dendrocerus carpenteri (Curtis), neotype $> Belgium: La Hulpe, 2.vi.66 (IRSNB, Brussels). Ceraphron
hyalinatus Thomson, lectotype <$, Sweden: 'Ld [Lund], 9.viii.6' (NR, Stockholm). Trichosteresis
punctatipennis Kieffer, holotype 9, Great Britain: 'Clober [Scotland, Central] 100' (Cameron) (BMNH).
Lygocerus rufiventris Kieffer, lectotype 9> Great Britain: '439' (Cameron) (BMNH). Lygocems aphidivorus
Kieffer, lectotype 9, Great Britain: '365 Kelvin [Scotland, Central Kelvinside] 10/6' (Cameron) (BMNH).
Lygocerus cameroni Kieffer, lectotype 9» Great Britain: 'York [= Yorkshire] 384 Cameron coll.' (BMNH).
Lygocerus thomsoni Kieffer, lectotype 9» Sweden: 'Ld [Lund] 8/6' (NR, Stockholm). Lygocerus inquilinus
Kieffer, holotype tf, Germany (West): 'Kol [or Rol] 4. 16/9/1 1 Iserlohn b F. rufa' (NHM, Maastricht).
738 cJ, 1029 9. Belgium: Moresnet. China: Foochow. Germany (West): Kehl; Westfallen, Plellenburg.
Great Britain. Bedfordshire; Berkshire; Buckinghamshire; Cambridgeshire; Cheshire; Cumbria; Devon;
Dorset; Durham; Essex; Hampshire; Hertfordshire; Highland; Isles of Scilly; Kent; Greater London;
Lancashire; Greater Manchester; West Midlands; Norfolk; Northamptonshire; Northumberland; North
Yorkshire; Oxfordshire; Powys; Somerset; South Glamorgan; Staffordshire; Strathclyde; Suffolk; Surrey;
Sussex (East); Sussex (West); West Yorkshire. India: Kashmir. Ireland: Antrim; Down; Dublin; Kildare;
Leitrim; Londonderry; Sligo; Wicklow. Italy: Dolomites, Seils am Schlern; Sicily, Villafranca; South Tirol.
Japan: Atami; Hokkaido, Sapporro; Honshu, Kure. Netherlands: Arnhem; Bennekom; Brandrvyk;
Dwingelo; Ede; Epen; Groningen; Gronsveld; Hazerswonde; Helmond; Hemmen; Kootwick; Langbrock;
Leersum; Lienden; Lunteren; Mheer; Oenkerk; Rhenen; Rijswizk; Schiermonnikoog; Tiel; Wageningen;
276 N. D. M. FERGUSSON
Wessem. Sweden: Skane, Rolsberga, Rovurekulan; S., Lund; Sm. Vaxjo S. Areda. Switzerland: Zurich.
U.S.S.R.: Vilnius, Verzuva.
FURTHER DISTRIBUTION. Europe: Austria (Dessart, 1972a: 120); Czechoslovakia (Stary, 1977: 2). North &
South America: Canada (Dessart, 19726: 239); Chile (Dessart, 1975: 267). Australasia: Australia (Dessart,
19726: 240).
Dendrocerus dubiosus (Kieffer) sp. rev.
(Figs 20, 46, 49, 58)
Ceraphron longicornis Thomson, 1858: 293. Holotype 9> SWEDEN: 'Malmo 7/56' (lost). [Primary homonym
of Ceraphron longicornis Boheman, 1832: 337.]
Lygocerus pallipes Kirchner, 1867: 193.Nomennudum.
Conostigmus dubiosus Kieffer, 1907: 152. Lectotype 9. GREAT BRITAIN (BMNH), designated by Masner
(1965: 15) [examined].
Lygocerus flavipes Kieffer, 1907: 54. Holotype 9> ITALY (MCSN, Genoa) [examined]. [Secondary homonym
of Dendrocerus flavipes Kieffer, 1907: 23.] Syn. n.
Lygocerus claripennis Kieffer, 1907: 43. Holotype $, ITALY: Genoa, 1894 (Solari) (MCSN, Genoa). Syn. n.
Lygocerus rectangularis Kieffer, 1907: 44-45. Lectotype $, GREAT BRITAIN (BMNH), designated by
Masner (1965: 20) [examined]. Syn. n.
Conostigmus alpestris Kieffer, 1907: 143-144. Holotype 9, ITALY (MCSN, Genoa) [examined]. Syn. n.
Dendrocerus navaensis Dessart, 1966: 12. [Replacement name for Lygocerus flavipes Kieffer, 1907: 54.]
Syn. n.
Dendrocerus dubiosus (Kieffer) Dessart, 1972a: 125.
Lygocerus flavus Hellen, 1966: 14. Holotype 9. FINLAND (ZMU, Helsinki) [examined]. Syn. n.
[Dendrocerus bicolor sensu auct, nee Kieffer. Misidentifications.j
FEMALE. Scape very long, 5.7-7.0 times longer than broad and much longer than head (except for one
brachypterous specimen in which it is 5.4 times longer than broad). AIII elongated, up to 4 times as long as
broad, clearly longer than All (Fig. 58). Segments AVI to AX 1.7-2.0 times longer than broad, AXI longer.
Ventrally Flagellar segments flat with central ridge. Antenna usually longer than body; weakly papillate;
pubescence short, dense; brown except distal end of All, which is slightly lighter, and base of scape, which is
yellow.
Head mostly alutaceous, frons often almost smooth; black or dark brown, mandibles yellow with red teeth,
palps colourless; pubescence short. Eyes large, almost as long as distance between torulus and preoccipital
crescent. Eye pubescence short but distinct. Interantennal carina distinctly developed and curved over central
swelling between toruli. A shallow depression present behind and inwards from each torulus. Frontal dent
shallow but can be conspicuous. Depressions near lateral ocelli shallow, that in front of median ocellus
usually large and conspicuous. Ocellar triangle broad. Lateral ocelli joined by distinct posterior horizontal
groove. Preoccipital crescent present but not strongly developed, not usually reaching eyes.
Thorax black or dark brown, mostly alutaceous or coriaceous, pubescence short, sometimes quite dense.
Anterior corners of mesonotum sharply angled. Notaulices complete, clearly curved outwards towards
anterior corners of mesonotum. Four secondary furrows often visible. Axillae mostly flat, only downcurved
in outer region. Metanotal and propodeal furrows strongly foveolate. Propodeal fovea small but prominent.
Y-shaped furrow on side of pronotum distinct. Legs entirely yellow, basal half of hind coxa very rarely dark.
Forewing with light but distinct brown area under pterostigma, veins often indicated as clear lines in brown
area. Pterostigma semiovoid, radius distinctly curved, about 1.3 times longer than pterostigma. Wing
pubescence normal.
Gaster smooth; 1.5-2.0 times longer than broad. Gastral collar long, well developed, with several parallel
carinae. Gastrocoeli visible in dark specimens as two lighter brown areas.
MALE. As for female except scape short (L/B 5.0 or more), nearly equal to length of head, nearly as long as
AH, AIII and AIV together. Basal flagellar segments elongate (AIII usually clearly the longest), clearly
asymmetrical (Fig. 49), pubescence longer than breadth of segments. Basal third of scape usually yellow,
flagellum brown or black but All distinctly lighter. Legs generally darker than in female, often mostly yellow.
REMARKS. As Lygocerus bicolor Kieffer, 1907 (but not bicolor of authors) is now synonymized
under D. aphidum (see p. 266) a new replacement name is required for Ceraphron longicornis
Thomson. Of the synonyms now assigned to this species the names published by Kieffer in 1907 have
precedence; Conostigmus dubiosus has been chosen because the lectotype is female and easily
accessible.
BRITISH SPECIES OF DENDROCERUS 211
Dessart (I912a: 143) considered D.flavus and D. dubiosus (= bicolor auct., nee Kieffer) to be
closely related but distinct; however D.flavus is just a brachypterous form of D. dubiosus. Many of
the stated differences are within the range of variation of D. dubiosus and the rest are consistent
with wing reduction. Both known specimens are female. The second specimen was collected at the
same time as a normal D. dubiosus, and I have seen a specimen of D. dubiosus with three-quarter-
length wings.
Females of D. dubiosus are separable from D. aphidum females by the long scape and long AIII
(compared with the length of All) of D. dubiosus. In large specimens of D. aphidum the scape is
sometimes long but the length of AIII is always about equal to AIL The legs of D. dubiosus females
are entirely yellow with the exception of two dark specimens which have a dark basal half to the
hind coxa. The legs of D. aphidum females always have some dark coloration, at the very least the
basal half of the hind coxa is dark. Small specimens of D. aphidum can often have entirely dark legs.
The males are more difficult to separate but generally the scape is shorter and the leg colour darker
in D. aphidum than in D. dubiosus. The males of D. aphidum and D. dubiosus could conceivably be
placed in the same species but the range of variation in the females is far too great for synonymy.
Previously I named a series of specimens from Akureyi, Iceland as D. bicolor (i.e. bicolor sensu
auct. = dubiosus) and Couchman (1977: 123) published this determination. After comparison with
types and other material, I have now decided that they are large specimens of/), aphidum.
Dessart has labelled a specimen from the Cameron collection as '? paralectotype of bicolor'; it is
not conspecific with the type-material, it is a specimen of D. dubiosus and has no type-status.
After D. carpenteri, D. dubiosus is one of the more common species ofDendrocerus.
BIOLOGY. Acyrthosiphon pisum I Aphidius ervi / Medicago saliva 17.vi.1967 and 18.vii.1969
(Stary, 1972: 2); A. pisum I Aphidius sp. / Medicago sativa (Boness, 1958: 322); 1 tf, A. sp. /
Praon dorsale I Trifolium pratense (Takada, 1973: 8); Amphorophora rubi /-/ Rubus sp.
21.vi.1914; 1 tf, 69, Aphidiid sp. / Praon sp. / Orixa japonica (Takada, 1973: 8); Aphis
craccivora I Binodoxys acalephae I Onobrychis sativa l.vi.1960 (Stary, 1977: 2); A. craccivora I
Lysiphlebus fabarum I Onobrychis sativa l.vi.1960 (Stary, 1977: 2)A.fabae /Binodoxys angelicae
1 Papaver sp. 18.vi.1959 (Stary, 1977: 2); 29, Aulacorthum magnoliae / Praon volucre I
Sambucus siebaldiana (Takada, 1973: 8); 1 9» -4- magnoliae I Praon volucre I- ll.v.1963; 1 <$,
2 9, A. syringae I Praon volucre I Syringa reticulata (Takada, 1973: 8); Cryptosiphum artemisiae
/ Ephedrus nacheri / Artemisia vulgaris 9.vi.l958 (Stary, 1977: 2); Galiobium langei /Aphidius
metricariae I Galium sp. 26.vi.1974 (Stary, 1977: 2); 1^, 19, Hyalopteroides humilis /-
Dactylis glomerata 28. v. 19 72; Macrosiphum rosae /Aphidius rosae I Rosa sp. 5.vi.l961 (Stary,
1977: 2); 1 9> Microlophium carnosum I Praon sp. / Armoracia rusticana 31.viii.1934; 1 9>
Rhopalosiphoninus deutzifoliae / Ephedrus sp. / Deutzia crenata (Takada, 1973: 8); 1 9«
Schizolachnus sp. / Pauesia unilachni I Pinus densiflora (Takada, 1973: 8); Sitobion avenae I
Aphidius ervi / Triticum vulgare 2.vii.l975 (Stary, 1977: 2); S. avenae I Aphidius uzbekistanicus I
Triticum vulgare vii. 1975 (Stary, 1977: 2); 1 tf, S. avenae 1-1 Hordeum sp. 26.vi.1972; S. equiseti
I Aphidius equiseticola I Equisetum silvaticum (Stary, 1977: 2); S. equiseti I Ephedrus plagiator /
Equisetum silvaticum (Stary, 1977: 2); 5". equiseti I Monoctonus caricis I Equisetum silvaticum
(Stary, 1977: 2); Therioaphis trifolii I Praon exoletum / Medicago sativa 23.V.1960 and
30.viii. 1967 (Stary, 1977: 2); Uroleucon aeneus I 'Ephedrus niger / Carduus acanthoides 28.vi.1961
(Takada, 1973: 8); -/-/ Medicago sativa (Eankov/ska, 1975: 312).
Dessart (1972a: 85) recorded D. dubiosus from Aphidius picipes, Diaeretiella rapae, Ephedrus
plagiator and Praon volucre. Donisthorpe (1927: 106) recorded it in association with Lasius
brunneus (Hymenoptera: Formicidae); as there is no other evidence of ant association it is probable
that this was just a chance encounter.
D. dubiosus is recorded from a considerable range of hosts. There are several records
characteristic of the Eurasian steppe complex, i.e. Acyrthosiphon spp. /Aphidius ervi, Aphis spp. /
Lysiphlebus fabarum, Aphis spp. / Binodoxys angelicae, Cryptosiphum artemisiae I Ephedrus
nacheri I Artemisia vulgaris and Galiobium langei / Aphidius matricariae. There are also records
characteristic of the European deciduous forest complex, i.e. Macrosiphum spp. / Aphidius rosae,
Sitobion spp. / Monoctonus caricis and Acyrthosiphon I Praon volucre. The Schizolachnus sp. /
278 N. D. M. FERGUSSON
Pauesia unilachni I Pinus record is the only one from the coniferous forest complex. The aphids listed
above are from diverse families (Lachnidae, Callaphididae and Aphididae). The primary parasites are
all Aphidiidae, and all the common genera are represented. D. dubiosus is recorded from a wide
diversity of plants, including trees, shrubs and herbs, and on three occasions from Equisetum (E-
quisetineae); it has been collected over much of the year and appears to be most abundant in July and
August. Takada (1973:15) listed D. dubiosus (as bicolor) under an 'immediate' type of habitat
(shrubs, orchards and gardens), but the above records indicate a much wider habitat range, from
forest to fields and wasteland.
MATERIAL EXAMINED
Conostigmus dubiosus Kieffer, lectotype 9> Great Britain: '356, Clober [Scotland, Central] l.vi.'
(Cameron). Lygocerus flavipes Kieffer, holotype 9, Italy: 'Nava 108, viii. 1902' (Solari) (MCSN, Genoa).
Lygocerus rectangularis Kieffer, lectotype $, Great Britain: '372, Dumfries' [Scotland, Borders] (Cameron).
Conostigmus alpestris Kieffer, holotype 9, Italy: 'Nava, viii 902' [1902] (Solari) (MCSN, Genoa).
Lygocerus flavus Hellen, holotype 9, Finland: Lemland, 1 l.viii.1953 (Hellen) (ZMU, Helsinki).
105 (5\ 99 9- Austria: Semmering region, Ruchenau dist. France: Pyrenees-Orientales, nr Vernet-les-
Bains. Great Britain: Bedfordshire: Dunstable Down, Steppingly; Berkshire: Streatley, Silwood Pk,
Thatcham Moor, Windsor, Wytham; Buckinghamshire: Burnham Beeches, Slough; Cambridgeshire; Devon:
Heathfield; Dorset: nr Sto borough; Dumfries and Galloway: Dumfries; Grampian: Elgin; Greater London:
Kew, New Maiden, Richmond; Gwent: Newton; Hampshire: Brockenhurst, Lyndhurst, New Forest;
Hertfordshire: Rothamsted; Leicestershire: Charnwood Forest; Lothian: Edinburgh: Mid Glamorgan:
Kenfig Pool; Northamptonshire: Spratton; Oxfordshire: Headington; Shetland: Fetlar, Unst, Haroldswick;
Somerset: Brompton Regis; Strathclyde: Aran, Catacol, Beinn Bhreac, Cawbber; Suffolk: Barton Mills;
Surrey: Ashtead, Bookham, Clandon Downs, Farnham, Oxshott; Western Isles: Lewis, Greeta, Ireland:
Dublin: Bohernabreena, Grand Canal, Mantfield, Phoenix Park; Kildare: R. Canal, Landenstown,
Trawallna; Longford: Castle Forbes; Wicklow: Ballyknockan, Dowry. Italy: Sicily, Villafranca. Japan:
Honshu, Tokyo. Sweden: Brakne, Hoby; B., Sjoard; Halleberg; Orebfo, Adby; Skane, Hoor dist.; S., Kivik; S.,
Loderup; S., Ring: S., Stensoffa.
FURTHER DISTRIBUTION. Algeria (Dessart, 1979: 33); Belgium (Dessart, 1972a: 87); Czechoslovakia
(Stary, 1977: 2); Faroes (Kryger & Schmiedeknecht, 1938: 76); Germany (West) (Dessart, 1972a: 87);
Iceland (Dessart, 1972a: 87); Norway (Dessart, 1972a: 87); Switzerland (Dessart, 1972a: 87).
Dendrocerus flavipes Kieffer
(Figs 30, 35, 62)
Dendrocerus flavipes Kieffer, 1907: 22-23. Holotype 9, ITALY (MCSN, Genoa) [examined].
Dendrocerus fuscipes Kieffer, 1907: 23-24, pi. 2, fig. 9. Holotype <?, ITALY (MCSN, Genoa) [examined].
[Synonymized by Dessart, 1978: 181.]
A tritomellus flavipes (Kieffer) Kieffer, 1914: 143-144.
FEMALE. Scape elongate, longer than maximum breadth of thorax or antennal segments II, III and IV
together. Scape brown but often lighter ventrally at base. AIII long, clearly longer than pedicel or AIV. Distal
flagellar segments approximately as broad as scape. Flagellar segments with median groove, flat ventrally.
Flagellum brown, papillate, with short pubescence.
Head alutaceous; brown but clypeus usually yellow, palps and mandibles yellow or colourless, teeth of
mandibles red-brown. Eyes large, 0.65-0.75 times as long as lateral length of head. Pubescence short.
Interantennal carina absent or indicated only by dark line. Frons alutaceous. Ocellar triangle broad, lateral
ocelli slightly closer to eyes than to median ocellus. Preoccipital crescent present, with a fine vertical furrow
centrally. Head rounded, not strongly transverse; pubescence short.
Thorax elongate, laterally compressed (L/B = 1-6— 1-9), clearly narrower than head; generally alutaceous,
brown to testaceous (sometimes bicoloured: brown with pronotum and mesonotum testaceous). Pronotum
always testaceous dorsally and testaceous laterally, at least in anterior region. Propleura testaceous. Anterior
margin of mesonotum clearly defined. Notaulices not reaching posterior mesonotal margin, fading out
immediately after bend although, in specimens with light coloured mesonotum, their continued path is
sometimes indicated by faint line. Secondary furrows distinct. Axillae strongly curved downwards. Scutellum
narrow, highly arched, sometimes flattened centrally Propodeal carinae united to form small median
projection. Legs pale yellow to light brown except claws and last tarsal segments which are brown.
Forewing with brown transverse band under pterostigma and radius. Pterostigma semioval, not broad.
Radius not sharply curved. Pubescence short; fringe occasionally hard to distinguish.
BRITISH SPECIES OF DENDROCERUS 279
Gastral collar colourless or light yellow, strongly contrasting with rest of gaster, which is brown; broad
with many fine carinae. Gaster long and narrow.
MALE. As for female except scape short, approximately same length as AIII, certainly shorter than All plus
AIII; broad, about 2.5 times as long as broad. Basal flagellar segments broad, strongly asymmetrical (Fig.
30), distal segments cylindrical. Antenna papillate, uniformly dark, pubescence shorter than maximum
breadth of segment, pubescence of scape very short. Eyes not elongated. Frons broad; preoccipital crescent
clearly developed, reaching ocelli and nearly reaching margin of eye. Body darker than female, black or
brown. Wings colourless. Legs brown, distal ends of femora, tibiae and tarsi sometimes lighter. Abdominal
collar dark.
REMARKS. D. flavipes is known only from females and D. fuscipes only from males. Dessart
(1972a: 153) described a 9 offuscipes which is a female of D. halidayi, as the wings are not clearly
smoked and the body coloration is uniform. The strong sexual bicoloration found in D. flavipes is
unusual in Dendrocerus but it does also occur in D. indicus and possibly some African species. D.
flavipes is similar in thoracic shape and to a lesser extent in general coloration to females of D.
halidayi, and to D. laticeps and D. laevis in having the notaulices only present anteriorly, but the
last two species are darker and have a much broader thorax. D. flavipes is here recorded for the
first time from Great Britain.
BIOLOGY. Dessart (1978: 177) recorded D. flavipes from Coniopteryx esbenpeterseni (Neuroptera).
D. flavipes has been found on Aesculus hippocastanum, Crataegus and Quercus robur, and under
Fagus sylvatica. Collected mostly in July and August.
MATERIAL EXAMINED
Dendrocerus flavipes Kieffer, holotype 9, Italy: Genoa, '75', vi.1900 (Borgioli) (MCSN, Genoa).
Dendrocerus fuscipes Kieffer, holotype $, Italy: Genoa, '105', vi.1900 (Borgioli) (MCSN, Genoa).
6 $, 10 9, Great Britain. Berkshire: Slough; Devon: Heathfield; Greater London: Kew; Hampshire:
Shawford; Hereford and Worcester: Burghill; Kent: Eynsford; Northamptonshire: Spratton; Surrey:
Ashtead, Oxshott Woods, Weybridge; Sussex: Bridge Park. Ireland: Wicklow, Dowry and Athdown.
Sweden: 6 1... [label illegible, slide No. 6911/062] (ZI, Lund). Yugoslavia: Dalmatia, Korcula.
FURTHER DISTRIBUTION. France (Dessart, 1972a: 156); Germany (Hellen, 1966: 14); Switzerland (Dessart,
1972a: 156); doubtfully (see Dessart, 1972a: 142) Finland (Hellen, 1966: 14).
Dendrocerus halidayi (Curtis)
(Figs 32, 45, 63)
Ceraphron halidayi Curtis, 1829a: 249, fig. Holotype $, IRELAND (NMI, Dublin) [examined].
Megaspilus halidayi (Curtis) Stephens, 1 829 : 40 1 .
Dendrocerus lichtensteinii Ratzeburg, 1852: 181. Syntype(s), no locality (lost). [Synonymized by Dessart,
1966:5.]
Ceraphron damicornis Foerster, 1856: 146. Nomen nudum. ['Synonymized' by Kieffer, 1907: 20.]
Ceraphron callicerus Thomson, 1858: 292. Holotype tf, SWEDEN (NR, Stockholm) [examined].
[Synonymized by Dessart, 1966: 5.]
Lygocerus halidayi (Curtis) Marshall, 1868: 158.
Lygocerus halidayii Marshall, 1873: 3. [Unjustified emendation.]
Dendrocerus halidayi (Curtis) Dessart, 1966: 5.
[Dendrocerus fuscipes Kieffer; Dessart, 1972a: 153. Misidentification.]
FEMALE. Scape (Fig. 63) long and thin (L/B 5.5-6.5), nearly as long as All, AIII, AIV and AV together.
AIII longer than AIV. All and AIII longer and less broad (L/B 2.0-3.5) than rest of flagellum, which is
relatively thick (L/B 1.2-1.5). Flagellar segments ventrally flattened. Antenna mostly coriaceous; brown,
base of scape clear brown, distal segments darker brown; pubescence short, dense and evenly distributed.
Head alutaceous pubescence short, eye pubescence very short; brown, slightly darker than thorax or base
of gaster. Mandibles brown with darker teeth, palps colourless. Head rounded, not transverse but clearly
broader (1.3—1.4 times broader) than thorax. Interantennal carina present, with horseshoe-shaped depression
linking toruli. Ocellar triangle broad and shallow, ocelli almost in line. Depression in front of median ocellus
conspicuous, lateral depressions joined by groove running behind ocelli. Preoccipital crescent present but not
strongly developed, not reaching margin of eye. Crescent with central furrow.
280 N. D. M. FERGUSSON
Thorax brown; alutaceous; with short pubescence except for tuft of long hairs on propodeum above hind
coxa; long and thin (L/B 1.5-2.0), clearly narrower than head. Notaulices distinct anteriorly before bend but
indistinct or, more often, absent posteriorly. Axillae strongly downcurved. Scutellum long, thin and highly
arched, 1.6-1.8 times longer than broad. Metanotum reduced; salient median fovea present between
metanotum and propodeum. Propodeum obliquely down-curved, shiny, with large carinae and fine alutaceous
sculpture. Legs brown, darkest on hind coxa and lightest on anterior tibia. Wings almost hyaline, pterostigma
semioval, radius thin, vannal lobe absent.
Gaster mostly smooth and brown, anterior half slightly lighter brown; long and thin (L/B 1.9-2.4), clearly
longer than thorax. Gastral collar dorsally with few short carinae, often only four, and with small tuft of long
hairs on each side.
MALE. As for female except scape short and broad (L/B 3.0-3.6). AIII and AIV short, even with All they
are shorter than scape (Fig. 32). AVI and AVII very long (L/B 6.0-9.5). Segments AIII to AVI each with
long projection, which is slightly swollen apically (Fig. 32). On AIII to AVI the projections are longer than
segments, but projection on AVII shorter than length of segment. AVII is clearly longest segment. AVIII
usually with small projection or hump. Antenna with long pubescence, especially on projections; brown but
basal half of scape sometimes lighter. Head 1.10-1.25 times as broad as thorax. Preoccipital crescent slightly
larger than in female, reaching eyes and ocelli. Notaulices continuous but sometimes faint posteriorly.
Pubescence not long but generally dense and obvious. Wings clear.
REMARKS. In his description of Ceraphron halidayi, Curtis (1829a: 249) stated that the insect was
taken on the 8th August near Holy wood (Ireland), that it was in the Haliday collection and that the
female was unknown. Dr J. P. O'Connor has searched the Haliday collection for this species and
found a specimen which closely corresponds with the original description and figure and also with
the accepted concept of halidayi. The specimen is unlabelled (except for the registration number
20.2.82) but this is quite normal in the Haliday collection. This specimen is undoubtedly the
holotype of C. halidayi; thus the neotype designation by Dessart (1972a: 161) was unnecessary.
In the Curtis collection there are two males of D. halidayi. As the style of card and label for one
specimen is almost certainly that of Haliday, G. E. J. Nixon has labelled it 'probably paratype of C.
halidayf.
The antennae of D. halidayi males are similar in form to those of D. ramicornis; the females of
D. halidayi and to a lesser extent the males have the thorax and coloration similar to those of D.
flavipes.
Although the syntype(s) of D. lichtensteinii (type-species of the genus Dendrocerus) are lost the
figure and description clearly show it to be a synonym of D. halidayi. Some authors have incorrectly
omitted the terminal '/' of the original spelling 'lichtensteinii'.
BIOLOGY. D. halidayi is a solitary, external parasite within the cocoon of Coniopterygidae
(Neuroptera).
Viggiani (1967: 173-175) bred D. halidayi from Semiadalis aleyrodiformis and the degree of
parasitism was 10% or less. He described the parasite larva as orange with a distal terminal
digitiform process.
Withycombe (1923: 590-591; 1924: 117) bred a Lygocerus sp. from Conwentzia psociformis.
Because of the close relationship of the two host genera and Withycombe's description of the larva
as orange-pink it seems likely that the Lygocerus sp. was D. halidayi. Withycombe stated that prior
to oviposition a small hole was usually torn in the host envelope by the parasite female, but this
hole was not always visible. When biting this hole the parasite may also bite or bruise the thorax of
the larva, 'probably to prevent pupation'. A single egg is laid alongside the larva, within the cocoon.
The parasite larva is fully fed by winter and pupates in early summer, the adult parasite emerging in
mid-summer of the second year. Withycombe thought the parasite was single brooded but there
was some evidence of two broods.
The neotype and neoparatypes of D. halidayi were all bred from Semiadalis aleyrodiformis
(Dessart, 1972a: 166).
Several specimens have been obtained from galls. Ratzeburg (1852: 181) obtained D.
lichtensteinii from galls of Biorhiza pallida (= Cynips terminalis} and specimens of D. halidayi
have been collected from second year galls of Biorhiza pallida on oak. As the Conwentzia and
Semiadalis species listed above are abundant on oak, it is likely that a host cocoon was present on
BRITISH SPECIES OF DENDROCERUS 281
or in the oak gall. As the second brood of C. psociformis often prefers to spin its cocoon in a
crevice, it may occasionally choose the inside of an empty gall.
D. halidayi has been collected as early as May, but from the records August appears to be the
month of maximum emergence.
MATERIAL EXAMINED
Ceraphron halidayi Curtis, holotype tf, Ireland: 20.11.82 (Haliday) [no further data] (NMI, Dublin).
Dendrocerus halidayi (Curtis), neotype tf, Italy: Portici, vi.-xi.1934 (IRSNB, Brussels). Ceraphron
callicems Thomson, holotype <$, Sweden: Dalarne (Boheman) (NR, Stockholm).
5 (5*, 6 9. Great Britain. Devon: Heathfield, Torquay dist.; Hampshire: New Forest, Denny Wood. Italy:
Portici (neoallotype and neoparatype of D. halidayi) (IRSNB, Brussels). Sweden: Kl [rest of label illegible]
(ZI, Lund); Or. 0. Mack, (neoparatype of D. halidayi) (ZI, Lund); [label illegible] (ZI, Lund); [label illegible]
A. J. [= (Jansson)] (allotype of D.fuscipes) (ZI, Lund).
FURTHER DISTRIBUTION. D. halidayi is also known from Germany (East) (Ratzeburg, 1852: 181).
Dendrocerus laevis (Ratzeburg)
(Figs 25, 36,48, 61)
Ceraphron laevis Ratzeburg, 1852: 180. Holotype 9, GERMANY (EAST) (IP, Eberswalde) [examined].
Ceraphron frontalis Thomson, 1858:293. Lectotype 9, NORWAY (NR, Stockholm), designated by Dessart
(1972a: 184) [examined]. [Synonymized by Dessart 1972a: 177.]
Megaspilus laevis (Ratzeburg) Kirchner, 1867: 191.
Atritomus coccophagus Foerster, 1878:56. Lectotype <$, GERMANY (WEST) (MNHU, Berlin), designated by
Dessart (1972c:235) [examined]. [Synonymized by Dessart 1972a: 177.]
Ceraphron levis Dalla Torre, 1898:526. [Unjustified emendation of laevis Ratzeburg. f
Atritomus levis: Kieffer, 1905:256. [Incorrect subsequent spelling of laevis Ratzeburg.]
Dendrocerus levis: Kieffer, 1907:20. [Incorrect subsequent spelling of laevis Ratzeburg.]
Atritomellus laevis (Ratzeburg) Kieffer, 1914: 142.
Atritomellus smirnoffi Ghesquiere, 1960:208, figs 1-3. Holotype <$, MOROCCO: Rabat, 8.vi.l949 (Smirnoff)
(lost). [Synonymized by Dessart, 1972a: 177.]
[Lygocerusflavipes Kieffer; Hellen, 1966: 14. Misidentification.]
Dendrocerus applanatus Dessart, 1972a:59, figs 25-30. Holotype c?, BELGIUM (IRSNB, Brussels)
[examined]. Syn. n.
Dendrocerus laevis (Ratzeburg); Dessart, 1972a: 176.
FEMALE. Scape clearly longer than combined length of All, AIII and AIV, and shorter than maximum
breadth of thorax. Pedicel at least as long as AIII, normally much longer. Flagellar segments extremely
short, AIV and AV approximately same length, segments AIV to AX only slightly longer than broad.
Segments All, AIII and AIV usually noticeably thinner than following segments. Flagellar segments flat
ventrally; black or brown. Scape usually completely dark but occasionally considerably lighter, never with
distinct light-coloured basal ring. Old specimens often uniformly light brown or yellow.
Head finely alutaceous or smooth; pubescence short; brown or black, mandible colour lighter than that of
head, mandibular teeth red or brown, palps testaceous. Head relatively broad, wider than thorax; transverse
when viewed anteriorly. Interantennal carina present but not developed. Groove directly behind lateral ocelli
almost absent. Preoccipital crescent always present and normally strongly developed, often reaching margin
of eye. Preoccipital crescent often gives apex of head, when viewed laterally, sharply angled appearance.
Thorax finely alutaceous; brown (testaceous in some faded specimens); pubescence short; not elongate
(L/B 1-1-1-4). Mesonotum very short, considerably wider than long. Anterior margin of mesonotum not
sharply angular. Median furrow present, notaulices not reaching posterior mesonotal margin, but fading out
near bend. Mesonotum sometimes flattened dorsally, especially in central region. Secondary furrows not
usually visible. Scutellum variable, often broad and shallowly arched, or narrow and almost flat; arching
occasionally more pronounced. Median propodeal projection weakly developed. Propodeum sharply sloping
laterally, posterior propodeal carina strongly developed. Leg coloration varies from dark black or brown on
proximal segments of posterior legs to light brown or yellow on distal segments of anterior legs. Tarsal
segments often slightly darker than preceding segments.
Wings clear, fringe obvious. Pterostigma often semiovoid, shape of pterostigma variable, especially in
depth. Radius shallowly curved.
282 N. D. M. FERGUSSON
Gaster broad (L/B 1-2-1-9), smooth. Gastral collar short, mid lateral length less than 0-04 mm.
Gastrocoeli often indistinct. Ovipositor relatively long.
Overall body length under 1-5 mm, often much smaller.
MALE. As for female except scape short, about equal to length of eye, longer than combined length of All
and AHI. Flagellar segments short (e.g., L/B for AIII 1-5-2-5). Segments AIV, AV, AVI and AVII approx-
imately equal in length, AIII slightly larger. Flagellar pubescence very long, often as much as twice breadth
of segment. Flagellum serrate, sometimes strongly serrate. Parameres short, thick, with blunt termination.
REMARKS. D. laevis is here recorded for the first time from Great Britain.
Dessart (1972a:59) described D. applanatus which, apart from a flattened thorax, was 'virtually
deprived of other salient characters'; he separated it from D. laevis by the flattened thorax, the
shape of the male genitalia and the degree of development of the preoccipital crescent. Such small
differences are of no more than infraspecific value. Flattening of the thorax can occur in D. laevis
(also to a much lesser extent in D. laticeps) but this character and the degree of development of the
preoccipital crescent are quite variable. The holotypes of D. laevis and D. applanatus are merely
opposite extremes of one species.
The type-locality of Ceraphron frontalis, Dovre fjell, is in Norway, but many authors give the
locality as being in Sweden.
D. laevis is closely related to D. laticeps; both are broad species with short notaulices. The
elongate antennae and greater size of D. laticeps distinguish it from D. laevis but small specimens
are difficult to recognize.
D. laevis has been only rarely collected but it is probably more common than the records
suggest. It is one of the smallest species in the genus.
BIOLOGY. The original host record published by Ratzeburg (1852:180) of Cecidomyia salicina
(= probably Rhabdophaga rosarid) is unlikely and the true host, a coccid or an aphid, may have
been overlooked.
D. laevis has been recorded from both the Coccoidea and the Aphidoidea.
Coccoidea records. Foerster (1878: 56) recorded Atritomus coccophagus from a coccid on Acer.
Ghesquiere (1960:206) gave the hosts of Atritomellus smirnqffi from Morocco as Eriococcus
araucariae (Eriococcidae), Planococcus citri and P. longispinus [=adonidum] (Pseudococcidae)
via the following primary parasites, Leptomastidea abnormis [—- aurantiaca] Tetracnemus
diversicornis and Microterys silvestrii [?] (Chalcidoidea, Encyrtidae). A. smirnqffi was most
common in the autumn and its life-cycle was 30-35 days. In the BMNH are 1 $ and 4 9 of D.
laevis bred from Heterococcus nudus (Pseudococcidae) on Holcus sp. Viggiani (1970:58) has
recorded D. laevis as a parasite ofPseudococcusfragilis.
Aphidoidea records. Ivanova-Kazas (1955:151) recorded D. frontalis from Hyalopterus pruni
(=H. arundinis) on Spanish reed. Evenhuis (1964:229; 1966:39; 1968:113) recorded D. frontalis
from Metopolophium albidum, Dysaphis angelicae, D. plantaginea, Aphis pomi, Rhopalosiphum
insertum and Sitobion avenae. In the case of Aphis pomi the primary parasite was Binodoxys
angelicae. The hyperparasites generally appeared late in the season.
Dessart (1972a: 149, 185) questioned all the above records from aphids and considered D. laevis
to be solely a hyperparasite of coccids. I have, however, seen specimens of D. laevis bred from
Aphis pomi I Binodoxys angelicae / apple and Tuberculoides annulatus I Trioxys pallidus /
Quercus robur, and Takada (1973:7) has recorded it from Shivaphis celti I Trioxys shivaphis I
Celtis sp.
The Shivaphis and Tuberculoides records are respectively typical of the Far Eastern and
European deciduous forest faunistic complexes. All the aphids listed above are recorded from
deciduous trees except Hyalopterus pruni on Spanish reed.
The aphids are Aphididae and Callaphididae and include gall-forming species. Apart from
Binodoxys listed above, Ephedrus persicae is a likely host for D. laevis, as it is a parasite of A.
pomi, H. pruni and Dysaphis spp.
D. laevis has been collected from May to September and the data labels of the material examined
indicate that emergence is at a maximum in July and August.
BRITISH SPECIES OF DENDROCERUS 283
MATERIAL EXAMINED
Ceraphron laevis Ratzeburg, holotype 9> Germany (East): '147d laevis Rtz' (IP, Eberswalde). Ceraphron
frontalis Thomson, lectotype 9» Norway: 'alp' (Boheman) (NR, Stockholm). Atritomus coccophagus
Foerster, lectotype <$, Germany (West): 'Atritomus coccophagus m, Cocc., Aceris' [no further data]
(MNHU, Berlin). Dendrocerus applanatus Dessart, holotype tf, Belgium: 'Knokhe, Fauchage prairie,
12.vu.1967' (Dessart) (IRSNB, Belgium).
11 J1, 24 9- Germany (West): same data as lectotype (MNHU, Berlin) (paralectotypes of Atritomus
coccophagus Foerster). Great Britain. Berkshire: Silwood Pk; Cheshire: Abbots Moss; Cumbria: Skirwith
(MM, Manchester); Greater London: Acton, New Maiden, Wimbledon Common; Hampshire: Heckfield
Heath. Ireland: Wicklow, Devil's Glen. Italy: Duino, Triest. Netherlands: Bennekom; Tiel. Sweden: Skane,
Rostanga; Norke, Markkarret (ZI, Lund), [label unreadable] (ZI, Lund) (paratypes of Dendrocerus
applanatus Dessart). Switzerland: Geneva (IRSNB, Brussels) (allotype of Ceraphron laevis Ratzeburg).
FURTHER DISTRIBUTION. Finland (Hellen, 1966:15); France (Dessart, 1972a:185); Japan (Takada,
1973:6); Morocco (Ghesquiere, 1960:208); U.S.S.R. (Ivanova-Kazas, 1955: 150).
Dendrocerus laticeps (Hedicke)
(Figs 15, 26, 34, 47, 60)
Atritomellus laticeps Hedicke, 1929:60-61, figs 1, 2. Holotype tf, GERMANY (EAST) (lost) (MNHU, Berlin).
Neotype tf, GERMANY (EAST), designated by Dessart (1972c: 235) (MNHU, Berlin) [examined].
Lygocerus laticeps (Hedicke) Hellen, 1966: 15.
Dendrocerus laticeps (Hedicke) Dessart, 1972a: 186.
FEMALE. Scape approximately equal to combined lengths of All, AIII and AIV, not as long as
maximum breadth of thorax and broader than rest of antenna. AIII longer than All (in female paratype
nearly equal). Pedicel long. AIII long and thin, about 3 times as long as broad, distal flagellar segments
broader, less elongate, with flat area ventrally. AIV sometimes slightly asymmetrical. Scape never completely
dark, normally (when viewed dorsally) with distinct light-coloured basal ring, but this sometimes indistinct.
Flagellum black or brown. Pubescence rather short.
Head broad; alutaceous; pubescence short; usually black, mouthparts brown or yellow, mandibular teeth
red. Eyes usually large. Interantennal carina distinct and sinuate. Ocellar triangle broad and short, lateral
ocelli well separated from eyes. Depression anterior to median ocellus large, lateral depressions often
extended to form groove running behind ocelli. Preoccipital crescent present but weakly developed, clearly
separated from margin of eye. Occipital carina strongly developed.
Thorax usually black, sometimes brown; broad (L/B 1-1-1-4). Notaulices very distinct but present only
anteriorly, disappearing immediately after bend. Median furrow distinct. Axillae not strongly curved
downwards. Scutellum broad, only strongly arched in small specimens. Anterior metanotal margin with
conspicuous furrow passing under apex of scutellum. Propodeum posteriorly with prominent, sparsely
crenulate furrow. Median propodeal projection weakly developed. Lateral thoracic segments bordered by
conspicuous foveolate or crenulate furrows. Legs usually yellow, sometimes darkened proximally, especially
on outside of hindlegs.
Wings clear, pubescence long, fringe long and obvious. Pterostigma noticeably semioval, long and thin.
Radius evenly curved.
Gaster broad (L/B 1-2-1-8), mostly smooth. Gastral collar large with prominent carinae. Gastrocoeli
visible as light or alutaceous patches.
MALE. As for female except interantennal carina and preoccipital crescent sometimes less distinct. Axillary
depression less obvious. Wings slightly and legs considerably darker. Scape always longer than length of eye;
alutaceous; pubescence short. Flagellum elongate (AIII long, L/B 2-4-3-0); weakly serrate (AIII almost
evenly cylindrical); papillate, especially distally; pubescence long, longer than breadth of segment, except on
terminal segment. Parameres long, thin and distally rounded.
REMARKS. Hedicke described this species from two males and two females, retained the holotype
male and the allotype in his own collection, and donated the two other paratypes to the MNHU,
Berlin. Dessart (1972c: 235), unable to find the holotype or allotype, designated a male paratype as
a neotype.
The paratypes do not agree entirely with the original description and figures as regards the length
of the scape, the termination of the radius and the shape of AIII in the female; these discrepancies
284 N. D. M. FERGUSSON
are nevertheless within the limits of variation of D. laticeps. The paratypes are lighter and slightly
smaller than is normal for the species.
D. laticeps is closely related to D. laevis; both are broad species with short notaulices. The
elongate antennae and greater size of D. laticeps distinguishes it from D. laevis, but small
specimens can be difficult to identify. D. laticeps is unusually variable in body length, ranging
between 1-2 mm. It is the most frequently collected species which has incomplete notaulices. D.
laticeps is here recorded as British for the first time.
BIOLOGY. The original record: 2 9» 2 <3*, Stenopelmus rufinasus [Coleoptera] /-/ Azolla
filliculoides, 25.x.-l.xi.l927 published by Hedicke (1929:59), has been strongly doubted by
Dessart (1972a: 192; 1972c:235). This single coleopterous record seems unlikely, and aphids were
probably present but overlooked.
2 <$, 3 9» Acyrthosiphon nipponicus / Aphidius amamioshimensis I Paederia scandens mairei
(Takada, 1973:6); 1 $, Aphidiid sp. /Aphidius sp. /Beta vulgaris rapa (Takada, 1973:6); Aphis
gossypii I Lysiphlebia japonica I- (Takada, 1976:237); 2 9> Aphis pomi / Binodoxys angelicae I
Mains sylvestris mitis, 25.vii.1962 and 28.viii.1963; Brevicoryne brassicae / Diaeretiella rapae /-
Takada, 1976:237); 19> Brachycaudus helichrysi I— I Chrysanthemum parthenium, 9.vi.l915; 1
$, Capitophorus sp. Ephedrus persicae I Elaeagnus umbellata (Takada, 1973:6); Cavariella
sp. / Aphidius salicis I Selinum carvifolia, 21.vi.1961 (Stary, 1977:3); C. aegopodii /-/ Daucus sp.,
ll.vi.1970 (Dessart, 1972c:236); C. theobaldi I Aphidius salicis / Heracleum sp., 16.vii.1974
(Stary, 1977:3); Chromaphis juglandicola / Trioxys pallidus / Juglans regia, 2.viii.l958 (Stary,
1977: 3); Coloradoa sp. / Lysaphidus arvensis I Artemisia vulgaris, 28.vii. 1960 (Stary, 1977: 3); 1
9, Lipaphis ?erysimi /-/-, 22-28.X.1971; L. erysimi I Diaeretiella rapae I Raphanus sativus
(Takada, 1973:6); 1 9> Macrosiphoniella grandicauda /Aphidius longipetiolus /Artemisia sp.
(Takada, 1973:6); 1 ^, 2 9» Macrosiphum akebiae / Aphidius picipes / Oryza sativa (Takada,
1973:6), M. euphorbiae /-/ Solanum tuberosum, 12.vi.1970 (Dessart, 1972c:236); 1 $, 1 9,
Microlophium carnosum I— I Urtica dioica, 25. vii. 1962— 28.viii. 1963; 3 $, 8 9> My zus persicae /
Aphidius gifuensis I Raphanus sativus (Takada, 1973:6); 4 <$, 3 9? M. persicae I Diaeretiella
rapae I Raphanus sativus (Takada, 1973:6); Periphyllus hirticornis I Trioxys falcatus / Acer
campestre, 25.vii.70 (Stary, 1977:3); 1 <$, 19, Rhopalosiphum padi I Aphidiid sp. / Triticum
aestivum (Takada, 1973:6); R.padi [Macrosiphum padi error!] / Aphelinus chaonia / Prunuspadus,
26.ix.1968 (Dessart, 1972c:236); R. padi / Aphelinus daucicola I Poa annua, 19.x. 1970 (Dessart,
1 972c: 236); R.padi I Aphidius sp. / Poa annua, 1 9.x. 1 970 (Dessart, 1 972c: 236); R . padi I Ephedrus
plagiator / Poa annua, 19.X.1970 (Dessart, 1972c:236); R. padi I Trioxys auctus I Poa annua
11.x. 1970 (Dessart, 1972c:236); 1 9, Staegeriella necopinata /-/ Galium verum, 7.vii.l915;
Sitobion avenae /-/- vi.-vii.1976; 1^, Toxoptera odinae / Ephedrus plagiator / Pittosporum tobira
(Takada, 1973:6); 9 tf, 14 9, -/ Binodoxys sinensis /-, 29.viii.1974; 1 9, -/-/ Crataegus sp.,
3.ix.l977; 6 9,-/-/ Vicia sp., l-9.viii.1974.
The host records Aphis pomi, Chromaphis juglandicola and Periphyllus, parasitized by Trioxys
or Binodoxys, are typical of the European deciduous forest faunistic complex; the two last-
mentioned aphids are the only non-Aphididae recorded. There are no conifer aphids in the host list
but the Aphididae are widely represented. The Far Eastern deciduous forest faunistic complex is
represented by the Myzus persicae I Aphidius gifuensis record. The Coloradoa sp. record is typical
of the Eurasian steppe faunistic complex, although Artemisia vulgaris is also found in hedgerows
and waste ground, etc. Several leaf curling aphids are hyperparasitized by D. laticeps, e.g.
Rhopalosiphum padi / Ephedrus plagiator. The two Aphelinus (Chalcidoidea) records are from late
in the year.
The plant records include trees, shrubs, herbs, crops and grasses, thus extending the field and
intermediate habitat distribution recorded by Takada (1973:15). One specimen of D. laticeps was
collected in a salt marsh.
From the material examined, July to September appears to be the time of maximum emergence.
MATERIAL EXAMINED
Atritomellus laticeps Hedicke, neotype $, Germany (East): 'Schonebecke [Elbe], aus Stenopelm,
rufinasus l.xi. 1927' (ManzecK) (MNHU, Berlin).
BRITISH SPECIES OF DENDROCERUS 285
25 c?, 569. China: Foochow. Channel Islands: Jersey, St. Aubins. Germany (East): same data as neotype
(MNHU, Berlin) (paratype of Atritomellus laticeps Hedicke). Great Britain: Bedfordshire: Steppingley
(Chambers coll.), White Lane, Odell (Chambers coll.); Berkshire: Hamm Wood; Devon: Newton Abbot;
Greater London: Battersea Fields (UM, Oxford), Coulsdon, Kew; Hertfordshire: Rothamsted (Rothamsted
Exp. Stn.); Royston; Humberside: Spurn (MM, Manchester); Isles of Scilly: Bryhr; Norfolk: Foulden
Common, North Wooton; Northamptonshire: Spratton; Oxfordshire: Oxford (UM, Oxford), Shotover Hill
(UM, Oxford); Suffolk: Barton Mills, Santon Downham; Surrey: Dorking, Esher Common; Sussex (West):
Littlehampton; no further data (UM, Oxford). Japan: Honshu, Niigata; Kyoto, Shimogamo. Netherlands:
De Loete, Hazerswoude (IPO, Wageningen); Tiel.
FURTHER DISTRIBUTION. Belgium (Dessart, 1972a: 192); Czechoslovakia (Stary, 1977:3); Finland (Hellen,
1966: 15); Germany (West) (Dessart, 1972a: 192); Sweden (Dessart, 1972a: 192).
Dendrocerus liebscheri Dessart
(Figs 17, 27, 42, 52, 56)
Ceraphron tenuicornis Thomson, 1858:291. Holotype 9, SWEDEN (ZI, Lund) [examined]. [Primary
homonym of Ceraphron tenuicornis Boheman, 1832:332.]
Lygocerus tenuicornis (Thomson) Dalla Torre, 1898:535.
Dendrocerus liebscheri Dessart, \912a: 193. [Replacement name for tenuicornis Thomson, 1858: 291.]
FEMALE. Scape long (L/B 5-0-7-0); longer than All, AIII and AIV together; longer than head. AH nearly as
long as AIII. AIII thin and relatively long (L/B 2-8-3-2). Rest of flagellar segments stout, relatively elongate,
about 1-8 times longer than broad. Flagellum flat ventrally. Pubescence short and dense. Antenna papillate,
completely black or at most with slightly light area at apex of All.
Head alutaceous; pubescence short; black, mandibles black with red teeth, palps clear brown. Depression
in front of median ocellus small but deep, depressions by lateral ocelli very small. Interantennal carina absent
or sometimes very faintly indicated near toruli. Ocellar triangle broad, short, ocelli almost in line. Vertical
furrow present from median ocellus to occipital carina. Preoccipital crescent present, not strongly developed,
not reaching eyes.
Thorax approximately 1-5 times longer than broad, slightly flat in profile. Thorax black, alutaceous.
Notaulices complete, sharply angled outwards anteriorly, convergent posteriorly, meeting or almost meeting
median furrow at scutal suture (Fig. 17).
Median furrow not as deeply marked as notaulices. Secondary furrows often present, relatively long.
Axillae strongly downcurved distally. Scutellum about 1-2 times longer than broad, central raised area
strongly arched. Propodeum slightly more coarsely sculptured than rest of thorax. Central propodeal fovea
normally not strongly developed. Pronotum with Y-shaped furrow but posterior arm of Y faint. Legs mostly
dark, joints and tibiae yellow-brown. Wings clear, fringe present, pterostigma large, semiovoid. Radius
distinctly curved (Fig. 42).
Gaster about 1-7 times longer than broad, brown or black. Collar prominent, with short carinae. Rest of
gaster smooth, becoming alutaceous distally. The gastrocoeli are visible.
MALE. As for female except scape short, very stout, 3-4 times longer than broad, not as long as All, AIII
and AIV together. Segments AIII to AX similar in length but varying in breadth (e.g. L/B for AIV 1-7 and
L/B for AX often over 2-5). Basal flagellar segments serrate (Fig. 52). Pubescence of scape short, pubescence
of basal flagellar segments much longer than breadth of segment. Antenna sparsely papillate.
Preoccipital crescent larger, almost reaching eyes. Parameres apically broad and truncate (Fig. 27).
REMARKS. This species is similar in appearance to D. carpenteri but much less common. D.
liebscheri is here recorded as British for the first time.
BIOLOGY. Cinara lands I Pauesia pint I Larix sp., 15.vii.1963 (Stary, 1977:3); C. nuda [C. pini
or C. eschlerichi ?] P. sp. / Pinus sylvestris (Dessart, 1972a:200); C. pectinatae I P. infulata I
Abies alba, 23. vi. 1960 and 22. vi. 19 74 (Stary, 1977:3); C. pilicornis / P. sp. / Picea excelsa
(Dessart, 1972a:200); 1 tf, 1 9, C. pineal P. sp. I Pinus sylvestris, 18.viii.1974; 3 tf, 9 9, C.pinea
/-/ Pinus sylvestris viii.1977 [5 from 1 mummy]; 1$, 49, C.pinea / -I Pinus sylvestris 16.vi.1978;
2 tf, 3 9, C. juniperi /-/ Juniperus communis 14.vii. and l.viii.1971; C. sp. / P. Pini I Pinus
sylvestris 22.vii.1959 and 13.vii.1974 (Stary, 1977:3); C. sp. / P. pini I Pinus sylvestris (Dessart,
1972a: 200); C. sp. / P. sp. / Pinus sylvestris (Dessart, 1972a: 200); Protolachnus agilis I Diaeretus
leucopterus I Pinus sylvestris, 22.vii.1959 and 13.vii.1974 (Stary, 1977:3); Protolachnus agilis I
286 N. D. M. FERGUSSON
Praon bicolor / Pinus sylvestris 22.vii.1959 and 13.vii.1974 (Stary, 1977:3); Protolachnus agilis I
Praon bicolor / Pinus sylvestris (Dessart, 19720:200); Schizolachnus pineti I Pauesia unilachni I
Pinus sylvestris, 13.vii.1974 (Stary, 1977:3); Schizolachnus pineti / Pauesia unilachni / Pinus
sylvestris (Dessart, 1972a:200).
Specimens of D. liebscheri have been collected in June, July and August and the data indicate a
peak in emergence in late July or early August. The above host records are typical of the East
Eurasian coniferous forest faunistic complex and to a lesser extent of the West Eurasian coniferous
complex. D. liebscheri has been recorded only from Aphidiinae parasitizing Lachnid aphids
(Lachnidae: Cinarinae) on conifers (Pinaceae or Cupressaceae) except for two records from Praon
bicolor (Ephedrinae). Although Praon bicolor is normally a deciduous forest species (Stary,
1970: 314) it is, when parasitic on Protolachnus species, a secondary element in the West Eurasian
coniferous complex. With the exception of Praon all the Aphidiidae listed are specialized parasites
of the Lachnidae.
D. liebscheri is the only known gregarious aphid hyperparasite in Dendrocerus. Stary (1977:7)
recorded as many as eight specimens from a single aphid mummy and I have seen five specimens
from one Cinara pinea mummy. Both sexes were present in these series and the specimens emerged
from the same exit hole in the mummy.
D. liebscheri is a specialized hyperparasite of coniferous forest aphids and although it has only
been rarely collected it is probably common where it occurs.
Several of the Aphididae/ Aphidiidae relationships recorded above as hosts for D. liebscheri are
known to occur in the coniferous forests of France, West Germany, Italy and Japan, thus D.
liebscheri may also be found in these countries.
MATERIAL EXAMINED
Ceraphron tenuicornis Thomson, holotype 9> Sweden: 'Rshm [= Rysjoholm] 20. vi. 1857' (ZI, Lund).
7 (51, 179. Germany (East): Mirow. Great Britain: South Glamorgan: Cardiff; Coed-y-wenaltt; Wiltshire:
Porton Blake.
FURTHER DISTRIBUTION. Czechoslovakia (Stary, 1977:3).
Dendrocerus punctipes (Boheman)
Ceraphron punctipes Boheman, 1832:332. Holotype 9, SWEDEN (NR, Stockholm).
Ceraphron parvulum Wollaston, 1858: 26, pi. 4, fig. 8. Lectotype 9, MADEIRA (BMNH), designated by
Dessart (1972a: 223) [examined]. [Synonymized by Dessart, 1972a:213.]
Conostigmus punctipes (Boheman) Kieffer, 1907: 137-138.
Dendrocerus punctipes (Boheman) Dessart, 1972a: 213.
FEMALE. Scape long (L/B over 4-0), slightly longer than All, AIII and AIV together; brown, lighter brown
basally; coarsely coriaceous. AIII clearly longer than AIV. Flagellum flat ventrally; brown; finely
coriaceous; pubescence very short.
Head coriaceous; pubescence extremely sparse and short except on clypeus; brown or almost balck, palps
colourless, mandibles brown-yellow, mandibular teeth red. Eyes large, occupying most of lateral length of
head. Eye pubescence very short. Ocular suture narrow. Head rounded, not conspicuously transverse but
clearly broader than thorax. Interantennal carina strongly developed, almost straight but slightly sinuate over
central hump. Toruli not prominent. Frons, above interantennal carina, almost flat with two shallow
depressions above toruli. Frontal dent shallow and indistinct. Depression in front of median ocellus distinct
but rather shallow, the depressions in front of lateral ocelli less evident. Depressions of lateral ocelli sometimes
linked by faint groove. Ocelli arranged in broad-based isosceles triangle. Preoccipital crescent almost absent,
vertical furrow present, starting within ocellar triangle.
Thorax brown, alutaceous, relatively long and narrow. Pubescence short and sparse. Notaulices complete,
sharply angular, converging posteriorly but not meeting median furrow. Secondary furrows visible in some
specimens. Axillae only strongly curved in outer region. Scutellum broad and flat. Propodeum without any
strongly projecting sculpture. Pronotum long with large crenulate furrow. Legs yellow to brown, darkest on
coxae. Long hairs present at junction of coxae and thorax. Most of distal half of forewing dark brown, rarely
lighter in colour. Apart from few hairs on costal vein, forewing totally without fringe. Hindwing with fringe
of long hairs on posterior edge, rest of wing pubescence ultra-short. Pterostigma long and thin, approximately
twice as long as broad, with blunt distal edge. Radius about as long as pterostigma. Brachypterous examples
have radius shorter than pterostigma.
BRITISH SPECIES OF DENDROCERUS 287
Gastral collar long with many distinct carinae and several long lateral hairs. Gaster slightly flat dorsally,
smooth or alutaceous. Gastrocoeli clearly marked as light patches on brown gaster.
MALE. Unknown.
REMARKS. Wollaston (1958:26) described Ceraphron parvulum from an unspecified number of
females. Masner (1965: 17) listed four female specimens of C. parvulum, 'one of which bears a
BMNH type label'; Dessart (1972a: 273) designated the latter specimen as lectotype and the three
others as paralectotypes.
Dendrocerus and Conostigmus are extremely difficult to separate and D. punctipes shows clear
affinities with both genera. The notaulices of D. punctipes indicate a place in Dendrocerus but its
flat scutellum and general appearance are similar to one group of species in Conostigmus.
Brachypterous forms are rare in Dendrocerus, only occurring occasionally in D. dubiosus and D.
punctipes, but are relatively common in Conostigmus. When the male is discovered the
characteristics of its antennae should enable the generic placement of the species to be clarified.
Like D. punctipes, D. wollastoni (Dodd) from St Helena has the forewing pubescence reduced, but
the two species are definitely not conspecific. D. punctipes is rarely collected; it is here recorded as
British for the first time
BIOLOGY. Unknown. Specimens have been collected from June to September.
MATERIAL EXAMINED
Ceraphron parvulum Wollaston, lectotype 9» Madeira: '55-7' (Wollaston).
1 9- Great Britain. Berkshire: Windsor Forest; Devon: Torquay dist.; Surrey: Oxshott. Madeira:
(paralectotypes of Ceraphron parvulum Wollaston). Sweden: Or. Sommarro (ZI, Lund).
FURTHER DISTRIBUTION. Algeria (Dessart, 1979:34); Finland (Hellen, 1966: 19); Germany (West) (Dessart,
1972a:223); Norway (Dessart, 1972a:223).
Dendrocerus pupparum (Boheman)
(Figs 8, 29, 44, 54)
Ceraphron pupparum Boheman, 1832:333-334. Holotype 9, SWEDEN (NR, Stockholm) [examined!.
Ceraphron ancyloneurus Ratzeburg, 1844:217. Syntype(s), [GERMANY]: ex Syrphus larvae (Saxesen) (lost).
[Synonymized by Dessart, 1972a: 224].
Ceraphron puparum: Thomson, 1868:292-293. [Incorrect subsequent spelling of pupparum Boheman.]
Lygocerus puparum (Boheman) Dalla Torre, 1898:534. [Unjustified emendation of pupparum Boheman.]
Lygocerus syrphidarum Kieffer, 1907:35-36. Lectotype 9> FRANCE: 'April ex Syrphidae pupae'
(Carpentier) (MP, Amiens), designated by Dessart (19746:401). [Synonymized by Dessart, 1972a:224.]
Dendrocerus puparum: Dessart, 19720:223. [Incorrect subsequent spelling of pupparum Boheman.]
FEMALE. Scape long (L/B over 4-0), nearly as long as All, AIII and AIV together; thin in basal third and
with central swelling; mostly coriaceous but also papillate. AIII longer than All or AIV. Rest of flagellar
segments short and broad (L/B 1-5-1-7), ventrally flat with longitudinal ridge and coriaceous. Antenna black
or brown; yellow or light brown within antennal socket; pubescence short and dense distally but longer on
three basal segments.
Head alutaceous; pubescence long, up to 0-07 mm (unusually long for this genus), eye pubescence
shorter; black or dark brown, mandibles brown with red teeth, palps clear-brown; strongly transverse, in
dorsal view slightly biconvex about midline. Frons behind interantennal carina with U-shaped depression (as
found in Conostigmus) but with central hump (unlike Conostigmus). Hairs in this area sparse and bent
outwards towards eyes. Interantennal carina incomplete; absent medially, or if complete then very weak and
indistinct centrally. A groove often present behind the carina, but also absent centrally. Frontal dent large
and deep (Fig. 8) with tendency to be linear. Depression in front of median ocellus deep, very close to median
ocellus, sometimes slightly prolonged anteriorly. Depressions by lateral ocelli obsolescent. Ocellar triangle
very broad, ocelli almost in straight line. Lateral ocelli without posterior groove. Preoccipital crescent present
but poorly developed, not reaching eyes or ocelli. Vertical furrow starts as prominent depression between
lateral ocelli. Occipital carina slightly prominent centrally.
Thorax alutaceous, black or dark brown, pubescence long. Mesonotum broad, anterior corners sharp, not
rounded. Notaulices sharply curving outwards anteriorly; converging posteriorly; deeply impressed (median
furrow not as deeply marked). Four very short secondary furrows present on mesonotum. Scutellum broad.
288 N. D. M. FERGUSSON
Transverse furrow of metanotum deep and foveolate. Propodeal carinae prominent with pronounced central
ridge and fovea. Y-shaped furrow on pronotum strongly developed. Legs proximally brown, becoming light
brown or yellow distally, hind coxa always brown.
Forewing with small brown area under radius. Radius very shallowly curved, almost straight. Pterostigma
long and broad (L/B about 1-7), distinctively rounded (Fig. 44). Wing pubescence long, especially on costal
vein, fringe absent centrally from hind margin of forewing.
Gaster brown-black, alutaceous, dorsally flat and ventrally swollen. Gastral collar broad, fluted, with short
strong carinae and long lateral hairs
MALE. As for female except scape about as long as All plus AIII. Flagellar segments not strongly serrate;
strongly papillate; densely pubescent, but hairs shorter than breadth of segment; broad, AIV to AIX about
twice as long as broad (Fig. 54). Radius slightly shorter than in females. Wings not coloured. Parameres long,
projecting from end of gaster and distinctively expanded upwards apically (Fig. 29).
REMARKS. D. pupparum is probably most closely related to D. basalts (unrecorded in Britain)
which is easily distinguished by coloration. The hind coxa of D. basalts females is black/brown on
the proximal half and yellow on the distal half. The scape of the males is yellow.
The holotype of Ceraphron pupparum Boheman was labelled in error by Dessart & Sundholm as
the 'lectotype of Ceraphron puparum Thomson'.
BIOLOGY. A parasite of Diptera, Syrphidae. Ratzeburg (1844:217) recorded C. ancyloneurus from
Syrphus larvae and Kieffer (1907:36) recorded L. syrphidarum from a Syrphid puparium as did
Dessart (1972a:232). Regnier (1923:174) listed Syrphus ribesii and Episyrphus balteatus
(= Syrphus balteatus) as hosts, and Boheman (1832:333) also gave Syrphus ribesii (=Scaeva
ribesii) as a host. One specimen in the BMNH was bred from an Episyrphus balteatus puparium.
Bankowska (1975:312) recorded D. pupparum on Medicago sativa. Both the above-mentioned
Syrphids are common in Britain. D. pupparum has been found on the following plants: Armoracia
rusticana, Crataegus oxyacanthoides, Pinus sp. Quercus robur and Sambucus niger, and has been
collected between May and September.
MATERIAL EXAMINED
Ceraphron pupparum Boheman, holotype 9> Sweden: 'Sm. Bhm' [Boheman] 'Lectotypus C. puparum
Thomson design. Dessart et Sundholm, 1965' (NR, Stockholm).
5 $, 17 9- Belgium: Kessel, Antwerp Prov. Great Britain. Bedfordshire; Buckinghamshire: Slough;
Cambridgeshire: Ent. Fid. Lab. Storeys Way (Varley)\ Cambridge; Greater London: Kew, Wimbledon;
Norfolk: Thuxton (plesiotype [?] of Ceraphron pupparum Boheman, designated by Dessart);
Northamptonshire: Spratton.
FURTHER DISTRIBUTION. Algeria (Dessart, 1979:34); France (Kieffer, 1907:35); Finland (Hellen, 1966:14);
Germany (Dessart, 1972a: 232).
Dendrocerus ramicornis (Boheman)
(Figs 7, 31, 37, 41)
Ceraphron ramicornis Boheman, 1832:329-330. Lectotype tf, SWEDEN (NR, Stockholm), designated by
Dessart (1972a: 243) [examined].
Ceraphron ramicornis Zetterstedt, 1838:413. Lectotype tf, SWEDEN (NR, Stockholm), designated by
Dessart (1972a: 234, 243) [examined]. [Synonymized by Dessart, 1972a: 234.]
Ceraphron glabriculus Thomson, 1858:291. Lectotype J1, SWEDEN (ZI, Lund), designated by Dessart
(1972a:234) [examined]. [Synonymized by Dessart, 1972a:243.]
Lygocerus ramicornis (Boheman) Marshall, 1868: 158.
Lygocerus japonicus Ashmead, 1904:70. Lectotype 9> JAPAN (USNM, Washington), designated by Masner
6 Muesebeck (1968: 1 12) [examined]. [Synonymized by Takada, 1973: 13.]
Lygocerus ratzeburgi Ashmead, 1904:70-71. Lectotype $, JAPAN (USNM, Washington), designated by
Masner & Muesebeck (1968: 113) [examined]. [Synonymized by Takada, 1973: 13.]
Dendrocerus ramicornis (Boheman) Dessart, 1966:5.
FEMALE. Scape long and broad (L/B 4-0-6-0), about as long as All, AIII and AIV together and i..uch
longer than eye; with median swelling and flat ventrally. AIII is longest flagellar segment although AXI can
be almost as long. Flagellar segments broad (L/B 1-5-2-0), except All and AIII (e.g. L/B for AIII 2-0-3-0).
BRITISH SPECIES OF DENDROCERUS 289
Antenna papillate; pubescence short, much less than breadth of segment; black (or brown in some old
specimens), distal end of All and base of scape often lighter.
Head alutaceous to coriaceous; black, mandibles basally black, yellow near apex, teeth red, palps yellow;
pubescence relatively long; strongly transverse. Eyes broad, almost as broad as long. Inter antennal carina
faintly indicated. Frontal dent and two depressions behind toruli shallow. Depressions by lateral ocelli
shallow, crescent-shaped. Median ocellus preceded by small shallow depression. Ocellar triangle very broad,
prominent, ocelli almost in line. Lateral ocelli joined posteriorly by groove. Preoccipital crescent strongly
developed, coming close to ocelli and close to but not quite reaching margin of eye, sharply angled, especially
behind ocelli. Vertical furrow present.
Thorax black (brown in old and faded specimens), alutaceous. Mesonotum broad, anterior mesonotal
corners sharply angled due to deep impressions of notaulices. Notaulices complete, but sometimes less
distinct posteriorly, sharply angled. Four secondary furrows visible. Scutellum broad but strongly arched.
Axillae only strongly curved in outer third. Axillary depression large. Posterior propodeal carina prominent.
Furrow present from top of anterior mesepisternal boundary to episternal scrobe. Short trace present of
smooth straight precoxal sulcus. Legs mostly light brown, coxae brown-black, often smooth and shiny or
weakly alutaceous.
Wings clear. Hindwing with distinct anal lobe (Fig. 37) but indentation of vannal fold smooth and
rounded. Pterostigma rounded, radius clearly angled.
Gaster broad but sometimes rather short; flattened dorsally and convex ventrally; smooth or alutaceous;
dark brown. Gastrocoeli unusually broad but only faintly indicated as light brown patches. Gastral collar
well developed.
A large species, body length 2- 1-2-8 mm.
MALE. As for female except scape black; short, about three times longer than broad. All, AIII, and AIV very
short, A VIII longest segment. AIII to A VII each with long projection, longer than segment (Fig. 31). AVIII
with small hump, AX and AXI cylindrical. Projection on AVII longer than segment AVIII. Pubescence long.
Gaster slightly less broad than in female. Parameres broad with long hairs.
REMARKS. The antennae of D. ramicornis males (Fig. 31) are obviously similar to those of D.
halidayi (Fig. 32), but the former is a darker and larger species.
D. ramicornis Boheman, 1832 was also described as new by Zetterstedt (1838:413). Dessart
(1972a:234, 243) has treated them as two synonymous species and designated a lectotype for each.
BIOLOGY. 1 $, Cinara sp. / Pauesia sp. / Abies firma (Takada, 1973: 14); 1 tf, 2 9, Cinara sp. /
Pauesia sp. / Chamaecyparis pisifirma (Takada, 1973:14); 1 $, 19> Cinara sp. / Pauesia
momicola I Abies firma (Takada, 1973: 14); 1 <$, C. lands I P. pini / Larix leptolepis (Takada,
1973:14); 3 9, C. longipennis / P. konoi / Abies firma (Takada, 1973:14); 1 tf, C. pilicornis / P.
abietis /-; 4 tf, 4 9, C. pinea I -I Pinus sylvestris; 13 <?, 25 9, C. pineti I P. abietis / Pinus
densifiora (Takada, 1973:14); 3 $, Lachniella costata I Pauesia jezoensis I Picea jezoensis
(Takada, 1973:14); 2 <$, 2 9? Lachnus tropicalis / Pauesia japonica I Quercus sp. (Takada,
1973:14); 2 9, Lachnus sp. /-/ Pinus sp. (Ashmead, 1904:70); [?] Pineus pini /-/- (Hellen,
1966: 1 1); Protolachnus thunbergii I Diaeretus leucopterus / Pinus thunbergii (Takada, 1973: 14);
5 <$, Schizolachnus sp. / P. unilachni / Pinus densifiora (Takada, 1973:14); Tuberolachnus
salignus I Euneura nawai /— (Yasumatsu et al. 1946:12); Tuberolachnus salignus I Pauesia
salignae I- (Yasumatsu et al. 1946: 12).
The above records are of Aphidoidea: Cinarinae in association with Eurasian coniferous forest
and Aphidoidea: Lachninae in association with deciduous forest, thus the host preference of D
ramicornis is linked to the Lachnidae rather than the host plant. The Pineus pini record seems
unlikely as this is the only record from the Adelgidae: also this is a very small aphid compared with
the other hosts and D. ramicornis is a large species. Ashmead (1904:71) recorded Z). ramicornis
(as ratzeburgi) from ''Aphis' but elsewhere he used this to mean just aphid. Yasumatsu et al.
(1946: 12) indicated that D. ramicornis (as ratzeburgi andjaponicus) could be a tertiary parasite of
Tuberolachnus salignus via Aphidiidae and Chalcidoidea.
From the material examined D. ramicornis seems to be most common in July.
MATERIAL EXAMINED
Ceraphron ramicornis Boheman, lectotype $, Sweden: 'Sc. Bhn.' [Scania (Boheman)} (NR, Stockholm),
Ceraphron ramicornis Zetterstedt, lectotype $, Sweden: Kengis (NR, Stockholm). Ceraphron glabriculus
Thomson, lectotype $, Sweden: 'Rhn' (ZI, Lund). Lygocerus japonicus Ashmead, lectotype 9, Japan: Atami
290 N. D. M. FERGUSSON
(Koebele) (USNM, Washington). Lygocerus ratzeburgi Ashmead, lectotype <5\ Japan: (Koebele) (USNM,
Washington).
12 (5, 9 9- Germany (West): Wiirbenthal. Great Britain. Buckinghamshire: Soulbury, Rammamere
Heath; Hampshire: New Forest; Highland: Caledonian Forest, Glen Afric; Strathclyde: Beinn Eigh; Surrey:
Esher Common. Japan: Honshu, Kobe; Honshu, Kyoto; no further data (paralectotype of Lygocerus
ratzeburgi Ashmead) (USNM, Washington). Netherlands: Wageningen. Spain: Castellon, Benicasim.
Sweden: Gtl, Ostergarn, Vike (ZI. Lund); [label illegible] (ZI, Lund).
FURTHER DISTRIBUTION. Finland (Hellen, (1966: 1 1).
Dendrocerus serricornis ( Boheman)
(Figs 43, 53)
Ceraphron serricornis Boheman, 1832:334—335. Holotype $, SWEDEN (ZI, Lund) [examined].
Ceraphron serricornis Zetterstedt, 1838:413. Holotype tf, SWEDEN (ZI, Lund) [examined, same specimen
as above]. [Synonymized by Dessart, 1972a: 253.]
Ceraphron piceae Ratzeburg, 1852:179. Syntype(s), GERMANY (lost). [Synonymized by Dessart,
1972a:253.]
Ceraphron lapponicus Thomson, 1858:290. Holotype 9, SWEDEN (NR, Stockholm) [examined].
[Synonymized by Dessart, 1972a:253.]
Lygocerus serricornis (Boheman) Marshall, 1868: 158.
Lygocerus semiramosus Kieffer, 1907:38. Lectotype tf, GREAT BRITAIN (BMNH), designated by
Masner (1 965 :21) [examined]. Syn. n.
Lygocerus subramosus Kieffer, 1907:39. Syntype(s), FRANCE (lost). [Synonymized by Dessart, 1972a: 253.]
[Lygocerus testaceimanus Kieffer; Wilson, 1938:378. Misidentification.]
Atritomellus zetterstedti Ghesquiere, 1960:208. [Replacement name for Ceraphron serricornis Zetterstedt,
1838.] [Synonymized by Dessart, 1972a:253.]
Ceraphron serraticornis: Ghesquiere, 1960:208. [Incorrect subsequent spelling of serricornis Zetterstedt.]
Dendrocerus serricornis (Boheman) Dessart, 1966: 13.
FEMALE. Scape short (L/B 4-0-4-5), slightly shorter than length of head; longer than All, AIII and AIV
together. In small specimens antenna! segments very short and scape often longer than next four segments
together. All about as long as AIII and often longer than AIV. AIII longer than any of following basal
segments which are stout (roughly 1-5 times longer than broad). Antenna black except for distal end of All,
which is lighter. Scape coriaceous to papillate, flagellum weakly papillate, pubescence short and dense.
Head coriaceous or alutaceous; pubescence not long but quite noticeable, clypeal hairs long, eye
pubescence short; black, mandibles brown with red-brown teeth, palps clear-yellow; rounded (especially in
dorsal view), less transverse than in many species of Dendrocerus. Interantennal carina variable, sometimes
almost absent or just complete, but usually present near toruli and absent over median hump. Depressions
behind and inwards from each torulus united to form large, slightly depressed area above interantennal
carina. Median hump between toruli gives this depression an appearance similar to the stronger, horseshoe-
shaped depression found in some Conostigmus species. Frontal dent absent. Depressions near ocelli usually
shallow but median depression often deeper in large specimens. Ocellar triangle broad, very short. Lateral
ocellar depressions sometimes linked by groove. Vertical furrow present on preoccipital crescent. Preoccipital
crescent very strongly developed, reaching ocelli and margin of eye; abruptly angled downwards from vertex.
Occipital carina foveolate.
Thorax black (or dark brown in old specimens); often long and narrow, 1-4-1-8 times longer than broad.
Anterior corners of mesonotum rounded. In lateral view mesonotum swollen anteriorly; this may be
enhanced by a backward slant of pronotum and propleuron. Notaulices complete, four secondary furrows
usually distinct. Notaulices very strongly angled anteriorly, almost straight posteriorly except for last fraction
where they curve towards midline. Scutellum rather narrow, raised region about 1 -4 times longer than wide,
clearly arched but sometimes slightly flat dorsally. Metanotal furrow strongly foveolate or canaliculate.
Propodeum coriaceous to rugose, appreciably more coarsely sculptured than rest of dorsal surface of thorax.
Propodeal fovea not prominent. Pleural suture foveolate, episternal scrobe distinct. Legs black-brown, lighter
or clear brown on tibia and tarsus (especially of forelegs). Wings clear, pubescence normal. Radius well
curved, about 1-3 times longer than pterostigma. Pterostigma characteristically shaped (Fig. 43), about 1-5
times longer than broad.
Gaster 1-5-2-0 times longer than broad. Gastral collar large with distinct carinae. Gaster mostly smooth
or finely alutaceous, gastrocoeli visible. Gaster lighter brown anteriorly, dark brown posteriorly.
BRITISH SPECIES OF DENDROCERUS 291
MALE. As for female except scape very short, much less than length of head and not as long as combined
length of All, AIII and AIV. Basal flagellar segments strongly serrate (Fig. 53), AIII characteristically
shaped, 1-0-1 -5 times longer than broad. AIV as long as broad (L/B 0-9—1-2). Flagellar pubescence much
longer than breadth of segment. Depression behind interantennal carina less obvious, head more transverse
and thorax less rounded than in female.
REMARKS. D. serricornis Boheman, 1832 was also described as new by Zetterstedt (1838:413).
Dessart (1972a:253, 261) has treated them as two synonymous species having the same holotype.
The specimens misidentified by Wilson (1938:378) as Lygocerus testaceimanus are in the
BMNH and are D. serricornis, as Dessart (1972a:254) suspected.
D. serricornis is recognizable by the shape of the male antenna, head and thorax, but females of
this species and D. carpenteri are sometimes hard to distinguish. The basal flagellar segments of D.
serricornis males are more strongly serrate than in other British species (except those with ramose
antennae, i.e. D. halidayi, Fig. 32 and D. ramicornis, Fig. 31).
BIOLOGY. Host information indicates that D. serricornis is chiefly a parasite of Chamaemyiidae
(Diptera) which prey on Adelgidae (Hemiptera).
Wilson (1938:378) reared D. serricornis (as Lygocerus testaceimanus) from pupae of Leucopis
obscura (Diptera: Chamaemyiidae) which were preying upon the Adelgids Pineus pint (on Pinus
sylvestris) and P. strobi (on Pinus strobus}. D. serricornis can subsequently emerge from the pupae
(collected in July) or can diapause and emerge in the following May. Melanips longitarsis
(Hymenoptera, Cynipoidea) and Aphidencyrtus aphidivorus (Hymenoptera, Chalcidoidea) were
also parasitic on L. obscura but D. serricornis was the most common parasite. These three
parasites seem to be most efficient as Wilson states that 'all the L. obscura puparia present in the
field from late summer to late winter are parasitized specimens'. In addition to Wilson's records, I
have seen a specimen of D, serricornis bred from a Leucopis sp. puparium found on Fagus
sylvatica.
Schremmer (1956:58) recorded D. serricornis (as Lygocerus piceae) as a parasite of Cremifania
nigrocellulata (Diptera: Chamaemyiidae) which was preying on Adelges (Dreyfusid) piceae
(Hemiptera, Adelgidae). Delucchi & Pschorn-Walcher (1954: 102) recorded D. serricornis
(=piceae) as a parasite of the 3rd larval stage of Cremifania nigrocellulata. Pachyneuron
vitodurense (=P.ferrierei Delucchi) and P. cremifaniae (Chalcidoidea, Pteromalidae) were given as
parasites of the puparia of Cremifania. D. serricornis and P. vitodurense were also listed as
parasites of Leucopis obscura and L. griseola. These Leucopis species and Cremifania
nigrocellulata are all predators on related Adelgids. Ratzeburg (1848:216) recorded D. serricornis
(as piceae) from Adelges piceae (C Hermes piceae).
According to Stary (1970:200) no Aphidiidae have been recorded as primary parasites of
Adelgids; if D. serricornis is a hyperparasite then it is likely to be via Chalcids, like those
mentioned above.
D. serricornis is known from other hosts. Lai (1934:328) recorded L. semiramosus as 'believed
to be' a hyperparasite of Prionomitus mitratus (Chalcidoidea, Encyrtidae) which was in turn a
primary parasite of Psylla peregrina on Crataegus. This chalcid has also been listed as a parasite of
other Psylla species. The hyperparasites appeared in mid September. One nymph contained three
recently hatched Dendrocerus larvae. Lai presumed that the Dendrocerus 'were acting as tertiary
parasites of a secondary parasite, or of their own species'. Haviland (1920ft: 103) recorded a similar
case. The egg of serricornis is long and elliptical, 0-67 x 0-28 mm, the surface sculptured with fine
longitudinal striae. The larva, which emerges through an operculum at the end of the egg, is small,
creamy white, and measures 0-45 x 0- 15 mm.
A male and female of D. serricornis in the BMNH were bred from second instars of
Parthenolecanium corni (Coccoidea) on Corylus avellana. D. serricornis has been found on the
following plants: Crataegus, Humulus lupulus, Populus nigra italica and Pinus sp. With the
exception of hop all the plant records are of trees or shrubs, and many of the trees are conifers.
Kieffer ( 1 907: 39) recorded L. subramosus from Cecidomyia pini (Diptera, Cecidomyiidae).
From the material examined D. serricornis seems to be most common in late July.
292 N. D. M. FERGUSSON
MATERIAL EXAMINED
Ceraphron serricornis Boheman and Ceraphron serricornis Zetterstedt, holotype $ (of both nominal
taxa], Sweden: 'Job No' (ZI, Lund). Ceraphron lapponicus Thomson, holotype 9, Sweden: Lappland
(Boheman) (NR, Stockholm). Lygocerus semiramosus Kieffer, lectotype $, Great Britain: 'Cad'
[Strathclyde, Gadder] (Cameron) (BMNH).
35 cf, 13 9» Great Britain. Bedfordshire (Chambers coll.); Buckinghamshire: Farnham House; Cheshire:
Delamere (MM, Manchester); Cumbria: Skirwith (MM, Manchester); Grampian: Elgin; Greater London:
Kew; Hertfordshire: Tring; Kent: Barnehurst, East Mailing, Eynsford; Norfolk: North Wooton;
Northamptonshire: Spratton; North Yorkshire: Malham Tarn (NM, Manchester); Oxfordshire: Oxford;
Suffolk: Santon Downham, Barton Mills; Wiltshire: Corsham. Ireland: Co. Dublin, Grand Canal; Leitrim; Co.
Wicklow, Killegar. Japan: Atami (paralectotype of Lygocerus koebelei) (USNM, Washington). Spain:
Valencia, El Perello. Sweden: Skane, Ring; Or (ZI, Lund).
FURTHER DISTRIBUTION. Finland (Hellen, 1966:11); France (Dessart, 1972a:266); Germany (Dessart,
1972a:266).
Dendrocerus spissicornis (Hellen)
(Fig. 19)
Lygocerus spissicornis Hellen, 1966: 12. Holotype 9> FINLAND, (ZMU, Helsinki) [examined].
Dendrocerus spissicornis (Hellen) Dessart, 1972a: 274—281.
FEMALE. Scape short and broad (L/B 3-6—5-0), thinner at base than apex, slightly longer than All, AIII and
AIV together; colour varying from entirely dark to orange-yellow, often lighter basally. AIII thinner (L/B 1 -4
or greater) than AIV to AX, which are quadrate. All and proximal part of AIII often orange-yellow, rest of
flagellum brown. Antennal insertion orange-yellow. Flagellar segments flat ventrally with single central ridge.
Antenna papillate with dense pubescence.
Head sculpture mostly rugose, large depression above interantennal carina sometimes weakly sculptured
but usually strongly strigose. Eye margin smooth. Head black, palps yellow or orange. Mandibles basally
black, yellow or orange medially, with red-black teeth. Toruli very short, especially internally. Interantennal
carina distinct, often slightly sinuate centrally over small hump. Large depression present above toruli.
Depression in front of median ocellus deep and distinct. Depressions near lateral ocelli linear and not joined
by ridge running behind ocelli. Preoccipital crescent strongly developed but not quite reaching margin of eye
or lateral ocelli, with central vertical furrow. Last segment of maxillary palps much longer than penultimate
segment. Hair round mandibles and on clypeus longer than on rest of head.
Thorax black, strongly sculptured, granulate, punctate and rugose; all furrows deeply foveolate.
Mesonotum and scutellum flattened but central area between notaulices slightly raised anteriorly.
Mesonotum with notaulices complete, reaching posterior margin and meeting median furrow. Notaulices not
deeply curved. Secondary furrows evident. Scutellum broad, very flat, with transverse carina apically.
Narrow band of strigose sculpture present running parallel to edge of scutellum from axillae to transverse
carina. Metanotum reduced, obscured by large bilobed transverse keel situated in middle of propodeum.
Femora brown, or black in fresh specimens, tibiae and tarsae lighter.
Wings almost clear but with feeble brown mark around radius; however, one old specimen (c. 1826) has
distinct brown cloud under radius. Pterostigma semioval, radius approximately as long as breadth of
pterostigma. Fringe short and fine.
Gastral collar broad, very short. Entire breadth of gaster covered by area of strigose sculpture radiating
from gastral collar and nearly reaching gastrocoeli (Fig. 19). Rest of gaster smooth or alutaceous.
MALE. As for female except scape longer than segments All, AIII and AIV together, just over 3 times as
long as broad. All shorter than AIII. AIII about 1-5 times longer than broad. AIV and AV as long as broad.
Rest of flagellar segments progressively less broad, AXI being 3 times longer than broad. Basal flagellar
segments clearly asymmetrical, especially segments AIV to AVII. Antenna brown except for All and small
light area at base of scape; papillate; with dense pubescence shorter than breadth of segment.
REMARKS. D. spissicornis is here recorded as British for the first time. It is generally more grossly
sculptured than is usual in Dendrocerus, also the notaulices are not as deeply curved as they are in
other species of the genus. D. spissicornis is rare, at least in collections.
BIOLOGY. Dessart (1972a: 281) has seen two specimens of D. spissicornis for which the puparia of
Cyclorrhapha (Diptera) are indicated as hosts. Half of the British specimens so far known were
collected on dead beech or oak trees. Dates of collection range from late June to early September.
BRITISH SPECIES OF DENDROCERUS
293
MATERIAL EXAMINED
Lygocerus spissicornis Hellen, holotype Q, Finland: Nystad (Hellen) (ZMU, Helsinki).
1 <3\ 11 9- Great Britain. Buckinghamshire: Shabbington Wood, nr Oakley; Cheshire: Abbots Moss
(MM, Manchester), Rostherne; Hampshire: W. of Winchester, Crab Wood; Surrey: Woodcote (MM,
Manchester); no further data (UM, Oxford). Sweden: [label illegible,] (ZI, Lund); Upl. Vallentuna (NR,
Stockholm) (male allotype).
FURTHER DISTRIBUTION. Belgium (Dessart, 1972a:281); France (Dessart, 1972a:281).
Review of the biology of Dendrocerus
The host records for the British species of Dendrocerus are here listed under host classification. The
basic host relationships of Dendrocerus are summarized in tabular form. The biology of the British
species is associated mostly with the Homoptera Sternorhyncha and is typical of the genus as a
whole. The faunistic complexes used in this work are briefly defined.
Host aphid list for Dendrocerus species
Host
APHIDOIDEA
LACHNIDAE
Cinarinae
Protolachnus
Schizolachnus
Cinara
Lachniella
Lachninae
Lachnini
Tuberolachnus
Lachnus
CHAITOPHORIDAE
Chaitophorinae
Periphyllus
Parachaitophorus
Siphinae
Sipha
CALLAPHIDIDAE
Callaphidinae
Callaphidini
Chromaphis
Myzocallidini
Myzocallis
Tuberculoides
Eucallipterus
Tinocallis
Shivaphis
Pterocallis
Phyllaphidinae
Phyllaphidini
Callipterinella
Betulaphis
Symydobius
Clethrobius
Eucer aphis
Parachaitophorus
Drepanosiphini
Drepanosiphum
Dendrocerus species
carpenteri, liebscheri, ramicornis
carpenteri, dubiosus, liebscheri,
ramicornis
carpenteri, liebscheri, ramicornis
ramicornis
carpenteri, ramicornis
ramicornis
carpenteri, laticeps
carpenteri
aphidum, carpenteri
laticeps
carpenteri
carpenteri, laevis
aphidum, carpenteri
carpenteri
laevis
carpenteri
carpenteri
carpenteri
carpenteri
aphidum
carpenteri
carpenteri
carpenteri
294
Therioaphidinae
Therioaphis
Neophyllaphidinae
Neophyllaphis
APHIDIDAE
Pterocommatinae
Pterocomma
Aphidinae
Hyalopterus
Rhopalosiphum
Schizaphis
Paraschizaphis
Melanaphis
Aphis
Toxoptera
Myzinae
Cryptosiphum
Ceruraphis
Dysaphis
Brachycaudus
Diuraphis
Hayhurstia
Brevicoryne
Pseudobrevicoryne
Lipaphis
Hyadaphis
Staegeriella
Toxopterella
Hyalopteroides
Coloradoa
Myzaphis
Chaetosiphon
Elatobium
Liosomaphis
Cavariella
Ovatus
Phorodon
Rhopalomyzus
Myzus
Galiobium
Tuberocephalus
Cryptomyzus
Capitophorus
Pleotrichophorus
Nasonovia
Hyperomyzus
Rhopalosiphoninus
Indomegoura
Macrosiphinae
Microlophium
Aulacorthum
Acyrthosiphon
Metopolophium
Corylobiwn
Staticobium
N. D. M. FERGUSSON
dubiosus
carpenteri
carpenter!
aphidum, carpenteri, laevis
aphidum, carpenteri, laevis,
laticeps
aphidum, carpenteri, laevis
aphidum, bifoveatus
carpenteri
aphidum, bifoveatus, carpenteri,
dubiosus, laevis, laticeps
carpenteri, laticeps
dubiosus
carpenteri
carpenteri, laevis
aphidum, carpenteri, laticeps
aphidum, carpenteri
carpenteri
carpenteri, laticeps
carpenteri
carpenteri, laticeps
carpenteri
?laticeps
carpenteri
dubiosus
carpenteri, laticeps
carpenteri
carpenteri
carpenteri
aphidum, carpenteri
aphidum, carpenteri, laticeps
carpenteri
carpenteri
carpenteri
aphidum, carpenteri, laticeps
dubiosus
carpenteri
carpenteri
carpenteri, laticeps
carpenteri
carpenteri
carpenteri
bifoveatus, carpenteri, dubiosus
carpenteri
aphidum, carpenteri, dubiosus,
laticeps
carpenteri, dubiosus
aphidum, bifoveatus, carpenteri,
dubiosus, laticeps
aphidum, carpenteri, laevis
carpenteri
carpenteri
BRITISH SPECIES OF DENDROCERUS 295
Macrosiphum carpenteri, dubiosus, laticeps
Sitobion aphidum, carpenteri, dubiosus,
laevis, laticeps
Titanosiphon carpenteri
Metopeurum carpenteri
Uroleucon aphidum, carpenteri, dubiosus
Macrosiphoniella aphidum, carpenteri, laticeps
Amphorophora carpenteri
THELAXIDAE
Thelaxinae
Mindarus carpenteri
Hormaphidinae
Hamamelistes carpenteri
PEMPHIGIDAE
Eriosomatinae
Pemphiginae
Prociphilus carpenteri
ADELGIDAE serricornis
Pineinae
Pineus ? ramicornis, serricornis
Adelginae
Adelges serricornis
* Stary (1977:4) recorded Dendrocerus sp. from Eriosoma ulmi but the specimens have not been made available for identification.
List of the Aphidiidae (Hymenoptera) parasitized by Dendrocerus
Host Dendrocerus species
APHIDIIDAE
Ephedrinae
Ephedrus californicus carpenteri
E. lacertosus carpenteri
E. minor carpenteri
E. nacheri carpenteri, dubiosus
E. niger carpenteri, dubiosus
E. persicae aphidum, carpenteri, laticeps
E. plagiator aphidum, carpenteri, dubiosus,
laticeps
E. salicicola carpenteri
E. sp. aphidum, carpenteri, dubiosus
Prainae
Areopraon nipponicum carpenteri
Praon abjectum aphidum, carpenteri
P. absinthii carpenteri
P. barbatum carpenteri
P. bicolor carpenteri, liebscheri
P. capitophori carpenteri
P. dorsale carpenteri, dubiosus
P. exsoletum dubiosus
P.flavinode carpenteri
P. gallicum carpenteri
P. necans aphidum, carpenteri
P. pequodorum carpenteri
P. silvestre carpenteri
P. volucre aphidum, carpenteri, dubiosus
P. sp. aphidum, carpenteri, dubiosus
Dyscritulus planiceps carpenteri
D. sp. carpenteri
296
N. D. M. FERGUSSON
Aphidiinae
Aphidiini
Lysiphlebus ambiguus
L. arvicola
L. dissolutus
L.fabarum
L.fritzmuelleri
L. testaceipes
Lysiphlebiajaponica
Pauesia abietis
P. akamatsucola
P. infulata
P.japonica
P.jezoensis
P. konoi
P. lands
P. momicola
P.piceaecollis
P. pint
P. salignae
P. unilachni
P.sp.
Metaphidius aterrimus
Diaeretus leucopterus
Aphidius absinthii
A . amamioshimensis
A. aquilus
A. areolatus
A. cingulatus
A . equiseticola
A. ervi
A.funebris
A. gifuensis
A. hieraciorum
A. hortensis
A. longipetiolus
A. matricariae
A. phalangomyzi
A . picipes
A.platensis
A. polygonaphis
A. rosae
A. salicis
A. schimitscheki
A . setiger
A. smithi
A. sonchi
A. tanacetarius
A. urticae
A. uzbekistanicus
A.sp.
Lysaphidus arvensis
L. erysimi
L. matsuyamensis
L. pleotrichophori
Diaeretiella rapae
carpenteri
carpenteri
carpenteri
carpenteri, dubiosus
carpenteri
carpenteri
carpenteri, laticeps
carpenteri, ramicornis
carpenteri
liebscheri
ramicornis
ramicornis
ramicornis
carpenteri
ramicornis
carpenteri
carpenteri, liebscheri, ramicornis
carpenteri, ramicornis
carpenteri, dubiosus, liebscheri,
ramicornis
carpenteri, liebscheri, ramicornis
carpenteri
carpenteri, liebscheri, ramicornis
carpenteri
laticeps
carpenteri
carpenteri
carpenteri
carpenteri, dubiosus
aphidum, carpenteri, dubiosus
carpenteri
carpenteri, laticeps
carpenteri
carpenteri
carpenteri, laticeps
carpenteri, dubiosus
carpenteri
aphidum, carpenteri, dubiosus,
laticeps
aphidum
carpenteri
carpenteri, dubiosus
carpenteri, laticeps
carpenteri
carpenteri
carpenteri
aphidum
carpenteri
bifoveatus, carpenteri
aphidum, carpenteri, dubiosus
aphidum, carpenteri, dubiosus,
laticeps
laticeps
carpenteri
carpenteri
carpenteri
aphidum, bifoveatus, carpenteri,
dubiosus, laticeps
BRITISH SPECIES OF DENDROCERUS
297
Trioxini
Monoctonus angustivalvus
M. caricis
M. cerasi
M. pseudoplatani
Lipo lexis gracilis
Calaphidius watanabei
Trioxys auctus
T. betulae
T. cirsii
T. curvicaudus
T. euceraphis
T.falcatus
T. pallidus
T. pannonicus
T. parauctus
T. shivaphis
Binodoxys acalephae
B. angelicae
B. brevicornis
B. centaureae
B. sinensis
carpenteri
dubiosus
carpenteri
carpenteri
carpenteri
carpenteri
laticeps
aphidum, carpenteri
carpenteri
carpenteri
carpenteri
carpenteri, laticeps
carpenteri, laevis, laticeps
carpenteri
carpenteri
laevis
aphidum, carpenteri, dubiosus
carpenteri, dubiosus, laevis,
laticeps
carpenteri
carpenteri
laticeps
Stary (1977: 5) recorded an unidentified species of Dendrocerus from Areopraon lepelleyi.
Other hosts
Host
HEMIPTERA
HOMOPTERA
PSYLLOIDEA
Psylla peregrina
COCCOIDEA
Coccidae
Parthenolecanium corni
Eriococcidae
Eriococcus araucariae
Pseudococcidae
Heterococcus nudus
Planococcus citri
P. longispinus
Pseudococcus fragilis
NEUROPTERA
SIALOIDEA
Coniopterygidae
Conwentzia psociformis
Semiadalis aleyrodiformis
Coniopteryx esbenpeterseni
Chrysopidae
Chrysopa virgestes
COLEOPTERA
Curculionidae
Erirhininae
Stenopelmus rufinasus
DIPTERA
Dendrocerus species
serricorms
serricornis
laevis
laevis
laevis
laevis
laevis
halidayi
halidayi
flavipes
? aphidum
? laticeps
? aphidum
298
N. D. M. FERGUSSON
NEMATOCERA
Cecidomyiidae
Rhabdophaga rosaria
Cecidomyia pint
CYCLORRHAPHA
Syrphidae
Syrphus ribesii
S.sp.
Episyrphus balteatus
Chamaemyiidae
Leucopis griseola
L. obscura
L. sp.
Cremifania nigrocellulata
Chloropidae
Meromyza saltatrix
HYMENOPTERA
FORMICOIDEA
Formica rufa
Lasius brunneus
L.fuliginosus
CYNIPOIDEA
Alloxystinae
Phaenoglyphis sp.
Alloxysta brassicae
A. victrix
A. sp.
Figitidae
Melanips longitarsis
Cynipidae
Gall of Biorhiza pallida
Gall ofDiplolepis rosae
CHALCIDOIDEA
Aphelinidae
Aphelinus abdominalis
A. chaonia
A. daucicola
Encyrtidae
Leptomastidea abnormis
Tetracnemus diversicornis
Microterys sp.
Aphidencyrtus aphidivorus
Prionomitus mitratus
Encyrtus sp.
Pteromalidae
Asaphes lucens
A. suspensus
A. vulgaris
Cyrtogaster vulgaris
Pachyneuron aphidis
P. cremifaniae
P. siphonophorae
P. vitodurense
Euneura nawai
Coruna clavata
? laevis
serricornis
pupparum
pupparum
pupparum
serricornis
serricornis
serricornis
serricornis
carpenteri
carpenteri
dubiosus
carpenteri
carpenteri
carpenteri
carpenteri
carpenteri
? serricornis
halidayi
carpenteri
carpenteri
laticeps
laticeps
laevis
laevis
laevis
carpenteri, ? serricornis
serricornis
carpenteri
carpenteri
carpenteri
carpenteri
carpenteri
carpenteri
serricornis
carpenteri
serricornis
ramicornis
carpenteri
Occasional fortuitous plant records may result from parasitized aphids moving from their normal
host plant before becoming mummified.
The host lists show that the British species of Dendrocerus are mostly aphid hyperparasites, but
BRITISH SPECIES OF DENDROCERUS
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some are parasitic on Diptera, some on Neuroptera and D. laevis is parasitic on coccids as well as
aphids. The biology of Dendrocerus is clearly very closely related to the Homoptera-Sternorhyncha
(i.e. Psylloidea, Aleyrodoidea, Aphidoidea and Coccoidea), and even the hosts listed from other
groups are often predators on aphids (e.g. Neuroptera; Diptera, Syrphidae, etc.).
Host information on some non-British species of Dendrocerus
The above pattern of host choice is typical of the genus as a whole. Host information about other
Dendrocerus species is briefly listed below.
Host
DIPTERA
Chamaemyiidae
Leucopis atrifacies
L. sp.
Cecidomyiidae
Mayetiola destructor
Dipterous larvae
NEUROPTERA
Coniopterygidae
Conwentzia hageni
Nimboa basipunctata
Hemerobiidae
Chrysopidae
Chrysopa madestes
C. sp.
HEMIPTERA
HOMOPTERA
APHIDOIDEA
Plocamaphis goernitzi
Macrosiphum euphorbiae
Macrosiphwn euphorbiae
Neuquenaphis similis
A wide range of aphids
COCCOIDEA
Saissetia oleae
COLEOPTERA
Coccinellidae
Pharoscymnus ovoideus
P. numidicus
Scymnus sp.
Chilocorus kuwanae
HYMENOPTERA
CHALCIDOIDEA
Encyrtidae
Homalotyus flaminius
Dendrocerus
species
D. pinicolus
D. leucopidis
D. triticum
D. koyamai
D. conwentziae
D. indicus
D. latifrons
D. propodealis
D. noumeae
D. remaudierei
D. attentus
D. incompletus
D. henkvlugi
D.floridanus
D. noumeae
D. ergensis
D. ergensis
D. ergensis
D. chilocori
D. ergensis
Country
U.S.A.
U.S.A.
U.S.A.
Italy, Japan
U.S.A.
India
U.S.A.
India
New Caledonia
Europe
U.S.A.
U.S.A.
Chile
Japan, U.S.A.
Reference
Muesebeck(1959:93)
Muesebeck(1959:92)
Ashmead (1893: 110)
Dessart(1972a:176)
Gahan (1919: 122)
Dessart(1972a:171)
Muesebeck(1959:94)
Dessart( 1973: 274)
Dessart( 1967: 346)
Dessart(1974a:82)
Muesebeck(1959:96)
Muesebeck(1959:96)
Dessart( 1975: 266)
Ashmead (1881: xxxiv)
New Caledonia Dessart (1967: 346)
Spain,
North Africa
North Africa
North Africa
Japan
Ghesquiere( 1960: 207)
Ghesquiere( 1960: 207)
Ghesquiere( 1960: 207)
Ishii( 195 1:93)
Ghesquiere( 1960: 207)
Homalotyus flaminius
Spain,
North Africa
D. chilocori Japan Ishii ( 1 95 1 : 94)
Conostigmus has often been reported as a parasite of aphids, but it is probable that these records
actually pertain to Dendrocerus. The record published by Jones (1972: 13) refers to a specimen of
Dendrocerus and not to Conostigmus (Jones, pers. comm.). According to Ashmead (1893:103)
Riley reared a Dendrocerus sp. from a tortricid larva but Haviland (19206: 102) considered this to
be unlikely.
In order that the biology of Dendrocerus can be easily related to that of the Aphidiidae, I have
used the faunistic complexes adopted by Stary (1970:201-202, 312-321), as very briefly indicated
below
BRITISH SPECIES OF DENDROCERUS 301
Boreal Europe faunis tic complex. Cooler climatic conditions, restricted to northern parts of
Europe. Some typical species: Decorosiphon spp. / Diaeretellus ephippium I mosses;
Rhopalosiphum nymphaeae I Praon necans I secondary host plants.
Eurasian steppe faunistic complex. Typical complex of steppe zone plus the cultivated steppe
landscape. Some typical species: Macrosiphoniella spp. / Aphidius absinthii /-; Acyrthosiphon spp.
/ Aphidius ervi /-; Cryptosiphum spp. / Ephedrus nacheri /-; Coloradoa spp. / Lysaphidus
arvensis /-; Uroleucon spp. / Praon dorsale /-; Aphis spp. / Binodoxys acalephae /—.
Deciduous forest (European) faunistic complex. Typical of the deciduous forests of Europe to
which they are mostly restricted. Some typical species: Liosomaphis berberidis I Aphidius
hortensis / Berberis; Macrosiphum spp. / Aphidius rosae /-; Macrosiphum spp. / Aphidius
lonicerae /-; Drepanosiphum spp. / Dyscritulus planiceps I Acer; Sitobion spp. / Monoctonus
caricis /-; Periphyllus spp. / Trioxys falcatus /—.
Deciduous forest (Far Eastern) faunistic complex. Typical of the deciduous forests of the Far
East and penetrating to the allied tropics but some elements have a wide trans-palaearctic distribu-
tion. Some typical species: Periphyllus spp. / Aphidius areolatus I Acer; Myzus spp. / Aphidius
gifuensis /-; Tuberolachnus salignae / Salix.
Coniferous forest (West Eurasian) faunistic complex. Members of this complex are distributed
all over Europe in coniferous forests. Some typical species: Cinara spp. / Metaphidius aterrimus I
conifers; Lachnidae aphids / Pauesia spp. / conifers.
Coniferous forest (East Eurasian) faunistic complex. Typical of the Far Eastern type of
coniferous forest but some species are widely distributed in Europe. Typical species: Lachnidae
aphids / Pauesia spp. / conifers; Eulachnus spp. / Diaeretus leucopterus I conifers.
Holarctic forest tundra faunistic complex. Typical of cool arctic conditions, it is a transitional
zone between forest-free tundra to the north and mostly coniferous forests to the south. Typical
trees: Salix, Alnus, Betula etc. Many elements of this complex are found further south. Typical
species: Pterocomma spp. / Aphidius cingulatus I Salix, Populus; Symydobius I Trioxys betulae.
Non-British species
The following species occur in Europe but have not been found in the British Isles.
D. basalis (Thomson, 1858) was mentioned by Marshall (1868: 158; 1873:3) as a synonym of
D. carpenteri but it has not been directly recorded as a British species. The most similar British
species to D. basalis is D. pupparum; in the discussion of the latter I have shown how the two
species can be separated.
D. ergensis (Ghesquiere, 1960) was described from North Africa, but I have seen specimens from
Spain. This species is easily identified as the radius is shorter than the pterostigma and the
notaulices are only present anteriorly; in males the basal flagellar segments are strongly serrate,
about as long as broad.
D. koyamai (Ishii, 1951). This species was known only from Japan until Dessart (1974a: 75)
recorded it from Italy. The radius is just shorter than the pterostigma, and in males the basal
flagellar segments have projections similar to those found in D. halidayi and D. ramicornis.
D. omostenus Dessart, 1979 is known only from the Algerian female holotype which I have
examined and believe to be a Conostigmus species.
D. remaudierei Dessart, 1974 was described from France. It has characters in common with
several different species of Dendrocerus, but it is distinguished by the following combination of
characters. The notaulices are complete, the interantennal carina is absent, the wings are hyaline, the
male antennae are only moderately serrate (L/B for AIII about 2-35), the flagellar pubescence in
males is longer than the breadth of the segment and the parameres are enlarged and truncate.
D. solarii (Kieffer, 1907) is known only from the Italian female holotype. It is distinguished from
other European species by the following characters. The apex of the scutellum has foveolate
sculpture which fades out laterally, the third antennal segment is elongate, about 3 times longer
than broad, and the median fovea on the propodeum is prominent.
302
N. D. M. FERGUSSON
Figs 2-12 2-4, Megaspilus sp., (2) scutellum and propodeal projection; (3) dorso-lateral view of
propodeal projection; (4) precoxal sulcus. 5, 6, heads of Conostigmus species. 7, 8, heads of (7)
Dendrocerus ramicornis; (8) D. pupparum. 9, Conostigmus sp., brachypterous forewing. 10-12,
Lagy nodes pallidus, (10) forewing of male; (1 1) tibial spurs of foreleg; (1 2) female.
BRITISH SPECIES OF DENDROCERUS
303
15
21
22
23 24
Figs 13—24 13, mesonotum of Conostigmus sp. 14—17, mesonota of (14) and (16) Dendrocerus
carpenteri; (15) D. laticeps; (17) D. liebscheri. 18-20, gastral collar of (18) Conostigmus
fasciatipennis; (19) Dendrocerus spissicornis; (20) D. dubiosus. 21-24, D. carpenteri, larval stages,
(21) first instar; (22) second instar; (23) third instar; (24) fourth instar.
304
N. D. M. FERGUSSON
25
30
31
Figs 25-32 Males. 25, 26, head of (25) Dendrocenis laevis; (26) D. laticeps. 27-29, gaster showing
parameres of (27) D. liebscheri; (28) D. carpenteri; (29) D. pupparum. 30-32, antennae of (30) D.
flavipes; (3 1) D. ramicornis; (32) D. halidayi.
BRITISH SPECIES OF DENDROCERUS
305
Figs 33-39 33-38, wings of (33) Dendrocerus aphidum; (34)£>. laticeps-,(35)D.flavipes;(36)D. laevis;
(3 7) D. ramicornis; (38) D. bifoveatus. 39, coastal vein and pterostigma of Conostigmusfasciatipennis.
306
N. D. M. FERGUSSON
40
Figs 40-48 40-46, pterostigma and radius of (40) Dendrocerus carpenteri; (41) D. ramicornis; (42) D.
liebscheri; (43) D. serricornis; (44) D.pupparum; (45) D. halidayi; (46) D. dubiosus. 47, 48, antennae
of males of (47) D. laticeps; (48) D. laevis.
BRITISH SPECIES OF DENDROCERUS
307
Figs 49-55 Antennae of males of (49) Dendrocerus dubiosus; (50) D. aphidum; (5 1)D. carpenter; (52)
D. liebscheri; (53) D. serricornis; (54) D. pupparum; (55) D. btfbveatus.
308
N. D. M. FERGUSSON
Figs 56-63 Antennae of females of (56) Dendrocerus liebscheri; (57) D. carpenteri; (58) D. dubiosus;
(59) D. aphidum; (60) D. laticeps; (61) D. laevis; (62) D.flavipes; (63) D. halidayi.
BRITISH SPECIES OF DENDROCERUS 309
Doubtfully placed species
Kieffer described the following species from specimens in the L. Carpentier collection: Lygocerus
antennalis Kieffer, 1907:46; L. antennalis subserratus Kieffer, 1907:62 and L. subtruncatus
Kieffer, 1907:44. The type-material cannot be found amongst the Carpentier material in the MP,
Amiens and the identity of these species cannot be ascertained from the descriptions alone.
The type ofCeraphron rosularum Ratzeburg is lost. Dessart(1972a:262) and Boucek (1964: 664)
have been unable to recognize the type from the mixed series of three specimens (one D. serricornis
and two D. carpenterf) remaining under this name in the Ratzeburg collection.
Tshumakova (1956: 1 14) described Lygocerus dauricus from the U.S.S.R. The type of this species
has not been located.
Acknowledgements
I am pleased to register my thanks to Mr T. Huddleston for his advice and criticism, to Dr V. F.
Eastop for identifying aphid mummies, and to Mr A. Sutton for drawing figures 1-8, 11, 12, 18-20
and 39. I would also like to thank Dr H. Takada and Dr H. H. Evenhuis for their gifts of bred
material.
I am grateful to the following persons for providing loans of specimens: Dr V. N. Alekseev
(ZMMLSU, Moscow); Dr C. Besuchet (MHN, Geneva); Mr A. Brindle (MM, Manchester); Dr V.
Chambers; Dr T. H. Chua (UM, Kuala Lumpur); Mr R. Danielsson (ZI, Lund); Dr P. Dessart
(IRSNB, Brussels); Dr F. N. Dingemans-Bakels (NHM, Maastricht); Dr H. H. Evenhuis (IPO,
Wageningen); Dr B. Gustafsson (NR, Stockholm); Dr K. J. Hedqvist (NR, Stockholm); Dr W.
Hellen (ZMU, Helsinki); Dr M. G. Jones (RES, Harpenden); Dr P. I. Persson (NR, Stockholm);
Mme S. Kelner Pillault (MNHN, Paris); Dr E. Konigsmann (MNHU, Berlin); Mrs S. Mascherini
(MZU, Florence); Dr P. M. Marsh (USNM, Washington); Prof. H. Morge (IP, Eberswalde); Dr A.
Neboiss (NM, Victoria); Dr J. P. O'Connor (NMI, Dublin); Dr G. Osella (MCSN, Verona); Mr C.
O'Toole (UM, Oxford); Dr R. Poggi (MCSN, Genoa); Dr G. Remaudiere (IP, Paris); Prof. G. C.
Varley (UM, Oxford) and Dr I. D. Wallace (MCM, Liverpool).
References
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1 904. Descriptions of new Hymenoptera from Japan - 1 . Jl. N. Y. ent. Soc. 12: 65-84.
Bankowska, R. 1975. Aphid-aphidophage community in alfalfa cultures (Medicago sativa L) in Poland. Pt. 1
Structure and phenology of the community. Annls zool. Warsz. 32: 299-345.
Bin, F. 1977. Preliminary report on a new structure in Trissolcus Ashm. (Hymenoptera, Scelionidae). Redia
60: 453-456.
Blanchard, M. E. 1840. Histoire naturelle des insectes 3: 672 pp. Paris.
Boheman, C. H. 1832. Forsok till beskrifning af de i Sverige funne Arter, horande till Insekt-slagtet
Ceraphron. K. svenska VetenskAkad. Handl. 1831: 322-339.
Boness, M. 1958. Biocoenotische Untersuchungen iiber die Tierwelt von Klee- und Luzernefeldern. Z. Morph.
'Okol.Tiere 47: 309-373.
Boucek, Z. 1964. Proctotrupoidea und Chalcidoidea aus den Resten der Ratzeburg-Sammlung. Beitr. Ent.
14: 663-673.
Buckton, G. B. 1876. Monograph of the British aphides 1: 193 pp., 38 pis. London.
Cameron, P. 1907. On the Scottish species of Oxyura (Proctotrypidae) - Part 2. Ann. Scot. nat. Hist. 63:
158-161.
Campbell, A. & Mackauer, M. 1975. Thermal constants for development of the pea aphid (Homoptera:
Aphididae) and some of its parasites. Can. Ent. 107: 419^423.
Chua, T. H. 1977. Population studies of Brevicoryne brassicae its parasites and hyperparasites in England.
Res.pop.Ecol. 19: 125-139.
310 N. D. M. FERGUSSON
Couchman, J. R. 1977. A collection of aphid parasites and hyperparasites (Hym.) from Iceland. Norw. J.
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Cumber, R. A. & Eyles, A. C. 1961. Insects associated with the major fodder crops in the North Island. IV
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Curtis, J. 1829a. British entomology. 6: folios 242-289. London.
18296. A guide to an arrangement of British insects 256 pp. London.
1860. Farm insects. 528 pp. London.
Dalla Torre, C. G. 1898. Catalogus Hymenopterorum 5: Chalcididae et Proctotrupidae. 598 pp.
Dean, G. J. 1974. Effects of parasites and predators on the cereal aphids Metopolophium dirhodum and
Macrosiphum avenae (Hem., Aphididae). Bull. ent. Res. 63: 41 1-422.
Delucchi, V. & Pschorn-Walcher, H. 1954. Cremifania nigrocellulata Czerny (Diptera, ?Chamaemyiidae),
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Dessart, P. 1965a. Contribution a Petude des Hymenopteres Proctotrupoidea (IX). Revision du genre
Macrostigma Rondani 1877 (Ceraphronidae, Megaspilinae). Redia 49: 157-163.
19656. Contribution a Petude des Hymenopteres Proctotrupoidea (VI). Les Ceraphroninae et Quelques
Megaspilinae (Ceraphronidae) du Musee Civique d'Histoire Naturelle de Genes. Bull. Annls Soc. r. ent.
Belg. 101: 105-192.
1966. Contribution a Petude des Hymenopteres Proctotrupoidea (XII). A propos des Ceraphronidae
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Index
Page numbers of principal references are in bold. Synonyms and unavailable names are in italics.
africanus 259 aphidorum26l,266
alpestris 262, 276 aphidum 259, 261, 262, 263, 264, 265, 266, 270,
ambianus262,21l 271, 276, 277, 293, 294, 295, 296, 297, 299,
ancyloneurus 262, 287 305, 307, 308
antennalis 309 applanatus 262, 28 1, 282
aphidivorus 262, 270 attentus 300
314
N. D. M. FERGUSSON
basalis 288
bicolor 261, 266, 267, 276, 277
bifoveatus 260, 261, 264, 269, 294, 296, 299, 305,
307
breadalbimensis 266, 267
britannicus 262, 271
callicerus 262, 279
cameroni 262, 270, 272
campestris 262, 270
carpenteri 260, 261, 264, 266, 270, 293, 294, 295,
296, 297, 298, 299, 303, 304, 306, 307, 308
chilocori 300
claripennis 262, 276
coccophagus 259, 262, 281
conwentziae 300
crispus 26 1,270, 271
damicornis 262, 279
dauricus 309
dubiosus 262, 263, 264, 276, 293, 294, 295, 296,
297, 298, 299, 306, 307, 308
dubitatus 259
elegans 26 1,270, 271
ergensis300, 301
flavipes 262, 263, 264, 276, 278, 281, 297, 299,
304, 305, 308
flavus 262, 276
floridanus 300
frenalis 26 1,266, 267
frontalis 262, 28 1,282
fuscipennis 261, 266
fuscipes 262, 270, 271, 278, 279
fusciventris 261, 266, 267
giraudi 262, 270
glabriculus 262, 288
halidayi 256, 259, 261, 262, 263, 264, 279, 297,
298, 299, 304, 306, 308
henkvlugi 300
26 1,270
incompletus 300
indicus 279, 300
inquilinus 262, 271
japonicus 262, 288
koebelei 26 1,266, 271
koyamai259,261,300, 301
laevis 259, 262, 263, 279, 281, 293, 294, 295, 297,
298, 304, 305, 306, 308
lapponicus 262, 290
laticeps 262, 263, 264, 279, 283, 293, 294, 295,
296, 297, 298, 299, 303, 304, 305, 306, 308
latifrons 300
leucopidis 300
levis 28 1
lichtensteinii 259, 262, 279, 280
liebscheri 262, 263, 264, 266, 271, 285, 293, 295,
296, 299, 303, 304, 306, 307, 308
longicornis 262, 276
lundensis 261,266
navaensis 262, 276
neglectus 261, 266
niger 262, 270
noumeae 300
omostenus 301
pallipes 262, 276
parvulus 262, 286
piceus 262, 290
pinicolus 300
propodealis 300
proximus 262, 270
punctatipennis 262, 270, 271
punctipes 258, 259, 261, 262, 263, 265, 286, 299
puparum 287
pupparum 259, 260, 262, 264, 265, 287, 298, 299,
301,302,304,306,307
ramicornis 256, 259, 260, 261, 262, 263, 265, 288,
293, 295, 296, 298, 299, 302, 304, 305, 306
ratzeburgi 259, 262, 288
rectangularis 262, 276
remaudierei 300, 301
rosularum 309
rufipes 26 1,266, 267
rufiventris 262, 270
semiramosus 262, 290
serraticornis 290
serricornis 260, 262, 263, 265, 267, 290, 295, 297,
298, 299, 306, 307
smirnoffi 262, 281
solarii 301
sordidipes 261,269
spissicornis 258, 260, 261, 262, 263, 264, 292,
299, 303
subquadratus 261, 266, 267
subramosus 262, 290
subtruncatus 309
syrphidarum 262, 287
tenuicornis 262, 285
testaceimanus 262, 270, 290
thomsoni262, 271
tischbeini 262, 271
triticum 300
wollastoni 287
zetterstedti 262, 290
British Museum (Natural History)
Publications
Catalogue of the Diptera
of the Afrotropical Region
Editor: R. W. Crosskey. Assistant editors: B. H. Cogan,
P. Freeman, A. C. Pont, K. G. V. Smith and the late H. Oldroyd.
British Museum (Natural History). 1980, 1437 pp. £55-00
The Diptera or two-winged flies are probably the most important
insects that affect man. Although most flies are harmless, some
have become transmitters of dangerous diseases to man and his
domestic animals, and others are important pests of agricultural
crops. Some flies are beneficial because they destroy large numbers
of plant-feeding insects through their parasitic or predacious habits.
Nowhere is their socio-economic and medical impact more sharply
felt than in tropical Africa, where fly-borne diseases are not only a
direct health hazard but can prevent or hinder development of the
land. The control of such diseases as sleeping sickness and
onchocerciasis depends in great measure upon controlling the flies
that carry them. This in turn requires a thorough appreciation of
all that is known about the insect vectors, including their basic
taxonomy, so that they can be correctly identified and their
geographical ranges accurately established.
This catalogue synthesizes the scattered basic taxonomic work on
the Diptera of tropical Africa and its islands by listing the known
16,500 species with their synonyms and known geographical
ranges within a comprehensive classification. A short introduction
is given to each family and a bibliography of 4,700 titles provides
references to the primary literature. Such a task has never before
been attempted for the region and its completion should greatly
stimulate taxonomic research. The Catalogue represents ten years'
careful work by a team of forty specialists, under the editorship of
six dipterists on the staff of the Natural History Museum,
themselves contributors with considerable expertise in the African fauna.
The Catalogue should serve for a long time as an indispensable tool
to the taxonomist and an essential source-work to anyone
concerned with African flies in the fields of medical, agricultural
and veterinary science.
Titles to be published in Volume 41
A revision of the western Palaearctic species of the genus Meteorus
(Hymenoptera : Braconidae). By T. Huddleston.
A revision of the Old World Polymorphanisini (Trichoptera : Hydropsychidae).
By P. C. Barnard.
A review of the genera of Neotropical Encyrtidae (Hymenoptera :
Chalcidoidea). By John S. Noyes.
A revision of the British species of Dendrocerus Ratzeburg (Hymenoptera
Ceraphronoidea) with a review of their biology as aphid hyperparasites.
By N. D. M. Fergusson.
A revision of the Pacific species of Conocephalus Thunberg (Orthoptera :
Tettigoniidae). By Linda M. Pitkin.
Printed by The Whitefriars Press Ltd, London and Tonbridge
Bulletin of the
British Museum (Natural Histo
A revision of the Pacific species of
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Linda M. Pitkin
Entomology series
Vol 41 No 5 18 December 1980
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©Trustees of the British Museum (Natural History), 1980
This number completes Volume 41
ISSN 0524-6431 Entomology series
Vol 41 No 5 pp 315-355
British Museum (Natural History)
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London SW7 5BD Issued 18 December 1980
A revision of the Pacific species of Conocephalus
Thunberg (Orthoptera: Tettigoniidae)
Linda M. Pitkin K
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Contents
Synopsis . . . . . . . . . . . . .315
Introduction . . . . . . . . . . . .315
Acknowledgements . . . . . . . . . . .316
Economic importance . . . . . . . . . . .316
Biogeography . . . . . . . . . . . .317
Material 319
Methods 320
Conocephalus Thunberg .......... 320
Checklist of the Pacific species and subspecies of Conocephalus . . . 323
Key to the Pacific species of Conocephalus 323
Descriptions of the Pacific species ........ 329
References . . . . . . . . . . . 353
Index -354
Synopsis
The species of Conocephalus Thunberg occurring on the Pacific islands are revised and three new species are
described. Keys are given to the eighteen species recognized, and to their subspecies and forms. Eleven
specific and subspecific synonyms are newly established, and a new name is proposed for a junior homonym.
Accounts are given of the economic importance and biogeography. The songs of two of the species are
described for the first time.
Introduction
I undertook this study as the result of an enquiry about the identity of Conocephalus species that
had been found to be potentially useful in the biological control of a rice pest in Papua New Guinea
(see p. 316). It soon became apparent that the specimens sent from Papua New Guinea could not
be reliably identified until all the species of the genus occurring in the South Pacific had been
revised. The geographical area covered by the study comprises the Pacific islands east of the
Philippines and the Moluccas, extending northwards to the Tropic of Cancer and eastwards to the
Marquesas Is. The countries and regions in this area from which I have examined material are
listed in Table 1 (p. 318). Australia has a large number of endemic species, which I have excluded
as being beyond the scope of the present study. I have examined 2150 Pacific specimens and many
specimens from outside the Pacific area.
The genus Conocephalus was erected by Thunberg (1815) for 24 species; one of these, Gryllus
Tettigonia Conocephalus L., is regarded as the type-species by tautonymy. Many of the other
species are now placed in the Copiphorinae. Conocephalus at present comprises approximately 100
valid named species in addition to the eighteen dealt with in the present study.
The prevalence of brachypterous and macropterous forms of the same species in Conocephalus
has been misinterpreted by many earlier authors, who often regarded them as separate species. This
source of confusion is evident in Redtenbacher's (1891) and Karny's (1907) revisions of the genus.
Since 1907 there has been no comprehensive revision of Conocephalus, although partial revisions
Bull. Br. Mas. not. Hist. (Ent.) 41 (5): 315-355 Issued 18 December, 1980
>r\ N f~\ ^ .
316 LINDA M.PITKIN
have covered the Neotropical (Rehn & Hebard, 1915a; 19156) and Palaearctic (Harz, 1969)
species, and new species have been described by various authors. Kaltenbach (1968) and Hudson
(1972) have dealt with the species of New Caledonia and New Zealand, respectively, but these
studies cover only a few of the species occurring in the Pacific; most of the remaining species have
not been discussed in the literature since their original description.
There is one New Guinea species of Conocephalus which I have been unable to include in this
study. This is Xiphidion consul Karny, 191 1: 344 (type-locality: Sattelburg), the holotype of which
is lost (pers. comm. from Dr A. Kaltenbach of the Naturhistorisches Museum, Vienna). Karny's
description does not entirely agree with any of the specimens I have examined, and no further
.descriptive information on the species has been published by subsequent authors. I am therefore
unable to determine the identity of X. consul Karny and consider it to be a nomen dubium.
Karny (1926: 181) records a specimen of C. melaenus (de Haan) from New Guinea. I consider
this is likely to have been misidentified, and I have not included this species in the present study
since its distribution is entirely extralimital to judge from the numerous specimens which I have
examined.
A number of subgenera have been described for the species of Conocephalus. Some of these
consist of only one or two species and some others are based on rather unreliable characters such
as the presence or absence of certain hind tibial spurs. I have found this character to be useful at
the specific level in the species I have studied, but even so it is subject to intraspecific variation.
Two of the three new species described in this paper are intermediate between two subgenera on the
basis of this character, and I have therefore decided not to assign any species to subgenera. (See
also p. 320.)
Acknowledgements
I would like to thank the following specialists who have lent me material for this study from their
respective institutions:
Dr C. Baroni Urbani, Dr P. H. van Doesburg, Dr M. Donskoff, Dr C. E. Dunn, Dr K. K.
Gunther, Dr B. Hauser, Dr A. Kaltenbach, Dr T. Kronestedt, Dr G. M. Nishida, Dr D. C. F.
Rentz, Dr J. N. L. Stibick, Miss A. K. Walker, and Dr F. Willemse.
I also particularly wish to thank Dr D. R. Ragge for his most helpful advice and criticism in the
course of this study; Dr M. H. Robinson for sending me live specimens, some of which were used
for making tape recordings of the songs; Mr W. J. Reynolds for invaluable assistance in making the
song recordings and preparing the oscillograms used in this study, and for testing the identification
keys; and Mr G. Young for information on the biology of some Conocephalus species.
Economic importance
Two species of Conocephalus, subsequently identified by me as C. redtenbacheri (Bolivar) and C.
semivittatus vittatus (Redtenbacher), have been found to be important predators of eggs and
nymphs of a serious rice pest, Leptocorisa oratorius (F.) (Hemiptera: Alydidae), in Papua New
Guinea. According to Sands (1977) these species do not appear to damage the developing rice
grains or eat any part of the rice plant apart from feeding to some extent on the stamens (together
with other grass flowers) and should therefore be considered distinctly beneficial. In New Guinea,
C. redtenbacheri is by far the more common of the two, judging from the specimens I have
examined. Both species have also been recorded as preying on eggs of other Hemipteran pests:
Riptortus annulicornis Bois (Alydidae) and Nezara viridula L. (Pentatomidae) (pers. comm. from
Dr G. Young of the Department of Primary Industry, Konedobu).
C. saltator (Saussure) has also been reported as a predator of pests, by Zimmerman (1948) and
Swezey (1905, under the synonym Xiphidium varipenne Swezey). It has been found in abundance
in the sugar cane fields of the Hawaiian Is., feeding largely on young and adult sugar cane
leafhoppers, Perkinsiella saccharicida Kirkaldy; the sugar cane bud moth, Decadarchis flavistriata
(Walsingham) (formerly in Ereunetis), and other caterpillars; the coccid Dymisococcus brevipes
(Cockerell) (formerly in Pseudococcus); and dipterous larvae. Swezey states that there is very little
evidence of this species eating cane, although it will eat pollen from the blossoms of Canna,
PACIFIC SPECIES OF Conocephalus 3 1 7
Lantana and other plants. C. saltator is also considered to be a valuable predator of pineapple
mealy bugs, and of the caterpillars and pupae of various bud moths which are pests of pineapples
(see Illingworth, 1929: 256).
Although C. saltator appears to be chiefly beneficial, it has been reported as causing some
damage to rice in the West Indies (Grist & Lever, 1969: 293). In the Pacific, it has been known to
do so only occasionally. It has also caused occasional damage to corn and pineapples; in the latter
case the damage is caused by laying its eggs in the flowers of the young fruit (Zimmerman, 1948).
C. longipennis (de Haan) and an unidentified Conocephalus species were recorded as feeding on
nymphs and probably eggs of the rice ear bug, Leptocorisa oratorius (F.), in Sarawak by
Rothschild (1970). He states, however, that in these particular species the beneficial effect is
probably largely nullified by their habit of feeding on developing rice grains. In fact, C. longipennis
has been recorded as a minor pest of rice in Sarawak and New Guinea by Grist & Lever
(1969: 293), although I have not seen any specimens, or other record, of the species from New
Guinea. It is common on rice in W. Java, where a specimen has been observed eating an unidenti-
fied insect (Kalshoven & van der Vecht, 1950: 137). This species also occurs on rice in Malaya
(Yunus, 1967), the Philippines, India and Bangladesh (specimens examined). I have seen speci-
mens of C. oceanicus (Le Guillou), also collected from rice in the Philippines.
Various Conocephalus species, not included in the present study, have been recorded as pests of
rice by Grist & Lever (1969:292-293). These are C. cinereus (Thunberg) in Surinam, C.
propinquus (Redtenbacher) in Guyana and Surinam, and C.fasciatus (De Geer) in Texas, U.S.A.
This last species is known to be insectivorous as well (Marshall, 1964). I have seen specimens of C.
Conocephalus (L.), collected from rice in West Africa, where it feeds on some plant material and
probably also eats the immature stages of a rice pest, Aleurocybotus ? indicus David (Hemiptera:
Aleyrodidae) (pers. comm. from Mr J. C. Deeming).
It would therefore appear that Conocephalus species are often both predacious and herbivorous.
Where they occur on plants of economic importance, species which are predominantly predacious
can be very beneficial, while predominantly herbivorous species may be pests.
Biogeography
The species of Conocephalus in the Pacific can be divided into two groups on the basis of two quite
different male characters. In one group the species have cerci with one internal spine and a small
structure slightly basad of this as shown in Figs 2-11, hereafter referred to as a tubercle. They also
have a stridulatory file with the row of teeth differentiated into large, well-spaced teeth at one end
and tiny dense teeth at the other end (Figs 45, 46). In the other group the species have one, or in
one case two, internal spines on the cerci but no tubercle; the stridulatory file is shaped as in Fig.
47, with comparatively uniform density and size of teeth. I am including C. upoluensis (Karny) and
C. albescens (Walker) in the first group on the basis of their stridulatory file shape, although the
tubercle of the cerci of these two species is extremely small or sometimes absent. One species, C.
tridens Hebard, is intermediate between the two groups in the stridulatory file structure and the
cerci are unique in the genus in having three internal spines. This species is endemic in the
Marquesas Is. and does not seem to be related to any of the other Pacific species.
The species of the first group are known only from the Pacific and nearby islands such as the
Moluccas (with the exception of a record of C. oceanicus from Vietnam) but most of the species in
the second group have distributions extending well outside the Pacific, in several cases in Asia, and
in two species also in tropical Africa, although they show no evidence of southern Gondwanaland
relationships. One of the species in the second group, C. saltator, is known to have been introduced
to the Pacific, and I think it is likely that most of the species in this group originated outside the
Pacific since they lack characters shared by the species known only from the Pacific.
The distribution patterns of the Conocephalus species in the Pacific (Table 1) suggest that their
dispersal has been largely achieved by flight. Most of the species that are at least sometimes
macropterous occur on two or more islands and some are widespread in the Pacific. One of these
species, C. angustivertex sp. n., is recorded only from New Guinea, but C. bispinatus sp. n. also
occurs outside the Pacific, and C. tridens, although restricted to the Marquesas, occurs on several
318 LINDA M. PITKIN
Table 1 The distribution of the species of Conocephalus in the Pacific
Hawaiian Is. +
Marquesas Is. +
Tubuai Is. +
Society Is. + +
Cook Is. + + +
Line Is. +
Tonga Is. + +
American Samoa + + + +
Western Samoa + + +
Fiji Is. + +
Ellice Is. +
Caroline Is. ^
New Zealand + + +
Kermadec Is. +
Norfolk I. +
Lord Howe I. + +
Loyalty Is.
New Caledonia + +
New Hebrides + + +
Solomon Is. + + + +
New Britain; New Ireland + + + +
Admiralty Is. + +
Southern Australia + + + +
Northern Australia + + +
Papua New Guinea + + + + + + + +
Irian Jaya + + + + + +
2
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1 § iti 1 1 i 1 1 1 i i H ^ ^^
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PACIFIC SPECIES OF Conocephalus 319
islands in this group. On the other hand, each of the three species known solely from brachypterous
specimens, C. starmuehlneri Kaltenbach, C. tumidus sp. n. and C. trivittatus (Stal), is recorded
from only one island. Some dispersal is still taking place, for example the spread of C. saltator to
the Line Is., Samoa and the Cook Is., following its introduction to the Hawaiian Is. from America
in c. 1890.
The distribution patterns of Pacific Conocephalus species do not support a rigid system of
biogeographical division of the Pacific. They show only slight evidence for the two Melanesian arcs
discussed by Ross (1956), Hennig (1966) and other authors. C. upoluensis has an outer Melanesian
arc distribution pattern in that it occurs from New Guinea to the Solomon Is., the New Hebrides,
Fiji and Samoa, but it also occurs in New Caledonia and elsewhere in the Pacific. C. redtenbacheri
and C. semivittatus vittatus occur from New Guinea to the Solomon Is. and to the New Hebrides
respectively, but do not extend further. None of the species have the inner Melanesian arc
distribution linking New Guinea, New Caledonia and New Zealand.
However, the distributions of the Conocephalus species do support the existence of some points
of partial discontinuity between certain islands or island groups. Lee (1975) and other authors
consider there to be a distinction between the biota of the Solomon Is. and the New Hebrides. This
is supported in the present study by C. redtenbacheri and C. infumatus (Redtenbacher) which
reach the eastern limit of their distribution in the Solomon Is. (including the Santa Cruz Is.) and C.
oceanicus which occurs in the New Hebrides but has not been found in the Solomon Is. The present
study corroborates the biogeographical inclusion of the Santa Cruz Is. in the Solomon Is. C.
upoluensis occurs in both the Solomon Is. and the New Hebrides; however, this species is very
widespread in the Pacific. C. semivittatus vittatus also occurs in both these island groups, but
although it occurs in most of the northern islands of the New Hebrides, it has not been found in
Erromango or the other islands south of Efate. This tends to corroborate Lee's suggestion of a
major disjunction between Efate and Erromango separating the northern and southern New
Hebrides.
There is also some differentiation between the Conocephalus occurring in northern Australia
(Northern Territory and Queensland) and southern Australia; all except the widespread C.
upoluensis are restricted to one or other region of Australia. This shows agreement with Gressitt
(1961) and Thorne (1963). All the Conocephalus species found in northern Australia also occur in
New Guinea. The species in southern Australia, other than C. upoluensis, also occur in New
Zealand.
New Caledonia has been noted as having a high degree of endemism by authors including van
Balgooy (1971) and Thorne (1963). In the present study one species, C. starmuehlneri Kaltenbach,
is endemic in New Caledonia and probably arose from the ancestry of its close relative C.
oceanicus, whose distribution includes New Caledonia.
Material
The material examined in this study is deposited in the following institutions.
BMNH British Museum (Natural History), London
DSIR, Auckland Department of Scientific and Industrial Research, Auckland
NM, Basle Naturhistorisches Museum, Basle
MNHU, Berlin Museum fur Naturkunde der Humboldt-Universitat, Berlin
ANIC, Canberra Australian National Insect Collection, Canberra
MHN, Geneva Museum d'Histoire Naturelle, Geneva
BPBM, Honolulu Bernice P. Bishop Museum, Honolulu
HSDA, Honolulu Hawaii State Department of Agriculture, Honolulu
DPI, Konedobu Department of Primary Industry, Konedobu, Papua New Guinea
RNH, Leiden Rijksmuseum van Natuurlijke Historic, Leiden
NM, Maastricht Natuurhistorisch Museum, Maastricht
MNHN, Paris Museum National d'Histoire Naturelle, Paris
ANS, Philadelphia Academy of Natural Sciences of Philadelphia
NR, Stockholm Naturhistoriska Riksmuseet, Stockholm
NM, Vienna Naturhistorisches Museum, Vienna
320 LINDA M. PITKIN
Methods
All the morphological data were taken from dried adult specimens. The measurements given for all
the previously described species were taken from Pacific specimens only (see Table 1 for summary
of provenances), although specimens from the complete ranges were examined, when available, for
other characters. In most species all the available Pacific specimens were measured; in the rest the
measurements were taken from samples of at least 20 specimens of each sex of every species,
subspecies or form, selected to cover a wide range of localities. Primary types from outside the
Pacific were measured, although not included in the samples. Any measurements of these types
outside the range given for any character has been noted under the relevant species.
Vernier callipers were used for all the measurements except the width of the fastigium, the male
cereal length and the length of the stridulatory area, which were measured using a microscope with
a moving stage vernier micrometer. Forewing length was measured on the flexed left wing, from
where the pronotum overlaps the veins Sc and R to the apex. Ovipositor length was measured
from the apex of the subgenital plate to the apex of the ovipositor. The length of the stridulatory
area was measured as indicated in Fig. 39; male cereal length was measured on the right cercus,
viewed laterally. Fastigium width was measured at the apex of the fastigium of the vertex, viewed
dorsally. All the measurements listed are given in millimetres and the number of specimens
measured is given in parentheses.
Pyroxylin replicas were made in order to examine the stridulatory structures of the under surface
of the left male forewing, following the method described by Ragge (1969: 172). Although it has
proved extremely useful in some recent taxonomic studies of Tettigoniidae, the stridulatory file was
of no help in the present study in distinguishing closely related species; however, the presence or
absence of spinules around the file was sometimes useful.
The drawings of stridulatory files and whole forewings were prepared by tracing the projected
image from a microprojector and a photographic enlarger respectively. A camera lucida
attachment to a microscope was used for all the other drawings.
In this study the term 'macropterous' refers to specimens in which the hind wings extend beyond
the forewings (when both pairs of wings are flexed); 'brachypterous' refers to those in which the
hindwings are shorter than the forewings. The wing-venation terminology was taken from Ragge
(1955).
All the distribution records are based on material I have examined, to avoid the possibility of
including incorrect records due to mistaken identity. For most of the species I have been able to
examine large numbers of specimens and in these cases I have abbreviated the data listed under
'Material examined', omitting collectors' names and abbreviating the dates to the months only.
Where the species has been previously recorded from the country or island group concerned, I have
given only the name of the country (or its major subdivision) or island without further details. For
new records of a species from a country or island group I have given the locality data in full, except
for sometimes abbreviating repeated localities to their initial letters. The data of type-material is
always given in full. The term 'Island' is omitted following individual island names listed after an
island group except in cases where this omission might lead to confusion.
For material from outside the main area studied I have listed only the countries or islands, or
sometimes only the zoogeographical region. The data of primary types from extralimital localities
is, however, always given in full.
Studio recordings were made of the songs of two of the species included in this study, by Mr W.
J. Reynolds, Dr D. R. Ragge and myself, using a Kudelski Nagra IVD tape recorder and
Sennheiser MKH 405 microphone. The oscillograms shown in Figs 48 and 49 were made from
these recordings using a Mingograf 34T.
CONOCEPHALUS Thunberg
Conocephalus Thunberg, 1815:214. Type-species: Gryllus Tettigonia conocephalus L., by tautonymy.
Ten subgenera of Conocephalus are currently regarded as valid by at least some authors. Of these
only the following three have had any of the species included in this study assigned to them. (See
also remarks on p. 316.)
PACIFIC SPECIES OF Conocephalus 321
Subgenus CONOCEPHALUS Thunberg, 1815:214.
Hebard (1933) states in his description of C. tridens that the species may be referable to this subgenus.
Subgenus ANISOPTERA Latreille, 1827:409. Type-species: Locusta dorsalis Latreille, by subsequent
designation (Kirby, 1906: 274).
Xiphidion Serville, 1831: 159. Type-species: Locusta fusca F., by subsequent designation (Kirby,
1906: 274). [Synonymized by Kirby, 1906: 274.]
Xiphidium Burmeister, [1838]: 707. [Unjustified emendation.]
Neoxiphidion Karny, 1912: 8. Type-species: Locusta fasciata De Geer, by subsequent designation (Rehn
& Hebard, 1915a: 157). [Synonymized by Rehn & Hebard, 1915a: 170.]
Thecoxiphidion Karny, 1912: 12. Type-species: Xiphidium strictum Scudder, by subsequent designation
(Rehn & Hebard, 1915a: 157). [Synonymized by Rehn & Hebard, 1915a: 170.]
Species included in the present study which have been assigned to the subgenus Anisoptera (or its junior
synonyms) by previous authors: C. oceanicus (Le Guillou): C. starmuehlneri Kaltenbach; C. semivittatus
(Walker); C. bilineatus (Erichson)1; C. maculatus (Le Guillou); C. laetus (Redtenbacher); C. saltator
(Saussure); C. longipennis (de Haan).
Subgenus CHLOROXIPHIDION Hebard, 1922:242. Type-species: Xiphidium javanicum Redtenbacher, by
original designation.
Only one of the species included in the present study has been assigned to this subgenus by previous authors:
C. upoluensis (Karny).
GENERIC DIAGNOSIS. <$ 9- Small to medium-sized Conocephalinae. Pronotum not produced backwards to
cover stridulatory file. Prosternum unarmed or bispinose. Macropterous or brachypterous. Stridulatory area
of left male forewing comparatively small. Fore and mid tibiae with 4-12 external (and similar number of
internal) ventral spurs (usually 6). Mid femora usually unarmed, but occasionally (C. bispinatus) with ex-
ternoventral spines. Hind femora unarmed or with externoventral (and occasionally also internoventral)
spines. Male tenth abdominal tergite usually with two median projections at apex; otherwise truncate. Male
cerci with 1—3 internal spines (one species, C. bituberculatum (Redtenbacher), is described as having no
internal cereal spines). Male subgenital plate with wide, usually shallow, V-shaped incision, or truncate.
Ovipositor straight or occasionally curved, of variable length.
DISCUSSION. Apart from Conocephalus, only two Conocephaline genera occur in the Pacific:
Fatuhivella and Nukuhivella, both originally described by Hebard (1933) from the Marquesas Is.,
where they are endemic. These two genera, unlike Conocephalus, have the pronotum produced
backwards in both sexes, so that in the male it covers the stridulatory organ; the males also have
exceptionally specialized cerci.
One of the most striking features of the genus is the variation in wing length. Of the 18 species in
the present study, nine are apparently always macropterous, three are known only from
brachypterous specimens, and the rest have both forms. Of the species in the last category, some
are strictly dimorphic without intermediates, notably C. angustivertex, while others show an almost
continuous range of wing length, as in C. redtenbacheri and C. saltator. In Conocephalus variation
in wing length within a species is in many cases correlated with variation in the size of the
stridulatory area and the size of the cells formed by the forewing venation.
The genus is moderately uniform in general appearance apart from the wing length, but there are
often clear differences between the species. Most of the Pacific species are easily distinguished from
all the others, but there are a few species which form close-knit groups on the basis of
morphological similarity. Two such species are C. upoluensis and C. albescens. A more complex
group consists of C. oceanicus, C. starmuehlneri, C. redtenbacheri and C. infumatus (see under the
discussions of these species). The taxa in this group, including the two forms of one of the species,
show quite a strong tendency to be allopatric (see Fig. 1 and Table 1) and could possibly all be
regarded as semispecies. The subspecies and forms of C. semivittatus (Walker) could also be
semispecies.
DISTRIBUTION (Table 1). Conocephalus has a worldwide distribution between approximately 60 °N
and 45 °S.
'Placed here by Karny (1912:11) but would now be assigned to Chloroxiphidion Hebard.
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PACIFIC SPECIES OF Conocephalus 323
Checklist of the Pacific species and subspecies of Conocephalus
oceanicus (Le Guillou)
ajfine Redtenbacher syn. n.
starmuehlneri Kaltenbach
redtenbacheri (Bolivar)
infumatus (Redtenbacher)
semivittatus semivittatus (Walker)
maoricum Walker
antipodum Scudder
b runner i Karny syn. n.
semivittatus vittatus (Redtenbacher) nom. rev., stat. n.
geniculare Redtenbacher
tumidus sp. n.
trivittatus (StSl)
upoluensis (Karny) nom. rev., stat. n.
modestum Redtenbacher (junior homonym)
brevixiphus Willemse syn. n.
albescens (Walker)
latifrons Redtenbacher syn. n.
bilineatus (Erichson)
immaculatum Karny syn. n.
maculatus (Le Guillou)
lepida de Haan
continuum Walker syn. n.
neglectum Bruner syn. n.
laetus (Redtenbacher)
dubius Willemse syn. n.
raggei Harz syn. n.
angustivertex sp. n.
saltator (Saussure)
meridionale Scudder
propinquum Redtenbacher
brachypterum Redtenbacher
varipenne Swezey
longipennis (de Haan)
spinipes Stal
longicorne Redtenbacher
carolinensis Willemse syn. n.
carolinensis f. macroptera Willemse syn. n.
willemsei nom. n.
ensiferus Willemse (junior homonym)
bispinatus sp. n.
tridens Hebard
Key to the Pacific species of Conocephalus
Measurements given in this key are taken from Pacific specimens and may differ for extralimital material.
1 Prosternum unarmed ............ 2
— Prosternum bispinose ............ 3
2 Hind femora armed ventrally with spines. Male cerci with 2 internal spines (Fig. 21). Ovipositor
straight (Fig. 25) ......... C. bispinatus sp. n. (p. 351)
Hind femora unarmed ventrally. Male cerci with 3 internal spines (Fig. 22). Ovipositor curved
(Fig. 27) C. tridens Hebard (p. 352)
3 Hind femora unarmed ventrally .......... 4
Hind femora with externoventral spines ......... 1 2
4 Forewings unicolorous. Ovipositor at least 13-7 mm long. Stridulatory area of left male forewing
large (at least 2-7 mm long, measured as in Fig. 39) . . C. laetus (Redtenbacher) (p. 345)
Forewings usually with some spots or other markings. If unicolorous, ovipositor not more than
13-2 mm long. Stridulatory area of left male forewing smaller (not more than 2-3 mm long) 5
"7\ (/ ^
a ^ 13^°
V
Figs 2-22 Dorsal view of the left male cercus, and in some cases (a) posterior view of the internal spine
and (b) externolateral view of the cercus of (2) Conocephalus oceanicus; (3) C. starmuehlneri; (4) C.
redtenbacheri; (5) C. infumatus (typical form); (6) C. infumatus (small New Ireland form); (7) C.
semivittatus semivittatus; (8) C. semivittatus vittatus (typical form); (9) C. semivittatus vittatus (small
form); (10) C. tumidus; (1 1) C. trivittatus; (12) C. upoluensis; (13) C. albescens; (14) C. bilineatus (15)
C. maculatus; (16) C. /ae/ws; (17) C. angustivertex; (18) C. saltator, (19) C. longipennis; (20) C.
willemsei; (21) C. bispinatus; (22) C. tridens.
PACIFIC SPECIES OF Conocephalus
325
23
24
25
26
Figs 23-27
Lateral view of the ovipositor of (23) Conocephalus tumidus; (24) C. angustivertex; (25) C.
bispinatus; (26) C. willemsei; (27) C. tridens.
10
11
12
13
14
Forewings with large dark spots; no markings in the costal or precostal areas (Fig. 31)
. C. maculatus (Le Guillou) (p. 344)
Forewings with dark pigmentation in the costal and precostal areas, with or without com-
paratively small spots on the rest of the wing (Figs 29, 30, 32); or unicolorous ... 6
Hind tibiae with 3 pairs of apical spurs (Fig. 40). Forewings unicolorous, or, if with dark
markings, cross-veins of the costal and precostal forewing areas regular, parallel . . 7
Hind tibiae usually with 2 pairs of apical spurs (ventral pair missing). If with 3 pairs of spurs:
forewings with dark markings at least in the costal and precostal areas, although occasionally
very faint; cross-veins of the costal and precostal forewing areas irregular, often reduced 8
Forewings unicolorous. Male cerci shaped as in Fig. 18, with one internal spine
C. saltator (Saussure) (p. 347)
Forewings with some darker pigmentation in the costal and precostal areas, as in Fig. 29,
although sometimes faint. Male cerci shaped as in Fig. 11, with one internal spine and one
tubercle C. trivittatus (Stal) (p. 340)
Brachypterous; forewing length not more than 6-1 mm ...... 9
Macropterous or brachypterous; forewing length at least 8 -Omm ..... 10
Male cerci with one internal spine and one tubercle (Fig. 10). Ovipositor length not more than
8-5 mm C. tumidus sp. n. (p. 339)
Male cerci with one internal spine (Fig. 14). Ovipositor length at least 10-5 mm
C. bilineatus (Erichson) (p. 343)
Fastigium of the vertex relatively narrow (at most 0-57 mm). Forewings with dark pigmentation
only in the costal and precostal areas, as in Fig. 29. Male cerci shaped as in Fig. 14
C. bilineatus (Erichson) (p. 343)
Fastigium of the vertex relatively wide (at least 0-59 mm). Forewings with dark pigmentation in
the costal and precostal areas, and usually with some small spots on the rest of the wing (Fig.
30). Male cerci shaped as in Figs 12, 13 11
Male cerci very slender in the apical part (Fig. 13). Male tenth abdominal tergite produced
perpendicularly downwards (Fig. 42). Ovipositor length at least 11-8 mm
C. albescens (Walker) (p. 342)
Male cerci less slender in the apical part (Fig. 12). Male tenth abdominal tergite nearly
unmodified, or slightly produced at apex but not bent downwards (Fig. 41). Ovipositor length
not more than 11 -Omm C. upoluensis (Karny) (p. 341)
Males
Females .........
Cerci with one internal spine (Figs 17-20) ....
Cerci with one internal spine and one tubercle (Figs 2-10)
Forewings unicolorous. Hind tibiae with 6 apical spurs (Fig. 40) . 15
Forewings with dark pigmentation as in Fig. 32. Hind tibiae usually with 5 apical spurs
(internodorsal spur missing; very rarely a small 6th spur is present)
C. angustivertex sp. n. (p. 346)
326 LINDA M. PITKIN
15 Cerci shaped as in Fig. 18, with internal spine near base. Width of fastigium of the vertex at least
0-42 mm ......... C. saltator (Saussure) (p. 347)
Cerci shaped as in Figs 19, 20, with internal spine in the middle or towards the apex. Width of
fastigium of the vertex not more than 0-3 7 mm . . .• . . . . 16
16 Internal cereal spine with a globular apex. Cerci comparatively slender, shaped as in Fig. 19
. C. longipennis (de Haan) (p. 349)
Internal cereal spine with a pointed apex. Cerci usually comparatively stout, shaped as in Fig. 20
. C. willemsei nom. n. (p. 350)
17 Forewings with dark pigmentation in the costal and precostal areas (Figs 28, 29). Fastigium of
the vertex comparatively wide (0-52-0-93 mm) . . . . . . . . 18
Forewings without dark pigmentation in the costal or precostal areas. Fastigium of the vertex
comparatively narrow (0-23-0-53 mm) . . . . . . . . 19
18 Cerci as in Figs 7-9; comparatively straight or only slightly bulging in profile. Hind tibiae with 6
apical spurs ......... C. semivittatus (Walker) (p. 335)
Cerci as in Figs 10, conspicuously bulging in profile. Hind tibiae usually with 4 apical spurs
(ventral pair missing) ........ C. tumidus sp. n. (p. 339)
19 Brachypterous ............. 20
Macropterous ............. 21
20 R and Sc of forewings at most only very slightly darkened; very rarely darkened towards the
base. Forewings comparatively wide, tapering comparatively abruptly towards the apex (Fig.
33). Length of cerci at least 1-8 mm; internal spine with a gradually curved apex as in Fig. 2a
C. starmuehlneri Kaltenbach (p. 330)
R and Sc of forewings usually darkened, at least towards the base. Forewings comparatively
narrow, tapering gradually towards the apex (Fig. 34). Length of cerci not more than 1 • 7 mm;
internal spine with a hook-shaped apex (Fig. 4a) . . C. redtenbacheri (Bolivar) (p. 331)
21 Forewings unicolorous. Internal cereal spine with a gradually curved apex (Fig. 2a). Width of
fastigium of the vertex at least 0-4 mm . . . . C. oceanicus (Le Guillou) (p. 329)
R and Sc of forewings usually darker than the rest of the wings, at least towards the base. If not,
internal cereal spine with a hook-shaped apex (Fig. 4a), or width of fastigium of the vertex not
more than 0-3 mm ............. 22
22 Internal cereal spine with a gradually curved apex (Figs 5 a, 6a). Stridulatory file usually with few
surrounding spinules, as in Fig. 45 . . . . C. infumatus (Redtenbacher) (p. 333)
Internal cereal spine with a hook-shaped apex (Fig. 4a). Stridulatory file surrounded by many
spinules, as in Fig. 46 ....... C. redtenbacheri (Bolivar) (p. 331)
23 Forewings with dark pigmentation at least in the costal and precostal areas as in Figs 28, 29, 32
... 24
Forewings unicolorous or with dark pigmentation only outside the costal and precostal areas 26
24 Ovipositor gently curved (Fig. 24). Hind tibiae with 5 apical spurs (internodorsal spur missing;
very rarely a small 6th spur is present). Width of fastigium of the vertex not more than
0-4 mm ......... C. angustivertex sp. n. (p. 346)
Ovipositor comparatively straight (Fig. 23). Hind tibiae with 3-6 apical spurs. Width of
fastigium of the vertex at least 0-5 mm . . . . . . . . .25
25 Hind tibiae with 6 apical spurs. Width of fastigium of the vertex 0-52-0-83 mm. Subgenital plate
with an excised or truncate apex; if truncate, width of fastigium of the vertex not more than
0-8 1mm c, semivittatus (Walker) (p. 335)
Hind tibiae usually with not more than 4 apical spurs (ventral pair missing or small). Width of
fastigium of the vertex 0-83-0-91 mm. Subgenital plate with a truncate apex
C. tumidus sp. n. (p. 339)
26 Forewing length at least 24 -Omm. Ovipositor comparatively broad as in Fig. 26
. C. willemsei nom. n. (p. 350)
Forewing length not more than 22-5 mm. Ovipositor comparatively narrow, not shaped as in
Fig 26 ' 27
27 R and Sc of forewings usually darkened at least towards the base; if brachypterous, forewings
comparatively narrow, tapering gradually towards the apex (Fig. 34) . ..,.,.. 28
Forewings unicolorous or almost so; R and Sc at most only very slightly darkened, very rarely
towards the base. If R and Sc slightly darkened; brachypterous, forewings comparatively
wide, tapering comparatively abruptly towards the apex (Fig. 33) .... 29
28 Ratio of the pronotum length to the width of fastigium of the vertex at most 8-8 mm.
Macropterous or brachypterous ..... C. redtenbacheri (Bolivar) (p. 331)
327
Figs 28-32 The left female forewing of (28) Conocephalus semivittatus vittatus (typical form)
(brachypterous); (29) C. tumidus; (30) C. upoluensis; (31) C. maculatus; (32) C. angustivertex
(macropterous). Macropterous and brachypterous specimens of the same species have a similar
pigmentation.
— Ratio of the pronotum length to the width of fastigium of the vertex at least 9-0 mm.
Macropterous C. infumatus (Redtenbacher) (p. 333)
29 Brachypterous. Subgenital plate with a slightly excised apex (Fig. 43) ....
C. starmuehlneri Kaltenbach (p. 330)
- Mostly macropterous. If brachypterous, subgenital plate with a truncate apex (Fig. 44) . 30
30 Width of fastigium of the vertex not more than 0-42 mm. Ovipositor length at least 12-5 mm.
Subgenital plate with a widely excised apex . . . C. longipennis (de Haan) (p. 349)
- Width of fastigium of the vertex more than 0-42 mm, or, if less than 0-42 mm, ovipositor length
not more than 11-5 mm. Subgenital plate with a truncate or shallow excised apex . 31
31 MA of forewings shaped as in Fig. 35. Macropterous. Hind femora gradually swollen towards
the base (Fig. 37). Ovipositor length 8-3-11-1 mm. Subgenital plate with a shallow excised
apex .......... C. oceanicus (Le Guillou) (p. 329)
- MA of forewings shaped as in Fig. 36 (macropterous). Macropterous or brachypterous. Hind
femora comparatively sharply swollen towards the base (Fig. 38). Ovipositor length
10-7-13-2 mm. Subgenital plate with a truncate apex (Fig. 44) C. saltator (Saussure) (p. 347)
Positive identification of C. oceanicus, C. starmuehlneri, C. redtenbacheri and C. infumatus may
prove to be very difficult in some cases owing to the variability of some of these species, and the
subtlety of the distinctions between them. Specimens of these species can be identified in many
cases by their geographical data (see under the descriptions of these species).
328
LINDA M. PITKIN
37
38
Figs 33-44 33-36, the left female forewing of (33) Conocephalus starmuehlneri; (34) C. redtenbacheri
(brachypterous); (35) C. oceanicus; (36) C. saltator (macropterous). 37, 38, the left hind femur of (37)
C. oceanicus; (38) C. saltator. 39, dorsal view of the left stridulatory area of C. laetus, indicating the
points between which measurements were made. 40, ventral view of the apex of the left hind tibiae of C.
saltator. 4 1 , 42, posterodorsal view of the male tenth tergite of (4 1) C. upoluensis; (42) C. albescens. 43,
44, ventral view of the female subgenital plate of (43) C. starmuehlneri; (44) C. saltator.
PACIFIC SPECIES OF Conocephalus 329
Descriptions of the Pacific species
Conocephalus oceanicus (Le Guillou)
(Figs 1, 2, 35, 37, 45)
Xiphidion oceanicum Le Guillou, 1841:294. LECTOTYPE tf, SAMOA (MNHN, Paris), here designated
[examined].
Xiphidium affine Redtenbacher, 1891:513. LECTOTYPE $, FIJI Is. (NM, Vienna), here designated
[examined]. Syn. n.
DIAGNOSIS. $ 9- Fastigium of vertex moderately narrow. Prosternum bispinose. Macropterous. Forewings
unicolorous. Cross-veins of costal and precostal areas of forewings mostly regular, parallel. MA of forewings
shaped as in Fig. 35. Stridulatory file of left male forewing shaped as in Fig. 45, with few surrounding
spinules (11 specimens examined). Hind femora not sharply swollen towards base (Fig. 37), with 1-7 ex-
ternoventral spines. Hind tibiae with 6 apical spurs. Male cerci as in Figs 2, 2a, with one well-developed
internal spine with gently curved apex, and one tubercle. Female subgenital plate with shallow excision at
apex. Ovipositor of medium length, relatively straight.
MEASUREMENTS
Males Females
Fastigium width (23): 0-40- 0-53 (0-47) (24): 0-42- 0-58 (0-50)
Median length of pronotum (26): 2-4 - 3-2 (2-94) (29): 2-5 - 3-6 (3-16)
Forewing length (23): 11-5 -18-6 (15-68) (27): 13-4 -20-7(16-80)
Hind femur length (20): 10-3 -13-9 (12-52) (21): 11-1 -15-4(13-54)
Cercus length (22): 1-47- 1-81 (1-65)
Ovipositor length (28): 8-3 -11-1 (9-63)
Pronotum length/fastigium width (22): 5-71-7-62 (6-29) (20): 5-52-7-27(6-30)
Hind femur length/ovipositor length (21): 1-29 -1-52(1-41)
DISCUSSION. C. oceanicus forms a very close group with C. starmuehlneri, C. redtenbacheri and C.
infumatus, differing from the first in wing length, and the others in having unicolorous forewings; it
also differs from C. redtenbacheri in the shape of the internal spine of the male cerci.
The similarity of C. oceanicus and C. redtenbacheri has led to them being confused in the literature.
Redtenbacher's description of Xiphidium affine refers to specimens with unicolorous forewings or
with the radial vein darker, which suggests that his syntype-series contained both species. I have
examined 5 <$, 2 9 from this series, all of which are C. oceanicus, and I have selected a lectotype from
these specimens, thus fixing the identity of X. affine as a synonym of C. oceanicus. The other
syntypes, which I have not seen, have the following data: Fiji: Ovalau. Samoa. Aru Is. (B runner).
I have selected a lectotype of Xiphidion oceanicum from the type-series of 1 <$, 3 9, all of which I
have examined. Redtenbacher (1891 :5 12) referred to Xiphidion oceanicum as a possible synonym of
Xiphidium longipenne de Haan. I have examined the types of both species and found them to be
distinct and very different species.
Further knowledge of C. oceanicus, C. starmuehlneri, C. redtenbacheri and C. infumatus may
show them to be no more than forms of a single species. I am regarding them as separate species here
as they have already been given specific status by previous workers, and there are distinct, although
slight, differences between them. They cannot be considered as four subspecies of one species as the
distribution of each overlaps with one or more of the others (see Fig. 1, Table 1, and the discussion on
p. 321).
MATERIAL EXAMINED (175 Pacific specimens)
Xiphidion oceanicum Le Guillou, lectotype tf, Samoa (Jacquinot) (MNHN, Paris). Xiphidium affine
Redtenbacher, lectotype tf, Fyi Is. ('Coll. Br. v. W. ex Mus. Godeffroy') (NM, Vienna).
Cook Is.: 1 J1, Aitutaki, v; 1 9, Raratonga, vii. Ellice Is.: 3 tf, 2 9, Funafuti I., Funafuti, 0-5 m, ii. Fyi
Is.: 1 $ (Thorey) (NM, Vienna) (paralectotype of Xiphidium affine Redtenbacher); 2 9 (NM, Vienna)
(paralectotypes of Xiphidium affine Redtenbacher); Avea; Fulanga; Komo; Lakemba; Matuku; Moala;
Oneata; Ovalau; Taveuni; Vanua Levu; Viti Levu. Marquesas Is.: Nuku Hiva. New Caledonia. New
Hebrides: Aneityum; Banks Is.; Efate; Erromango; Espiritu Santo; Malekula; Oba; Vanua Lava. New
Guinea: Papua New Guinea. Samoa: 2 9 (Jacquinot) (MNHN, Paris) (paralectotypes of Xiphidion
oceanicum Le Guillou); American, Swains; A., Tutuila; Western, Alafua; W., Savaii; W., Upolu. Society Is.:
330
LINDA M. PITKIN
Bora Bora; Huahine; Nt Raiatea; Tahiti. Tonga Is.: 1 <$, Niuatoputapu, v; 2 <^, Tongatapu, vii; 1 ^ » 1 9» T.,
Nukualofa, ii-x. Tubuai Is. (Austral Is.): 1 <?, Rurutu, iii. (BMNH; DSIR, Auckland; ANIC, Canberra;
BPBM, Honolulu.)
Extralimital material. Aru Is.: 1 9 (MNHN, Paris) (paralectotype of Xiphidion oceanicum Le Guillou).
Philippines: 3 $ (MHN, Geneva) (paralectotypes of Xiphidium qffine Redtenbacher); Babuyan Is.; Luzon.
Vietnam.
DISTRIBUTION (Fig. 1 and Table 1). This species is comparatively widely distributed in the Pacific.
45
itilllMIIUl •••••••
•* -° ,-, •* ^ -8 -9 ^ -»
-9 -a
-e -o
46
47
Figs 45—47 Diagrams showing the arrangement of teeth in the stridulatory file on the under surface of
the left male forewing, and the associated spinules of (45) Conocephalus oceanicus; (46) C.
redtenbacheri; (47) C. willemsei. Other species, similar to these in the arrangement of the stridulatory
teeth, may differ in the size of the file and in the presence or absence of surrounding spinules.
Conocephalus starmuehlneri Kaltenbach
(Figs 1, 3, 33, 43)
Conocephalus (Xiphidion) starmuhlneri Kaltenbach, 1968: 548. Holotype $, NEW CALEDONIA (NM,
Vienna) [examined].
DIAGNOSIS. $ 9- Fastigium of vertex narrow. Prosternum bisipinose. Brachypterous. Forewings
unicolorous, or at most veins R and Sc only slightly darkened (very rarely towards base). Forewings com-
paratively wide, tapering abruptly towards apex (Fig. 33). Cross-veins of costal and precostal areas of
forewings regular, parallel. Stridulatory file of left male forewing shaped as in Fig. 45, with few surrounding
spinules (5 specimens examined). Hind femora with 4-8 externoventral spines. Hind tibiae with 6 apical
spurs. Male cerci as in Fig. 3, with one well-developed internal spine with gently curved apex, similar to Fig.
2a, and one tubercle. Female subgenital plate with shallow excision at apex (Fig. 43). Ovipositor of medium
length, relatively straight.
331
MEASUREMEMENTS
Males Females
Fastigium width (8): 0-39-0-42 (0-41) (11): 0-42-0-52 (0-46)
Median length of pronotum (10): 2-9 - 3-2 (3-04) (13): 3-1 - 3-7 (3-29)
Forewing length (9): 7-8 -10-7 (9-11) (12): 6-9 - 8-3 (7-44)
Hind femur length (6): 11-2 -13-0 (11-92) (13): 12-1 -14-6 (12-90)
Cercus length (8): 1-81- 1-91 (1-84)
Ovipositor length (13): 10-1 -12-2 (10-90)
Pronotum length/fastigium width (8): 6-8 - 8-1 (7-53) (11): 6-8 - 7-9 (7-29)
Hind femur length/ovipositor length (13): 1-11- 1-26 (1-19)
DISCUSSION. C. starmuehlneri closely resembles C. oceanicus, C. redtenbacheri and C. infumatus.
It is distinguished from all except C. redtenbacheri by being brachypterous, and differs from
brachypterous C. redtenbacheri in the shape of the male internal cereal spine, in the forewing
shape, and in most cases in the fainter forewing pigmentation. C. starmuehlneri differs from C.
oceanicus and C. infumatus in the ratio of the hind femur to ovipositor length, and in addition
differs from C. infumatus in having fainter forewing pigmentation. If it were not for these
differences C. starmuehlneri might be regarded as a brachypterous form of one of these species. I
have seen a macropterous female specimen from New Caledonia: Bourail, which possibly belongs
to this species. This specimen differs from both C. oceanicus and C. infumatus in having a ratio of
hind femur to ovipositor length of 1-24, and has almost unicolorous forewings.
I have found a character used by Kaltenbach (1968) to separate C. starmuehlneri and C.
oceanicus to be unreliable in the material I have examined; this is a longitudinal brown stripe on
each side of the pronotum. These stripes are present in the type-specimens I have seen, although
they are very faint in the holotype. They are not present, however, in the other material I have
examined, which is otherwise identical with the type-material. Stripes on the sides of the pronotum,
although not found in C. oceanicus, are present (usually faintly) in some specimens of another
closely related species, C. redtenbacheri.
MATERIAL EXAMINED
Holotype g, New Caledonia: R. Negropo, near Canala, bank ['Uferregion'], 29.vii.1965 (Qsterreichische
Neukaledonien-Expeditiori) (NM, Vienna).
New Caledonia: 7 tf, 9 9 (BMNH); 1 tf, col. d'Amieu, 650 m, (BPBM, Honolulu); 2 9, La Crouen, iii
(BPBM, Honolulu); 1 tf, upper course of R. Negropo, near Koh, 29.vii.1965 (Qsterr. Neukaled.-Exped.)
(NM, Vienna) (paratype); 2 9, R. Negropo, near Canala, bank, 29.vii.1965 (Qsterr. Neukaled.-Exped.) (NM,
Vienna) (paratypes).
DISTRIBUTION (Fig. 1 and Table 1). Known only from New Caledonia.
Conocephalus redtenbacheri (Bolivar)
(Figs 1, 4, 34, 46, 48, 49)
Xiphidium redtenbacheri Bolivar, 1905:389. Syntypes tf, 9, NEW GUINEA: Papua New Guinea, Sattelberg,
1898 (Biro) (lost). NEOTYPE <?, NEW GUINEA (BMNH), here designated [examined].
DIAGNOSIS. $ 9. Fastigium of vertex narrow. Prosternum bispinose. Brachypterous to macropterous.
Forewings with veins R and Sc darkened, at least towards base, sometimes only faintly. Forewings (of
brachypterous form) shaped as in Fig. 34. Cross-veins of costal and precostal areas of forewings regular,
parallel. Stridulatory file of left male forewing shaped as in Fig. 46, surrounded by many spinules (14
specimens examined). Hind femora with 1-7 externoventral spines; very rarely unarmed. Hind tibiae with 6
apical spurs. Male cerci as in Figs 4, 4a, with one well-developed spine with hook-shaped apex, and one
tubercle. Female subgenital plate with shallow excision at apex. Ovipositor moderately short, relatively
straight. Song as in oscillograms, Figs 48, 49, consisting of chirps lasting 1-3 seconds, repeated at a variable
rate (usually c. 9 per minute), with a regular rate of c. 120 syllables per second. The songs of brachypterous
and macropterous specimens were similar.
332 LINDA M. PITKIN
MEASUREMENTS
Males Females
Fastigium width (35): 0-26- 0-50 (0-36) (27): 0-31- 0-55 (0-43)
Median length of pronotum (31): 2-2 - 3-1 (2-74) (33): 2-4 - 3.5 (2-99)
Forewing length (42): 6-6 -17-6 (12-81) (42): 5-5 -20-6 (13-09)
Hind femur length (30): 9-9 -14-3 (12-10) (23): 10-6 -16-2 (13-65)
Cercus length (36): 1-29- 1-66 (1-48)
Ovipositor length (31): 6-8 -10-7 (8-97)
Pronotum length/fastigium width (31): 5-8 -9-3 (7-71) (26): 5-5-8-8 (7-09)
Hind femur length/ovipositor length (22): 1-37- 1-70 (1-51)
DISCUSSION. C. redtenbacheri closely resembles C. oceanicus, C. starmuehlneri and C. infumatus.
It differs from all these species in the shape of the internal spine of the male cerci, and from the first
two, in most cases, in the forewing pigmentation. Many spinules surround the stridulatory file of C.
redtenbacheri whereas these are fewer and more restricted in the other three species (except
occasionally in C. infumatus).
To judge from the material examined C. redtenbacheri is the commonest species of the genus in
the Pacific. The large number of specimens examined showed considerable variation, particularly in
forewing length, which ranged continuously from brachypterous to macropterous, although the in-
termediates were in the minority. Specimens from the Solomon Is. were always macropterous.
Those from Solomon Is.: Ontong Java were usually particularly small, resembling the small New
Ireland form of C. infumatus. The width of the fastigium of the vertex was also variable,
particularly in New Guinea specimens, while some small populations elsewhere in the range were
comparatively uniform.
The forewings were rarely as strongly pigmented around the veins R and Sc as those of C.
infumatus, and were occasionally extremely faintly pigmented. It is therefore possible that some
females of this species might not be distinguishable with certainty from females of C. oceanicus on
morphology alone. Approximately 1% of the specimens examined lacked spines on one hind femur,
and 1 c?, 1 9 had no spines on either hind femur. I examined one specimen, which I believe to be an
aberration of this species, with very tiny cerci; these differed in shape from those of the other adult
males, although they were similar to those of nymphs of the species.
The syntypes of Xiphidium redtenbacheri, from New Guinea, were lost when the Orthopteroid
collection of the Termeszettudomanyi Muzeum, Budapest, was burnt in 1956 (pers. comm. from
Dr H. Steinmann). I have selected a male neotype, also from New Guinea, for which I have a song
recording.
MATERIAL EXAMINED (1077 Pacific specimens)
Xiphidium redtenbacheri Bolivar, neotype $, New Guinea: Papua New Guinea, Morobe Province, Wau, c.
1200 m, Ecology Institute grounds, 3.vii.l979 (Robinson) (BMNH), associated with tape recording no. 301
(recorded by W. J. Reynolds and L. M. Pitkin; tape in BMNH Library of Recorded Insect Sounds).
Admiralty Is.: 1 tf, Manus, 6 km SE. of Lorengau, Rossum, 180 m, xii. Australia: 1 9> Queensland, vi-vii;
1 9, Q., Aratula, xii; 2 <$, Q., Cairns, iii-xi; 4 tf, 1 9, Q., Cairns, Freshwater Creek, ii; 1 9, Q., Kuranda,
200 m, iii. New Britain: 1 $, Gazelle Pen., Keravat, 60 m, ix; 1 tf, G. P., Keravat, ii; 1 tf, G. P., Warongoi
Val., 100m, v; 2 <$, 5 9, Nakanai Mts, Silanga, 150m, vii-viii; 1 tf, Vudal, vi. New Guinea: Irian Jaya;
Papua New Guinea (and associated tape recordings). New Hanover: 1 <$, \. Solomon Is.: I $,l Q;4 $,6
9, Bellona, x-xi; 2 9» B., Henuangoto, v-x; 1 9> Biawa, iv; 2 $, 1 9» Bougainville, v-vii; 1 9? B., Buin, vi; 1
<?, B., Buin (Kangu), 1-50 m, v; 2 9» B., Buin area, Konga Village, ii-iii; 1 $, 3 9> B., Buin area, Kugukai
Village, 150 m, xi-i; 1 tf, 1 9, B., Arigua, vi; 2 tf, 1 9, B., Kieta, xi; 3 tf, B., Kokure, 690 m, vi; 1 J1* 3 9, B.,
Kokure, near Crown Prince Ra., 900 m, vi; 2 9> B., Mumurai, vi; 2 9» B., Mumurai, 400 m, vi; 1 9> B.,
Numa Numa, vi; 1 $, B., Sovele Mission, 250 m, vi; 2 tf, 2 9, B., Teopasino, c. 5° 40' S, 155° 07' E, vi; 1
9, B., Tokinoitu, 20m, vi; 1 tf, Buka, Gagan, 40m, vi; 1 9» Choiseul, Malangono, viii; 1 $, 2 9> C.,
Malangono, 10 m, iii; 3 tf, 3 9, Guadalcanal, ii-xii; 3 tf, 1 9, G., 900-1500 m, xii; 2 tf, 1 9, G., Betikama
R., viii-ix; 2 <$, 2 9, G., Bonegi R., 210 m, xii; 1 9, G., Gold Ridge, iv; 2 tf, 1 9, G., Gold Ridge, 500 m, vi;
1 9, G., Gold Ridge, 800 m, vi; 1 tf, 1 9, G., Honiara, v-x; 2 tf, 3 9, G., Honiara District, Tenaru, vii-x; 1
9, G., Tenaru, 0-100 m, i; 1 <J, G., Tenaru R. (mouth), viii; 1 9, G., 30 km W. of Honiara, Tambalia, v; 1
c?, G., 35 km W. of Honiara, Tambalia, 30 m, v; 1 tf, 1 9, G., H. D., Poha R., vii; 2 tf, G., Ilu Farm, near
Nalimbu R., 5 m, vii; 3 tf, 2 9, G., Ilu, ii-vi; 1 9, G., Kiwi Creek, viii; 3 tf, 3 9, G., Kukum, i-vii; 2 tf, 1 9,
G., Lunga R. (mouth), v-vi; 1 g, 1 9, G., Metanikan R. (mouth), v; 12 tf, 6 9, G., Nuhu, x; 1 9, G., Rua
PACIFIC SPECIES OF Conocephalus
333
Vatu, xi; 2 tf, G., Suta, vi; 3 tf, 3 9, G., Suta, 500-1200 m, vi; 1 9, G., Sutakiki R., vi; 1 9, G., Sutakiki R.,
610m, iv; 1 $, G., Tadhimboko, 0-100 m, x; 1 $, 1 9, G., Tapenanje, xii; 1 $, 3 9, G., near Terere,
Roroni, 1 9, G., Tinahula R., iii; 1 9, G., Umasani R., 10 km inland, i-vii; 1 9, G., Wright's Creek, viii; 1 $,
2 9, Malaita, Auki, 2-20 m, ix-xii; 1 9, M., Kwailasi, Fulisango, v; 1 tf, 1 9, M., Rai'ako, v; 1 tf, 2 9, M.,
Sikiana, iii-x; 1 9> M., Su'u, iv; 1 9> Ndai, Bethlehem, 0-10 m, xii; 2 9, New Georgia Group, Gizo I., 30 m,
vii; 1 tf, N. G. G., G., 50-120 m, iv; 1 rf, 1 9, N. G. G., G., Gizo 0-100 m, xi; 1 9, N. G. G., G., Loga I., x;
1 tf, 1 9, N. G. G., Kolombangara, viii; 4 $, 2 9, N. G. G., K., Kuzi, x; 1 g, 1 9, N. G. G., K., Pepele,
30 m, ii; 2 9, N. G. G., New Georgia I., Munda, 1-30 m, vii; 2 9, N. G. G., N. G., Munda and district, viii;
1 cf , 3 9, N. G. G., N. G., Segi, Maravo, v; 1 tf, 2 9, Ontong Java, Kepae, i; 1 1 g, 9 9, O. J., Kiloma, i; 2
O*, O. J., Leuaniua, i; 5 <?, 10 9, O. J., Peku, 0-10 m, xii; 6 J, 8 9, O. J., Pelau, ii; 3 tf, 3 9, Rennell,
Hutuna, x-xi; 2 9, Russell Is., Yandina, ix; 1 $, San Cristobal, Hawa, v; 3 $, S. C., Huni R. (mouth), viii; 1
o% S. C., Makina, v; 1 tf, S. C., Waimamura, iv; 1 tf, S. C., Wainoni, vii; 3 9, Santa Cruz Is., Utupua, xi; 6
c?, 1 9, S. C., Vanikoro Is., ii-xi; 1 9, S. C., V., Buma, ix; 1 $, Santa Isabel, iii; 1 9, S. I., Allardyce Hbr, ii;
1 9, S. I., Gatere, ii; 1 9, S. I., Maringe Lagoon ('Lgu'?), ii; 1 9, S. I., Rasa, v; 1 9, S. I., Sisaga, ii; 2 9, S. I.,
Tatamba, ix; 1 $, Vella Lavella, Kow, 30 m, xi; 4 $, 6 9, V. L., Ulo Crater, 10 m, xii. (BMNH; ANIC,
Canberra; BPBM, Honolulu; DPI, Konedobu.)
Extralimital material. Philippines: Mindanao; Palawan.
DISTRIBUTION (Fig. 1 and Table 1). This species is very common and widespread in the western
Pacific.
Conocephalus injumatus (Redtenbacher)
(Figs 1, 5, 6)
Xiphidium infumatum Redtenbacher, 1891: 512. Holotype 9, DUKE OF YORK GROUP: Mioko (between
New Britain and New Ireland) (lost). NEOTYPE tf, NEW BRITAIN (BPBM, Honolulu), here designated
[examined],
DIAGNOSIS. $ 9- Fastigium of vertex narrow. Prosternum bispinose. Macropterous. Forewings with veins/?
and Sc darkened. Cross-veins of costal and precostal areas of forewings regular, parallel. Stridulatory file of
left male forewing shaped as in Fig. 45, mostly with few surrounding spinules. Hind femora with 2—9 ex-
ternoventral spines. Hind tibiae with 6 apical spurs. Male cerci as in Figs 5, 5a, 6, 6a, with one well-developed
internal spine with moderately gradually curved apex, and one tubercle. Female subgenital plate with
moderately shallow excised apex. Ovipositor moderately short to medium length, relatively straight.
MEASUREMENTS
Typical form
Fastigium width
Median length of pronotum
Forewing length
Hind femur length
Cercus length
Ovipositor length
Pronotum length/fastigium width
Hind femur length/ovipositor length
SP .dll New Ireland form (see below)
Fastigium width
Median length of pronotum
Forewing length
Hind femur length
Cercus length
Ovipositor length
Pronotum length/fastigium width
Hing femur length/ovipositor length
Males
(10): 0-31- 0-39 (0-35)
(10): 3-1 - 3-5 (3-24)
(10): 13-4 -18-5 (16-37)
(9): 13-4 -15-8 (14-18)
(10): 1-68- 1-84 (1-77)
(10): 8-8 -10-3 (9-38)
Males
(6): 0-23- 0-29 (0-27)
(6): 2-2 - 3-0 (2-44)
(6): 9-3 -13-3 (10-53)
(4): 9-9 -12-5 (10-89)
(5): 1-29- 1-64 (1-44)
(6): 8-0 -10-5 (9-22)
Females
(33): 0-26- 0-42 (0-34)
(33): 3-2 - 4-0 (3-56)
(33): 15-9 -22-1 (18-60)
(28): 14-1 -17-0 (15-56)
(31): 9-7 -11-5 (10-56)
(33): 9-0 -12-5 (10-63)
(26): 1-35- 1-57 (1-48)
Females
(3): 0-28- 0-29 (0-29)
(3): 2-6 - 2-8 (2-70)
(3): 11-7 -13-1 (12-35)
(3): 11-6 -12-3 (11-81)
(3): 8-2 - 9-1 (8-57)
(3): 9-3 - 9-6 (9-41)
(3): 1-36- 1-41 (1-38)
DISCUSSION. C. infumatus closely resembles C. oceanicus, C. starmuehlneri and C. redtenbacheri,
in particular the last of these. It can be distinguished from these species by the following
combination of characters: the strongly darkened R and Sc in the forewings (except for small New
Ireland form - see below); the fairly gently curved apex of the internal spine of the male cerci, and
the high ratio of pronotum length to width of fastigium. The internal spine of the male cerci is
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PACIFIC SPECIES OF Conocephalus 335
intermediate in shape between C. oceanicus and C. redtenbacheri, although slightly closer to the
former.
Most of the specimens examined from New Ireland seemed to belong to a small form, dis-
tinguishable from the typical form by the following key.
1 Comparatively large specimens with dark pigmentation around R and Sc of the forewings. Hind
femur length at least 13-0 mm. Forewing length at least: <$, 13-4 mm; 9> 15-9 mm
. typical form
- Comparatively small specimens with fairly weak pigmentation around R and Sc of the
forewings. Hind femur length at most 12-5 mm. Forewing length at most: tf, 13-3 mm; 9>
13- 1mm small New Ireland form
The existence of two forms of of this species is a further complication in considering its status in
relation to its three close relatives. Where C. infumatus comes in contact with C. redtenbacheri in
New Britain, the two species are distinct. At the eastern end of the range, however, the speciation
appears to be less complete. The small New Ireland form of C. infumatus is very similar to some
specimens of C. redtenbacheri in the Solomon Is., in particular Ontong Java. Most specimens of
the small New Ireland form have been found off New Ireland in the Faed Is. of the Nuguria Is.,
which, like the Ontong Java Is. in the Solomon Is., are a very small, outlying easterly group.
The female holotype of Xiphidium infumatum is lost (see Weidner, 1966:243). I have selected a
male neotype, as the males of this species are more easily distinguished from related species than
the females. The neotype-locality is in New Britain, not far from the original type-locality, the
island of Mioko.
One specimen of C. infumatus was found to have an internoventral spine on one hind femur.
MATERIAL EXAMINED
Xiphidium infumatum Redtenbacher, neotype tf, New Britain: Gazelle Peninsula, Raining, St Paul's, 350 m,
5.ix.l955 (Gressiti) (BPBM, Honolulu).
Typical form. New Britain: 7 tf, 9 9, Gazelle Peninsula, Baining, St Paul's, 350 m, ix; 1 9, G. P., Keravat,
ii; 1 9, G. P., Upper Warongoi, Illugi, 230m, xii; 1 9, G. P., Warongoi Valley, 100m, v; 3 9, Mosa
Plantation, iv-vi; 2 9, Mosa Village, iv; 2 9, Nakanai Mts, Gisiluve, 1050 m, vii; 1 $, 3 9, N. M., Silanga,
150m, vii-viii; 2 9, N. M., Talalo, 900m, vii; 1 $, N. M., Ti, vii; 1 9, S. coast, Rano Pl'n, iv; 1 9, Mt
Sinewit, 1070 m, viii; 1 9> N. coast, Walo, vii; 2 9, W. of Willaumez Peninsula, Linga Linga Pl'n, 1 m, iv.
New Ireland: 1 9> Kandan, xii. Solomon Is.: 1 9» Bougainville, Kokure, near Crown Prince Range, 900 m,
vi. (BMNH; ANIC, Canberra; BPBM, Honolulu; DPI, Konedobu.)
Small New Ireland form. New Ireland: 5 g, 3 9, Faed Is., x (BMNH; DPI, Konedobu); 1 tf, Gilingil Pl'n,
2 m, vii (BPBM, Honolulu).
DISTRIBUTION (Fig. 1 and Table 1). This species only occurs in New Britain, New Ireland and the
northernmost of the Solomon Is. Previous records of the species from New Guinea refer to
specimens which I have not seen but consider likely to be C. redtenbacheri.
Conocephalus semivittatus (Walker)
(Figs 7-9, 28, 50-53)
Decticus semivittatus Walker, 1869:263.
DIAGNOSIS. $ 9- Fastigium of vertex wide. Prosternum bispinose. Macropterous or brachypterous.
Forewings with dark pigmentation in costal and precostal areas, and usually with spot at posterior margin
(Fig. 28). Cross-veins of costal and precostal areas of forewings irregular or reduced. Stridulatory file of left
male forewing shaped as in Figs 45 and 46. Hind femora with 1—9 externoventral spines. Hind tibiae with 6
apical spurs. Male cerci shaped as in Figs 7-9, with one well-developed internal spine and one tubercle.
Female subgenital plate with excised or truncate apex. Ovipositor of short to medium length, relatively
straight.
DISCUSSION. C. semivittatus resembles C. tumidus but differs from it in the shape of the male cerci
and usually in the number of hind tibial spurs. Nearly all the females examined had a narrower
fastigium of the vertex than those of C. tumidus. The excised apex of the female subgenital plate of
the nominate subspecies and the typical form of subspecies vittatus also distinguishes them from
those of C. tumidus.
336 LINDA M. PITKIN
Specimens occurring in the north of the species range differ in some characters from those in the
south, and I consider them to belong to a distinct subspecies. The geographical ranges of the two
subspecies do not overlap, although they approach each other in Australia, where C. 5. semivittatus
occurs in the south and C. s. vittatus is apparently confined to the north. I have only seen two
specimens of the latter subspecies from Australia, both males, but these have particularly stout
cerci, suggesting that this character may be more pronounced in the region of contact between the
two subspecies. If this is taken as evidence of character displacement, it suggests that the two
groups, which have long been regarded as a single species, might be specifically distinct. More
specimens are required before a firm judgement can be made on this possibility.
The right fore wing of the brachypterous males was usually slightly longer (by 0-3 mm on
average) than the left forewing.
DISTRIBUTION (Table 1). This species is common and widespread in the western Pacific.
Key to the subspecies and forms of C. semivittatus
1 Left forewing, and sometimes right, with spot at posterior margin as in Fig. 28. Male cerci
shaped as in Figs 7, 8. Female subgenital plate with excised apex .... 2
Forewings without spot at posterior margin, or at most with diffuse trace. Male cerci shaped as
in Fig. 9. Female subgenital plate with truncate apex ......
C. s. vittatus (Redtenbacher) (small form) (p. 338)
2 Dorsum of pronotum patterned as in Fig. 52. Male cerci, from internal spine to apex, com-
paratively long and slender in proportion to size (Fig. 7). Ovipositor length at least 10-5 mm
. C. s. semivittatus (Walker) (p. 336)
Dorsum of pronotum patterned as in Fig. 53. Male cerci, from internal spine to apex, com-
paratively short and robust, narrowing sharply at apex (Fig. 8). Ovipositor length not more
than 9-5 mm • • • • C. s. vittatus (Redtenbacher) (typical form) (p. 338)
Conocephalus semivittatus semivittatus (Walker)
(Figs 7, 52)
Decticus semivittatus Walker, 1869:263. Holotype 9, NEW ZEALAND (BMNH) [examined].
Xiphidium maoricum Walker, 1869:276. LECTOTYPE 9, NEW ZEALAND (BMNH), here designated
[examined]. [Synonymized by Hutton, 1898:137.]
Xiphidium antipodum Scudder, 1875:460. LECTOTYPE tf, NEW ZEALAND (ANS, Philadelphia), here
designated [examined]. [Synonymized by Hutton, 1898:137.]
Xiphidium (Xiphidion) brunneri Karny, 1907:94. LECTOTYPE 9, NEW ZEALAND (NM, Vienna), here
designated [examined]. Syn. n.
DIAGNOSIS. $ 9- Dark stripe of dorsum of pronotum shaped as in Fig. 52. Left forewing, and sometimes
right, with spot on posterior margin as in Fig. 28. Male cerci from internal spine to apex, comparatively long
and uniformly slender, in proportion to size (Fig. 7). Female subgenital plate with excised apex. Ovipositor of
medium length.
MEASUREMENTS
Males Females
Fastigium width (20): 0-55-0-80 (0-65) (40): 0-57-0-81 (0-71)
Median length of pronotum (20): 3-0 - 3-9 (3-38) (42): 3-2 - 4-1 (3-68)
Forewing length (macropterous) (5):20-1 -21-2 (20-73) (21): 20-7 -23-8 (21-91)
(brachypterous) (15): 4-6 - 7-8 (5-94) (19): 2-5 - 5-9 (3-70)
Hind femur length (14): 11-6 -14-2 (13-15) (33): 12-3 -16-5 (14-70)
Ovipositor length (41): 9-9 -12-7 (11-49)
DISCUSSION. The nominate subspecies differs from C. 5. vittatus in the shape of the male cerci and
the longer ovipositor. It also differs, at least from the typical form of C. s. vittatus, in the shape of
the dorsal pronotal stripe.
The lectotype of Xiphidium maoricum was selected from the type-series of 2 9» both of which I
have examined. A lectotype was also selected from the 1 ^, 1 9 adult syntypes of Xiphidium
antipodum, which I have examined; there are several nymphal syntypes which I have not seen.
55
58
Figs 52-59 52, 53, dorsal view of the pronotum showing the outline of the area of dark pigmentation of
(52) Conocephalus semivittatus semivittatus; (53) C. semivittatus vittatus (typical form). 54-59, dorsal
view of the right titillator of (54) C. tumidus; (55) C. angustivertex; (56) C. bispinatus (New Guinea);
(57) C. bispinatus (Thailand); (58) C. bispinatus (Malaya); (59) C. bispinatus (Borneo). Surface
sculpturing (other than ridges) is omitted.
I have compared the type-series of Xiphidium (Xiphidion) brunneri with the holotype and other
material of C. s. semivittatus and consider them to be conspecific. The type-series of brunneri
consists of 1 $, 1 9> of which I have selected the female as the lectotype as it is in a better state of
preservation than the male.
Brachypterous specimens were more common than macropterous specimens by 37 to 26 and
usually had rounded forewing apices. One brachypterous female from New Zealand had distinctly
pointed forewings though, and the wings were slightly longer than those of the other brachypterous
females. This feature has been described by Hudson (1972).
Three of the specimens examined were found to have one or two internoventral spines on one
hind femur.
MATERIAL EXAMINED (63 specimens, brachypterous unless otherwise stated)
Decticus semivittatus Walker, holotype 9> New Zealand: Auckland (Sinclair) (BMNH). Xiphidium
maoricum Walker, lectotype 9» macropterous, New Zealand (Boltori) (BMNH). Xiphidium antipodum
Scudder, lectotype $, New Zealand (Edwards) (ANS, Philadelphia). Xiphidium (Xiphidion) brunneri Karny,
lectotype 9» New Zealand (Mutton) (NM, Vienna).
Australia: macropterous and brachypterous, New South Wales. Kermadec Is.: Raoul. Lord Howe I.: 1 <$,
2 9, macropterous, 2 tf, 2 9, NSW., ii-iii. New Zealand: 1 9, macropterous (Boltori) (BMNH)
338 LINDA M. PITKIN
(paralectotype of Xiphidium maoricum Walker); 1 9> macropterous (Colenso) (BMNH) (Xiphidium
maoricum Walker, var. B); 1 9 (Edwards) (ANS, Philadelphia) (paralectotype of Xiphidium antipodum
Scudder); 1 $ (Huttori) (NM, Vienna) (paralectotype of Xiphidium (Xiphidion) brunneri Karny);
macropterous and brachypterous, North I. (BMNH; BPBM, Honolulu.)
DISTRIBUTION (Table 1). This subspecies occurs in the south of the species range, extending
northwards to Australia: New South Wales.
Conocephalus semivittatus vittatus (Redtenbacher) nom. rev., stat. n.
(Figs 8, 9, 50, 51, 53)
Xiphidium vittatum Redtenbacher, 1891:513. LECTOTYPE, $, ARU Is. (NM, Vienna), here designated
[examined]. [Synonymized with Conocephalus maoricus (Walker) by Karny, 1912:11.]
Xiphidium geniculare Redtenbacher, 1891:527. LECTOTYPE, $, ARU Is. (NM, Vienna), here designated
[examined]. [Synonymized with Xiphidium semivittatum (Walker) by Hutton, 1898:137.]
DIAGNOSIS. $ 9- Dark stripe of dorsum of pronotum shaped as in Fig. 53. Forewings with or without spot
on posterior margin (Fig. 28). Male cerci shaped as in Figs 8, 9. Female subgenital plate with excised or
truncate apex. Ovipositor short. Song (of typical form) as in oscillograms (Figs 50, 51), consisting of a burst
of sound usually lasting 5—6 seconds, occasionally up to 15 seconds. Each burst of sound consists of two
alternating types of sound, the longer parts of which have a rate of c. 50 syllables per second, and the
considerably shorter parts of which have a rate of 16-20 syllables per second. The songs of brachypterous
and macropterous specimens were similar.
MEASUREMENTS
Typical form Males Females
Fastigium width (47): 0-52-0-76 (0-65) (36): 0-52-0-83 (0-68)
Median length of pronotum (49): 2-8-3-9 (3-31) (37): 3-0-4-0 (3-37)
Forewing length (macropterous) (19): 18-0 -21-0 (19-35) (14): 17-3 -21-3 (19-15)
(brachypterous) (30): 4-0 - 7-5 (5-39) (21): 2-1 - 4-3 (3-07)
Hind femur length (39): 13-0 -16-4 (14-70) (34): 14-2 -16-6 (15-46)
Ovipositor length (37): 7-2 - 9-3 (8-69)
Small form Males Females
Fastigium width (14): 0-57-0-73 (0-66) (24): 0-63-0-81 (0-73)
Median length of pronotum (14): 2-5 - 3-1 (2-89) (24): 2-9 - 3-3 (3-11)
Forewing length (macropterous) (2): 18-0 -18-1 (18-05) (2): 19-1 -19-5 (19-30)
(brachypterous) (12): 2-5 - 3-6 (3-25) (19): 1-9 - 3-3 (2-39)
Hind femur length (12): 11-7 -14-9 (13-22) (24): 13-0 -15-1 (14-26)
Ovipositor length (22): 7-3 - 8-3 (7-85)
DISCUSSION. C. s. vittatus differs from the nominate subspecies in the shape of the male cerci and
the shorter ovipositor. The shape of the dorsal pronotal stripe of the typical form of C. s. vittatus
also differs from the nominate subspecies; that of the small form is more variable and tends to be
intermediate.
The specimens I am referring to as the small form show small but consistent differences from the
rest of the subspecies. It is possible that they may prove to belong to a separate species, but this
cannot be firmly established on the basis of present knowledge.
This form differs from the typical form of the subspecies, and also from C. s. semivittatus, in
having no spot of pigmentation on the posterior forewing margin, or at most only a diffuse trace. It
also differs from both in the shorter forewings of brachypterous males, the truncate apex of the
female subgenital plate and the shape of the male cerci. The male cerci of the typical form of C. s.
vittatus are more robust than in the others, while those of the small form are on average smaller
than those of the other two, and to some extent intermediate in shape between the typical form and
C. tumidus. The internal cereal spine of the small form is comparatively wider at the base than in
the rest of the species.
The small form shares with C. tumidus most of the characters distinguishing it from the typical
form, while sharing with the typical form the diagnostic features which separate C. semivittatus
from C. tumidus.
PACIFIC SPECIES OF Conocephalus 339
Brachypterous specimens of C. s. vittatus were very much more common than macropterous, in
the ratio of 7 to 1.
Redtenbacher (1891) gives Xiphidium antipodum and X bilineatum as possible synonyms of X.
geniculare. X, antipodum is in fact a synonym of C. s. semivittatus, and C. bilineatus is a valid
species. C. semivittatus has been regarded as a synonym of C. bilineatus by several authors prior to
Hudson, 1972, who rightly separated the two species, although she leRX. geniculare as a synonym
of C. bilineatus.
Xiphidium geniculare was synonymized with Xiphidium semivittatum by Hutton, 1898, but has
since been placed as a synonym of C. bilineatus. I have examined 4 $ adults, 1 9 nymph from the
type-series of X. geniculare, and I agree with the earlier synonymy. In addition to the syntypes
examined, from which I have selected a male lectotype, there are specimens from Mioko, New
Zealand and 1 $ from the Aru Is. which I have not seen. I have also selected a male lectotype from
the type-series of Xiphidium vittatum, consisting of 1 $, 1 9 which I have examined, and one or
more specimens from New Zealand which I have not seen.
One of the apical spurs was missing on one hind tibia in two specimens examined.
MATERIAL EXAMINED
Xiphidium vittatum Redtenbacher, lectotype <$, macropterous, Aru Is. (Ribbe) (NM, Vienna). Xiphidium
geniculare Redtenbacher, lectotype tf, Aru Is. (Ribbe) (NM, Vienna).
Typical form (273 Pacific specimens, brachypterous unless otherwise stated). Admiralty Is.: 1 tf, Manus,
Lorengau, 1-75 m, vi; 1 9. M., near Lorengau, vi; 2 9, M., c. 8 km E. of Lorengau, near Lombrum, vi; 2 9,
M., 6 km SE. of Lorengau, Rossum, 180 m, xii; 1 $, M., Rossum, 32-125 m, vi; 1 <$, 1 9, M., Momote, xii.
Australia: N. Queensland. Duke of York Group: 1 tf, Mioko (between New Britain and New Ireland) (NM,
Vienna) (paralectotype of Xiphidium geniculare Redtenbacher). New Britain: 2 <$, Akanglo I., near
Kandrian, S. coast, iv; 1 $, Baronga I., near Lindenhafen, iv; 2 tf, Gazelle Peninsula, Baining, St Paul's,
350 m, ix; 2 tf, G. P., Keravat, 60 m, viii; 1 tf, G. P., 10 km E. of Keravat, Vunabakan, 180 m, xi; 1 tf, G.
P., Upper Warongoi, 250-600 m, xi; 1 $, 2 9, macropterous, G. P., Warongoi Valley, 100m, v. New
Guinea: macropterous and brachypterous, Irian Jaya; macropterous and brachypterous, Papua New
Guinea, and associated tape recordings. New Hebrides: Banks Is.; Epi; Espiritu Santo; Malekula; Pentecost;
Vanua Lava. N-* Ireland: 1 9, SW., Gilingil Pl'n, 2m, vii. Solomon Is.: 1 9, Bellona, xi; 1 9, B.,
Henuangoto, x; 1 9» macropterous, Biawa, iv; 1 $, macropterous, 1 <$, Bougainville, near Crown Prince
Range, Kokure, 900 m, vi; 1 <$, B., S., Kieta, xi; 1 $, B., Mumurai, vi; 2 9, B., Teopasino, c. 5° 40' S, 155°
07' E, vi; 1 9. Choiseul, Malangona, 10 m, iii; 1 tf, Guadalcanal, xii; 1 J1, G., Gold Ridge, iii; 1 $, G.,
Honiara District, Kukum, viii; 2 $, G., H. D., Poha R., vii-x; 1 <$, macropterous, 2 tf, 2 9, G., H. D.,
Tenaru, v-x; 1 tf, G., Ruavatu, iv; 4 tf, 1 9, G., Suta, 500-1200 m, vi; 1 9, G., Sutakiki R., iv; 2 9, G.,
Tapenanje, xii; 1 9, G., Tapenanje, c. 335 m, xii; 1 <$, Malaita, Auki, 2-20 m, ix; 1 $, M., Fulisango-
Maelegwasu, v; 1 $. Rendova, x; 2 <$, 1 9' San Cristobal, Huni R. Mouth, viii; 1 tf, S. C., Kira-Kira, grass,
xi; 1 9. Santa Cruz Is., coconut, xi; 1 <$, 1 9, Santa Isabel, Gatere, ii; 1 9» S. I., Maringe Lagoon (Lgu' ?), ii;
1 c?, Western Group, Kolombangara, Kuli, x; 1 <$, K., 2 km inland from Kuzi, by Kolombangara R., ix.
(BMNH; DSIR, Auckland; ANIC, Canberra; BPBM, Honolulu; DPI, Konedobu.)
Extralimital material. Aru Is.: 2 $ (Ribbe) (NM, Vienna) (paralectotypes of Xiphidium geniculare
Redtenbacher); 1 9> macropterous (Ribbe) (NM, Vienna) (paralectotype of Xiphidium vittatum
Redtenbacher). Kai Archipelago: Taam. Moluccas. Philippines. Sulawesi: Cape Dulang? ('Ke Dulan').
Small form (38 Pacific specimens). New Guinea: macropterous and brachypterous, Irian Jaya;
macropterous and brachypterous, Papua New Guinea. (BMNH; ANIC, Canberra; BPBM, Honolulu.)
Extralimital material. Philippines: macropterous and brachypterous.
DISTRIBUTION (Table 1). This subspecies occurs in the north of the species range extending
southwards to Australia: N. Queensland and the New Hebrides. The two forms of C. s. vittatus are
sympatric, although the small form has a more restricted distribution.
Conocephalus tumidus sp. n.
(Figs 10, 23, 29, 54)
DESCRIPTION AND DIAGNOSIS. $ 9- Fastigium of vertex very wide, usually slightly more than width of basal
antennal segment. Prosternum bispinose. Brachypterous. Forewings with dark pigmentation in costal and
precostal areas, without spot at posterior margin (Fig. 29). Cross-veins of costal and precostal areas of
forewings reduced, irregular. Stridulatory file of left male forewing shaped as in Figs 45 and 46. Hind femora
340 LINDA M. PITKIN
usually with 1-4 externoventral spines; one femur sometimes unarmed. Hind tibiae usually with not more
than 4 apical spurs (ventral pair missing). In some cases one, or occasionally both, internodorsal apical spurs
also missing. Male cerci as in Fig. 10, swollen in middle, with one well-developed internal spine and one
tubercle. Titillators as in Fig. 54. Female subgenital plate with truncate apex. Ovipositor short, relatively
straight (Fig. 23).
General coloration: mixture of green, reddish brown and dark brown. Head and pronotum with dark
brown, dorsal, longitudinal stripe, with light margins on the pronotum. Femora usually with small dark spots.
Abdomen sometimes with wide, longitudinal stripes, faintly darker than ground colour.
MEASUREMENTS
Males Females
Totallength (5): 11-6 -13-9 (12-48) (4): 12-8 -16-1 (14-40)
Fastigium width (6): 0-62-0-93 (0-72) (4): 0-83-0-91 (0-85)
Median length of pronotum (6): 2-7 - 3-3 (2-97) (4): 3-1 - 3-3 (3-19)
Forewing length (6): 4-9 - 6-1 (5-30) (4): 2-6 - 3-0 (2-89)
Hind femur length (5): 10-5 -13-5 (11-85) (3): 12-7 -14-2 (13-40)
Ovipositor length (4): 7-4 - 8-4 (7-98)
DISCUSSION. C. tumidus is similar to C. semivittatus, particularly the small form of C. s. vittatus.
The males can be distinguished by the shape of the cerci, and in most cases by the hind tibial apical
spurs. The females can be distinguished by the hind tibial apical spurs together with the wide
fastigium of the vertex. The ventral hind tibial apical spurs were absent in all except one of the
specimens examined and were often replaced by hairs. The right forewing of the brachypterous
males was usually slightly longer (by 0-3 mm on average) than the left forewing.
MATERIAL EXAMINED
Holotype tf, New Guinea: Papua New Guinea, NE., 48km E. of Kainantu, Kassam, 1350m, 28.X.1959
(Mad) (BPBM, Honolulu).
Paratypes. New Guinea: 1 <3\ 1 9» Papua New Guinea, Eastern Highlands District, 25 km ENE. of
Kainantu, Kassam Pass, 16.ii.1976 (Farrow) (ANIC, Canberra); 1 tf, 1 9, P. N. G., Eastern Highlands
District, 25 km ENE. of Kainantu, Kassam Pass, 16.ii.1976 (Farrow) (BMNH); 1 tf, P. N. G., NE., 48 km
E. of Kainantu, Kassam, 1350m, 28.X.1959 (Maa) (BMNH); 1 g, P. N. G., NE., 48 km E. of Kainantu,
Kassam, 1350m, 30.x. 1959 (Maa) (BPBM, Honolulu); 1 9, P. N. G., Morobe District, Ramu-Markham
Valley, Gusap Downs Sta., 13.ii.1976 (Farrow) (ANIC, Canberra); 1 $, 1 9, P. N. G., Morobe District,
Ramu-Markham Valley, Gusap Downs Sta., 14.ii.1976 (Farrow) (ANIC, Canberra).
DISTRIBUTION (Table 1). Known only from north-eastern New Guinea.
Conocephalus trivittatus (Stal)
(Fig. 11)
Xiphidium trivittatum Stal, 1860: 323. LECTOTYPE J1, SOCIETY Is. (NR, Stockholm), here designated
[examined].
DIAGNOSIS. J1 9- Fastigium of vertex moderately wide. Prosternum bispinose. Brachypterous. Forewings
with darker pigmentation, sometimes faint, towards the base of the costal and precostal areas. Cross-veins of
costal and precostal areas of forewings regular, parallel. Stridulatory file of left male forewing shaped as in
Figs 45 and 46. Hind femora unarmed ventrally. Hind tibiae with 6 apical spurs. Male cerci as in Fig. 11,
with one well-developed internal spine and one tubercle. Ovipositor of medium length, relatively straight.
MEASUREMENTS
Male Female
Fastigium width 0-60 0-69
Median length of pronotum 2-8 3-1
Forewing length 5-2 4-6
Length of Stridulatory area 1 -04
Hind femur length 10-5 11-3
Ovipositor length 10-7
DISCUSSION. This species resembles C. semivittatus in the form of the male cerci, and forewing
pigmentation, and C. oceanicus and related species in the subcostal forewing venation and to a lesser
PACIFIC SPECIES OF Conocephalus 341
extent, the male cerci. It can be distinguished from these species, however, by the unarmed hind
femora. The tubercle of the male cerci is comparatively small.
The lectotype of Xiphidium trivittatum was selected from a syntypic series of 1 c?, 1 9-
MATERIAL EXAMINED
Lectotype tf, Society Is.: Tahiti (Kinb.) (NR, Stockholm).
Society Is.: 1 9, Tahiti (Kinb.) (NR, Stockholm) (paralectotype).
DISTRIBUTION (Table 1). Known only from the type-locality.
Conocephalus upoluensis (Karny) nom. rev., stat. n.
(Figs 12, 30, 41)
Xiphidium modestum Redtenbacher, 1891:510. LECTOTYPE tf, AUSTRALIA (NM, Vienna), here
designated [examined]. [Homonym of Xiphidium modestum Bruner, 1891:56.]
Xiphidium (Xiphidion) modestum upoluense Karny, 1907:95. LECTOTYPE <$, SAMOA (NM, Vienna), here
designated [examined]. [Synonymized with Xiphidium modestum Redtenbacher by Holdhaus, 1908: 11.]
Conocephalus brevixiphus Willemse, 1942:95. Holotype 9, AUSTRALIA (NM, Basle) [examined]. Syn. n.
DIAGNOSIS. <$ 9- Fastigium of vertex wide. Prosternum bispinose. Macropterous. Forewings with dark
pigmentation in costal and precostal areas and comparatively small dark spots elsewhere; with or without
dark line or spot towards base of posterior margin, usually faint if present (Fig. 30); light and dark
pigmentation usually not strongly contrasted. Cross-veins of costal and precostal areas of forewings
irregular. Stridulatory file of left male forewing shaped as in Figs 45 and 46. Hind femora unarmed ventrally.
Hind tibiae usually with 4 apical spurs (ventral pair missing). Male tenth abdominal tergite nearly unmodified
or slightly produced at apex but not bent downwards (Fig. 41). Male cerci moderately slender in apical part,
with one internal spine and usually with one minute tubercle or slight swelling (Fig. 12). Ovipositor
moderately short, relatively straight.
MEASUREMENTS
Males Females
Fastigium width (21): 0-59-0-86 (0-70) (20): 0-59-0-81 (0-74)
Median length of pronotum (25): 2-6 - 3-8 (3-09) (21): 2-7 - 3-7 (3-18)
Forewing length (25): 13-3 -22-2 (17-13) (22): 14-2 -23-0 (18-31)
Length of Stridulatory area (20): 1-55- 2-19 (1-84)
Hind femur length (20): 10-8 -15-1 (12-92) (22): 11-8 -16-5 (14-26)
Ovipositor length (22): 6-8 -11-0 (8-93)
DISCUSSION. C. upoluensis very closely resembles C. albescens, but may be distinguished from it
by the shape of the male cerci and tenth abdominal tergite, and the fairly short ovipositor. The dark
pigmentation of the forewings of C. upoluensis varies from being very faint to being as conspicuous
as in C. albescens, although the latter species usually has more strongly contrasted light and dark
pigmentation. Both species have one internal spine and a minute tubercle on the male cerci, thus
being intermediate between species with one internal spine and species with one internal spine and a
tubercle. This tubercle is sometimes virtually absent in C. albescens, and very occasionally absent
in C. upoluensis.
The species was originally described as Xiphidium modestum by Redtenbacher in July, 1891. I
have found this to be a primary homonym of a species described by Bruner in March, 1891, from
U.S.A.
Comparison of the type-specimens and other material of C. brevixiphus and C. upoluensis has
shown them to be conspecific. Willemse's description of C. brevixiphus refers to a female type, and
I therefore regard this specimen as the holotype, despite the fact that it is labelled 'Allotype' and a
male paratype is labelled 'Type'. I have selected a lectotype from the 1 $, 3 9 syntypes of
Xiphidium modestum upoluense, all of which I have examined. I have also selected a lectotype from
the syntype-series of Xiphidium modestum. In addition to the specimens listed under material
examined, there are syntypes which I have not examined with the following data: Australia: Cape
York (Brunner). Fiji: - — ; Ovalau. Tonga Is.: Tongatapu. The syntypes with the following data
are lost (see Weidner, 1966:243): Australia: Peak Downs; Sydney. Samoa. There are also 2 $
syntypes which are C. albescens and I have listed them under that species.
342 LINDA M. PITKIN
MATERIAL EXAMINED (175 Pacific specimens)
Xiphidium modestum Redtenbacher, lectotype <$, Australia: Sydney (Trauenfeld) (NM, Vienna). Xiphidium
(Xiphidion) modestum upoluense Karny, lectotype tf, Samoa: Western, Upolu (Rechinger) (NM, Vienna).
Conocephalus brevixiphus Willemse, holotype 9' Australia: Northern Territory, Marrakai, v.1931
(Handschiri) (NM, Basle) (mislabelled 'Allotype').
Australia: A. C. T.: New South Wales; Northern Territory; 1 tf, 1 9, N. T., Burnside, v.1931 (Handschiri)
(NM, Basle) (paratypes of C. brevixiphus Willemse); 1 tf, N. T., Katherine, v. 193 1 (Handschiri) (NM, Basle)
(paratype of C. brevixiphus Willemse, mislabelled 'Type'); Queensland; S. Australia; Western Australia.
Cook Is.: 1 (51, Niue, Kaimiti Forest, on the firebreak, xi; 1 9. N., Vaiea Farm, vii. Fyi Is.: 1 $ (Thorey)
(NM, Vienna) (paralectotype of Xiphidium modestum Redtenbacher); Aiwa; Kandavu; Komo; Lomaloma;
Mango; Matuku; Oneata; Ovalau; Vanua Levu; Vanua Mbalavu; Vanua Vatu; Viti Levu; Waya; Yasawa
Group. Loyalty Is.: Ouvea. New Britain: 1 9» Gazelle Peninsula, Baining, St Paul's, 350m, ix; 1 <$, G. P.,
Gaulim, 140 m, x; 2 <$, G. P., Keravat, Lowl Agr. Exp. Sta., at light in oilpalm block, v; 1 9, Nakanai Mts,
Gisiluve, 1050 m, vii. New Caledonia: 1 9 (Deyrolle) (NM, Vienna) (paralectotype of Xiphidium modestum
Redtenbacher). New Guinea: Irian Jaya; Papua New Guinea; 1 <$, P. N. G., Katow (MHN, Geneva)
(paralectotype of Xiphidium modestum Redtenbacher). New Hebrides: Aneityum; Efate; Erromango;
Espiritu Santo; Malekula; Pentecost. New Ireland: 1 9, Anir I., x. Norfolk I.: 1 $; 2 tf, 2 9, ii-xii; 1 9,
76 m, flying over creek, wet weather, iv; 1 9> Emily Bay, ii; 1 $, Kingatan, ii. Samoa: American, Tutuila;
Western, Alafua; W., Savaii; W., Upolu; 3 9» w-» Upolu (Rechinger) (NM, Vienna) (paralectotypes of
Xiphidium (Xiphidion) modestum upoluense Karny). Solomon Is.: 1 (51, vii-viii; 1 tf, Bougainville, N. end, vi;
1 9, B., Buin, vi; 1 rf, 1 9, B., Buin (Kangu), 1-50 m, v; 4 9, B., Numa Numa, c. 5° 53' S, 155° 15' E, vi; 1
9, B., Tokinoitu, 20 m, vi; 1 9, Buka, Agric. Sta., xii; 1 1 tf, 5 9, Guadalcanal, xi-xii; 1 tf, G., Honiara, ii; 1
9, G., Honiara District, viii; 7 tf, 3 9, G., H. D., Ilu, ii-v; 1 tf, 1 9, G., H. D., Tenaru, ii-vii; 2 tf, G.,
Kukum, xi-xii; 1 9» G., Lunga, viii; 1 tf, G., R. Poha, 5 m, vii; 1 $, G., Savo I., xii; 1 $, Malaita, Auki,
2-20 m, ix; 6 tf, 6 9, New Georgia Group, Gizo, 30 m, vii; 2 <$, 2 9, Nggela, Tulagi, ii-iii; 1 tf, Olu Malau
Is. (Three Sisters Is.), iv; 1 $, 1 9» Ontong Java, Peku, 0-10 m, xii; 1 9, Russell Is., Lingatu, vi; 1 9, Small
Malaita, near Maramasike Passage, xi. Tonga Is.: 1 ^ (MHN, Geneva) (paralectotype of Xiphidium
modestum Redtenbacher); Eua; Niuatoputapu; Tongatapu. (BMNH; DSIR, Auckland; ANIC, Canberra;
BPBM, Honolulu; DPI, Konedobu.)
Extralimital material. Cocos Keeling Is. Lesser Sunda Is.
DISTRIBUTION (Table 1). This species occurs widely in the Pacific. I have found a record of the
species from New Zealand to be a misidentification for C. albescens, from the measurements and
other characters given.
Conocephalus albescens (Walker)
(Figs 13, 42)
Xiphidium albescens Walker, 1869: 275. Holotype 9, AUSTRALIA (BMNH) [examined].
Xiphidium latifrons Redtenbacher, 1891:526. LECTOTYPE 9, AUSTRALIA (MHN, Geneva), here
designated [examined]. Syn. n.
[Conocephalus modestus (Redtenbacher); Hudson, 1972:254. Misidentification.]
DIAGNOSIS. $ 9- Fastigium of vertex wide. Prosternum bispinose. Macropterous or brachypterous.
Forewings with dark pigmentation in costal and precostal areas, with or without comparatively small dark
spots elsewhere (similar to Fig. 30); light and dark pigmentation strongly contrasted. Left forewing, and
sometimes right, with small dark line or spot towards base of posterior margin. Cross-veins of costal and
precostal areas of forewings irregular. Stridulatory file of left male forewing shaped as in Figs 45 and 46.
Hind femora unarmed ventrally. Hind tibiae usually with 4 apical spurs (ventral pair missing). Male tenth
abdominal tergite with apex produced perpendicularly downwards (Fig. 42). Male cerci very slender in apical
part, with one internal spine and one minute tubercle or swelling which is sometimes absent (Fig. 13).
Ovipositor long, relatively straight.
MEASUREMENTS
Males Females
Fastigium width (30): 0-65-0-77 (0-71) (28): 0-68-0-95 (0-77)
Median length of pronotum (31): 2-9 - 3-4 (3-12) (27): 2-8 - 3-6 (3-23)
Forewing length (macropterous) (27): 15-1 -18-5 (16-48) (22): 15-7 -19-6 (17-44)
(brachypterous) (4): 8-5 -11-0 (9-95) (5): 8-0 - 9-7 (8-68)
PACIFIC SPECIES OF Conocephalus 343
Males Females
Length of stridulatory area (macropterous) (26): 1-87— 2-14 (1-99)
(brachypterous) (4): 1-58- 1-85 (1-70)
Hind femur length (26): 11-6 -13-9 (12-54) (27): 12-3 -17-0 (14-21)
Ovipositor length (26): 11-8 -17-1 (13-95)
DISCUSSION. This species is very similar to C. upoluensis, differing, however, in the shape of the
male cerci and tenth abdominal tergite, and the long ovipositor. Two of the syntypes ofXiphidium
modestum (name replaced by C. upoluensis q. v.) are C. albescens, and C. albescens has been
mistaken in the subsequent literature for C. upoluensis.
Of the specimens examined 84% were macropterous, including the holotype of Xiphidium
albescens. The type-specimens ofXiphidium latifrons are brachypterous. I have synonymized these
two names since their type-specimens are otherwise morphologically identical. I have selected a
lectotype from the two female syntypes of X. latifrons I have examined. The rest of the syntype-
series, consisting of at least one male from Australia: Sydney, is lost (see Weidner, 1966:243).
MATERIAL EXAMINED (macropterous unless otherwise stated)
Xiphidium albescens Walker, holotype 9> Australia: South Australia (Bakewell) (BMNH). Xiphidium
latifrons Redtenbacher, lectotype 9> brachypterous, Australia: New South Wales, Clarence River (MHN,
Geneva).
Australia: 1 $, New South Wales (Thorey) (NM, Vienna) (paralectotype of Xiphidium modestum
Redtenbacher); 22 tf, 30 9, N. S. W., Cabramatta, i-xii; 1 tf, 2 9, N. S. W., Cabramatta, Georges R.
Valley, i; 1 $, N. S. W., Blue, Wentworth Falls, iii; 1 tf, brachypterous, N. S. W., Canley Vale, ii; 1 9, N. S.
W., Casula, iii; 1 9> brachypterous, N. S. W., Clarence River (MHN, Geneva) (paralectotype ofXiphidium
latifrons Redtenbacher); 1 <$, brachypterous, N. S. W., Coonabarabran, i; 1 9> N. S. W., Deep Creek,
Waitpinga, ii; 1 tf, 1 9» brachypterous, N. S. W., Narrabri, iii; 2 9, brachypterous, N. S. W., near Sydney,
Gundaman, National Park, iv; 1 ^, N. S. W., Villawood, iii; 1 <$, brachypterous, Queensland, near Brisbane,
Mogill, ix (BPBM, Honolulu). Lord Howe I.: 1 $ (Stevens) (NM, Vienna) (paralectotype of Xiphidium
modestum Redtenbacher). New Zealand: 1 9» brachypterous, Northland, 48 km W. of Whangarei,
Tangowahine, iv (BPBM, Honolulu). (In BMNH unless otherwise stated.)
DISTRIBUTION (Table 1). Known only from east to south-eastern Australia, Lord Howe I. and
New Zealand.
Conocephalus bilineatus (Erichson)
(Fig. 14)
Xiphidium bilineatum Erichson, 1842:249. Holotype tf, AUSTRALIA (MNHU, Berlin) [examined].
Xiphidion immaculatum Karny, 1911: 343. Holotype 9> AUSTRALIA (NM, Vienna) [examined]. Syn. n.
DIAGNOSIS. $ 9- Fastigium of vertex of medium width. Prosternum bispinose. Macropterous or
brachypterous. Forewings with darker pigmentation in the costal and precostal areas as in Fig. 29. Cross-
veins of costal and precostal areas of forewings fairly irregular. Hind femora unarmed ventrally. Hind tibiae
with 4 apical spurs (ventral pair missing). Male cerci as in Fig. 14, with one internal spine. Ovipositor of
medium length, relatively straight.
MEASUREMENTS
Males Females
Fastigium width (3): 0-47-0-57 (0-51) (3): 0-52-0-55 (0-53)
Median length of pronotum (2): 2-8 - 3-0 (2-85) (3): 2-8 - 3-3 (3-03)
Forewing length (macropterous) (1): 15-0 (2): 14-5 -17-0 (15-75)
(brachypterous) (2): 5-0 - 5-2 (5-10) (1): 2-3
Length of stridulatory area (brachypterous) ( 1 ) : 1-7
Hind femur length (2): 10-5 -11-6 (11-05) (3): 11-7 -12-7 (12-20)
Ovipositor length (3): 10-5 -10-6 (10-57)
DISCUSSION. This species has features in common with C. albescens and C. upoluensis, but it is
distinguished from them by the narrower fastigium of the vertex together with the forewing
pigmentation, and by the shape of the male cerci. The forewing pigmentation is sometimes very
faint. Macropterous and brachypterous specimens were equally represented in the small sample
examined.
344 LINDA M. PITKIN
Comparison of the holotype and other material of C. bilineatus and Xiphidion immaculatum
has shown them to be conspecific, differing only in that the former is brachypterous and the latter is
macropterous.
MATERIAL EXAMINED (macropterous unless otherwise stated)
Xiphidium bilineatum Erichson, holotype $, brachypterous, Australia: Tasmania (Schayer) (MNHU,
Berlin). Xiphidion immaculatum Karny, holotype i , Australia: Victoria, Dandenong Ranges (NM, Vienna).
Australia: 1 9» Capital Territory, Mt Gingera, Snowy Flat, 1520 m, i (BMNH); 1 tf, brachypterous, New
South Wales, Forest Reefs (BMNH); 1 9, brachypterous, Tasmania, Swansea (BMNH). New Zealand: 1 $,
South I., Tasman Valley, Glentanner Station, 610m, tussock grass and low plants on arid hillside, ii
(BMNH).
DISTRIBUTION (Table 1). This species is known only from south-eastern Australia and New
Zealand.
Conocephalus maculatus (Le Guillou)
(Figs 15, 31)
Xiphidion maculatum Le Guillou, 1841:294. Holotype or syntypes, SULAWESI: Makassar (lost). NEOTYPE
c?, WEST MALAYSIA (BMNH), here designated [examined].
Locusta (Xiphidium) lepida de Haan, 1842: 189. LECTOTYPE tf, JAVA (RNH, Leiden), here designated
[examined]. [Synonymized by Redtenbacher, 1891:515.]
Xiphidium continuum Walker, 1869:271. Holotype 9, SIERRA LEONE (BMNH) [examined]. Syn. n.
Xiphidion neglectum Bruner, 1920: 123, Holotype tf, CAMEROUN(ANS, Philadelphia) [examined]. Syn. n.
DIAGNOSIS. $ 9- Fastigium of vertex moderately wide. Prosternum bispinose. Macropterous. Forewings
pigmented with comparatively large dark spots; no markings in costal and precostal areas (Fig. 31). Cross-
veins of costal and precostal areas of forewings irregular to almost regular. Stridulatory file of left male
forewing shaped as in Fig. 47. Hind femora unarmed ventrally. Tibiae marked with spot towards base, dark
on mid tibiae, often faint on others. Hind tibiae with 6 apical spurs. Male cerci as in Fig. 15, with one internal
spine. Ovipositor very short, relatively straight.
MEASUREMENTS
Males Females
Fastigium width (20): 0-51- 0-72 (0-59) (21): 0-62- 0-72 (0-66)
Median length of pronotum (20): 2-7 - 3-3 (2-97) (22): 2-9 - 3-4 (3-13)
Forewing length (20): 13-0 -19-0 (15-70) (21): 14-9 -21-4 (17-64)
Length of Stridulatory area (23): 1-49- 1-92 (1-75)
Hind femur length (20): 11-2 -14-3 (12-84) (20): 12-6 -15-8 (14-02)
Ovipositor length (22): 6-6 - 8-2 (7-33)
DISCUSSION. C. maculatus may be easily recognized by the distinctive pigmentation, particularly
of the forewings.
I believe the type-specimen(s) of Xiphidion maculatum to be lost since they are not in the original
depository (MNHN, Paris). Supposed type-specimens of this were sent to me by Dr Donskoff of
the MNHN, Paris, but differed irreconcilably from the description. I have selected a male neotype
from West Malaysia in preference to any specimens available from nearer the original type-locality
in Sulawesi, since the West Malaysian specimen is in a better state of preservation.
The type-series of Locusta (Xiphidium) lepida consists of 1 <$, 1 9> °f which I have examined
the male and selected it as the lectotype.
Redtenbacher (1891:515) listed Xiphidium continuum as a possible synonym of Xiphidium
maculatum. I have compared the holotype of X. continuum with the neotype and many other
specimens of C. maculatus and confirm the synonymy, although the holotype of X. continuum has
a narrower fastigium of the vertex (0*49 mm) than the specimens examined from the Pacific.
Comparison of the holotype of Xiphidion neglectum with the neotype of C. maculatus has shown
them to be conspecific, confirming the unpublished opinion of my colleague Mr J. Huxley. X.
neglectum has a slightly narrower fastigium (0-46 mm) than the Pacific material I have examined,
and the pigmentation is rather pale but there are no differences warranting a separate specific
identity.
PACIFIC SPECIES OF Conocephalus 345
Xiphidium sinense Walker (1871:35) was synonymized with C. maculatus by Hebard
(1922a:243). I have examined the two male syntypes of the former, from Hong Kong and China,
and found one (from Hong Kong) to be C. maculatus and the other (from China) to be a clearly
different species. I am here designating the male from China as LECTOTYPE, so that X. sinense
(sp. rev.) is no longer synonymous with C. maculatus.
MATERIAL EXAMINED
Xiphidion maculatum Le Guillou, neotype tf, West Malaysia: Klang Gates, 25.ii.1934 (Miller) (BMNH).
Locus ta (Xiphidium) lepida de Haan, lectotype J1, Java (RNH, Leiden). Xiphidium continuum Walker,
holotype 9> Sierra Leone ('pres. Morgan') (BMNH). Xiphidion neglectum Bruner, holotype <$, Cameroun:
Batanga, iv.1914 (Hope) (ANS, Philadelphia).
Australia: 1 tf, Northern Territory, 7km ESE. of Nhulunbuy, 12° 17' S, 136° 50' E, v; 1 tf, 4 9,
Queensland, Cairns; 1 <$, Q., Cairns, Freshwater Creek, ii; 1 <$, Q., Redlynch, ix. New Guinea: 2 <$, 1 9,
Irian Jaya, Kulima, 1400 m, ii; 1 tf, 1 9, 1. J., Swart Valley, W. side, 1400-2000 m, xi; 1 tf, 2 9, 1. J., W. of
Swart Valley, Kutsime, 1500 m, xi; 1 tf, 2 9, 1. J., Wamena, 1700 m, ii; 1 9, Papua New Guinea, Aiyura,
xii; 1 tf, 1 9, P. N. G., Asaro V., Miramar, 1800 m, vi; 2 9, P. N. G., Bena R., 1500-1550 m, ix; 1 tf, P. N.
G., Bisianumu, viii; 1 9, P. N. G., Central District, 2 km NW. of Bomana War Cemetery, 9° 24' S, 147° 14'
E, iv; 1 9, P. N. G., C. D., Brown River, Karema, 9° 12' S, 147° 14' E, viii; 1 tf, P. N. G., C. D., Idlers Bay
(picnic area), 9° 29' S, 147° 05' E, iv; 1 <$, P. N. G., C. D., E. of Port Moresby, Otomata Plantation, 1 m,
xi; 1 tf, 2 9, P. N. G., C. D., Sogeri Plateau, 4 km SSE. of lorowari, 9° 27' S, 147° 26' E, iv-vii; 1 tf, 1 9,
P. N. G., Chimbu District, Bomai, vi; 1 9, P. N. G., Finschhafen, iv; 1 9, P. N. G., Garaina, 830 m, i; 3 9,
P. N. G., Goroka, 1550m, v-vi; 2 9, P. N. G., 24km E. of Kainantu, viii; 1 9, P. N. G., 30km S. of
Kainantu, Sinofi, 1590 m, x; 1 tf, P. N. G., Lae, vii; 1 9, P. N. G., Maprik, x; 4 3, 3 9, P. N. G., Morobe
District, Gusap Downs Sta., Ramu-Markham Valley, ii; 1 9, P. N. G., M. D., Wau, 1200 m, iv; 1 9, P. N.
G., Musgrave R., near Javarere, Daradae, x; 1 tf, 1 9, P. N. G., 9 km N. of Port Moresby, 9° 25' S, 147°
08' E, iv; 1 9, P. N. G., Port Moresby, Lawes Rd, at lights, iii; 1 9, P. N. G., Sepik District, Dreikikir,
350m, vi; 1 tf, P. N. G., Subitana-Musgrave District, iv; 5 tf, 3 9, P. N. G., Swart Valley, Karubaka,
1450 m, xi; 1 tf, P. N. G., Western District, Daru, 9° 05' S, 143° 12' E, v; 1 <$, 1 9, P. N. G., W. D., Lake
Murray (Patrol Post), iv; 1 tf, P. N. G., W. D., Mabaduan, x; 1 9, P. N. G., W. D., 2-3 km S. of Morehead,
8° 44' S, 141° 38' E, viii; 1 rf, P. N. G., W. D., 6 km N. of Morehead, 8° 39' S, 141° 38' E, ix; 1 9, P. N.
G., W. D., R. Morehead, Rouku, iii-v; 1 tf, P. N. G., W. D., Oriomo Govt Sta., x; 3 $, 1 9, P. N. G., W.
Highlands, Mt Hagen Valley, Keltiga, 1710m, ix-x. (BMNH; ANIC, Canberra; BPBM, Honolulu; DPI,
Konedobu.)
Extralimital material. Ethiopian Region. Malagasian Region. Hong Kong, 1 $ ('pres. Bowring') (BMNH)
(paralectotype of Xiphidium sinense Walker). West Malaysia. Indonesia. Philippines.
DISTRIBUTION (Table 1). This species is widely distributed throughout the tropical regions of the
Old World, its range extending eastwards to Australia and New Guinea.
Conocephalus laetus (Redtenbacher)
(Figs 16, 39)
Xiphidium laetum Redtenbacher, 1891:514. Holotype 9, AUSTRALIA (NM, Vienna) [examined].
Conocephalus dubius Willemse, 1942:95. Holotype <$, AUSTRALIA (NM, Basle) [examined]. Syn. n.
Conocephalus raggei Harz, 1970: 194. Holotype tf, UGANDA (BMNH) [examined]. Syn. n.
DIAGNOSIS. $ 9- Fastigium of vertex moderately narrow. Prosternum bispinose. Macropterous. Forewings
unicolorous. Cross-veins of costal and precostal areas of forewings irregular. Stridulatory area of left male
forewing large (Fig. 39). Stridulatory file of left male forewing shaped as in Fig. 47. Hind femora unarmed
ventrally. Hind tibiae with 6 apical spurs. Male cerci as in Fig. 16, with one internal spine. Ovipositor long,
straight.
MEASUREMENTS
Males Females
Fastigium width (10): 0-42-0-52 (0-46) (12): 0-47-0-52 (0-51)
Median length of pronotum (11): 3-1-3-4 (3-26) (14): 3-3-4-2 (3-68)
Forewing length (11): 19-1 -22-2 (20-75) (14): 21-7 -26-6 (24-03)
Length of Stridulatory area (10): 2-72- 3-20 (3-00)
Hind femur length (9): 12-6 -15-5 (13-94) (12): 14-7 -18-5 (16-29)
Ovipositor length (14): 13-7 -17-7 (15-30)
346 LINDA M. PITKIN
DISCUSSION. The large stridulatory area of the left male forewing and the long ovipositor
distinguish C. laetus from the other species included in the present study with unicolorous
forewings and unarmed hind femora.
The type-specimens of C. dubius and C. raggei were compared with the holotype and other
material of C. laetus, and were found to be conspecific with them, despite the wide geographical
separation. The holotype of C. raggei has a forewing length of 18-7 mm, slightly shorter than in
any of the Pacific specimens examined, but shows no other morphological differences.
C. laetus closely resembles C. gladiatus (Redtenbacher), an oriental species which does not
occur in the Pacific. I have found no consistent character for separating these two species. The
main difference between them is in the ovipositor, which is considerably longer in C. gladiatus, and
wing length, which is shorter in C. gladiatus. Many Oriental and Austro-oriental specimens which I
have examined have both short wings (as in C. gladiatus) and a short ovipositor (as in C. laetus) or
vice versa. The fastigium of the vertex of these specimens varies in width. In C. laetus the fastigium
is moderately narrow, and it appears to be variable in C. gladiatus. I have omitted the specimens of
doubtful identity from the list of material examined.
A minute, barely discernible, pointed swelling is sometimes present on the male cerci, at the base
of the internal spine.
MATERIAL EXAMINED (25 Pacific specimens)
Xiphidium laetum Redtenbacher, holotype 9> Australia: Queensland, Port Curtis ('Darnel ded., coll. Br. v.
W.') (NM, Vienna). Conocephalus dubius Willemse, holotype $, Australia: Northern Territory, Burnside,
iv.1931 (Handschiri) (NM, Basle). Conocephalus raggei Harz, holotype $, Uganda: Lake George, short
grass plains, iii.1936 (Johnston) (BMNH).
Australia: 1 9> Northern Territory, Burnside, iv.1931 (Handschiri) (NM, Basle) (allotype of Conocephalus
dubius Willemse); Queensland (BMNH; BPBM, Honolulu). New Guinea: 1 <$, 1 9» I"an Jaya, Sentani,
90+ m, vi (BMNH); 1 9, Papua New Guinea, Bulolu, 800-900 m, xi (BPBM, Honolulu); 1 <J, P. N. G.,
Kokoda, 370m, viii (BMNH); 2 $, 1 9, P. N. G., Morobe District, Gusap Downs Sta., Ramu-Markham
Valley, ii (ANIC, Canberra); 1 $, P. N. G., Morobe District, Gusap Downs Sta., Ramu-Markham Valley, ii
(BMNH); 1 tf, P. N. G., Tapini, v (BPBM, Honolulu); 2 rf, 1 9, P. N. G., Western District, Daru, 9° 05' S,
143° 12' E, viii (ANIC, Canberra); 1 9, P. N. G., W. D., Morehead, 8° 43' S, 141° 38' E, viii (ANIC,
Canberra).
Extralimital material. Celebes. N. Moluccas. Philippines. Central African Republic. Mali (including 6 $, 1
9, paratypes of C. raggei Harz). Nigeria (including 1 9 paratype of C. raggei Harz). Uganda (including 4 $,
6 9 paratypes of C. raggei Harz). Zaire (1 $, 8 9 paratypes of C. raggei Harz).
DISTRIBUTION (Table 1). In the area of the present study, this species was previously known from
Australia, and has been recorded from the New Hebrides although I have not seen specimens from
there; it has now also been found in New Guinea. It occurs west of the Pacific in the Austro-
Oriental Region, in tropical Africa, and possibly in the Oriental Region. This is a similar
distribution pattern .to that of C. maculatus which is, however, a much more common species.
Conocephalus angustivertex sp. n.
(Figs 17, 24, 32, 55)
DESCRIPTION AND DIAGNOSIS. $ 9. Fastigium of vertex narrow, slightly less than half width of basal
antennal segment. Prosternum bispinose. Macropterous or brachypterous. Forewings with broad band of
dark pigmentation as in Fig. 32, and pigmentation in costal and precostal areas. Cross- veins of costal and
precostal areas of forewings moderately irregular. Stridulatory file of left male forewing shaped as in Fig. 47.
Hind femora with 1-8 externoventral spines, and frequently with 1-4 internoventral spines. Hind tibiae
usually with 5 apical spurs (internodorsal spur missing or, exceptionally, present but small). Male tenth
abdominal tergite with two comparatively large median projections at apex. Male cerci as in Fig. 1 7, with one
internal spine. Titillators as in Fig. 55; weakly sclerotized. Female subgenital plate with excised apex.
Ovipositor moderately short, gently curved (Fig. 24).
General coloration brown, with wide dark brown stripe along top of body from fastigium of vertex to end
of abdomen. Abdomen, and usually pronotum, with lateral stripes, occasionally fusing with dorsal stripe;
lateral stripes often slightly lighter than dorsal stripe although darker than ground colour. Lateral pronotum
stripes narrower than dorsal stripe. Femora frequently with small dark brown or reddish spots.
PACIFIC SPECIES OF Conocephalus 347
MEASUREMENTS
Males Females
Total length (macropterous) (4): 26-2 -29-5 (28-25) (9): 28-3 -31-9 (29-71)
(brachypterous) (6): 11-3 -15-0 (12-60) (7): 13-5 -14-7 (14-01)
Fastigium width (11): 0-27-0-36 (0-31) (16): 0-27-0-38 (0-33)
Median length of pronotum (11): 2-8 - 3-2 (2-95) (16): 3-0 - 3-3 (3-10)
Forewing length (macropterous) i (4): 18-4 -21-2 (20-14) (9): 20-0 -23-8 (21-21)
(brachypterous) (7): 3-1 - 4-1 (3-69) (7): 2-1 - 4-1 (2-48)
Hind femur length (10): 12-5 -14-4 (13-39) (14): 13-7 -14-7 (14-16)
Ovipositor length (16): 7-9 - 9-2 (8-42)
DISCUSSION. C. angustivertex may be distinguished from all other species of the genus by the
absence of the internodorsal hind tibial spurs, together with the shape of the male cerci and female
ovipositor, and the forewing pigmentation. Macropterous and brachypterous specimens were
nearly equally represented in the sample studied. The right forewing of the brachypterous males
was usually slightly longer (by 0-4 mm on average) than the left forewing. The brachypterous
specimens had rounded forewing apices with the exception of one female with sharply pointed
forewings which were longer than those of the other brachypterous females. One shrivelled,
possibly teneral, male specimen had colourless forewings.
MATERIAL EXAMINED
Holotype J1, macropterous, New Guinea: Papua New Guinea, 24 km S. of Paup, Torricelli Mts, 520 m,
4-1 l.iii. 1939 (Moore) (BMNH).
Paratypes. New Guinea: 1 9> macropterous, Irian Jaya, 40 km N. of Baliem Valley, Bokondini, c. 1300 m,
16-23.xi.1961 (Quote) (BPBM, Honolulu); 1 tf, 1 9, brachypterous, I. J., Mt Cyclops, 1070m, iii.1936
(Cheesman) (BMNH); 2 9, brachypterous, I. J., Cyclops Mts, Mt Lina, 1070m, iii.1936 (Cheesman)
(BMNH); 2 tf, 4 £, brachypterous, I. J., Cyclops Mts, Mt Lina, 1070-1370 m, iii.1936 (Cheesman)
(BMNH); 1 (51, brachypterous, Papua New Guinea, Eastern Highlands District, 25 km ENE. of Kainantu,
Kassam Pass, 16.ii.1976 (Farrow) (ANIC, Canberra); 1 <$, brachypterous, P. N. G., NE., Swart Valley,
Karubaka, 1450 m, 5.xi.l958 (Gressitf) (BPBM, Honolulu); 1 9, macropterous, P. N. G., NE., Upper Jimmi
Valley, Tsenga, 14.vii.1955 (Gressitf) (BPBM, Honolulu); 1 tf, brachypterous, P. N. G., NE., Upper Jimmi
Valley, Tsenga, 1200 m, 15.vii.1955 (Gressitf) (BPBM, Honolulu); 1 $, macropterous, P. N. G., NE., Upper
Jimmi Valley, Wana, 1500 m, ll.vii.1955 (Gressitf) (BPBM, Honolulu); 1 tf, brachypterous, P. N. G., NE.,
Wana, Korop, 1500 m, 12.vii.1955 (Gressitf) (BPBM, Honolulu); 2 tf, 6 9, macropterous, P. N. G., 24 km
S. of Paup, Torricelli Mts, 520 m, 4-1 l.iii. 1939 (Moore) (BMNH); 1 9, macropterous, P. N. G., Torricelli
Mts, Afua, 15-30 m, 2.iv.l939 (Moore) (BMNH).
DISTRIBUTION (Table 1). Known only from New Guinea.
Conocephalus saltator (Saussure)
(Figs 18, 36, 38, 40, 44)
Xiphidium saltator Saussure, 1859:208. Holotype 9, GUYANA (MHN, Geneva) [examined].
Xiphidiwn meridionale Scudder, 1875:460. Holotype 9» BRAZIL (ANS, Philadelphia) [examined].
[Synonymized by Rehn & Hebard, 19156:269.]
Xiphidium propinquum Redtenbacher, 1891:522. LECTOTYPE tf, ST VINCENT (BMNH), here designated
[examined]. [Synonymized by Rehn & Hebard, 19156:269.]
Xiphidium brachypterum Redtenbacher, 1891:523. LECTOTYPE g, BRAZIL (NM, Vienna), here
designated [examined]. [Synonymized by Rehn & Hebard, 19156:269.]
Xiphidium varipenne Swezey, 1905:212. LECTOTYPE tf, HAWAIIAN Is. (HSDA, Honolulu), here
designated [examined]. [Synonymized by Hebard, 19226:347.]
DIAGNOSIS. $ 9. Fastigium of vertex of medium width. Prosternum bispinose. Macropterous or
brachypterous. Forewings unicolorous. Cross-veins of costal and precostal areas of forewings moderately
regular and parallel. MA of forewings shaped as in Fig. 36. Stridulatory file of left male forewing shaped as in
Fig. 47. Hind femora comparatively sharply swollen towards base (Fig. 38); with 1-6 externoventral spines,
or occasionally unarmed. Hind tibiae with 6 apical spurs (Fig. 40). Male cerci shaped as in Fig. 18, with one
internal spine. Female subgenital plate with truncate apex (Fig. 44). Ovipositor of medium length, relatively
straight.
348 LINDA M. PITKIN
MEASUREMENTS
Males Females
Fastigium width (31): 0-42-0-63 (0-51) (20): 0-44-0-63 (0-56)
Median length of pronotum (31): 2-7 - 3-4 (3-04) (20): 3-1 - 3-9 (3-41)
Forewing length (macropterous) (9): 13-9 -20-0 (18-06) (4): 17-0 -20-1 (18-99)
(brachypterous) (19): 4-5 - 8-2 (6-06) (14): 3-5 -12-3 (5-28)
Length of stridulatory area (macropterous) (11): 1-37- 1-47 (1-39)
(brachypterous) (18): 1-01- 1-27 (1-16)
Hind femur length (23): 10-7 -14-8 (12-89) (18): 11-0 -15-4 (14-51)
Ovipositor length (21): 10-7 -13-2 (11-79)
DISCUSSION. C. saltator differs from all the other species in the present study in the shape of the
male cerci, with the internal spine positioned towards the base. The species is of Neotropical origin
and has no close relatives in the Pacific. The females, however, are similar in appearance to those of
C. longipennis and C. oceanicus. They may be distinguished particularly from the former by the
truncate apex of the subgenital plate. Females of C. saltator also differ from C. longipennis in the
fastigium width, and from C. oceanicus in the shape of the hind femora and the forewing venation.
The male cerci of C. saltator show some variation in length. The shape of the male cerci gives
rise to a slight rounded swelling above the base of the internal spine. One specimen examined had
only five apical spurs on one hind tibia due to the fusion of one ventral and one lateral spur.
I have selected a lectotype from the 2 $ syntypes of Xiphidium propinquum I have examined.
There are additional syntypes from Guatemala (Dohrri) and Venezuela: Merida (B runner), which I
have not seen. I have also selected a male lectotype from the 2 $, 1 9 syntypes of Xiphidium
brachypterum I have examined. The length of the stridulatory area of this specimen is 1-38 mm,
longer than in the brachypterous Pacific specimens examined. There are other syntypes of this
species which I have not examined: Brazil: 1 <$; Colombia: 1 9; Venezuela: 1 9 (all NM, Vienna);
Peru: 1 9 (MHN, Geneva).
I have selected a lectotype from 3 (^, 1 9 specimens examined which I consider to be probable
syntypes of Xiphidium varipenne. This was described from 'numerous specimens collected in
Honolulu and elsewhere in the Hawaiian Islands'.
The holotype of Xiphidium saltator has an ovipositor length of 10-0 mm, shorter than in the
Pacific specimens examined.
MATERIAL EXAMINED (52 Pacific specimens, brachypterous unless otherwise stated)
Xiphidium saltator Saussure, holotype 9» macropterous, Guyana (MHN, Geneva). Xiphidium meridionale
Scudder, holotype 9> Brazil (Janson) (ANS, Philadelphia). Xiphidium propinquum Redtenbacher, lectotype
cf, Windward Is.: St Vincent, Windward side (Smith) (BMNH). Xiphidium brachypterum Redtenbacher,
lectotype $, Brazil: Rio de Janeiro, Cachoeira (Meyer-Dury) (NM, Vienna). Xiphidium varipenne Swezey,
lectotype tf, Hawaiian Is.: Oahu, Mt Tantalus, 400m, vii-x.1904 (Giffard) (HSDA, Honolulu).
Cook Is.: 1 <5, macropterous, Mangaia, coast; 1 $, macropterous, M., Makatea; 1 9» macropterous, M.,
Makatea, coast; 1 $, Rarotonga, Ngatangiia, x; Temakatea. Hawaiian Is.: macropterous and
brachypterous, Oahu; 2 $, 1 9» macropterous, O., Mt Tantalus, 400m, vii-x.1904 (Giffard) (HSDA,
Honolulu) (paralectotypes of Xiphidium varipenne Swezey). Line Is.: 2 <$, 1 9> Palmyra, camp and airstrip
area, 2-3 m, iii. Samoa: 1 tf, American, Manua Is., Tau I., E. Tau Village (Luma), 50-200 m, ii; 1 9, A.,
Tutuila, viii; 1 $, A., T., near Alao, vi; 1 <$, A., T., Aua-Afono Trail, ii; 1 9> A., T., Fagatoga Tr., reservoir,
iii; 1 9> A., T., Pango Pango, north side, viii; 1 $, Western, Alafua, ii; 1 $, W., Savaii, Asau, i; 1 $, W.,
Upolu, ii; 1 9, macropterous, 1 tf, 1 9, W., U., Apia, ii-vi; 1 tf, 1 9, W., U., Sliding Rock, iv; 1 9,
macropterous, 1 9, W., U., Utumapu, xii. (BMNH; DSIR, Auckland; ANIC, Canberra; BPBM, Honolulu.)
Extralimital material. Neotropical Region, macropterous and brachypterous; Brazil: 1 $ (NM, Vienna)
(paralectotype of Xiphidium brachypterum Redtenbacher); Peru: 1 9 (MHN, Geneva) (paralectotype of
Xiphidium brachypterum Redtenbacher); Windward Is.: 1 $, St Vincent (Smith) (BMNH) (paralectotype of
Xiphidium propinquum Redtenbacher).
DISTRIBUTION (Table 1). This species was introduced from America to the Hawaiian Is., originally
in the Honolulu area, c. 1890. It was first recorded from the Hawaiian Is. by Bruner in 1895
(Swezey, 1905: 212), whence it has spread gradually, now occurring also in the Line Is., Samoa and
Cook Is.
PACIFIC SPECIES OF Conocephalus 349
Conocephalus longipennis (de Haan)
(Fig. 19)
Locusta (Xiphidium) longipennis de Haan, 1842:189. LECTOTYPE $, SUMATRA (RNH, Leiden), here
designated [examined].
Xiphidium spinipes Stal, 1877:47. Lectotype 9, PHILIPPINES (NR, Stockholm), designated by Sjostedt
(1933:29) [examined]. [Synonymized by Hebard, 1922a:245.]
Xiphidium longicorne Redtenbacher, 1891:513. LECTOTYPE tf, CAROLINE Is. (NM, Vienna), here
designated [examined]. [Synonymized by Hebard, 1922a:245.]
Conocephalus carolinensis Willemse, 1942:98. Holotype 9, CAROLINE Is. (BPBM, Honolulu) [examined].
Syn. n.
Conocephalus carolinensis f. macroptera Willemse, 1942:99. Holotype 9, CAROLINE Is. (BPBM, Honolulu)
[examined]. Syn. n.
DIAGNOSIS. <$ 9- Fastigium of vertex narrow. Prosternum bispinose. Macropterous (occasionally
brachypterous outside Pacific). Forewings unicolorous. Cross-veins of costal and precostal areas of
forewings regular, parallel, varying to fairly irregular in some females. Stridulatory file of left male forewing
shaped as in Fig. 47. Hind femora with 3-7 externoventral spines. Hind tibiae with 6 apical spurs. Male tenth
abdominal tergite nearly unmodified, with apical projections very slight and broadly rounded. Male cerci
shaped as in Fig. 19, with one internal spine with a glofeuiar apex. Female subgenital plate with excised apex.
Ovipositor long, relatively straight.
MEASUREMENTS
Males Females
Fastigium width (6): 0-30-0-37 (0-32) (4): 0-34-0-42 (0-38)
Median length of pronotum (6): 2-8 - 3-3 (3-03) (4): 3-2 - 3-6 (3-31)
Forewing length (6): 9-6-18-2 (12-89) (4): 13-3 -17-6 (14-58)
Hind femur length (5): 12-1 -14-2 (12-91) (3): 13-5 -14-5 (14-03)
Ovipositor length (4): 12-6 -15-9 (14-10)
DISCUSSION. C. longipennis differs from all the other species included in the present study in the
shape of the male cercus and the globular apex of its internal spine. The females can be distin-
guished from the other species with unicolorous forewings and hind femora with spines, by a
combination of the following characters: the long, relatively straight ovipositor, the narrow
fastigium, and the excised apex of the female subgenital plate.
All the Pacific specimens examined were macropterous, although some were only just so. They
showed a considerable variation in wing length and C. carolinensis was originally described as two
forms on the basis of this. Comparison of the type-series of C. carolinensis and C. carolinensis f.
macroptera with the lectotype and other material of C. longipennis has shown them to be
conspecific; C. carolinensis f. macroptera is merely a longer-winged variant. Willemse (1942) gives
a photograph of what he refers to as the female 'type' of C. carolinensis. When I examined this
specimen, which can be easily recognized from the photograph, I found it to be labelled as
'Allotype', while a male specimen in the type-series was labelled as 'Type'. I am nevertheless taking
the photographed female specimen as being the holotype by Willemse's published designation.
One specimen of C. longipennis was found to have an internoventral spine on one hind femur, and
one specimen lacked the internodorsal apical spur on one hind tibia.
I have selected a male lectotype of Locusta (Xiphidium) longipennis from 1 cT» 1 9 syntypes I have
examined. This lectotype has a forewing length of 20 mm, which is longer than in the Pacific
specimens I have seen. One or more syntypes from Borneo: Bandjarmasin are missing.
I have also selected a male lectotype from the 3 <$, 1 9 syntypes of Xiphidium longicorne I have
examined. Other syntypes which I have not seen have the following data: Caroline Is.: — — ; Yap
(lost, see Weidner, 1966:243); Yap (Brunner); Cook Is.: Rarotonga; East Indies.
The lectotype of Xiphidium spinipes exceeds the measurements of the Pacific specimens in the
following: fastigium width 0-53 mm; median length of pronotum 3-8 mm; forewing length at least
21-8 mm (slightly damaged); hind femur length 17-6 mm and 17-9 mm; ovipositor length 16-8 mm.
MATERIAL EXAMINED
Locusta (Xiphidium) longipennis de Haan, lectotype J1, Sumatra: Padang (RNH, Leiden). Xiphidium spinipes
StSl, lectotype 9, Philippines (Semper) (NR, Stockholm). Xiphidium longicorne Redtenbacher, lectotype tf,
350 LINDA M. PITKIN
Caroline Is.: Palau Is. (Semper) (NM, Vienna). Conocephalus carolinensis Willemse, holotype 9, Caroline Is.:
Palau Is. Melokeiok, 6.iv. 1936 (Ono) (BPBM, Honolulu) (mislabelled 'Allotype'). Conocephalus carolinensis f.
macroptera Willemse, holotype 9» Caroline Is.: Ponape, Wone, ll.ii.1936 (Kondo) (BPBM, Honolulu).
Caroline Is.: 1 9» Ponape, 6.iii.l936 (Ono) (NM, Maastricht) (paratype of Conocephalus carolinensis
Willemse); 1 g, P., Roi, 14.ii.1936 (Ono) (BPBM, Honolulu) (allotype of Conocephalus carolinensis f.
macroptera Willemse); 1 9, P., Ronkiti, 4.ii.l936 (Ono) (NM, Maastricht) (paratype of Conocephalus
carolinensis Willemse); 1 $, Truk Is., Dublon, 25.xii. 1 935 (Ono) (NM, Maastricht) (paratype of Conocephalus
carolinensis Willemse); 1 tf, T., Moen, 27.xii.1935 (Ono) (NM, Maastricht) (paratype of Conocephalus
carolinensis Willemse); 1 tf, T., Tank, 3.J.1936 (Ono) (BPBM, Honolulu) (paratype, mislabelled Type', of
Conocephalus carolinensis Willemse). Samoa: 1 <$, American, Tutuila, Fagatoga, i (BPBM, Honolulu).
Extralimital material. Oriental Region; Borneo: 1 J1, 1 9 (Boucard) (NM, Vienna) (paralectotypes of
Xiphidium longicorne Redtenbacher); Java: 1 $ (Candeza) (NM, Vienna) (paralectotype of Xiphidium
longicorne Redtenbacher); Philippines: 1 9 (Semper) (NR, Stockholm) (paralectotype of Xiphidium spinipes
Stal); Sulawesi; Sumatra: 1 9, Padang (RNH, Leiden) (paralectotype ofLocusta (Xiphidium) longipennis de
Haan); West Malaysia.
DISTRIBUTION (Table 1). This species is very restricted in the Pacific, specimens having been seen by
me only from the Caroline Is. and Samoa. The species is, however, widespread in the Oriental and
Austro-Oriental Regions.
Conocephalus willemsei nom. n.
(Figs 20, 26, 47)
Conocephalus ensiferus Willemse, 1942:97. Holotype 9, AUSTRALIA (NM, Basle) [examined]. [Homonym of
Conocephalus ensiferus Bolivar, 1884:88.]
DIAGNOSIS. ^ 9- Fastigium of vertex very narrow. Prosternum bispinose. Macropterous. Forewings
unicolorous. Cross-veins of costal and precostal areas of forewings regular and parallel to rather irregular in
some females. Stridulatory file of left male forewing shaped as in Fig. 47. Hind femora with 1-6 externoventral
spines. Hind tibiae with 6 well-developed apical spurs. Male tenth abdominal tergite with two comparatively
large median projections at apex. Male subgenital plate sometimes with small median notch at apex. Male cerci
shaped as in Fig. 20, with one internal spine with pointed apex. Ovipositor moderately short to long, broad,
shaped as in Fig. 26.
MEASUREMENTS
Males Females
Fastigium width (5): 0-21- 0-36 (0-29) (4): 0-24- 0-38 (0-32)
Median length of pronotum (5): 3-0 - 3-3 (3-09) (5): 3-2 - 4-1 (3-62)
Forewing length (5): 20-8 -25-0 (21-95) (5): 24-2 -30-0 (26-39)
Hind femur length (5): 14-0 -16-4 (15-03) (4): 14-8 -18-0 (16-51)
Ovipositor length (5): 9-3 -15-1 (12-01)
DISCUSSION. C. willemsei differs from all the other species with unicolorous forewings in the present
study in the shape of the male cerci, and the ovipositor. It is similar in some respects to C. longipennis,
from which it differs additionally in forewing length, and it is the longest-winged Conocephalus
species in the Pacific. The few specimens I have been able to examine show considerable variation in
ovipositor length, both extremes being in Australia. The shape of the male cerci also varies from
slender to stout. This species was originally described from a single female as Conocephalus ensiferus
by Willemse in 1942. 1 have found this to be a primary homonym of a species described by Bolivar in
1884, from Peru.
MATERIAL EXAMINED
Holotype 9, Australia: Northern Territory, Brock's Creek, 22.iv.1932 (NM, Basle).
Australia: 1 tf, 2 9, Northern Territory, 22 km WSW. of Borroloola, 16° 08' S, 136° 06' E, iv; 1 tf, N. T.,
5 km NNW. of Cahills Crossing, East Alligator R., 12° 23' S, 132° 57' E, vi; 1 tf, N. T., Humpty Doo, Hi. New.
Guinea: 1 tf, 2 9, Papua New Guinea, Western District, 1-3 km E. of Morehead, 8° 43' S, 14 1 ° 39' E, v; 1 J,
P. N. G., W. D., 2-3 km S. of Morehead, 8° 44' S, 141° 38' E, iv. (BMNH; ANIC, Canberra).
DISTRIBUTION (Table 1). This species was previously known only from N. Australia, but has now
also been found to occur in Papua New Guinea, in the south-western area near to Australia.
PACIFIC SPECIES OF Conocephalus 351
Conocephalus bispinatus sp. n.
(Figs 2 1,25, 56-59)
DESCRIPTION AND DIAGNOSIS. $ 9- Fastigium of vertex very narrow, slightly less than half width of basal
antennal segment. Prosternum unarmed. Macropterous. Forewings faintly smoked towards apex and around
stridulatory area; otherwise unicolorous. Cross-veins of costal and precostal areas of forewings irregular, or
sometimes moderately regular and parallel. Stridulatory file of left male forewing shaped as in Fig. 47.
Tympanal region of fore tibiae tending to be bulbous. Mid femora usually with 1-4 externoventral spines. Hind
femora with 3-8 externoventral spines, and 0-5 internoventral spines (0 in most Borneo specimens). Hind
tibiae with 5-6 apical spurs (internodorsal spur missing, or present but very small). Male tenth abdominal
tergite with 2 slight, broadly rounded, apical projections. Male subgenital plate with V-shaped incision. Female
subgenital plate with approximately truncate apex. Male cerci as in Fig. 21, with 2 internal spines. Titillators as
in Figs 56-59. Ovipositor very short, relatively straight (Fig. 25).
General coloration green, brown, or occasionally straw-coloured. Dorsum of head and pronotum of nearly
all females examined with wide, dark stripe, with narrow, light, lateral borders; also with thin dark line at outer
edge of light borders, on vertex. This pattern very faint or occasionally absent in males. Frons with large, black-
patterned area in nearly all females, absent in males. Hind femora usually with dark brown or red 'herring-bone'
pattern externally. Fore and mid femora usually with spots.
MEASUREMENTS
Type-series from New Guinea
Male Females
Totallength 27-6 (5):32-2 -35-7 (33-54)
Fastigium width 0-26 (6): 0-31- 0-36 (0-34)
Median length of pronotum 3-1 (6): 3-5 - 4-3 (3-68)
Forewing length 19-5 (6): 22-7 -28-2 (24-60)
Hind femur length 12-1 (6): 14-2 -17-7 (15-47)
Ovipositor length (6): 6-7 - 8-5 (7-35)
Extralimital material (excluded from type-series)
Males Females
Totallength (5): 32-6 -37-0 (33-78) (4): 35-8 -37-5 (36-78)
Fastigium width (5): 0-28- 0-32 (0-31) (4): 0-37- 0-40 (0-38)
Median length of pronotum (5): 3-2 - 3-6 (3-42) (4): 3-6 - 4-0 (3-75)
Forewing length (5): 22- 8 -25-2 (23-66) (4): 25-2 -26-7 (25-88)
Hind femur length (4): 13-4 -14-9 (13-91) (4): 15-2 -16-0 (15-50)
Ovipositor length (4): 6-8-7-9 (7-30)
DISCUSSION. C. bispinatus appears to be unique in the genus in almost always having ventral spines
on the mid femora. It is one of the few species of the genus with an unarmed prosternum, and differs
from all the other species of the genus with this feature by the following characters: bispinose male
cerci, 5-6 hind tibial apical spurs, and at least in Pacific specimens of C. bispinatus, internoventral
hind femoral spines. This last character is rarely found in Conocephalus.
The titillators of C. bispinatus were examined in the holotype, the single male available from New
Guinea, and were found to be slender structures with a spinose apex, of a general form commonly
found in Conocephalus. The titillators of specimens from Malaya and Thailand were larger and
distinctly different in shape from those of the holotype, and those from Borneo were even larger and
again strikingly different.
Other differences were evident between the specimens from New Guinea; Malaya, Thailand and
Vietnam; and Borneo. Most specimens from Borneo lacked internoventral hind femoral spines while
these were present in all the other specimens examined. The apex of the male cerci curved downwards
in New Guinea specimens, slightly less so in Borneo specimens, and curved outwards slightly, not
downwards, in specimens from Malaya and Thailand. The venation of the costal and precostal areas
of the forewing was irregular in New Guinea specimens, although fairly regular in males and variable
in females from outside the Pacific area. The tympanal region of the fore tibiae, already bulbous in
females from New Guinea, was even more bulbous in both sexes outside the Pacific, particularly in
specimens from Thailand and Vietnam. One specimen, from Thailand, lacked spines on one mid
femur.
The specimens examined from outside the Pacific region have been excluded from the type-series
352 LINDA M. PITKIN
because of these differences, particularly in the titillators. The specimens from New Guinea and the
two groups of specimens from outside the Pacific may be geographical variants, or they may
represent three distinct species, although closely related in many morphological characters. The
information available, based on small samples of specimens and without song data, is insufficient to
confirm the status of the specimens from outside the Pacific.
MATERIAL EXAMINED
Holotype j, New Guinea: Papua New Guinea, Western District, Morehead, 8° 43' S, 141° 38' E, 29.iv.1971
(Balderson) (A NIC, Canberra).
Paratypes. New Guinea: 1 9> Papua New Guinea, Morobe District, Wau, 1200 m, 15. v. 1962 (Sedlacek)
(BMNH); 1 9, P. N.G., Morobe District, Wau, 1700-1800 m, 17.xi.1961 (Sedlacek) (BPBM, Honolulu); 2 9,
P. N. G., Port Moresby, Mt Lawes, 400 m, 5.iii-12.v.l963 (Brandt) (ANIC, Canberra); 1 9, P. N. G., Port
Moresby, Mt Lawes, 400 m, iv.1963 (Brandt) (BMNH); 1 9, P. N. G. Western District, Morehead, 8° 43' S,
141° 38' E, 28.iv.1971 (Balderson & Baker) (ANIC, Canberra).
Material excluded from the type-series. Borneo: 2 $, Sabah, 1 1 km SSE. of Telupid, R. Karamuak, 60 m,
l-7.ix. 1977 (Bacchus) (BMNH); 1 9, Sabah, 13 km S. of Telupid, Tawai Plateau, 400 m, 8.ix. 1977 (Bacchus}
(BMNH); 1 9, Sarawak, Gunong Mulu National Park, ii-vii.1978 (Collins (R.G.S. Exped. 1977-8))
(BMNH); 1 9» Sarawak, Gunong Mulu National Park, Long Pala (base), 70 m, alluvial, secondary forest, MV
light trap, on batu, canopy, iii.1978 (Holloway et al. (R.G.S. Exped. 1977-8)) (BMNH); 1 J1, Sarawak,
Gunong Mulu National Park, 150 m, helipad, mixed dipterocarp forest, mainly canopy, MV light trap, ii. 1978
(Holloway et al. (R.G.S. Exped. 1977-8)) (BMNH). Thailand: 1 tf, in black light trap, vi.1963 (BMNH).
Vietnam: 1 9, DiLinh, 27.ix- 14.x. 1960 (Yoshimoto) (BPBM, Honolulu). West Malaysia: 1 <$, Kuala Lumpur,
24-31.xii.1958 (Quota) (BPBM, Honolulu).
DISTRIBUTION. This species, known in the Pacific only from Papua New Guinea, extends westwards
to Thailand.
Conocephalus tridens Hebard
(Figs 22, 27)
Conocephalus tridens Hebard, 1933: 127. Holotype c?, MARQUESAS Is. (BPBM, Honolulu) [examined].
DIAGNOSIS. $ 9- Fastigium of vertex narrow. Prosternum unarmed. Macropterous. Forewings often with
narrow band of rather faint pigmentation just below R, together with spots above this vein; sometimes
unicolorous. Cross-veins of costal and precostal areas of forewings regular, parallel. Stridulatory file of left
male forewing intermediate in shape between those shown in Figs 45, 46, and 47. Hind femora unarmed
ventrally. Hind tibiae with 6 apical spurs. Male tenth abdominal tergite widely truncate at apex. Male cerci as in
Fig. 22, with 3 internal spines. Ovipositor short and curved as in Fig. 27.
MEASUREMENTS
Males Females
Fastigium width (7): 0-21- 0-37 (0-28) (7): 0-21- 0-40 (0-32)
Median length of pronotum (8): 3-4 - 4-1 (3-76) (7): 3-6 - 4-2 (3-91)
Forewing length (8): 14-4 -18-7 (16-54) (7): 15-1 -21-0 (17-66)
Hind femur length (8): 11-5 -14-3 (12-55) (7): 12-2 -14-6 (13-06)
Ovipositor length (6): 7-3 - 8-4 (7-77)
DISCUSSION. This species has several distinctive characters which separate it from all the other
Pacific species of Conocephalus, including the form of the male cerci and the strongly curved
ovipositor. Only one of these species, C. bispinatus, shares with C. tridens the unarmed prosternum.
MATERIAL EXAMINED
Holotype $, Marquesas Is.: Eiao, Vaituha, 370m, on Cassia occidentalis, 3.x. 1929 (Adamson) (BPBM,
Honolulu).
Marquesas Is.: 2 $, Eiao, above Vaituha, 240 m, 29.ix.1929 (Adamson) (ANS, Philadelphia) (paratypes); 1
<$, 1 9> E., above Vaituha, 340 m, on Dodonaea viscosa, 2.x. 1929 (Adamson) (ANS, Philadelphia) (paratypes);
1 9, E., 520 m, 16.iv.1931 (Le Bronnec & Tauraa) (ANS, Philadelphia) (paratype); 1 9, E., 520 m, iv (ANS,
Philadelphia); 1 tf, E., c. 550m, on Sida sp., 22.iv.1931 (Le Bronnec & Tauraa) (ANS, Philadelphia)
(paratype); 1 $ nymph, E., 550 m, iv (ANS, Philadelphia); 1 9, E., near centre, 400 m, 1.x. 1929 (Adamson)
(ANS, Philadelphia) (paratype); 1 g, Fatu Hiva, i (BMNH); 1 tf, Hiva Oa, i (BMNH); 1 9, H., Kaava Ridge,
PACIFIC SPECIES OF Conocephalus 353
Kakahopuanui, 610m, x (ANS, Philadelphia); 1 tf, Tahuata, Amatea, 790m, 27.vi.1930 (Le Bronnec &
Tauraa) (ANS, Philadelphia) (paratype); 1 9, T., Amatea, 820 m, 7.vii.l930 (Le Bronnec & Taurad) (ANS,
Philadelphia) (paratype); 1 9 nymph, T., Vaitupaahei, 700 m, vii (ANS, Philadelphia).
DISTRIBUTION (Table 1). This species is endemic in the Marquesas Is.
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Index
Principal references are in bold; invalid names are in italics. References to the Checklist (p. 323) are not
included.
affine 329
albescens 317, 321, 325, 341, 342, 343
Aleurocybotus 317
angustivertex 317, 321, 325, 326, 346
Anisoptera 321
annulicornis 316
antipodum 336, 339
bilineatus 32 1,325, 339, 343
bispinatus 317, 321, 323, 351, 352
bituberculatus 321
brachypterum 347
brevipes 316
brevixiphus 34 1
brunneri 336
carolinensis 349
Chloroxiphidion 321
cinereus 3 1 7
Conocephalus 320, 321
conocephalus 315, 320
consul 316
continuum 344
Decadarchis 316
dorsalis 321
dubius 345
Dymisococcus 316
ensiferus 350
Ereunetis 316
fasciata 321
fasciatus 3 1 7
Fatuhivella321
flavistriata 316
fusca321
geniculare 338
gladiatus 346
immaculatum 343
indicus 3 1 7
infumatus 319, 321, 326, 327, 329, 331, 332, 333
javanicum 321
laetus 32 1,323, 345
latifrons 342
lepida 344
Leptocorisa 316, 317
longicorne 349
longipennis 317, 321, 326, 327, 329, 348, 349, 350
macroptera 349
maculatus 321, 325, 344, 346
maoricum 336, 338
melaenus 316
meridionale 347
modestum 341, 343
modestus 342
PACIFIC SPECIES OF Conocephalus
neglectum 344
Neoxiphidion 321
Nezara 3 1 6
Nukuhivella321
355
oceanicus 317, 319, 321, 326, 327, 329, 331, 332,
333, 335, 340, 348
oratorius 316, 317
Perkinsiella 316
propinquum 347
Pseudococcus 316
raggei 345
redtenbacheri 316, 319, 321, 326, 327, 329, 331,
333,335
Riptortus 316
saccharicida 316
saltator 3 16, 3 17, 3 19, 321, 325, 326, 327, 347
semivittatus 321, 326, 335, 336, 338, 339, 340
sinense 345
spinipes 349
starmuehlneri 319, 321, 326, 327, 329, 330, 332,
333
strictum 321
Thecoxiphidion 321
tridens317, 321,323, 352
trivittatus319, 325, 340
tumidus 319, 325, 326, 335, 338, 339
upoluensis 317, 319, 321, 325, 341, 343
varipenne 316, 347
viridula 316
vittatus 316, 319, 335, 336, 338, 340
willemsei 326, 350
Xiphidion 321
Xiphidium 321
537
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A revision of the western Palaearctic species of the genus Meteorus
(Hymenoptera : Braconidae). By T. Huddleston.
A revision of the Old World Polymorphanisini (Trichoptera : Hydropsychidae).
By P. C. Barnard.
A review of the genera of Neotropical Encyrtidae (Hymenoptera :
Chalcidoidea). By John S. Noyes.
A revision of the British species of Dendrocerus Ratzeburg (Hymenoptera
Ceraphronoidea) with a review of their biology as aphid hyperparasites.
By N. D. M. Fergusson.
A revision of the Pacific species of Conocephalus Thunberg (Orthoptera :
Tettigoniidae). By Linda M. Pitkin.
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