Bulletin of the

British Museum (Natural History)

Entomology series Vol 45 1982

British Museum (Natural History)
London 1982

Dates of publication of the parts

No 1 27 May 1982

No 2 .... 24 June 1982

No 3 . 26 August 1982

No 4 ' . ... 30 September 1982

ISSN 0524-6431

Printed in Great Britain by Henry Ling Ltd, at the Dorset Press, Dorchester, Dorset

Contents
Entomology Volume 45

Page

No 1 A catalogue and reclassification of the Ichneumonidae (Hymen-
optera) described by C. G. Thomson
M. G. Fitton . . 1

No 2 A taxonomic review of the genus Phlebotomus (Diptera: Psychodidae)

D. J. Lewis 121

No 3 Stenomine moths of the Neotropical genus Timocratica (Oecophoridae)

Vitor O. Becker 211

No 4 Afrotropical species of the myrmicine ant genera Cardiocondyla,
Leptothorax, Melissotarsus, Messor and Cataulacus (Formicidae)
Barry Bolton 307

Bulletin of the

British Museum (Natural Hisfory)

A catalogue and reclassification of the
Ichneumonidae (Hymenoptera) described by
C. G. Thomson

M. G. Fitton

Entomology series

Vo\ 45 No 1 21 May 1982

The Bulletin of the British Museum (Natural History), instituted in 1949, is issued in four
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Papers in the Bulletin are primarily the results of research carried out on the unique and
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World List abbreviation: Bull. Br. Mus. nat. Hist. (Ent.)

Trustees of the British Museum (Natural Histo

The Entomology series is produced under the general editorship of the

Keeper of Entomology : Laurence A. Mound

Assistant Editor : W. Gerald Tremewan

ISSN 0524-6431

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

Entomology series
Vol45No 1 pp 1-119

Issued 27 May 1982

/* GENERAL +
* 3 JUN nm

A catalogue and reclassification of the Ichneumonjda£ARY
(Hymenoptera) described by C. G. Thomson \^L #£

M. G.

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

Contents

Synopsis 1

Introduction

C.G.Thomson

Acquisition of Thomson's collections by the University of Lund . 3

Manuscript and other material associated with the collections . . 3

The collection of Ichneumonidae 3

Labelling of specimens 4

Notes on the recognition of type-material and on the selection and designation of

lectotypes g

Thomson's use of names for subgeneric categories 9

Format and arrangement of catalogue 10

Catalogue 10

Nomenclatural summary g7

Species incorrectly attributed to Thomson 100

Acknowledgements 100

References 100

Index 105

Synopsis

The 957 nominal species of Ichneumonidae (all from the western Palaearctic region) described by C. G.
Thomson are catalogued. An attempt is made to account for the type-material of all species and the generic
placements of the species to which the names apply are established after study of the types. Types of 74
species are lost and 9 names remain nomina dubia. Lectotypes are designated for 116 species and 103 new
combinations are established. One neotype is designated and one replacement name is proposed.

Introduction

The Ichneumonidae is one of the largest families of animals. More than 10000 species have been
described from the western Palaearctic region alone. Because of their parasitic habits they are of
great economic importance and biological interest. However, studies of their 'biology' depend
upon a sound and accurate knowledge of their taxonomy. It is unfortunate that the taxonomy of
the western Palaearctic fauna is currently more confused and in need of attention than that of any
other zoogeographical region. The main reason for this is that the results of the outstanding work
over the past forty years by Henry Townes and his co-workers, on the taxonomy and classifi-
cation of the family, have now been applied to all other regions and have wrought order from
chaos. There is a firm base for future systematic studies on the family in these areas. A similar
base, in the form of comprehensive modern 'catalogues', is needed for the western Palaearctic.
The word catalogue is used with some reservation because it tends to convey the wrong im-
pression of the kind and quality of studies needed to produce such works, for a group as large
and as difficult as the Ichneumonidae. This paper on the Thomson species is intended as a
contribution to a complete catalogue of the western Palaearctic Ichneumonidae.

Bull Br. Mus. not. Hist. (Ent.) 45(1): 1-119 Issued 27 May 1982

2 M. G. FITTON

C. G. Thomson is generally acknowledged to have been one of the most able hymenopterists of
his period. He had a talent for distinguishing closely related species and he described a very large
number of new species, including 957 Ichneumonidae, all from Europe and mainly from Sweden.
However, his ability is not fully demonstrated in his publications; he lacked a type-concept; and
he neglected the proper labelling of material. The existence of these deficiencies perhaps helps to
explain how he was able to be so prolific; and, together with the recent revolutionary changes in
the classification and taxonomy of the Ichneumonidae, they now limit seriously the use which can
be made of his work. This paper attempts to place all of the species of Ichneumonidae described
by Thomson in the currently-accepted generic classification of the family. This sort of work must
precede revisionary studies because, if such studies of genera or higher taxa are to have a lasting
value, one of the essential prerequisites is a knowledge of the described species which belong to
them. Because of the vast literature this problem has bedevilled taxonomic work on many groups
of European insects, but it is especially severe in the large and difficult families of parasitic
Hymenoptera such as the Ichneumonidae.

That the work of correctly placing the already-described species of western Palaearctic Ichneu-
monidae cannot be achieved successfully, as revisionary studies are undertaken, can be dem-
onstrated easily by reference to the Thomson species. For instance, in a revision of Dichrogaster,
a small distinctive genus with nine species in Europe, Horstmann (19736) included only two of the
four Thomson species which belong in it (Horstmann, 19766). Thomson originally described
three of these species in Hemiteles and one in Phygadeuon. Recognition of the genuine types of
Thomson's species has also caused problems (the reasons for which are fully explained in the
sections on labelling of specimens and recognition of types). Of about 400 specimens designated
as lectotypes or recognised as holotypes of Thomson species between 1966 and 1978 more than
twenty-five can now be shown not to have been original material of the species concerned and
therefore to be invalid. For example, Aubert (19766: 271) designated as lectotype of Mesoleius
frontatus Thomson a specimen labelled '50', the significance of which was not stated. However,
Aubert had the handwritten label upside down; it was really 'OG', an abbreviation for
Ostergotland. Since the species was described from Ystad in Skane this specimen could not be a
type. These sorts of problems can only be solved by comprehensive studies of all species described
by an author and of his methods, collections and idiosyncracies.

The generic classification of the Ichneumonidae which is the basis of the placements given in
this paper is that published by Townes (1969; 19700; 19706; 1971). This work does not cover the
subfamily Ichneumoninae, in which case Townes, Momoi & Townes (1965) and Perkins (1959;
1960) are followed. Placements of species of Anomaloninae and Ophioninae were made by I. D.
Gauld and follow his work on these groups (Gauld, 1976; 1979). The classification of parts of the
Phygadeuontinae and Tersilochinae takes into account some changes and new genera proposed
by Horstmann (19716; 19746; 1976a; 1978). Aubert (19766), Frilli (1973) and Horstmann (1979a)
have made particular studies of the Thomson species originally described in Mesoleius, Phyga-
deuon and Hemiteles respectively. In these three genera, where I have not felt the need to check,
the generic placements are credited to these authors. All species synonymies are given on the
basis of the published opinions of competent workers (to which references are given).

C. G. Thomson

The following biographical information, relating particularly to Thomson's work on the Hyme-
noptera, is taken mainly from the obituary by Bengtsson (1900).

Carl Gustav Thomson was born in the province of Skane on 13 October 1824 and died in
Lund on 20 September 1899. He succeeded Dahlbom as curator of the entomological collections
at the University of Lund. He was extremely productive: his first paper appeared in 1851 and his
total entomological publications exceeded 8800 pages. Coleoptera were his initial interest but he
soon became involved with the work on Hymenoptera which occupied him until his death. He
was a popular teacher of entomology and students were sometimes given specimens, from his
collections, of the species dealt with in his lectures.

The Proctotrupoidea was the subject of his first important work on Hymenoptera. Between

THE ICHNEUMONIDAE DESCRIBED BY C. G. THOMSON 3

1871 and 1879 he published the five volumes of Hymenoptera Scandinaviae. The Opuscula En-
tomologica, issued in 22 parts between 1869 and 1897, included all of his major work on the
groups of Hymenoptera not covered in Hymenoptera Scandinaviae. He paid for the printing of the
Opuscula Entomologica himself. Publication ceased in 1897 because problems with his eyesight
put an end to his taxonomic work. Of the Hymenoptera, only the Formicidae and Mymaridae
did not receive his attention.

Although he described over 2400 new species (including more than 2100 Hymenoptera) he
apparently regarded his work on higher classification as more important. He dealt mainly with
the Swedish fauna and collected most of the material on which he worked himself. The 'biology'
as well as the morphology of the species interested him and he spent a lot of time in the field. In
summer he went on walking tours, mainly in southern Sweden (including most parts of Skane).
He also visited Blekinge, Halland, Smaland, Oland and Gotland. He was often accompanied by
C. D. E. Roth. At the end of the 1860s he went twice to Norrland and in 1871 visited Jamtland.
He travelled abroad to Germany and in 1872 made a long trip to Germany, Austria, Switzerland
and France, during which he visited many museums and saw important collections (including
those of Fabricius and Hartig).

Thomson did not spend much time preparing specimens, which were often pinned alive in the
field. At the time of his death his collection of Hymenoptera comprised about 80 000 specimens
representing about 7000 species and was housed in 78 cabinet drawers.

Acquisition of Thomson's collections by the University of Lund

Thomson's collections were his own private property. He himself sold his collection of Coleopt-
era (but not the 'duplicate' collection) to Berlin (see Charpentier, 1972). In November 1899, after
his death, his daughter offered the remaining collections for sale to the university. She said that
Thomson had valued the collections at between 20000 and 30000 Kr. but she asked for only
8000 Kr. Together with the written offer to the university she included a synopsis of the col-
lections. The Ichneumonidae occupied 35 cabinet drawers and there were about 30 boxes of
'duplicate' material. The collections were purchased by the university on 23 January 1900.

Manuscript and other material associated with the collections

Thomson's correspondence is deposited in the main university library in Lund. He was in contact
with workers in Sweden and in Europe, including most contemporary ichneumonid specialists.
Many of the letters are accompanied by lists of species.

The library of the Zoological Institute has Thomson's personal copies of the Opuscula En-
tomologica, etc. They contain marginal notes made by Thomson. The notes are more numerous
in the earlier parts and include new synonymy and descriptions of new species. Unfortunately,
they have not proved helpful in tracing the type-material that is apparently missing from the
collection.

The Entomology Museum of the Zoological Institute has little manuscript material that is
relevant to the Ichneumonidae. It includes the letter from Thomson's daughter offering the
collections to the university (see above) and a few lists, including one of ichneumonids from
Holmgren's collection.

The collection of Ichneumonidae

Thomson's 'main' collection of Ichneumonidae is contained at present in parts of two cabinets
(numbered 395 and 396). It occupies 50 drawers (numbered 31 to 80). The arrangement of the
collection follows the Opuscula Entomologica, thus: Ichneumonidae (drawers 31-41), Cryptidae
(41-50), Pimplidae (50-56), Agriotypidae (56), Ophionidae (56-65) and Tryphonidae (66-80).

The 'duplicate' ichneumonid material is contained in various cabinet drawers (in cabinets 398,
399, 403 and 404) and in numerous separate boxes (cigar boxes etc.). The boxes are kept in
cupboard 324. Parts of the duplicate collection (boxes as well as drawers) are arranged taxo-

4 M. G. FITTON

nomically, with labels for genera and species. In some parts the arrangement is tidy and it is
possible to relate specimens to particular labels. In other parts there is a confusion of material.
Some boxes contain material from a single collector (e.g. Lethierry); others contain an assort-
ment. The contents of some boxes are partly sorted and named. The duplicate collection includes
Dahlbom, Ljungh, Holmgren, Wesmael and Zetterstedt material. There is type-material of
Thomson species and there may be type-material of other workers species (notably Holmgren
and perhaps Wesmael). Thomson apparently received a collection of Wesmael ichneumonids
(currently in two drawers in cabinet 399). It is still 'as received', each specimen bearing a number
(1-249). No key to the numbers, and thus Wesmael's identifications, has been found.

The 'main', formal collection was arranged in its present form by Simon Bengtsson. Bengtsson
was appointed curator of the entomological collections in 1900 and one of his first duties was to
take care of the then newly-acquired Thomson collections and transfer the Hymenoptera to three
new cabinets. It is known that the formal collection corresponds to Thomson's own 'main'
collection but that the arrangement may have been changed (if necessary) to correspond with the
Opuscula Entomologica. The 'duplicate' collection appears to be as Thomson left it.

The only significant curatorial work on the collection since Bengtsson's time has been the
recognition and labelling of type-material by specialists and, more recently, the addition of labels
(in the form '1978 329') to specimens sent out on loan. These labels are not removed when the
specimens are returned to the collection and they form, in conjunction with the 'loan journal', a
useful record of borrowers of material. Some years ago a few specimens were labelled 'typ' or
'typi' (e.g. Cteniscus genalis) as part of a curatorial exercise attempting to identify types/syntypes
in the museum collections (not just the Thomson collection) (H. Andersson, pers. comm.).

There are surprisingly large numbers of specimens missing from the collection (deduced from
information on localities, sexes and specimens given in the Opuscula Entomologica). There are
several possible explanations for this but none is supported by more than circumstantial evi-
dence.

There is some Anthrenus damage in the collection but very little evidence of attacks in the
present cabinets. Perhaps badly damaged specimens, including type-material, were discarded at
some time by Thomson, or by Bengtsson at the time of the transfer to the present cabinets.

The collection was undoubtedly a 'working' collection and Thomson may have redetermined
material at various stages, changing its position in the collection. Some such displaced specimens
have already been identified as types.

It is probable that Thomson exchanged material with other European workers. As far as is
known, however, he did not give any of his Swedish material to other Swedish workers (H.
Andersson, pers. comm.), with the possible exception of specimens of common species given to
students (Bengtsson, 1900). He may have returned to other Swedish workers specimens which
they sent to him. He returned to Jensen and to Drewsen material, including types now in
Copenhagen, which they collected in Jutland and Zealand.

It is difficult to assess the effects of the transfer to new cabinets and associated curatorial work
by Bengtsson. Thomson's arrangement was almost certainly not as precise and neat as the
present one. Bengtsson replaced Thomson's handwritten 'cabinet' labels by type-written ones. In
the case of generic names Thomson's labels were concealed beneath the new ones. The species
name labels were folded and transferred to the pin of the first specimen in the species series. The
type-written labels follow the Opuscula Entomologica exactly and include the typographical
errors, e.g. Microcryptus 'arrideus' instead of 'arridens' as on Thomson's own cabinet label and
Exetastes 'guttiferri instead ot'guttifer'. In some cases Thomson's cabinet label name differs from
the published one, e.g. Catoglyptus fusiventris" instead oi'fusiformis1 — presumably he changed his
mind between writing the label and writing the manuscript for publication.

Labelling of specimens

Apart from Thomson's cabinet labels (added to specimen pins by Bengtsson (see note above) and
of no significance whatsoever in the recognition of types) material in the collection is usually very
poorly labelled. The specimen labels are generally small squares of paper with an abbreviated

THE ICHNEUMONIDAE DESCRIBED BY C. G. THOMSON 5

locality name and sometimes a date. The locality labels may be handwritten or printed. Oc-
casionally the locality name is given in full. Some specimens have no locality name or abbrevi-
ation but instead have a small square of coloured paper. Unfortunately, the meaning of only one
colour is known: green indicates Ringsjon. There are two kinds of green squares, very small dark
ones and slightly larger (up to about 3x4 mm) paler, brighter ones. Interpretation of the small
dark labels as indicating Ringsjon is now generally accepted (for example, Huggert, 1973: 107;
Aubert, 1976a: 154) and I have discovered specimens bearing both a green square and a printed
label 'Ringsio'. Similar systems of coloured labels were used by other contemporary and earlier
workers, for example Zetterstedt (R. Danielsson, pers. comm.).

Specimens sometimes also have other labels, usually of one or more of the following four
kinds: a sex sign (printed); the name of a collector or collection; an additional locality label
giving a province or country; an identification.

The style of label with a locality abbreviation used by Thomson was also popular with other
contemporary collectors — most notably C. D. E. Roth who often accompanied Thomson on his
collecting trips. Thomson's and Roth's handwriting styles were similar and most labels are

Ibrekovl* B5stad« •Osb-a Karup

narcjret'el'orp

• Rossjoholm

Kungsmarken* •Reuen

-•Lund •frS3elsan3

Kallby^RSby •TornaHallestod'Ovedskl aster
•Atnarp •Dolby

yddingesj6n/^»Holmeja
rsjo* '-'•Bokeberg

Map 1 The Swedish province of Skane showing the type-localities of species of Ichneumonidae
described by C. G. Thomson.

6 M. G. FITTON

difficult to identify with certainty as the work of Thomson, although Roth's labels usually have a
date (day/month) below the locality abbreviation. The labels are poorly written and hard to
interpret until one is familiar with the forms of individual letters and the locality names from
which the abbreviations are derived.

A list of abbreviations used for Swedish localities is given below. It is not exhaustive and relates
mainly to ichneumonid type-material. The spelling used by Thomson is given first followed by
the modern equivalent where this differs. A [?] indicates that there is doubt about the form of the

abbreviation, its equivalence to the locality given or both. [Note. The Swedish letters a, a and 6
properly follow z in the Swedish alphabet but for the purposes of alphabetical order are treated as
a, a and o respectively in this list.] The localities in Skane are shown on Map 1.

Abbreviation Locality

Alnp Alnarp, Skane

Alp see 'Alnp'

Ar Arrie, Skane

Are Areskutan, Jamtland

Bast Bastad, Skane

Bgs Bogestad = Bokestad, Skane

Bkbg Bokeberg, Skane

Bl Blekinge

Boh Bohuslan

Bohl see 'Boh'

Bok see 'Bkbg'

Boks see 'Bgs'

Bor Borringe, Skane

Bs see 'Bgs'

Dby Dalby, Skane

Deg Degeberga, Skane

Dg see 'Deg'

Dgb see 'Deg'

Ekh Ekeshult, Skane

Esp Esperod = Asperod, Skane

Fg Fogelsang = Fagelsang, Skane

Fsg see 'Fg'

G Gottland = Gotland

Gbg Goteborg

Gotl see 'G'

Gott see 'G'

Hall Halland

Hbg Helsingborg, Skane

Hels see 'His'

Hg see 'Hbg'

Hhm [?] Hassleholm [?], Skane

Hkl Herrevadskloster, Skane

Him Holmia = Stockholm area

His Halsingland

Hma Holmeja, Skane

Hme see 'Hma'

Holm see 'Him'

lisp Ilstorp, Skane

Jtl Jemtland = Jamtland

Kalm Kalmar, Smaland

Kas Kaseberga, Skane

Kfge Kjeflinge = Kavlinge, Skane

Kgsm Kungsmarken, Skane

Kpe Kempinge = Kampinge, Skane

Krp Ostra Karup, 'Halland' [ = Skane]

La Lomma, Skane

Lap Lappland = Lapland [usually assumed to be Swedish Lapland]

Lapp see 'Lap'

THE ICHNEUMONIDAE DESCRIBED BY C. G. THOMSON 7

Ld Lund, Skane

Lhn Lindholmen, Skane

Loma see 'La'

Lop Loparod, Skane

Lpl see 'Lap'

Marg Margretetorp, 'Halland' [= Skane]

Mark Markaryd, Smaland

Mol Molle, Skane

Mrki Markiehage, Skane

Norl Norrland

Norr see 'Norl' [This abbreviation is easily confused with 'Norv' = Norvegica =

Norway.]

O Oland

Oel see 'O'

O.G. Ostergothland = Ostergotland

O Got see 'O.G.'

Oke Ofvedskloster = Ovedskloster, Skane

Ort Ortofta, Skane

Pal Palsio = Palsjo, Skane

Rab Raby, Skane

Raml Ramlosa, Skane

Rfn Reften, Skane

Rhm Ryssjoholm = Rossjoholm, Skane

Ron Ronnemolla, Skane

Rost Rostanga, Skane

Rota see 'Rost'

Rshm see 'Rhm'

Rsio Ringsion = Ringsjon, Skane

Rsjo see 'Rsio'

Sbg Sofdeborg = Sovdeborg, Skane

Scan Skane

Sk see 'Scan'

Skan Skanor, Skane

Skb Skabersjo, Skane

Sm Smaland

Smol see 'Sm'

Snor see 'Skan'

Ste Stehag, Skane

Steh see 'Ste'

Stkm Stockholm

Tbg Trelleborg, Skane

Tkov, Torekov, Skane

Tn [?] Torringelund [?], Skane

Tor Torringe, Skane

Trkv see Tkov'

Tve Tvedora = Torna Hallestad, Skane

V.G. Vestergothland = Vastergotland

V.W. Vestra Wram = Vastra Vram, Skane

Witt Wittsio = Vittsjo, Skane

Wml Vermland = Varmland

W.W. see 'V.W.'

Yd Yddinge, Skane

Ydd see 'Yd'

Ys Ystad, Skane

When labelling material Thomson apparently 'bracketed' together adjacent localities. Thus,
specimens of a species stated in the description to come from Palsjo may be labelled 'Hbg'
(= Helsingborg), Palsjo being a district of Helsingborg. (Other specimens are actually labelled
'Pal'.) The terms Norrland and Lappland were used rather imprecisely and sometimes inter-
changeably by Thomson. Norrland is the area of Sweden north of and including the provinces

8 M. G. FITTON

Harjedalen and Halsingland. Species stated to come from Lappland are often labelled Norrland
and vice versa. A list of the localities which it is assumed Thomson 'bracketed' is given below.

Helsingborg includes Palsjo

fKlinta
Rmgsjon includes < _ ,

(Stehag

... . fHolmeja

Yddinge includes < „

[Bokeberg

Norrland includes

Lappland

Jamtland

Halsingland

Specimens from particular collectors or collections were known by Thomson to come from a
particular locality. For instance, Ljungh specimens come mainly from Smaland and if Smaland is
given as a locality for a species the relevant specimens may actually be labelled 'Col. L-gh' but
without a locality label. Rudolphi collection specimens originate mainly from Halsingland and
Fallen specimens from Asperod (near Kivik).

Notes on the recognition of type-material and on the
selection and designation of lectotypes

Thomson did not have a type concept in any modern nomenclatural sense. He made no attempt
to preserve or label in any particular way the specimens which were the bases of his descriptions
of new species.

Usually Thomson gives no direct details of specimens with original descriptions, only the
localities or more general areas where the species had been found, such as 'Funnen vid Holmeja i
narheten af Yddingesjon' and 'Funnen vid Degeberga i Skane'. For several species much less
precise locality information is given, for example, 'Ej sallsynt i norra och medlersta Europa', or
sometimes none at all. Localities outside Skane are often only given at the level of province
('Oland' or 'Norrland', for example) or country. In the latter case Thomson may not have been
able to decipher abbreviations, read handwritten labels or ascertain the correct geographical
positions of the localities of material obtained from foreign workers. Information in addition to
the locality, when any is given, includes habitat, date, collector, an indication of abundance and
host data. Examples, 'Funnen vid Kjeflinge i barrplanteringen', 'Funnen i September vid Ortofta
nara Lund', 'Funnen talrikt vid Ilstorp i Skane af Conservator C. D. E. Roth', 'Sallsynt; funnen
pa sandmarker pa Oland' and 'I Munchen utklackt ur Thecla Betulae af D : r Kriechbaumer'.
More precise information about the specimens themselves is given only rarely, and often when
there was only one, for example, 'Exemplaret, en hona, ar funnet pa Gottland' and 'Endast ett ex.
fran sodra Frankrike (Coll. Lethierry)'.

Thus Thomson's lack of attention to original material; the inadequate published information;
the poor labelling of specimens; and changes in the arrangement of the collection subsequent to
publication all hinder recognition of type-material. Indeed, for most species it is impossible to be
absolutely certain which specimens are types. On the other hand this combination of poor
information gives wide scope in deciding which specimens are possible types. It also leaves open
the possibility that a specimen chosen as a primary type may be shown, at some later date and in
the light of further evidence, not to have been used as the basis of an original description.
However, practical considerations justify the selection of single primary type-specimens (usually
lectotypes) to serve as stable bases for the nomenclature of the species.

Of the specimens standing under the name of a Thomson species in his collection I have
recognised as syntypes all those which are in agreement with the description and with the other
information (on localities, etc.) given in the original publication. This latter qualification would
perhaps be better expressed as lack of disagreement because, for example, where Thomson cites a

THE ICHNEUMONIDAE DESCRIBED BY C. G. THOMSON 9

locality within a province and there are specimens labelled only with the name of the province
they have been accepted as syntypes. Thomson's subsequent references to his own species are
sometimes helpful in deciding on the limits of a syntype series. Syntypes of some species are in
other collections. For a few species specimens under other names have been regarded as syntypes,
usually on the basis of precise agreement with descriptions and other information, and evidence
of changes in that part of the collection subsequent to the relevant publication.

It is not always easy, using the above criteria, to decide whether or not a particular specimen
should be included in a syntype series. Problems are caused by minor disagreements with
descriptions; illegibility of labels; interpretation of locality abbreviations on labels and of col-
oured tags; and differences between localities as published and as given on labels. Agreement of
specimens with descriptions calls for a subjective judgement, sometimes open to alternative
opinions. The questions relating to labelling and localities are dealt with in the previous section
on labelling of specimens. Each case has to be judged on its individual merits, paying attention to
the utility of the particular situation.

Thomson described many new species from single specimens (holotypes). In some cases there
can be no doubt, for example, 'Ett exemplar fran Smaland'; in others it must be inferred from the
published information together with the fact that there is only one specimen in the collection.
Some workers object to recognition of specimens of the latter kind as holotypes. In cases which I
consider doubtful I have cited such specimens as single surviving syntypes.

Any member of a syntype series is eligible for designation as lectotype of the nominal species
concerned. Lectotypes have already been designated (published) for a lot of Thomson's species.
Many others have been selected (labelled) but not yet published. Lectotypes already selected but
not published are designated in the present work, but where no lectotype has been selected details
of the syntype series are given. Only a few lectotypes have been selected and designated by me,
usually in cases where previous designations are invalid for one reason or another. Lectotype
designations are best made in the context of a complete revision of the group to which the species
concerned belongs. However, for reasons of practicality it was thought desirable to publish here
the selections already made by other workers (some as long ago as 1954).

A number of the lectotype designations already published call for comment. Several are casual
in the extreme and need careful consideration of their validity in relation to the relevant provis-
ions of the International Code of Zoological Nomenclature. Others, such as Aubert's lists (1966;
1968; 1972) of indiscriminately chosen species, have been published as ends in themselves. Much
more attention must be paid to the publication of lectotype designations. There are too many
references such as 'lecto. des Townes, 1958', not meaning that there is a designation published by
Townes in 1958 but that the specimen was labelled by Townes in that year. It often happens that
such a reference to the label is the first publication of a lectotype for that species. Are such
references valid designations? For practical reasons they are best accepted as such but the
situation is far from satisfactory. Editors of journals, as well as authors, must take some of the
blame for this state of affairs.

Thomson's use of names for subgeneric categories

Thomson's use of names for 'subgenera' was inconsistent and is difficult to interpret. This incon-
sistency is not surprising in work published over a long period but, as a consequence, determining
the original subgeneric (and sometimes generic) placements of species is not always easy.

Even when Thomson used subgenera clearly he often prefixed the name of each species with
the initial letter of the subgeneric and not the generic name, for example, in Mesochorus (1886a:
327-344). Workers who have not studied Thomson's work properly have been misled by this
practice into citing incorrectly original generic placements.

Of more importance is Thomson's habit of giving genus-group names in parentheses at various
points in his keys to species. The names used in this way are mainly those originating from
Foerster's work (see Perkins, 1962: 387). In some cases the rank of such names as subgenera is
clear because Thomson gives a 'Conspectus subgenerum' at the beginning of the genus, some-
times including synonymy, for example, Megastylus (18886: 1310). In other cases Thomson does

10 M. G. FITTON

not give such an indication or the names are given so as to apply to groups of species within
subgenera, for example, the use of Myriarthrus within Megastylus subgenus Megastylus
(1888b: 1314). Thus, it is not always clear whether the names apply to formal subgenera; formal
infra-subgeneric categories ; informal groups of species ; or are included for purposes of synony-
my. They are important because they are often the first association of species with Foerster
generic names (Perkins, 1962). Subsequently the names have been cited most frequently as
subgenera. Except for those used by Thomson infrasubgenerically they are treated uniformly as
subgenera in the present work, but because of the doubt about many of them the catalogue
section is arranged in alphabetical order of binominal, and not trinominal, combinations.

Format and arrangement of catalogue

The catalogue section is arranged in alphabetical order of the original binominal combination
(that is, disregarding any subgeneric component of the name). The nomenclatural summary
which follows the catalogue and the index provide entry into the catalogue via species names,
subgeneric placements and current combinations.
For each nominal species the entry is arranged in the following sequence :

Name; date and page reference of original publication; status and sex of primary type(s); type-
locality; type-depository; lectotype designation or reference to previous valid type-restriction
(when necessary).

Details of the labels on the specimen(s).

Notes.

A statement, prefixed 'Identity', on the generic placement and synonymy of the species.

The following points should be noted with regard to these data.

The name is given as published except that the orthography is altered to comply with Articles
26, 27, 28 and 32 of the Code when necessary (in which cases the form of the name as published is
also given).

Swedish type-localities are given as cited by Thomson with the addition of the names of the
country and province (when necessary) and the modern spelling (when different).

Names of type-depositories are abbreviated as in the list below. Where types are lost this is
stated in place of a depository. The collections from which Thomson described species are given
after each depository.

CM, Norwich Castle Museum, Norwich, England (Bridgman collection)
NM, Goteborg Naturhistoriska Museet, Goteborg, Sweden (G. F. Moller collection)
NR, Stockholm Naturhistoriska Riksmuseet, Stockholm, Sweden (Holmgren collection)
UZI, Lund Universitetets Zoologiska Institutionen, Lund, Sweden (Thomson collection)

ZM, Copenhagen Zoologisk Museum, Copenhagen, Denmark (Drewsen, Jensen and Wustnei

collections)
ZSBS, Munich Zoologische Sammlung des Bayerischen Staates, Munich, West Germany

(Kriechbaumer and Foerster collections)

For specimens from Thomson's own collection details are not given of labels added since
Thomson's death (except for the cabinet labels which were transferred to the first specimen in
each series by Bengtsson). For each label an indication is given (in square brackets) of whether it
is handwritten or printed. Except for a few species with large syntype series, all primary type-
specimens have individual modern labels showing their identity and status.

Catalogue

Acanthocryptus nigriceps, 1883: 868. Syntype 1 cJ, SWEDEN: Smaland, Calmar [= Kalmar], Hossmo (UZI,
Lund).

Labels. Hossmo 4/6 70 [hand] ; Kalmar [hand] ; nigriceps [Thomson cabinet label].

Gravenhorst's specimen (1829ft: 676. Phygadeuon quadrispinus Var. 1. £) [not examined] is also a
syntype of this species. Thomson mis-spelled the name of the Gravenhorst species as quadrispinosus
instead of quadrispinus.

Identity. ? Rhembobius nigriceps (Thomson).

THE ICHNEUMONIDAE DESCRIBED BY C. G. THOMSON 1 1

Acanthocryptus nigricollis, 1883: 868. Lectotype $, SWEDEN: Skane, Bastad (UZI, Lund), by designation of
Aubert, 1966: 129.

Labels. Bast [hand] ; nigricollis [Thomson cabinet label].
Identity. ? Rhembobius nigricollis (Thomson).

Adelognathus (Adelognathus) aciculatus, 1883: 879. Type(s) 9, SWEDEN : Skane, Stehag (lost).
Identity. Adelognathus aciculatus Thomson.

Adelognathus (Adelognathus) dlmidiatus, 18886: 1276. Lectotype 9, FRANCE: Raismes (UZI, Lund), by
designation of Aubert, 1972: 147.
Labels. Raismes. [hand] ; dimidiatus [hand].
Identity. Adelognathus dimidiatus Thomson.

Adelognathus (Adelognathus) facialis, 1883: 880. Holotype?, SWEDEN : Norrland (UZI, Lund).
Labels. Norl. [printed] ; facialis [Thomson cabinet label].
Identity. Adelognathus facialis Thomson.

Adelognathus (Adelognathus) fasciatus, 1883: 878. Holotype?, SWEDEN : Skane, Sofdeborg [= Sovdeborg]
(UZI, Lund).

Label. Sbg 23/7 [hand].

Identity. Adelognathus fasciatus Thomson.

Adelognathus (Adelognathus) laevicollis, 1883: 878. Syntypes 4 ?, 1 <$, SWEDEN: Skane, Ringsion [ =
Ringsjon] (UZI, Lund).

Labels. Rsio [printed] (2$). Scan [printed] (1$). [small green square]; laevicollis [Thomson cabinet
label] (1 9)- [small green square]; <$ [printed] (1 $).
Identity. Adelognathus laevicollis Thomson.

Adelognathus (Adelognathus) limbatus, 1888ft: 1275. LECTOTYPE 9, SWEDEN: Skane, Palsjo (UZI, Lund),
here designated (selected by J. F. Aubert).
Label. Pal [hand].
Identity. Junior synonym of Adelognathus brevicornis Holmgren (Perkins, 1943ft: 95, 104).

Adelognathus (Adelognathus) nigriceps, 1888ft: 1274. Type(s)$, FRANCE (lost).

From the description it seems likely that there was only one specimen, which may have been returned
to Lethierry. There are no specimens under this name in the Thomson collection.
Identity. Adelognathus nigriceps Thomson (Perkins, 1943ft: 99).

Adelognathus (Adelognathus) nigricornis, 1888ft: 1276. Type(s) 9, FRANCE (lost).
The comments on A. nigriceps apply to this species also.
Identity. Adelognathus nigricornis Thomson (Perkins, 1943ft: 100).

Adelognathus (Cnemischus) pilosus, 1888ft: 1277. Holotype9, SWEDEN : Skane, Alnarp (UZI, Lund).
Labels. Alnarp [printed] ; pilosus [Thomson cabinet label].
Identity. Adelognathus pilosus Thomson.

Adelognathus (Adelognathus) puncticoltis, 1883: 877. Holotype9, SWEDEN : Smaland (UZI, Lund).
Labels. Smol [printed] ; puncticollis [Thomson cabinet label].
Identity. Adelognathus puncticollis Thomson.

Adelognathus (Adelognathus) punctiventrls, 1883: 877. Lectotype 9, SWEDEN: Skane, Torekov (UZI, Lund),
by designation of Jussila, 1965: 31.
Label. Tkov 23/7 [hand] [not Tkro' as stated by Jussila].
Identity. Adelognathus punctiventris Thomson.

Adelognathus (Adelognathus) punctulatus, 1883: 879. Syntype 1 9, SWEDEN: Skane, Ringsion [= Ringsjon]
(UZI, Lund).

Label. Rsio [printed].

Identity. Adelognathus punctulatus Thomson.

b,

Adelognathus (Adelognathus) scabriculus, 1883: 877. Lectotype 9, SWEDEN: Lappland (UZI, Lund), by
designation of Jussila, 1965: 30.
Label. Lpl. [printed].
Identity. Junior synonym of Adelognathus tetracinctorius (Thunberg) (Jussila, 1965 : 30).

12 M. G. FITTON

Aethecerus graniger, 1891 : 1641. Syntype 1 9, SWEDEN: Skane, Ringsjon (UZI, Lund).
Label, [small green square].
The head is missing from the syntype 9-
Identity. Aethecerus graniger Thomson.

Aethecerus palttcoxa, 1891 : 1640. Syntypes 4 9, 8 & SWEDEN: Skane (UZI, Lund).

Labels. Hbg [hand] ; pallicoxa [Thomson cabinet label] (1 9, 2 $ all on one pin). Pal [hand] (3 9, 6<J).
The pin bearing three specimens also bears Aubert's lectotype label but without any indication of the
specimen to which it applies.

Identity. Aethecerus pallicoxa Thomson.

Allomacrus pimplarius, 18886: 1282. LECTOTYPE ?, SWEDEN: Skane, Esperod [= Asperod] (UZI, Lund),
here designated (selected by H. K. Townes).
Label. Esp [printed].
Identity. Allomacrus pimplarius Thomson.

Amblyteles (Amblyteles) anurus, 1888a: 114. Syntypes 9 d, SWEDEN (lost).
There are no specimens under this name in the Thomson collection.
Identity. Unknown, the name remains a nomen dubium.

Amblyteles (Ctenichneumon) circulator, 1894: 2085. Syntypes 1 $, 1 <$, SWEDEN: ut'd Goteborg (9) and pa
Gottland (rf) (UZI, Lund).

Labels. Boh. Skarg [printed] [ = Bohuslan skargard, islands off the coast at Goteborg] ; Circulator m
[Thomson cabinet label] (9). G [hand] (<$).

Identity. Ctenichneumon circulator (Thomson).

Amblyteles (Amblyteles) limnophilus, 1888a: 1 19. Syntypes 3 9, 4 <$, SWEDEN (UZI, Lund).

Labels. L-d [printed] (1 9)- Yddinge [printed]; limnophilus [hand]; Ammonius [Thomson cabinet
label] (1 9). Ringsio [printed] (1 <J). Col. Hgn. [printed] [= Col. Holmgren] (1 £). [No labels] (1 9, 2 rf).

With the original description Thomson merely cited the locality as 'Suecia'. Later (1894: 2089) he
commented 'Sallsynt pa fuktiga stallen'. In view of this, it is almost certain that not all of the 38 specimens
standing in the collection are syntypes. The few specimens which best agree with the description have been
labelled as syntypes.

Identity. Spilichneumon limnophilus (Thomson) comb. n.

Amblyteles (Amblyteles) longigena, 1888a: 112. Lectotype 9, SWEDEN: Skane, Palsjo (UZI, Lund), by
designation of Townes, Momoi & Townes, 1965: 491.
Label. Palsio [printed].
Identity. Diphyus longigena (Thomson).

Amblyteles (Platylabus) opaculus, 1888a: 124. Syntypes 3 9, 2 <J, SWEDEN: Skane, Palsjo and Alnarp (UZI,
Lund).

Labels. Pal [hand] (3 9, 1 rf). Alp. [hand] (1 <J).

Although Thomson only gave 'Suecia' as the locality in the original description, he later (1894: 2108)
gave more precise information : 'vid Palsjo och Alnarp i Skane'. Three further specimens are from other
localities and were, therefore, probably added to the collection after 1894 and unlikely to be syntypes.

Identity. Platylabus opaculus (Thomson).

Amblyteles (Anisobas) parviceps, 1888a: 122. Syntype 1 9, SWEDEN : Skane, Lund (UZI, Lund).

Labels. Lund [printed] ; parviceps [hand] ; sublae vis [Thomson cabinet label].

The syntype stands in the collection under '? hostilis'. Thomson may have had only one original
specimen (holotype). In his later treatment of Anisobas (1894: 2099) he does not mention parviceps but
gives characters of both sexes under hostilis. The label 'Lund', present when I first examined the specimen
in 1978, was missing when I re-examined it in 1980.

Identity. Anisobas parviceps (Thomson).

Amblyteles (Anisobas) platy stylus, 1888a: 122. Lectotype^, SWEDEN : Skane, Rossjoholm (UZI, Lund), by
designation of Aubert, 1966: 128.

Label. Rshm 16/6 [hand].

Thomson (1894: 2099) referred to the species as Anisobas platylabus (incorrect subsequent spelling), the
name also used by Aubert when designating the lectotype. The locality label was wrongly interpreted by
Aubert as Raby.

Identity. Anisobas platystylus (Thomson).

THE ICHNEUMONIDAE DESCRIBED BY C. G. THOMSON 13

Amblyteles (Platylabus) punctifrons, 1888a: 124. Syntypes 2 $, ? syntypes 2 ?, SWEDEN (UZI, Lund).

Labels. Scan [printed] (syntype). Sk. [hand]; punctifrons m [hand] (syntype). 12 [ + two words (illeg-
ible)] (? syntype). I. antennator ? [hand] (? syntype).

Thomson only gave 'Suecia' as the locality in the original description. Later (1894: 2108) he stated
'Funnen i Skane vid Esperod.' The labels of the specimens labelled as syntypes or possible syntypes do not
bear any data which conflict with this locality information. Further specimens are from Ringsjon and,
therefore, probably post-date the 1894 publication and unlikely to be syntypes. The specimen with
Thomson's label 'punctifrons m' has the apex of the gaster missing.

Identity. Platylabus punctifrons (Thomson).

Amblyteles (Amblyteles) simplicidens, 1888a: 120. Holotype ?, SWEDEN : Skane, Stehag (UZI, Lund).

Labels. Ste CM [hand] [CM = Carl Moller]; simplicidens [hand]; simplicidens [Thomson cabinet
label].

Thomson's original description did not include any characters of the male and the single female
specimen is presumed to be a holotype. Thomson later (1894: 2090) gave characters of both sexes and the
locality Ringsjon. It is assumed that the three male specimens in the collection post-date the original
description. One of these males is of particular interest because its pin bears both a green square (of the
paler, brighter kind) and a printed label 'Ringsio', further evidence that the green squares mean that the
specimens with them were collected in the vicinity of Ringsjon.

Identity. Spilichneumon simplicidens (Thomson).

Amblyteles (Amblyteles) stagnicola, 1888a: 119. Lectotype $, SWEDEN: Smaland (UZI, Lund), by des-
ignation of Townes, Momoi & Townes, 1965: 505.

Labels. Smoland [printed] ; stagnicola [hand] ; laeviventris [Thomson cabinet label].
Thomson gave only 'Suecia australis' as the locality in the original description but later (1894: 2089-
2090) he stated 'Funnen vid Bastad och Ringsjon i Skane'. Although the locality of the lectotype is not in
agreement with this later information there is nothing positive to invalidate its selection as lectotype.
However, in cases of this kind it is prudent to recognise as syntypes (and select lectotypes from) material
from the localities mentioned by Thomson. Material from other localities probably post-dates the later
publication and is therefore unlikely to include syntypes.
Identity. Spilichneumon stagnicola (Thomson).

Amblyteles (Spilichneumon) triplicates, 1894: 2088. Syntypes 2 $, 2 <$, SWEDEN: Stockholm and Halland
(UZI, Lund).

Labels. 106. [hand]; Stkm. [hand]; Amb. 7-guttatus [hand]; 3-plicatus m [Thomson cabinet label]
(1 ?). Holm [printed] [ = Holmia] ; Hallstr. [hand] (1 ?). Halland [hand] ; Rui [hand] (2 <J).

Identity. Diphyus triplicatus (Thomson) comb. n.

Amblyteles (Amblyteles) truncicola, 1888a: 115. LECTOTYPE ?, SWEDEN : Skane, Lund (UZI, Lund), here
designated (selected by J. F. Aubert).

Labels. Lund [printed]; $ [printed] ; truncicola [Thomson cabinet label].

Thomson gave only Suecia as the locality in the original description but later (1894: 2092) stated
'funnen vid Lund'.

Identity. Spilothyrateles truncicola (Thomson) comb. n.

Angitia annulicrus, 1887c: 1155. Lectotype $, SWEDEN: Skane, Palsjo (UZI, Lund), by designation of Aubert,
1966:130.

Labels. Pal [hand]; $ [printed] ; annulicrus [Thomson cabinet label].
Identity. Diadegma annulicrus (Thomson).

Angitia anura, 1887c: 1164. Syntypes 1 $, SWEDEN: Gottland [= Gotland] (UZI, Lund); 1 $, ? GERMANY
(ZSBS, Munich).

Labels. G [hand] ; anura [Thomson cabinet label] (Lund specimen). 56. 2. [hand] ; 40. anura Thms.
[hand] (Munich specimen).

Aubert (1972: 150) was incorrect in referring to the Lund specimen as 'holotype'. Thomson gives clear
evidence of a syntype series. The Munich specimen (from the Kriechbaumer collection) is accompanied by
two cocoons (each labelled '56. 2.' and on separate pins).

Identity. Diadegma anura (Thomson).

Angitia brevivalvis, 1887c: 1163. Lectotype $, SWEDEN: Skane, Ringsion [= Ringsjon] (UZI, Lund), by
designation of Horstmann, 1969: 436.
Labels. Rsio [printed] ; brevivalvis [Thomson cabinet label].
Identity. Diadegma brevivalvis (Thomson).

14 M. G. FITTON

Angitia claripennis, 1887c: 1161. Lectotype <$, SWEDEN: Skane, Bastad (UZI, Lund), by designation of
Aubert, 1966: 130.
Label. Bast [hand].
Identity. Junior synonym of Diadegma majalis (Gravenhorst) (Horstmann, 1969: 461).

Angitia crassiseta, 1887c: 1162. Type(s)9, SWEDEN : Skane, Lund (lost).

NEOTYPE 9, SWEDEN: Skane, Ringsjon (UZI, Lund), here designated (selected by J. F. Aubert, 1966:
130).

Aubert 's original publication of the neotype (1966: 130) was not valid because it did not comply with
the provisions of Article 75(c) of the Code. Horstmann (1969: 457) cited the neotype and gave characters
to differentiate the species, but otherwise did not satisfy the conditions of Article 75(c).

Thomson's original material of this species cannot be found either in his main collection or among the
duplicate material. It is therefore believed to be lost or destroyed. The neotype stands in Thomson's
collection under the name Angitia crassiseta and is in agreement with the original description.

Labels. Rsio [printed] ; crassiseta [Thomson cabinet label].

Identity. Diadegma crassiseta (Thomson).

Angitia crataegellae, 1887c: 1164. Lectotype £, GERMANY (WEST): Bavaria (UZI, Lund), by designation of
Townes, Momoi & Townes, 1965: 293.

Labels. 5650 [yellow, hand] ; Germ, [hand] ; Crataegellae [Thomson cabinet label].

Identity. Probable junior synonym of Enytus apostatus (Gravenhorst) (Horstmann, 1969: 441). Horst-
mann (1969) regards Dioctes (= Enytus) as a junior synonym of Diadegma and not as a distinct genus
(Townes, 19706: 177).

Angitia elongata, 1887c: 1155. Lectotype $, SWEDEN: Skane, Palsjo (UZI, Lund), by designation of Horst-
mann, 1969:429.

Labels. Pal [hand] ; elongata [Thomson cabinet label].
Identity. Diadegma elongata (Thomson).

Anffitia holopyga, 1887c: 1160. LECTOTYPE 9, SWEDEN: Skane, Ortofta (UZI, Lund), here designated
(selected by Aubert and Horstmann).

Aubert's published lectotype designation (1966: 130) and Horstmann's citation of the lectotype
(1969: 436) are invalid because there are two females on the pin bearing Aubert's label (dated 1963 and
not indicating which specimen is lectotype). Aubert (1966) stated 'No. 1' ['indique le numero d'ordre de
ITnsecte dans la serie originate'] but it is impossible to decide whether the upper or lower specimen is the
first in the series. The pin also bears Horstmann's label 'Lektotypus das untere9 Horstm. del. 1970' which
postdates his publication on this species. The lower of the two specimens is here designated lectotype.

Labels.Qrtofta. [printed] ; holopyga [Thomson cabinet label].

Identity. Diadegma holopyga (Thomson).

Angitia lacticrus, 1887c: 1163. Lectotype 9, SWEDEN: Skane, Ringsion [= Ringsjon] (UZI, Lund), by
designation of Horstmann, 1969: 443.

Labels, [small green square] ; lacticrus [Thomson cabinet label].
Identity. Diadegma lacticrus (Thomson).

Angitia latungula, 1887c: 1 165. Lectotype 9, FRANCE (UZI, Lund), by designation of Horstmann, 1969: 442.
Label. Gall [hand] [ = Gallia].
Identity. Diadegma latungula (Thomson).

Angitia macrostoma, 1887c: 1166. Holotype9, SWEDEN: Skane, Ortofta (UZI, Lund).
Labels. Ort [hand] ; macrostoma [Thomson cabinet label].
Identity. Lathrostizus macrostoma (Thomson).

Angitia melania, 1887c: 1 160. Lectotype 9, SWEDEN: Skane, Yddinge (UZI, Lund), by designation of Horst-
mann, 1969: 438.

Labels. Yd. [hand] ; melania [Thomson cabinet label].
Identity. Diadegma melania (Thomson).

Angitia micrura, 1887c: 1164. Syntypes 2 9, 2 ^, SWEDEN : Skane, Palsjo and FRANCE: Emmerin and Scarpe
(UZI, Lund).

Labels. Pal. [hand]; Var [printed] (1 9). Emmerin. [hand]; 9 [printed] (1 9). Emmerin. [hand] (1^).
Scarpe [hand] ; micrura [Thomson cabinet label] (1 $).

The male specimen published by Aubert (1966: 131) as lectotype is from Ortofta [label: 'Ort'] not
Palsjo as stated by Aubert. It has no type status.

Identity. Diadegma micrura (Thomson).

THE ICHNEUMONIDAE DESCRIBED BY C. G. THOMSON 15

Angitia monilicornis, 1887c: 1167. Lectotype 9, SWEDEN: Lappland (UZI, Lund), by designation of Horst-
mann, 1969: 446.

Labels. Lpl. [printed] ; crassinervis [hand] ; monilicornis [Thomson cabinet label].
Identity. Lathrostizus monilicornis (Thomson).

Angitia monospila, 1 887c : 1157. Lectotype 9, SWEDEN : Skane, Bastad (UZI, Lund), by designation of Aubert,
1966: 131.

Labels. Bast [hand] ; monospila [Thomson cabinet label].

Although Townes, Momoi & Townes (1965: 297) were almost certainly correct in recognising this as
the type ( = holotype, the single original specimen) Aubert chose to designate it as lectotype. He presented
no evidence of a syntype series.

Identity. Diadegma monospila (Thomson).

Angitia parvicanda, 1887c: 1163. Lectotype 9, SWEDEN: Skane, Palsjo (UZI, Lund), by designation of
Horstmann, 1969: 437.

Labels. Pal. [hand] ; Var [printed] ; parvicauda [Thomson cabinet label].

Authors since Thomson (for example, Dalla Torre, 1901 : 130; Horstmann, 1969: 437) have chosen to
alter the spelling of the name to parvicauda and there is evidence (Thomson's own cabinet label) that this
is what was intended. However, a strict interpretation of Article 32(aXii) of the Code (as amended, Bull.
zoo/. Nom. 31 (1974): 83) suggests that the original spelling should be retained.

Identity. Diadegma parvicanda (Thomson).

Angitia polyzona, 1887c: 1159. Lectotype 9, FRANCE: Marseille (UZI, Lund), by designation of Aubert,
1966: 131.

Labels. Marseille [hand] ; polyzona [Thomson cabinet label].
Identity. Junior synonym of Diadegma maculata (Gravenhorst) (Horstmann, 1969: 454).

Angitia punctipes, 1887c: 1166. Lectotype $, SWEDEN : Skane, Palsjo (UZI, Lund), by designation of Townes,
Momoi & Townes, 1965: 303.

Labels. Pal. [hand] ; punctipes [Thomson cabinet label].
Identity. Lathrostizus punctipes (Thomson).

Angitia rimator, 1 887c : 1 1 56. Syntype 1 9, SWEDEN : Skane, Torekov (UZI, Lund).
Label. Sk [hand].

The specimen published by Aubert (1966: 131) as lectotype, and cited by Horstmann (1969: 459), is
from Lomma [label : 'Loma 20/7'] not Torekov as stated by Aubert. It has no type status.
Identity. Diadegma rimator (Thomson).

Angitia sordipes, 1887c: 1156. Lectotype 9, SWEDEN: Skane, Palsjo (UZI, Lund), by designation of Aubert,
1966: 131.

Labels. Pal. [hand] ; sordipes [Thomson cabinet label].
Identity. Diadegma sordipes (Thomson).

Angitia specularis, 1887c: 1162. Lectotype 9, SWEDEN: Skane, Palsjo (UZI, Lund), by designation of Aubert,
1966:131.

Label. Pal [hand].

Although Townes, Momoi & Townes (1965: 298) were almost certainly correct in recognising this as
the type ( = holotype, the single original specimen) Aubert chose to designate it as lectotype. He presented
no evidence of a syntype series.

Identity. Diadegma specularis (Thomson).

Angitia subbuccata, 1887c: 1156. Lectotype 9, SWEDEN : Skane, Lund (UZI, Lund), by designation of Aubert,
1966:131.

Label. L-d [printed].

The lectotype is the upper of three specimens on one pin. The middle specimen has the gaster missing
and the lower one is a male.

Identity. Junior synonym of Diadegma truncata (Thomson) (Horstmann, 1969: 459).

Angitia tenuipes, 1887c: 1158. Lectotype 9, SWEDEN: Skane, Lund (UZI, Lund), by designation of Aubert,
1966: 131.

Labels. Ld [hand]; tenuipes [Thomson cabinet label].
Aubert (1966: 131) cited the locality, incorrectly, as Ortofta.
Identity. Diadegma tenuipes (Thomson).

16 M. G. FITTON

Angitia trochanterata, 1887c: 1157. Lectotype 9, SWEDEN: Skane, Degeberga (UZI, Lund), by designation of
Aubert, 1966: 131.
Label. Dgb. [hand].
Identity. Diadegma trochanterata (Thomson).

Angitia truncata, 1887c: 1155. Lectotype 9, SWEDEN: Skane, Palsjo (UZI, Lund), by designation of Aubert,
1966: 131.

Labels. Pal [hand] ; truncata [Thomson cabinet label].
Identity. Diadegma truncata (Thomson).

Anilasta albicrus, 1887c: 1171. Lectotype $, GERMANY (WEST): Munich (UZI, Lund), by designation of
Aubert, 1972: 150.

Labels. 63.3. [hand] ; Germ [hand] ; albicrus [Thomson cabinet label].
Identity. Junior synonym of Hyposoter vividus (Holmgren) (Aubert, 1972: 150, 151).

Anilasta boops, 1887c: 1173. Lectotype 9, FRANCE: near Lille, Emmerin (UZI, Lund), by designation of
Aubert, 1972: 150.

Labels. Emmerin [hand]; 9 [printed].
Identity. Hyposoter boops (Thomson).

Anilasta coxator, 1887c: 1 173. Syntypes 2 9, SWEDEN : Skane, Ringsion [= Ringsjon] and GERMANY (WEST):
Munich (UZI, Lund).

Labels, [small green square]; 3 [hand] ; Coxator [Thomson cabinet label] (1 9)- 75.528. [hand]; Germ.
[hand](l 9).

Identity. Hyposoter coxator (Thomson) comb. n.

Anilasta faciaUs, 1887c: 1 174. Lectotype 9, FRANCE (UZI, Lund), by designation of Aubert, 1972: 150.
Label. Gallia [hand].
Identity. Hyposoter facialis (Thomson).

Anilasta leucomera, 1887c: 1172. Lectotype 9, SWEDEN: Skane, Yddinge (UZI, Lund), by designation of
Aubert, 1966: 131.
Label. Yddinge [printed].
Identity. Hyposoter leucomerus (Thomson).

Anilasta longula, 1887c: 1171. LECTOTYPE 9, SWEDEN : Skane, Ryssjoholm [= Rossjoholm] (UZI, Lund),
here designated (selected by R. Hinz).

Labels. Rshm 16/6 [hand] ; longula [hand] ; longula [Thomson cabinet label].
Identity. Hyposoter longulus (Thomson) comb. n.

Anilasta pectinata, 1887c: 1171. Lectotype 9, SWEDEN: Skane, Esperod [= Asperod] (UZI, Lund), by
designation of Aubert, 1972: 151.
Label. Scan [hand].
Identity. Hyposoter pectinatus (Thomson).

Anilasta picticollis, 1887c: 1174. ?Holotype9, ?lTALY(UZI, Lund).

Labels. +125 [hand]; Dalm. [printed] [= Dalmatia]; picticollis [Thomson cabinet label].
Thomson gave the locality for his single specimen as 'norra Italien'. Because the specimen is labelled
'Dalm.' there must be doubt about its status or the type-locality or both.
Identity. Hyposoter picticollis (Thomson).

Anilasta quadrinotata [as 4-notatd], 1887c: 1 174. Lectotype 9, FRANCE: Ostricourt (UZI, Lund), by designa-
tion of Aubert, 1972: 151.
Label. Ostricourt. [hand].
Ostricourt is near Phalempin (south of Lille).
Identity. Echthronomas quadrinotata (Thomson).

Anilasta ruficrus, 1887c: 1172. Lectotype 9, SWEDEN: Skane, Helsingborg (UZI, Lund), by designation of
Aubert, 1966: 131.

Labels. Hbg. [hand] ; ruficrus [Thomson cabinet label].
Identity. Hyposoter ruficrus (Thomson).

Anilasta tenuicosta, 1887c: 1170. Lectotype $, SWEDEN: Norrland (UZI, Lund), by designation of Aubert,
1972: 151.

Labels. Norl. [printed] ; tenuicosta [Thomson cabinet label].
Identity. Hyposoter tenuicosta (Thomson).

THE ICHNEUMONIDAE DESCRIBED BY C. G. THOMSON 17

Anilasta varicoxa, 1887c: 1173. Lectotype 9, SWEDEN: Skane, Palsjo (UZI, Lund), by designation of Aubert,
1972: 151.

Labels. Pal [hand] ; varicoxa [Thomson cabinet label].
Identity. Junior synonym ofHyposoter neglectus (Holmgren) (Aubert, 1972: 151).

Anomalon (Anomalon) annulitarse, 1892a: 1764. Syn types 9 (J, SWEDEN and GERMANY (lost).

The syntype series of this species comprises the specimens in Thomson's own collection (from 'med-
lersta Sverige') together with the specimens (from Germany and Sweden) which were the bases of the
descriptions of Gravenhorst and Holmgren to which Thomson refers. All of the syntypes are lost.

The seven specimens in Thomson's collection are from Germany (label 'Germ.') and France (label
'Gall.') and therefore cannot be his syntypes. One of them has been labelled incorrectly as 'lectotype'. The
part of the Gravenhorst collection including Anomalon fibulator is lost (Townes, 1959: 77). The collections
of the Naturhistoriska Riksmuseum, Stockholm include only two specimens referable to Anomalon fib-
ulator sensu Holmgren. These do not correspond to Holmgren's 'Var. 1' and are therefore not syntypes of
Thomson's species.

Identity. Junior synonym of Gravenhorstia (Erigorgus) fibulator (Gravenhorst) (det. I. D. Gauld).

Anomalon (Anomalon) claripennis, 1892a: 1764. Lectotype <J, SWEDEN: Ostergotland (UZI, Lund), by
designation of Aubert, 1966: 130.

Labels. O Got [printed] ; Col. Hgn. [printed] ; claripenne m [Thomson cabinet label].
Aubert (1966: 130) cited the locality, incorrectly, as Gotland.
Identity. Gravenhorstia (Erigorgus} claripennis (Thomson) (det. I. D. Gauld).

Anomalon (Barylypa) genalls, 1892a: 1767. LECTOTYPE $, SWEDEN: Skane, Ringsjon (UZI, Lund), here
designated (selected by H. K. Townes).

Labels, [green square] ; genalis [hand] ; genalis m. [Thomson cabinet label].
Identity. Junior synonym of Barylypa delictor (Thunberg) (det. I. D. Gauld).

Anomalon (Anomalon) lapponicum, 1892a: 1763. Holotype 9, SWEDEN: Lappland (UZI, Lund).
Labels. Lap [hand] ; Lapponicum m. [Thomson cabinet label].
Identity. Junior synonym of Gravenhorstia (Erigorgus) cerinops (Gravenhorst) (det. I. D. Gauld).

Anomalon (Barylypa) laticeps, 1892a: 1766. Lectotype <$, SWEDEN: Oland (UZI, Lund), by designation of
Viktorov & Athanasov, 1974: 376.
Labels. 0 [hand] ; laticeps n. [Thomson cabinet label].
Identity. Junior synonym of Barylypa pallida (Gravenhorst) (det. I. D. Gauld).

Anomalon (Anomalon) orbitale, 1892a: 1764. Holotype 9, SWEDEN : Skane, Lund (UZI, Lund).

Labels, [red square] [= Lund, Zetterstedt collection]; O. vagator ? Zett. 9- ex duf [illegible] 15 Mars
1869 [hand]; orbitale m [Thomson cabinet label].
Identity. Gravenhorstia (Erigorgus) orbitale (Thomson) (det. I. D. Gauld).

Anomalon (Exochilum) pyramidatus, 1894: 2118. Holotype (J, SWEDEN: Ostergothland [= Ostergotland]
(UZI, Lund).

Labels. O Got [printed] ; Goes [hand] ; pyramidatus m. [Thomson cabinet label].
Identity. Junior synonym of Therion giganteum (Gravenhorst) (det. I. D. Gauld).

Anomalon (Agrypon) rugifer, 1894: 2119. LECTOTYPE 9, SWEDEN: Ostergothland [= Ostergotland]
(UZI, Lund), here designated (selected by G. A. Viktorov).
Labels. O Got [printed] ; rugifer m [Thomson cabinet label].
Identity. Agrypon rugifer (Thomson) (det. I. D. Gauld).

Anomalon (Agrypon) stenostigma, 1892a: 1771. Lectotype 3, SWEDEN: Skane, Palsjo (UZI, Lund), by
designation of Townes, Momoi & Townes, 1965: 378.
Label. Pal [hand].
Identity. Agrypon stenostigma (Thomson) (det. I. D. Gauld).

Anomalon (Anomalon) varicorne, 1894: 2119. LECTOTYPE 9, SWEDEN: Norrland (UZI, Lund), here
designated (selected by H. Schnee).

Labels. [Grey-green diamond] ; varicorne m [Thomson cabinet label].
Identity. Gravenhorstia (Erigorgus) varicorne (Thomson) (det. I. D. Gauld).

Asthenarus crassifemur, 1889: 1437. LECTOTYPE 9, SWEDEN: Skane, Vestra Wram [= Vastra Vram]
(UZI, Lund), here designated (selected by H. K. Townes).
Labels. V. W. [hand] ; crassifemur [Thomson cabinet label].
Identity. Asthenara crassifemur (Thomson).

18 M. G. FITTON

Atractodes (Polyrhembia) alutaceus, 1884: 1026. Lectotype $, SWEDEN: Skane, Lund (UZI, Lund), by
designation of Jussila, 1979: 40.
Labels. L-d [printed] ; $ [printed].
Identity. Atractodes alutaceus Thomson.

Atractodes (Atractodes) breviscapus, 1884: 1023. Lectotype $, SWEDEN: Skane, Ortofta (UZI, Lund), by
designation of Jussila, 1979: 19.
Labels. Ort [hand]; $ [printed].

Jussila (1979: 19) incorrectly cites the type-locality as Ostergotland. Also, Horstmann's label is dated
1965 not 1956.
Identity. Junior synonym of Atractodes pauxillus Foerster (Jussila, 1979: 19).

Atractodes (Atractodes) compressus, 1884: 1023. Lectotype $, SWEDEN: Halland (UZI, Lund), by des-
ignation of Jussila, 1979: 38.

Labels. Halland [printed] ; compressus [Thomson cabinet label].
Identity. Junior synonym of Atractodes croceicornis Haliday (Jussila, 1979: 37-38).

Atractodes (Atractodes) crassicornis, 1884: 1025. Syntype 1 $, SWEDEN : Skane, Lund (UZI, Lund).
Label. Lund [printed].

The specimen labelled and published by Jussila (1979: 21) as lectotype is from 'Palsio' and therefore
cannot have been a syntype. It has no type status.
Identity. Atractodes crassicornis Thomson, ? junior synonym of Atractodes intersectus Foerster.

Atractodes (Exolytus)fiticornis, 1884: 1020. Syntypes $ ^, SWEDEN: Skane, Bokeberg (lost).
Identity. Mesoleptusfilicornis (Thomson) (based on material in the collection).

Atractodes (Atractodes) flavicoxa, 1884: 1024. Lectotype 9, SWEDEN : Skane, Lund (UZI, Lund), by designa-
tion of Jussila, 1979: 28.
Label. L-d [printed].
Identity. Junior synonym of Atractodes angustipennis Foerster (Jussila, 1979: 28).

Atractodes (Exolytus) flavipes, 1884: 1021. Syntypes 1 $, 6 cJ, SWEDEN: Skane, Palsio [= Palsjo] (UZI,
Lund).

Labels. Pal [hand] (1 $ 4 £). Palsio [printed] (1 J). Pal [hand] ; flavipes [Thomson cabinet label] (1 J).
Identity. Mesoleptus flavipes (Thomson) comb. n.

Atractodes (Atractodes) ttogaster, 1884: 1023. Lectotype $, SWEDEN: Skane, Ortofta (UZI, Lund), by
designation of Aubert, 1966: 129.
Label. Ortofta [printed].
Identity. Junior synonym of Atractodes pusillus Foerster (Jussila, 1979: 19).

Atractodes (Exolytus) marginatus, 1884: 1020. Lectotype ?, SWEDEN: Goteborg (UZI, Lund), by des-
ignation of Aubert, 1966: 130.

Labels. Suecia [printed] ; marginatus [Thomson cabinet label].
Identity. Mesoleptus marginatus (Thomson).

Atractodes (Atractodes) parallel™, 1884: 1024. Lectotype $ [not 9 as stated by Jussila, 1979: 39], SWEDEN:
Skane, Bastad (UZI, Lund), by designation of Jussila, 1979: 39.
Label. Bastad [printed].
Identity. Junior synonym of Atractodes eryptobius Foerster (Jussila, 1979: 39).

Atractodes (Exolytus) petiolaris, 1884: 1020. Lectotype $, SWEDEN : Skane, Raby near Lund (UZI, Lund), by
designation of Aubert, 1966: 130.

Labels. Rab 1/7 [hand] ; petiolaris [Thomson cabinet label].
Identity. Mesoleptus petiolaris (Thomson).

Atractodes (Exolytus) ripicola, 1884: 1021. Lectotype $, SWEDEN : Skane, Ortofta (UZI, Lund), by designati-
on of Townes, Momoi & Townes, 1965: 149.
Label. Ortofta [printed].
Identity. Mesoleptus ripicola (Thomson).

Atractodes (Asyncrita) rufipes, 1884: 1025. Holotype $, SWEDEN: Skane, Ringsjon (UZI, Lund).
Labels. Sk. [hand] ; rufipes [Thomson cabinet label].

Identity. Junior primary homonym of Atractodes rufipes Provancher, 1874: 151 and of Atractodes
rufipes Foerster, 1876: 151 [not 135 as stated by Jussila, 1979: 21] and junior secondary homonym of

THE ICHNEUMONIDAE DESCRIBED BY C. G. THOMSON 19

Atractodes rufipes (Foerster, 1876: 30 (Asyncrita)). Replacement name Atractodes thomsoni (Dalla Torre,
1902: 739 (Asyncrita)) comb. n. The replacement name is itself a secondary homonym of Atractodes
thomsonii Dalla Torre, 1902: 725. However, these latter two names were published simultaneously and,
acting as first reviser (Article 24{a) of the Code), I hereby reject Atractodes thomsonii Dalla Torre, 1902:
725 as the junior name.

Unfortunately, the above names were not taken into account by Jussila (1979) in his revision of western
Palaearctic Atractodes and as a result Atractodes thomsoni Jussila, 1979: 14 and Atractodes rufipes
(Foerster, 1876: 30 [not 14 as stated by Jussila, 1979: 31]) are junior primary and secondary homonyms
respectively, and need replacement names. Replacement names are not here proposed but the problems
have been drawn to the attention of Jussila.

Atractodes (Atractodes) tenuipes, 1884: 1022. Holotype 9, SWEDEN : Skane, Ortofta (UZI, Lund).
Labels. Ort. [hand] ; tenuipes [Thomson cabinet label].
Identity. Atractodes tenuipes Thomson.

Baeosomus oenescens, 1891 : 1615. Syntype 1 9, FRANCE: Furca (UZI, Lund).

Labels. Furca [hand] ; oenescens m [Thomson cabinet label].

It is not clear whether the male mentioned by Thomson (which is in the Kriechbaumer collection,
ZSBS, Munich) should be considered a syntype or excluded from the type-series under Article 72(b) of the
Code.

Some authors (for example, Dalla Torre, 1902: 753) have unjustifiably emended the name to aenescens.

Identity. Baeosemus oenescens (Thomson).

Bane has femor alts, 1897: 241 1. Lectotype 9, SWEDEN: Skane (UZI, Lund), by designation of Townes, Momoi
&Townes, 1965:237.
Label. Scania [printed].
Identity. Junior synonym of Banchus hastator (Fabricius).

Bassus deletus, 1890: 1466, 1471. LECTOTYPE 9, SWEDEN: Lappland (UZI, Lund), here designated (selec-
ted by K. Horstmann).
Label. Lpl [printed].
Identity. Diplazon deletus (Thomson).

Bassus varicoxa, 1890: 1466, 1468. Holotype 9, SWEDEN: Norrland (UZI, Lund).
Labels. Norl. [printed] ; varicoxa n. [Thomson cabinet label].
Identity. Diplazon varicoxa (Thomson).

Blapticus (Blapticus) crassulus, 18886: 1289. Lectotype <$, SWEDEN: Skane, Yddingesjon (UZI, Lund), by
designation of Aubert, 1972: 147-148.
Label. Yddinge [printed].
Identity. Blapticus crassulus Thomson.

Blapticus (Blapticus) dentifer, 18886: 1288. LECTOTYPE £, SWEDEN: Skane, Palsjo (UZI, Lund), here
designated (selected by H. K. Townes).
Label. Pal. [hand].
Identity. Blapticus dentifer Thomson.

Brachycryptus erythrocerus, 1873: 488. Lectotype 9, SWEDEN : Skane, Ringsjon (UZI, Lund), by designation
of Townes & Townes, 1962: 319.
Label. Ringsio [printed].
Identity. Junior synonym of Hidryt afrater (Cresson) (Townes & Townes, 1962: 319).

Brachycryptus fusiventris, 1873: 489. Syntype 1 9, DENMARK : Jutland, Horsens (ZM, Copenhagen).
Labels. 9 23/7 1870 Horsens O. Jensen [hand] ; Danmark ex coll. Schi0dte [printed].
Identity. Hidrytafusiventris (Thomson).

Brachycryptus sordidulus, 1873: 488. Syntypes 2 9, SWEDEN : Skane, Ringsjon (UZI, Lund).
Labels, [small green square] (2 $).
Identity. Hidryta sordidula (Thomson).

Cacotropa sericea, 18886: 1259. Syntype 1 9, ? syntype 1 9, SWITZERLAND: Biel and ? SWEDEN:
Ostergothland [ = Ostergotland] (UZI, Lund).

Labels. 17.v.85/Biel [printed, date hand]; 3235 [printed] ; sericea [hand] (syntype). OG [hand, illegible
and possibly not 'OG'] ; Sp. ign. 9- [hand] ; Col Zet [printed] (? syntype).

Identity. Junior synonym of Sphecophaga vesparum (Curtis).

20 M. G. FITTON

Caenocryptus apum, 1873: 497. Syntype 1 9, DENMARK : Zealand, Dyrehaven (ZM, Copenhagen).
Labels. 9 Dyrehave Drewsen [hand]; Danmark ex coll. Schi0dte [printed].
Identity. Xylophrurus apum (Thomson) comb. n.

Caenocryptus dentifer, 1896: 2362. Holotype 9, SWEDEN : Stockholm (UZI, Lund).

Labels. Col. Hgn. [printed] ; dentifer [hand].

Identity. Junior synonym of Xylophrurus lancifer (Gravenhorst) (Townes, Momoi & Townes, 1965:
202). [Note. Xylophrurus dispar (Thunberg (Ichneumon)) is a junior primary homonym of Ichneumon
dispar Gmelin in Linnaeus.]

Caenocryptus inflatus, 1873 : 497. Syntype 1 9, SWEDEN: Skane, Ringsjon (UZI, Lund).
Label, [small green square].

There are 3 $ in the Zoological Museum, Copenhagen which might also be syntypes, but although they
are from the Drewsen collection they were not collected by Drewsen.
Identity. Enclisis inflatus (Thomson) comb. n.

Caenocryptus laticrus, 1896: 2362. Lectotype 9, SWEDEN: Skane, Ringsjon (UZI, Lund), by designation of
Aubert, 1966: 128.

Label. Ringsio [printed].

Identity. Enclisis laticrus (Thomson) comb. n.

Caenocryptus nubifer, 1896: 2361. Holotype 9, SWEDEN: Vermland [= Varmland] (UZI, Lund).
Labels. Wml [printed] ; nubifer [Thomson cabinet label].
Identity. Enclisis nubifer (Thomson) comb. n.

Caenocryptus pubiventris, 1873: 497. Syntypes 1 9, 1 c? (ZM, Copenhagen), 1 £ (UZI, Lund), DENMARK:
Zealand, Strandmollen.

Labels. 9 8/50 Strandmo'l Drewsen [hand] ; Danmark ex coll. Schicdte [printed] (9). c? Strandmo'l
Drewsen [hand]; Danmark ex coll. Schi0dte [printed] (Copenhagen cJ). pubiventris [Thomson cabinet
label] (Lund £).

Identity. Enclisis pubiventris (Thomson) comb. n.

Caenocryptus striolatus, 1896: 2362. Syntype 1 9, SWEDEN : Skane, Ringsjon (UZI, Lund).
Labels. Scan sylv [printed] ; striolatus [Thomson cabinet label].
Identity. Enclisis striolatus (Thomson) comb. n.

Caenocryptus tener, 1873:496. Syntypes 1 9 (UZI, Lund), 29 9, 14 <J (ZM, Copenhagen), DENMARK:
Zealand.

Labels. Dan [printed] ; tener [Thomson cabinet label] (Lund 9)- Danmark ex coll. Schiedte [printed]
(Copenhagen 99 cJ<J).

The Copenhagen specimens are on pins identical to that of the Lund specimen and obviously from the
same series. They stand with the labels 'Caenocryptus Thorns.' and Tener Thorns' under Caenocryptus
vindex Tschek. Although it is not certain that Thomson saw all of the Copenhagen specimens it is perhaps
desirable that a lectotype is selected from them since the Lund specimen has the gaster missing.

Identity. Enclisis tener (Thomson) comb. n.

Callidora annellata, 1887c: 1136. LECTOTYPE 9, SWEDEN: Skane, Palsjo (UZI, Lund), here designated
(selected by J. F. Aubert).

Labels. Hbg. [hand] ; annellata [Thomson cabinet label].

Identity. Junior synonym of Callidora albovincta (Holmgren) (Townes, 19706: 165).

Campoplex angustatus, 1887c: 1061. Lectotype 9, SWEDEN: Skane, Palsjo (UZI, Lund), by designation of
Aubert, 1966: 130.
Label. Pal [hand].
Identity. Dusona angustata (Thomson).

Campoplex bifidus, 1887c: 1063. Holotype 9, CZECHOSLOVAKIA: Bohemia, Chodau [=Chodov] (UZI,
Lund).

Labels. 26/5 85 Ch [hand] ; bifidus [Thomson cabinet label].
Identity. Dusona bifida (Thomson).

Campoplex castanipes, 1887c: 1063. Holotype 9, CZECHOSLOVAKIA: Bohemia, Chodau [= Chodov] (UZI,
Lund).

Labels. 9/6/76 Ch [hand] ; castanipes [Thomson cabinet label].
Identity. Dusona castanipes (Thomson) comb. n.

THE ICHNEUMONIDAE DESCRIBED BY C. G. THOMSON 21

Campoplex crassipes, 1887c: 1075. Lectotype 9, GERMANY (WEST): Harz (UZI, Lund), by designation of
Aubert, 1968:195.

Labels. Harz [hand] ; crassipes [Thomson cabinet label].

Although Hinz (1963: 339) was almost certainly correct in recognising this as 'Das typische 9'
( = holotype, the single original specimen) Aubert chose to designate it as lectotype. He presented no
evidence of a syntype series.

Identity. Dusona crassipes (Thomson).

Campoplex flaviscapus, 1887c: 1061. Holotype $, FRANCE: Tias (UZI, Lund).
Labels. Tias [hand] ; flaviscapus [Thomson cabinet label].
Identity. Dusona flaviscapus (Thomson).

Campoplex genalis, 1887c: 1070. Lectotype 9, POLAND: Schlesien [= Silesia] (UZI, Lund), by designation of
Aubert, 1968: 195.

Labels. 25 [hand] ; Germ [hand] ; genalis [Thomson cabinet label].

Although Hinz (1963: 337) was almost certainly correct in recognising this as 'Das typische 9'
( = holotype, the single original specimen) Aubert chose to designate it as lectotype. He presented no
evidence of a syntype series.

Identity. Dusona genalis (Thomson).

Campoplex latungula, 1887c: 1065. Holotype 9, U.S.S.R.: Tartu [not 'norra Tyskland' as stated by Thom-
son] (UZI, Lund).

Labels. 21/5 83 [hand]; Dorpat [hand] ; latungula [Thomson cabinet label].

The specimen matches the description and is undoubtedly the holotype. The locality given by Thomson
is obviously a mistake (which he also made in the case of Megastylus pleuralis).

Identity. Junior synonym of Dusona polita (Foerster) (Hinz, 1963: 338).

Campoplex limnobius, 1887c: 1088. Lectotype 9, GERMANY (EAST): Schwerin (UZI, Lund), by designation of
Hinz, 1963:338.

Labels. Schwerin 7.85 [locality printed, date hand] ; limnobius [Thomson cabinet label].
Identity. Dusona limnobia (Thomson).

Campoplex luteipes, 1887c: 1089. Holotype $, SWEDEN : Skane, Ryssjoholm [= Rossjoholm] (UZI, Lund).
Labels. 81. [hand]; Rshm 16/6 [hand] ; luteipes [Thomson cabinet label].
Identity. Dusona luteipes (Thomson).

Campoplex opacus, 1887c: 1074. Holotype 9, GERMANY (WEST): Harz (UZI, Lund).
Labels. Harz [hand] ; opacus [Thomson cabinet label].
Identity. Dusona opaca (Thomson).

Campoplex rectus, 1887c: 1086. Syntypes 9, GERMANY (lost).

There are two specimens in the collection from Chodov (label 'Ch', see Campoplex castanipes above).
Identity. Dusona recta (Thomson) (on the basis of the material in the collection) comb. n.

Campoplex spinipes, 1887c: 1076. Lectotype 9, GERMANY (EAST): Quedlingburg (UZI, Lund), by designation
of Townes, Momoi & Townes, 1965: 289.
Labels. Quedlinburg [hand] ; spinipes [Thomson cabinet label].
Identity. Dusona spinipes (Thomson).

Campoplex splendent, 1887c: 1064. Holotype 9, SWEDEN : Skane, Reften (UZI, Lund).

Labels. Reften 20/vi 82 [hand]; splendens [Thomson cabinet label]. The date '21 Juni' given in the
published description is probably a typographical or copying error.

The holotype was designated 'lectotype' by Hinz (1963: 337). There can be no doubt that the specimen
is a holotype because Thomson specifies 'ett ex'.

Identity. Junior synonym of Dusona polita (Foerster) (Hinz, 1963: 337).

Campoplex stenocarus, 1887c: 1083. Holotype 9, SWEDEN : Gotland (lost).

The specimen labelled and published by Hinz (1963: 339) as holotype is from Smaland (label 'Sm') and
therefore cannot be the type.
Identity. Dusona stenocarus (Thomson).

Canidia anura, 1887c: 1113. Lectotype 9, SWEDEN: Skane, Ortofta (UZI, Lund), by designation of Townes,
Momoi & Townes, 1965 : 282.

Labels. Ort. [hand] ; anura [Thomson cabinet label].
Identity. Bathyplectes anurus (Thomson).

22 M. G. FITTON

Canidia balteata, 1887c: 1114. Lectotype ?, FRANCE: Scarpe near Lille (UZI, Lund), by designation of
Aubert, 1968:195.

Labels. Scarpe [hand] ; balteata [Thomson cabinet label].
Thomson incorrectly attributed authorship of this species to Bridgman.
Identity. Bathyplectes balteatus (Thomson).

Canidia contracta, 1887c: 1113. Lectotype ?, GERMANY (UZI, Lund), by designation of Horstmann,
1974a: 66.
Label. Mormal.
Identity. Junior synonym of Bathyplectes anurus (Thomson) (Horstmann, 1974a: 66).

Canidia corvina, 1887c: 1111. Lectotype ?, SWEDEN: Skane, Lund (UZI, Lund); by designation of Horst-
mann, 19 74a: 70.
Label. L-d [printed].
Identity. Bathyplectes corvinus (Thomson).

Canidia curculionis, 1887c: 1113. Lectotype 9, SWEDEN: Skane, Stehag (UZI, Lund), by designation of
Aubert, 1960:490.

Labels. Steh 7/59 [hand] ; Curculionis [Thomson cabinet label].
Identity. Bathyplectes curculionis (Thomson).

Canidia rostrata, 1887c: 1112. Lectotype 9, SWEDEN: Skane, Bastad (UZI, Lund), by designation of Aubert,
1972: 148.
Label. Bast [hand].
Identity. Bathyplectes rostratus (Thomson).

Canidia stenostigma, 1887c: 1114. Lectotype <$, GERMANY (WEST): Aachen (UZI, Lund), by designation of
Aubert, 1970:276.

Labels. <$ 21 gl. [hand] ; Germ, [hand] ; stenostigma [Thomson cabinet label].
Identity. Bathyplectes stenostigma (Thomson).

Canidia trochantella, 1887c: 1114. Lectotype 9, FRANCE/SPAIN: Pyrenees (UZI, Lund), by designation of
Horstmann, 1974a: 77.

Labels. Pyr. [hand] ; trochantella [Thomson cabinet label].

Aubert's lectotype designation (1966: 130) is not considered valid because he did not indicate (in the
publication or on the labels) which of the two specimens on the mount was designated. The lectotype is
the left hand specimen.

Identity. Junior synonym otBiolysia tristis (Gravenhorst) (Horstmann, 1974a: 77).

Casinaria alboscutellaris, 1887c: 1098. Holotype £, GERMANY (UZI, Lund).

Labels. Oerfelse, Aug [hand, ?? first word] ; Casin. orbital, var. Scut. albo. [hand].
Identity. Junior synonym of Casinaria orbitalis (Gravenhorst).

Casinaria alpina. 1887c: 1 100. Holotype & TYROLEN' (UZI, Lund).
Labels. S6./273. ['86' hand, '273' printed] ; 42. [hand].
Identity. Casinaria alpina Thomson.

Casinaria ischnogaster, 1887c: 1101. LECTOTYPE 9, SWEDEN: Skane, Loparod (UZI, Lund), here des-
ignated (selected by R. Hinz).

Labels. Lop [hand] ; 9 [printed] ; ischnogaster [Thomson cabinet label].
Identity. Casinaria ischnogaster Thomson.

Casinaria monticola, 1887c: 1 100. Holotype <J, FRANCE: Pyrenees (UZI, Lund).
Labels. Berck. [hand]; Gall, [hand] ; monticola [Thomson cabinet label].
Identity. Casinaria monticola Thomson.

Casinaria protensa, 1887c: 1 102. LECTOTYPE <£ GERMANY (UZI, Lund), here designated (selected by J. F.
Aubert).

Label, protensa [Thomson cabinet label].
Identity. Casinaria protensa Thomson.

Casinaria scabra, 1887c: 1099. Lectotype 9, FRANCE (UZI, Lund), by designation of Townes, Momoi &
Townes, 1965:280.

Labels. Berck. [hand]; Gall, [hand] ; scabra [Thomson cabinet label] ; scabra m [hand].
Identity. Casinaria scabra Thomson.

THE ICHNEUMONIDAE DESCRIBED BY C. G. THOMSON 23

Casinaria subglabra, 1887c: 1101. LECTOTYPE 9, ITALY: Trieste (ZSBS, Munich), here designated (selec-
ted by K. Horstmann).

Labels. Triest 22.5.71. Krchb. [hand]; 536. [hand]; rufimanus £ Gr. subglabra Thms. O.E. 1101.9.
[hand].

Identity. Casinaria subglabra Thomson.

Catoglyptus (Stiphrosomus) canattculatus, 1894: 1973. Type(s) 9, GERMANY (lost).

Aubert's publication of a neotype female (1972: 147) for this species is not valid because it does not
comply with the provisions of Article 75(c) of the Code. No attempt is made to validate that 'neotype' here
because there has been no recent revisionary work on this group.

Identity. Sympherta canaliculata (Thomson) (Aubert, 1972: 147, on the basis of the invalid 'neotype').

Catoglyptus (Asthenarus)fusiformis, 1894: 1975. Syntypes 1 9, 1 <J, FRANCE (UZI, Lund).
Labels. Oignies [hand] ; fusiventris [Thomson cabinet label] ($). Oignies [hand] ($).
Identity. Syntactusfusiformis (Thomson) comb. n.

Catoglyptus (Asthenarus) scabriculus, 1894: 1975. Holotype <$ [not 9 as stated by Thomson], SWEDEN:
Skane, Yddinge (UZI, Lund).

Label. Ydd. [hand].

Thomson was misled into believing the specimen to be a female by an elongate piece of debris attached
beneath the apex of the gaster.

Identity. Syntactus scabriculus (Thomson) comb. n.

Catoglyptus (Stiphrosomus) sulcatus, 1894: 1974. Lectotype <J, SWEDEN: Norrland (UZI, Lund), by des-
ignation of Jussila, 1967: 110.
Labels. Norl. [printed] ; sulcatus m [hand].
Identity. Sympherta sulcata (Thomson).

Catomicrus trichops, 18886: 1293. Lectotype $, SWEDEN: Norrland, Lappland (UZI, Lund), by designation
of Aubert, 1972:147.
Label. Lpl. [printed].
Identity. Junior synonym ofEusterinx pusilla (Zetterstedt) (Townes, 1971 : 203).

Centeterus (Eparces) grandiceps, 1891: 1638. LECTOTYPE <?, SWEDEN: Skane, Palsjo (UZI, Lund), here
designated (selected by H. K. Townes).
Label. Pal [hand].
Identity. Eparces grandiceps (Thomson).

Centeterus (Centeterus) nigricornis, 1891 : 1638. Holotype 9, FRANCE: Avignon (UZI, Lund).
Labels. Avignon [hand] ; nigricornis [Thomson cabinet label].
Identity. Centeterus nigricornis Thomson.

Chorinaeus australis, 18876: 201. Type(s) <J, ITALY: Trieste (lost).

Identity. Chorinaeus australis Thomson nomen dubium (Aeschlimann, 1975 : 725).

Chorinaeus brevicalcar, 18876: 200. Lectotype 9, SWEDEN: Skane, Palsjo (UZI, Lund), by designation of
Aeschlimann, 1975: 735.

Labels. Pal. [hand] ; brevicalcar [Thomson cabinet label].
Identity. Chorinaeus brevicalcar Thomson.

Chorinaeus facialis, 18876: 202. Lectotype 9, SWEDEN (UZI, Lund), by designation of Aeschlimann,
1973a:982.

Labels. Coll. L-gh. [printed] ; facialis [Thomson cabinet label].
Identity. Trieces facialis (Thomson).

Chorinaeus longicalcar, 18876: 201. Lectotype 9, GERMANY (EAST): Schwerin (UZI, Lund), by designation of
Townes, Momoi & Townes, 1965: 347.

Labels, f. 12/5. 85 [hand]; Germ, [hand] ; longicalcar [Thomson cabinet label].
Identity. Chorinaeus longicalcar Thomson.

Chorinaeus longicornis, 18876: 201. Lectotype 9, SWEDEN: Skane, Yddinge (UZI, Lund), by designation of
Aeschlimann, 1975: 739.

Labels. 9 [printed]; Yd [hand] [not 'Apd' as stated by Aeschlimann (1975: 739)]. A third, pencil-
written, label is illegible.

Identity. Chorinaeus longicornis Thomson.

24 M. G. FITTON

Chorinaeus nitifrons, 1 8876 : 202. Lectotype 9, SWEDEN : Skane, Arrie (UZI, Lund), by designation of Aeschli-
mann, 1973a: 986.

Labels. Ar [hand] ;9 [printed] ; nitifrons [Thomson cabinet label].
Identity. Trieces nitifrons (Thomson).

Coleocentrus heteropus, 1894: 2122. Holotype 9, SWEDEN : Smaland (UZI, Lund).
Label, heteropus m [Thomson cabinet label].
Identity. Coleocentrus heteropus Thomson.

Colpognathus armatus, 1891 : 1636. Syntypes 1 9, 2 <J, FRANCE (UZI, Lund).

Labels. Avignon, [hand] (9). Toulouse, [hand]; armatus [Thomson cabinet label] (1 $). 29.6.89
Marg [illegible] [hand] (1 <J).

The male specimen from Montpellier, which Thomson mentions, is in the collection of the Zoological
Museum, Copenhagen. It is labelled: Montpellier [hand]; [green square]; <J [printed]; armatus Thms
[hand]; Coll. Wiistnei [printed]. It should perhaps be excluded from the syntype series under Article
72(b)oftheCo<fe.

Identity. Colpognathus armatus Thomson.

Colpognathus divisus, 1891 : 1636. Syntypes 4 9, 4 <J, SWEDEN: Skane (UZI, Lund).

Labels. Hbg [hand] (3 9, 2 <J). Ortofta [printed]; divisus m [Thomson cabinet label] (1 9). Scan
[printed] (1 £). Ld [rest illegible] [hand] (1 £).
Identity. Colpognathus divisus Thomson.

Colpognathus pentagonus, 1891 : 1637. Holotype 9, GREECE: Corfu (ZSBS, Munich).
Labels. Graecia. [printed] ; h. [hand] ; Colpognathus [hand].
Schmiedeknecht (1903 : 301-302) gives further details of the holotype.
Identity. Dicaelotus pentagonus (Thomson) comb. n. (det. E. Diller).

Cratocryptus annulitarsis, 1873: 526. Syntypes 1 9, 2 3, SWEDEN : Skane, Arrie and DENMARK : Zealand (UZI,
Lund).

Labels, annulitarsis [Thomson cabinet label] (9). Ar 6/56 [hand] (1 <$). Dan [hand] (1 J).
Identity. Cubocephalus annulitarsis (Thomson).

Cratocryptus bispinus, 1894: 21 17. Holotype <J, SWEDEN : Norrland (UZI, Lund).
Labels. Norrl. [printed] ; bispinus m [Thomson cabinet label].
Identity. Junior synonym of Amphibulus gracilis Kriechbaumer (teste J. Sawoniewicz).

Cratocryptus femoralis, 1873: 527. Syntypes 1 9, DENMARK: Zealand, Dyrehaven (ZM, Copenhagen); 1 <£
SWEDEN: Skane, Skabersjo (UZI, Lund).

Labels. 9 5/1860 Dyrehav. Drewsen [hand]; Danmark ex coll. Schi0dte [printed] (9). Skb [hand];
femoralis [Thomson cabinet label] (<$).

Identity. Cubocephalus femoralis (Thomson) comb. n.

Cratocryptus opacus, 1873 : 523. Syntypes 2 9, 1 <J, SWEDEN : Skane, Molle and Ringsjon (UZI, Lund).
Labels. Mol [hand] ; opacus [Thomson cabinet label] (1 9). [small green square] (1 9 1 £).
Identity. Javra opaca (Thomson).

Cratocryptus pleuralis, 1873: 526. Lectotype 9, SWEDEN: Norrland (UZI, Lund), by designation of Aubert,
1972: 148.

Label. Norl. [printed].
Identity. Parmortha pleuralis (Thomson).

Cratocryptus ruficoxis, 1873 : 525. Syntypes 5 ?, 3 & SWEDEN : Skane, Palsjo and Ringsjon (UZI, Lund).
Labels. Pal. [hand] (392 <J). Ringsio [printed] (1 9). [green square] (1 9). Scan med. [printed] (1 <J).
Identity. Cubocephalus ruficoxis (Thomson) comb. n.

Cratocryptus sternocerus, 1873: 523. Lectotype 9, SWEDEN : Skane, Skabersjo (UZI, Lund), by designation of
Townes, Momoi & Townes, 1965: 157.
Labels. Skb [hand] ; sternocerus [Thomson cabinet label].
Identity. Cubocephalus sternocerus (Thomson).

Cremastus crasskornis, 1890: 1445, 1454. LECTOTYPE £, GERMANY (UZI, Lund), here designated.
Labels. 538 [hand]; Germ, [hand] ; crassicornis [Thomson cabinet label].

The lectotype is the specimen presumed by Sedivy (1970: 22) to be the holotype. However, Thomson
must have had more than one specimen because he gives a range of length ('long. 2^-3 lin.') for the species.
Identity. Cremastus crassicornis Thomson.

THE ICHNEUMONIDAE DESCRIBED BY C. G. THOMSON 25

Cremastus guttifer, 1890: 1444, 1449. Lectotype^, SWEDEN : Skane, Kjeflinge[= Kavlinge] (UZI, Lund), by
designation of Townes, Momoi & Townes, 1965: 312.
Labels. Kfge [hand] ; guttifer [Thomson cabinet label].
Identity. Temelucha guttifer (Thomson).

Cremastus laeviusculus, 1890: 1445, 1454. Lectotype 9, SWEDEN: Oland (UZI, Lund), by designation of
Sedivy, 1970: 33-34.

Labels. 0. [printed] ; sublaevis [Thomson cabinet label].
Identity. Junior synonym of Cremastus pungens Gravenhorst(Sedivy, 1970: 33).

Cremastus macrostigma, 1890: 1444, 1448. Lectotype $, FRANCE: Lille (UZI, Lund), by designation of
Aubert, 1968: 196.
Label. Gall [hand].
Identity. Junior synonym of Temelucha ophthalmica (Holmgren) (Sedivy, 1971 : 25).

Cremastus radialis, 1890: 1445, 1453. Holotype 9, SWEDEN : Gotland (UZI, Lund).

Labels. G. [hand] [not f ' as stated by Sedivy, 1970: 32] ; radialis [Thomson cabinet label].
Identity. Junior synonym of Cremastus lineatus Gravenhorst (Sedivy, 1970: 31).

Cremastus schoenobius, 1890: 1444, 1446. Lectotype?, SWEDEN : Skane, Kjellby[ = Kallby] (UZI, Lund), by
designation of Townes, Momoi & Townes, 1965: 313.
Label. Sk. [hand] [not 'St' as stated by Sedivy, 1971 : 28].
Identity. Temelucha schoenobia (Thomson).

Cremastus subnasutus, 1890: 1445, 1450. Lectotype ?, GERMANY (UZI, Lund), by designation of Aubert,
1966: 131.

Labels. 535. [hand]; Germ, [hand] ; subnasutus [Thomson cabinet label].
Identity. Temelucha subnasuta (Thomson).

Cryptus arenicola, 1873: 484. Lectotype 9, SWEDEN: Skane, Ilstorp (UZI, Lund), by designation of van
Rossem, 19696: 323.

Labels. lisp 6/7 [hand] ; arenicola [Thomson cabinet label].
Identity. Itamoplex arenicola (Thomson) comb. n.

Cryptus borealis, 1873: 484. Lectotype 9, SWEDEN: Norrland (UZI, Lund), by designation of van Rossem,
19696: 337.
Label. Norl. [printed].
Identity. Junior synonym of Itamoplex dianae (Gravenhorst) (van Rossem, 19696: 336).

Cryptus curvicauda, 1896: 2350. Lectotype 9, SWEDEN: Ostergothland [= Ostergotland] (UZI, Lund), by
designation of van Rossem, 1971 : 203.

Labels. O.G. [printed] ; bellitarsis m. [Thomson cabinet label].
Identity. Junior synonym of Buathra tarsoleuca (Schrank) (van Rossem, 1971 : 203).

Cryptus infumatus, 1873: 481. Lectotype 9, SWEDEN: Skane, Palsjo (UZI, Lund), by designation of van
Rossem, 19696: 343.
Label. Pal [hand].
Identity. Junior synonym of Itamoplex titubator (Thunberg) (van Rossem, 19696: 341).

Cryptus latitarsis, 1873: 483. Lectotype 9, SWEDEN: Oland, Borgholm (UZI, Lund), by designation of van
Rossem, 1969a: 177.
Label. Oel [printed].
Identity. Junior synonym ofMeringopus titillator (Linnaeus) (van Rossem, 1969a: 174).

Cryptus serratus, 1873: 478. Holotype 9, SWEDEN: Skane, Bogestad [= Bokestad] (UZI, Lund).
Labels. Bgs. [hand] ; serratus [Thomson cabinet label].
Identity. Junior synonym of Meringopus nigerrimus (Fonscolombe) (van Rossem, 1969a: 191).

Cryptus subquadratus, 1873: 478. Lectotype 9, SWEDEN: Skane, Esperod [= Asperod] (UZI, Lund), by
designation of van Rossem, 19696: 364.

Label. Scania [printed].

The lectotype was labelled 'type' (= holotype) by Townes and Townes, Momoi & Townes (1965: 184)
were almost certainly correct in recognising it as such. However, van Rossem chose to designate it as
lectotype. He did not present any evidence of a syntype series.

Identity. Itamoplex subquadratus (Thomson).

26 M. G. FITTON

Cteniscus albicoxa, 1883: 891. Holotype 9, SWEDEN : Skane, Ryssioholm [ = Rossjoholm] (UZI, Lund).
Labels. Rshm 16/6 [hand] ; albicoxa [Thomson cabinet label].
Identity. Eridolius albicoxa (Thomson).

Cteniscus brevigena, 1883: 893. Holotype 9, SWEDEN: Skane, Stehag(UZI, Lund).
Label, [small green square].
Identity. Eridolius brevigena (Thomson).

Cteniscus breviventris, 1883: 890. Lectotype $, SWEDEN: Skane, Torekov (UZI, Lund), by designation of
Kerrich, 1952: 439.
Label. Tkov 7/60 [hand].
Identity. Junior synonym of Eridolius rufonotatus (Holmgren) (Kerrich, 1952: 437).

Cteniscus deletus, 1883: 894. Holotype 9, SWEDEN : Norrland (UZI, Lund).
Labels. Norl. [printed] ; deletus [Thomson cabinet label].
Identity. Eridolius deletus (Thomson).

Cteniscus genab's, 1883: 894. Holotype?, SWEDEN: Skane, Alnarp(UZI, Lund).
Labels. Scan [printed] ; genalis [Thomson cabinet label].
Identity. Eridolius genalis (Thomson) comb. n.

Cteniscus lineiger, 1883: 894. LECTOTYPE & SWEDEN: Ostergotland, Mjolsefall (NR, Stockholm), here
designated.

Labels. 5 ug 5/8 Mjfall. [hand] ; <$ [hand] ; extirpatorius Gr sec. Hgn. [hand].

The lectotype is the specimen recognised by Kerrich (1952: 425) as 'type'. It is not clear from Kerrich's
statement whether he intended to make a type restriction or merely recognised the specimen as one of the
syntypes.

Identity. Eridolius lineiger (Thomson).

Cteniscus marginatus, 1883: 892. LECTOTYPE 9, SWEDEN: Skane, Ortofta (UZI, Lund), here designated
(selected by G. J. Kerrich).
Label. Ort. [hand].
Identity. Eridolius marginatus (Thomson).

Cteniscus punctipes, 1883: 892. LECTOTYPE 9, SWEDEN: Lappland (UZI, Lund), here designated (selected
by G. J. Kerrich).
Label. Lpl. [printed].
Identity. Eridolius punctipes (Thomson).

Cteniscus punctipleuris, 1883: 893. Syntypes ?£, SWEDEN: Skane, Kjellby [= Kallby] (lost).
Identity. ? Eridolius punctipleuris (Thomson) comb. n.

Cteniscus quadrinotatus [as 4-notatus~\, 1883: 892. Holotype 9, SWEDEN: Lappland (UZI, Lund).
Labels. Lpl. [printed] ; 4-notatus [Thomson cabinet label].
Identity. Eridolius quadrinotatus (Thomson) comb. n.

Cteniscus signifer, 1883: 893. LECTOTYPE 9, SWEDEN: Skane, Lindholmen (UZI, Lund), here designated
(selected by G. J. Kerrich).
Labels. Lhn 15/7 [hand]; 109. [hand].
Identity. Eridolius signifer (Thomson).

Cteniscus t-nigrum, 1883: 891. LECTOTYPE 9, SWEDEN: Skane, Klinta (UZI, Lund), here designated
(selected by G. J. Kerrich [labelled as 'holotype']).

Labels. Scan [printed]; 9 [printed].

Identity. Eridolius t-nigrum (Thomson).
Ctenopelma verticina, 1883: 925. Syntypes 5 9, SWEDEN: Skane, Alnarp (UZI, Lund).

Labels. Scan [printed] ; verticina [Thomson cabinet label] (1 9)- Scan [printed] (49).

Identity. Ctenopelma verticinum Thomson.

Cymodusa longicalcar, 1887c: 1096. Type(s) [?sex], SWEDEN: Skane, Esperod[= Asperod] (lost).
Identity. Cymodusa longicalcar Thomson.

Deloglyptus punctiventris, 1891 : 1623. LECTOTYPE 9, SWEDEN: Skane, Lund (UZI, Lund), here designated
(selected by H. K. Townes).

Labels. Scan [printed] ; punctiventris m [Thomson cabinet label].
Identity. Dicaelotus punctiventris (Thomson).

THE ICHNEUMONIDAE DESCRIBED BY C. G. THOMSON 27

Delotomus auriculatus, 1883 : 884. Syntypes 3 $, 2 cJ, SWEDEN : Skane, Palsjo and Ilstorp (UZI, Lund).

Labels. Pal [hand]; auriculatus [Thomson cabinet label] (1 £). Pal [hand] (2 ?). lisp 26/6 [hand]

(1 ? 1 <J).

Identity. Junior synonym of Acrotomus lucidulus (Gravenhorst) (Kerrich, 1952: 452).

Delotomus binotatus, 1883: 886. Lectotype 9, SWEDEN: Oland (UZI, Lund), by designation of Kerrich,
1952: 328.

Labels. 0. [printed] ; binotatus [Thomson cabinet label].
Identity. Junior synonym ofCycasis rubiginosus (Gravenhorst) (Kerrich, 1952: 328).

Delotomus calcaratus, 1883: 885. Holotype $, SWEDEN : Norrland (UZI, Lund).
Labels. Norl. [printed] ; calcaratus [Thomson cabinet label].
Identity. Junior synonym of Kristotomus laetus (Gravenhorst) (Kerrich, 1952: 347).

Delotomus mar ginatus, 1883: 885. Syntypes 1 ?, 1 <$, SWEDEN : Oland, Isgarde(UZI, Lund).
Label. 0. [printed].
Both specimens are on the same pin.
Identity. Junior synonym of Kristotomus laetus (Gravenhorst) (Kerrich, 1952: 346).

Delotomus parvulus, 1883: 886. Type(s) [? sex], SWEDEN: Oland (lost).

Identity. Junior synonym of Cycasis rubiginosus (Gravenhorst) (Kerrich, 1952: 328).

Demophorus annellatus, 1890: 1458. LECTOTYPE £, SWEDEN: Skane, Kjeflinge [= Kavlinge] (UZI,
Lund), here designated.

Labels. Kfge [hand] ; annellatus [Thomson cabinet label].

There are already three published mentions of a lectotype for this species but none constitutes a proper
designation. The earliest (Aubert, 1959: 163) was apparently not intended to be a designation and in-
cludes no details of any particular syntype specimen.The second (Aubert, 1966: 131) specifies 'cT and 'No.
1' but the pin (the first in the series) bears three males (and an additional empty card point) and there is no
indication either in the publication or on Aubert's label which specimen is intended to be the lectotype.
The third (Sedivy, 1970: 6) refers to Aubert's first publication (incorrectly as 1958); does not specify a
particular specimen ; and gives the sex, incorrectly, as female.

In order to clarify the situation a lectotype is designated here. It is the lowest of three male specimens on
the pin bearing Aubert's lectotype label.

Identity. Junior synonym ofDimophora evanialis (Gravenhorst) (Sedivy, 1970: 5).

Demophorus arenicola, 1890: 1457. Lectotype 9, SWEDEN: Skane, Ilstorp (UZI, Lund), by designation of
Sedivy, 1970: 7.
Label. lisp 15/7 [hand].
Identity. Junior synonym ofDimophora robust a Brischke (Sedivy, 1970: 6).

Diaborus filipalpis, 1883: 889. Lectotype $, SWEDEN: Skane, Holmeja (UZI, Lund), by designation of Ker-
rich, 1953: 154 [as 'type'].
Label. Hme [hand].
Identity. Junior synonym of Cteniscus pedatorius (Panzer) (Kerrich, 1953: 158).

Diaborus frontalis, 1883: 889. Syntypes 2 9, 4 <J, SWEDEN : Skane, Ortofta (UZI, Lund).

Labels. Ort [hand]; frontalis [Thomson cabinet label] (1 <J). Ort. [hand] (1 ? 2<J). Ortofta [printed]
(1 cJ). [hand, illegible] ; Ort. [hand] (1 ?).

Identity. Cteniscus frontalis (Thomson) comb. n.

Diaborus nigrifrons, 1883: 889. Type(s), SWEDEN : Skane, Esperod [ = Asperod] (lost).

Neotype $, SWEDEN: Skane, Palsjo (UZI, Lund), by designation of Kerrich, 1953: 153-154.
Labels. Pal [hand] ; nigrifrons [Thomson cabinet label].
Identity. Cteniscus nigrifrons (Thomson).

Diaborus pallit arsis, 1883: 889. Lectotype ?, SWEDEN: Skane, Esperod [= Asperod] (UZI, Lund), by
designation of Kerrich, 1953: 152.

Labels. Scan [printed] ; pallitarsis [Thomson cabinet label].
Identity. Cteniscus pallitarsis (Thomson).

Diadromus (Diadromus) arcticus, 1891 : 1634. Syntypes 2 ?, 1 <$, SWEDEN: Lappland (UZI, Lund).

Labels, [two small squares of paper]; Lpl. [printed]; arcticus [Thomson cabinet label] (1 $). Afolu 29
Jul. [hand] ; Lpl. [printed] (1 $). [small square of paper] ; Col. Ros [printed] (1 (J).
Identity. Diadromus arcticus Thomson.

28 M. G. FITTON

Diadromus (Diadromus) medialis, 1891 : 1634. Syn types 2 ?, 1 cJ, SWEDEN: Lappland (UZI, Lund).

Labels, [small square of paper]; Col Ros [printed]; medialis m [Thomson cabinet label] (1 $). Norl.
[printed] (1 ?). [small square of paper]; Col Ros [printed]^).

The two females belong to different species, the one labelled 'Norl.' has the gaster missing and the other
has the head missing.

Identity. Diadromus medialis Thomson.

Dicoelotus [lapsus for Dicaelotus~\ (Cinxaelotus) annellatus, 1891: 1621. Holotype 9, SWEDEN :0land (UZI,
Lund).

Label. O. [printed].
Identity. Dicaelotus annellatus Thomson.

Dicoelotus [lapsus for Dicaelotus] (Cinxaelotus) crassifemur, 1891: 1620. Lectotype 9, SWEDEN: Skane,
Ilstorp (UZI, Lund), by designation of Townes, Momoi & Townes, 1965: 423.
Labels. lisp 13/7 [hand] ; crassifemur m [Thomson cabinet label].
Identity. Dicaelotus crassifemur Thomson.

Dicoelotus [lapsus for Dicaelotus'] (Dicoelotus) [lapsus] inflexus, 1891 : 1619. Syntypes 2$, SWEDEN: Skane,
Kjeflinge [= Kavlinge] (UZI, Lund).

Labels. Kfge [hand] ; inflexus [Thomson cabinet label] (1 $). Kfge [hand] (1 $).
Identity. Dicaelotus inflexus Thomson.

Dicoelotus [lapsus for Dicaelotus] (Dicoelotus) [lapsus] orbitalis, 1891: 1621. Holotype 9, SWEDEN: Skane,
Stehag (UZI, Lund).

Labels. Steh 2/5 [hand] ; orbitalis m [Thomson cabinet label].
Identity. Dicaelotus orbitalis (Thomson).

Ephialtes abbreviatus, 1877: 740. Syntypes 7 9, 1 <J, SWEDEN: Skane (UZI, Lund).

Labels. Pal [hand]; abbreviatus [hand] (1 9). Pal [hand] (3 ?). [small green square] (2 $). Scan
[printed] (1 ?). Tkov 7/60 [hand] ; abbreviatus [hand] (1 <J).
Identity. Junior synonym of Dolichomitus populneus (Ratzeburg) (Oehlke, 1967: 14).

Ephialtes antefurcalis, 1877: 741. Holotype 9, SWEDEN: Skane (UZI, Lund).
Labels. Sk [printed].

Townes, Momoi & Townes (1965: 17) say Types' ( = syntypes) but there is no reason to suppose that
the one female from Skane should not be the only original specimen (= holotype).
Identity. Junior synonym of Townesia tenuiventris (Holmgren) (Oehlke, 1967 : 1 1).

Ephialtes crassiceps, 1877: 739. Type(s) [? sex], SWEDEN: Lappland (lost).

Perkins (1943a: 255, 256) indicated that he had seen 'the type or the type series' of this species but there
are no specimens in the collection which can be types.

Identity. Junior synonym of Dolichomitus dux (Tschek) (Oehlke, 1967: 13).

Ephialtes crassiseta, 1877: 743. Holotype 9, SWEDEN : Norrland (UZI, Lund).
Label. Norl. [printed].
Identity. Liotryphon crassisetus (Thomson).

Ephialtes gnathaulax, 1877: 739. Lectotype 9, SWEDEN: Skane (UZI, Lund), by designation of Townes,
Momoi & Townes, 1965: 18.
Label. Scan bor. [printed].
Identity. Paraperithous gnathaulax (Thomson).

Ephialtes heteropus, 1888ft: 1249. Holotype 9, SWEDEN: Skane, Lund (UZI, Lund).

Labels. Lund [printed] ; exclusus e Cerambyx moschata [hand] ; heteropus n. sp. [hand].
Identity. Junior synonym of Dolichomitus messor (Gravenhorst) (Oehlke, 1967: 13).

Ephialtes luteipes, 1877: 740. Holotype 9, SWEDEN: Skane (UZI, Lund).
Label. Scan [printed].

There is a second female in the collection (from Bokestad in Skane [label 'Bgs']) which could also be the
type but which is not in such good agreement with the original description.
Identity. Paraperithous luteipes (Thomson).

Ephialtes macrurus, 1894: 2123. Holotype 9, SWEDEN: Stockholm (UZI, Lund).
Labels. Holm [printed] ; macrurus m [hand].
Identity. Junior synonym of Dolichomitus cognator (Thunberg) (Oehlke, 1967: 12).

THE ICHNEUMONIDAE DESCRIBED BY C. G. THOMSON 29

Ephialtes parallelus, 18886: 1248. Lectotype 9, SWEDEN: Skane, Farhult (UZI, Lund), by designation of
Townes, Momoi & Townes, 1965: 22.

Labels. E. tubercula. tus. 9- 3. 228 105. O. foveolata Fall, [hand]; parallelus m Farhult [hand].
Identity. Junior synonym of Dolichomitus tuberculatus (Geoffroy) (Oehlke, 1967: 15).

Ephialtes planifrons, 1877: 741. Holotype 9, SWEDEN: Norrland (UZI, Lund).
Label. Norl. [printed].
Identity. Junior synonym of Dolichomitus terebrans (Ratzeburg) (Oehlke, 1967: 15).

Ephialtes pleuralis, 1877: 744. Holotype 9, SWEDEN: Lappland (UZI, Lund).
Labels, [small square of paper]; Col Ros [printed] ; pleuralis [hand].
Identity. Junior synonym of Liotryphon crassisetus (Thomson) (Oehlke, 1967 : 9).

Ephialtes scutellaris, 1877: 738. Type(s) 9, SWEDEN: Skane (lost).

Perkins (1943a: 256) indicated that he had seen 'the type or the type series' of this species but there are
no specimens in the collection which can be types.

Identity. Dolichomitus scutellaris (Thomson) (Oehlke, 1967: 14).

Epitomus parvus, 1891: 1626. Lectotype $, SWEDEN: Skane, Lund (UZI, Lund), by designation of Aubert,
1966: 128.

Label. L-d [printed].
Identity. Epitomus parvus Thomson.

Erromenus are nicola, 1883: 905. Syntype 1 <$, SWEDEN : Skane, Degeberga (UZI, Lund).
Label. Dgb. [hand].
Identity. Junior synonym of Erromenus junior (Thunberg) (Kasparyan, 1973 : 296).

Erromenus brevitarsis, 1883 : 904. Syntypes 4 9, SWEDEN : Skane, Klinta (UZI, Lund).

Labels. Rsio [printed]; brevitarsis [Thomson cabinet label] (1 $). Rsio [printed] (1 $). [small green
square] (2 ?).

Identity. Junior synonym of Erromenus plebejus (Woldstedt) (Kasparyan, 1973 : 299).

Erromenus cavigena, 1883: 904. Lectotype <J, SWEDEN: Skane, Ronnemolla (UZI, Lund), by designation of
Aubert, 1968: 195.

Labels. Ron [hand] ; <J [printed] ; cavigena [Thomson cabinet label].
Identity. Junior synonym of Erromenus melanotus (Gravenhorst) (Kasparyan, 1973 : 298).

Erromenus simplex, 1883: 905. Lectotype $, SWEDEN: Norrland (UZI, Lund), by designation of Townes,
Momoi & Townes, 1965: 104.

Labels. Norl. [printed] ; simplex [Thomson cabinet label].
Identity. Junior synonym of Erromenus punctatus (Woldstedt) (Kasparyan, 1973 : 301).

Eurylabus vinulator, 1894: 2102. LECTOTYPE <?, SWEDEN: Skane, Stehag (NM, Goteborg), here
designated.

Labels. Ste CM [hand].

The lectotype is in the G. F. M oiler collection and is one of the two specimens specifically mentioned by
Thomson. The original syntype series of this species consists of the lectotype and a male in Marklins
collection (there are two males from Marklins collection in Uppsala) together with the material which
formed the bases of the publications of Degeer, Wesmael and Holmgren cited by Thomson.

Identity. Junior synonym of Eurylabus larvatus (Christ).

Euryproctus (Syndipnus) atricornis, 1883 : 928. Syntypes 8 9, 5 <£ SWEDEN : Skane, Lund (UZI, Lund).

Labels. L-d [printed]; atricornis [Thomson cabinet label] (3 9 on one pin). Ld [hand] (1 9)- L-d
[printed] ; Synoditus [hand] (2 $ 1 <$ on one pin). [No label] (29 2<J on one pin). [No label] (2$ on one
pin).

Identity. Syndipnus atricornis (Thomson).

Euryproctus (Himertus) bisannulatus, 1883: 928. Syntypes 5 9, 4 3, SWEDEN: Skane (UZI, Lund).

Labels, [green square] ; 9 [printed] ; bisannulatus [Thomson cabinet label] (1 9)- Scania [printed] (1 $).
Ort. [hand] (2 £). Ort. [hand]; 9 [printed] (1 9). [green square] (19 1 c?)- Ringsio [printed] (1 9)- Rsio
[printed] (1 rf).

Identity. Himerta bisannulata (Thomson).

Euryproctus (Euryproctus) crassicornis, 1889: 1433. LECTOTYPE 9, SWEDEN: Skane, Palsjo (UZI, Lund),
here designated (selected by M. Idar).

Labels. Palsio [printed] ; crassicornis [Thomson cabinet label].
Identity. Euryproctus crassicornis Thomson.

30 M. G. FITTON

Euryproctus (Euryproctus) exareolatus, 1889: 1435. LECTOTYPE <J, SWEDEN: Skane, Ringsjon (UZI,
Lund), here designated (selected by M. Idar).
Label. Rsio [printed].
Identity. Euryproctus exareolatus Thomson.

Euryproctus (Euryproctus) inferus, 1889: 1435. Holotype 9, SWEDEN : Skane, Yddinge (UZI, Lund).
Labels. Yddinge [printed] ; inferus [Thomson cabinet label].
Identity. Euryproctus inferus Thomson.

Euryproctus (Phobetus) Kopleuris, 1889: 1431. Syntype 1 [? sex], SWEDEN : Skane, Ortofta (UZI, Lund).
Label. Ort [hand].

The gaster is missing from the specimen.
Identity. Phobetes liopleuris (Thomson).

Euryproctus (Syndipnus) macrocerus, 1883:928. LECTOTYPE ?, SWEDEN: Skane, Ryssjoholm
[ = Rossjoholm] (UZI, Lund), here designated (selected by H. K. Townes).
Label. Rshm 16/6 [hand].
Identity. Syndipnus macrocerus (Thomson).

Euryproctus (Euryproctus) nitidulus, 1889: 1436. Holotype 9, SWEDEN : Skane, Arrie (UZI, Lund).
Labels. Ar [hand] ; nitidulus [Thomson cabinet label].
Identity. Euryproctus nitidulus Thomson.

Euryproctus (Euryproctus) parvulus, 1883: 926. LECTOTYPE ?, SWEDEN: Skane, Fogelsang
[ = Fagelsang] (UZI, Lund), here designated (selected by M. Idar).
Labels. Fsg 1/7 [hand] ; 9 [printed].
Identity. Euryproctus parvulus Thomson.

Exenterus (Exenterus) claripennis, 1883 : 887. Syntypes $ <£, SWEDEN: Skane, Wittsio [ = Vittsjo] (lost).
Neotype 9, SWEDEN: Dalarne, Fulufjall (NR, Stockholm), by designation of Kerrich, 1952: 360.
Identity. Exenterus claripennis Thomson.

Exenterus (Exenterus) flavellus, 1883: 887. Holotype 9, SWEDEN: Skane Kjeflinge [= Kavlinge] (UZI,
Lund).

Labels, [note, in Swedish, describing capture in pine plantation 19.ix.1839] ; flavellus [Thomson cabinet
label].

Identity. Junior synonym of Exenterus oriolus Hartig (Kerrich, 1952: 357).

Exenterus laricinus, 18886: 1254. Holotype 9, SWEDEN: Skane, Ramlosa (UZI, Lund).
Labels. Raml. 7.85 [hand] ; Laricinus [Thomson cabinet label].

Identity. Exenterus laricinus Thomson. Placed as a synonym of Exenterus adspersus Hartig by Kerrich
(1952: 360), for which a lectotype has since been designated (Townes, Momoi & Townes, 1965: 114).

Exenterus (Exenterus) simplex, 1883 : 887. Syntypes 2 9, SWEDEN: Gottland [ = Gotland] (UZI, Lund).
Labels. Gott [printed] (1 9)- [red square] ; [grey square] ; Suecia [printed] ; muticus [hand] (1 $).
Identity. Exenterus simplex Thomson.

Exetastes guttifer, 1897: 2417. Lectotype 9, SWEDEN: Gottland [= Gotland] (UZI, Lund), by designation of
Aubert, 1972: 146.
Label. Gott [printed].
Identity. Exetastes guttifer Thomson.

Exochus annutttarsis, 18876: 215. Syntype 1 9, SWEDEN: Skane, Rostanga (UZI, Lund).

Label. Rota [hand, partly illegible].

Thomson only gave 'Suecia' as the locality in the original description. Later (1894: 2136) he stated
'Funnen vid Rostanga: Skane'. Seven other specimens are from other localities and were, therefore,
probably added to the collection after 1894 and unlikely to be syntypes.

Identity. Exochus annulitarsis Thomson.

Exochus anospilus, 18876: 217. Type(s) [? sex], GERMANY (lost).

There are two specimens in the collection from Dalmatia (label 'Dalm').
Identity. Exochus anospilus Thomson.

Exochus australis, 1894: 2137. Lectotype 9, ITALY: Trieste (UZI, Lund), by designation of Townes, Momoi
&Townes, 1965: 358.

Label. 8.IX Triest. [date, hand; locality, printed].
Identity. Exochus australis Thomson.

THE ICHNEUMONIDAE DESCRIBED BY C. G. THOMSON 31

Exochus citripes, 18876: 213. Lectotype $, FRANCE: Lille, Oisy (UZI, Lund), by designation of Aubert,
1972: 147.

Labels. Oisy. [hand]; 9 [printed] ; citripes [Thomson cabinet label].
Identity. Exochus citripes Thomson.

Exochus crassicornis, 1894: 2134. Holotype 9, SWEDEN: Dalsland (lost).
Identity. Exochus crassicornis Thomson.

Exochus incidens, 18876: 208. Syntypes 4 9, 9 <J, 2 [? sex], SWEDEN (UZI, Lund).

The syntype series includes material from the following localities : Skane, Ortofta [label 'Ort'] ; Rings-
jon [label, green square] ; Kavlinge [label 'Kfge'] ; Vastra Vram [label 'V.W.']. Oland [label '0']. Gotland
[label 'G.']. Ostergotland [label 'OG'].
Identity. Exochus incidens Thomson.

Exochus lineifrons, 1887ft : 213. Syntypes 4 9, 5 & SWEDEN: Skane, Palsjo (UZI, Lund).

Thomson only gave 'Suecia' as the locality in the original description but later (1894: 2135) gave the
more precise information 'vid Palsjo'. All of the specimens in the collection are from this locality [labels
'Palsio' (printed) and 'Pal' (hand)].

Identity. Exochus lineifrons Thomson.

Exochus longicornis, 18876: 214. Lectotype 9, SWEDEN: Skane, Palsjo (UZI, Lund), by designation of
Aubert, 1966: 128.

Labels. Palsio [printed] ; longicornis [Thomson cabinet label].
Identity. Exochus longicornis Thomson.

Exochus nigripalpis, 18876: 209. Lectotype 9, SWEDEN: Skane, Ortofta (UZI, Lund), by designation of
Townes & Townes, 1959: 217.
Label. Ort. [hand].
Identity. Exochus nigripalpis Thomson.

Exochus parvispina, 18876: 216. Syntype 1 9, SWEDEN : Skane, Bokeberg (UZI, Lund).

Labels. Bok 8/78 [hand] ; parvispina [Thomson cabinet label].

Thomson only gave 'Suecia' as the locality in the original description. Later (1894: 2136) he stated
'Funnen vid Bokeberg i Skane'. Therefore, a second female in the collection from Ostergotland (label
'OG') probably post dates 1894 and is unlikely to be a syntype.

Identity. Exochus parvispina Thomson.

Exochus signifrons, 18876: 216. LECTOTYPE 9, SWEDEN: Jamtland, Areskutan (UZI, Lund), here des-
ignated (selected by J. F. Aubert).

Labels. Areskut. lapp 2 Ag. [hand].

Thomson (1894: 2136) later referred to this species as nigrifrons. There is no evidence that this was a
deliberate change and it is here regarded as an incorrect subsequent spelling (Article 33(b) of the Code).

Identity. Exochus signifrons Thomson.

Exyston brevipetiolatus, 1883: 883. Syntypes 7 9, 5 <J, SWEDEN: Stockholm area and Smaland, Anneberg
(NR, Stockholm).

Labels. Him [printed] ; Bhn [= Boheman] [printed] (493 <£). Sm [printed] ; Bhn [printed] (39 2<$).

The syntype series of this species comprised Thomson's own specimen(s) from 'Vestergothland' (which
are lost) together with the material which was the basis of Holmgren's misidentification of Exenterus
triangulatorius.

Identity. Junior synonym of Exyston pratorum (Woldstedt) (Kerrich, 1952: 385).

Exyston calcaratus, 1883: 883. LECTOTYPE 9, SWEDEN: Skane, Palsio [= Palsjo] (UZI, Lund), here
designated (selected by H. K. Townes).

Labels. Pal. [hand] ; calcaratus [Thomson cabinet label].
Identity. Exyston calcaratus Thomson.

Exyston carinatus, 1883: 882. Lectotype 9, SWEDEN: Skane, Palsjo (UZI, Lund), by designation of
Kasparyan, 1975:304.

Labels. Pal [hand] ; carinatus [Thomson cabinet label].
Identity. Junior synonym of Exyston sponsorius (Fabricius) (Kasparyan, 1975: 304).

Exyston genalis, 1883: 883. Syntype 1 9, SWEDEN: Lappland (UZI, Lund).
Labels, [square of paper]; Col Ros [printed].
Identity. Exyston genalis Thomson.

32 M. G. FITTON

Gambrus inferus, 1896: 2375. Syntype 1 £, SWEDEN : Skane, Lund (UZI, Lund).
Label. Ld [hand].
Identity. Gambrus inferus Thomson.

Gambrus superus, 1896: 2375. Syntypes 9 <$, SWEDEN: Skane, Ortofta (lost).

Aubert's publication of a male neotype (1966: 128) for this species is not valid because it does not
comply with the provisions of Article 75(c) of the Code. No attempt is made to validate that 'neotype' here
because there has been no recent revisionary work on the Palaearctic species of Gambrus.

Identity. Gambrus superus Thomson.

Glypta (Glypta) brevipetiolata, 1889: 1327, 1344. Lectotype 9, SWEDEN: Skane, Kjeflinge [= Kavlinge]
(UZI, Lund), by designation of Aubert, 19786: 38.
Label. Kfge [hand].
Identity. Glypta brevipetiolata Thomson.

Glypta (Glypta) breviventris, 1889: 1327, 1347. Holotype?, SWEDEN: Lappland (UZI, Lund).
Labels. Lap [hand] ; breviventris [Thomson cabinet label].
Identity. Junior synonym of Glypta crassitarsis Thomson (Aubert, 19786: 46).

Glypta (Glypta) caudata, 1889: 1326, 1337. Holotype 9, SWEDEN: Skane, Ringsjo (UZI, Lund).
Labels. Scania [printed] ; caudata [Thomson cabinet label].
Identity. Glypta caudata Thomson.

Glypta (Glypta) crassitarsis, 1889: 1327, 1346. Lectotype ?, SWEDEN: Norrland (UZI, Lund), by designation
ofTownes, Momoi & Townes, 1965: 211.
Label. Norl. [printed].
Identity. Glypta crassitaris Thomson.

Glypta (Glypta) crenulata, 1889: 1325, 1334. Lectotype <J, FRANCE (UZI, Lund), by designation of Aubert,
1972: 146.

Labels. Gall [hand] ; crenulata [Thomson cabinet label].
Identity. Junior synonym ofApophua cicatricosa (Ratzeburg) (Aubert, 19786: 34).

Glypta (Glypta) dentifera, 1889: 1350. Syntypes 6$, GERMANY (UZI, Lund).

Labels. Kosen 9.8.51 [hand]; Germ [hand]; dentifera [Thomson cabinet label] (1 9)- Koesen 7.88
[hand] ; Germ [hand] (3 ?). Germ [hand] (2 ?).
Identity. Glypta dentifera Thomson.

Glypta (Glypta) filicornis, 1889: 1328, 1351. Lectotype 9, SWEDEN: Skane (UZI, Lund), by designation of
Townes, Momoi & Townes, 1965: 212.
Label. Sk [hand].
Identity. Junior synonym of Glypt afemorator Desvignes (Aubert, 19786: 40).

Glypta (Glypta) fractigena, 1889: 1325, 1334. Lectotype 9, FRANCE: Lille (UZI, Lund), by designation of
Townes, Momoi & Townes, 1965: 212.
Labels. Lille [hand]; Gall. [hand].
Identity. Junior synonym of Glypta nigrina Desvignes (Aubert, 19786: 44).

Glypta (Glypta) heterocera, 1889: 1326, 1337. Lectotype 9, SWEDEN: Skane, Palsjo (UZI, Lund), by des-
ignation of Aubert, 1968: 194.
Label. Pal [hand].
Identity. Glypta heterocera Thomson.

Glypta (Glypta) microcera, 1889: 1328, 1350. Syntypes 4 9, GERM ANY (WEST): Harz (UZI, Lund).
Labels. Harz 8.85 [hand] ; microcera [Thomson cabinet label] (1 9). Harz 8.85. [hand] (39).
Identity. Glypta microcera Thomson.

Glypta (Glypta) nigricornis, 1889: 1328, 1353. Lectotype 9, SWEDEN: Skane, Palsjo (UZI, Lund), by des-
ignation of Aubert, 1972: 146.

Labels. Pal [hand] ; nigricornis [Thomson cabinet label].

There is a male specimen (paralectotype) on the same pin as the lectotype.

Identity. Glypta nigricornis Thomson.

Glypta (Glypta) nigriventris, 1889: 1325, 1336. Lectotype 9, SWEDEN: Skane, Ringsjon (UZI, Lund), by
designation of Aubert, 1976a: 154.
Label, [green square].

THE ICHNEUMONIDAE DESCRIBED BY C. G. THOMSON 33

Aubert's prior publication of a neotype (1968: 194) for this species is, fortunately, not valid because it
does not comply with the provisions of Article 75(c) of the Code. Therefore, no reference to the Commis-
sion is needed (Article 75(f )).

Identity. Junior synonym of Glypta extincta Ratzeburg (Aubert, 1978ft: 60).

Gfypta (Glypta) nigroplica, 1889: 1326, 1341. Lectotype 9, SWEDEN: Skane, Wittsjo [= Vittsjo] (UZI,
Lund), by designation of Aubert, 1978ft: 45.
Labels. Scan [hand] ; nigroplica [Thomson cabinet label].
Identity. Glypta nigroplica Thomson.

Gfypta (Glypta) rufipes, 1889: 1328, 1353. Lectotype 9, SWEDEN: Oland (UZI, Lund), by designation of
Aubert, 1966: 127.

Labels. O. [printed] ; rufipes [Thomson cabinet label].

Although Townes, Momoi & Townes (1965: 212) were almost certainly correct in recognising this as
Type' ( = holotype, the single original specimen) Aubert chose to designate it as lectotype. He presented
no evidence of a syntype series.

Identity. Junior primary homonym of Glypta rufipes Spinola, 1851 and of Glypta rufipes Brischke, 1865.
Replacement name Glypta thomsonii Dalla Torre, 1901: 416. Junior synonym of Glypta similis Bridgman
(Aubert, 1978ft: 52).

Glypta (Glypta) solids, 1889: 1328, 1348. Lectotype 9, GERMANY (UZI, Lund), by designation of Aubert,
1978ft: 50.

Labels. 46 [hand, on red paper] ; Germ. [hand].

Thomson attributed this name to Hartig but it was never published by him.
Identity. Glypta salicis Thomson.

Gfypta (Glypta) scutellaris, 1889: 1327, 1344. Lectotype 9, SWEDEN: Skane, Alnarp (UZI, Lund), by des-
ignation of Townes, Momoi & Townes, 1965: 213.

Labels. ALNARP [printed] ; scutellaris [Thomson cabinet label].

The lectotype was labelled by Hinz, 1962 not Townes, 1965 as stated by Townes, Momoi & Townes
(1965:213).

Identity. Glypta scutellaris Thomson.

Glypta (Glypta) tegularis, 1889: 1325, 1335. Lectotype 9, FRANCE: Pyrenees (UZI, Lund), by designation of
Aubert, 1972: 146.

Labels. Pyren [hand] ; tegularis [Thomson cabinet label].
Identity. Glypta tegularis Thomson.

Gfypta (Glypta) tenuicornis, 1889: 1326, 1340. Lectotype 9, GERMANY (WEST): Munich (UZI, Lund), by
designation of Townes, Momoi & Townes, 1965: 214.
Label. Germ. [hand].
Identity. Glypta tenuicornis Thomson.

Gfypta (Glypta) tenuitarsis, 1889: 1327, 1346. LECTOTYPE 9, SWEDEN: Lappland (UZI, Lund), here
designated (selected by J. F. Aubert).

Labels. Lap [hand] ; tenuitarsis [Thomson cabinet label].

Aubert (1972: 146) published a lectotype designation for 'Glypta tenuiventris Ths. (? recte crassitarsis
Ths. ?)'. Thomson did not describe a species tenuiventris, so Aubert's tenuiventris may have been a lapsus
for tenuitarsis.

Identity. Junior synonym of Glypta crassitarsis Thomson (Aubert, 1978ft: 46).

Gfypta (Glypta) varicoxa, 1889: 1328, 1348. Lectotype 9, SWEDEN: Skane, Ringsion [= Ringsjon] (UZI,
Lund), by designation of Aubert, 1972: 146.
Labels. Scan [printed] ; varicoxa [Thomson cabinet label].
Identity. Glypta varicoxa Thomson.

Gfypta (Glypta) xanthognatha, 1889: 1325, 1335. Lectotype 9, SWEDEN: Skane, Skanor (UZI, Lund), by
designation of Townes, Momoi & Townes, 1965: 214.
Label. Skan [hand].
Identity. Junior synonym of Glypta consimilis Holmgren (Aubert, 1978ft: 59).

Goniocryptus annulicornis, 1896: 2357. Holotype 9, SWEDEN : Skane, Ringsjon (UZI, Lund).
Labels. Ringsio [printed] ; annulicornis [Thomson cabinet label].
Identity. Junior synonym of Trychosis mesocastanus (Tschek) (van Rossem, 1966: 33).

34 M. G. FITTON

Goniocryptus annutttarsis, 1873: 492. Lectotype 9, SWEDEN: Skane, Bogestad [= Bokestad] (UZI, Lund), by
designation of van Rossem, 1966: 12.
Late/. Boks 2 1/6 [hand].
Identity. Junior synonym of Trychosis neglectus (Tschek) (van Rossem, 1966: 10).

Goniocryptus clypearis, 1873: 494. Lectotype 9, SWEDEN: Skane, Degeberga (UZI, Lund), by designation of
van Rossem, 1966: 33.

Labels. Dgb [hand] ; clypearis [Thomson cabinet label].
The lectotype is the upper of two specimens on the same pin.
Identity. Junior synonym of Trychosis legator (Thunberg) (van Rossem, 1966: 24).

Goniocryptus glabriculus, 1873 : 491. Lectotype 9, SWEDEN : Norrland (UZI, Lund), by designation of Aubert,
1966: 128.

Label. Norl. [printed].
Identity. Trychosis glabriculus (Thomson).

Goniocryptus lapponicus, 1894: 2116. Lectotype ?, SWEDEN: Lappland (UZI, Lund), by designation of van
Rossem, 1966:22.

Labels. Lapp [printed] ; lapponicus [Thomson cabinet label].
Identity. Junior synonym of Trychosis pauper (Tschek) (van Rossem, 1966: 20).

Goniocryptus macrourus, 1873: 492. Lectotype ?, DENMARK: Zealand, Strandmellen (ZM, Copenhagen), by
designation of van Rossem, 1966: 38.

Labels. 9 Strandmo Drewsen [hand] ; Danmark ex coll. Schiodte [printed].
Identity. Junior synonym of Trychosis ingratus (Tschek) (van Rossem, 1966: 37).

Goniocryptus nitidulus, 1896: 2359. Holotype 9, SWEDEN : Skane, Degeberga (UZI, Lund).
Labels. Dgb. [hand] ; nitidulus [Thomson cabinet label].
Identity. Junior synonym of Trychosis gradarius (Tschek) (van Rossem, 1966: 16).

Goniocryptus pictus, 1873: 494. Lectotype 9, DENMARK : Jutland, Horsens (ZM, Copenhagen), by designation
of van Rossem, 1966: 32.

Labels. 9 29/5 1870 Horsens O. Jensen [hand] ; Danmark ex coll. Schi0dte [printed].
Identity. Junior synonym of Trychosis legator (Thunberg) (van Rossem, 1966: 24).

Goniocryptus pleur alts, 1896: 2358. Holotype 9, GERMANY (WEST): Bavaria (UZI, Lund).

Labels. 871 [hand]; 21. [hand]; G. tristator for pleuralis m. [hand] [This last is an original label and is
not in Roman's handwriting as suggested by van Rossem (1966: 14)].

Identity. Junior synonym of Trychosis tristator (Tschek) (van Rossem, 1966: 12).

Gonotypa melanostoma, 1887c: 1137. LECTOTYPE 9, SWEDEN: Skane, Lund (UZI, Lund), here designated
(selected by H. K. Townes).
Label. L-d [printed].
Identity. Gonotypus melanostoma (Thomson).

Grypocentrus apicaUs, 1883: 905. Lectotype 9, SWEDEN: Skane, Palsio [= Palsjo] (UZI, Lund), by des-
ignation of Kasparyan, 1973: 207.
Label. Hbg. [hand].
Identity. Grypocentrus apicalis Thomson.

Habrocryptus orbitatorius, 1896: 2364. Holotype 9, YUGOSLAVIA : Dalmatia (UZI, Lund).
Labels. +129 [hand] ; Dalm [hand] ; orbitatorius [hand].
Identity. Ischnus orbitatorius (Thomson).

Habrocryptus punctiger, 1896: 2364. Lectotype 9, SWEDEN: Lappland (UZI, Lund), by designation of
Townes, Momoi & Townes, 1965: 176.
Label. Lpl. [printed].
Identity. Ischnus punctiger (Thomson).

Hadrodactylus albicoxa, 1883: 921. Lectotype 9, SWEDEN: Skane, Torringe (UZI, Lund), by designation of
Idar, 1973:24.

Labels. Sk [printed] ; 9 [printed] ; albicoxa [Thomson cabinet label].
Identity. Junior synonym of Hadrodactylus confusus (Holmgren) (Idar, 1973 : 24).

Hadrodactylus bidentulus, 1883: 919. Lectotype 9, SWEDEN: Skane, Palsjo (UZI, Lund), by designation of
Idar, 1973:24.

THE ICHNEUMONIDAE DESCRIBED BY C. G. THOMSON 35

Label. Pal. [hand].

Identity. Hadrodactylus bidentulus Thomson.

Hadrodactylus genalis, 1883: 921. Lectotype 9, SWEDEN: Skane, Ortofta (UZI, Lund), by designation of Idar
1973:24.

Labels. Ort [hand] ; genalis [Thomson cabinet label].
Identity. Hadrodactylus genalis Thomson.

Hadrodactylus gracilipes, 1883: 920. Lectotype 9, SWEDEN: Skane, Yddinge (UZI, Lund), by designation of
Idar, 1973:24.
Label. Ydd [hand].
Identity. Hadrodactylus gracilipes Thomson.

Hadrodactylus laticeps, 1883: 920. Lectotype 9, SWEDEN: Skane, Ilstorp (UZI, Lund), by designation of Idar
1973:24.

Labels. lisp 28/7 [hand] ; 204. [hand] ; 9 [printed] ; laticeps [Thomson cabinet label].
Identity. Junior synonym of Hadrodactylus tiphae (Geoffroy).

Hadrodactylus nigrifemur, 1883: 920. Lectotype $, SWEDEN: Lappland (UZI, Lund), by designation of Idar
1974:113.

Labels. Lap [hand] ; nigrifemur [Thomson cabinet label].
Identity. Hadrodactylus nigrifemur Thomson.

Hadrodactylus tarsator, 1883: 919. Lectotype 9, SWEDEN: Skane, Yddinge (UZI, Lund), by designation of
Idar, 1973:24.

Label. Yddinge [printed].

Identity. Hadrodactylus tarsator Thomson.

Hadrodactylus villosulus, 1883: 919. Lectotype 9, SWEDEN: Skane, Ryssioholm [= Rossjoholm] (UZI,
Lund), by designation of Idar, 1973: 24.
Labels. Rshm 16/6 [hand]; 9 [printed].
Identity. Hadrodactylus villosulus Thomson.

Hemichneumonfuscipes, 1891 : 1612. Holotype 9, SWEDEN: Oland (UZI, Lund).
Labels. C). [printed] ; fuscipes m [Thomson cabinet label].
Identity. Hemichneumonfuscipes Thomson.

Hemiteles aeneus, 1884: 982. Lectotype 9, NORWAY: Hjerkinn (UZI, Lund), by designation of Horstmann,
1979a:297.

Labels. Jerkin. 3.8.77 [locality, printed ; date, hand] ; 234 [hand] ; aeneus [Thomson cabinet label].
Thomson gave the locality as 'Lappland' so it is not absolutely certain that the lectotype is an original
specimen. Hjerkinn is not really far enough north to be considered Lappland.
Identity. Junior synonym ofGelis glacialis (Holmgren) (Horstmann, 1979a: 297).

Hemiteles albipalpus, 1884: 981. Lectotype 9, SWEDEN: Skane, Ringsjon (UZI, Lund), by designation of
Aubert, 1966: 129.

Labels. Rsio [printed] ; albipalpus [Thomson cabinet label].
Identity. Gelis albipalpus (Thomson) (Horstmann, 1979a: 297).

Hemiteles alpinus, 1884: 997. Lectotype 9, SWEDEN: Jamtland, Areskutan (UZI, Lund), by designation of
Jussila, 1965: 160.

Labels. Are [hand]; Thms [printed].
Identity. Phygadeuon alpinus (Thomson) (Horstmann, 1979a: 297).

Hemiteles apertus, 1884: 990. Lectotype 9, SWEDEN: Oland (UZI, Lund), by designation of Horstmann,
19790:297.
Label. 0 [hand].
Identity. Junior synonym of Gnypet omorpha obscura (Bridgman) (Horstmann, 1979a: 297-298).

Hemiteles arcticus, 1884: 998. Syntypes 9, NORWAY: Skalstugan (lost).
Identity. Phygadeuon arcticus (Thomson) (Horstmann, 1979a: 298).

Hemiteles areolaris, 1884: 986. Syntypes 9, SWEDEN: Skane, Bastad (lost).

The specimen published as lectotype by Horstmann (1979a: 298) is labelled 'B6r'[= Borringe] not 'Ba'
and therefore cannot be a syntype.

Identity. Charitopes areolaris (Thomson) (Horstmann, 1979a: 298, on the basis of the invalid 'lec-
totype').

36 M. G. FITTON

Hemiteles auriculatus, 1884: 977. Lectotype 9, SWEDEN: Skane, Palsjo (UZI, Lund), by designation of
Horstmann, 1972:222.
Label. Pal. [hand].
Identity. Junior synonym of Zoophthorus graculus (Gravenhorst) (Horstmann, 1979a: 298).

Hemiteles australis, 1885: 26. Type(s) 9, FRANCE: Avignon (lost).
Identity. Unknown, the name remains a nomen dubium.

Hemiteles balteatus, 1885: 28. Syntypes 9 c?, FRANCE (lost).

Identity. Gelis balteat us (Thomson) (Horstmann, 1979a: 298).

Hemiteles bellicornis, 18886: 1243. Lectotype 9, SWEDEN: Skane, Palsjo (UZI, Lund), by designation of
Horstmann, 1979a: 298.
Label. Pal [hand].
Identity. Handaoia bellicornis (Thomson) (Horstmann, 1979a: 298).

Hemiteles bidentulus, 1884: 971. Lectotype 9, SWEDEN: Skane, Ofvedskloster [ = Ovedskloster] (UZI,
Lund), by designation of Aubert, 1966: 129.
Labels. Oke A [hand] ; bidentulus [Thomson cabinet label].
Identity. Junior synonym of Isadelphus armatus (Gravenhorst) (Horstmann, 1979a: 298).

Hemiteles breviareolatus, 1884: 995. Lectotype 9, SWEDEN: Skane, Stehag (UZI, Lund), by designation of
Horstmann, 1979a:298.

Labels. Scan sylv [printed] ; breviareolatus [Thomson cabinet label].
Identity. ? Stibeutes breviareolatus (Thomson) (Horstmann, 1979a: 298).

Hemiteles brevicauda, 1884: 984. Syntype 1 <$, SWEDEN: Skane, Loparod (UZI, Lund).

Label. Lop [hand].

Although Loparod has not been located, it is difficult to see how the designation as lectotype of a
specimen from Ringsjon (Horstmann, 1979a: 298) can be justified, especially as an undoubted syntype is
present in the collection. However, there is nothing positive to invalidate the lectotype. Its status must
remain in doubt until Loparod is identified.

Identity. Gelis brevicauda (Thomson).

Hemiteles capra, 1884: 974. Lectotype 9, SWEDEN: Skane, Reften (UZI, Lund), by designation of Aubert,
1972:148.

Labels. Sk [printed] ; Capra [Thomson cabinet label].
Identity. Mastrulus capra (Thomson) (Horstmann, 1979a: 298).

Hemiteles capreolus, 1884: 970. Lectotype 9, SWEDEN: Skane (UZI, Lund), by designation of Horstmann,
1972:220.

Label. Scan [printed].
Identity. Junior synonym of Acrolyta rufocincta (Gravenhorst) (Horstmann, 1979a : 298).

Hemiteles clausus, 18886: 1245. Holotype 9, SWEDEN : Skane, Ortofta (UZI, Lund).
Labels. Ort IX/86 [hand] ; n sp [hand].
Identity. Charitopes clausus (Thomson) (Horstmann, 1979a: 298).

Hemiteles constrictus, 1884: 997. Lectotype 9, SWEDEN: Skane, Torekov (UZI, Lund), by designation of
Horstmann, 1974c: 344.
Label. Trkv [hand].
Identity. Xiphulcus constrictus (Thomson) (Horstmann, 1979a: 298).

Hemiteles costalis, 1884: 984. Lectotype 9, SWEDEN: Skane, Lund, (UZI, Lund), by designation of Horst-
mann, 1979a: 298.

Labels. L-d [printed] ; costalis [Thomson cabinet label].
Identity. Mastrus costalis (Thomson) (Horstmann, 1979a: 298).

Hemiteles cyclogaster, 1884: 992. Type(s) 9, SWEDEN: Skane, Ryssioholm [ = Rossjoholm] (lost).
Identity. Pleurogyrus cyclogaster (Thomson) (Horstmann, 1979a: 298).

Hemiteles cynipinus, 1884: 977. Lectotype 9, SWEDEN: Skane (UZI, Lund), by designation of Aubert, 1966:
129.

Labels. Scan [printed] ; Cynipinus [Thomson cabinet label].
Identity. Zoophthorus cynipinus (Thomson) (Horstmann, 1979a: 298).

THE ICHNEUMONIDAE DESCRIBED BY C. G. THOMSON 37

Hemiteles dispar, 1885: 28. Syntypes 9 J, FRANCE: Libercourt (lost).

Identity. Junior primary homonym of Hemiteles dispar Ratzeburg. Replacement name Hemiteles thom-
soni Schmiedeknecht. The identity of the species is unknown and the name remains a nomen dubium.

Hemiteles distans, 1884: 978. Holotype 9, SWEDEN: Skane, Klinta (UZI, Lund).
Labels, [small green square] ; distans [Thomson cabinet label].
Identity. Clypeoteles distans (Thomson) (Hofstmann, 1979a: 298).

Hemiteles elymi, 1884: 981. Lectotype $, SWEDEN: Skane, Skanor (UZI, Lund), by designation of Horst-
mann, 1979a: 299.

Labels. Snor [hand]; Elymi [Thomson cabinet label].
Identity. Gelis elymi (Thomson) (Horstmann, 1979a: 299).

Hemiteles falcatus, 1884: 999. Lectotype 9, SWEDEN: Skane, Fogelsang [= Fagalsang] (UZI, Lund), by
designation of Horstmann, 1976a: 24.
Label. Fsg [hand].
Identity. Tropistesfalcat us (Thomson) (Horstmann, 19790:299).

Hemiteles fasciatus, 1884: 995. Lectotype 9, SWEDEN: Skane, Stehag (UZI, Lund), by designation of Horst-
mann, 1979a: 299.

Label, [small green square].

Identity. Junior primary homonym of Hemiteles fasciatus Heer. Replacement name Theroscopus fas-
ciatulus Horstmann, 1979a: 299.

Hemiteles fumipennis, 1884: 984. Holotype 9, SWEDEN : Skane, Lund (UZI, Lund).
Labels. Lund [printed] ; fumipennis [Thomson cabinet label].
Identity. Mastrus fumipennis (Thomson) (Horstmann, 1979a: 299).

Hemiteles fuscicarpus, 1885: 29. Type(s) 9, FRANCE: Libercourt (lost).
Identity. Unknown, the name remains a nomen dubium.

Hemiteles geniculatus, 1884: 989. Lectotype 9, SWEDEN: Skane, Klinta (UZI, Lund), by designation of
Aubert, 1966: 129.

Labels, [small green square] ; 7 [hand] ; geniculatus [Thomson cabinet label].
Identity. Junior synonym of Dichrogaster aestivalis (Gravenhorst) (Horstmann, 1979a: 299).

Hemiteles gibbifrons, 1884: 980. Holotype 9, SWEDEN : Smaland (UZI, Lund).
Labels. Col Ljgh [printed] ; gibbifrons [Thomson cabinet label].
Identity. Gelis gibbifrons (Thomson) (Horstmann, 1979a: 299).

Hemiteles glyptonotus, 1885: 32. Type(s) 9, FRANCE (lost).

Identity. Junior synonym ofChirotica maculipennis (Gravenhorst) (Horstmann, 1979a: 299).

Hemiteles gracilipes, 1884: 992. Lectotype 9, SWEDEN: Skane, Ryssioholm [= Rossjoholm] (UZI, Lund), by
designation of Aubert, 1966: 129.

Labels. Rshm 16/6 [hand] ; gracilipes [Thomson cabinet label].
Identity. Oecotelma gracilipes (Thomson) (Horstmann, 1979a: 299).

Hemiteles gracilis, 1884: 989. Lectotype 9, SWEDEN: Skane, Lund (UZI, Lund), by designation of Townes,
Momoi & Townes, 1965: 138.
Label. Lund [printed].
Identity. Aclast us gracilis (Thomson) (Horstmann, 1979a: 299).

Hemiteles hadrocerus, 1884: 991. Lectotype 9, SWEDEN: Skane, Esperod [= Asperod] (UZI, Lund), by
designation of Horstmann, 1979a: 299.
Label. Esp [hand].
Identity. Orthizema hadrocerum (Thomson) (Horstmann, 1979a: 299).

Hemiteles hirticeps, 1885: 27. Type(s) 9, FRANCE: Pyrenees (lost).

Identity. Zoophthorus hirticeps (Thomson) (Horstmann, 1979a: 299).

Hemiteles homocerus, 1885 : 29. Syntypes 9 c?, FRANCE: Libercourt (lost).

Identity. ? Junior synonym ofSulcarius biannulatus (Gravenhorst) (Horstmann, 1979a: 299).

Hemiteles inflates, 1884: 992. Lectotype 9, SWEDEN: Skane, Lund (UZI, Lund), by designation of Aubert,
1966: 129.

Label. Lund [printed].
Identity. Platyrhabdus inflat us (Thomson) (Horstmann, 1979a: 299).

38 M. G. FITTON

Hemiteles infumatus, 1884: 983. Lectotype 9, SWEDEN: Skane, Lund, Raby (UZI, Lund), by designation of
Horstmann, 1979a: 299.

Labels. Rab 1/7 [hand] ; infumatus [Thomson cabinet label].
Identity. Gelis infumatus (Thomson) (Horstmann, 1979a: 299).

Hemiteles ischnocerus, 18886: 1246. Lectotype 9, SWEDEN: Skane, Ortofta (UZI, Lund), by designation of
Horstmann, 1976a: 23.
Label. Ort. 2/VI [hand].
Identity. Tricholinum ischnocerum (Thomson) (Horstmann, 1979a: 299).

Hemiteles liambus, 1885 : 25. Type(s) 9, FRANCE: Avignon (lost).
Identity. Unknown, the name remains a nomen dubium.

Hemiteles liostylus, 1885: 30. Syntypes 9 <3, FRANCE: Libercourt (lost).
Identity. Dichrogaster liostylus (Thomson) (Horstmann, 1979a: 299).

Hemiteles lissonotoides, 1885: 30. Lectotype 9, SWEDEN: Skane, Palsjo (UZI, Lund), by designation of
Horstmann, 1979a: 299.

Labels. Palsio [printed] ; Lissonotoides m [hand].
Identity. Junior synonym oiAteleute linearis Foerster (Horstmann, 1979a: 299).

Hemiteles longicauda, 1884: 980. Lectotype 9, SWEDEN: Skane, Palsio [Palsjo] (UZI, Lund), by designation
of Aubert, 1966: 129.
Label. Palsio [printed].
Identity. Gelis longicauda (Thomson) (Horstmann, 1979a: 299).

Hemiteles longicaudatus, 1884: 989. Holotype 9, SWEDEN : Smaland (UZI, Lund).
Label. Sm [hand].
Identity. Dichrogaster longicaudatus (Thomson) (Horstmann, 1979a: 300).

Hemiteles longulus, 1884: 997. Lectotype 9, SWEDEN: Skane, Lund (UZI, Lund), by designation of Aubert,
1966! 129.

Label Lund [printed].
Identity. Junior synonym of Xiphulcusfloricolator (Gravenhorst) (Horstmann, 1979a: 300).

Hemiteles macrurus, 1884: 985. Lectotype 9, SWEDEN: Skane, Bastad (UZI, Lund), by designation of Aubert,
1966: 129.
Label. Bast [hand].
Identity. Charitopes macrurus (Thomson) (Horstmann, 1979a: 300).

Hemiteles magnicornis, 1884:994. Lectotype cJ, SWEDEN: Skane, Yddingesion [= Yddingesjon] (UZI,
Lund), by designation of Horstmann, 1979a: 300.
Labels. Yd. [hand] ; magnicornis [Thomson cabinet label].
Identity. Phygadeuon magnicornis (Thomson) (Horstmann, 1979a: 300).

Hemiteles melanogaster, 1884. 982. Lectotype 9, SWEDEN: Skane, Klinta (UZI, Lund), by designation of
Jussila, 1965: 155.
Label. Scan [printed].
Identity. Gelis melanogaster (Thomson) (Horstmann, 1979a: 300).

Hemiteles microstomus, 1884: 969. Lectotype 9, SWEDEN: Skane, Ryssioholm [= Rossjoholm] (UZI, Lund),
by designation of Jussila, 1965: 152.
Labels. Rhm [hand] ; microstomus.
Identity. Zoophthorus microstomus (Thomson) (Horstmann, 1979a: 300).

Hemiteles monodon, 1884: 991. Lectotype 9, SWEDEN: Skane, Yddingesjon (UZI, Lund), by designation of
Aubert, 1966: 129.

Labels, [small pale blue-green square] ; monodon [Thomson cabinet label].

There is no evidence that the blue-green square indicates that the specimen was collected at Yddinges-
jon, and if it is later shown to indicate another locality then the lectotype designation will be invalid.

I do not consider the earlier reference~by Aubert (1964: 154) to be a valid lectotype designation because
he gave no indication of which specimen was selected.

Identity. Platyrhabdus monodon (Thomson) (Horstmann, 1979a: 300).

Hemiteles nigricornis, 1884: 987. Lectotype 9, SWEDEN: Lappland (UZI, Lund), by designation of Horst-
mann, 1979a: 300.

Labels. Lpl. [printed] ; nigricornis [Thomson cabinet label].
Identity. Sulcarius nigricornis (Thomson) (Horstmann, 1979a: 300).

THE ICHNEUMONIDAE DESCRIBED BY C. G. THOMSON 39

Hemiteles nigriventris, 1884: 975. Lectotype 9, SWEDEN: Skane, Vittsjo (UZI, Lund), by designation of
Townes, Momoi & Townes, 1965: 129.

Labels. Witt 10/6 [hand] ; nigriventris [Thomson cabinet label].

Identity. Junior synonym oflsadelphus gallicola (Bridgman) (Horstmann, 1979a: 300).

Hemiteles notaticrus, 18886: 1244. Lectotype 9, SWEDEN: Skane, Palsjo (UZI, Lund), by designation of
Horstmann, 1979a: 300.

Labels. Pal [hand] ; n. sp [hand] ; notaticrus [hand].
Identity. Zoophthorus notaticrus (Thomson) (Horstmann, 1979a: 300).

Hemiteles obliquus, 1885: 24. Syntypes 9 $, FRANCS (lost).
Identity. Unknown, the name remains a nomen dubium.

Hemiteles obscuripes, 1884: 976. Lectotype 9, SWEDEN: Skane, Palsio [= Palsjo] (UZI, Lund), by des-
ignation of Townes, Momoi & Townes, 1965: 130.
Labels. Pal. [hand] ; obscuripes [Thomson cabinet label].
Identity. Junior synonym oflsadelphus inimicus (Gravenhorst) (Horstmann, 1979a: 300).

Hemiteles opaculus, 1884: 975. Lectotype 9, SWEDEN: Skane, Esperod [= Asperod] (UZI, Lund), by des-
ignation of Aubert, 1966: 129.

Label. Col Zet [printed].

Horstmann (1976a: 29) casts some doubt on Aubert's designation of this specimen as lectotype. How-
ever, in the absence of specimens labelled with the type-locality and in the absence of evidence that the
Zetterstedt specimen did not come from Asperod it can be accepted as an original specimen.

Identity. Diaglyptellana opacula (Thomson) (Horstmann, 1979a: 300).

Hemiteles ornatulus, 1884: 980. Holotype 9, SWEDEN: Skane, Kaseberga (UZI, Lund).
Labels. Kas [hand] ; ornatulus [Thomson cabinet label].
Identity. Gelis ornatulus (Thomson) (Horstmann, 1979a: 300).

Hemiteles pallicarpus, 1884: 970. Lectotype 9, SWEDEN: Skane, Reften (UZI, Lund), by designation of
Aubert, 1961:197.
Label. Rfn 12/7 [hand].
Identity. Junior synonym of Eudelus simillimus (Taschenberg) (Horstmann, 1979a: 300).

Hemiteles plumbeus, 1884: 979. Holotype 9, SWEDEN : 'Halland' [Skane], Margretetorp (UZI, Lund).
Label. Marg.

Margretetorp is in northern Skane, not southern Halland as stated by Thomson. It is near the bound-
ary between the two provinces, so Thomson's error is easily explained.

Identity. Zoophthorus plumbeus (Thomson) (Horstmann, 1979a: 300).

Hemiteles punctiventris, 1884: 977. Lectotype <J, SWEDEN: Skane, Bokeberg (UZI, Lund), by designation of
Horstmann, 1979a: 300.
Label. Bok [hand].
Identity. Zoophthorus punctiventris (Thomson) (Horstmann, 1979a: 300).

Hemiteles rubricollis, 1884: 979. Holotype 9, SWEDEN : Skane, Stehag(UZI, Lund).

Labels. Stehag 28/V 82 in truncus Quercus [hand] ; rubricollis [Thomson cabinet label].

Identity. Gelis rubricollis (Thomson) (Horstmann, 1979a: 300-301).

Hemiteles rubripes, 1884: 976. Lectotype 9, SWEDEN: Lappland (UZI, Lund), by designation of Horstmann,
1979a: 301.

Labels. Lpl. [printed] ; rubripes [Thomson cabinet label].

Identity. Junior synonym oflsadelphus inimicus (Gravenhorst) (Horstmann, 1979a: 301).

Hemiteles rubrotinctus, 1885: 31. Type(s) 9, FRANCE: Avignon (lost).

Identity. Chirotica rubrotincta (Thomson) (Horstmann, 1979a: 301).

Hemiteles rufulus, 1884: 972. Lectotype 9, SWEDEN: Skane, Lund or Palsio [= Palsjo] (UZI, Lund), by
designation of Aubert, 1966: 129.
Label. Scan [printed].
Identity. Mastrus rufulus (Thomson) (Horstmann, 1979a: 301).

Hemiteles rugifer, 1884: 983. Type(s) 9, SWEDEN : Norrland (lost).

The specimen recognised as 'holotype' by Horstmann (1979a: 301) is from Lillehammer in Norway and
therefore cannot be the type. It is labelled 'Norv' and 'Lhmr 24/6 77'. There are no other specimens in the
collection.

Identity. Gelis rugifer (Thomson) (Horstmann, 1979a: 301, on the basis of the invalid 'holotype').

40 M. G. FITTON

Hemiteles rugifrons, 1884: 978. Lectotype 9, SWEDEN: Skane, Stehag (UZI,Lund), by designation of Horst-
mann, 1979a: 301.

Labels. Rsio [printed] ; rugifrons [Thomson cabinet label].

Identity. Junior synonym of Clypeoteles distans (Thomson) (Horstmann, 1979a: 301).

Hemiteles scabriculus, 1884: 969. Lectotype 9, SWEDEN: Skane, Holmeja (UZI, Lund), by designation of
Horstmann, 1979a: 301.

Labels. Yd [hand] ; scabriculus [Thomson cabinet label].

There is a paralectotype male (? not conspecific) on the same pin, and above, the lectotype.
Identity. Junior synonym ofEudelus simillimus (Taschenberg) (Horstmann, 1979a: 301).

Hemiteles solutus, 1884: 990. Lectotype (J, SWEDEN: Skane, Ortofta (UZI, Lund), by designation of Aubert,
1966: 129.

Label. Ortofta [printed].
Identity. Aclastus solutus (Thomson) (Horstmann, 1979a: 301).

Hemiteles stagnalis, 1884: 987. Lectotype 9, SWEDEN: Skane, Ringsjon (UZI, Lund), by designation of
Horstmann, 1976a: 26.

Labels. Rsio [printed] ; stagnalis [Thomson cabinet label].

The lectotype is on the same pin as four other specimens (1 $, 3 cJ); it is the second from the top.
Identity. Junior synonym ofAgasthenes varitarsus (Gravenhorst) (Horstmann, 1979a: 301).

Hemiteles triannulatus, 1884: 991. Lectotype 9, SWEDEN: Skane, Holmeja (UZI, Lund), by designation of
Aubert, 1968:195.
Label. Yddinge [printed].
Identity. Orthizema triannulatum (Thomson) (Horstmann, 1979a: 301).

Hemiteles trochanteratus, 1884: 994. Syntype 1 J, SWEDEN : Skane, Ortofta (UZI, Lund).

Label. Ortofta [printed].

Despite Horstmann's view (1979a: 301) this male specimen cannot be excluded from Thomson's type-
series and must be a syntype. Thomson expressly included the male (Code, Article 72(b)) even though his
description may not apply to it.

Identity. Theroscopus trochanteratus (Thomson).

Hemiteles trochanteratus, 1885: 26. ? Syntype 1 3, FRANCE (UZI, Lund).

Label. Gallia [hand].

It is possible, perhaps probable, that this is a specimen sent by Lethierry to Thomson after 1885 and
therefore not a type.

Identity. Junior primary homonym of Hemiteles trochanteratus Thomson, 1884. Replacement name
Hemiteles trochanteralis Dalla Torre, 1902: 668. ? Theroscopus trochanteralis (Dalla Torre) comb. n.

Hemiteles ungularis, 1884: 994. Lectotype 9, SWEDEN: Skane, Ortofta (UZI, Lund), by designation of Horst-
mann, 1979a: 301.

Labels. Ort. [hand] ; ungularis [Thomson cabinet label].
Identity. Theroscopus ungularis (Thomson) (Horstmann, 1979a: 301).

Hemiteles unicolor, 1884: 974. Lectotype 9, SWEDEN: Oland (UZI, Lund), by designation of Townes, Momoi
&Townes, 1965: 137.

Labels. 0. [printed] ; unicolor [Thomson cabinet label].
The lectotype is the lower of two specimens on one pin.
Identity. Junior synonym of Hemiteles similis (Gmelin) (Horstmann, 1979a: 301).

Hemiteles validicornis, 1884: 995. Lectotype 9, SWEDEN: Norrland (UZI, Lund), by designation of Aubert,
1972: 148.

Labels. Norl. [printed] ; validicornis [Thomson cabinet label].
Identity. Junior synonym of Theroscopus melanopygus (Gravenhorst) (Horstmann, 1979a: 301).

Hodostatus brevis, 1883: 929. LECTOTYPE 9, SWEDEN: Skane, Lund (UZI, Lund), here designated (selec-
ted by H. K. Townes).

Labels. LD 4/8. ['LD' printed, date hand] ; brevis [Thomson cabinet label].
Identity. Hodostates brevis (Thomson).

Holocremna annulitarsis, 1887c: 1 179. Syntypes 1 9, 1 £, 2 ? sex, SWEDEN : Skane, Ringsjon (UZI, Lund).
Labels, [small green square] (1 <$, 2 ? sex), [small green square] ; 9 [printed] (1 9).
Aubert 's publication of a neotype (1966: 131) for this species is, fortunately, not valid because it does

THE ICHNEUMONIDAE DESCRIBED BY C. G. THOMSON 41

not comply with the provisions of Article 75(c) of the Code. The discovery of syntypes does not, therefore,
need a reference to the Commission (Article 75(/)).
Identity. Olesicampe annulitarsis (Thomson) comb. n.

Holocremna bergmanni, 1887c: 1182. Lectotype 9, SWEDEN: Skane, Lund (UZI, Lund), by designation of
Aubert, 1972: 149.

Labels. Lund [printed] ; Bergmani [Thomson cabinet label].
Identity. Olesicampe bergmanni (Thomson) comb. n.

Holocremna buccata, 1887c: 1180. LECTOTYPE 9, SWEDEN: Skane, Palsjo (UZI, Lund), here designated
(selected by R. Hinz).

Labels. Hbg [hand]; 9 [printed].

Identity. Olesicampe buccata (Thomson) (det. R. Hinz) comb. n.

Holocremna curtigena, 1887c: 1179. Lectotype 9, SWEDEN: Skane, Palsjo (UZI, Lund), by designation of
Aubert, 1966: 131.

Labels. Pal. [hand] ; curtigena [Thomson cabinet label].
The lectotype is the upper of two specimens on the same pin.
Identity. Olesicampe curtigena (Thomson) comb. n.

Holocremna frutetorum, 1887c: 1178. Lectotype 9, GERMANY (EAST): Dresden (UZI, Lund), by designation
of Townes, Momoi & Townes, 1965: 302.
Labels. Dresden [hand] ; frutetorum m [hand].
Identity. Olesicampe frutetorum (Thomson).

Holocremna heterogaster, 1887c: 1 178. Lectotype 9, GERMANY (UZI, Lund), by designation of Aubert, 1972:
150.

Label, f. 3. 84. [hand].
Identity. Olesicampe heterogaster (Thomson) comb. n.

Holocremna melanogaster, 1887c: 1181. LECTOTYPE 9, SWEDEN: Skane; Palsjo (UZI, Lund), here des-
ignated (selected by R. Hinz).

Labels. Hbg [hand] ; 9 [printed] ; melanogaster [Thomson cabinet label].
Identity. Olesicampe melanogaster (Thomson) comb. n.

Holocremna sinuata, 1 887c : 11 80. Type(s) 9, SWEDEN : Skane, Ringsjon (lost).

The specimen labelled and published by Aubert (1966: 131) as lectotype is fromOrtofta [label 'Ort'],
not Ringsjon, and is not a type. The lectotype designation is therefore invalid.

Identity. Olesicampe sinuata (Thomson) comb. n. (on the basis of material in the collection).

Holocremna spireae, 1887c: 1 182. Holotype 9, GERMANY (WEST): Munich (UZI, Lund).
Labels. 1. [hand] ; Spireae [Thomson cabinet label].
Identity. Olesicampe spireae (Thomson) comb. n.

Holocremna tarsator, 1887c: 1 180. LECTOTYPE 9, GERMANY (UZI, Lund), here designated (selected by R.
Hinz).
No labels.
Identity. Olesicampe tarsator (Thomson) comb. n.

Homoporus brevicornis, 1890: 1507. Syntype 1 9, FRANCE: Lille (UZI, Lund).

Labels. St. Germ, de Princay. [hand] ; brevicornis m (Thomson cabinet label].

Identity. Junior synonym of Syrphoctonus crassicornis (Thomson). Thomson published the two names
Homoporus crassicornis (see below) and H. brevicornis simultaneously for the same species. The former
name was used in the key (page 1490) and the latter with the description (page 1507). Stelfox (1941 : 117)
made a first reviser choice between the two names. The female noted above is the only surviving syntype
of both nominal species.

Homoporus brevitarsis, 1890: 1489, 1495. Lectotype $ [not 9 as stated by Aubert, 1966: 128], SWITZERLAND:
Chur (UZI, Lund), by designation of Aubert, 1966: 128.
Labels. Chur [printed] ; Suisse. [hand].
Identity. Daschia brevitarsis (Thomson).

Homoporus caudatus, 1890: 1490, 1499. Syntypes 59, Id1, SWEDEN: Skane, Ringsjon and FRANCE: near Lille
(UZI, Lund).

Labels, [small green square]; 9 [printed]; caudatus m. [Thomson cabinet label] (1 9)- Phalempin

42 M. G. FITTON

[hand] (1 ?). Libercourt. [hand]; Gall [hand] (2 ?). Fives, [hand, ?some letters] ; Gall [hand] (1 ?). Lille,
[hand]; Gall [hand] (1 rf).
Identity. Campocraspedon caudatus (Thomson).

Homoporus crassicornis, 1890: 1490. Syntype 1 9, FRANCE: Lille (UZI, Lund).

Labels. St. Germ, de Princay. [hand] ; brevicornis m [Thomson cabinet label].

Identity. Syrphoctonus crassicornis (Thomson) comb. n. See notes under Homoporus brevicornis above.

Homoporus crassicrus, 1890: 1491, 1516. Lectotype ?, SWEDEN: Oland (UZI, Lund), by designation of
Jussila, 1966: 319.

Labels. O. [printed] ; crassicrus m [Thomson cabinet label].

The lectotype is on the same pin as another specimen (a tryphonine).

Identity. Syrphoctonus crassicrus (Thomson).

Homoporus hygrobius, 1890: 1491, 1524. Syntypes 6 9, 5 & SWEDEN: Skane (UZI, Lund).

Labels. Lund [printed]; hygrobius m [Thomson cabinet label] (1 9)- Lund [printed] (1 $ 1 J on one
pin). Ort. [hand] (3 rf). Ld [hand] (1 9). [green square] (1 9). Bor [hand]; 9 [printed] (1 9). Bok 9/76
[hand] (1 9). Bo [illegible letters] 7/88 [hand] (1 rf).

Identity. Junior synonym of Syrphoctonus signatus (Gravenhorst) (Carlson, 1979: 718).

Homoporus incisus, 1890: 1511. Holotype 9, SWEDEN: Skane, Ringsjon (UZI, Lund).
Labels. Ringsio [printed] ; incisus m [Thomson cabinet label].
Identity. Syrphoctonus incisus (Thomson) comb. n.

Homoporus longiventris, 1890: 1491, 1514. Lectotype 9, SWEDEN : Skane, Palsjo (UZI, Lund), by designation
of Townes, Momoi & Townes, 1965: 404.
Labels. Pal. [hand] ; longiventris m [Thomson cabinet label].
Identity. Syrphoctonus longiventris (Thomson).

Homoporus megaspis, 1890: 1491, 1516. Holotype 9, GERMANY (WEST): Bavaria (ZSBS, Munich).

Labels. Rsh. Hst. 11.9.73. A.Krchb. [hand]; 1 185. [hand]; ex. typ. [hand]; B. megaspism 9 [hand].
Identity. Syrphoctonus megaspis (Thomson) comb. n.

Homoporus nigricornis, 1890: 1490, 1506. Holotype 9, SWEDEN: Skane, Palsjo(UZI, Lund).
Labels. Hbg. [hand] ; nigricornis n [Thomson cabinet label].
Identity. Enizemum nigricornis (Thomson).

Homoporus punctiventris, 1890: 1490, 1500. Type(s)9, DENMARK : Zealand, Strandm011en (lost).

In the collection of the Zoological Museum, Copenhagen there is a label 'Punctiventris Thoms. Origin.
Type $ 9' but there are no specimens which could be types. Also, there are no specimens from the
type-locality in the Thomson collection despite the indication by Dasch (1964: 267) and the statement by
Beirne (1941 : 683) that Perkins had compared a specimen with the type.

Identity. Sussaba punctiventris (Thomson) (on the basis of material in the Thomson collection).

Homoporus xanthaspis, 1890: 1491, 1518. Syntypes 19, 1 c?» DENMARK: Zealand, Strandmellen (ZM,
Copenhagen).

Labels. 9 Strandm. Drewsen [hand] ; Danmark ex coll. Schi0dte [printed] (9). 3 Strandm. Drewsen
[hand] ; Danmark ex coll. Schi0dte [printed] (<J).

Identity. Phthorima xanthaspis (Thomson) comb. n.

Hoplocryptus binotatulus [as 2-notatulus~], 1873: 512. LECTOTYPE 9, SWEDEN: Smaland (UZI, Lund), here
designated (selected by H. K. Townes).
Labels, [pink square]; Smol [printed].
Identity. Junior synonym of Aritranisfugitivus (Gravenhorst) (Townes & Townes, 1962: 99).

Hoplocryptus elegans, 1873: 511. Syntypes 1 9, 1 <£, SWEDEN: Skane, Reften (UZI, Lund).

Labels. Rfn 5/7 [hand] (9). Rfn 7/7 [hand] (£).

Thomson (1896: 2371) synonymised this species with Hoplocryptus confector (Gravenhorst) and the
syntypes stand in his collection under the name confector in a series beginning with a specimen bearing
Thomson's cabinet label 'elegans'.

Identity. Aritranis elegans (Thomson) comb. n.

Hoplocryptus (Aritranis) graefei [as grafef], 1896: 2373. Syntypes 2 9, 1 cJ, ITALY: Trieste (UZI, Lund).

Labels. Aiis Rubus St. Triest. ['Triest.' printed, remainder hand] ; Hoplocryptus graefei typer [hand]
(1 9)- 2. VI Triest. Zaiile Wiesen. ['Triest.' printed, remainder hand];(l 9)- Aiis Rubus Stengel gezogen.
Triest. ['Triest.' printed, remainder hand] (1 <$).
Identity. Aritranis graefei (Thomson) comb. n.

THE ICHNEUMONIDAE DESCRIBED BY C. G. THOMSON 43

Hoplocryptus mesoxanthus, 1873 : 509. Syntypes 3 9, 1 c?, SWEDEN: Skane, Oland and Smaland (UZI, Lund).
Labels. 1190 [hand]; Scan [printed]; mesoxanthus [Thomson cabinet label] (1 9)- Col Ljgh [printed]
(2 9)- [small dark blue square] (1 <£).

Identity. Aritranis mesoxanthus (Thomson) comb. n.

Hoplocryptus pulcher, 1873: 509. Syntypes 3 ?, 6 <& SWEDEN: Skane, Lund and Ilstorp (UZI, Lund).

Labels. lisp 13/8 [hand] ; pulcher [Thomson cabinet label] (1 9). lisp 13/7 [hand](l 9). lisp 7/8 [hand]
(1 9). Hsp 18/7 [hand] (1 rf). lisp 5/7 [hand] (1 <J). lisp 30/7(1 <?), Ld [hand] (3 rf).
Identity. Aritranis pulcher (Thomson) comb. n.

Hygrocryptus (Hygrocryptus) brevispina, 1896: 2377. Holotype <3, ITALY: Trieste (UZI, Lund).
Labels. 6. VIII Triest. ['Triest.' printed, date hand] ; brevispina m [Thomson cabinet label].
Identity. Thrybius brevispina (Thomson) comb. n.

Hygrocryptus drewseni, 1873 : 514. Syntypes 4 9, 3 (J, DENMARK: Zealand, Leerso (ZM, Copenhagen).

Labels. 9 7/1870 Lerso Drewsen [hand]; Danmark ex coll. Schiodte [printed] (1 9). Danmark ex coll.
Schiodte [printed] (392 £). £ 7/1870 Lerso Drewsen [hand] ; Danmark ex coll. Schiedte [printed] (1 <J).
Identity. Junior synonym of Thrybius leucopygus (Gravenhorst) (Kerrich, 1938 : 174).

Hygrocryptus palustris, 1873 : 514. Syntypes 2 9, SWEDEN : Skane, Alnarp (UZI, Lund).

Labels, palustris [Thomson cabinet label] (1 9). Alnp 8/57 [hand] (1 9).

In addition to the two syntypes in Lund there are six females in the collection of the Zoological
Museum, Copenhagen; some or all of which might be Danish specimens (syntypes) mentioned by Thom-
son. However, because of the existence of syntypes in Lund and some doubt that the Danish specimens
are the ones mentioned by Thomson they are not positively included in the syntype series here recognised.

Identity. Gambrus palustris (Thomson) comb. n.

Ichcunwn (Ichneumon) acuticornis, 1896: 2396. Holotype 9, SWEDEN : Goteborg (UZI, Lund).
Labels, [blue square]; Gbg. [hand] ; acuticornis m [Thomson cabinet label].
Identity. Ichneumon acuticornis Thomson.

Ichneumon aequicalcar, 1888b: 1231. Holotype 9, SWEDEN: Jemtland [= Jamtland], Areskutan (UZI, Lund).
Labels. Lpl. [printed] ; Ths [printed] ; aequicalcar [Thomson cabinet label].
Identity. Ichneumon aequicalcar Thomson.

Ichneumon (Cratichneumon) albiscuta, 1893: 1946. Syntypes 1 $, 2 c?, SWEDEN: Gottland [= Gotland] and
FRANCE (UZI, Lund),

Labels. Gotl. [hand]; Col. Hgn. [printed]; albiscuta [Thomson cabinet label] (1 9)- Gotl. [hand]; Col.
Hgn. [printed] (1 £). 2. [hand] ; Gall [hand] ; 6-armillatus Krchb [hand] (1 rf).

Identity. Cratichneumon albiscuta (Thomson).

Ichneumon (Ichneumon) anospilus, 18866: 15. Syntype 19,? syntype 1 9, SWEDEN: Smaland and [?] Skane
(UZI, Lund).

Labels. Smoland [printed]; anospilus [Thomson cabinet label] (syntype). Scan [handL; Col. Hgn.
[printed] (? syntype).

Thomson gave only 'Suecia australis' as the locality in the original description but later (1893: 191 1) he
stated 'Funnen i Smaland'. The Skane specimen may therefore postdate 1893 and if so cannot be a
syntype.

Identity. Coelichneumon anospilus (Thomson) comb. n.

Ichneumon (Cratichneumon) anotylus, 1896: 2403. Holotype 9, SWEDEN: Skane (UZI, Lund).
Label. Scania [hand].
Identity. Cratichneumon anotylus (Thomson) comb. n.

Ichneumon (Ichneumon) arctobius, 1896: 2399. ? Holotype 9, SWEDEN (UZI, Lund).

Labels. Him [printed] ; Rudolphi [hand] ; arctobius m [Thomson cabinet label].

The specimen tentatively regarded as holotype is the only one in the collection under this name. It is a
Rudolphi specimen from the Stockholm area (label 'Him'). Most Rudolphi material comes from Halsin-
gland (part of 'Norrland') and Thomson may have misread 'Him' as 'His'. It is perhaps significant that the
preceding species, in both the collection and the publication, /. monospilus is also from 'Norrland' and its
holotype is a Rudolphi specimen from Halsingland (see below).

Identity. Ichneumon arctobius Thomson.

Ichneumon (Ichneumon) boreellus, 1896: 2396. Syntypes 2 9, SWEDEN : 'Norrland', Halsingland (UZI, Lund).
Labels. 3. [hand]; Norl [printed]; boreellus m [Thomson cabinet label] (1 9). His [printed]; Rui
[hand] [ = Rudolphi] ; No. 5 n sp [hand] (1 9).
Identity. Ichneumon boreellus Thomson.

44 M. G. FITTON

Ichneumon (Ichneumon) brevigena, 18866: 19. Syntypes 2 9, GERMANY (WEST): Birkenfeld (UZI, Lund).

Labels. Birkenfeld [hand]; 4. [hand]; Brevigena [Thomson cabinet label] (1 $). Birkenfeld Tischbein
[hand] (1 ?).
Identity. Junior synonym of Ichneumon inquinatus Wesmael (Perkins, 1953 : 113).

Ichneumon (Ichneumon) captorius, 1887a: 7. Syntypes [? number, see notes below] 9 c?, SWEDEN (UZI,
Lund).

The series of specimens representing this species in the collection has been combined with that of /.
xanthognathus (see below). The labels 'captorius' and 'xanthognathus' are placed together and 12 $ and 1 1
cJ represent both species. One of the females is lectotype of /. xanthognathus. A lectotype for /. captorius
needs to be selected carefully from the remaining specimens, which are all eligible in terms of type-locality.
It is not considered worthwhile to attempt to delimit the syntype series of captorius. Perkins' (1953: 114)
mention of 9 $ and 2 <$ syntypes may have been the result of such an attempt but this is not clear.

Identity. Junior synonym of Ichneumon minutorius Desvignes (Perkins, 1953: 113).

Ichneumon (Ichneumon) chrysostomus, 1896: 2400. Holotype $, SWEDEN: Jemtland [= Jamtland] (UZI,
Lund).

Labels. Jtl [printed] ; No 5 [hand] ; chrysostomus m [Thomson cabinet label].
Identity. Ichneumon chrysostomus Thomson.

Ichneumon (Coelichneumon) coactus, 1893: 1908. Syntypes 1 $, 1 cJ, SWEDEN: Skane, Ringsjon (UZI, Lund).
Labels. Ringsio [printed] ; breviscuta m [Thomson cabinet label] (9). Ringsio Uprinted] (cJ).
Identity. Coelichneumon coactus (Thomson).

Ichneumon corfitzi, 1890: 1530. LECTOTYPE 9, SWEDEN: Skane, Stehag (UZI, Lund), here designated
(selected by R. Hinz).

Label. Shg. Haslsm vii.88 CM [hand].
Identity. Ichneumon corfitzi Thomson.

Ichneumon (Ichneumon) crassifemur, 18866: 18. Lectotype 9, GERMANY (WEST): Aachen (UZI, Lund), by
designation of Townes, Momoi & Townes, 1965: 463.

Labels. 10/496 ['10' hand, '496' printed] ; Aachen [printed].
Identity. Ichneumon crassifemur Thomson.

Ichneumon (Ichneumon) crassitarsis, 1893: 1925. LECTOTYPE 9, SWEDEN: Skane, Ringsjon (UZI, Lund),
here designated (selected by R. Hinz).
Label. Ringsio [printed].
Identity. Ichneumon crassitarsis Thomson.

Ichneumon (Ichneumon) decrescens, 18866: 13. Syntypes 1 9, 2 $, SWEDEN: Skane and Kalmar (UZI, Lund).
Labels. Calmar [printed]; decrescens [Thomson cabinet label] (1 9)- Scania [printed] (1 ^). Col. Hgn.
[printed] (1 rf).
Identity. Coelichneumon decrescens (Thomson).

Ichneumon eurycerus, 1890: 1528. Lectotype 9, SWEDEN: Skane, Stehag (UZI, Lund), by designation of
Townes, Momoi & Townes, 1965 : 465.
Labels. Rsio [printed]; Eurycerus [Thomson cabinet label].

A second lectotype designation (for a different specimen and invalid) was published by Aubert (1966:
128).
Identity. Ichneumon eurycerus Thomson.

Ichneumon (Ichneumon) gibbulus, 18866: 21. Syntypes 1 $, 1 <J, SWEDEN: Skane (UZI, Lund).

Labels. Scan occi [printed] ; gibbulus Ths. [Thomson cabinet label] (9). Scan occi [printed] (^).
Identity. Ichneumon gibbulus Thomson.

Ichneumon (Ichneumon) grandiceps, 1887a: 13. Holotype 9, SWEDEN : Skane, Fagelsang (UZI, Lund).
Labels. Fogelsang 2 Jl 35 [hand] ; grandiceps [Thomson cabinet label].

Thomson gave the locality as only 'Suecia australis' in the original description but later (1893: 1953)
gave more details.

Identity. Cratichneumon grandiceps (Thomson) comb. n.

Ichneumon (Ichneumon) grandicornis, 18866: 24. LECTOTYPE 9, SWEDEN: Lappland (UZI, Lund), here
designated (selected by G. H. Heinrich).
Label. Norrl. [printed].
Identity. Ichneumon grandicornis Thomson.

THE ICHNEUMONIDAE DESCRIBED BY C. G. THOMSON 45

Ichneumon hypolius, 18886: 1226. Holotype 9, SWEDEN: Norrland, Halsingland (UZI, Lund).

Labels. Hels Ri [hand] ; [small gold square] [= G. F. Moller] ; hypolius [Thomson cabinet label].
Identity. Ichneumon hypolius Thomson.

Ichneumon (Ichneumon) jesperi, 1893: 1925. LECTOTYPE 9, SWEDEN: Skane, Stehag (UZI, Lund), here
designated.

Label. Shg. Haslsm vii.88 CM [hand].

The lectotype is also the lectotype of Ichneumon corfitzi Thomson. Thomson (1894: 2080) says that he
needs to correct a few wrong names, that number 29 [1893: 1925 I. jesperi] should be corfitzi and that
number 79 should be melanopygus. It is clear therefore that jesperi is an error for corfitzi and should have
the same type-specimen.

Identity. Junior objective synonym of Ichneumon corfitzi Thomson.

Ichneumon leucopeltis, 18886: 1230. Holotype $, SWEDEN : Jemtland [= Jamtland] (UZI, Lund).
Label. Jtl [printed].
Identity. Ichneumon leucopeltis Thomson.

Ichneumon liocnemis, 18886: 1220. Syntypes 1 ?, 2 cJ, SWEDEN: Gotheborg [= Goteborg] and Stockholm
area (UZI, Lund).

Labels. Gbg [hand] (2 rf). Him [printed] ; P. Wg. [printed] ; Col. Hgn. [printed] (1 ?).
Identity. Coelichneumon liocnemis (Thomson).

Ichneumon (Ichneumon) ttostylus, 1887a: 12. Syntypes 2 $, 3 <$, SWEDEN: Skane, Palsjo and Degeberga
(UZI, Lund).

Labels. Hbg. [hand]; liostylus [Thomson cabinet label] (1 ?). Degeberga [printed] (1 $). Pal. [hand]
(2 c?).Hbg. [hand] (!(?).

It is possible that the males are not syntypes (see Thomson, 1893: 1950).

Identity. Cratichneumon liostylus (Thomson).

Ichneumon (Ichneumon) longeareolatus, 18866: 21. Lectotype ?, SWEDEN: Skane (UZI, Lund), by des-
ignation of Townes, Momoi & Townes, 1965: 471.
Label. Sc. [hand].
Identity. Ichneumon longeareolatus Thomson.

Ichneumon (Ichneumon) macrocerus, 18866: 20. Syntypes 49, 6 <$, SWEDEN: Skane, Ringsjon, Palsjo, Reften
and Fagelsang and GERMANY (UZI, Lund).
No notes were made of individual specimen labels.
Identity. Ichneumon macrocerus Thomson.

Ichneumon (Ichneumon) mesostilpnus, 1888a: 107. Holotype 9, GERMANY (WEST): Aachen (UZI, Lund).

Labels. 8/525 ['8' hand, '525' printed]; AACHEN Aug. 75 [printed]; Tb [printed] ; Ichneumon albosig-
natus Grv. 9 [hand] ; mesostilpnus Th [Thomson cabinet label].
Identity. Barichneumon mesostilpnus (Thomson) comb. n.

Ichneumon (Ichneumon) micropnygus, 1893: 1927. Replacement name for Ichneumon (Ichneumon) spiracu-
laris Thomson (see below), junior primary homonym oflchneumon spiracularis Tischbein.

Ichneumon (Ichneumon) monospilus, 1896: 2398. Holotype 9, SWEDEN : Halsingland (UZI, Lund).

Labels. His [printed]; Rud [hand]; gravipes [hand]; Frei? [hand]; monospilus m [Thomson cabinet
label].

Identity. Ichneumon monospilus Thomson.

Ichneumon (Ichneumon) nereni [as nereni], 1887a: 8. Lectotype 9, SWEDEN: Skane (UZI, Lund), by des-
ignation of Townes, Momoi & Townes, 1965: 474.

Labels. Scan occi [printed] ; Nereni [Thomson cabinet label].
Identity. Ichneumon nereni Thomson.

Ichneumon (Ichneumon) nordenstromi [as nordenstromi], 1896: 2399. Holotype 9, NORWAY: Dovre (UZI,
Lund).

Labels. 33. [hand]; Dovre Nord-strom [hand] ; Nordenstromi m [Thomson cabinet label].
Identity. Ichneumon nordenstromi Thomson.

Ichneumon (Ichneumon) nudicoxa, 1888a: 107. Syntypes 2 9, 2 $, SWEDEN: Stockholm area and Skane,
Asperod (UZI, Lund).
Labels. Esp. [hand] (1 9). Col. Hgn. [printed] (1 9). Holm [printed] ; Col. Hgn. [printed] (2 <J).

46 M. G. FITTON

Although Thomson only gave 'Suecia' as the locality in the original description, he later (1893: 1958)
gave more details: 'funnen vid Torekov och Esperod i Skane'. The three syntypes originating from the
Holmgren collection are considered to be the specimens of albosignatus sensu Holmgren. Two other
specimens in the collection are not considered to be syntypes.

Identity. Junior synonym of Barichneumon digrammus (Gravenhorst) (Perkins, 1953 : 135).

Ichneumon (Ichneumon) pallitarsis, 1887a: 11. Syntypes 11 9» 3 <J» SWEDEN (UZI, Lund).

No notes were made of individual specimen labels. The syntype series includes specimens from the
following localities: Skane, Torekov (label Tkov'); Lappland (label 'LpF); and Norrland (label 'NorF).
Identity. Cratichneumon pallitarsis (Thomson).

Ichneumon ( Cratichneumon) parviscopa, 1893: 1950. Syntypes 1 9, 5 <?, SWEDEN: Skane, Ringsjon^^ only)
(UZI, Lund).

Labels, [green square] (5 $). Col. Hgn. [printed] (1 ?).

The female specimen originating from the Holmgren collection is assumed to be nigritarius sensu
Holmgren. It does not have a locality.

Identity. Cratichneumon parviscopa (Thomson) comb. n.

Ichneumon (Ichneumon) quadriannellatus, 1893: 1929. Unjustified emendation of Ichneumon (Ichneumon)
quadriannulatus Thomson, 1887a (see entry below).

Ichneumon (Ichneumon) quadriannulatus, 1887a: 10. Syntype 1 9, SWEDEN: Lappland (UZI, Lund).

Label. Norrl. [hand].

Identity. Junior primary homonym of Ichneumon quadriannulatus Gravenhorst, 1829a. Thomson
(1893: 1929) changed the name to quadriannellatus and maintained this change when he referred to the
species for a third time (1896: 2395). However, in neither of these subsequent references did Thomson
mention the homonymy or the original spelling. The name quadriannellatus is therefore probably best
treated as an unjustified emendation rather than a replacement name or incorrect subsequent spelling. As
such it is a junior objective synonym of /. quadriannulatus Thomson and has the same type. It is the oldest
available name for this species, which belongs in the genus Ichneumon.

Ichneumon (Ichneumon) quinquenotatus [as 5-notatus], 1893: 1936. Syntypes 1 9, 1 <$, SWEDEN: Uppland
and Skane, Ringsjon (UZI, Lund).

Labels. Col. Hgn. [printed]; 5-notatus Ths. [Thomson cabinet label] (9). [green square] (£).

Identity. Ichneumon quinquenotatus Thomson. Dalla Torre (1902: 977) gives an erroneous reference to
an earlier publication by Tischbein of the same name. Tischbein did not describe an Ichneumon quinque-
notatus.

Ichneumon (Ichneumon) simulosus, 18866: 16. Syntype 1 $, SWEDEN : Skane, Ramlosa(UZI, Lund).

Labels. Hbg. [hand] ; simulosus [the first letter could be 's' or 'r'] [Thomson cabinet label].

Thomson gave only 'Suecia australi' as the locality in the original description but later (1893: 1966) he
stated 'Funnen i bada konen vid Ramlosa'. A male specimen (from Palsjo) in the collection probably
post-dates 1893 and is not therefore regarded as a syntype.

In the 1893 reference to the species Thomson altered the spelling of the name to rimulosus. This spelling
has been widely used since, but it is incorrect.

Identity. Stenichneumon simulosus (Thomson).

Ichneumon (Ichneumon) spiracularis, 18866: 22. Holotype 9, SWEDEN: Norrland (UZI, Lund).
Labels. Norl. [printed] ; spiracularis [Thomson cabinet label].

Identity. Junior primary homonym of Ichneumon spiracularis Tischbein. Replacement name Ichneumon
(Ichneumon) micropnygus Thomson (see above). The species belongs in the genus Ichneumon.

Ichneumon (Ichneumon) stenocarus, 1887a: 13. Lectotype 9, SWEDEN : Skane, [? Asperod] (UZI, Lund), by
designation of Townes, Momoi & Townes, 1965: 446.

Labels. 29 Asperod [hand, not clear, could easily be '29 Aug ']; stenocarus [Thomson cabinet

label].

Identity. Cratichneumon stenocarus (Thomson).

Ichneumon (Ichneumon) stenocerus, 1887a: 7. Syntypes 6 9, SWEDEN: Goteborg, Skane, Ringsjo and
? Skane, Hassleholm (UZI, Lund).

Labels. Gbg [hand]; Stenocerus [Thomson cabinet label] (1 9)- Ringsio [printed] (1 9)- Scania
[printed] (2 9). Scan med [printed] (1 9). Hhm [hand, ? individual letters] [? = Hassleholm] (1 9).

Identity. Ichneumon stenocerus Thomson.

THE ICHNEUMONIDAE DESCRIBED BY C. G. THOMSON 47

Ichneumon (Ichneumon) subquadratus, 1887a: 9. Lectotype 9, SWEDEN: Skane, Palsjo (UZI, Lund), by
designation of Townes, Momoi & Townes, 1 965 : 48 1 .
Labels. Pal [hand] ; subquadratus [Thomson cabinet label].
Identity. Ichneumon subquadratus Thomson.

Ichneumon (Coetichneumon) tenuitarsis, 1893: 1907. Holotype?, SWEDEN: ? Ostergotland (UZI, Lund).
Labels, [small blue square]; Col. Hgn. [printed] ; tenuitarsis [Thomson cabinet label].
Thomson expressed doubt about the locality, presumably he was guessing at the meaning of the blue
square.

Identity. Coelichneumon tenuitarsis (Thomson) comb. n.

Ichneumon trispilus [as 3-spilus\, 18886: 1228. Syntypes 6 $, SWEDEN : Skane, Palsjo (UZI, Lund).
Labels. Pal. [hand] ; trispilus [Thomson cabinet label] (1 ?). Pal [hand] (5 9).
Identity. Ichneumon trispilus Thomson.

Ichneumon (Ichneumon) truncatulus, 18866: 15. Syntypes 1 9, 1 $, SWEDEN : Skane, Reften (UZI, Lund).

Labels. Rfn 27/6 [hand] ; truncatulus [Thomson cabinet label] (£). Rfn 10/7 [hand] (?).

Thomson gave only 'Suecia australi' as the locality in the original description but later (1893: 1911) he
stated 'Funnen vid Reften nara Lund'. Specimens in the collection from other localities probably post-
date 1893 and therefore cannot be syntypes.

Identity. Coelichneumon truncatulus (Thomson).

Ichneumon (Eupalamus) wesmaeli, 18866: 12. Lectotype 9, SWEDEN: Skane, Lindholmen (UZI, Lund), by
designation of Townes, Momoi & Townes, 1965: 448.

Label. Lhn 8/8 [hand].

Townes, Momoi & Townes (1965: 448) gave the original combination incorrectly as Eupalamus wes-
maeli.

Identity. Eupalamus wesmaeli (Thomson).

Ichneumon (Ichneumon) xanthognathus, 1887a: 8. Lectotype 9, SWEDEN: Skane, Ronnemolla (UZI, Lund),
by designation of Aubert, 1966: 128.
Label. Ron [hand].
Identity. Ichneumon xanthognathus Thomson.

Ischnus (Ischnus) coxator, 1891: 1624. LECTOTYPE <J, SWITZERLAND: Zermatt (UZI, Lund), here des-
ignated (selected by J. F. Aubert).
Labels. Zermatt [hand] ; coxator [hand].
Identity. Heterischnus coxator (Thomson) comb. n.

Ischnus (Ischnus) pulchellus, 1891: 1625. Syntypes 1 9, 1 <J, YUGOSLAVIA: Dalmatia(^) and [?](9) (UZI,
Lund).

Labels. +158 [hand]; Dalm. [printed] (<$). +139 [hand]; Buss [printed, ? a locality]; pulchellus m
[Thomson cabinet label] (9).

Identity. Heterischnus pulchellus (Thomson) comb. n.

Lathrolestus caudatus, 1883 : 917. Type(s) 9 [and ? <?], SWEDEN : Norrland (lost).

Identity. Lathrolestes caudatus (Thomson) (on the basis of a specimen in the collection).

Lathrolestus luteolus, 1883:917. Holotype <$, SWEDEN : Skane, Lund (UZI, Lund).
Label. L-d [printed].
Identity. Lathrolestes luteolus (Thomson).

Lathrolestus marginatus, 1883: 917. Lectotype 9, SWEDEN: Skane, Fogelsang[= Fagelsang] (UZI, Lund), by
designation of Aubert, 1972: 147.

Labels. Sk. [hand] ; marginatus [Thomson cabinet label].
Identity. Lathrolestes marginatus (Thomson).

Lathrolestus pie uralis, 1883: 916. LECTOTYPE 9, SWEDEN: Norrland (UZI, Lund), here designated (selec-
ted by R. Hinz).

Labels. 238 [hand]; Norr [hand] ; pleuralis [Thomson cabinet label].
Identity. Lathrolestes pleuralis (Thomson).

Lathrolestus ungularis, 1 883 : 9 1 8. Syntype 1 9, SWEDEN : Skane, Palsio [ = Palsjo] (UZI, Lund).
Labels. Pal. [hand] ; ungularis [Thomson cabinet label].
Identity. Lathrolestes ungularis (Thomson).

48 M. G. FITTON

Lathroplex clypearis, 1887c: 1135. Lectotype 9, SWEDEN: Skane, Ringsion [ = Ringsjon] (UZI, Lund), by
designation of Horstmann, 1977: 68.
Label. Rsio [printed].

Identity. Campoplex clypearis (Thomson). Horstmann (1977) considers Lathroplex distinct from Cam-
poplex.

Lathrostiza forticanda, 1887c: 1153. Lectotype $, SWEDEN: Lappland (UZI, Lund), by designation of Horst-
mann, 1971a: 11.

Labels. Lpl. [printed] ; Ths [printed] ; forticauda [Thomson cabinet label].

Authors since Thomson (including Horstmann, 1971) have chosen to alter the spelling of the name to
forticauda and there is evidence (Thomson's own cabinet label) that this is what was intended. However, a
strict interpretation of Article 32(aXii) of the Code (as amended, Bull. zoo/. Norn. 31 (1974): 83) suggests
that the original spelling should be retained.

Identity. Lathrostizus forticanda (Thomson).

Lathrostiza sternocera, 1887c: 1152. Lectotype 9, SWEDEN: Skane, Stehag (UZI, Lund), by designation of
Horstmann, 1971a: 10.
Label. Ste 5/81 [hand].
Identity. Lathrostizus sternocerus (Thomson).

Leptocryptus brevis, 1884: 965. Lectotype 9, SWEDEN: Skane, Lund (UZI, Lund), by designation of Sawonie-
wicz, 1978: 126.

Labels. Lund [printed] ; brevis [Thomson cabinet label].
Identity. Junior synonym of Bathythrix aereus (Gravenhorst) (Sawoniewicz, 1978 : 126).

Leptocryptus clavipes, 18886: 1243. Lectotype 9, SWEDEN: Skane, Ortofta (UZI, Lund), by designation of
Townes, Momoi & Townes, 1965: 127.
Labels. Ortofta [printed] ; clavipes [Thomson cabinet label].

I do not consider that the earlier publication by Aubert (1964: 61), cited by Horstmann (1976a: 28),
constitutes a valid lectotype designation.
Identity. Leptocryptoides clavipes (Thomson) (Horstmann, 1976a: 27).

Leptocryptus collaris, 1896: 2388. Lectotype 9, SWEDEN: Skane, Rostanga (UZI, Lund), by designation of
Sawoniewicz, 1980: 356.
Labels. Rost. [hand] ; collaris n [hand].
Identity. Bathythrix collaris (Thomson).

Leptocryptus geniculosus, 1884: 966. Lectotype 9, SWEDEN: Smaland (UZI, Lund), by designation of Aubert,
1966: 129.

Labels. Sm. [printed] ; geniculosus [Thomson cabinet label].
Identity. Junior synonym of Bathythrix fragilis (Gravenhorst) (Sawoniewicz, 1978: 127).

Leptocryptus heteropus, 1884: 1040. Lectotype 9, SWEDEN: Skane, Bokeberg (UZI, Lund), by designation of
Hinz in Sawoniewicz, 1980: 360.

Labels. Yddinge [printed] ; heteropus [Thomson cabinet label].
Identity. Junior synonym of Bathythrix linearis (Gravenhorst) (Sawoniewicz, 1980: 360).

Leptocryptus lamina, 1884: 965. Lectotype $, SWEDEN: Skane, Yddinge (UZI, Lund), by designation of
Aubert, 1972: 148.
Label. Yd. [hand].
Identity. Bathythrix laminus (Thomson).

Leptocryptus rugulosus, 1884: 966. Lectotype 9, SWEDEN: Skane, Ortofta (UZI, Lund), by designation of
Sawoniewicz, 1980: 329.

Labels. Ort. [hand]; rugulosus [Thomson cabinet label].
Identity. Bathythrix rugulosus (Thomson).

Leptocryptus strigosus, 1884: 964. Lectotype 9, SWEDEN: Skane, Helsingborg (UZI, Lund), by designation of
Aubert, 1972: 148.
Label. Hbg. [hand].
Identity. Bathythrix strigosus (Thomson).

Limneria costalis, 1887c: 1106. Lectotype 9, SWEDEN: Norrland (UZI, Lund), by designation of Aubert,
1966: 130.
Label. Norl. [printed].

THE ICHNEUMONIDAE DESCRIBED BY C. G. THOMSON 49

Although Townes, Momoi & Townes (1965: 272) were almost certainly correct in recognising this as
the type ( = holotype, the single original specimen) Aubert chose to designate it as lectotype. He presented
no evidence of a syntype series.

Identity. Sinophorus costalis (Thomson).

Limm'ria crassifemur, 1887c: 1106. Lectotype 9, SWEDEN: Skane, Ringsion [= Ringsjon] (UZI, Lund), by
designation of Townes, Momoi & Townes, 1965: 272.
Labels, [green square] ; crassifemur [Thomson cabinet label].
Identity. Sinophorus crassifemur (Thomson).

Limneria fuscicarpus, 1887c: 1104. Lectotype <$, SWEDEN: Skane, Palsjo (UZI, Lund), by designation of
Townes, Momoi & Townes, 1965 : 272.
Label. Pal. [hand].
Identity. Sinophorus fuscicarpus (Thomson).

Limneria hyperborea, 1887c: 1 106. Holotype 9, NORWAY : Tromse (UZI, Lund).
Labels. Tromso 14.6.77 [hand] ; hyperborea [Thomson cabinet label].
Identity. Tranosema hyperborea (Thomson) (Horstmann, 1977: 78).

Limneria nigritella, 1887c: 1 107. Type(s) 9, SWEDEN : Skane, Sjobo (lost).
Identity. ? Sinophorus nigritellus (Thomson) comb. n.

Limneria pine ticola, 1887c: 1108. Lectotype 9, SWEDEN: Skane, Kavlinge (UZI, Lund), by designation of
Aubert, 1968: 195.
Label. Kfge [hand].
Identity. Sinophorus pineticola (Thomson).

Limneria planiscapus, 1887c: 1105. Lectotype 9, SWEDEN: Skane, Lund (UZI, Lund), by designation of
Aubert, 1966: 130.

Labels. Lund 24/5 [hand] ; planiscapus [Thomson cabinet label].
Identity. Sinophorus planiscapus (Thomson).

Limneria pleuralis, 1887c: 1 105. Syntypes 2 9, SWEDEN: Skane, Fogelsang [= Fagelsang] (UZI, Lund).
Labels. Scan occi [printed] (2 9)-
Identity. Sinophorus pleuralis (Thomson) comb. n.

Limneria rufifemur, 1887c: 1106. Lectotype 9, SWEDEN: Skane, Torekov (UZI, Lund), by designation of
Townes, Momoi & Townes, 1965: 273.

Labels. Tkov 18/6 [hand] ; rufifemur [Thomson cabinet label] ; 72. [hand].
Identity. Sinophorus rufifemur (Thomson).

Limneria tegularis, 1887c: 1 107. Holotype 9, SWEDEN: Skane, Skanor (UZI, Lund).
Labels. Skanor [hand] ; tegularis [Thomson cabinet label].
Identity. Sinophorus tegularis (Thomson) comb. n.

Liocryptus tenuicornis, 1896: 2356. Holotype 9, SWEDEN : Norrland (UZI, Lund).
Label. Norrl [printed].
Identity. Idiolispa tenuicornis (Thomson).

Lissonota antennaKs, 1877: 765. LECTOTYPE 9, SWEDEN: Skane, Ilstorp (UZI, Lund), here designated
(selected by R. Hinz).
Label. lisp 15/8 [hand].
Identity. Lissonota antennalis Thomson.

Lissonota basalts, 1889: 1424. Lectotype 9, SWEDEN: Skane, Ringsjon (UZI, Lund), by designation of
Aubert, 1972: 146.

Labels. Rsio [printed] ; basalis m [Thomson cabinet label].

Identity. Junior primary homonym of Lissonota basalis Brischke, 1865. Replacement name Lissonota
mutanda Schmiedeknecht, 1900: 377. Junior synonym of Lissonota saturator (Thunberg) (Aubert,
19786: 110).

Lissonota carinifrons, 1877: 768. Syntypes 2 9, SWEDEN: Skane, Asperod (UZI, Lund).

Labels. Esp [printed] ; carinifrons [Thomson cabinet label] (1 9). Fall, [hand] ; Col Zet [printed] (1 9).
Identity. Junior synonym of Lissonota quadrinotata Gravenhorst (Aubert, 19786: 108).

50 M. G. FITTON

Lissonota clypealis, 1877: 769. LECTOTYPE 9, SWEDEN: Skane, Helsingborg, Palsjo (UZI, Lund), here
designated (selected by J. F. Aubert).

Label. Pal. [hand].

Aubert's mention (19786: 87) of 'lectotype inedit, Palsjo 9' is not a valid lectotype designation because
he does not indicate to which of three species it applies.

Identity. Lissonota clypealis Thomson.

Lissonota crassipes, 1877: 772. Lectotype 9, SWEDEN: Skane, Lindholmen (UZI, Lund), by designation of
Aubert, 1966: 127.
Label. Lhn 14/8 [hand].
Identity. Junior synonym of Lissonota biguttata Holmgren (Aubert, 19786: 84).

Lissonota folii, 1877: 771. Lectotype 9, SWEDEN: Skane, Ringsjon (UZI, Lund), by designation of Townes,
Momoi &Townes, 1965: 217.

Labels. Ringsjo 16/7 [hand]; uv Cyn. qu. g . . . . [hand, partly illegible] ; folii [Thomson cabinet label].
Identity. Lissonota folii Thomson.

Lissonota genalis, 1877: 760. Syntypes2 9, SWEDEN : Norrland (UZI, Lund).

Labels. Norl. [printed] ; genalis [Thomson cabinet label] (1 9)- Norl. [printed] (1 $).
Aubert (1972: 146) probably only saw one specimen, which he assumed (quite reasonably, but incor-
rectly) to be a holotype.

Identity. Cryptopimpla genalis (Thomson).

Lissonota gracittpes, 1877: 770. Lectotype 9, SWEDEN : Skane, Bastad (UZI, Lund), by designation of Aubert,
1966: 127.
Label. Bast [hand].
Identity. Lissonota gracilipes Thomson.

Lissonota hians, 1877: 762. Lectotype 9, SWEDEN: Skane, Lund (UZI, Lund), by designation of Aubert,
19786: 91.

Label. Ld 27/5 [hand].
Identity. Junior synonym of Lissonota digestor (Thunberg) (Aubert, 19786: 91).

Lissonota humerella, 1877: 771. Lectotype 9, SWEDEN: Oland (UZI, Lund), by designation of Aubert,
1966: 127.

Labels. 0. [printed] ; humerella [Thomson cabinet label].
Identity. Lissonota humerella Thomson.

Lissonota impressifrons, 1889: 1419. Lectotype 9, FRANCE (UZI, Lund), by designation of Aubert, 1972: 146.
Labels. Gall, [hand] ; impressifrons m [hand].
Identity. Lissonota impressifrons Thomson.

Lissonota irrigua, 18886: 1248. Lectotype 9, SWEDEN: Skane, Ortofta (UZI, Lund), by designation of
Townes, Momoi & Townes, 1965: 218.

Labels. Ort. [hand] ; irrigua [Thomson cabinet label].

Identity. Junior synonym of Lissonota coracina (Gmelin) (Aubert, 19786: 89).

Lissonota nigridens, 1889: 1425. Lectotype 9, SWEDEN: Skane, Palsjo (UZI, Lund), by designation of Aubert,
1972: 146.

Labels. Pal [hand] ; nigridens [Thomson cabinet label].
Identity. Lissonota nigridens Thomson.

Lissonota palpalis, 1889: 1422. Lectotype 9, SWEDEN: Skane, Ringsjon (UZI, Lund), by designation of
Aubert, 1968: 194.
Label. Rsio [printed].
Identity. Lissonota palpalis Thomson.

Lissonota punctiventris, 1877: 769. Replacement name for Lissonota errabunda Holmgren, 1860, junior
secondary homonym of Lissonota errabunda (Gravenhorst, 18296) in Thomson's treatment (1877: 759-
772) of this group.

The Thomson name is a junior objective synonym of the Holmgren name, and as such must have the
same type. The lectotype designations of Townes, Momoi & Townes (1965: 219) and Aubert (1966: 127)
for punctiventris are invalid. A lectotype for Lissonota errabunda Holmgren, and thus also for Lissonota
punctiventris Thomson, 1877, was designated by Aubert (1968: 187).

Identity. Under the provisions of Article 59(bXi) of the Code (as amended, Bull. zool. Norn. 31 (1974): 83)

THE ICHNEUMONIDAE DESCRIBED BY C. G. THOMSON 51

the International Commission must decide whether punctiventris or errabunda should be used, because
the rejected homonym (errabunda) continued to be and is currently in use as the name of this species.
However, this might be an academic point as Aubert (19786: 85, 93) suggests that the species might be a
junior synonym of Lissonota buccator (Thunberg).

Lissonota (Syzeucta) punctiventris, 1894: 2128. Holotype 9, ITALY: Trieste (UZI, Lund).
Label. 29/VI Triest. [date, hand; locality, printed].

It is difficult to see why Aubert (19786: 133) stated that this species was described as a variety of
Lissonota punctiventris Thomson, 1 877 and that the name is 'sans valeur systematique'. After describing a
variety of S. maculatoria from Holmgren's collection Thomson (1894: 2128) immediately described the
new species as follows :

'Anm. Fran Triest har jag erhallit en hona, som synes tillhora en annan art:
Ib. S. punctiventris m.

Praecedentis varietati similis, sed major, abdomine segmentis anterioribus parcius subtiliter punc-
tatis'.

He in no way referred to L. punctiventris. Schmiedeknecht (1900: 345) gave a clear account of the
situation.

Identity. Junior primary homonym of Lissonota punctiventris Thomson, 1877. Replacement name
Syzeuctus tenuifasciatus Schmiedeknecht, 1900: 345.

Lissonota rimator, 1877: 762. Syntypes 1 ?, 3 & SWEDEN : Nerike [= Narke] and Skane, Lund (UZI, Lund).

Labels. Ner. [hand] (1 $, 2 <J). L-d [printed] (1 £).

The specimen published as lectotype by Aubert (1972: 146) is labelled 'Gbg'(= Goteborg) and cannot
therefore be one of the syntypes. The four syntypes noted above were found in the duplicate collection (the
fourth drawer in cabinet 404) under the name Lissonota rimator. They have been transferred to the main
collection.

Identity. Lissonota rimator Thomson.

Lissonota subfumata, 1877: 760. Lectotype 9, SWEDEN: Skane, Ilstorp (UZI, Lund), by designation of
Aubert, 1972: 146.

Labels. lisp 25/7 [hand] ; subfumata [Thomson cabinet label].
Identity. Cryptopimpla subfumata (Thomson).

Lissonota tenerrima, 1877: 766. Lectotype 9, SWEDEN: Smaland (UZI, Lund), by designation of Aubert,
1972: 146.

Label. Coll. L-gh [printed].
Identity. Lissonota tenerrima Thomson.

Lissonota varicoxa, 1877: 768. Holotype 9, SWEDEN: Skane, Markiehage (UZI, Lund).
Labels. Mrki [hand] ; varicoxa [hand].
Identity. Junior synonym of Lissonota buccator (Thunberg) (Aubert, 19786: 85).

Macrochasmus alysiina, 18886: 1279. LECTOTYPE [?sex], SWEDEN: Lappland (UZI, Lund), here
designated (selected by H. K. Townes).

Label. Lap [hand].

The lectotype is badly damaged and lacks head and gaster. No other syntypes are present in the
collection.

Identity. Idiogramma alysiina (Thomson).

Macrocryptus coraebi, 1885: 19. Syntypes 9 c£, FRANCE (lost).

Thomson attributed this species to Regimbart as 'Cryptus Coraebi Regimb.' but said that it belonged in
the genus Macrocryptus. The name was never published by Regimbart and therefore Thomson is the
author.

The name has often been misspelled coroebi.

Identity. Xylophrurus coraebi (Thomson).

Megastylus (Helictes) pilicornis, 18886: 1312. Holotype 9, GERMANY (WEST) : Aachen (ZSBS, Munich).
Labels. 9 30 .... Aachen [hand, partly illegible] ; Pilicornis Thorns [hand] ; invalidus Frst. [hand].
Identity. Helictes pilicornis (Thomson).

Megastylus (Megastylus) pleuralis, 18886: 1313. Syntypes 1 ?, 1 & U.S.S.R.: Tartu [not 'norra Tyskland' as
stated by Thomson] (UZI, Lund).

Labels. 26/8 85. [hand]; Dorpat [hand]; pleuralis [Thomson cabinet label] (9). 26/8 85. [hand]; 926
[hand](<J).

52 M. G. FITTON

The specimens match the description and are almost certainly the syntypes. The locality given by
Thomson is obviously a mistake (which he also made in the case of Campoplex latungula).
Identity. Megastylus pleuralis Thomson.

Meloboris hygrobia, 1 887c : 1151. Lectotype 9, SWEDEN : Skane, Lund (UZI, Lund), by designation of Horst-
mann, 1969: 432.

Labels. Ld [printed] ; hygrobia [Thomson cabinet label].
Identity. Diadegma hygrobia (Thomson).

Meloboris ischnocera, 1887c: 1151. Lectotype 9, SWEDEN: Skane, Lund (UZI, Lund), by designation of
Horstmann, 1969: 432.
Label. Ld [hand].
Identity. Junior synonym of Diadegma hygrobia (Thomson) (Horstmann, 1969: 432).

Mesochorus (Mesochorus) acuminatus, 1886a: 343. LECTOTYPE £, SWEDEN: Skane, Yddinge (UZI,
Lund), here designated (selected by W. Schwenke).
Labels. Yd. [hand] ; acuminatus [Thomson cabinet label].
Identity. Mesochorus acuminatus Thomson.

Mesochorus (Mesochorus) albipes, 1886a: 341. LECTOTYPE 9, SWEDEN: Skane, Lund (UZI, Lund), here
designated (selected by R. Hinz).

Labels. Ld [hand] ; albipes [Thomson cabinet label].
Identity. Mesochorus albipes Thomson.

Mesochorus (Mesochorus) angustatus, 1886a: 343. LECTOTYPE 9, SWEDEN: Skane, Lund (UZI, Lund),
here designated (selected by W. Schwenke).
Labels. Ld [hand] ; angustatus [Thomson cabinet label].
Identity. Mesochorus angustatus Thomson.

Mesochorus (Stictopisthus) bilineatus, 1886a: 344. LECTOTYPE <J, SWEDEN: Skane, Lund (UZI, Lund),
here designated (selected by W. Schwenke).
Labels. Lund [printed] ; bilineatus [Thomson cabinet label].
Identity. Stictopisthus bilineatus (Thomson).

Mesochorus (Mesochorus) brevicollis, 1886a: 335. LECTOTYPE 9, SWEDEN: Skane, Helsingborg (UZI,
Lund), here designated (selected by W. Schwenke).
Label. Hbg [hand].
Identity. Mesochorus brevicollis Thomson.

Mesochorus (Mesochorus) brevigena, 1886a: 338. LECTOTYPE 9, SWEDEN: Skane, Helsingborg (UZI,
Lund), here designated (selected by K. Horstmann).
Labels. Hg [hand] ; brevigena [Thomson cabinet label].
Identity. Mesochorus brevigena Thomson.

Mesochorus (Astiphrommus) buccatus, 1886a: 329. LECTOTYPE 9, SWEDEN : Skane, Tvedora(UZI, Lund),
here designated (selected by W. Schwenke).

Labels. Tve 6/78 [hand] ; [small gold square] ; buccatus [Thomson cabinet label].
Identity. Astiphromma buccatum (Thomson).

Mesochorus (Stictopisthus) convexicollis, 1886a: 344. LECTOTYPE 9, SWEDEN: Skane, Helsingborg (UZI,
Lund), here designated (selected by W. Schwenke).
Labels. Hbg. [hand] ; convexicollis [Thomson cabinet label].
Identity. Stictopisthus convexicollis (Thomson).

Mesochorus (Mesochorus) crassicrus, 1886a: 339. Syntypes 3 9, 3 & 1 ?sex, SWEDEN: Skane (UZI, Lund).

Labels. Pal. [hand] ; crassicrus [Thomson cabinet label] (1 9). Ort. [hand] (2 <J). Pal. [hand](l 9, 1 rf).
Rsio [printed] (1 9). Sbg 23/7 [hand] (1 ?sex).
Identity. Mesochorus crassicrus Thomson.

Mesochorus (Mesochorus) curvicauda, 1886a: 335. LECTOTYPE 9, SWEDEN: Oland (UZI, Lund), here
designated (selected by W. Schwenke).
Labels. O. [printed] ; curvicauda [Thomson cabinet label].
Identity. Mesochorus curvicauda Thomson.

Mesochorus (Mesochorus) curvulus, 1886a: 343. LECTOTYPE £, SWEDEN: Skane, Ortofta (UZI, Lund),
here designated (selected by W. Schwenke).
Label. Ort. [hand].
Identitv. Mesochorus curvulus Thomson.

THE ICHNEUMONIDAE DESCRIBED BY C. G. THOMSON 53

Mesochorus (Mesochorus) fulvus, 1886a: 336. Lectotype $, SWEDEN: Skane, Palsjo (UZI, Lund), by
designation of Townes, Momoi & Townes, 1965: 342.
Labels. Pal [hand] ; fulvus [Thomson cabinet label].
Identity. Mesochorus fulvus Thomson.

Mesochorus (Astiphrommus) graniger, 1886a: 328. LECTOTYPE?, SWEDEN: Skane, Ortofta (UZI, Lund),
here designated (selected by R. Hinz).
Labels. Ort. [hand] ; graniger [Thomson cabinet label].
Identity. Astiphromma graniger (Thomson).

Mesochorus (Astiphrommus) hamulus, 1886a: 330. ? Syntype 1 £, DENMARK: Zealand, Strandmollen (ZM,
Copenhagen).

Labels. <$ Strandm Drewsen [hand] ; Danmark ex coll. Schiodte [printed] ; Hamulus Thorns: [hand].

There are no syntypes in the Thomson collection in Lund. There is, unfortunately, no direct evidence
that the specimen in Copenhagen is a syntype.

Identity. Astiphromma hamulum (Thomson).

Mesochorus (Astiphrommus) incident, 1886a: 331. Type(s) 9, ENGLAND (lost).

There are no specimens of this species in the Bridgman collection (A. G. Irwin, pers. comm.) (see
Bridgman, 1886: 335, 353 and 354). The only specimen in the Thomson collection is from Palsjo and
cannot be a type.

Identity. Astiphromma incidens (Thomson).

Mesochorus (Mesochorus) lapponicus, 1886a: 336. LECTOTYPE $, SWEDEN: Lappland (UZI, Lund), here
designated (selected by W. Schwenke).
Labels. Lap [hand] ; Lapponicus [Thomson cabinet label].
Identity. Mesochorus lapponicus Thomson.

Mesochorus (Stictopisthus) laticeps, 1886a: 344. LECTOTYPE 9, SWEDEN: Skane, Bokeberg (UZI, Lund),
here designated (selected by W. Schwenke).
Labels. Bok 8/84 [hand] ; laticeps [Thomson cabinet label].
Identity. Stictopisthus laticeps (Thomson).

Mesochorus (Mesochorus) longicauda, 1886a: 338. Lectotype 9, SWEDEN: Skane, Palsjo (UZI, Lund), by
designation of Aubert, 1972: 152.
Label. Pal [hand].
Identity. Mesochorus longicauda Thomson.

Mesochorus (Mesochorus) macrurus, 1886a: 342. LECTOTYPE 9, SWEDEN: Lappland (UZI, Lund), here
designated (selected by W. Schwenke).

Labels, [small paper square] ; [small paper square] ; Lpl. [hand] ; macrurus [Thomson cabinet label].
Identity. Mesochorus macrurus Thomson.

Mesochorus (Astiphrommus) mandibularis, 1886a: 330. Lectotype $, SWEDEN: Skane, Yddinge (UZI, Lund),
by designation of Townes, Momoi & Townes, 1965: 340.

Labels. Yd [hand] ; mandibularis [Thomson cabinet label].

Townes, Momoi &Townes (1965: 340) gave the original combination incorrectly as Astiphrommus
mandibularis.

Identity. Astiphromma mandibulare (Thomson).

Mesochorus (Mesochorus) marginatus, 1886a: 339. Lectotype 9, SWEDEN: Skane (UZI, Lund), by
designation of Aubert, 1966: 131.
Label. Scan [printed].
Identity. Mesochorus marginatus Thomson.

Mesochorus (Mesochorus) nigriceps, 1886a: 334. LECTOTYPE 9, SWEDEN : Skane, Lund (UZI, Lund), here
designated (selected by W. Schwenke).
Label. L-d [printed].
Identity. Mesochorus nigriceps Thomson.

Mesochorus (Mesochorus) pectinipes, 1886a: 336. Lectotype 9, SWEDEN: Skane (UZI, Lund), by designation
of Townes, Momoi &Townes, 1965: 344.
Label. Scan [printed].
Identity. Junior primary homonym of Mesochorus pectinipes Bridgman, 1883. Replacement name

Mocnfhmie cfi/>s.iV.fio Thalia Tr.rr» 1QH1- ^8

54 M. G. FITTON

Mesochorus (Mesochorus) picticrus, 1886a: 340. LECTOTYPE 9, SWEDEN: Skane, Palsjo (UZI, Lund), here
designated (selected by R. Hinz).
Label. Pal [hand].
Identity. Mesochorus picticrus Thomson.

Mesochorus (Astiphrommus) plagiatus, 1886a: 332. Syntypes 1 ^, SWEDEN: Skane, Helsingborg (UZI,
Lund); 1 3, ENGLAND (CM, Norwich).

Labels. Hbg. [hand] ; plagiatus [Thomson cabinet label] (Lund specimen). 546 [hand, on the specimen
mount]; G. C. Bignell April 1882 from Apanteles from Odontopera bidentata [hand, on the underside of
the specimen mount] ; plagiatus Thorn [hand] ; 3 [hand] (Norwich specimen).

The Norwich specimen is in the J. B. Bridgman collection. From Bridgman's paper (1886: 335, 353 and
354) and the label with the number (3) it is virtually certain that this specimen was sent to Thomson and is
a syntype.

Identity. Astiphromma plagiatum (Thomson).

Mesochorus (Mesochorus) punctipleuris, 1886a: 334. Lectotype 9, SWEDEN: Skane, Ringsjon (UZI, Lund),
by designation of Aubert, 1966: 131.
Label. Rsio [printed].
Identity. Mesochorus punctipleuris Thomson.

Mesochorus (Mesochorus) solids, 1886a: 338. LECTOTYPE 9, SWEDEN: Skane, Ringsjon (UZI, Lund),
here designated (selected by W. Schwenke).
Labels. Rsio [printed] ; Salicis [Thomson cabinet label].
Identity. Mesochorus salicis Thomson.

Mesochorus (Astiphrommus) simplex, 1886a: 334. LECTOTYPE 9, SWEDEN: Skane, Yddinge (UZI, Lund),
here designated (selected by W. Schwenke).
Labels. Yd [hand] ; simplex [Thomson cabinet label].
Identity. Astiphromma simplex (Thomson).

Mesochorus (Mesochorus) stigmaticus, 1886a: 341. LECTOTYPE 9, DENMARK: Maribo (UZI, Lund), here
designated (selected by W. Schwenke).

Labels. 28/7 77 Maribo ex Microgaster [hand] ; stigmaticus [Thomson cabinet label].

Identity. Junior primary homonym of Mesochorus stigmaticus Brischke, 1880. Replacement name
Mesochorus orgyiae Dalla Torre, 1901 : 56.

Mesochorus (Mesochorus) temporalis, 1886a: 336. Syntype 1 9, ENGLAND (CM, Norwich).

Labels. Bred from filipendulae 25.7.78 G. C. Bignell [hand, on the underside of the specimen mount] ; 48
[hand] ; temporalis Thn [hand].

The syntype is in the Bridgman collection. For the reasons why it is considered as such see the notes
under M . plagiatus above.

Identity. Mesochorus temporalis Thomson.

Mesochorus (Astiphrommus) tenuicornis, 1886a: 332. LECTOTYPE 9, SWEDEN: Skane, Palsjo (UZI, Lund),
here designated (selected by R. Hinz).
Labels. Pal [hand] ; tenuicornis [Thomson cabinet label].
Identity. Astiphromma tenuicornis (Thomson).

Mesochorus (Mesochorus) tenuiscapus, 1886a: 341. LECTOTYPE 9, SWEDEN: Lappland, Lund (UZI,
Lund), here designated (selected by W. Schwenke).

Labels. Lund 3 Ag. [hand] ; Lpl. [printed] ; tenuiscapus [Thomson cabinet label].
Identity. Mesochorus tenuiscapus Thomson.

Mesochorus (Mesochorus) tuberculiger, 1886a: 333. Lectotype ^, SWEDEN: Skane, Torekov (UZI, Lund), by
designation of Townes, Momoi & Townes, 1965: 345.

Labels. Trkv [hand] ; tuberculiger [Thomson cabinet label].
Identity. Mesochorus tuberculiger Thomson.

Mesocryptus nigriventris, 1896: 2384. Holotype 9, SWEDEN : 'Halland' [Skane], Margretetorp (UZI, Lund).
Labels. Hall, [printed] ; nigriventris m [Thomson cabinet label].

Margretetorp is in northern Skane, not southern Halland as stated by Thomson. It is near the bound-
ary between the two provinces, so Thomson's error is easily explained.
Identity. Oresbius nigriventris (Thomson) comb. n.

THE ICHNEUMONIDAE DESCRIBED BY C. G. THOMSON 55

Mesocryptus ochrostomus, 1896: 2384. Holotype 9, SWEDEN : Skane, Palsjo (UZI, Lund).
Labels. Pal [hand] ; ochrostomus m [Thomson cabinet label].
Identity. Aptesis ochrostomus (Thomson) comb. n.

Mesoleius (Alexeter) albilabris, 1894: 2025. Syntypes 3 9, 3 <J, SWEDEN : Skane, Palsjo (UZI, Lund).

Labels. Pal. [hand]; 9 [printed] (3 9). Palsio [printed] (1 <J). Hbg [hand]; scutellaris [Thomson
cabinet label] (1 £). Hbg. [hand] (1 <J).
Identity. Alexeter albilabris (Thomson).

Mesoleius (Barytarbus) annulipes, 1 883 : 932. Holotype <$, SWEDEN : Gotland (UZI, Lund).
Labels. Got [printed] ; annulipes [Thomson cabinet label].
Identity. Barytarbes annulipes (Thomson).

Mesoleius (Mesoleius) brachypus, 1894: 2054. Lectotype9, SWEDEN: Norrland (UZI, Lund), by designation
of Aubert, 19766:269.

Labels. Col. Rud. [hand] ; brachypus m [Thomson cabinet label].
Identity. Anoncus brachypus (Thomson).

Mesoleius (Mesoleius) brevipalpis, 1894: 2047. Lectotype 9, SWEDEN: Oland (UZI, Lund), by designation of
Aubert, 19766:269.

Labels. 0. [printed]; 9 [printed] ; brevipalpis [Thomson cabinet label].
Identity. Mesoleius brevipalpis Thomson.

Mesoleius (Saotus) brevispina, 1883: 934. Syntypes 8 9, SWEDEN: Skane, Lund (UZI, Lund).

Labels. Lund [printed] ; brevispina [Thomson cabinet label] (4 9, all on one pin). Lund [printed] (4 9,
two on one pin).

Identity. Saotis brevispina (Thomson).

Mesoleius (Mesoleius) brevitarsis, 1894: 2037. Lectotype 9, SWEDEN: Norrland (UZI, Lund), by designation
of Aubert, 19766:269.

Labels. Norl. [printed] ; brevitarsis m [Thomson cabinet label].
Identity. Anoncus brevitarsis (Thomson).

Mesoleius (Lamachus) castaneiventris, 1894: 2023. ? Syntypes 2 9, SWEDEN: Vestergotland [ =
Vastergotland] (UZI, Lund).

Labels. Col. Hgn. [printed] ; castaneiventris [Thomson cabinet label] (1 9)- Col. Hgn. [printed] (1 $).
The two females originating from Holmgren's collection are the only specimens under this name. They
have no indication of locality but they may well be syntypes.
Identity. Lamachus castaneiventris (Thomson).

Mesoleius (Mesoleius) clypealis, 1894: 2077. Lectotype 9, SWEDEN: 'Halland' [Skane], Margretetorp (UZI,
Lund), by designation of Aubert, 1966: 127.
Label. Hall, [printed].

Margretetorp is in northern Skane, not southern Halland as stated by Thomson. It is near the bound-
ary between the two provinces, so Thomson's error is easily explained.
Identity. Campodorus clypealis (Thomson).

Mesoleius (Saotus) compressiusculus, 1883: 934. Syntypes 1 9, 3 <J, SWEDEN: Skane, Ringsion[= Ringsjon]
(UZI, Lund).

Labels, [small green square]; 146. [hand]; compressiusculus [Thomson cabinet label] (1 9)- [small
green square] (3 <$).

Identity. Saotis compressiusculus (Thomson).

Mesoleius (Spudaeus) confusus, 1883: 932. Syntypes 4 9, 3 cJ, SWEDEN: Skane, Ringsjon and Lindholmen
(UZI, Lund).

Labels, [small green square]; 9 [printed]; confusus [Thomson cabinet label] (1 9)- [small green
square]; 9 [printed] (2 9). Lhn 29/5 [hand]; 82. [hand] (1 rf). Lhn 12/6 [hand]; 81. [hand](l J). Scan
[printed] (1 <^). [small green square] (1 $).

Identity. Rhinotorus confusus (Thomson) comb. n.

Mesoleius (Mesoleius) crassipes, 1894: 2060. Lectotype 9, SWEDEN: Norrland (UZI, Lund), by designation
of Aubert, 19766:270.

Labels. Col. Rud. [hand] ; crassipes [Thomson cabinet label].
Identity. Campodorus crassipes (Thomson).

56 M. G. FITTON

Mesoleius (Mesoleius) crassitarsis, 1883: 935. Type(s) 9, SWEDEN: Skane, Ryssjoholm [= Rossjoholm]
(lost).

The male from Palsjo (Thomson, 18886: 1262) published as lectotype by Aubert (19766: 270) cannot be
a syntype. The lectotype designation is therefore invalid. The female mentioned by Aubert is from
Skanes-Fagerhult and, also, cannot be a syntype.

Identity. Campodorus crassitarsis (Thomson) (Aubert, 19766: 270, on the basis of the invalid 'lectotype').

Mesoleius (Mesoleius) curtitarsis, 1894: 2038. Lectotype $, SWEDEN: Ostergotland (UZI, Lund), by
designation of Aubert, 19766: 270.

Labels. 86. [hand]; OG. [hand] ; curtitarsis [Thomson cabinet label].
Identity. Campodorus curtitarsis (Thomson).

Mesoleius (Mesoleius) deletus, 1894: 2069. Lectotype 9, SWEDEN: 'Halland' [Skane], Margretetorp (UZI,
Lund), by designation of Aubert, 19766: 270.

Label. Hall [printed].

Margretetorp is in northern Skane, not Halland as stated by Thomson. It is near the boundary between
the two provinces, so Thomson's error is easily explained.

Identity. Campodorus deletus (Thomson).

Mesoleius (Scopesus) depressus, 1894: 2030. Holotype 9, SWEDEN (UZI, Lund).
Labels. Col Ljgh [printed] ; depressus [Thomson cabinet label].
Identity. Scopesis depressus (Thomson).

Mesoleius (Lagarotus) didymus, 1894: 2024. Type(s) 9, GERMANY (WEST): Bavaria (lost).

There are no specimens under this name in the Thomson collection and Diller (pers. comm.) has been
unable to trace it in the Kriechbaumer collection in Munich.
Identity. ? Lagarotis didymus (Thomson).

Mesoleius (Saotus) dorsatus, 18886: 1264. Holotype 9, SWEDEN : Skane, Palsjo (UZI, Lund).
Labels. Pal [hand] ; dorsatus [Thomson cabinet label].
Identity. Saotis dorsatus (Thomson).

Mesoleius (Saotus) emarginatus, 1883: 933. Syntypes 29, 1 <$, SWEDEN: Skane, Ortofta (UZI, Lund).

Labels, pil [hand]; Ort. [hand]; emarginatus [Thomson cabinet label] (1 9)- Ort. [hand] (1 9)- Ort.
[hand] ; £ [printed] (1 <$).
Identity. Saotis emarginatus (Thomson).

Mesoleius (Mesoleius) femorator, 1894: 2047. Lectotype 9, SWEDEN: Norrland (UZI, Lund), by designation
of Aubert, 19766:271.
Label. Col. Rui [hand].
Identity. Anoncus femorator (Thomson).

Mesoleius (Barytarbus) flavicornis, 18926: 1875. Holotype 9, FRANCE (UZI, Lund).
Labels. FT. merid.l [hand] ; Gall [hand] ; flavicornis [Thomson cabinet label].
Identity. Mesolept idea flavicornis (Thomson) comb. n.

Mesoleius (Perispudus)flavitarsis, 1894: 2023. Holotype cJ, FRANCE: Libercourt (UZI, Lund).
Labels. Libercourt. [hand] ; Gallia [printed] ; flavitarsis [Thomson cabinet label].
Identity. Perispuda flavitarsis (Thomson).

Mesoleius (Barytarbus)flavoscutellatus, 18926: 1876. Holotype <£ FRANCE: Lapugnoy (UZI, Lund).
Labels. Lapugnoy. [hand]; Gall [hand] ; flavoscutellat [Thomson cabinet label].
Identity. Barytarbesflavoscutellatus (Thomson).

Mesoleius (Mesoleius) frenalis, 1894: 2047. Lectotype 9, SWEDEN: Skane, Palsjo (UZI, Lund), by des-
ignation of Aubert, 19766: 271.
Label. Pal [hand].
Identity. Mesoleius frenalis Thomson.

Mesoleius (Mesoleius) frontatus, 1894: 2069. Syntype 1 (J, SWEDEN: Skane, Ystad (UZI, Lund).

Label. Ys [hand].

The female specimen published as lectotype by Aubert (19766: 271) is from Ostergotland and cannot be
a syntype. It is labelled 'OG', which Aubert misread (upside down) as '50'. The lectotype designation is
therefore invalid.

Identity. Mesoleius frontatus Thomson.

THE ICHNEUMONIDAE DESCRIBED BY C. G. THOMSON 57

Mesoleius (Mesoleius) galticus, 1894: 2041. Lectotype 9, FRANCE: Phalempin (UZI, Lund), by designation
of Aubert, 19766:272.

Labels. Phalempin. [hand]; Gall [hand] ; Gallicus m [Thomson cabinet label].
Identity. Campodorus gallicus (Thomson).

Mesoleius (Mesoleius) glyptus, 1894: 2076. Lectotype 9, SWEDEN: Skane, Palsjo (UZI, Lund), by des-
ignation of Aubert, 19766: 272.
Label. Pal [hand].
Identity. Campodorus glyptus (Thomson).

Mesoleius (Protarchus) grandis, 18886: 1260. Lectotype 9, SWEDEN: Skane (UZI, Lund), by designation of
Aubert, 1966: 127.

Labels. Sk. [hand] ; grandis [Thomson cabinet label].
Identity. Protarchus grandis (Thomson).

Mesoleius (Saotus) heteropus, 1883: 934. LECTOTYPE 9, SWEDEN : Lappland (UZI, Lund), here designated
(selected by R. Hinz).

Labels. Lpl. [printed] ; heteropus [Thomson cabinet label].
Identity. Saotis heteropus (Thomson).

Mesoleius (Mesoleius) humerellus, 1894: 2042. Lectotype 9, SWEDEN: Skane, Lund (UZI, Lund), by des-
ignation of Aubert, 19766: 272.
Label. L-d [printed].
Identity. Campodorus humerellus (Thomson).

Mesoleius (Otlophorus) hypomelas, 1894: 2027. LECTOTYPE 9, GERMANY (UZI, Lund), here designated
(selected by R. Hinz).
Label. 6. 296 [hand].
Identity. Otlophorus hypomelas (Thomson).

Mesoleius (Mesoleius) immarginatus, 1894: 2037. Lectotype 9, SWEDEN: Skane, Ringsjon (UZI, Lund), by
designation of Aubert, 19766: 272.
Labels. Rsio [printed] ; imarginatus [hand].
Identity. ? Mesoleius immarginatus Thomson.

Mesoleius (Mesoleius) incident, 1894: 2077. Lectotype 9 [not^ as stated by Aubert, 19766: 272], SWEDEN:
Skane, Molle (UZI, Lund), by designation of Aubert, 19766: 272.
Label. M61 [hand].
Identity. Campodorus incidens (Thomson).

Mesoleius (Mesoleius) incisus, 1894: 2064. Holotype 9, SWEDEN: Norrland (UZI, Lund).
Labels. Col. Rud. [hand] ; incisus m [Thomson cabinet label].
Identity. Mesoleius incisus Thomson.

Mesoleius (Mesoleius) laevipectus, 1894: 2041. Lectotype ^, SWEDEN: Skane, Molle (UZI, Lund), by
designation of Aubert, 19766: 273.
Label. M61 [hand].
Identity. Campodorus laevipectus (Thomson).

Mesoleius (Barytarbus) laeviusculus, 1883: 931. LECTOTYPE <J, SWEDEN: Oland (UZI, Lund), here
designated (selected by H. K. Townes).

Labels. 0. [printed] ; laeviusculus [Thomson cabinet label].

This is the specimen regarded as holotype by Aubert (1972: 147). However, it cannot be a holotype
because Thomson gave a range of length and must, therefore, have had a syntype series. The specimen was
labelled as lectotype by Townes, and not holotype as implied by Aubert. Aubert also interpreted the
locality label '0.' as meaning Ortofta (which, if correct, would have excluded the specimen from type
status!).

Identity. Barytarbes laeviusculus (Thomson).

Mesoleius (Mesoleius) latiscapus, 1894: 2060. Lectotype 9, SWEDEN: Vestergothland [= Vastergotland]
(UZI, Lund), by designation of Aubert, 1 9766 : 273.

Labels. V.G. [printed] ; latiscapus [Thomson cabinet label].
Identity. Campodorus latiscapus (Thomson).

Mesoleius (Saotus) liopleuris, 18886: 1263. LECTOTYPE 9, SWEDEN: Skane, Palsjo (UZI, Lund), here
designated (selected by R. Hinz).

58 M. G. FITTON

Labels. Pal. [hand] ; liosternus [Thomson cabinet label].

Saotus liosternus (Thomson, 1894: 2018) is an incorrect subsequent spelling of liopleuris, and has no
status in nomenclature.

Identity. Saotis liopleuris (Thomson).

Mesoleius (Mesoleius) liosternus, 1894: 2078. Syntypes 9 <?, SWEDEN: Jemtland [= Jamtland], Areskutan
(lost).

Aubert's publication of a neotype female (1976ft: 273) for this species is not valid because it does not
comply with the provisions of Article 75(c) of the Code. No attempt is made to validate that 'neotype' here
because there has been no recent revisionary work on this group.

Identity. Campodorus liosternus (Thomson) (Aubert, 19766: 273, on the basis of the invalid 'neotype').

Mesoleius (Mesoleius) lobatus, 1894: 2072. Lectotype $, SWEDEN: Skane, Ryssjoholm [= Rossjoholm]
(UZI, Lund), by designation of Aubert, 19766: 273.

Labels. Rshm 16/6 [hand] ; lobatus m [Thomson cabinet label].
Identity. Campodorus lobatus (Thomson).

Mesoleius (Scopesus) longigena, 1894: 2031. Syntypes 3 9, 3 <$, NORWAY: Dovre and FRANCE: Libercourt
and Ostricourt (UZI, Lund).

Labels. Libercourt [hand]; longigena [Thomson cabinet label] (1 £). Ostricourt [hand] (2 ?). Liber-
court [hand] (2 £). [small paper square]; Dovre. [printed] (1 $).

Identity. Neostroblia longigena (Thomson) comb. n.

Mesoleius (Saotus) longiventris, 18886: 1263. Holotype 9, SWEDEN: Skane, Ortofta (UZI, Lund).
Labels. Ort 28/V [hand] ; longiventris [Thomson cabinet label].
Identity. Saotis longiventris (Thomson).

Mesoleius (Lamachus) longiventris, 1894: 2023. Holotype 9, SWEDEN (UZI, Lund).
Labels. Col. Hgn. [printed] ; longiventris [Thomson cabinet label].

Identity. Junior primary homonym of Mesoleius longiventris Thomson, 18886. Replacement name here
proposed Lamachus thomsoni nom. n.

Mesoleius (Scopesus) macropus, 1894: 2030. Syntypes 2 <£, SWEDEN : Skane, Ringsjon (UZI, Lund).

Labels, [small green square] (2 £).

Aubert's publication of a neotype (1966: 127) for this species is, fortunately, not valid because it does
not comply with the provisions of Article 75(c) of the Code. The discovery of syntypes does not, therefore,
need a reference to the Commission (Article 75(f)).

Identity. Scopesis macropus (Thomson).

Mesoleius (Otlophorus) melanocarus, 1894: 2027. Holotype 9, GERMANY (UZI, Lund).
Labels. 5 992 [hand] ; nigrifrons [Thomson cabinet label].
Identity. Otlophorus melanocarus (Thomson).

Mesoleius (Protarchus) melanurus, 1894: 2020. Type(s) [? sex], GERMANY (WEST): Harz (lost).
Identity. Protarchus melanurus (Thomson).

Mesoleius (Perispudus) mesoxanthus, 1894: 2022. Holotype 9, FRANCE: Vosges (UZI, Lund).

Labels, [small paper square]; Vosges. [hand]; Gall, [hand]; mesoxanthus [Thomson cabinet
label].

Identity. I have not been able to satisfactorily place this species in any of the genera (as defined by
Townes, 19706) of the Mesoleiini.

Mesoleius (Mesoleius) nemati, 1894: 2067. Lectotype 9, DENMARK : S0nderborg (UZI, Lund), by designation
of Aubert, 19766:274.

Labels. Sdbg 25.5.81 [hand] ; Nematus [hand, first word illegible] ; 774 [printed].

Identity. Campodorus nemati (Thomson).

Mesoleius (Saotus) nigriscuta, 18886: 1264. Holotype 9, SWEDEN : Skane, Palsjo(UZI, Lund).
Labels. Pal. [hand] ; nigriscuta [Thomson cabinet label].
Identity. Saotis nigriscuta (Thomson).

Mesoleius (Mesoleius) obliquus, 1894: 2070. Syntype 1 <$, SWEDEN: 'Halland' [Skane], Magretetorp (UZI,
Lund).

Labels. Halland [printed] ; obliquus [Thomson cabinet label].
See note on locality under Mesoleius deletus.

THE ICHNEUMONIDAE DESCRIBED BY C. G. THOMSON 59

Aubert (19766: 274) recognised this specimen as a holotype, but Thomson specified 'J1 9'; showing that
he had a syntype series.

Identity. Mesoleius obliquus Thomson.

Mesoleius (Mesoleius) or bit alls. 1894: 2050. Lectotype 9, NORWAY: Forsa (UZI, Lund), by designation of
Aubert, 19766:275.

Labels. Forssa 24 Juli [hand]; Lpl. [printed] [not 'Lap' as stated by Aubert]; orbitalis [Thomson
cabinet label].

Identity. Junior synonym ofHyperbatus segmentator (Holmgren) (Aubert, 19766: 275).

Mesoleius (Mesoleius) picticoxa, 1894: 2072. Lectotype 9, GERMANY (WEST): Bavaria (UZI, Lund), by
designation of Aubert, 19766: 275.

Labels. Germ, [hand]; = 67 [hand].

The specimen designated as lectotype does not agree precisely with the original description in the
colour of the coxae. There is therefore the possibility that it is not an original specimen. There are no other
specimens in the collection which could be types and Diller (pers. comm.) has been unable to trace the
species in the Kriechbaumer collection in Munich.

Identity. Mesoleius picticoxa Thomson.

Mesoleius (Mesoleius) pineti, 1894: 2071. Lectotype 9, SWEDEN: Skane, Fagelsang (UZI, Lund), by
designation of Aubert, 19766: 275.
Labels. Fsg 12/7 [hand]; 9 [printed].

The lectotype was selected and labelled by K. Horstmann, not Townes as indicated by Aubert.
Identity. Campodorus pineti (Thomson).

Mesoleius (Mesoleius) pleuralis, 1894: 2076. Lectotype 9, SWEDEN: Skane, Palsjo (UZI, Lund), by des-
ignation of Aubert, 19766: 275.

Labels. Pal. [hand] ; 9 [printed] ; pleuralis [Thomson cabinet label].
Identity. Campodorus pleuralis (Thomson).

Mesoleius (Lathiponus) pule her rimus, 18886: 1261. LECTOTYPE 9, SWEDEN: Skane, Palsjo (UZI, Lund)
here designated (selected by H. K. Townes).
Labels. Pal [hand] ; pulcherrimus [Thomson cabinet label].
Identity. Junior synonym of Lathiponus frigidus (Woldstedt) (Townes, 19706: 86).

Mesoleius (Mesoleius) rubidus, 1883: 935. Lectotype 9, NORWAY: Dovre (UZI, Lund), by designation of
Aubert, 19766:276.
Labels, none.
Identity. Mesoleius rubidus Thomson.

Mesoleius (Mesoleius) sinuatus, 1894: 2040. Lectotype 9, SWEDEN: Skane, Palsjo (UZI, Lund), by
designation of Aubert, 19766: 276.

Labels. Pal [hand] ; sinuatus [Thomson cabinet label].
Identity. Mesoleius sinuatus Thomson.

Mesoleius (Mesoleius) stenostigma, 1894: 2042. Holotype 9, SWEDEN : Norrland (UZI, Lund).
Labels. Col. Rui [hand] ; stenostigma [Thomson cabinet label].
Identity. Mesoleius stenostigma Thomson.

Mesoleius (Mesoleius) subroseus, 18886: 1262. Lectotype 9, SWEDEN: Skane, Palsjo (UZI, Lund), by
designation of Aubert, 19766: 277.
Label. Pal. [hand].
Identity. Mesoleius subroseus Thomson.

Mesoleius (Scopesus) tegular is. 1894: 2031. Lectotype 9, SWEDEN: Stockholm (UZI, Lund), by designation
of Townes, Momoi & Townes, 1965: 259.
Labels. Him [printed] ; DeV [printed].
Identity. Scopesis tegularis (Thomson).

Mesoleius (Mesoleius) tenuitarsis. 1894: 2039. Holotype 9, SWEDEN: Lappland (UZI, Lund).

Labels, [square of paper]; [square of paper]; Lap [hand] ; tenuitarsis m [Thomson cabinet label].
Identity. Campodorus tenuitarsis (Thomson).

Mesoleius (Saotus) tricolor, 1883: 933. Syntypes 3 9, SWEDEN: Skane, Lund (UZI, Lund).

Labels. L-d [printed] ; tricolor [Thomson cabinet label] (1 9). Ld [hand](l 9). L-d [printed] (1 9).
Identity. Saotis tricolor (Thomson).

60 M. G. FITTON

Mesoleius (Mesoleius) varicoxa, 1894: 2044. Lectotype $, SWEDEN: Skane, Ringsjon (UZI, Lund), by
designation of Aubert, 1976ft: 278.
Label. Rsio [printed].
Identity. Mesoleius varicoxa Thomson.

Mesoleptus (Mesoleptus) holmgreni, 1894: 1982. Syntypes 3 9, 7 <£ SWEDEN: Skane, Palsjo (UZI, Lund).
Labels. Pal [hand] ; Holmgreni [Thomson cabinet label] (1 9). Pal. [hand] (2 9 7 £, on 7 pins).
Identity. Mesoleptidea holmgreni (Thomson) comb. n.

Mesoleptus (Hadrodactylus) nigricoxa, 1894: 1979. Lectotype 9, DENMARK: Satrupholz (UZI, Lund), by
designation of Idar, 1973: 24.
Label. Satruph. 15.VI.93 [hand].

Idar gave the original combination incorrectly as Hadrodactylus nigricoxa.
Identity. Junior synonym of Hadrodactylus femoralis (Holmgren) (Idar, 1975: 184).

Mesoleptus (Hadrodactylus) varicoxa, 1894: 1979. Lectotype <£ SWEDEN: Skane, Ryssjoholm [ =
Rossjoholm] (UZI, Lund), by designation of Idar, 1973 : 24.
Label. Rshm 16/6 [hand].

Idar gave the original combination incorrectly as Hadrodactylus varicoxa.
Identity. Junior synonym of Hadrodactylus insignis (Kriechbaumer) (Idar, 1975: 187).

Mesostenus crassifemur, 18886: 1237. Lectotype 9, SWEDEN: Skane, Kjeflinge [= Kavlinge] (UZI, Lund), by
designation of Aubert, 1966: 128.

Labels. Scan [printed] ; crassifemur m [Thomson cabinet label].
Identity. Mesostenus crassifemur Thomson.

Mesostenus (Stenaraeus) dentifer, 1896: 2381. Syntypes 3 9, 1 <$, SWEDEN : Skane, Degeberga (UZI, Lund).
Labels. Dg [hand] ; dentifer Thorns [Thomson cabinet label] (1 9). Dg [hand] (2 9 1 rf).
Identity. Mesostenus dentifer Thomson.

Mesostenus (Mesostenus) subcircularis, 1896: 2379. Holotype 9, SWEDEN: Vermland [= Varmland] (UZI,
Lund).

Labels. Wml [printed] ; subcircularis Ths [hand].
Identity. Junior synonym of Mesostenidea obnoxius (Gravenhorst) (Horstmann, 1968: 121).

Mesostenus subovalis, 1873: 516. Lectotype 9, SWEDEN: Skane, Palsjo (UZI, Lund), by designation of
Horstmann, 1968: 121.

Labels. Pal [hand] ; subovalis [Thomson cabinet label].
Identity. Junior synonym of Mesostenidea obnoxius (Gravenhorst) (Horstmann, 1968: 121).

Metopius (Metopius) brevispina, 1887ft: 195. Syntypes 3 9, SWEDEN: Skane, Ringsjon and Ronnemolla (UZI,
Lund).

Labels. Ron [hand] ; brevispina [Thomson cabinet label] (1 9)- Ringsio [printed] (2 $).
Identity. Metopius brevispina Thomson.

Metopius (Metopius) clypealis, 1887ft: 196. Holotype <$, GERMANY (UZI, Lund).
Labels. Germ Ichn. [hand] ; clypealis [Thomson cabinet label].
Identity. Metopius clypealis Thomson.

Metopius (Peltocarus) croceicornis, 1887ft: 196. Syntype 1 <$, ? syntype 1 9, GERMANY and ? SWEDEN:
Gotland (UZI, Lund).

Labels, [square of paper] ; Germ Ichn. [hand] (<J). [square of red paper] ; [square of greyish paper] ;
324. [hand] ; croceicornis [Thomson cabinet label] (9).
Identity. Metopius croceicornis Thomson.

Metopius (Peltocarus) interruptus, 1887ft: 197. Syntype 1 3, SWEDEN : Smaland Markaryd (UZI, Lund).
Labels. Mark 13/6 [hand] ; interruptus [Thomson cabinet label].
Identity. Metopius interruptus Thomson.

Microcryptus alutaceus, 1883: 863. Syntypes 2 $, SWEDEN : Norrland (UZI, Lund).
Labels. Norl. [printed] (2 (J).

The specimen published by Aubert (1972: 148) as lectotype almost certainly came from Smaland (as
indicated by him!) and therefore cannot be a syntype. The lectotype designation is thus invalid.
Identity. ? Pleolophus alutaceus (Thomson). The two syntypes are not conspecific.

THE ICHNEUMONIDAE DESCRIBED BY C. G. THOMSON 61

Microcryptus areolaris, 1883 : 858. Syntypes 1 $, 2 cJ, SWEDEN: Skane, Loparod and Yddinge (UZI, Lund).
Labels. Lop [hand] (1 $). Yd [hand] (2 <J).
Identity. Javra areolaris (Thomson) comb. n.

Microcryptus aries, 1883: 851. Syntypes 5 ?, 2 <£, SWEDEN: Lappland; Norrland; Smaland; and Skane,
Ryssioholm [ = Rossjoholm] (UZI, Lund).

Labels. Smoland [printed] ; Aries [Thomson cabinet label] (1 $). Smoland [printed] (2 $). Rhm [hand]
(1 (J). Norl. [printed] (1 $). [square of paper] ; Smoland [printed] (1 $). Lap [hand] ; Alcis [hand] (1 ?).

Identity. Schenkia aries (Thomson) comb. n.

Microcryptus borealis, 1883: 862. Syntypes 1 9, 1 $, SWEDEN: Lappland (UZI, Lund).
Labels. Lpl. [printed] ; borealis [Thomson cabinet label] ($). Lpl. [printed] (^).
Identity. Aptesis borealis (Thomson) comb. n.

Microcryptus distant, 1883: 864. Lectotype 9, SWEDEN: Oland (UZI, Lund), by designation of Aubert,
1966: 129.

Labels. 0. [printed] ; distans [Thomson cabinet label].
Identity. Aptesis distans (Thomson).

Microcryptus femor -alls, 1883: 853. Lectotype 9, SWEDEN: Norrland (UZI, Lund), by designation of Aubert,
1972: 148.

Labels. Norl. [printed] ; femoralis [Thomson cabinet label].
Identity. Aptesis femoralis (Thomson).

Microcryptus gravenhorsti, 1883: 854. Syntype 1 $, SWEDEN: Skane, Ringsion [= Ringsjon] (UZI, Lund).
Label. Rsio [printed].
Identity. Polytribax gravenhorsti (Thomson) comb. n.

Microcryptus lapponicus, 1883: 862. Syntypes 2 9, SWEDEN: Lappland (UZI, Lund).

Labels, [square of paper] ; [square of paper] ; Lpl. [printed] ; Lapponicus [Thomson cabinet label]
(1 9). Lap [hand] (19).

Identity. Aptesis lapponica (Thomson) comb. n.

Microcryptus longicauda, 1883: 862. LECTOTYPE 9, SWEDEN: Lappland, Lycksele (UZI, Lund), here
designated (selected by J. F. Aubert).

Labels. Lycksele Lapp [hand] ; longicauda [Thomson cabinet label].
Identity. Cubocephalus longicauda (Thomson) comb. n.

Microcryptus nigricornis, 1883 : 860. Holotype 9, SWEDEN : Skane, Lund (UZI, Lund).
Labels. Lund [printed] ; nigricornis [Thomson cabinet label].
Identity. Oresbius nigricornis (Thomson) comb. n.

Microcryptus nigrit ulus, 1885: 23. Syntypes 9 d1, FRANCE (lost).
This species is not present in the collection.

Identity. Aptesis nigritula (Thomson) (on the basis of specimens in the BMNH collection determined as
this species by Schmiedeknecht).

Microcryptus opaculus, 1883 : 851. Type(s) 9, SWEDEN : Skane, Wittsio [ = Vittsjo] (lost).
Identity. Schenkia opacula (Thomson).

Microcryptus orbitalis, 1883: 856. Syntypes 1 9, 1 cT, SWEDEN: Skane, Lund and Ringsion [= Ringsjon]
(UZI, Lund).

Labels. LD 28/6 ['Lo' printed, date hand] (9). [small green square] ($).
Identity. Aptesis orbitalis (Thomson) comb. n.

Microcryptus ornaticeps, 1885: 23. Type(s) 9, FRANCE: Paris (lost).
This species is not present in the collection.
Identity. Unknown, the name remains a nomen dubium.

Microcryptus pectoralis, 1888ft: 1237. Syntypes 1 9, 1 <$, SWEDEN : Skane, Palsjo (UZI, Lund).
Labels. Pal. [hand] ; pectoralis [Thomson cabinet label] (9). Pal. [hand] (c?).
Identity. Aptesis pectoralis (Thomson) comb. n.

Microcryptus puncticoltis, 1883: 866. LECTOTYPE £, SWEDEN: Skane, Ringsjon (UZI, Lund), here
designated (selected by J. F. Aubert).
Label. Rsio [printed].
Identity. Aptesis puncticollis (Thomson) comb. n.

62 M. G. FITTON

Microcryptus punctifer, 1883: 860. Lectotype 9, SWEDEN: Skane, Lund (UZI, Lund), by designation of
Aubert, 1966: 129.

Labels. L-d [printed]; punctifer [Thomson cabinet label].
Identity. Oresbius punctifer (Thomson) comb. n.

Microcryptus rubricollis, 1883: 853. Holotype?, NORWAY: Lillehammer (UZI, Lund).

Labels. Lhmr. 25.6.77 ['Lhmr.', printed ; date, hand]; 24 [hand] ; rubricollis [Thomson cabinet label].
Identity. Schenkia rubricollis (Thomson).

Microcryptus septentrionalis, 1883 : 863. Holotype 9, SWEDEN : Lappland (UZI, Lund).
Labels. Norl. [printed] ; septentrionalis [Thomson cabinet label].
Identity. Oresbius septentrionalis (Thomson) comb. n.

Miomeris glabriventris, 18886: 1317. Syntypes9 <J, SWEDEN : Skane, Yddinge (lost).

Identity. Microleptes glabriventris (Thomson) comb. n. (on the basis of material in the collection).

Miomeris rectangulus, 18886: 1317. LECTOTYPE & FRANCE: Bar-s-Seine (UZI, Lund), here designated
(selected by J. F. Aubert).

Labels. Cartereau Bar-s-Seine [printed] ; Gallia [printed] ; rectangulus <£. [hand] [on reverse of cabinet
label: 'Aquisgrana'].

The type-locality is in northern rather than southern France as stated by Thomson. Presumably he
made a mistake, if he did not the specimen cannot be a type.

Identity. Microleptes rectangulus (Thomson) comb. n.

Monoblastus angulatus, 18886: 1256. Syntypes 3 9, 3 J. SWEDEN : Skane, Palsjo (UZI, Lund).

Labels. Pal. [hand]; angulatus [Thomson cabinet label] (1 9). Pal. [hand]; 9 [printed] (1 9). Pal.
[hand] (1 9, 1 <J). Palsio [printed] (1 £). Pal. [hand];^ [printed] (1 £).
Identity. Rhorus angulatus (Thomson).

Monoblastus longigena, 1883: 903. LECTOTYPE 9, SWEDEN: Lappland (UZI, Lund), here designated
(selected by R. Hinz).

Labels. Lpl. [printed] ; longigena [Thomson cabinet label].
Identity. Rhorus longigena (Thomson).

Nemeritis caudatula, 1887c: 1119. Lectotype 9, [? locality] (UZI, Lund), by designation of Aubert,
• 1972: 149.

Labels, f. 584 [hand] ; caudatula [Thomson cabinet label].
Identity. Nemeritis caudatula Thomson.

Nemeritis convergens, 1887c: 1 120. LECTOTYPE 9, RUMANIA : Tasnad (UZI, Lund), here designated (selec-
ted by K. Horstmann).

Labels. Tasnad 1 1.5.83 [locality, printed; date, hand] ; convergens [Thomson cabinet label].
Identity. Cymodusa convergens (Thomson).

Nemeritis lativentris, 1887c: 1119. Lectotype 9, SWEDEN: Gotland (UZI, Lund), by designation of Horst-
mann, 1973a: 11.
Label. G [hand].
Identity. Nemeritis lativentris Thomson.

Nemeritis stenura, 1887c: 1119. Lectotype 9, SWEDEN: Oland (UZI, Lund), by designation of Aubert,
1972: 149.

Labels. 0. [printed] ; stenura [Thomson cabinet label].
Identity. Nemeritis stenura Thomson.

Nepiesta marginella, 1887c: 1117. Lectotype 9, SWEDEN: Ostergotland (UZI, Lund), by designation of
Aubert, 1968: 195.

Labels. 153. [hand] ; OG. [hand] ; marginella [Thomson cabinet label].

Identity. Junior synonym of Biolysia immolator (Gravenhorst) (Horstmann, 1974a: 78). Townes
(19706: 164) places Biolysia as a synonym of Bathyplect es.

Nepiesta subclavata, 1887c: 1116. Lectotype 9, FRANCE: Mt Noir (UZI, Lund), by designation of Horst-
mann, 1973c: 737.

Labels. Mt. Noir. [hand] ; subclavata [Thomson cabinet label].
Identity. Nepiesta subclavata Thomson.

THE ICHNEUMONIDAE DESCRIBED BY C. G. THOMSON 63

Notopygus mordax, 1883 : 925. Holotype ?, SWEDEN : Smaland (UZI, Lund).
Labels. Smol [printed] ; mordax [Thomson cabinet label].
Identity. Xenoschesis mordax (Thomson).

Notopygus (Homaspis) robustus, 1894: 1984. Holotype ?, POLAND : Silesia (UZI, Lund).
Labels. Silesia [hand] ; robusta [Thomson cabinet label].
The holotype lacks the gaster.
Identity. Homaspis robustus (Thomson).

Notopygus (Homaspis) varicolor, 1894: 1984. Holotype?, POLAND: Silesia (UZI, Lund).
Labels. Silesia Becker [hand] ; varicolor [Thomson cabinet label].
Identity. Homaspis varicolor (Thomson).

Nyxeophilus nigricornis, 1885: 18. Type(s) ?, FRANCE (lost).
This species is not present in the collection.
Identity. Xylophrurus nigricornis (Thomson) comb. n.

Odontomerus pinetorum, 1877: 777. Lectotype ?, SWEDEN: Vastergotland (UZI, Lund), by designation of
Townes, Momoi & Townes, 1965 : 1 19.
Label. V.G. [printed].
Identity. Junior synonym of Odontocolon dentipes (Gmelin) (Townes, Momoi & Townes, 1965 : 119).

Odontomerus punctulatus, 1877: 777. Holotype?, SWEDEN : Smaland (UZI, Lund).

Labels. Coll. L-gh. [printed] ; punctulatus n [hand] ; gracilis [Thomson cabinet label].
Identity. Odontocolon punctulatum (Thomson).

Odontomerus quercinus, 1877: 777. Lectotype ?, SWEDEN: Oland (UZI, Lund), by designation of Townes,
Momoi & Townes, 1965: 120.
Label. O. [printed].
Identity. Odontocolon quercinum (Thomson).

Oedimopsis [lapsus for Oedemopsis] limbata, 1883: 907. Holotype?, SWEDEN : Skane, Esperod [= Asperod]
(UZI, Lund).

Labels. Esp [printed] ; limbata [Thomson cabinet label].
Identity. Oedemopsis limbata Thomson.

Olesicampa alboplica, 1887c: 1141. Syntype 1 <$, SWITZERLAND (UZI, Lund).
Labels. 84.6. 599 [hand] ; alboplica [Thomson cabinet label].

This specimen is not a holotype (Type unique') as stated by Aubert (1966: 130). Thomson gives a range
of length. Other syntypes may be in Kriechbaumer's collection.
Identity. Olesicampe alboplica (Thomson).

Olesicampa basalis, 1887c: 1 143. Holotype ?, SWEDEN : Smaland, Kalmar (UZI, Lund).
Labels. Sm [hand] ; ? [printed] ; basalis [Thomson cabinet label].
Identity. Olesicampe basalis (Thomson).

Olesicampa binotata, 1887c: 1141. LECTOTYPE ?, GERMANY (WEST): Aachen (UZI, Lund), here des-
ignated (selected by R. Hinz).

Label. Germ [hand].

The lectotype lacks the gaster. It was not missing when the lectotype was examined by Hinz in 1954
(R. Hinz, pers. comm.).

Identity. Olesicampe binotata (Thomson).

Olesicampa cavigena, 1887c: 1140. Lectotype ?, SWEDEN: Skane, Torringe (UZI, Lund), by designation of
Aubert, 1966: 130.

Labels. Tor [hand] ; ? [printed] ; cavigena [Thomson cabinet label].
Identity. Olesicampe cavigena (Thomson).

Olesicampa crassitarsis, 1887c: 1146. LECTOTYPE ?, SWEDEN: Skane, Ortofta (UZI, Lund), here des-
ignated (selected by R. Hinz).

Labels. Ort. [hand] ; crassitarsis [Thomson cabinet label].
Identity. Olesicampe crassitarsis (Thomson).

Olesicampa femorella, 1887c: 1144. LECTOTYPE ?, SWEDEN: Skane, Esperod [= Asperod] (UZI, Lund),
here designated (selected by R. Hinz).

Labels. Fall, [hand] ; femorator [hand] ; femorella [Thomson cabinet label].
Identity. Olesicampe femorella (Thomson).

64 M. G. FITTON

Olesicampa flavicornis, 1887c: 1 143. LECTOTYPE 9, SWEDEN: Skane, Palsjo (UZI, Lund), here designated
(selected by R. Hinz).
Label. Palsio [printed].
Identity. Olesicampeflavicornis (Thomson).

Olesicampafulcrans, 1887c: 1 145. LECTOTYPE 9, SWEDEN: Skane, Asperod (UZI, Lund), here designated
(selected by R. Hinz).

Labels. Esp 20/6 [hand] ; fulcrans [Thomson cabinet label].
Identity. Olesicampe fulcrans (Thomson).

Olesicampa geniculella, 1887c: 1 144. Holotype 9, SWEDEN : Smaland, Kalmar (UZI, Lund).
Labels. Kalm. [printed] ; geniculella [Thomson cabinet label].
Identity. Olesicampe geniculella (Thomson).

Olesicampa gracilipes, 1887c: 1143. LECTOTYPE 9, SWEDEN: Norrland (UZI, Lund), here designated
(selected by R. Hinz).

Labels. Norl. [printed] ; 9 [printed].
Identity. Olesicampe gracilipes (Thomson).

Olesicampa luteipes, 1887c: 1147. LECTOTYPE $, SWEDEN: ? Blekinge (UZI, Lund), here designated
(selected by R. Hinz).

Labels. Col Ljgh [printed]; $ [printed] ; luteipes [Thomson cabinet label].
Identity. Olesicampe luteipes (Thomson).

Olesicampa nigricoxa, 1887c: 1145. LECTOTYPE 9, SWEDEN: Skane, Ringsjon (UZI, Lund), here des-
ignated (selected by R. Hinz).

Labels. Scan [printed] ; 9 [printed] ; nigricoxa [Thomson cabinet label].
Identity. Olesicampe nigricoxa (Thomson).

Olesicampa nigroplica, 1887c: 1143. LECTOTYPE 9, GERMANY (UZI, LunJ), here designated (selected by
R. Hinz).

Labels. I. 80 5.8 [hand] ; nigroplica [Thomson cabinet label].
Identity. Olesicampe nigroplica (Thomson).

Olesicampa patellana, 1887c: 1140. LECTOTYPE 9, FRANCE (UZI, Lund), here designated (selected by
R. Hinz).

Labels. 9 [printed] ; Gall. [hand].
Identity. Olesicampe patellana (Thomson).

Olesicampa punctitarsis, 1887c: 1146. LECTOTYPE 9, GERMANY (WEST): Bavaria (UZI, Lund), here
designated (selected by R. Hinz).

Labels. 84. 537. [hand] ; punctitarsis [Thomson cabinet label].
Identity. Olesicampe punctitarsis (Thomson).

Olesicampa radiella, 1887c: 1147. LECTOTYPE 9, SWEDEN: Norrland (UZI, Lund), here designated (selec-
ted by R. Hinz).

Labels. Norl. [printed]; 9 [printed].
Identity. Olesicampe radiella (Thomson).

Olesicampa return, 1887c: 1144. Lectotype 9, SWEDEN: Skane, Yddinge (UZI, Lund), by designation of
Aubert, 1972: 149.

Labels. Yd. [hand] ; retusa [hand] ; retusa [Thomson cabinet label].
Identity. Olesicampe retusa (Thomson).

Olesicampa simplex, 1887c: 1147. Lectotype 9, SWEDEN: Skane, Palsjo (UZI, Lund), by designation of
Aubert, 1966: 130.
Label. Pal [hand].
Identity. Olesicampe simplex (Thomson).

Olesicampa sternella, 1887c: 1146. Lectotype 9, SWEDEN: Skane, Palsjo (UZI, Lund), by designation of
Aubert, 1972: 149.
Label. Pal [hand].
Identity. Olesicampe sternella (Thomson).

Olesicampa subcallosa, 1887c: 1146. Lectotype 9, SWEDEN: Skane, Alnarp (UZI, Lund), by designation of
Aubert, 1972: 149.

THE ICHNEUMONIDAE DESCRIBED BY C. G. THOMSON 65

Labels. Alp. [hand] ; subcallosa [Thomson cabinet label].
Identity. Olesicampe subcallosa (Thomson).

Omorga angulata, 1887c: 1129. Lectotype 9, SWEDEN: Skane, Kungsmarken (UZI, Lund), by designation of
Aubert, 1966: 130.

Label. Kgsm 10/7 [hand].

Identity. Campoplex angulatus (Thomson).

Omorga biloba, 1887c: 1126. Lectotype 9, SWEDEN: Skane, Palsjo (UZI, Lund), by designation of Aubert,
1972: 149.
Label. Pal [hand].
Identity. Campoplex bilobus (Thomson).

Omorga continua, 1887c: 1132. Lectotype ?, SWEDEN: Gotland (UZI, Lund), by designation of Aubert,
1966: 130.

Labels. G. [hand] ; continua [Thomson cabinet label].
Identity. Campoplex continuus (Thomson).

Omorga coracina, 1887c: 1130. Lectotype 9, SWEDEN: Skane, Fogelsang [= Fagelsang] (UZI, Lund), by
designation of Jussila, 1965: 81.

Labels. Fogelsang [printed] ; coracina [Thomson cabinet label].
Identity. Campoplex coracinus (Thomson).

Omorga exoleta, 1887c: 1127. Lectotype cJ, SWEDEN: Skane, Ryssjoholm [= Rossjoholm] (UZI, Lund), by
designation of Townes, Momoi & Townes, 1965: 275.
Labels. Rhm [hand] ; exoleta [Thomson cabinet label].
Identity. Tranosema exolet a (Thomson) (Horstmann, 1977: 77).

Omorga forticosta, 1887c: 1131. Lectotype 9, SWEDEN: Skane, Palsio [= Palsjo] (UZI, Lund), by des-
ignation of Aubert, 1966: 130.
Label. Pal [hand].
Identity. Campoplex forticosta (Thomson).

Omorga fusciplica, 1887c: 1127. Lectotype 9, SWEDEN : Skane, Lund (UZI, Lund), by designation of Aubert,
1968: 195.

Labels. L-d [printed] ; fusciplica [Thomson cabinet label].
Identity. Campoplex fusciplica (Thomson).

Omorga fusicornis, 1887c: 1132. LECTOTYPE 9, GERMANY (WEST): Aachen (UZI, Lund), here designated
(selected by K. Horstmann).

Labels. Germ [printed] ; fusicornis [Thomson cabinet label].
Identity. Campoplex fusicornis (Thomson) comb. n.

Omorga hadrocera, 1887c: 1134. Lectotype 9, GERMANY (WEST): Aachen (UZI, Lund), by designation of
Aubert, 1966: 130.

Labels. 19/8 38 [hand]; Germ. [hand].
Identity. Campoplex hadrocerus (Thomson).

Omorga liogaster, 1887c: 1 130. Type(s) 9, SWEDEN: Dalsland (lost).

The specimen recognised byAubert (1966: 130) as holotype (Type unique') cannot be a type because it
comes from Bohuslan (label 'Bohl.'), as stated by Aubert !
Identity. Campoplex liogaster (Thomson) (Aubert, 1966: 130, on the basis of the supposed type).

Omorga litorea, 1887c: 1134. LECTOTYPE 9, SWEDEN: Skane, Lomma (UZI, Lund), here designated
(selected by K. Horstmann).

Labels. Loma 20/7 [hand] ; litorea [Thomson cabinet label].
Identity. Campoplex litoreus (Thomson).

Omorga lyrata, 1887c: 1128. Lectotype 9, SWEDEN : Skane, Ringsjon (UZI, Lund), by designation of Aubert,
1966: 130.

Label. Scan [printed].
Identity. Campoplex lyratus (Thomson).

Omorga melampus, 1887c: 1131. LECTOTYPE 9, SWEDEN: Skane, Degeberga (UZI, Lund), here designated
(selected by R. Hinz).

Labels. Dgb. [hand] ; melampus [Thomson cabinet label].
Identity. Campoplex melampus (Thomson) comb. n.

66 M. G. FITTON

Omorga nigridens, 1887c: 1 130. Lectotype 9, SWEDEN: Skane, Kungsmarken (UZI, Lund), by designation of
Horstmann, 1977: 78.

Labels. Kgsm 10/7 [hand] ; nigridens [Thomson cabinet label].
Identity. Tranosema nigridens (Thomson) (Horstmann, 1977: 78).

Omorga picticrus, 1887c: 1128. Lectotype 9, SWEDEN: Skane, Lund (UZI, Lund), by designation of Horst-
mann, 1977: 75.

Labels. Scan [printed] ; picticrus [Thomson cabinet label].

Identity. Junior synonym of Campoplex cerophagus Gravenhorst (Horstmann, 1969: 421). Horstmann
(1977) separates Sesioplex (including cerophagus) from Campoplex.

Omorga ruficoxa, 1887c: 1 127. Lectotype 9, HUNGARY [? CZECHOSLOVAKIA] (UZI, Lund), by designation of
Aubert, 1968: 195.

Labels. -Ujhely. 13.6 [locality, printed ; date, hand] ; ruficoxa [Thomson cabinet label].

The locality on the label is probably Satoraljaujhely, which is on the border between Czechoslovakia
and Hungary. Its Czechoslovakian name is Slovenske Nove Mesto.

Identity. Campoplex ruficoxa (Thomson).

Omorga scaposa, 1887c: 1128. LECTOTYPE 9, SWEDEN: Skane, Palsjo (UZI, Lund), here designated
(selected by R. Hinz).

Labels. Pal. [hand] ; scaposa [Thomson cabinet label].
Identity. Campoplex scaposus (Thomson) comb. n.

Omorga striolata, 1887c: 1131. Lectotype <$, SWEDEN: Skane, Ringsjon (UZI, Lund), by designation of
Aubert, 1966: 130.
Label. Rsio [printed].
Identity. Junior synonym of Tranosema nigridens (Thomson) (Horstmann, 1977: 78).

Ophion (Ophion) distant, 18886: 1191. Lectotype 9, SWEDEN: Stockholm (UZI, Lund), by designation of
Townes, Momoi & Townes, 1965: 316.

Labels. Him. [printed]; Musko'n 8.1885 [hand]; Mortu [hand];9 [hand]; 7. [hand].
Identity. Ophion distans Thomson.

Ophion (Ophion) longigena, 1888b: 1191. Lectotype <$, SWEDEN: Skane, Torringelund (UZI, Lund), by
designation of Aubert, 1972: 148.

Label. Tn [hand, ? first letter].

The label on the lectotype is certainly not 'Sn' as stated by Aubert (1972: 148) and Sn was not used by
Thomson as an abbreviation for Skane, as far as is known.

Identity. Ophion longigena Thomson.

Ophion (Ophion) scutellaris, 1888ft: 1192. Lectotype 9, SWEDEN : Goteborg (UZI, Lund), by designation of
Aubert, 1972: 148.

Labels. Gbg [hand] ; scutellaris [Thomson cabinet label].
Identity. Ophion scutellaris Thomson.

Orthocentrus (Stenomacrus) compressus, 1897: 2436. Holotype 9, SWEDEN : Skane, Palsjo (UZI, Lund).
Labels. Hbg. [hand] ; compressus Ths. [Thomson cabinet label].
Identity. Neurateles compressus (Thomson) comb. n.

Orthocentrus (Stenomacrus) crassicornis, 1897: 2434. Holotype 9, GERMANY (UZI, Lund).
Labels. Mdsk. 21.V.84. [hand]; 28-39 [hand].
Identity. Neurateles crassicornis (Thomson) comb. n.

Orthocentrus (Stenomacrus) cubiceps, 1897: 2447. LECTOTYPE 9, SWEDEN: Skane, Palsjo (UZI, Lund),
here designated (selected by J. F. Aubert).
Label. Pal. [hand].
Identity. Stenomacrus cubiceps (Thomson).

Orthocentrus (Stenomacrus) curvulus, 1897: 2443. LECTOTYPE 9, SWEDEN: Skane, Fogelsang [ =
Fagelsang] (UZI, Lund), here designated (selected by J. F. Aubert).
Labels. Fg 556 [hand] ; 9 [printed] ; curvulus [Thomson cabinet label].
Identity. Stenomacrus curvulus (Thomson).

Orthocentrus (Stenomacrus) deletus, 1897: 2442. LECTOTYPE 9, SWEDEN: Skane, Ortofta (UZI, Lund),
here designated (selected by J. F. Aubert).

THE ICHNEUMONIDAE DESCRIBED BY C. G. THOMSON 67

Labels. Ort. [hand] ; deletus [Thomson cabinet label].
Identity. Stenomacrus deletus (Thomson).

Orthocentrus (Stenomacrus) exserens, 1897: 2448. Lectotype 9, SWEDEN: Skane, Yddinge (UZI, Lund), by
designation of Aubert, 1968: 195.
Labels. Yd. [hand] ; exserens m [Thomson cabinet label].
Identity. Stenomacrus exserens (Thomson).

Orthocentrus (Stenomacrus) falcatus, 1897: 2435. LECTOTYPE 9 SWEDEN: Skane, Ringsjon (UZI, Lund),
here designated (selected by J. F. Aubert).

Labels. Rsio [printed] ; $ [printed] ; falcatus [Thomson cabinet label].
Identity. Neurateles falcatus (Thomson) comb. n.

Orthocentrus (Stenomacrus) flavicornis, 1897: 2439. Type(s) 9, SWEDEN : Ostergotland (lost).
Identity. ? Leipaulus flavicornis (Thomson) comb. n.

Orthocentrus (Stenomacrus) fortipes, 1897: 2442. LECTOTYPE 9, SWEDEN: Skane, Palsjo (UZI, Lund),
here designated (selected by J. F. Aubert).
Label. Palsio [printed].
Identity. Stenomacrus fortipes (Thomson).

Orthocentrus (Stenomacrus) innotatus, 1897: 2449. Lectotype 9, SWEDEN: Skane, Degeberga (UZI, Lund),
by designation of Aubert, 1968: 195.

Labels. Deg. [hand] ; Deg. [hand] ; innotatus [Thomson cabinet label].
Identity. Stenomacrus innotatus (Thomson).

Orthocentrus (Orthocentrus) petiolaris, 1897: 2428. Lectotype 9, SWEDEN : Skane, Ringsjon (UZI, Lund), by
designation of Aubert, 1968: 195.

Labels. Rsio [printed] ; 9 [printed] ; petiolaris m [Thomson cabinet label].
Identity. Orthocentrus petiolaris Thomson.

Orthocentrus (Orthocentrus) radialis, 1897: 2430. Lectotype 9, SWEDEN: Skane, Ortofta (UZI, Lund), by
designation of Aubert, 1978a: 24.
Label. Ortofta [printed].
Identity. Orthocentrus radialis Thomson.

Orthocentrus (Picrostigeus) recticauda, 1897: 2431. LECTOTYPE 9, SWEDEN: Jemtland [= Jamtland],
Areskutan (UZI, Lund), here designated (selected by J. F. Aubert).
Labels. Norl. [printed] ; anomalus H [Thomson cabinet label].
Identity. Picrostigeus recticauda (Thomson).

Orthocentrus (Stenomacrus) superus, 1897: 2443. LECTOTYPE 9, SWEDEN: Skane, Trelleborg (UZI,
Lund), here designated (selected by J. F. Aubert).
Labels. Tbg 9/76 [hand] ; superus [Thomson cabinet label].
Identity. Stenomacrus superus (Thomson).

Orthocentrus (Stenomacrus) ungula, 1897: 2436. Holotype 9, SWEDEN : Skane, Palsjo (UZI, Lund).
Labels. Pal [hand] ; Ungula Ths [Thomson cabinet label].
Identity. Stenomacrus ungula (Thomson).

Oxytorus armatus, 1883: 910. Lectotype 9, SWEDEN: Skane, Palsio [= Palsjo] (UZI, Lund), by designation
ofKerrich, 1939: 127.
Label. Pal. [hand].
Identity. Oxytorus armatus Thomson.

Pachymerus puncticeps, 1877: 734. Lectotype 9, SWEDEN: Skane, Lund (UZI, Lund), by designation of
Townes, Momoi & Townes, 1965: 398.
Label. Lund [printed].

Identity. Junior synonym of Collyria coxator (Villers) (Townes, Momoi & Townes, 1965: 397,
398).

Pachymerus trichophthalmus, 1877: 734. Lectotype & SWEDEN: Skane, Ringsjon (UZI, Lund), by designation
of Aubert, 1966: 127.

Label, [small green square].

Identity. Collyria trichophthalma (Thomson).

68 M. G. FITTON

Paniscus brachycerus, 1888fc : 1201. Lectotype ?, SWEDEN: Skane, Ilstorp (UZI, Lund), by designation of
Townes, Momoi & Townes, 1965 : 87.

Labels. lisp 28/6 [hand] ; brachycerus [Thomson cabinet label].
Identity. Junior synonym of Netelia dilatatus (Thomson) (Delrio, 1975: 47).

Paniscus dilatatus, 18886: 1200. Lectotype ? [not (J as stated by Aubert, 1972: 146], SWEDEN: Skane,
Degeberga (UZI, Lund), by designation of Aubert, 1972: 146.
Labels. Deg [hand] ; dilatatus [Thomson cabinet label].
Identity. Netelia dilatatus (Thomson).

Paniscus gracilipes, 1888ft: 1201. Lectotype £, SWEDEN: Skane, Palsjo (UZI, Lund), by designation of
Townes, Momoi & Townes, 1965: 88.
Label. Pal [hand].
Identity. Junior synonym of Netelia fuscicornis (Holmgren) (Delrio, 1975: 51).

Paniscus melanurus, 18886: 1199. Lectotype <£, SWEDEN: Skane, Palsjo (UZI, Lund), by designation of
Townes, Momoi & Townes, 1965 : 89.
Label. Pal [hand].
Identity. Netelia melanurus (Thomson).

Paniscus ocellaris, 18886: 1199. Lectotype ?, SWEDEN: Ostergothland [= Ostergotland], (UZI, Lund), by
designation of Townes, Momoi & Townes, 1965: 90.

Labels. O.G. Bh [hand] ; ocellaris m [hand] ; ocellaris [Thomson cabinet label].
Identity. Netelia ocellaris (Thomson).

Paniscus opaculus, 18886: 1199. Lectotype 9, SWEDEN: Skane, Lindholmen (UZI, Lund), by designation of
Townes, Momoi & Townes, 1965: 92.
Labels. Lhn 23/7 [hand]; 360. [hand].
Identity. Netelia opaculus (Thomson).

Parabatus cristatus, 18886: 1197. Lectotype 9, SWEDEN: Skane, Palsjo (UZI, Lund), by designation of
Townes, Momoi & Townes, 1965: 96.
Label. Pal [hand].
Identity. Netelia cristatus (Thomson).

Parabatus latungula, 18886: 1196. Lectotype 9, SWEDEN: Skane, Palsjo (UZI, Lund), by designation of
Townes, Momoi & Townes, 1965: 98.
Label. Pal [hand].
Identity. Netelia latungulus (Thomson).

Parabatus nigricarpus, 18886: 1196. Lectotype $, SWEDEN: Skane, Ringsjon (UZI, Lund), by designation of
Delrio, 1975: 39.

Labels. Pal [hand] ; nigricarpus [Thomson cabinet label].
Identity. Netelia nigricarpus (Thomson).

Peritissus (Ecclinops) albitarsis, 1883: 914. Syntypes 1 9, 2 <J, SWEDEN: Skane, Ringsion [= Ringsjon]
(UZI, Lund).

Labels, [small green square] (1 9 1 J). [small green square]; 330. [hand](l ^).
Identity. Perilissus albitarsis Thomson.

Perilissus (Ecclinops) compressus, 1883: 914. Holotype 9, SWEDEN: Skane, Sofdeborg [= Sovdeborg] (UZI,
Lund).

Labels. Sbg 28/7 [hand] ; compressus [Thomson cabinet label].
Identity. Perilissus compressus Thomson.

Perilissus (Spanotecnus) coxalis, 1883: 912. Syntype 1 $, SWEDEN : Skane, Arrie (UZI, Lund).
Label. Ar [hand].
Identity. Perilissus coxalis Thomson.

Perilissus (Ecclinops) emarginatus, 1883: 914. Syntype 1 & SWEDEN : Skane, Ringsion [= Ringsjon] (UZI,
Lund).

Labels, [small green square]; 186. [hand].
Identity. Perilissus emarginatus Thomson.

Perilissus (Ecclinops) frontator, 1883: 914. Type(s) [? sex]. SWEDEN : Skane, Holmeja (lost).
Identity. Perilissus frontator Thomson.

THE ICHNEUMONIDAE DESCRIBED BY C. G. THOMSON 69

Perilissus (Polyoncus) grandiceps, 1883: 913. Lectotype c?, SWEDEN: Skane, Arrie (UZI, Lund), by des-
ignation of Aubert, 1966: 127.

Labels. Ar 6/56 [hand] ; grandiceps [Thomson cabinet label].
Identity. Lathrolestes grandiceps (Thomson).

Perilissus (Luphyroscopus) nigricollis, 1883: 915. Syntypes 3 $, 3^, SWEDEN: Skane, Helsingborg, Ortofta,
Arrie and Palsjo (UZI, Lund).

Labels. Hbg. [hand] ; nigricollis [Thomson cabinet label] (1 9). Ort. [hand] (2 9 1 <$). Ar. [hand](l cJ).
Pal. [hand] (1 <£).

Identity. Lathrolestes nigricollis (Thomson) comb. n.

Perilissus (Perilissus) spiniger, 1883: 912. Holotype^, SWEDEN : Skane, Holmeja(UZI, Lund).
Label. Yd [hand].
Identity. Perilissus spiniger Thomson.

Pezomachus (Pezomachus) breviceps, 1884: 1017. Syntypes 9 <$, SWEDEN : Skane, Skanor (lost).
Identity. Gelis breviceps (Thomson).

Pezomachus (Pezomachus) gonatopinus, 1884: 1008. Lectotype $, SWEDEN: Skane, Palsjo (UZI, Lund), by
designation of Aubert, 1972: 148.
Label Pal [hand].

Aubert incorrectly referred to the species as Gelis gonatopinus.
Identity. Gelis gonatopinus (Thomson).

Pezomachus (Pezomachus) grandiceps, 1884: 1007. Syntype 1 <$, ? syntype 1 9, SWEDEN : Skane, Degeberga
(UZI, Lund).

Labels. Deg [hand] ; grandiceps [Thomson cabinet label] (^). 1 72. [hand] ; Scania [printed] ($).
The female specimen is only tentatively recognised as a syntype because it lacks precise locality data.
Identity. Gelis grandiceps (Thomson).

Pezomachus (Pezomachus) mandibularis, 1884: 1009. Syntypes 14 $, 10 $, SWEDEN: Skane, Bokeberg,
Helsingborg, Lindholmen, Palsjo and Yddinge (UZI, Lund).
No notes were made of individual specimen labels.
Identity. Gelis mandibularis (Thomson).

Pezomachus (Pezomachus) myrmecinus, 1884: 1001. Syntypes 11 9, 1 c?, SWEDEN : Skane, Skanor, Lund,
Lomma and Fagelsang; and Oland (UZI, Lund).
No notes were made of individual specimen labels.
Identity. Gelis myrmecinus (Thomson).

Pezomachus numidicus, 1885: 32. Type(s) 9, ALGERIA (lost).

Horstmann (1979a: 297) has searched for this species in the Fairmaire collection in Paris, but without
success.

Identity. Unknown, the name remains a nomen dubium.

Pezomachus (Pezolochus) pilosulus, 1884: 1003. LECTOTYPE 9, SWEDEN : Skane, Lund (UZI, Lund), here
designated (selected by K. Horstmann).
Labels. Lund [printed] ; pilosulus [Thomson cabinet label].
Identity. Gelis pilosulus (Thomson) comb. n.

Pezomachus (Pezomachus) spinula, 1884: 1006. Syntype 1 <$, SWEDEN : Skane, Lund (UZI, Lund).
Labels. $ [printed]; L-d [printed].
Identity. Gelis spinulus (Thomson).

Phaeogenes (Phaeogenes) crassidens, 1891: 1644. Syntypes 2 9, SWEDEN: Skane, Ryssjoholm [ =
Rossjoholm] (UZI, Lund).

Labels. Rshm 16/6 [hand] (1 9). Rhm [hand](l 9).
Identity. Phaeogenes crassidens Thomson.

Phaeogenes (Proscus) elongatus, 1891: 1651. Lectotype 9, DENMARK: Senderborg (UZI, Lund), by des-
ignation of Aubert, 1966: 128.

Labels. Sandb'g. 19.5.86 [hand] ; elongatus m [Thomson cabinet label].
Identity. Phaeogenes elongatus Thomson.

70 M. G. FITTON

Phaeogenes ( Phaeogenes ) montanus, 1891: 1652. Holotype?, CZECHOSLOVAKIA: Altvater [= PradSd] (UZI,
Lund).

Labels. Altvater [hand] ; montanus m [Thomson cabinet label].
Identity. Phaeogenes montanus Thomson.

Phaeogenes (Phaeogenes) ruficoxa, 1891: 1648. Lectotype 9, SWEDEN: Skane, Ringsjon (UZI, Lund), by
designation of Townes, Momoi & Townes, 1965: 416.
Label. Ringsio [printed].

A second lectotype designation (of another specimen) published by Aubert (1966: 128) is invalid.
Identity. Dirophanes ruficoxa (Thomson).

Phaeogenes (Phaeogenes) tegularis, 1891 : 1656. Holotype 9, SWEDEN: Lappland (UZI, Lund).
Labels. Lap [hand] ; tegularis [Thomson cabinet label].
Identity. Phaeogenes tegularis Thomson.

Phaestus heterocerus, 1894: 2017. LECTOTYPE 9, SWEDEN: Skane, Palsjo (UZI, Lund), here designated
(selected by H. K. Townes).

Labels. Pal. [hand] ; heterocerus [Thomson cabinet label].

Identity. Junior synonym of Phaestus anomalus (Brischke) (Townes, 19706: 74).

Phobetus (Phobetus) femorator, 1894: 1986. Lectotype 9, DENMARK: Senderborg (UZI, Lund), by des-
ignation of Aubert, 1966: 127.
Label. Sandb'g. VI.91. [hand].
Identity. Phobetes femorator (Thomson).

Phobetus (Ipoctonus) fulviventris, 1894: 1987. Holotype?, SWEDEN : Bohuslan (UZI, Lund).
Labels. Bohl. [printed] ; fulviventris [Thomson cabinet label].
Identity. Phobetes fulviventris (Thomson) comb. n.

Phobetus (Ipoctonus) latipes, 1894: 1987. Holotype^, SWEDEN : Skane, Kjellby[= Kallby] (UZI, Lund).
Labels. Kallby 2 Jul. 30. [hand] ; latipes [hand].
Identity. Phobetes latipes (Thomson) comb. n.

Phobetus (Ipoctonus) rufipes, 1894: 1987. Syntypes 9 <$, SWEDEN: Skane, Palsjo (lost).

Identity. Phobetes rufipes (Thomson) comb. n. (on the basis of specimens in the collection).

Phobocampa alticollis, 1887c: 1121. Lectotype 9, GERMANY (UZI, Lund), by designation of Aubert, 1968:
195.

Labels. Germ [hand] ; alticollis [Thomson cabinet label].
Identity. Phobocampe alticollis (Thomson).

Phobocampa confusa, 1887c: 1122. ? Syntype 1 9, ? GERMANY (UZI, Lund).

Label. Kalnh. 27/6. 84. [hand].

This specimen is probably a syntype but it has not been possible to discover the meaning of the locality
abbreviation 'Kalnh.' (the last letter could be 'p'). The species was described from Germany. The only
other specimen in the collection is from Loos (near Lille in France) and cannot therefore be a syntype.

Identity. Phobocampe confusa (Thomson).

Phobocampa flavicincta, 1887c: 1122. LECTOTYPE 9, SWEDEN: Skane, Ringsjon (UZI, Lund), here des-
ignated (selected by R. W. Carlson).
Label. Scan lac [printed].
Identity. Phobocampe flavicincta (Thomson).

Phobocampa pulchella, 1887c: 1121. LECTOTYPE 9, SWEDEN: Skane, Fogelsang [=Fagelsang] (UZI,
Lund), here designated (selected by R. W. Carlson).
Label. Lund [printed].
Identity. Phobocampe pulchella (Thomson).

Phygadeuon acutipennis, 1884: 954. Syntypes 9 cJ, SWEDEN : Skane, Stehag (lost).
Identity. ? Phygadeuon acutipennis Thomson (Frilli, 1973: 95).

Phygadeuon annulicornis, 1884: 947. Lectotype 9, SWEDEN: Skane, Ringsjon (UZI, Lund), by designation of
Aubert, 1972: 148.

Labels, [small green square] ; annulicornis [Thomson cabinet label].
Identity. Theroscopus annulicornis (Thomson) (Frilli, 1973 : 95).

THE ICHNEUMONIDAE DESCRIBED BY C. G. THOMSON 71

Phygadeuon anurus, 1884: 946. Lectotype 9, SWEDEN: Skane, Ortofta (UZI, Lund), by designation of Frilli,
1973:95.

Labels. Ortofta [printed] ; 9 [printed] ; anurus [Thomson cabinet label].
Identity. Ceratophygadeuon anurus (Thomson).

Phygadeuon armatulus, 18886: 1240. Lectotype 9, SWEDEN: Skane, Fogelsang [= Fagelsang] (UZI, Lund),
by designation of Frilli, 1973: 96.

Labels. Fogelsang [printed] ; armatulus n [Thomson cabinet label].
Identity. Medophron armatulus (Thomson).

Phygadeuon bidens, 1884: 958. Lectotype 9, SWEDEN: Skane, Ringsion [= Ringsjon] (UZI, Lund), by des-
ignation of Frilli, 1973: 96.

Labels, [small green square] ; bidens [Thomson cabinet label].
Identity. Phygadeuon bidens Thomson.

Phygadeuon brachyurus, 1884: 955. Lectotype 9, SWEDEN: Skane, Ortofta (UZI, Lund), by designation of
Jussila, 1965: 141.
Label. Ort. [hand].
Identity. Phygadeuon brachyurus Thomson.

Phygadeuon brevitarsis, 1884: 959. Lectotype 9, SWEDEN: Skane, Ringsion [= Ringsjon] (UZI, Lund), by
designation of Frilli, 1973: 97.
Label. Rsio [printed].
Identity. Phygadeuon brevitarsis Thomson.

Phygadeuon canaliculatus, 1889: 1406. Syntypes 9 3, SWEDEN : Skane, Palsjo (lost).
Identity. ? Phygadeuon canaliculatus Thomson (Frilli, 1973: 97).

Phygadeuon caudatus, 1884: 946. Lectotype 9, SWEDEN: Lappland (UZI, Lund), by designation of Frilli,
1973:97.

Labels. Lap [hand] ; caudatus [Thomson cabinet label].

Identity. Junior secondary homonym of Medophron caudatus (Provancher). Replacement name
Medophron caudatulus (Dalla Torre).

Phygadeuon cubiceps, 1884: 961. Syntypes 9, SWEDEN : Skane, Torringe (lost).

The specimen designated as lectotype by Horstmann (1967a: 15) (and also recognised as such by
Aubert (1968: 195) and Frilli (1973: 97)) is from Yddinge (label 'Yd'). Although Thomson often 'bracketed'
together adjacent localities it seems to be unlikely that he would have done so in this case, especially as he
states 'vid Torringe nara Malmo'. Torringe and Yddinge are about 6 km apart. Therefore, I do not think
the 'lectotype' can have been a syntype, unless Thomson misquoted the locality.

Identity. Phygadeuon cubiceps Thomson.

Phygadeuon curviscapus, 1889: 1405. Holotype 9, SWEDEN : Skane, Palsjo (lost).

The female specimen designated as lectotype by Frilli (1973: 98) is on the same pin as a male specimen.
Since Thomson specified 'Ett exemplar' (that is, a holotype) it seems highly unlikely that the specimen
concerned was on the same pin as a male. Therefore, I do not think the 'lectotype' can be the original
specimen.

Identity. Phygadeuon curviscapus Thomson (on the basis of the invalid 'lectotype').

Phygadeuon curvispina, 1884: 948. Lectotype cJ, SWEDEN: Skane, Lund (UZI, Lund), by designation of Frilli,
1973:98.

Label. Lund [printed].

I do not regard the lectotype designation published by Aubert (1966: 129) as valid because he failed to
indicate (in both his publication and labels attached to the specimen) which syntype was selected. There
are five specimens (and the remains of a sixth) on one pin. From the top of the pin the specimens are: 1,
paralectotype <J; 2, remains (head) of a paralectotype [? sex]; 3, lectotype <$; 4, paralectotype 9; 5,
paralectotype <$; 6, paralectotype cJ.

Identity. Stibeutes curvispina (Thomson).

Phygadeuon dimidiatus, 1884: 963. Lectotype 9, SWEDEN: Skane, Klinta (UZI, Lund), by designation of
Frilli, 1973:98.

Labels, [small green square] ; dimidiatus [Thomson cabinet label].
Identity. Phygadeuon dimidiatus Thomson.

72 M. G. FITTON

Phygadeuon facialis, 1884: 952. Lectotype 9, SWEDEN: Skane, Esperod [= Asperod] (UZI, Lund), by des-
ignation of Frilli, 1973: 98-99.

Labels. Esp [printed] ; facialis [Thomson cabinet label].

Identity. Junior primary homonym of Phygadeuon facialis Gravenhorst. Replacement name Thero-
scopusfaciator (Aubert) (Frilli, 1973: 98).

Phygadeuon flavicans, 1884: 961. Lectotype J, SWEDEN : Skane, Lund (UZI, Lund), by designation of Aubert,
1966: 129.
Label Ld [hand].
Identity. Phygadeuon flavicans Thomson.

Phygadeuon flavipes, 18886: 1238. Lectotype 9, SWEDEN: Skane, Ortofta (UZI, Lund), by designation of
Frilli, 1973:99.

Label. Ort [hand].

Identity. Junior secondary homonym of Phygadeuon flavipes (Provancher) (described in Mesostenus
and currently placed in Grypocentrus). Replacement name Medophronflavitarsis (Dalla Torre).

Phygadeuon grandiceps, 1884: 950. Lectotype ^ [not 9 as stated by Townes, Momoi & Townes, 1965: 145],
SWEDEN: Skane, Palsio [= Palsjo] (UZI, Lund), by designation of Townes, Momoi & Townes, 1965:
145.

Label. Pal [hand].

Identity. Phygadeuon grandiceps Thomson.

Phygadeuon grandis, 1884: 940. Lectotype 9, SWEDEN: Norrland (UZI, Lund), by designation of Frilli, 1973:
100.

Label. Norl. [printed].
Identity. Pygocryptus grandis (Thomson).

Phygadeuon heterogaster, 1885 : 22. Type(s) 9, FRANCE (lost).
Identity. ? Phygadeuon heterogaster Thomson.

Phygadeuon heteropus, 1896: 2387. Lectotype 9, SWEDEN: Skane, Palsjo (UZI, Lund), by designation of
Frilli, 1973: 100.
Label. Pal [hand].
Identity. Dichrogaster heteropus (Thomson).

Phygadeuon inflates, 1884: 959. Lectotype 9, SWEDEN: Skane, Lund (UZI, Lund), by designation of Townes,
Momoi & Townes, 1965: 145.

Label. L-d [printed].

Identity. Junior secondary homonym of Phygadeuon inftatus (Provancher) (described in Ichneumon and
currently placed in Endasys). Replacement name Phygadeuon infelix Dalla Torre. Use of this replacement
name would be contrary to existing usage (which is Phygadeuon inflatus (e.g. Horstmann, 1967a: 10)),
which should be maintained pending reference to the International Commission under Article 59(bXO of
the Code (as amended, Bull. zool. Norn. 31 (1974): 83).

Phygadeuon laeviventris, 1884: 955. Lectotype 9, SWEDEN: Skane, Lund (UZI, Lund), by designation of
Jussila, 1965:141.

Labels. Ld [hand] ; laeviventris [Thomson cabinet label].
Identity. Phygadeuon laeviventris Thomson.

Phygadeuon lapponicus, 1884: 952. Lectotype 9, SWEDEN: Lappland (UZI, Lund), by designation of Jussila,
1965:143.
Label. Lap [hand].
Identity. Phygadeuon lapponicus Thomson.

Phygadeuon Kogaster, 1884: 949. Lectotype 9, NORWAY (UZI, Lund), by designation of Frilli, 1973: 101.
Labels. 205 [hand] ; Norv [hand] ; liogaster [Thomson cabinet label].
Identity. Phygadeuon liogaster Thomson.

Phygadeuon liosternus, 1884: 1040. Syntype 1 <J, SWEDEN : Skane, Ortofta (UZI, Lund).
Label. Ort. [hand].

Frilli (1973: 101-102) considered that the male from Ortofta did not agree with the original description
and could not, therefore, be a syntype. I believe that there is sufficient agreement for it to be a syntype.
Identity. Phygadeuon liosternus Thomson.

THE ICHNEUMONIDAE DESCRIBED BY C. G. THOMSON 73

Phygadeuon longiceps, 1884: 946. Syntype 1 3, SWEDEN : Skane, Lund (UZI, Lund).

Label. L-d 14/6 [hand].

Frilli (1973: 102) considered that the male recognised here as a syntype was not in agreement with the
original description and he designated a neotype 9 for this species. If Frilli's neotype designation is
considered to fulfil the provisions of Article 75 of the Code and to be 'valid' then the case must be re^rred
to the International Commission (Article 75(f)).

Identity. Ceratophygadeuon longiceps (Thomson).

Phygadeuon longigena, 1884: 947. Lectotype $, SWEDEN: Skane (UZI, Lund), by designation of Frilli,
1973:102.

Labels. Scan [printed] ; longigena [Thomson cabinet label].
Identity. Phygadeuon longigena Thomson.

Phygadeuon monodon, 1884: 950. Syntype 1 9, SWEDEN: Skane, near Hellestad [= Hallestad] (UZI, Lund).

Label. Dahlby [hand].

Dahlby [ = Dalby] is not far from Hallestad and assuming that the specimen was not collected in the
town itself it seems reasonable to suppose that it is a syntype. Aubert's publication of a 'neotype' for this
species (1966: 129) is not valid because it does not comply with the provisions of Article 75(c) of the
Code.

Identity. Phygadeuon monodon Thomson.

Phygadeuon ochrogaster, 18886: 1241. Syntypes 9, SWEDEN: Skane, Ryssjoholm [= Rossjoholm] (lost).

The specimen designated as lectotype by Frilli (1973: 103) cannot be a syntype — it is from Kungsmar-
ken near Lund and is labelled 'Kgsm' not 'Rysm' as stated by Frilli.

Identity. Theroscopus ochrogaster (Thomson) (Frilli, 1973: 103, on the basis of the invalid 'lecto-
type').

Phygadeuon ocularis, 1889: 1405. Lectotype $, SWEDEN: Skane, Palsjo (UZI, Lund), by designation of
Horstmann, 1967a: 11.

Labels. Pal [hand] ; ocularis m [Thomson cabinet label].
Identity. Phygadeuon ocularis Thomson.

Phygadeuon oppositus, 1884: 960. Lectotype 9, SWEDEN : Skane, Lund (UZI, Lund), by designation of Jussila,
1965: 142.

Label. Lund [printed].
Identity. Phygadeuon oppositus Thomson.

Phygadeuon ovalis, 1884: 963. Lectotype 9, SWEDEN: Skane, Stehag (UZI, Lund), by designation of Aubert,
1966: 129.

Labels, [green square] ; ovalis [Thomson cabinet label].

Identity. Junior primary homonym of Phygadeuon ovalis Provancher. Replacement name Phygadeuon
ovaliformis Dalla Torre.

Phygadeuon pallicarpus, 1884: 947. Lectotype 9, SWEDEN : Skane, Fogelsang [= Fagelsang] (UZI, Lund), by
designation of Frilli, 1973: 104.

Labels. Fsg 18/5 [hand] ; pallicarpus [Thomson cabinet label].

Frilli (1973: 104) misspelled the name pallidicarpus. This unjustified emendation was first used by Dalla
Torre (1902: 690).

Identity. Phygadeuon pallicarpus Thomson.

Phygadeuon parvicauda, 1885: 20. Syntypes 9, FRANCE: Marchiennes (lost).

Aubert's publication of a neotype female (1966: 129) for this species is not valid because it does not
comply with the provisions of Article 75(c) of the Code.

Identity. Junior synonym of Ceratophygadeuon anurus (Thomson) (Horstmann, 19796: 45).

Phygadeuon parvipennis, 1884: 944. Lectotype (J, SWEDEN: Skane, Lund (UZI, Lund), by designation of
Frilli, 1973:104.

Label. Lund [printed].

Identity. Arotrephes parvipennis (Thomson).

Phygadeuon pimplarius, 1884: 941. Lectotype 9, SWEDEN: Skane, Ofvedskloster [= Ovedskloster] (UZI,
Lund), by designation of Frilli, 1973: 105.
Label. Oke A [hand].
Identity. Lochetica pimplaria (Thomson).

74 M. G. FITTON

Phygadeuon punctigena, 1884: 953. Lectotype ?, SWEDEN: Skane, Alnarp (UZI, Lund), by designation of
Frilli, 1973: 105.

Labels. Alp [hand] ; punctigena [Thomson cabinet label].
Identity. Phygadeuon punctigena Thomson.

Phygadeuon punctipleuris, 1884: 962. Lectotype ?, SWEDEN: Skane, Palsio [= Palsjo] (UZI, Lund), by
designation of Frilli, 1973: 105.
Label. Pal [hand].
Identity. Phygadeuon punctipleuris Thomson.

Phygadeuon punctiventris, 1884: 955. Lectotype 9, SWEDEN: Skane, Klinta (UZI, Lund), by designation of
Aubert, 1966: 129.

Label, [small green square].

I do not agree with Frilli that an earlier publication by Aubert (1965: 564) constitutes a valid lectotype
designation.

Identity. Phygadeuon punctiventris Thomson.

Phygadeuon recurvus, 1884: 943. Syntypes ?, SWEDEN : Skane, Klinta (lost).

The specimen designated as lectotype by Frilli (1973: 106) is from Herrevadskloster (label 'Hkl 6/74')
and it cannot, therefore, be a syntype. A male in the collection could be from the type-locality but the
original description is restricted to females.

Identity. Medophron recurvus (Thomson) (Frilli, 1973 : 106, on the basis of the invalid 'lectotype').

Phygadeuon ripicola, 1885: 19. Lectotype £, SWEDEN: Skane, Ortofta (UZI, Lund), by designation of Frilli,
1973: 106.

Label. Ortofta [printed].

Thomson 'redescribed' this species in the Opuscula Entomologica(188Sb: 1242).
Identity. Phygadeuon ripicola Thomson.

Phygadeuon rotundipennis, 1884: 963. Lectotype $, SWEDEN: Skane, Ortofta (UZI, Lund), by designation of
Horstmann, 1967a: 15.
Label. 6rt. [hand].
Identity. Phygadeuon rotundipennis Thomson.

Phygadeuon rugipectus, 1884: 1040. Lectotype ?, SWEDEN: Skane, Ortofta (UZI, Lund), by designation of
Frilli, 1973: 107.

Labels. Ortofta [printed] ; rugipectus [Thomson cabinet label].
Identity. Phygadeuon rugipectus Thomson.

Phygadeuon scaposus, 1884: 961. Lectotype ?, SWEDEN: Skane, Palsio [= Palsjo] (UZI, Lund), by des-
ignation of Aubert, 1966: 129.

Labels. Hbg [hand] ; scaposus [Thomson cabinet label].
Identity. Phygadeuon scaposus Thomson.

Phygadeuon stilpninus, 18886: 1239. Lectotype $, SWEDEN: Skane, Palsjo (UZI, Lund), by designation of
Frilli, 1973: 107.

Labels. Pal. [hand] ; Stilpninus [Thomson cabinet label].
Identity. Phygadeuon stilpninus Thomson.

Phygadeuon submuticus, 1884: 962. Holotype $, SWEDEN: Skane, Stehag (UZI, Lund).
Labels. Rsio [printed] ; submuticus [Thomson cabinet label].

The holotype is the specimen referred to as 'neotype' by Aubert (1966: 129) and Frilli (1973: 107).
Identity. Phygadeuon submuticus Thomson.

Phygadeuon tenuicosta, 1884: 957. Lectotype ?, SWEDEN: Skane, Ringsion [= Ringsjon] (UZI, Lund), by
designation of Frilli, 1973: 108.

Labels. Rsio [printed] ; tenuicosta [Thomson cabinet label].
Identity. Phygadeuon tenuicosta Thomson.

Phygadeuon tenuiscapus, 1884: 960. Lectotype ?, SWEDEN: Skane, Ortofta (UZI, Lund), by designation of
Aubert, 1968: 195.

Labels. Ort. [hand] ; tenuiscapus [Thomson cabinet label].
Identity. Phygadeuon tenuiscapus Thomson.

THE ICHNEUMONIDAE DESCRIBED BY C. G. THOMSON 75

Phygadeuon trie/tops, 1884: 962. Lectotype 9, SWEDEN: Skane, Lund (UZI, Lund), by designation of Horst-
mann, 1967a: 11.
Label. L-d [printed].
Identity. Phygadeuon trichops Thomson.

Phygadeuon ungularis, 1884: 951. Syntypes 1 9, 7 $, SWEDEN: Skane, Ortofta and Skabersio [ = Skabersjo]
(UZI, Lund).

Labels. Ortofta [printed] (2 <J). Ort. [hand] (1 9 3 <?). Ort 5/VI [hand] (1 rf). Skb [hand] (1 J).

Frilli (1973: 108-109) thought that none of the specimens standing under this name agreed with the
original description. I believe that the eight specimens noted above are in sufficient agreement with the
description to be considered syntypes.

Identity. I believe this species belongs in Theroscopus. However, because of some uncertainty and
because it would create problems of secondary homonymy I am leaving it in Phygadeuon.

Phygadeuon varicornis, 1885 : 21. Syntypes 9 <3, FRANCE: le Crotoy (lost).
Identity. ? Phygadeuon varicornis Thomson.

Phytodietus continuus, 1877: 773. Lectotype 9, SWEDEN: Skane, Fagelsang (UZI, Lund), by designation of
Kerrich, 1962: 50-51.

Labels. Fsg 7/7 [hand] ; continuus [Thomson cabinet label].
Identity. Junior synonym of Phytodietus obscurus Desvignes (Kerrich, 1962: 50).

Phytodietus crassitarsis, 1877: 774. Lectotype $, SWEDEN: Skane, Ilstorp (UZI, Lund), by designation of
Sedivy, 1961:41.

Labels. lisp 12/7 [hand] ; crassitarsus [Thomson cabinet label].
Identity. Phytodietus crassitarsis Thomson.

Phytodietus geniculatus, 1877: 774. Lectotype $, SWEDEN: Skane, Bastad (UZI, Lund), by designation of
Sedivy, 1961 : 41.
Label, [small pinkish square].
Identity. Phytodietus geniculatus Thomson.

Phytodietus rubricosus, 1877: 773. Lectotype 9, SWEDEN : Skane, Lindholmen (UZI, Lund), by designation of
Tolkanitz, 1973: 880.

Labels. Lhn 9/8 [hand]; 192. [hand].

Identity. Junior synonym of Phytodietus ornatus Desvignes (Tolkanitz, 1973 : 880).

Pimpla brachycera, 1894: 2126. Lectotype $, ITALY : Trieste (UZI, Lund), by designation of Townes, Momoi
&Townes, 1965: 9.

Labels. 9.X Triest. [date, hand ; locality, printed] ; brachycera m [Thomson cabinet label].
Identity. Junior synonym of Exeristes roborator (Fabricius) (Townes, Momoi & Townes, 1965 : 9).

Pimpla (Itoplectis) clavicornis, 1889: 1409. Holotype 9, SWEDEN : Skane, Palsjo (UZI, Lund).
Labels. Pal. [hand] ; clavicornis [Thomson cabinet label].
Identity. Itoplectis clavicornis (Thomson).

Pimpla flavicoxis, 1877: 747. Lectotype 9, SWEDEN: Norrland (UZI, Lund), by designation of Townes,
Momoi & Townes, 1965 : 47.
Label. Norl. [printed].
Identity. Pimpla flavicoxis Thomson.

Pimpla laevifrous, 1877: 750. Lectotype 9, SWEDEN: Norrland (UZI, Lund), by designation of Aubert,
1972: 145.

Labels. Norl [printed] ; laevifrons [Thomson cabinet label].

Authors since Thomson (for example, Oehlke, 1967: 34; Aubert, 1969: 98, 1972: 145) have chosen to
alter the spelling of the name to laevifrons and there is evidence (Thomson's own cabinet label) that this is
what was intended. However, a strict interpretation of Article 32(aXii) of the Code (as amended, Bull. zool.
Norn. 31 (1974): 83) suggests that the original spelling should be retained.

Identity. Delomerista laevifrous (Thomson).

Pimpla longiceps, 1877: 746. Syntypes 2 $, 1 cJ, SWEDEN: Lappland (UZI, Lund).

Labels. Lpl. [printed] ; longiceps [Thomson cabinet label] (1 9)- LpL [printed] (1 $ 1 <J).
Identity. Junior synonym of Pimpla sodalis Ruthe (Perkins, 1941 : 645).

76 M. G. FITTON

Pimpla nigricans, 1877 : 754. Lectotype 9, SWEDEN : Skane (UZI, Lund), by designation of Townes, Momoi &
Townes, 1965: 11.
Label. Scan [printed].
Identity. Scambus nigricans (Thomson).

Pimpla nigriscaposa, 1877: 755. Syntypes 15 ?, 7 cJ, SWEDEN: [various localities] (UZI, Lund).

All specimens standing under this name except for two from Trieste (almost certainly added to the
collection after 1877) and a braconid (on the same pin as a female syntype) are regarded as syntypes.
Details of individual labels were not noted.

Identity. Junior synonym of Scambus brevicornis (Gravenhorst) (Perkins, 1943a: 268).

Pimpla ovalis, 1877: 748. Holotype 9, SWEDEN: Skane, Ilstorp (UZI, Lund).
Labels. lisp 12/7 [hand]; 90. [hand] ; ovalis [Thomson cabinet label].
Identity. Junior synonym ofltoplectis viduata (Gravenhorst) (Perkins, 1941 : 646).

Pimpla parallela, 1877: 752. Holotype 9, SWEDEN : Nerike [= Narke] (lost).

Identity. Junior synonym of Tromatobia ovivora (Boheman) (Oehlke, 1967: 18).

Pimpla pictifrons, 1877: 757. Lectotype 9, SWEDEN: Skane, Torekov (UZI, Lund), by designation ofSedivy,
1963: 246.

Label Tkov 7/60 [hand]
Identity. Dreisbachia pictifrons (Thomson).

Pimpla punctata, 1894: 2126. Syntype 1 9, AUSTRIA : 'Steiermarks alper'(UZI, Lund).
Label. Alpes styriaiae [hand].
Identity. Junior synonym ofExeristes roborator (Fabricius) (Perkins, 1943a: 261).

Pimpla punctiventris, 1877: 756. Syntypes 16 9, 11 g, 3 ?sex, SWEDEN: Skane, [various localities] (UZI,
Lund).

Thomson stated that this species was 'Sallsynt i Skane'. It therefore seems unlikely that all the Skane
specimens standing under this name date from 1877. However, in the absence of any evidence with which
to distinguish the syntypes, all are regarded as such. Details of individual labels were not noted. In
addition to the syntypes there is one female fromOstergotland.
Identity. Junior synonym of Scambus brevicornis (Gravenhorst) (Perkins, 1943a: 268).

Pimpla quadridentata, 1877: 749. Lectotype 9, SWEDEN: Skane (UZI, Lund), by designation of Townes,
Momoi & Townes, 1965: 45.

Labels. Scan [printed] ; 4-dentata [Thomson cabinet label].
Identity. Apechthis quadridentata (Thomson).

Pimpla stenostigma, 1877: 755. Syntypes 2 9, 2 <J, SWEDEN: Oland and Skane, Ringsjon (UZI, Lund).
Labels, [small green square] (1 9). O. [printed] (1 9 2 <£).
Identity. Junior synonym ofAcropimpla pictipes (Gravenhorst) (Perkins, 1943a: 267).

Pimpla strigipleuris, 1877: 747. Syntypes 12 9,4 cJ, SWEDEN: Skane, [various localities] (UZI, Lund).

Similar comments apply as for Pimpla punctiventris (see above). Details of individual labels were not
noted. In addition to the syntypes there are two males (one without locality, the other from Uppland) and
one female (from Trieste).

Identity. Junior synonym of Pimpla spuria Gravenhorst (Perkins, 1941 : 645).

Pimpla tricineta, 1877: 748. Lectotype 9, SWEDEN: Skane, Reften (UZI, Lund), by designation of Aubert,
1968: 194.

Labels. Rfn 10/7 [hand] ; 3-cincta [Thomson cabinet label].

Subsequent authors (for example, Dalla Torre, 1901:420; Perkins, 1941: 646; Oehlke, 1967:27;
Aubert, 1968: 194; 1969: 78) have chosen to alter the spelling of the name to tricineta and there is
evidence (Thomson's own reference (1890: 1408) and cabinet label) that this is what was intended.
However, a strict interpretation of Article 32(aXii) of the Code (as amended, Bull. zoo/. Norn. 31 (1974): 83)
suggests that the original spelling should be retained. Pimpla tricineta Thomson would, in any case, be a
junior primary homonym of Pimpla tricineta Cresson.

Identity. Junior synonym ofltoplectis alternans (Gravenhorst) (Perkins, 1941 : 646).

Platylabus concinnus, 18886: 1235. Syntype 1 9, SWEDEN: Skane, Palsjo (UZI, Lund).
Label. Palsjo [hand].
Identity. Platylabus concinnus Thomson.

THE ICHNEUMONIDAE DESCRIBED BY C. G. THOMSON 77

Platylabus (Platylabus) cyaneoviridis, 1894: 2105. Syntypes 2 ^, SWEDEN: Uppland, Upsala [= Uppsala]
(UZI, Lund).

Labels. Upsal 30/VI 91. [hand] ; cyaneoviridis m [Thomson cabinet label] (1 <$). Upsal [hand](l ^).
Identity. Cratichneumon cyaneoviridis (Thomson).

Platylabus (Tricholabus)femoratis, 1894: 2114. Syntype 1 9, SWEDEN : Skane, Palsjo (UZI, Lund).
Labels. Hbg. [hand] ; femoralis m [Thomson cabinet label].
Identity. Tricholabus femoralis (Thomson).

Platylabus (Platylabus) latiscapus, 1894: 2110. LECTOTYPE 9, SWEDEN: Stockholm (UZI, Lund), here
designated (selected by H. K. Townes).

Labels. 209 [hand] ; Col. Hgn. [printed] ; latiscapus [Thomson cabinet label].
Identity. Asthenolabus latiscapus (Thomson).

Platylabus (Platylabus) lativentris, 1894: 2109. Type(s) 9, SWEDEN: Skane, Ringsjon (lost).
Identity. Junior synonym of Platylabus transversus Bridgman (Perkins, 1953 : 1 15).

Platylabus (Platylabus) muticus, 1894: 2112. Syntypes 1 9, 1 c?, SWEDEN: Smaland and Vermland [ =
Varmland] (UZI, Lund).

Labels. Sm. [printed] ; Bhn [printed] ; Col. Hgn. [printed] ; muticus m [Thomson cabinet label] (9).
Verml [printed] ; Col. Hgn. [printed] (<?).
Identity. Platylabus muticus Thomson.

Plectiscus (Plectiscus) bistriatus, 18886: 1299. Syntype 1 $, SWEDEN: Skane, Ortofta (UZI, Lund).
Label Ort. [hand].
Identity. Plectiscidea bistriatus (Thomson).

Plectiscus (Dialipsis) crassipes, 18886: 1304. Syntypes 7 9, 1 c?» SWEDEN: [various localities] (UZI, Lund).
No notes were made of individual specimen labels.
Identity. Junior synonym of Dialipsis exilis Foerster (Townes, 1971 : 196).

Plectiscus (Plectiscus) curticauda, 18886: 1302. LECTOTYPE 9, GERMANY (WEST): ? Kaltenkirchen (UZI,
Lund), here designated (selected by J. F. Aubert).
Labels. Kalkh. 23/8.86. [hand] ; curticauda [Thomson cabinet label].
Identity. Plectiscidea curticauda (Thomson) comb. n.

Plectiscus (Plectiscus) eurystigma, 18886: 1301. Lectotype 9, SWEDEN: Skane, Esperod[= Asperod] (UZI,
Lund), by designation of Townes, Momoi & Townes, 1965 : 396.
Labels. Esp [printed] ; eurystigma [Thomson cabinet label].
Identity. Plectiscidea eurystigma (Thomson).

Plectiscus (Proclitus) heterocerus, 18886: 1307. Lectotype 9, SWEDEN: Skane, Palsjo (UZI, Lund), by des-
ignation of Townes, Momoi & Townes, 1965: 396.
Labels. Pal [hand] ; heterocerus [Thomson cabinet label].
Identity. Proclitus heterocerus (Thomson).

Plectiscus (Proclitus) longitarsis, 18886: 1306. Holotype 9, GERMANY (EAST): Rostock (UZI, Lund).
Labels. Rostock 5.10.87 [hand] ; longitarsis [Thomson cabinet label].
Identity. Proclitus longitarsis (Thomson).

Plectiscus (Aperileptus) obliquus, 18886: 1298. LECTOTYPE 9, SWEDEN: Skane, Vastra Vram (UZI, Lund),
here designated (selected by R. Hinz).
Label. W.W. [hand].
Identity. Aperileptus obliquus (Thomson).

Plectiscus (Plectiscus) subteres, 18886: 1300. Holotype 9, GERMANY (UZI, Lund).
Labels, f. 22/5. 86. [hand]; subteres [Thomson cabinet label].
Identity. Plectiscidea subteres (Thomson).

Plectocryptus pectoralis, 1896: 2383. Syntypes 1 9, 2 <$, SWEDEN : Skane, Ringsjon (UZI, Lund).
Labels, [green square] ; pectoralis m [Thomson cabinet label] (1 #). [green square] (1 9 1 c?)-
Identity. Aconias pectoralis (Thomson) comb. n.

Plectocryptus scansor, 1890: 1532. Holotype 9, SWEDEN : Goteborg (UZI, Lund).
Labels. Gbg [hand] ; Scansor m [Thomson cabinet label].
Identity. Giraudia scansor (Thomson) comb. n.

Polyblastus (Nemioblastus) albicoxa, 1883: 901. Holotype 9, SWEDEN : Skane, Arrie (UZI, Lund).
Label. Ar [hand].

78 M. G. FITTON

The specimen designated as lectotype by Kasparyan (1973 : 233) is from Ringsjon (label, a green square)
and cannot, therefore, be a syntype.

Identity. Polyblastus albicoxa Thomson.

Polyblastus (Scopiorus) angulatus, 1883: 902. Lectotype $, SWEDEN: Skane, Ringsion [= Ringsjon] (UZI,
Lund), by designation of Kasparyan, 1973: 249.
Label, [small green square].
Identity. Ctenochira angulata (Thomson).

Polyblastus (Scopiorus) fusicornis, 1883: 903. Lectotype?, SWEDEN: Skane, Palsio [ = Palsjo] (UZI, Lund),
by designation of Kasparyan, 1973: 284.
Labels. Hbg [hand] ; fusicornis [Thomson cabinet label].
Identity. Junior synonym of Ctenochira validicornis (Brischke) (Kasparyan, 1973: 284).

Polyblastus (Ctenacmus) genalis, 1883 : 902. Type(s) 9, SWEDEN: Skane, Torekov (lost).

The specimen designated as lectotype by Kasparyan (1973 : 259) is from Ringsjon (label, a green square)
and cannot, therefore, be a syntype.
Identity. Ctenochira genalis (Thomson) (Kasparyan, 1973 : 259, on the basis of the invalid 'lectotype').

Polyblastus macrocentrus, 18886: 1257. Lectotype 9, SWEDEN: Skane, Ortofta (UZI, Lund), by designation of
Aubert, 1966: 127.

Labels. Ort. [hand]; 9 [printed].

Identity. Polyblastus macrocentrus Thomson.

Polyblastus (Ctenacmus) nigripalpis, 1883:902. Lectotype & SWEDEN: Skane, Palsio [= Palsjo] (UZI,
Lund), by designation of Kasparyan, 1973: 262.

Labels. Hbg [hand] ; vertic A [hand] ; nigripalpis [Thomson cabinet label].
Identity. Junior synonym of Ctenochira haemosternus (Haliday) (Kasparyan, 1973: 262).

Polyblastus pallicoxa, 18886: 1257. Syntypes 5 ?, 1 <J, SWEDEN: Skane, Palsjo (UZI, Lund).
Labels. Pal [hand] ; pallicoxa [Thomson cabinet label] (1 9). Pal. [hand] (4 9 1 £).
Identity. Polyblastus pallicoxa Thomson.

Polyblastus (Ctenacmus) scutellaris, 1883: 901. Lectotype 9, SWEDEN: Skane, Lund (UZI, Lund), by des-
ignation of Kasparyan, 1973: 253.

Labels. Lund [printed] ; scutellatus [Thomson cabinet label].

Identity. Junior primary homonym of Polyblastus scutellaris Holmgren. Replacement name
Polyblastus scutellatus Thomson, 18886: 1257. Junior synonym of Ctenochira bisinuata Foerster
(Kasparyan, 1973: 253).

Polyblastus scutellatus, 18886: 1257. Replacement name for Polyblastus scutellaris Thomson (see entry
above).

Polyblastus (Polyblastus) subtitis, 1883: 900. Lectotype 9, SWEDEN: Oland (UZI, Lund), by designation of
Kasparyan, 1970: 863.

Labels, [large pink diamond] ; subtilis [Thomson cabinet label].
Identity. Junior synonym of Polyblastus varitarsus (Gravenhorst) (Kasparyan, 1973 : 227).

Polysphincta (Polysphincta) caudata, 18886: 1253. Holotype9, SWEDEN : Skane, Ronnemolla (UZI, Lund).
Labels. Ron. [hand] ; caudata [Thomson cabinet label].
Identity. Junior synonym ofSinarachna nigricornis (Holmgren) (Oehlke, 1967: 25).

Polysphincta (Polysphincta) picticollis, 18886:1254. Lectotype <J, SWEDEN: Gottland [= Gotland] (UZI,
Lund), by designation of Aubert, 1966: 127.
Label. G. [hand].
Identity. Zatypota picticollis (Thomson).

Polysphincta pulchrator, 1877: 757. Lectotype 9, SWEDEN: Skane, Holmeja (UZI, Lund), by designation of
Aubert, 1966: 127.

Labels. Hma [hand] ; pulchrator [hand].

Identity. Junior synonym of Zatypota percontatoria (Miiller) (Oehlke, 1967 : 26).

Porizon (Barycnemis) anurus, 1889: 1365. Lectotype 9, SWEDEN: Oland (UZI, Lund), by designation of
Horstmann, 1981: 62.
Label. 0. [printed].
Identity. Barycnemis anurus (Thomson).

THE ICHNEUMONIDAE DESCRIBED BY C. G. THOMSON 79

Portion (Cratophion) caudatulus, 1889: 1364. Holotype 9, SWEDEN: Norrland (UZI, Lund).
Labels. Norl. [printed] ; longicauda [Thomson cabinet label].
Identity. Barycnemis caudatulus (Thomson).

Porizon (Leptopygus)filicornis, 1889 : 1366. Holotype 9, GERMANY (WEST): Bavaria (UZI, Lund).
Labels. 73./311. ['73', hand; '3 11 ', printed]; Germ, [hand] ; filicornis [Thomson cabinet label].
Identity. Barycnemis filicornis (Thomson).

Porizon (Barycnemis) gracillimus, 1889: 1365. Lectotype 9, SWEDEN: Skane, Lund (UZI, Lund), by des-
ignation of Aubert, 1966: 131.
Label. Lund [printed].
Identity. Barycnemis gracillimus (Thomson).

Porizon (Barycnemis) laeviceps, 1889: 1365. Lectotype $, SWEDEN: Skane, Bogestad [= Bokestad] (UZI,
Lund), by designation of Aubert, 1968: 196.
Label Bs. [hand].
Identity. Barycnemis laeviceps (Thomson).

Pristomerus pallidus, 1890: 1456. Lectotype 9, YUGOSLAVIA: Dalmatia (UZI, Lund), by designation of
Townes, Momoi & Townes, 1965: 306.

Labels. Dalm [printed] ; sulphureus [Thomson cabinet label].
Identity. Pristomerus pallidus Thomson.

Promethus albicoxa, 1890: 1476, 1479. Lectotype d, SWEDEN: Skane, Palsjo (UZI, Lund), by designation of
Townes, Momoi & Townes, 1965: 414.
Label. Pal. [hand].
Identity. Junior synonym ofSussaba cognata (Holmgren) (Townes, Momoi & Townes, 1965: 414).

Promethus laticarpus, 1890: 1476, 1481. Lectotype 9, SWEDEN: Skane, Lund (UZI, Lund), by designation of
Diller, 1980: 60.
Label. Ld [hand].

There is a paralectotype male on the same pin as the lectotype.
Identity. Junior synonym ofSussaba pulchella (Holmgren) (Diller, 1980: 59).

Promethus melanaspis, 1890: 1477. Holotype 9, GERMANY (WEST): Bavaria, near Munich (ZSBS,
Munich).

Labels. 85. 818. [hand] ; melanaspis [hand]; Th . . . [hand, partly illegible].
Identity. Promethes melanaspis (Thomson).

Promethus nigriventris, 1890: 1476. Lectotype 9, SWEDEN: Halland, Ostra Karup (UZI, Lund), by des-
ignation of Townes, Momoi & Townes, 1965: 413.

Labels. Krp 14/6 [hand] ; nigriventris m [Thomson cabinet label].

Townes labelled the lectotype as 'Promethus albicoxd" (for which species he also labelled a lectotype).
This is undoubtedly a mistake and I have added my own label clarifying the status of the specimen as
lectotype of nigriventris.

Identity. Promethes nigriventris (Thomson).

Pyracmon lateralis, 1887c: 1109. Lectotype 9, SWEDEN: Skane, Ringsjon (UZI, Lund), by designation of
Horstmann, 1977: 73.

Labels. Rsjo 24/6 [printed] ; lateralis [Thomson cabinet label].
Identity. Junior synonym of Pyracmon truncicola Thomson (Horstmann, 1977: 73).

Pyracmon truncicola, 1887c: 1109. Lectotype 9, SWEDEN: Skane, Ekeshult (UZI, Lund) by designation of
Aubert, 1966: 130.

Labels. Ekh 16/6 [hand] ; truncicola [Thomson cabinet label].
Identity. Pyracmon truncicola Thomson.

Rhaestus (Rhaestus)femoralis, 1894: 1976. Holotype 9, FRANCE: Oignies (UZI, Lund).
Label. Oignies. [hand].
Identity. Rhaestus femoralis Thomson.

Rhaestus punctatus, 1890: 1533. Lectotype ^, SWEDEN: Oland (UZI, Lund), by designation of Townes,
Momoi & Townes, 1965: 243.
Label. O. [printed].
Identity. Glyptorhaestus punctatus (Thomson).

80 M. G. FITTON

Rhaestus (Glyptorhaestus) wuestneii [as wustneii], 1894: 1977. Lectotype 9, DENMARK: S0nderborg (ZM,
Copenhagen), by designation of Hinz, 1975: 44.
Labels. Sdb'g 23.V.83 [hand]; Coll. Wustnei. [printed].
Identity. Junior synonym of Glyptorhaestus punctulatus (Woldstedt) (Hinz, 1975: 44).

Sagaritis brachycera, 1887c: 1091. LECTOTYPE 9, SWEDEN: Oland, Borgholm (UZI, Lund), here des-
ignated (selected by R. Hinz).
Label.Q. [printed].
Identity. Campoletis brachycera (Thomson).

Sagaritis erythropus, 1887c: 1093. Lectotype $, SWEDEN: Skane, Ringsjon (UZI, Lund), by designation of
Aubert, 1972: 148.

Labels. Rsio [printed] ; erythropus [Thomson cabinet label].
Identity. Campoletis erythropa (Thomson).

Sagaritis macroura, 1887c: 1093. Syntype 1 ^, SWEDEN : Skane, Ortofta (UZI, Lund).
Label. Ortofta [printed].
Identity. Campoletis macroura (Thomson).

Sagaritis mucronella, 1887c: 1095. Holotype $, SWEDEN : Skane, Arrie (UZI, Lund).
Labels. Ar. [hand] ; mucronella [Thomson cabinet label].
Identity. Campoletis mucronella (Thomson).

Sagaritis varians, 1887c: 1095. LECTOTYPE 9, SWEDEN : Skane, Lund (UZI, Lund), here designated (selec-
ted by R. Hinz).
Label. L-d [printed].
Identity. Campoletis varians (Thomson).

Saotus nigriventris, 1894: 2019. Holotype 9, GERMANY (UZI, Lund).

Labels. Mdrk 20.VI.83 [hand] ; nigriventris [Thomson cabinet label].
Identity. Saotis nigriventris (Thomson).

Saotus varicoxa, 1894: 2019. Lectotype 9, SWEDEN: Skane, Palsjo (UZI, Lund), by designation of Kerrich,
1942: 70.

Labels. Pal [hand] ; varicoxa [Thomson cabinet label].
Identity. Saotis varicoxa (Thomson).

Smicroplectrus costulatus, 1883: 888. Holotype 9, SWEDEN: Lappland (UZI, Lund).
Labels. Lpl. [printed] ; costulatus [Thomson cabinet label].
Identity. Junior synonym of Smicroplectrus jucundus (Holmgren) (Kerrich, 1952: 409).

Spilocryptus dispar, 1873: 504. Syntypes 7 9, 14 <£ SWEDEN: [various localities] (UZI, Lund).

All specimens standing under this name (except 1 <$ with a white annulus on each antenna and differing
in other ways from the description) are regarded as syntypes. Some were probably added to the collection
after 1873 but there is no way of differentiating the original syntype series. The information given by
Thomson subsequently (1896: 2367-2368) needs to be taken into consideration when a lectotype is
selected.

Details of individual specimen labels were not noted.

Identity. Junior synonym of Agrothereut es abbreviator (Fabricius) (Roman, 1939: 187).

Spilocryptus nasutus, 1 873 : 505. Syntypes 4 ?, 3 (J, SWEDEN : Skane, Gotland and Oland (UZI, Lund).

Labels. Rsio [printed]; nasutus [Thomson cabinet label] (1 9). Gott. [hand] (1 9). Gott [hand]; Col
Dbm [printed] (1 9). Pal. [hand] (1 9). 0. [printed] (2 £). [small green square] (1 £).
Identity. Agrothereutes nasutus (Thomson) comb. n.

Spilocryptus ornatulus, 1873: 507. Lectotype 9, SWEDEN: Skane, Dalby (UZI, Lund), by designation of
Townes, Momoi & Townes, 1965: 171.
Label. Dby 9/53 [hand].
Identity. Gambrus ornatulus (Thomson).

Spilocryptus tibialis, 1873: 503. Lectotype 9, SWEDEN: Oland (UZI, Lund), by designation of Horstmann,
1968: 127.

Labels. 0. [printed] ; tibialis [Thomson cabinet label].

The lectotype designation published by Aubert (1966: 128) is invalid because the specimen concerned is
not a syntype (see Horstmann, 1968 : 127).

THE ICHNEUMONIDAE DESCRIBED BY C. G. THOMSON 81

Spilocryptus zygaenarum, 1873: 504. Lectotype 9, DENMARK : Zealand (ZM, Copenhagen), by designation of
Horstmann, 1968: 124.

Labels. 9 7/1858 Af Zygona filipend. Drewsen [hand] ; Danmark ex coll. Schi0dte [printed].

The lectotype designation published by Aubert (1966: 128) is invalid because the specimen concerned is
not a syntype (see Horstmann, 1968 : 125).

Identity. Junior synonym of Agrothereutesfumipennis (Gravenhorst) (Horstmann, 1968: 124).

Spudaeus facialis, 1894: 2014. Lectotype $, SWEDEN: Lappland (UZI, Lund), by designation of Aubert,
1966: 127.

Labels, [yellow square] ; Col Ros [printed] ; facialis [Thomson cabinet label].
Identity. Synodit es facialis (Thomson) comb. n.

Spudaeus mandibularis, 1894: 2013. Lectotype 9, SWEDEN: Norrland (UZI, Lund), by designation of Aubert,
1976ft: 273.

Labels. Norl. [printed] ; $ [printed] ; mandibularis [Thomson cabinet label].
Identity. Campodorus mandibularis (Thomson).

Spudaeus mesocastanus, 1894: 201 1. Type(s) 9, SWEDEN: Norrland (lost).

There is one female specimen in the collection originating from the Rudolphi collection, labelled 'Him'
[= Holmia]. Most Rudolphi specimens come from Halsingland (part of 'Norrland'). It is, therefore,
possible that this specimen is the type and that Thomson misread 'Him' as 'His'.

Identity. Rhinotorus mesocastanus (Thomson) comb. n. (on the basis of the specimen in the collection).

Spudaeus nigridens, 1894: 2013. Lectotype 9, FRANCE: Phalempin (UZI, Lund), by designation of Aubert,
19766:274.

Labels. Phalempin [hand] ; Gall. [hand].
Identity. Campodorus nigridens (Thomson).

Spudaeus sanguinipes, 1894: 2012. Syntype 1 <J, ? syntype 1 9, SWEDEN : Norrland (UZI, Lund).

Labels. Col. Rud. [hand] (cJ). [square of paper]; Col. Hgn. [printed] ; sanguinipes m [Thomson cabinet

label] (9).

There is no evidence that the female came from Norrland so it is only tentatively regarded as a syntype.
Identity. ? Arbelus sanguinipes (Thomson) comb. n.

Spudaeus stenocerus, 1894: 2013. Syntypes 9 <$, SWEDEN: Norrland and Skane, Ringsjon (lost).
Identity. ? Campodorus stenocerus (Thomson) comb. n.

Spudaeus subimpressus, 1894: 201 1. Syntype 1 9, SWEDEN: Skane, Bastad (UZI, Lund).
Labels. Bast [hand] ; 9 [printed] ; subimpressus [Thomson cabinet label].
Identity. Rhinotorus subimpressus (Thomson) comb. n.

Spudastica petiolaris, 1887c: 1123. LECTOTYPE 9, SWEDEN: Skane, Ringsion [= Ringsjon] (UZI, Lund),
here designated (selected by H. K. Townes).
Label, [small green square].
Identity. Junior synonym of Spudastica kriechbaumeri (Bridgman) (Townes, 1970ft: 169).

Stenocryptus nigriventris, 1874: 604. Lectotype 9, SWEDEN: Lappland (UZI, Lund), by designation of
Townes, Momoi & Townes, 1965: 156.
Label. Lap [hand].
Identity. Cubocephalus nigriventris (Thomson).

Stilpnus angustatus, 1884: 1027. Holotype 9, SWEDEN : Skane, Ronnemolla (UZI, Lund).
Labels. Ron [hand] ; alutaceus [hand] ; angustatus [Thomson cabinet label].
Identity. Stilpnus angustatus Thomson.

Stilpnus crassicornis, 1884: 1027. Syntype 1 9, SWEDEN: Skane, Bastad (UZI, Lund).
Labels. Scan [printed] ; crassicornis [Thomson cabinet label].
Identity. Stilpnus crassicornis Thomson.

Stilpnus tenuipes, 1884: 1028. Syntype 1 <J, SWEDEN : Skane, Palsjo (UZI, Lund).
Label. Palsio [printed].
Identity. Stilpnus tenuipes Thomson.

Stylocryptus (Stylocryptus) analis, 1883: 871. Lectotype 9, SWEDEN: Skane, Fogelsang [ = Fagelsang] (UZI,
Lund), by designation of Townes, Momoi & Townes, 1965 : 139.
Label. Fg. [hand].

82 M. G. FITTON

Stylocryptus (Gnathocryptus) clypealis, 1883: 870. Lectotype <$, SWEDEN: Skane, Ortofta (UZI, Lund), by
designation of Aubert, 1966: 129.
Label. Ort [hand].
Identity. Glyphicnemis clypealis (Thomson).

Stylocryptus eurycerus, 1896: 2386. Holotype 9, SWEDEN : Stockholm (UZI, Lund).

Labels. Him [printed] ; De V [printed] ; Col. Hgn. [printed] ; eurycerus m [Thomson cabinet label].
Identity. Endasys eurycerus (Thomson) (teste J. Sawoniewicz).

Stylocryptus (Stylocryptus) minutulus, 1883:872. Syntypes 2$, 6 <$, SWEDEN: Skane, Ringsion [ =
Ringsjon] and Bastad (UZI, Lund).

Labels, [small green square] (1 9 4 <J). Rsio [printed] (2 <£, on one pin). Bast [hand](l $).
Identity. Endasys minutulus (Thomson).

Symplecisfacialis, 18886: 1286. LECTOTYPE 9, SWEDEN: Skane, Degeberga (UZI, Lund), here designated
(selected by G. van Rossem).

Labels. Dgb. [hand] ; facialis [Thomson cabinet label].
Identity. Symplecisfacialis Thomson.

Syndipnus (Polyrhysius) anterior, 1894: 1999. LECTOTYPE $, SWEDEN: Skane, Palsjo (UZI, Lund), here
designated (selected by R. Hinz).

Labels. Pal [hand] ; 9 [printed] ; anterior m [Thomson cabinet label].
Identity. Synocoetes anterior (Thomson) comb. n.

Syndipnus (Trophoctonus) curvulus, 1894: 2000. Syntypes 1 $, 1 <J, SWEDEN : Norrland (UZI, Lund).
Labels. Rud. [hand] ; curvulus [Thomson cabinet label].
The two syntypes are on one pin.
Identity. Synomelix curvulus (Thomson).

Syndipnus (Hypamblys) lineiger, 1894: 2007. Syntype 1 9, SWEDEN : Skane, Palsjo (UZI, Lund).
Labels. Pal. [hand] ; lineiger n. [Thomson cabinet label].
The syntype lacks the gaster.
Identity. Synodites lineiger (Thomson) comb. n.

Syndipnus (Synodytes) orbitaKs, 1894: 2002. Holotype?, SWEDEN: Oland (UZI, Lund).
Labels. 0. [printed] ; brevicalcar [Thomson cabinet label].
Identity. Synodites orbitalis (Thomson).

Syndipnus (Smicrottus) parvicalcar, 1894: 2008. LECTOTYPE $, DENMARK : Sonderborg (UZI, Lund), here
designated (selected by H. K. Townes).
Label. Sondb'g [hand].
Identity. Smicrolius parvicalcar (Thomson).

Syndipnus (Synodytes) par viceps, 1894: 2002. Holotype ^, SWEDEN: Lappland (UZI, Lund).
Labels. Lpl. [printed] ; parviceps [Thomson cabinet label].
Identity. Synodites parviceps (Thomson).

Syndipnus (Syndipnus) pectoraKs, 1894: 2006. Syntypes 3 $, FRANCE: Libercourt and Thumeries (UZI,
Lund).

Labels. Libercourt. [hand] ; flavipectus [Thomson cabinet label] (1 $). Thumeries. [hand] (1 9)- Liber-
court, [hand] ; Gall, [hand] (1 9).

Identity. Syndipnus pectoralis Thomson.

Syndipnus (Syndipnus) punctiscuta, 1894: 2005. Syntypes 8 $, 1 & SWEDEN: Skane, Ilstorp; FRANCE: Fortif ;
GERMANY (WEST): Gotteskoog See and Gliicksburg; and DENMARK: Sandacker (UZI, Lund).

Labels. lisp 15/7 [hand] (1 $). Fortif. [hand]; punctiscuta m [Thomson cabinet label] (1 9)- Fortif.
[hand] (4 9). Gotteskog See. 7.8.87 [hand] (1 9). Glucksb'g 8.8.92. [hand] (1 9). Sandack. 27.7.88 [hand]
(19).

Identity. Syndipnus punctiscuta Thomson.

Syndipnus (Synodytes) subscaber, 1894: 2002. Syntype 1 9, SWEDEN : Norrland (UZI, Lund).

Labels. Nod. [printed] ; subscaber [Thomson cabinet label].

Aubert (1972: 147) was incorrect in regarding this specimen as holotype. Thomson mentions material
from Norrland and a specimen from Germany sent by Kriechbaumer. The latter specimen is not in Lund
but may be in the Kriechbaumer collection in Munich.

Identity. Synodites subscaber (Thomson).

THE ICHNEUMONIDAE DESCRIBED BY C. G. THOMSON 83

Synetaeris carbonella, 1887c: 1115. Syntype 1 9, SWEDEN : Skane, Yddinge (UZI, Lund).
Labels. Bkbg [hand] ; carbonella [Thomson cabinet label].

Identity. Synetaeris carbonella Thomson. Horstmann (1977) has synonymised Synetaeris with Pyr-
acmon.

Synetaeris heteropus, 1887c: 1115. Lectotype 9, GERMANY (UZI, Lund), by designation of Horstmann,
1977: 73.

Labels. Germ [hand] ; heteropus [Thomson cabinet label].

The lectotype was selected and labelled by Townes and not Aubert as stated by Horstmann (1977: 73).
Identity. Synetaeris heteropus Thomson. Horstmann (1977) has synonymised Synetaeris with Pyracmon.

Thersilochus (Thersilochus) apertus, 1889: 1382. Lectotype 9, SWEDEN: Ostergotland, Skeninge [ =
Skanninge] (UZI, Lund), by designation of Townes, Momoi & Townes, 1965: 314.

Labels. 104. [hand]; OG [hand] [not '50' as stated by Horstmann (1971b: 99) !]; apertus [Thomson
cabinet label].

Identity. N anodiaparsis apertus (Thomson).

Thersilochus (Thersilochus) brevicauda, 1889: 1382. Lectotype 9, SWEDEN : Skane, Helsingborg (UZI, Lund),
by designation of Horstmann, 19676: 129.
Labels. Hbg [hand] ; brevicauda [Thomson cabinet label].

The lectotype is the uppermost of three specimens on one pin. There is a fourth card point (without a
specimen) above the lectotype.

Identity. Aneuclis brevicauda (Thomson).

Thersilochus (Thersilochus) carinifer, 1889: 1392. Lectotype 9, FRANCE (UZI, Lund), by designation of
Horstmann, 19676: 127.

Labels. Gall, [hand] ; carinifer [Thomson cabinet label].
Identity. Diaparsis carinifer (Thomson).

Thersilochus (Thersilochus) crassicauda, 1889: 1396. Lectotype 9, GERMANY (UZI, Lund), by designation of
Horstmann, 19716:58.

Labels. 381 [hand] ; [small yellowish triangle] ; Germ, [hand] ; crassicauda [Thomson cabinet label].
Identity. Junior synonym of Pectinolochus ensifer (Brischke) (Horstmann, 19716: 58).

Thersilochus (Thersilochus) crassipes, 1889: 1400. Lectotype 9, FRANCE: Lille, Raismes (UZI, Lund), by
designation of Horstmann, 19676: 127.
Labels. Raismes. [hand]; Gall [hand].
Identity. Rugodiaparsis crassipes (Thomson).

Thersilochus (Thersilochus) decrescens, 1889: 1386. Lectotype 9, GERMANY (UZI, Lund), by designation of
Horstmann, 19716: 77.

Labels, [grey triangle] ; Germ, [hand] ; decrescens [Thomson cabinet label].
Identity. Phradis decrescens (Thomson).

Thersilochus (Diaparsus) fenestralis, 1889: 1370. Lectotype 9, GERMANY (EAST): Rostock (UZI, Lund), by
designation of Horstmann, 19716: 112.

Labels. Rostock 27.4.85 [hand]; Germ, [hand] ; fenestralis [Thomson cabinet label].
Identity. Gonolochus fenestralis (Thomson).

Thersilochus (Thersilochus) filicornis, 1889: 1393. Lectotype 9, SWEDEN: Skane, Alnarp (UZI, Lund), by
designation of Horstmann, 19716: 121.

Labels. Alnarp [printed] ; filicornis [Thomson cabinet label].
There is a paralectotype $ on the same pin as (and above) the lectotype.
Identity. Tersilochus filicornis (Thomson).

Thersilochus (Thersilochus) flavicornis, 1889: 1391. Lectotype 9, SWEDEN: Skane, Lund (UZI, Lund), by
designation of Horstmann, 19716: 106.
Labels. Ld [hand] ; ruficornis [Thomson cabinet label].
Identity. Junior synonym of Diaparsis stramineipes (Brischke) (Horstmann, 19716: 106).

Thersilochus (Diaparsus) genalis, 1889: 1373. Lectotype & SWEDEN: Skane, Yddinge (UZI, Lund), by
designation of Aubert, 1966: 131.

Label. Bok 8/78 [hand].

I do not agree with Horstmann (19716: 105) that the earlier publication by Aubert (1964: 63) consti-
tutes a valid lectotype designation.

Identity. Junior synonym of Diaparsis nutritor (Fabricius) (Horstmann, 19716: 104-105).

84 M. G. FITTON

Thersilochus (Thersilochus) heterocerus, 1889: 1383. Lectotype 9, SWEDEN: Skane (UZI, Lund), by des-
ignation of Horstmann, 1967 'b: 127.

Labels. Scan [printed]; heterocerus [Thomson cabinet label].

The lectotype is the lower of two females on one pin. There is a third card point (without a specimen)
below the lectotype.

Identity. Tersilochus heterocerus (Thomson).

Thersilochus (Thersilochus) incident, 1889: 1382. Lectotype 9, SWEDEN: Skane, Trelleborg (UZI, Lund), by
designation of Horstmann, 19716: 61.
Label. Tbg 6/84 [hand].
Identity. Aneuclis incidens (Thomson).

Thersilochus (Thersilochus) inter stitialis, 1889: 1389. Lectotype <$, SWEDEN : Skane, Lomma (UZI, Lund), by
designation of Horstmann, 19676: 128.

Labels. L-a [printed] ; incidens [Thomson cabinet label].

The lectotype is the lowest of three specimens on one pin. The middle specimen is represented only by
part of a wing.

Identity. Phradis interstitialis (Thomson).

Thersilochus (Thersilochus) liopleuris, 1889: 1398. Lectotype 9, SWEDEN: Norrland (UZI, Lund), by de-
signation of Horstmann, 19716: 129.
Labels. Norl [printed] ; liopleuris [Thomson cabinet label].
Identity. Tersilochus liopleuris (Thomson).

Thersilochus (Thersilochus) longicornis, 1889: 1384. Type(s) 9, SWEDEN : Skane, Trelleborg (lost).

The specimen designated as lectotype by Horstmann (19716: 129) is from Tvedora [as stated by
Horstmann !] and cannot, therefore, be from the type-series.

Identity. Tersilochus longicornis (Thomson) (Horstmann, 19716: 129, on the basis of the invalid 'lec-
totype').

Thersilochus (Thersilochus) maritimus, 1889: 1381. Holotype?, SWEDEN: Skane, Kempinge[= Kampinge]
(UZI, Lund).

Labels. Kpe 8/74 [hand] ; maritimus [Thomson cabinet label].
Identity. Aneuclis maritimus (Thomson).

Thersilochus (Thersilochus) melanogaster, 1889: 1392. Lectotype 9, SWEDEN: Skane, Tvedora (UZI, Lund),
by designation of Aubert in Aubert & Jourdheuil, 1959: 187.
Label. Tve 5/80 [hand].
Identity. Tersilochus melanogaster (Thomson).

Thersilochus (Thersilochus) monticola, 1889: 1388. Lectotype & FRANCE: Mt des Cattes (UZI, Lund), by
designation of Horstmann, 19716: 65.

Labels. Mt. des Cattes. [hand]; Gall [hand] ; montanus [Thomson cabinet label].
Identity. Junior synonym ofHeterocola proboscidalis (Thomson) (Horstmann, 19716: 65).

Thersilochus (Thersilochus) obliquus, 1889: 1392. Lectotype ?, SWEDEN: Skane, Ortofta (UZI, Lund), by
designation of Horstmann, 19716: 131.
Label. Ort. [hand].
Identity. Tersilochus obliquus (Thomson).

Thersilochus (Thersilochus) pallicarpus, 1889: 1387. Lectotype 9, SWEDEN: Stockholm (UZI, Lund), by
designation of Horstmann, 19676: 129.

Labels. Him [printed] ; Stal [printed] ; pallicarpus [Thomson cabinet label].
Identity. Junior synonym ofHeterocola proboscidalis (Thomson) (Horstmann, 19716: 65).

Thersilochus (Diaparsus) parviceps, 1889: 1376. Holotype 9, FRANCE (UZI, Lund).
Labels. Gall, [hand] ; parviceps [Thomson cabinet label].
Identity. Junior synonym of Microdiaparsis versutus (Holmgren) (Horstmann, 19716: 81).

Thersilochus (Thersilochus) proboscidalis, 1889: 1388. Lectotype 9, GERMANY (WEST) : Bavaria (UZI, Lund),
by designation of Horstmann, 19716: 65.
Label. Germ. [hand].
Identity. Heterocola proboscidalis (Thomson).

THE ICHNEUMONIDAE DESCRIBED BY C. G. THOMSON 85

Thersilochus (Thersilochm) striola, 1889: 1396. Lectotype 9, NORWAY (UZI, Lund), by designation of
Horstmann, 19716:56.

Labels. 287 [hand]; Norv. [hand] ; Striola [Thomson cabinet label].
Identity. Pectinolochus striolus (Thomson).

Thersilochus (Thersilochus) subdepressus, 1889: 1396. Lectotype 9, SWEDEN: Skane, Ringsion [= Ringsjon]
(UZI, Lund), by designation of Horstmann, 19716: 132.

Labels, [small green square] ; subdepressus [Thomson cabinet label].
Identity. Tersilochus subdepressus (Thomson).

Thersilochus (Thersilochus) temporally, 1889: 1387. Lectotype 9, GERMANY (UZI, Lund), by designation of
Horstmann, 19716: 73.
Labels, [grey triangle] ; Germ [hand].
Identity. Junior synonym ofPhradis brevis (Brischke) (Horstmann, 19716: 73).

Thymarus collaris, 1883 : 909. Syntype 1 & SWEDEN: Gottland [ = Gotland] (UZI, Lund).
Label. Gott [printed].
Identity. Thymaris collaris (Thomson).

Thymarus compressus, 1883: 909. LECTOTYPE 9, SWEDEN: Skane, Ringsion [Ringsjon] (UZI, Lund), here
designated (selected by H. K. Townes).
Labels. Rsio [printed] ; compressus [Thomson cabinet label].
Identity. Thymaris compressa (Thomson).

Trachyarus corvinus, 1891: 1612. LECTOTYPE 9, SWEDEN: Skane, Palsjo (UZI, Lund), here designated
(selected by H. K. Townes).

Labels. Pal [hand] ; corvinus m [Thomson cabinet label].
Identity. Trachyarus corvinus Thomson.

Tranosema arenicola, 1887c: 1 138. Lectotype 9, SWEDEN: Skane, Kungsmarken (UZI, Lund), by designation
of Horstmann, 1977: 78.

Labels. Kgsm 30/7 [hand] ; arenicola [Thomson cabinet label].
Identity. Junior synonym of Tranosema rostralis (Brischke) (Horstmann, 1977: 78).

Tranosema latiuscula, 1887c: 1 138. Type(s) 9, SWEDEN: Skane, Lund (lost).

A lectotype designation has been published by Hortsmann (1977: 78) for this species. I regard it as
invalid because I consider the specimen not to be a syntype. It disagrees with the original description in a
number of important characters. Of the five specimens standing under this name one female from
Skabersjo [not a type and labelled 'Skb' and 'latiuscula'] agrees very well with the description and is the
basis of the generic placement given here.

Identity. Dolophron latiusculus (Thomson) comb. n.

Trematopygus curvispina, 1883: 930. Holotype9, SWEDEN : Skane, Stehag(UZI, Lund).
Labels. Rsio [printed] ; 9 [printed] ; curvispina [Thomson cabinet label].
Identity. Lethades curvispina (Thomson) comb. n.

Trematopygus kriechbaumeri, 1894: 2015. Syntypes 1 9 (UZI, Lund), 1 9 (ZSBS, Munich), GERMANY
(WEST): Bavaria.

Labels. Alpes [printed] ; 2307. [hand] ; Kriechbaumeri m [Thomson cabinet label] (Lund specimen).
Mon. 9.5.56. A.Krchb [hand]; 8800. [hand] (Munich specimen).

Identity. Junior synonym of Trematopygus melanocerus (Gravenhorst) (Townes, 19706: 71).

Trematopygus lethierryi, 1894: 2016. Lectotype 9, FRANCE: Lille, Libercourt (UZI, Lund), by designation of
Aubert, 1966: 127.

Labels. Libercourt. [hand] ; Lethierryi [Thomson cabinet label].
Identity. Trematopygus lethierryi Thomson.

Trematopygus scabriculus, 1883: 930. Lectotype 9, SWEDEN: Jamtland, Areskutan (UZI, Lund), by desig-
nation of Aubert, 1972: 147.
Label. Are [hand].
Identity. Lethades scabriculus (Thomson).

Trichocryptus aquaticus, 1874: 611. Syntypes 9 9, 4 ^, SWEDEN: Skane, [various localities] (UZI, Lund).
Details of individual labels were not noted.

It is unlikely that all the specimens under this name date from 1874. However, as there is no means of
differentiating the original series all are regarded as syntypes.
Identity. Apsilops aquaticus (Thomson) comb. n.

86 M. G. FITTON

Tricttstus albicinctus, 18876: 206. Type(s) 9, SWEDEN (lost).

Neotype 9, GERMANY (UZI, Lund), by designation of Aeschlimann, 19736: 249.
Identity. Triclistis albicinctus Thomson.

Triclistus areolatus, 18876: 203. Lectotype 9, SWEDEN: Skane (UZI, Lund), by designation of Aubert,
1966: 128.

Label. Scan camp [printed].
Identity. Triclistus areolatus Thomson.

Triclistus facialis, 18876: 205. Holotype 9, SWEDEN : Gotland (UZI, Lund).
Labels. Gott [printed] ; facialis [Thomson cabinet label].
Identity. Triclistus facialis Thomson.

Tricttstus lativentris, 18876: 203. Lectotype 9, SWEDEN: Skane, Ortofta (UZI, Lund), by designation of
Aubert, 1966: 128.

Labels. Ort. [hand] ; lativentris [Thomson cabinet label].
Identity. Triclistus lativentris Thomson.

Triclistus longicalcar, 18876: 205. Lectotype 9, SWEDEN: Skane, Lund (UZI, Lund), by designation of
Aubert, 1966: 128.
Label. L-d [printed].
Identity. Triclistus longicalcar Thomson.

Triclistus nitifrons, 18876: 204. Lectotype 9 FRANCE: Phalempin (UZI, Lund), by designation of Aeschli-
mann, 19736: 238.

Labels. Phalempin [hand] ; Gall, [hand] ; nitifrons [Thomson cabinet label].
Identity. Junior synonym of Triclistus pallipes Holmgren (Aeschlimann, 19736: 236).

Triclistus pubiventris, 18876: 205. Lectotype 9, SWEDEN: Skane, Palsjo (UZI, Lund), by designation of
Aeschlimann, 19736: 249.

Labels. Pal [hand] ; pubiventris [Thomson cabinet label].

Aubert's lectotype designation (1966: 128) is not considered valid because he did not indicate (in either
the publication or by labelling) which of two specimens in one pin was selected. The lectotype is the upper
specimen.

Identity. Triclistus pubiventris Thomson.

Triclistus spiracularis, 18876: 205. Lectotype 9, SWEDEN: Norrland (UZI, Lund), by designation of Aubert,
1966: 128.

Label. Norl. [printed].
Identity. Triclistus spiracularis Thomson.

Tryphon auricularis, 1883: 897. Lectotype cJ, SWEDEN: Skane, Fagelsang (UZI, Lund), by designation of
Townes, Momoi & Townes, 1965: 106.
Label. Fsg 1/7 [hand].
Identity. Tryphon auricularis Thomson.

Tryphon bidentulus, 1883: 897. Lectotype ^, SWEDEN: Skane, Ringsion [= Ringsjon] (UZI, Lund), by
designation of Kasparyan, 1969: 655.
Label, [small green square].
Identity. Tryphon bidentulus Thomson.

Tryphon ceratophorus, 18886: 1256. Lectotype 9, SWEDEN: Skane, Lund (UZI, Lund), by designation of
Kasparyan, 1971: 293.
Label. Lund [printed].
Identity. Cosmoconus ceratophorus (Thomson).

Tryphon erythrogaster, 1883: 897. Lectotype 9, SWEDEN: Skane, Skabersjo (UZI, Lund), by designation of
Kasparyan, 1969: 658.
Label. Skb [hand].

The lectotype is the lower of two specimens on the same pin.
Identity. Junior synonym of Tryphon relator (Thunberg) (Kasparyan, 1969: 658).

Tryphon pleuralis, 1883: 897. Syntype 1 £, SWEDEN : Skane, Ringsjon (UZI, Lund).
Label, [small green square].

Identity. Junior primary homonym of Tryphon pleuralis Cresson. Replacement name Tryphon abditus
Kasparyan, 1969: 654.

THE ICHNEUMONIDAE DESCRIBED BY C. G. THOMSON

87

Xylonomus glyptus, 1877: 776. Syntype 1 ^, SWEDEN: Oland (UZI, Lund).
Labels.Q. [printed], [paper square].
Identity. Xorides glyptus (Thomson).

Nomenclatural summary

A systematic list of the species treated in this paper is given below. The classification is basically
that of Townes (as outlined in the introduction) but the family-group names (subfamilies, tribes
and subtribes) are those correct under the Code (see Fitton & Gauld, 1976; 1978).

Subfamily PIMPLINAE
Tribe EPHIALTINI

EXERISTES Foerster
roborator (Fabricius)

brachycera (Thomson)
punctata (Thomson)

SCAMBUSHeirtig

brevicornis (Gravenhorst)
nigriscaposa (Thomson)
punctiventris (Thomson)
nigricans (Thomson)
LIO TR YPHON Ashmead
crassisetus (Thomson)
pleuralis (Thomson)

TOWN ESI A Ozols
tenuiventris (Holmgren)
antefurcalis (Thomson)

PARAPERITHOUS Haupt
gnathaulax (Thomson)
luteipes (Thomson)

DOL1CHOMITUS Smith
cognator (Thunberg)

macrurus (Thomson)
dux (Tschek)

crassiceps (Thomson)
messor (Gravenhorst)

heteropus (Thomson)
populneus (Ratzeburg)

abbreviates (Thomson)
scutellaris (Thomson)
terebrans (Ratzeburg)

planifrons (Thomson)
tuberculatus (Geoffroy)

parallelus (Thomson)

ACROPIMPLA Townes
pictipes (Gravenhorst)

stenostigma (Thomson)

TROMATOBIA Foerster
ovivora (Boheman)

parallela (Thomson)

Tribe POLYSPHINCTINI

DREISBACHIA Townes
pictifrons (Thomson)

SINARACHNA Townes
nigricornis (Holmgren)
caudata (Thomson)

ZATYPOTA Foerster
percontatoria (Miiller)

pulchrator (Thomson)
picticolUs (Thomson)

Tribe PIMPLINI

ITOPLECT1S Foerster
alternant (Gravenhorst)

tricineta (Thomson)

tricincta (Thomson) (unjustified

emendation)
clavicornis (Thomson)
viduata (Gravenhorst)

ovalis (Thomson)

APECHTHIS Foerster
quadridentata (Thomson)

PIMPLA Fabricius
flavicoxis Thomson
sodalis Ruthe

longiceps Thomson
spurla Gravenhorst

strigipleuris Thomson

Tribe DELOMERISTINI
DELOMERISTA Foerster
laevifrous (Thomson)

laevifrons (Schmiedeknecht) (unjustified
emendation)

Subfamily TRYPHONINAE
Tribe PHYTODIETINI

PHYTODIETUS Gravenhorst
crassitarsls Thomson
geniculatus Thomson
obscurus Desvignes

continuus Thomson
ornatus Desvignes

rubricosus Thomson

NETELIA Gray
cristatus (Thomson)
dilatatus (Thomson)

brachycerus (Thomson)
fuscicornis (Holmgren)

gracilipes (Thomson)
latungulus (Thomson)
melanurus (Thomson)
nlgricarpus (Thomson)
ocellaris (Thomson)
opaculus (Thomson)

88

M. G. FITTON

Tribe THYMARIDINI
THYMARISFoerster
collaris (Thomson)
compressa (Thomson)

OEDEMOPSIS Tschek
limbata Thomson

Tribe TRYPHONINI
GR YPOCENTRUS Ruthe
apicalis Thomson

POL YBLASTUS Hartig
albicoxa Thomson
macrocentrus (Thomson)
palttcoxa Thomson
varitarsus (Gravenhorst)

subtilis Thomson
CTENOCHIRA Foerster
angulata (Thomson)
bisinuata Foerster

scutellaris (Thomson) (homonym)

scutellatus (Thomson)
genalis (Thomson)
haemosternus (Haliday)

nigripalpis (Thomson)
validlcornis (Brischke)

fusicornis (Thomson)

ERROMENUS Holmgren
junior (Thunberg)

arenicola Thomson
melanotus (Gravenhorst)

cavigena Thomson
plebejus (Woldstedt)

brevitarsis Thomson
punctatus (Woldstedt)
simplex Thomson
COSMOCONUS Foerster
ceratophorus (Thomson)
TRYPHON Fallen
abditus Kasparyan

pleuralis Thomson (homonym)
auricularix Thomson
bidentulus Thomson
relator (Thunberg)

erythrogaster Thomson

Tribe EXENTERINI
KRISTOTOMUS Mason
laetus (Gravenhorst)

calcaratus (Thomson)
marginatus (Thomson)

CFC4S/,STownes
rubiginosus (Gravenhorst)
binotatus (Thomson)
parvulus (Thomson)

EXYSTONSchi0dte
calcaratus Thomson
genalis Thomson
pratorum (Woldstedt)

brevipetiolatus Thomson

sponsorius (Fabricius)
carinatus Thomson

SMICROPLECTRUS Thomson
jucundus (Holmgren)

costulatus Thomson

ACROTOMUS Holmgren
lucidulus (Gravenhorst)
auriculatus (Thomson)

CTENISCUS Haliday
frontalis (Thomson) comb. n.
nigrifrons (Thomson)
paltitarsis (Thomson)
pedatorius (Panzer)

filipalpis (Thomson)

EXENTERUS Hartig
claripennis Thomson
laricinus Thomson

? adspersus Hartig
oriolus Hartig

flavellus Thomson
simplex Thomson

ERIDOLWS Foerster
albicoxa (Thomson)
brevigena (Thomson)
deletus (Thomson)
genalis (Thomson) comb. n.
lineiger (Thomson)
marginatus (Thomson)
punctipes (Thomson)
1 punctipleuris (Thomson) comb. n.
quadrinotatus (Thomson) comb. n.
rufonotatus (Holmgren)

breviventris (Thomson)
signifer (Thomson)
t-nigrum (Thomson)

Tribe IDIOGRAM MATINI

IDIOGRAMMA Foerster
alysiina (Thomson)

Subfamily ADELOGNATHINAE

ADELOGNA THUS Holmgren
aciculatus Thomson
brevicornis Holmgren

limbatus Thomson
dimidiatus Thomson
facialis Thomson
fasciatus Thomson
laevicollis Thomson
nigriceps Thomson
nigricornis Thomson
pilosus Thomson
puncticollis Thomson
punctiventris Thomson
punctulatus Thomson
tetracinctorius (Thunberg)

scabriculus Thomson

THE ICHNEUMONIDAE DESCRIBED BY C. G. THOMSON

89

Subfamily XORIDINAE

ODONTOCOLON Cushman
dentipes (Gmelin)

pinetorum (Thomson)
punctulatum (Thomson)
quercinum (Thomson)

XORIDESLatreille
glyptus (Thomson)

Subfamily PHYGADEUONTINAE
Tribe PHYGADEUONTINI

Subtribe CHIROTICINA
HANDAO1A Seyrig
bellicornis (Thomson)

CH IRQ TIC A Foerster
maculipennis (Gravenhorst)
glyptonotus (Thomson)
rubrotincta (Thomson)

Subtribe ACROLYTINA
EUDELUS Foerster
simillimus (Taschenberg)
pallicarpus (Thomson)
scabriculus (Thomson)

ACROLYTA Foerster

rufocincta (Gravenhorst)
capreolus (Thomson)
DIAGLYPTELLANA Horstmann

opacula (Thomson)

Subtribe HEMITELINA
PLEUROGYRUS Townes
cyclogaster (Thomson)

ARO TREPHES Townes
parvipennis (Thomson)

XIPHULCUSTov/nes
constrictus (Thomson)
floricolator (Gravenhorst)
longulus (Thomson)

HEMITELES Gravenhorst
similis (Gmelin)

unicolor Thomson

ACLASTUS Foerster
gracilis (Thomson)
solutus (Thomson)

Subtribe GELINA
DICHROGASTER Doumerc
aestivalis (Gravenhorst)

geniculatus (Thomson)
heteropus (Thomson)
liostylus (Thomson)
longicaudatus (Thomson)

CEL/SThunberg
albipalpus (Thomson)
balteatus (Thomson)
brevicauda (Thomson)
breviceps (Thomson)
elynu (Thomson)

gibbifrons (Thomson)
gladalis (Holmgren)

aeneus (Thomson)
gonatopinus (Thomson)
grandiceps (Thomson)
infumatus (Thomson)
longicauda (Thomson)
mandibularis (Thomson)
melanogaster (Thomson)
myrmecinus (Thomson)
ornatulus (Thomson)
pilosus (Thomson) comb. n.
rubricollis (Thomson)
rugifer (Thomson)
spinulus (Thomson)

AGASTHENES Foerster
varitarsus (Gravenhorst)
stagnalis (Thomson)

Subtribe GNYPETOMORPHINA
GNYPETOMORPHA Foerster
obscura (Bridgman)
apertus (Thomson)

Subtribe MASTRINA
PYGOCR YPTUS Roman

grandis (Thomson)
MA SIR US Foerster

costalis (Thomson)

fumipcnnis (Thomson)

rufulus (Thomson)

ISADELPHUS Foerster
armatus (Gravenhorst)

bidentulus (Thomson)
gallicola (Bridgman)

nigriventris (Thomson)
inimicus (Gravenhorst)

obscuripes (Thomson)

rubripes (Thomson)

ZOOPHTHORUS Foerster
cynipinus (Thomson)
graculus (Gravenhorst)

auriculatus (Thomson)
hirticeps (Thomson)
microstomus (Thomson)
notaticrus (Thomson)
plumbeus (Thomson)
punctiventris (Thomson)

MASTRULUS Horstmann
capra (Thomson)

CL YPEO TELES Horstmann

distant (Thomson)

rugifrons (Thomson)
LOCHETICA Kriechbaumer

pimplaria (Thomson)

Subtribe ETHELURGINA
RHEMBOBIUS Foerster
? nigriceps (Thomson)
? nigricollis (Thomson)

90

M. G. FITTON

Subtribe ENDASEINA
CHARITOPES Foerster
areolaris (Thomson)
clausus (Thomson)
macrurus (Thomson)
MEDOPHRON Foerster
armatulus (Thomson)
caudatulus (Dalla Torre)

caudatus (Thomson) (homonym)
flavitarsis (Dalla Torre)

flavipes (Thomson) (homonym)
recurvus (Thomson)

ENDASYS Foerster
analis (Thomson)
eurycerus (Thomson)
minutulus (Thomson)
GL YPHICNEMIS Foerster

clypealis (Thomson)
AMPHIBULUS Kriechbaumer
gradtts Kriechbaumer
bispinus (Thomson)

Subtribe BATHYTRICHINA
BA THYTHRIX Foerster
aereus (Gravenhorst)
brevis (Thomson)
collaris (Thomson)
fragilis (Gravenhorst)

geniculosus (Thomson)
linearis (Gravenhorst)

heteropus (Thomson)
laminm (Thomson)
rugulosus (Thomson)
strigosus (Thomson)

Subtribe PHYGADEUONTINA
OECOTELMA Townes
gracilipes (Thomson)
PLA TYRHABDUS Townes
inflatus (Thomson)
monodon (Thomson)

SULCARIUS Townes

biannulatus (Gravenhorst)
homocerus (Thomson)

nigricornis (Thomson)
TROPISTES Gravenhorst

falcatus (Thomson)

ORTH1ZEMA Foerster
hadrocerum (Thomson)
triannulatum (Thomson)

TR1CHOLINUM Foerster
ischnocerum (Thomson)

S TIBEUTES Foerster

? breviareolatus (Thomson)

curvispina (Thomson)
THEROSCOPUS Foerster

annulicornis (Thomson)

faciator (Aubert)

facialis (Thomson) (homonym)

fasciatulus Horstmann

fasciatus (Thomson) (homonym)
melanopygus (Gravenhorst)

validicornis (Thomson)
ochrogaster (Thomson)
? trochanteralis (Dalla Torre) comb. n.

trochanteratus (Thomson) (homonym)
trochanteratus (Thomson)
ungularis (Thomson)

PHYGADEUON Gravenhorst
? acutipennis Thomson
alpinus (Thomson)
arcticus (Thomson)
bidens Thomson
brachyurus Thomson
brevitarsis Thomson
? canattculatus Thomson
cubiceps Thomson
curviscapus Thomson
dimidiatus Thomson
fluvicuns Thomson
grandiceps Thomson
? heterogaster Thomson
inflatus Thomson

infelix Dalla Torre
laeviventris Thomson
lapponicus Thomson
Kogaster Thomson
Kosternus Thomson
longigena Thomson
magnicornis (Thomson)
monodon Thomson
ocularis Thomson
oppositus Thomson
ovaliformis Dalla Torre

ovalis Thomson (homonym)
pallicarpus Thomson

pallidicarpus Dalla Torre (unjustified

emendation)
punctigena Thomson
punctipleuris Thomson
punctiventris Thomson
ripicola Thomson
rotundipennis Thomson
rugipectus Thomson
scaposus Thomson
stilpninus Thomson
submuticus Thomson
tenuicosta Thomson
tenuiscapus Thomson
trichops Thomson
? ungularis Thomson
? varicornis Thomson

CERA TOPHYGADEUON Viereck
anurus (Thomson)

parvicauda (Thomson)
longiceps (Thomson)

LEPTOCR YPTOIDES Horstmann
clavipes (Thomson)

THE ICHNEUMONIDAE DESCRIBED BY C. G. THOMSON

91

Subtribe STILPNINA
STILPNUS Gravenhorst
angustatus Thomson
crassicornis Thomson
tenuipes Thomson

MESOLEPTUS Gravenhorst
filicornis (Thomson)
flavipes (Thomson) comb. n.
marginatus (Thomson)
petiolaris (Thomson)
ripicola (Thomson)

A TRACTODES Gravenhorst
alutaceus Thomson
angustipennis Foerster

flavicoxa Thomson
crassicornis Thomson

? intersectus Foerster
croceicornis Haliday

compressus Thomson
eryptobius Foerster

parallelus Thomson
pauxillus Foerster

breviscapus Thomson
pusillus Foerster

liogaster Thomson
tenuipes Thomson
thomsoni (Dalla Torre)

rufipes Thomson (homonym)

Tribe PSEUDOCRYPTINI

JA VRA Cameron

areolaris (Thomson) comb. n.
opaca (Thomson)

PARMORTHA Townes
pleuralis (Thomson)

CUBOCEPHA L US Ratzeburg
annulitarsis (Thomson)
femoralis (Thomson) comb. n.
longicauda (Thomson) comb. n.
nigriventris (Thomson)
ruficoxis (Thomson) comb. n.
sternocerus (Thomson)

ORESBIUS Marshall

nigricornis (Thomson) comb. n.
nigriventris (Thomson) comb. n.
punctifer (Thomson) comb. n.
septentrionalis (Thomson) comb. n.

POL YTRIBAX Foerster
gravenhorsti (Thomson) comb. n.

A CO NIA S Cameron

pectoralis (Thomson) comb. n.

GIRAUDIA Foerster

scansor (Thomson) comb. n.

SCHENKIA Foerster
aries (Thomson) comb. n.
opacula (Thomson)
rubricollis (Thomson)

PLEOLOPHUS Townes
? alutaceus (Thomson)

A PTESIS Foerster

borealis (Thomson) comb. n.
distans (Thomson)
femoralis (Thomson)
lapponica (Thomson) comb. n.
nigritula (Thomson)
ochrostomus (Thomson) comb. n.
orbitalis (Thomson) comb. n.
pectoralis (Thomson) comb. n.
puncticollis (Thomson) comb. n.

Tribe MESOSTENINI
Subtribe AGROTHEREUTINA
AP SHOPS Foerster

aquaticus (Thomson) comb. n.

THRYBIUS Townes
brevispina (Thomson) comb. n.
leucopygus (Gravenhorst)
drewseni (Thomson)

AGRO THEREUTES Foerster
abbreviator (Fabricius)

dispar (Thomson)
fumipennis (Gravenhorst)

zygaenarum (Thomson)
nasutus (Thomson) comb. n.
tibialis (Thomson)

MESOSTENIDEA Viereck
obnoxius (Gravenhorst)

subcircularis (Thomson)
subovalis (Thomson)

GAMBRUS Foerster
inferus Thomson
ornatulus (Thomson)
palustris (Thomson) comb. n.
superus Thomson

ARITRANIS Foerster

elegans (Thomson) comb. n.
fugitivus (Gravenhorst)

binotatulus (Thomson)
graefei (Thomson) comb. n.
mesoxanthus (Thomson) comb. n.
pulcher (Thomson) comb. n.

HWRYTA Foerster
frater (Cresson)

erythrocerus (Thomson)
fusiventris (Thomson)
sordidula (Thomson)

IDIOLISPA Foerster
tenuicornis (Thomson)

TR YCHOSIS Foerster
glabriculus (Thomson)
gradarius (Tschek)

nitidulus (Thomson)
ingratus (Tschek)

macrourus (Thomson)

92

M. G. FITTON

legator (Thunberg)

clypearis (Thomson)

pictus (Thomson)
mesocastanus (Tschek)

annulicornis (Thomson)
neglectus (Tschek)

annulitarsis (Thomson)
pauper (Tschek)

lapponicus (Thomson)
tristator (Tschek)

pleuralis (Thomson)

Subtribe HEDYCRYPTINA
EWCL/S/STownes
inflatus (Thomson) comb. n.
laticrm (Thomson) comb. n.
nubifer (Thomson) comb. n.
pubiventris (Thomson) comb. n.
striolatus (Thomson) comb. n.
tener (Thomson) comb. n.

ISCHNUS Gravenhorst
orbitatorius (Thomson)
punctiger (Thomson)

BUA THRA Cameron
tarsoleuca (Schrank)

curvicauda (Thomson)

ITAMOPLEX Foerster

arenicola (Thomson) comb. n.
dianae (Gravenhorst)

borealis (Thomson)
subquadratus (Thomson)
titubator (Thunberg)

infumatus (Thomson)

XYLOPHRURUS Foerster
apum (Thomson) comb. n.
coraebi (Thomson)

coroebi misspelling
lancifer (Gravenhorst)

dispar (Thunberg) (homonym)

dentifer (Thomson)
nigricornis (Thomson) comb. n.

MERINGOPUS Foerster
nigerrimus (Fonscolombe)

serratus (Thomson)
titillator (Linnaeus)

latitarsis (Thomson)

Subtribe MESOSTENINA
MESOSTENUS Gravenhorst
crassifemur Thomson
dentifer Thomson

Subtribe ATELEUTINA
A TELEUTE Foerster
lincaris Foerster

lissonotoides (Thomson)

Subtribe SPHECOPHAGINA
SPHECOPHAGA Westwood
vesparum (Curtis)

sericea (Thomson)

Nomina dubia in PHYGADEUONTINAE

HEMITELES Gravenhorst

australis Thomson (nomen dubium)
fuscicarpus Thomson (nomen dubium)
liambus Thomson (nomen dubium)
obtiquus Thomson (nomen dubium)
thomsoni Schmiedeknecht (nomen dubium)
dispar Thomson (homonym)

MICROCR YPTUS Thomson

ornaticeps Thomson (nomen dubium)

PEZOMACHUS Gravenhorst

numidicus Thomson (nomen dubium)

Subfamily BANCHINAE
Tribe GLYPTINI

APOPHUA Morley
cicatricosa (Ratzeburg)
crenulata (Thomson)

GLYPTA Gravenhorst
brevipetiolata Thomson
caudata Thomson
consimilis Holmgren

xanthognatha Thomson
crassitarsis Thomson

breviventris Thomson

tenuitarsis Thomson
dentifera Thomson
extincta Ratzeburg

nigriventris Thomson
femorator Desvignes

filicornis Thomson
heterocera Thomson
microcera Thomson
nigricornis Thomson
nigrina Desvignes

fractigena Thomson
nigroplica Thomson
solids Thomson
scutellaris Thomson
similis Bridgman

rufipes Thomson (homonym)

thomsonii Dalla Torre
tegularis Thomson
tenuicornis Thomson
varicoxa Thomson

Tribe LISSONOTINI

LISSONOTA Gravenhorst
antennalis Thomson
biguttata Holmgren

crassipes Thomson
buccator (Thunberg)

varicoxa Thomson
clypealis Thomson
coradna (Gmelin)

irrigua Thomson
digestor (Thunberg)

hians Thomson

THE ICHNEUMONIDAE DESCRIBED BY C. G. THOMSON

93

crrahunda Holmgren

punctiventris Thomson

? buccator (Thunberg)
folii Thomson
gracilipes Thomson
humerella Thomson
impressifrons Thomson
nigridens Thomson
palpalis (Thomson)
quadrinotata Gravenhorst

carinifrons Thomson
rimator Thomson
saturator (Thunberg)

basalts Thomson (homonym)

mutanda Schmiedeknecht
tenerrima Thomson
CRYPTOPIMPLA Taschenberg
genatts (Thomson)
subfumata (Thomson)

SYZEUCTUS Foerster
tenuifasciatus Schmiedeknecht

punctiventris (Thomson) (homonym)

Tribe BANCHINI

EXETASTES Gravenhorst
guttifer Thomson

BANCHUSFabricius
hastator (Fabricius)

femoralis Thomson

Subfamily CTENOPELMATINAE

Tribe CTENOPELMATINI

CTENOPELMA Holmgren
verticinum Thomson

XENOSCHESIS Foerster
mordax (Thomson)

HOMASPIS Foerster
robustus (Thomson)
varicolor (Thomson)

Tribe PIONINI

LETHADES Davis

curvispina (Thomson) comb. n.

scabriculus (Thomson)
HODOSTA TES Foerster

brevis (Thomson)

TREMA TOPYGUS Holmgren
lethierryi Thomson
melanocerus (Gravenhorst)
kriechbawneri Thomson

GL YPTORHAESTUS Thomson
punctatus (Thomson)
punctulatus (Woldstedt)
wuestneii (Thomson)

RHORUS Foerster
angulatus (Thomson)
longigena (Thomson)

PHA ES TVS Foerster
anomalus (Brischke)

heterocerus Thomson

SYNTACTUS Foerster
fusiformis (Thomson) comb. n.
scabriculus (Thomson) comb. n.

RHAESTUS Thomson
femoralis Thomson

SYMPHERTA Foerster
canaliculata (Thomson)
sulcata (Thomson)

ASTHENARA Foerster
crassifemur (Thomson)

Tribe PERILISSINI

LA THIPONUS Foerster
frigidus (Woldstedt)

pulcherrimus (Thomson)

SYNOCOETES Foerster
anterior (Thomson) comb. n.

PERILISSUS Holmgren
albitarsis Thomson
compressus Thomson
coxalis Thomson
emarginatus Thomson
frontator Thomson
spiniger Thomson

LA THROLESTES Foerster
caudatus (Thomson)
grandiceps (Thomson)
luteolus (Thomson)
marginatus (Thomson)
nigricollis (Thomson) comb. n.
pleuralis (Thomson)
ungularis (Thomson)

Tribe MESOLEHNI

O TLOPHOR US Foerster
hypomelas (Thomson)
melanocarm (Thomson)

LAGARO TIS Foerster
? didymus (Thomson)

BARYTARBES Foerster
annulipes (Thomson)
flavoscutellatus (Thomson)
laeviusculus (Thomson)

ALEXETER Forester
albilahris (Thomson)

PRO TA RCHUS Foerster
grandis (Thomson)
melanurus (Thomson)

PERISPUDA Foerster
flavitarsis (Thomson)

LAMACHUS Foerster
castaneiventris (Thomson)
thomsoni Fitton nom. n.

longiventris (Thomson) (homonym)

94

M. G. FITTON

NEOSTROBLIA Heinrioh
longigena (Thomson) comb. n.

SCOPESIS Foerster
depressus (Thomson)
macropus (Thomson)
tegularis (Thomson)

HIMERTA Foerster

bisannulata (Thomson)
RHINO TORUS Foerster

confusus (Thomson) comb. n.

mesocastanus (Thomson) comb. n.

subimpressus (Thomson) comb. n.

ARBELUSTownes

? sanguinipes (Thomson) comb. n.

CAMPODORUS Foerster

clypeatis (Thomson)

crassipes (Thomson)

crassitarsis (Thomson)

curtitarsis (Thomson)

deletus (Thomson)

galKcus (Thomson)

glyptus (Thomson)

humerellus (Thomson)

incident (Thomson)

laevipectus (Thomson)

latiscapus (Thomson)

liosternus (Thomson)

lobatus (Thomson)

mandihularis (Thomson)

netnati (Thomson)

nigridens (Thomson)

pineti (Thomson)

pleuraKs (Thomson)

? stenocerus (Thomson) comb. n.

tenuitarsis (Thomson)
SMICROLIUS Thomson

parvlcalcar (Thomson)

MESOLEWS Holmgren
brevipalpis Thomson
frenalis Thomson
frontatus Thomson
? immarginatus Thomson
incisus Thomson
obliquus Thomson
picticoxa Thomson
sinuatus Thomson
stenostigma Thomson
subroseus Thomson
rubidus Thomson
varicoxa Thomson

HYPERBA TVS Foerster
segmentator (Holmgren)
orbitalis (Thomson)

SAOT1S Foerster
brevispina (Thomson)
compressiusculus (Thomson)
dorsatus (Thomson)
emarginatus (Thomson)

heteropus (Thomson)

Kopleuris (Thomson)

longiventris (Thomson)

nigriscuta (Thomson)

nigriventris (Thomson)

tricolor (Thomson)

varicoxa (Thomson)
ANONCUSTownes

brachypus (Thomson)

brevitarsis (Thomson)

femorator (Thomson)

Nomen dubium in MESOLEIINI
MESOLEWS Holmgren

mesoxanthus Thomson (nomen dubium)

Tribe EURYPROCTINI
SYNOMELIX Foerster

curvulus (Thomson)
SYNODITES Foerster

facialis (Thomson) comb. n.

lineiger (Thomson) comb. n.

orbitalis (Thomson)

parviceps (Thomson)

subscaber (Thomson)

MESOLEPTIDEA Viereck
flavicornis (Thomson) comb. n.
holmgreni (Thomson) comb. n.

HA DRODA CTYL US Foerster
bidentulus Thomson
confusus (Holmgren)

albicoxa Thomson
femoralis (Holmgren)

nigricoxa (Thomson)
genalis Thomson
gracilipes Thomson
insignis (Kriechbaumer)

varicoxa (Thomson)
nigrifemur Thomson
tarsator Thomson
tiphae (Geoffroy)

laticeps Thomson
villosulus Thomson

SYNDIPNUS Foerster
atricornis (Thomson)
macrocerus (Thomson)
pectoralis Thomson
punctiscuta Thomson

PHOBETES Foerster

femorator (Thomson)

fulviventris (Thomson) comb. n.

latipes (Thomson) comb. n.

liopleuris (Thomson)

rufipes (Thomson) comb. n.
EUR YPROCTUS Holmgren

crassicornis Thomson

exareolatus Thomson

inferus Thomson

nitidulus Thomson

parvulus Thomson

THE ICHNEUMONIDAE DESCRIBED BY C. G. THOMSON

95

Subfamily CAMPOPLEGINAE

Tribe CAMPOPLEGINI
SINOPHORUS Foerster
costalis (Thomson)
crassifemur (Thomson)
fuscicarpus (Thomson)
? nigritellus (Thomson) comb. n.
pineticola (Thomson)
planiscapus (Thomson)
pleuralis (Thomson) comb. n.
rufifemur (Thomson)
tegularis (Thomson) comb. n.

CAMPOPLEX Gravenhorst

angulatus (Thomson)

bilobus (Thomson)

cerophagus Gravenhorst
picticrus (Thomson)

clypearis (Thomson)

contlnuus (Thomson)

cor acinus (Thomson)
forticosta (Thomson)
fuscipKca (Thomson)
fusicornis (Thomson) comb. n.

hadrocerus (Thomson)

liogaster (Thomson)

litoreus (Thomson)

lyratus (Thomson)

melampm (Thomson) comb. n.

ruficoxa (Thomson)

scaposus (Thomson) comb. n.

CASINARIA Holmgren
alpina Thomson
ischnogaster Thomson
monticola Thomson
orbitalis (Gravenhorst)

alboscutellaris Thomson
protensa Thomson
scabra Thomson
subglabra Thomson

Tribe LIMNERIINI

NEMERITIS Holmgren
caudatula Thomson
lativentris Thomson
stenura Thomson

BA THYPLECTES Foerster
anurus (Thomson)

contracta (Thomson)
balteatus (Thomson)
corvinus (Thomson)
curculionis (Thomson)
rostratus (Thomson)
stenostigma (Thomson)

BIOLYSIA Schmiedeknecht
immolator (Gravenhorst)

marginella (Thomson)
tristis (Gravenhorst)

trochantella (Thomson)

CALLIDORA Foerster

albovincta (Holmgren)

annellata (Thomson)
NEPIESTA Foerster

subclavata Thomson
GONOTYPUS Foerster

melanostoma (Thomson)
PYRACMON Holmgren

truncicola Thomson

lateralis Thomson
SPUDASTICA Foerster

kriechbaumeri (Bridgman)

petiolaris Thomson
SYNETAERIS Foerster

carbonella Thomson

heteropus Thomson
CAMPOLETIS Foerster

brachycera (Thomson)

erythropa (Thomson)

macroura (Thomson)

mucronella (Thomson)

variant (Thomson)
DUSONA Cameron

angustata (Thomson)

hifida (Thomson)

castanipes (Thomson) comb. n.

crassipes (Thomson)

flaviscapus (Thomson)

genalis (Thomson)

limnobia (Thomson)

luteipes (Thomson)

opaca (Thomson)

polita (Foerster)

latungula (Thomson)
splendens (Thomson)

recta (Thomson) comb. n.

spinipes (Thomson)

stenocarus (Thomson)
CYMODVSA Holmgren

convergens (Thomson)

longicalcar Thomson
DOLOPHRON Foerster

latiusculus (Thomson) comb. n.
PHOBOCAMPE Foerster

alticollis (Thomson)

confusa (Thomson)

flavicincta (Thomson)

pulchella (Thomson)
TRANOSEMA Foerster

exoleta (Thomson)

hyperborea (Thomson)

nigridens (Thomson)
striolata (Thomson)

rostralis (Brischke)

arenicola Thomson
ENYTUS Cameron

apostatus (Gravenhorst)

? crataegellae (Thomson)

96

M. G. FITTON

DIADEGMA Foerster
annulicrus (Thomson)
anura (Thomson)
brevivalvis (Thomson)
crassiseta (Thomson)
elongata (Thomson)
holopyga (Thomson)
hygrobia (Thomson)

ischnocera (Thomson)
lacticrus (Thomson)
latungula (Thomson)
maculata (Gra venhorst )

polyzona (Thomson)
majalix (Gravenhorst)

claripennis (Thomson)
mi'lania (Thomson)
micrura (Thomson)
monospila (Thomson)
parvicanda (Thomson)

parvicauda misspelling
rimator (Thomson)
sordipes (Thomson)
specularis (Thomson)
tenuipes (Thomson)
trochanterata (Thomson)
truncata (Thomson)

subbuccata (Thomson)

HYPOSOTER Foerster

hoops (Thomson)

coxator (Thomson) comb. n.

facialis (Thomson)

leucomerus (Thomson)

longulus (Thomson) comb. n.

neglectus (Holmgren)
varicoxa (Thomson)

pectinatus (Thomson)

picticollis (Thomson)

ruficrus (Thomson)

tenuicosta (Thomson)

vividus (Holmgren)

albicrus (Thomson)
OLESICAMPE Foerster

alboplica (Thomson)

annulitarsis (Thomson) comb. n.

basalts (Thomson)

bergmanni (Thomson) comb. n.

binotata (Thomson)

buccata (Thomson) comb. n.

cavigena (Thomson)

crassitarsis (Thomson)

curtigena (Thomson) comb. n.

femoretta (Thomson)

flavicornis (Thomson)

frutetorum (Thomson)

fuUrans (Thomson)

geniculella (Thomson)

gracittpes (Thomson)

heterogaster (Thomson) comb. n.

luteipes (Thomson)

melanogaster (Thomson comb. n.

nigricoxa (Thomson)
nigroplica (Thomson)
patellana (Thomson)
punctitarsis (Thomson)
radiella (Thomson)
retusa (Thomson)
simplex (Thomson)
sinuata (Thomson) comb. n.
spireae (Thomson) comb. n.
sternella (Thomson)
subcallosa (Thomson)
tarsator (Thomson) comb. n.

LA THROSTIZUS Foerster

forticanda (Thomson)

forticauda misspelling

macrostoma (Thomson)

monilicornis (Thomson)

punctipes (Thomson)

sternocerus (Thomson)
ECHTHRONOMAS Foerster

quadrinotata (Thomson)

Subfamily CREMASTINAE

PRISTOMERUS Curtis
palli Jus Thomson

DIMOPHORA Foerster
evanialis (Gravenhorst)

annellatus (Thomson)
rohusta Brischke

arenicola (Thomson)

CREMASTUS Gravenhorst
crassicornis Thomson
Kneatus Gravenhorst

radialis Thomson
pungens Gravenhorst

laeviusculus Thomson
TEMELUCHA Foerster
guttifer (Thomson)
ophthalmica (Holmgren)

macrostigma (Thomson)
schoenobia (Thomson)
subnasuta (Thomson)

Subfamily TERSILOCHINAE

MICRODIAPARSIS Horstmann

versutus (Holmgren)

parviceps (Thomson)
BARYCNEMIS Foerster

anurus (Thomson)

caudatulus (Thomson)

filicornis (Thomson)

gracillimus (Thomson)

laeviceps (Thomson)
GONOLOCHUS Foerster

fenestralis (Thomson)
PECTINOLOCHUS Aubert

ensifer (Brischke)

crassicauda (Thomson)

THE ICHNEUMONIDAE DESCRIBED BY C. G. THOMSON

97

striolus (Thomson)
RUGODIAPARS1S Horstmann

crassipes (Thomson)
TERSILOCHUS Holmgren

filicornis (Thomson)

heterocerus (Thomson)

liopleuris (Thomson)

longicornis (Thomson)

melanogaster (Thomson)

obliquus (Thomson)

subdepressus (Thomson)

PHRADIS Foerster
brevis (Brischke)

temporalis (Thomson)
decrescent (Thomson)
interstitialis (Thomson)
HETEROCOLA Foerster
proboscidalis (Thomson)
monticola (Thomson)
pallicarpus (Thomson)

DIAPARSIS Foerster
carinifer (Thomson)
nutritor (Fabricius)

genalis (Thomson)
stramineipes (Brischke)
flavicornis (Thomson)

NANODIAPARSIS Horstmann
apertus (Thomson)

ANEUCLIS Foerster
brevicauda (Thomson)
incident (Thomson)
maritimus (Thomson)

Subfamily OPHIONINAE

OPHION Fabricius
distant Thomson
longigena Thomson
scutellaris Thomson

Subfamily MESOCHORINAE

ASTIPHROMMA Foerster
buccatutn (Thomson)
graniger (Thomson)
hamulum (Thomson)
incidens (Thomson)
mundibulare (Thomson)
plagiatum (Thomson)
simplex (Thomson)
tenuicornis (Thomson)

MESOCHORUS Gravenhorst
acuminatus Thomson
albipes Thomson
angustatus Thomson
brevicollis Thomson
brevigena Thomson
crassicrus Thomson
curvicauda Thomson
curvulus Thomson

fulvus Thomson
lapponicus Thomson
longicauda Thomson
macrurus Thomson
marginatus Thomson
nigriceps Thomson
orgyiae Dalla Torre

stigmaticus Thomson (homonym)
picticrus Thomson
punctipleuris Thomson
salicis Thomson
suedcus Dalla Torre

pectinipes Thomson (homonym)
temporalis Thomson
tenuiscapus Thomson
tuberculiger Thomson

STICTOPISTHUS Thomson
hilineatm (Thomson)
convexicollis (Thomson)
laticeps (Thomson)

Subfamily METOPIINAE

CHOR1NAEUS Holmgren
australis Thomson (nomen dubium)
brevicalcar Thomson
longicalcar Thomson
longicornis Thomson

77?/£C£STownes

facialis (Thomson)
nitifrons (Thomson)

METOPWS Panzer
brevispina Thomson
clypealis Thomson
croceicornis Thomson
interruptus Thomson

TRICLISTUS Foerster
albicinctus Thomson
areolatus Thomson
facialis Thomson
lativentris Thomson
longicalcar Thomson
pallipes Holmgren

nitrifrons Thomson
pubiventris Thomson
spiracularis Thomson

EXOCHUS Gravenhorst
annulitarsis Thomson
anospilus Thomson
australis Thomson
citripes Thomson
crassicornis Thomson
incidens Thomson
lineifrons Thomson
longicornis Thomson
nigripalpis Thomson
parvispina Thomson
signifrons Thomson

nigrifrons misspelling

98

M. G. FITTON

Subfamily ANOMALONINAE
Tribe THERIONINI

BARYLYPA Foerster

delict or (Thunberg)

genalis (Thomson)
pallida (Gravenhorst)

laticeps (Thomson)
THERION Curtis
giganteum (Gravenhorst)

pyramidatus (Thomson)

GRAVENHORSTIA Boie (ERIGORGUS
Foerster)
cerinops (Gravenhorst)

lapponicum (Thomson)
claripennis (Thomson)
fibulator (Gravenhorst)

annulitarse (Thomson)
orhitale (Thomson)
varicorne (Thomson)

AGRYPON Foerster
rugifer (Thomson)
stenostigma (Thomson)

Subfamily ACAENITINAE

COLEOCENTRUS Gravenhorst
heteropus Thomson

Subfamily OXYTORINAE

MICROLEPTES Gravenhorst
glabriventris (Thomson) comb. n.
rectangulus (Thomson) comb. n.

OXYTORUS Foerster
armatus Thomson

ALLOMACRUS Foerster
pimplarius Thomson

PROCLITUS Foerster
heterocerus (Thomson)
longitarsis (Thomson)

DIALIPSIS Foerster

exilis Foerster

crassipes (Thomson)
PLECTISCIDEA Viereck

bistriatus (Thomson)

curticauda (Thomson) comb. n.

eurystigma (Thomson)

subteres (Thomson)

APERILEPTUS Foerster
obliquus (Thomson)

BLAPTICUS Foerster
crassulus Thomson
dentifer Thomson

SYMPLEC1S Foerster
facialis Thomson

EUSTERINX Foerster
pusilla (Zetterstedt)

trichops (Thomson)

HELICTESHaliday

pilicornis (Thomson)

MEGASTYLUS Schiodte
pleuralis Thomson

Subfamily COLLYRIINAE

COLLYRIASchi0dle
coxator (Villers)

puncticeps (Thomson)
trichophthalma (Thomson)

Subfamily ORTHOCENTRINAE

OR THOCENTR US Gravenhorst
petiolaris Thomson
radialis Thomson

PICROSTIGEUS Foerster
recticauda (Thomson)

STENOMA CR US Foerster
cubiceps (Thomson)
curvulus (Thomson)
deletus (Thomson)
exserens (Thomson)
fortipes (Thomson)
innotatus (Thomson)
superus (Thomson)
ungula (Thomson)

LEIPAULUSTownes

Iflavicornis (Thomson) comb. n.

NEURA TELES Ratzeburg
compressus (Thomson) comb. n.
crassicornis (Thomson) comb. n.
falcatm (Thomson) comb. n.

Subfamily DIPLAZONTINAE

SYRPHOCTONUS Foerster

crassicornis (Thomson) comb. n.
brevicornis (Thomson)

crassicrus (Thomson)

incisus (Thomson) comb. n.

longiventris (Thomson)

megaspis (Thomson) comb. n.

signatus (Gravenhorst)

hygrobius (Thomson)
ENIZEMUM Foerster

nigricornis (Thomson)
CAMPOCRASPEDON Uchida

caudatus (Thomson)

PHTHORIMA Foerster

xanthaspis (Thomson) comb. n.

DASCH1A Diller
brevitarsis (Thomson)

DIPLAZONNees

deletus (Thomson)
varicoxa (Thomson)

PROMETHES Foerster
melanaspis (Thomson)
nigriventris (Thomson)

THE ICHNEUMONIDAE DESCRIBED BY C. G. THOMSON

99

SUSS ABA Cameron
cognata (Holmgren)

albicoxa (Thomson)
pulchella (Holmgren)

laticarpus (Thomson)
punctiventris (Thomson)

Subfamily ICHNEUMONINAE
Tribe PROTICHNEUMONINI
COELICHNEUMON Thomson

anospllus (Thomson) comb. n.

coactus (Thomson)

decrescens (Thomson)

tiocnemis (Thomson)

tenuitarsis (Thomson) comb. n.

truncatulus (Thomson)

Tribe LISTRODROMINI
ANISOBASWesmad

parviceps (Thomson)
platystylus (Thomson)

Tribe ICHNEUMONINI
STENICHNEUMON Thomson
simulosus (Thomson)

rimulosus misspelling

CR A TICHNEUMON Thomson
albiscuta (Thomson)
anotylus (Thomson) comb. n.
cyaneoviridis (Thomson)
grandiceps (Thomson) comb. n.
liostylus (Thomson)
pallitarsis (Thomson)
parviscopa (Thomson) comb. n.
stenocarus (Thomson)

EUPALAMUS Wesmael
wesmaeli (Thomson)

BARICHNEUMON Thomson
digrammus (Gravenhorst)
nudicoxa (Thomson)
mesostilpnus (Thomson) comb. n.

ICHNEUMON Linnaeus
acuticornis Thomson
acquit alcar Thomson
arctobius Thomson
boreellus Thomson
chrysostomus Thomson
corfitzi Thomson

jesperi Thomson
crassifemur Thomson
crassitarsis Thomson
eurycerus Thomson
gibbulus Thomson
grandicornis Thomson
hypolius Thomson
inquinatus Wesmael

brevigena Thomson
leucopeltis Thomson
longeareolatus Thomson
macrocerus Thomson

micropnygus Thomson

spiracularis Thomson (homonym)
minutorius Desvignes

captorius Thomson
monospilus Thomson
nereni Thomson
nordenstromi Thomson
quadriannellatus Thomson

quadriannulatus Thomson (homonym)
quinquenotatus Thomson
stenocerus Thomson
suhquadratus Thomson
trispilus Thomson
xanthognathus Thomson

CTENICHNEUMON Thomson

circulator (Thomson)
SPILICHNEUMON Thomson

limnophilus (Thomson) comb. n.

simplicidens (Thomson)

stagnlcola (Thomson)
SPILOTHYRA TELES Heinrich

truncicola (Thomson) comb. n.
DIPHYUS Kriechbaumer

longigena (Thomson)

triplicates (Thomson) comb. n.

AMBL YTELES Wesmael

anurus Thomson (nomen dubium)

7 R1C HO LA BUS Thomson
femoralis (Thomson)

Tribe EURYLABINI
EUR YLABUS Wesmael
larvatus (Christ)

vinulator Thomson

Tribe PLATYLABINI

PLA TYLABUS Wesmael
concinnus Thomson
muticus Thomson
opaculus (Thomson)
punctifrons (Thomson)
transversus Bridgman
lativentris Thomson

ASTHENOLABUS Heinrich
latiscapus (Thomson)

Tribe PHAEOGENINI

HETER1SCHNUS Wesmael

coxator (Thomson) comb. n.

pulchellus (Thomson) comb. n.
HEMICHNEUMON Wesmael

fuscipes Thomson
TRACHYARUS Thomson

corvinus Thomson
DIC A ELO TUS Wesmael

annellatus Thomson

crassifemur Thomson

inflexus Thomson

orbitalis Thomson

100

M. G. FITTON

pentagonus (Thomson) comb. n.
punctiventris (Thomson)

EPITOMUS Foerster
parvus Thomson

DIADROMUS Wesmael
arctlcus Thomson
medialis Thomson

COLPOGNA THUS Wesmael
armatus Thomson
divisus Thomson

CENTETERUS Wesmael
nigricornis Thomson

EPARCES Foerster
grandiceps Thomson

AETHECERUS Wesmael
graniger Thomson
pallicoxa Thomson

DIROPHANES Foerster
ruficoxa (Thomson)

BAEOSEMUS Foerster
oenescens (Thomson)

aenescens misspelling

PHAEOGENES Wesmael
crassidens Thomson
elongatus Thomson
montanus Thomson
tegularis Thomson

Species incorrectly attributed to Thomson

For various reasons the following species have been incorrectly attributed to Thomson.

Campoplex auriculatus Foerster (Thomson, 1887c: 1071). For reasons not stated Aubert (1966: 130) at-
tributed this species to Thomson and 'designated' a 'lectotype' for it.

Glypta tenuiventris. See catalogue entry for G. tenuitarsis.

Microcryptus nigriventris. Aubert's reference (1966: 129) to this species is presumably an error for Meso-
cryptus nigriventris (Thomson).

Pimplu lapponica Zetterstedt (Thomson, 1877: 746). This is technically an 'incorrect subsequent spelling'
(Code, Article 33) of Pimpla arctica Zetterstedt. Thomson himself (18886: 1250) drew attention to the
error.

Trichomastix pallipe s Holmgren (Thomson, 1890: 1473). This is technically an 'incorrect subsequent spell-
ing' of Bassusflavipes Holmgren. Dalla Torre (1901 : 242) first drew attention to the error.

Acknowledgements

For the loan of specimens in their care and for the provision of information I am grateful to the
following: H. Andersson, R. W. Carlson, E. H. Diller, K. J. Hedqvist, R. Hinz, K. Horstmann. M.
Idar, A. G. Irwin, B. Petersen, G. van Rossem, J. Sawoniewicz, H. K. Townes, H. W. Walden and
C. J. Zwakhals. I am deeply indebted to Roy Danielsson for his help and hospitality during my
visits to Lund in 1978 and 1980. He made available a great deal of information on Thomson's life,
methods and collections which otherwise would have been difficult or impossible for me to
obtain. His advice and opinions on matters such as the interpretation of labels were invaluable. I
also benefited from discussion of various points with my colleagues at the British Museum
(Natural History), particularly Ian D. Gauld. Finally, I wish to thank the late Miss V. Dick for
typing the manuscript.

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- 1960. Les Ichneumonides des Pyrenees-Orientales. Vie Milieu 11 : 473-493.

THE ICHNEUMONIDAE DESCRIBED BY C. G. THOMSON 101

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1968. Revision einiger Arten der Gattungen Mesostenus Gravenhorst, Agrothereutes Foerster und

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1969. Typenrevision der europaischen Arten der Gattung Diadegma Foerster (syn. Angitia Holm-

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1973a. Revision der westpalaarktischen Arten der Gattung Nemeritis Holmgren (Hymenoptera,

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19736. Revision der europaischen Arten der Gattung Dichrogaster Doumerc (Hym. Ichneumon-

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1973c. Revision der Gattung Nepiesta Foerster (mit einer Obersicht iiber die Arten der Gattung

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1974a. Revision der westpalaarktischen Arten der Schlupfwespen-Gattungen Bathyplectes und Bioly-

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19746. Typenrevision der von Strobl in der Gattung Hemiteles Gravenhorst s.l. beschriebenen

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1974c. Typenrevision der von E. Zilahi-Kiss beschriebenen Hemitelinen mit Bemurkungen zu den

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19766. Nachtrag zur Revision der europaischen Dichrogaster-Arten (Hymenoptera, Ichneumon-

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1977. Bemerkungen zur Systematik einiger Gattungen der Campopleginae II (Hymenoptera, Ich-

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1978. Revision der Gattungen der Mastrina Townes (Hymenoptera, Ichneumonidae, Hemitelinae).

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THE ICHNEUMONIDAE DESCRIBED BY C. G. THOMSON 103

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Jussila, R. 1965. The Ichneumonidae of the Kevojoki area in Inari Lapland (Finland). Annls Univ. turku
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1942. Second review of literature concerning British Ichneumonidae (Hym.), with notes on Palaearc-

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1962. Systematic notes on Tryphonine Ichneumonidae (Hym.). Opusc. ent. 27: 45-56.

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104 M. G. FITTON

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THE ICHNEUMONIDAE DESCRIBED BY C. G. THOMSON

105

Index

abbreviator (Agrothereutes) 80, 91

abbreviatus (Ephialtes, Dolichomitus) 28, 87

abditus (Tryphon) 86, 88

Acaenitinae 98

Acanthocryptus 10, 11

aciculatus (Adelognathus) 1 1, 88

Aclastus 37, 40, 89

Aconias 77,91

Acrolyta 36, 89

Acrolytina 89

Acropimpla 76, 87

Acrotomus 27, 88

acuminatus (Mesochorus) 52, 97

acuticornis (Ichneumon) 43, 99

acutipennis (Phygadeuon) 70, 90

Adelognathinae 88

Adelognathus 11,88

adspersus (Exenterus) 30, 88

aenescens (Baeosomus, Baeosemus) 19, 100

aeneus (Hemiteles, Gelis) 35, 89

aequicalcar (Ichneumon) 43, 99

aereus (Bathythrix) 48, 90

aestivalis (Dichrogaster) 37, 89

Aethecerus 12, 100

Agasthenes 40, 89

Agrothereutes 80, 8 1,91

Agrothereutina91

Agrypon 17,98

albicinctus (Triclistus) 86, 97

albicoxa (Cteniscus, Eridolius) 26, 88

albicoxa (Hadrodactylus) 34, 94

albicoxa (Polyblastus, Nemioblastus) 77, 78, 8i

albicoxa (Promethus, Sussaba) 79, 99

albicrus (Anilasta, Hyposoter) 16, 96

albilabris (Mesoleius, Alexeter) 55, 93

albipalpus (Hemiteles, Gelis) 35, 89

albipes (Mesochorus) 52, 97

albiscuta (Ichneumon, Cratichneumon) 43, 99

albitarsis (Perilissus, Ecclinops) 68, 93

alboplica (Olesicampa, Olesicampe) 63, 96

alboscutellaris (Casinaria) 22, 95

albosignatus (Ichneumon) 46

albovincta (Callidora) 20, 95

Alexeter 55, 93

Allomacrusl2,98

alpina (Casinaria) 22, 95

alpinus (Hemiteles, Phygadeuon) 35, 90

alternans (Itoplectis) 76, 87

alticollis (Phobocampa, Phobocampe) 70, 95

alutaceus (Atractodes, Polyrhembia) 18, 91

alutaceus (Microcryptus, Pleolophus) 60, 91

alysiina (Macrochasmus, Idiogramma) 51, 88

Ambly teles 12,99

Amphibulus 24, 90

analis (Stylocryptus, Endasys) 81, 90

Aneuclis 83, 84, 97

Angitia 13, 14, 15, 16

angulata (Omorga, Campoplex) 65, 95
angulatus (Monoblastus, Rhorus) 62, 93
angulatus (Polyblastus, Scopiorus, Ctenochira) 78,

88

angustatus (Campoplex, Dusona) 20, 95
angustatus (Mesochorus) 52, 97
angustatus (Stilpnus) 81,91
angustipennis (Atractodes) 18, 91
Anilasta 16, 17
Anisobas 12, 99
annellata (Callidora) 20, 95
annellatus (Demophorus, Dimophora) 27, 96
annellatus (Dicoelotus, Dicaelotus, Cinxaelotus)

28,99

annulicornis (Goniocryptus, Trychosis) 33, 92
annulicornis (Phygadeuon, Theroscopus) 70, 90
annulicrus (Angitia, Diadegma) 13, 96
annulipes (Mesoleius, Barytarbus, Barytarbes) 55,

93
annulitarse (Anomalon, Gravenhorstia, Erigorgus)

17,98

annulitarsis (Cratocryptus, Cubocephalus) 24, 91
annulitarsis (Exochus) 30, 97
annulitarsis (Groniocryptus, Trychosis) 34, 92
annulitarsis (Holocremna, Olesicampe) 40, 96
Anomalon 17
Anomaloninae 98
anomalus (Phaestus) 70, 93
Anoncus 55, 56, 94
anospilus (Exochus) 30, 97
anospilus (Ichneumon, Coelichneumon) 43, 99
anotylus (Ichneumon, Cratichneumon) 43, 99
antefurcalis (Ephialtes, Townesia) 28, 87
antennalis (Lissonota) 49, 92
anterior (Syndipnus, Polyrhysius, Synocoetes) 82,

93

anura (Angitia, Diadegma) 13, 96
anura (Canidia, Bathyplectes) 21, 22, 95
anurus (Amblyteles) 12, 99
anurus (Phygadeuon, Ceratophygadeuon) 71, 73,

90

anurus (Porizon, Barycnemis) 78, 96
Apechthis 76, 87
Aperileptus 77, 98

apertus (Hemiteles, Gnypetomorpha) 35, 89
apertus (Thersilochus, Nanodiaparsis) 83, 97
apicalis (Grypocentrus) 34, 88
Apophua 32, 92

apostatus (Enytus, Diadegma) 14, 95
Apsilops85,91
Aptesis55,61,91

apum (Caenocryptus, Xylophrurus) 20, 92
aquaticus (Trichocryptus, Apsilops) 85, 91
Arbelus81,94
arctica (Pimpla) 100
arcticus (Diadromus) 27, 100
arcticus (Hemiteles, Phygadeuon), 35, 90

106

INDEX

arctobius (Ichneumon) 43, 99

arenicola (Cryptus, Itamoplex) 25, 92

arenicola (Demophorus, Dimophora) 27, 96

arenicola (Erromenus) 29, 88

arenicola (Tranosema) 85, 95

areolaris (Hemiteles, Charitopes) 35, 90

areolaris (Microcryptus, Javra) 61,91

areolatus (Triclistus) 86, 97

aries (Microcryptus, Schenkia) 61, 91

Aritranis42,43,91

armatulus (Phygadeuon, Medophron) 71, 90

armatus (Colpognathus) 24, 100

armatus (Isadelphus) 36, 89

armatus (Oxytorus) 67, 98

Arotrephes 73, 89

Asthenara 17,93

Asthenarus 17, 23

Asthenolabus 77, 99

Astiphromma 52, 53, 54, 97

Astiphrommus 52, 53, 54

Asyncrita 18

Ateleute 38, 92

Ateleutina 92

Atractodes 18, 19,91

atricornis (Euryproctus, Syndipnus) 29, 94

auricularis (Tryphon) 86, 88

auriculatus (Campoplex) 100

auriculatus (Delotomus, Acrotomus) 27, 88

auriculatus (Hemiteles, Zoophthorus) 36, 89

australis (Chorinaeus) 23, 97

australis (Exochus) 30, 97

australis (Hemiteles) 36, 92

Baeosemus 19, 100

Baeosomus 19

balteata (Canidia, Bathyplectes) 22, 95

balteatus (Hemiteles, Gelis) 36, 89

Banchinae 92

Banchini 93

Banchus 19, 93

Barichneumon 45, 46, 99

Barycnemis 78, 79, 96

Barylypal7,98

Barytarbes 55, 56, 57, 93

Barytarbus 55, 56, 57

basalis Brischke (Lissonota) 49

basalis Thomson (Lissonota) 49, 93

basalis (Olesicampa, Olesicampe) 63, 96

Bassus 19, 100

Bathyplectes 21, 22, 62, 95

Bathythrix 48, 90

Bathytrichina 90

bellicornis (Hemiteles, Handaoia) 36, 89

bergmanni (Holocremna, Olesicampe) 41, 96

biannulatus (Sulcarius) 37, 90

bidens (Phygadeuon) 71, 90

bidentulus (Hadrodactylus) 34, 94

bidentulus (Hemiteles, Isadelphus) 36, 89

bidentulus (Tryphon) 86, 88

bifidus (Campoplex, Dusona) 20, 95

biguttata (Lissonota) 50, 92

bilineatus (Mesochorus, Stictopisthus) 52, 97

biloba (Omorga, Campoplex) 65, 95

binotata (Olesicampa, Olesicampe) 63, 96

binotatulus (Hoplocryptus, Aritranis) 42, 91

binotatus (Delotomus, Cycasis) 27, 88

Biolysia 22, 62, 95

bisannulatus (Euryproctus, Himertus, Himerta) 29,

94

bisinuata (Ctenochira) 78, 88
bispinus (Cratocryptus, Amphibulus) 24, 90
bistriatus (Plectiscus, Plectiscidea) 77, 98
Blapticusl9,98

boops (Anilasta, Hyposoter) 16, 96
borealis (Cryptus, Itamoplex) 25, 92
borealis (Microcryptus, Aptesis) 61, 91
boreellus (Ichneumon) 43, 99
brachycera (Pimpla, Exeristes) 75, 87
brachycera (Sagaritis, Campoletis) 80, 95
brachycerus (Paniscus, Netelia) 68, 87
Brachycryptus 19

brachypus (Mesoleius, Anoncus) 55, 94
brachyurus (Phygadeuon) 71, 90
breviareolatus (Hemiteles, Stibeutes) 36, 90
brevicalcar (Chorinaeus) 23, 97
brevicauda (Hemiteles, Gelis) 36, 89
brevicauda (Thersilochus, Aneuclis) 83, 97
breviceps (Pezomachus, Gelis) 69, 89
brevicollis (Mesochorus) 52, 97
brevicornis (Adelognathus) 11, 88
brevicornis (Homoporus, Syrphoctonus) 41, 42,

98

brevicornis (Scambus) 76, 87
brevigena (Cteniscus, Eridolius) 26, 88
brevigena (Ichneumon) 44, 99
brevigena (Mesochorus) 52, 97
brevipalpis (Mesoleius) 55, 94
brevipetiolata (Glypta) 32, 92
brevipetiolatus (Exyston) 31, 88
brevis (Hodostatus, Hodostates) 40, 93
brevis (Leptocryptus, Bathythrix) 48, 90
brevis (Phradis) 85, 97
breviscapus (Atractodes) 18, 91
brevispina (Hygrocryptus, Thrybius) 43, 91
brevispina (Mesoleius, Saotus, Saotis) 55, 94
brevispina (Metopius) 60, 97
brevitarsis (Erromenus) 29, 88
brevitarsis (Homoporus, Daschia) 41, 98
brevitarsis (Mesoleius, Anoncus) 55, 94
brevitarsis (Phygadeuon) 71, 90
brevivalvis (Angitia, Diadegma) 13, 96
breviventris (Cteniscus, Eridolius) 26, 88
breviventris (Glypta) 32, 92
Buathra 25, 92

buccata (Holocremna, Olesicampe) 41, 96
buccator (Lissonota) 51, 92, 93
buccatus (Mesochorus, Astiphrommus,

Astiphromma) 52, 97

INDEX

107

Cacotropa 19

Caenocryptus 20

calcaratus (Delotomus, Kristotomus) 27, 88

calcaratus (Exyston) 31, 88

Callidora 20, 95

Campocraspedon 42, 98

Campodorus 55, 56, 57, 58, 59, 81, 94

Campoletis 80, 95

Campopleginae 95

Campoplegini 95

Campoplex 20, 21, 48, 65, 66, 95, 100

canaliculatus (Cataglyptus, Stiphrosomus,

Sympherta) 23, 93
canaliculatus (Phygadeuon) 71, 90
Canidia21,22

capra (Hemiteles, Mastrulus) 36, 89
capreolus (Hemiteles, Acrolyta) 36, 89
captorius (Ichneumon) 44, 99
carbonella (Synetaeris, Pyracmon) 83, 95
carinatus (Exyston) 31, 88
carinifer (Thersilochus, Diaparsis) 83, 97
carinifrons (Lissonata) 49, 93
Casinaria 22, 23, 95

castaneiventris (Mesoleius, Lamachus) 55, 93
castanipes (Campoplex, Dusona) 20, 95
Catoglyptus 23
Catomicrus 23
caudata (Glypta) 32, 92
caudata (Polyspincta, Sinarachna) 78, 87
caudatula (Nemeritis) 62, 95
caudatulus (Medophron) 71, 90
caudatulus (Porizon, Cratophion, Barycnemis) 79,

96

caudatus (Homoporus, Campocraspedon) 41, 98
caudatus (Lathrolestus, Lathrolestes) 47, 93
caudatus (Phygadeuon, Medophron) 71, 90
cavigena (Erromenus) 29, 88
cavigena (Olesicampa, Olesicampe) 63, 96
Centeterus 23, 100

ceratophorus (Tryphon, Cosmoconus) 86, 88
Ceratophygadeuon 71, 73, 90
cerinops (Gravenhorstia, Erigorgus) 17, 98
cerophagus (Campoplex, Sesioplex) 66, 95
Charitopes 35, 36, 38, 90
Chirotica 37, 39, 89
Chiroticina 89
Chorinaeus 23, 24, 97
chrysostomus (Ichneumon) 44, 99
cicatricosa (Apophua) 32, 92
Cinxaelotus 28

circulator (Amblyteles, Ctenichneumon) 12, 99
citripes (Exochus) 31, 97
claripennis (Angitia, Diadegma) 14, 96
claripennis (Anomalon, Gravenhorstia, Erigorgus)

17,98

claripennis (Exenterus) 30, 88
clausus (Hemiteles, Charitopes) 36, 90
clavicornis (Pimpla, Itoplectis) 75, 87
clavipes (Leptocryptus, Leptocryptoides) 48, 90

clypealis (Lissonota) 50, 92

clypealis (Mesoleius, Campodorus) 55, 94

clypealis (Metopius) 60, 97

clypealis (Stylocryptus, Gnathocryptus,
Glyphicnemis) 82, 90

clypearis (Goniocryptus, Trychosis) 34, 92

clypearis (Lathroplex, Campoplex) 48, 95

Clypeoteles 37, 40, 89

Cnemischus 1 1

coactus (Ichneumon, Coelichneumon) 44, 99

Coelichneumon 43, 44, 45, 47, 99

cognata (Sussaba) 79, 99

cognator (Dolichomitus) 28, 87

Coleocentrus 24, 98

collaris (Leptocryptus, Bathythrix) 48, 90

collaris (Thymarus, Thymaris) 85, 88

Collyria 67, 98

Collyriinae 98

Colpognathus 24, 100

compressiusculus (Mesoleius, Saotus, Saotis) 55,
94

compressus (Atractodes) 18, 91

compressus (Orthocentrus, Stenomacrus,

Neurateles) 66, 98

compressus (Perilissus, Ecclinops) 68, 93
compressus (Thymarus, Thymaris) 85, 88
concinnus (Platylabus) 76, 99
confusa (Phobocampa, (Phobocampe) 70, 95
confusus (Hadrodactylus) 34, 94
confusus (Mesoleius, Spudaeus, Rhinotorus) 55, 94
consimilis (Glypta) 33, 92
constrictus (Hemiteles, Xiphulcus) 36, 89
continua (Omorga, Campoplex) 65, 95
continuus (Phytodietus) 75, 87
contracta (Canidia, Bathyplectes) 22, 95
convergens (Nemeritis, Cymodusa) 62, 95
convexicollis (Mesochorus, Stictopisthus) 52, 97
coracina (Lissonota) 50, 92
coracina (Omorga, Campoplex) 65, 95
coraebi (Macrocryptus, Xylophrurus) 51, 92
corfitzi (Ichneumon) 44, 45, 99
coroebi (Macrocryptus, Xylophrurus) 51, 92
corvina (Canidia, Bathyplectes) 22, 95
corvinus (Trachyarus) 85, 99
Cosmoconus 86, 88
costalis (Hemiteles, Mastrus) 36, 89
costalis (Limneria, Sinophorus) 48, 95
costulatus (Smicroplectrus) 80, 88
coxalis (Perilissus, Spanotecnus) 68, 93
coxator (Anilasta, Hyposoter) 16, 96
coxator (Collyria) 67, 98
coxator (Ischnus, Heterischnus) 47, 99
crassicauda (Thersilochus, Pectinolochus) 83, 96
crassiceps (Ephialtes, Dolichomitus) 28, 87
crassicornis (Atractodes) 18, 91
crassicornis (Cremastus) 24, 96
crassicornis (Euryproctus) 29, 94
crassicornis (Exochus) 31, 97
crassicornis (Homoporus, Syrphoctonus) 41, 42, 98

108

INDEX

crassicornis (Orthocentrus, Stenomacrus,

Neurateles) 66, 98
crassicornis (Stilpnus) 81, 91
crassicrus (Homoporus, Syrphoctonus) 42, 98
crassicrus (Mesochorus) 52, 97
crassidens (Phaeogenes) 69, 100
crassifemur (Asthenarus, Asthenara) 17, 93
crassifemur (Dicoelotus, Dicaelotus, Cinxaelotus)

28,99

crassifemur (Ichneumon) 44, 99
crassifemur (Limneria, Sinophorus) 49, 95
crassifemur (Mesostenus) 60, 92
crassipes (Campoplex, Dusona) 21, 95
crassipes (Lissonota) 50, 92
crassipes (Mesoleius, Campodorus) 55, 94
crassipes (Plectiscus, Dialipsis) 77, 98
crassipes (Thersilochus, Rugodiaparsis) 83, 97
crassiseta (Angitia, Diadegma) 14, 96
crassiseta (Ephialtes, Liotryphon) 28, 29, 87
crassitarsis (Glypta) 32, 33, 92
crassitarsis (Ichneumon) 44, 99
crassitarsis (Mesoleius, Campodorus) 56, 94
crassitarsis (Olesicampa, Olesicampe) 63, 96
crassitarsis (Phytodietus) 75, 87
crassulus (Blapticus) 19, 98
crataegellae (Angitia, Diadegma, Enytus, Dioctes)

14,95

Cratichneumon 43, 45, 46, 77, 99
Cratocryptus 24
Cratophion 79
Cremastinae 96
Cremastus 24, 25, 96
crenulata (Glypta, Apophua) 32, 92
cristatus (Parabatus, Netelia) 68, 87
croceicornis (Atractodes) 18, 91
croceicornis (Metopius, Peltocarus) 60, 97
Cryptopimpla 50, 51,93
Cryptus 25
Ctenacmus 78
Ctenichneumon 12,99
Cteniscus 26, 27, 88
Ctenochira 78, 88
Ctenopelma 26, 93
Ctenopelmatinae 93
Ctenopelmatini 93

cubiceps (Orthocentrus, Stenomacrus) 66, 98
cubiceps (Phygadeuon) 71, 90
Cubocephalus 24, 61, 81, 91
curculionis (Canidia, Bathyplectes) 22, 95
curticauda (Plectiscus, Plectiscidea) 77, 98
curtigena (Holocremna, Olesicampe) 41, 96
curtitarsis (Mesoleius, Campodorus) 56, 94
curvicauda (Cryptus, Buathra) 25, 92
curvicauda (Mesochorus) 52, 97
curviscapus (Phygadeuon) 71, 90
curvispina (Phygadeuon, Stibeutes) 71, 90
curvispina (Trematopygus, Lethades) 85, 93
curvulus (Mesochorus) 52, 97
curvulus (Orthocentrus, Stenomacrus) 66, 98

curvulus (Syndipnus, Trophoctonus, Synomelix)

82,94

cyaneoviridis (Platylabus, Cratichneumon) 77, 99
Cycasis 27, 88

cyclogaster (Hemiteles, Pleurogyrus) 36, 89
Cymodusa 26, 62, 95
cynipinus (Hemiteles, Zoophthorus) 36, 89

Daschia41,98

decrescens (Ichneumon, Coelichneumon) 44, 99

decrescens (Thersilochus, Phradis) 83, 97

deletus (Bassus, Diplazon) 19, 98

deletus (Cteniscus, Eridolius) 26, 88

deletus (Mesoleius, Campodorus) 56, 94

deletus (Orthocentrus, Stenomacrus) 66, 98

delictor (Barylypa) 17, 98

Deloglyptus 26

Delomerista 75, 87

Delomeristini 87

Delotomus 27

Demophorus 27

dentifer (Blapticus) 19, 98

dentifer (Caenocryptus, Xylophrurus) 20, 92

dentifer (Mesostenus, Stenaraeus) 60, 92

dentifera (Glypta) 32, 92

dentipes (Odontocolon) 63, 89

depressus (Mesoleius, Scopesus, Scopesis) 56, 94

Diaborus 27

Diadegma 13, 14, 16,52,96

Diadromus 27, 28, 100

Diaglyptellana 39, 89

Dialipsis 77, 98

dianae (Itamoplex) 25, 92

Diaparsis 83, 97

Diaparsus 83, 84

Dicaelotus 24, 26, 28, 99

Dichrogaster 37, 38, 72, 89

Dicoelotus 28

didymus (Mesoleius, Lagarotus, Lagarotis) 56, 93

digestor (Lissonota) 50, 92

digrammus (Barichneumon) 46, 99

dilatatus (Paniscus, Netelia) 68, 87

dimidiatus (Adelognathus) 11, 88

dimidiatus (Phygadeuon) 71, 90

Dimophora 27, 96

Dioctes 14

Diphyus 12,99

Diplazon 19, 98

Diplazontinae 98

Dirophanes 70, 100

dispar Ratzeburg (Hemiteles) 37

dispar Thomson (Hemiteles) 37, 92

dispar Gmelin (Ichneumon) 20

dispar Thunberg (Ichneumon, Xylophrurus) 20, 92

dispar (Spilocryptus, Agrothereutes) 80, 91

distans (Hemiteles, Clypeoteles) 37, 40, 89

distans (Microcryptus, Aptesis) 61,91

distans (Ophion) 66, 97

divisus (Colpognathus) 24, 100

INDEX

109

Dolichomitus 28, 29, 87

Dolophron 85, 95

dorsatus (Mesoleius, Saotus, Saotis) 56, 94

Dreisbachia 76, 87

drewseni (Hygrocryptus, Thrybius) 43, 91

Dusona20,21,95

dux (Dolichomitus) 28, 87

Ecclinops 68

Echthronomas 16,96

elegans (Hoplocryptus, Aritranis) 42, 91

elongata (Angitia, Diadegma) 14, 96

elongatus (Phaeogenes, Proscus) 69, 100

elymi (Hemiteles, Gelis) 37, 89

emarginatus (Mesoleius, Saotus, Saotis) 56, 94

emarginatus (Perilissus, Ecclinops) 68, 93

Enclisis 20, 92

Endaseina 90

Endasys72,81,82,90

Enizemum 42, 98

ensifer (Pectinolochus) 83, 96

Enytus 14,95

Eparces 23, 100

Ephialtes 28, 29

Ephialtini 87

Epitomus29, 100

Eridolius 26, 88

Erigorgusl7,98

errabunda (Lissonota) 50, 93

Erromenus 29, 88

eryptobius (Atractodes) 18, 91

erythrocerus (Brachycryptus, Hidryta) 19, 91

erythrogaster (Tryphon) 86, 88

erythropus (Sagaritis, Campoletis) 80, 95

Ethelurgina 89

Eudelus 39, 40, 89

Eupalamus 47, 99

eurycerus (Ichneumon) 44, 99

eurycerus (Stylocryptus, Endasys) 82, 90

Eurylabini 99

Eurylabus 29, 99

Euryproctini 94

Euryproctus 29, 30, 94

eurystigma (Plectiscus, Plectiscidea) 77, 98

Eusterinx 23, 98

evanialis (Dimophora) 27, 96

exareolatus (Euryproctus) 30, 94

Exenterini 88

Exenterus 30, 31,88

Exeristes 75, 76, 87

Exetastes 30, 93

exilis (Dialipsis) 77, 98

Exochilum 17

Exochus 30, 31,97

exoleta (Omorga, Tranosema) 65, 95

Exolytus 18

exserens (Orthocentrus, Stenomacrus) 67, 98

extincta (Glypta) 33, 92

Exyston31,88

facialis (Adelognathus) 1 1, 88

facialis (Anilasta, Hyposoter) 16, 96

facialis (Chorinaeus, Trieces) 23, 97

facialis Gravenhorst (Phygadeuon) 72

facialis Thomson (Phygadeuon, Theroscopus) 72,
90

facialis (Spudaeus, Synodites) 81, 94

facialis (Symplecis) 82, 98

facialis (Triclistus) 86, 97

faciator (Theroscopus) 72, 90

falcatus (Hemiteles, Tropistes) 37, 90

facatus (Orthocentrus, Stenomacrus, Neurateles)
67,98

fasciatulus (Theroscopus) 37, 90

fasciatus (Adelognathus) 1 1, 88

fasciatus Heer (Hemiteles) 37

fasciatus Thomson (Hemiteles, Theroscopus) 37,
90

femoralis (Banchus) 19, 93

femoralis (Cratocryptus, Cubocephalus) 24, 91

femoralis (Hadrodactylus) 60, 94

femoralis (Microcryptus, Aptesis) 61, 91

femoralis (Platlyabus, Tricholabus) 77, 99

femoralis (Rhaestus) 79, 93

femorator (Glypta) 32, 92

femorator (Mesoleius, Anoncus) 56, 94

femorator (Phobetus, Phobetes) 70, 94

femorella (Olesicampa, Olesicampe) 63, 96

fenestralis (Thersilochus, Diaparsus, Gonolochus)

83,96
fibulator (Anomalon, Gravenhorstia, Erigorgus)

17,98

filicornis (Atractodes, Exolytus, Mesoleptus) 18, 91
filicornis (Glypta) 32, 92
filicornis (Porizon, Leptopygus, Barycnemis) 79,

96

filicornis (Thersilochus, Tersilochus) 83, 97
filipalpis (Diaborus, Cteniscus) 27, 88
flavellus (Exenterus) 30, 88
flavicans (Phygadeuon) 72, 90
flavicincta (Phobocampa, Phobocampe) 70, 95
flavicornis (Mesoleius, Barytarbus, Mesoleptidea)

56,94

flavicornis (Olesicampa, Olesicampe) 64, 96
flavicornis (Orthocentrus, Stenomacrus,

Leipaulus) 67, 98

flavicornis (Thersilochus, Diaparsis) 83, 97
flavicoxa (Atractodes) 18, 91
flavicoxis (Pimpla) 75, 87

flavipes (Atractodes, Exolytus, Mesoleptus) 18, 91
flavipes (Bassus) 100
flavipes Provancher (Mesostenus, Phygadeuon,

Grypocentrus) 72
flavipes Thomson (Phygadeuon, Medophron) 72,

90

flaviscapus (Campoplex, Dusona) 21, 95
flavitarsis (Medophron) 72, 90
flavitarsis (Mesoleius, Perispudus, Perispuda) 56,
93

110

INDEX

flavoscutellatus (Mesoleius, Barytarbus,

Barytarbes) 56, 93
floricolator (Xiphulcus) 38, 89
folii (Lissonota) 50, 93

forticanda (Lathrostiza, Lathrostizus) 48, 96
forticauda (Lathrostiza, Lathrostizus) 48, 96
forticosta (Omorga, Campoplex) 65, 95
fortipes (Orthocentrus, Stenomacrus) 67, 98
fractigena (Glypta) 32, 92
fragilis (Bathythrix) 48, 90
frater(Hidryta)19,91
frenalis (Mesoleius) 56, 94
frigidus (Lathiponus) 59, 93
frontalis (Diaborus, Cteniscus) 27, 88
frontator (Perilissus, Ecclinops) 68, 93
frontatus (Mesoleius) 56, 94
frutetorum (Holocremna, Olesicampe) 41, 96
fugitivus (Aritranis) 42, 91
fulcrans (Olesicampa, Olesicampe) 64, 96
fulviventris (Phobetus, Ipoctonus, Phobetes) 70, 94
fulvus (Mesochorus) 53, 97
fumipennis (Agrothereutes) 81,91
fumipennis (Hemiteles, Mastrus) 37, 89
fuscicarpus (Hemiteles) 37, 92
fuscicarpus (Limneria, Sinophorus) 49, 95
fuscicornis (Netelia) 68, 87
fuscipes (Hemichneumon) 35, 99
fusciplica (Omorga, Campoplex) 65, 95
fusicornis (Omorga, Campoplex) 65, 95
fusicornis (Polyblastus, Scopiorus, Ctenochira) 78,

88
fusiformis (Catoglyptus, Asthenarus, Syntactus)

23,93
fusiventris (Brachycryptus, Hidryta) 19, 91

gallicola (Isadelphus) 39, 89

gallicus (Mesoleius, Campodorus) 57, 94

Gambrus32,43,80,91

Gelina 89

Gelis 35, 36, 37, 38, 39, 69, 89

genalis (Anomalon, Barylypa) 17, 98

genalis (Campoplex, Dusona) 21, 95

genalis (Cteniscus, Eridolius) 26, 88

genalis (Exyston) 31, 88

genalis (Hadrodactylus) 35, 94

genalis (Lissonota, Cryptopimpla) 50, 93

genalis (Polyblastus, Ctenacmus, Ctenochira) 78,

88

genalis (Thersilochus, Diaparsus, Diaparsis) 83, 97
geniculatus (Hemiteles, Dichrogaster) 37, 89
geniculatus (Phytodietus) 75, 87
geniculella (Olesicampa, Olesicampe) 64, 96
geniculosus (Leptocryptus, Bathythrix) 48, 90
gibbifrons (Hemiteles, Gelis) 37, 89
gibbulus (Ichneumon) 44, 99
giganteum (Therion) 17, 98
Giraudia77, 91

glabriculus (Goniocryptus, Trychosis) 34, 91
glabriventris (Miomeris, Microleptes) 62, 98

glacialis(Gelis)35,89

Glyphicnemis 82, 90

Glypta 32, 33, 92, 100

Glyptini 92

glyptonotus (Hemiteles, Chirotica) 37, 89

Glyptorhaestus 79, 80, 93

glyptus (Mesoleius, Campodorus) 57, 94

glyptus (Xylonomus, Xorides) 87, 89

gnathaulax (Ephialtes, Paraperithous) 28, 87

Gnathocryptus 82

Gnypetomorpha 35, 89

Gnypetomorphina 89

gonatopinus (Pezomachus, Gelis) 69, 89

Goniocryptus 33, 34

Gonolochus 83, 96

Gonotypa 34

Gonotypus 34, 95

gracilipes (Hadrodactylus) 35, 94

gracilipes (Hemiteles, Oecotelma) 37, 90

gracilipes (Lissonota) 50, 93

gracilipes (Olesicampa, Olesicampe) 64, 96

gracilipes (Paniscus, Netelia) 68, 87

gracilis (Amphibulus) 24, 90

gracilis (Hemiteles, Aclastus) 37, 89

gracillimus (Porizon, Barycnemis) 79, 96

graculus (Zoophthorus) 36, 89

gradarius (Trychosis) 34, 91

graefei (Hoplocryptus, Aritranis) 42, 91

grandiceps (Centeterus, Eparces) 23, 100

grandiceps (Ichneumon, Cratichneumon) 44, 99

grandiceps (Perilissus, Polyoncus, Lathrolestes) 69,

93

grandiceps (Pezomachus, Gelis) 69, 89
grandiceps (Phygadeuon) 72, 90
grandicornis (Ichneumon) 44, 99
grandis (Mesoleius, Protarchus) 57, 93
grandis (Phygadeuon, Pygocryptus) 72, 89
graniger (Aethecerus) 12, 100
graniger (Mesochorus, Astiphrommus,

Astiphromma) 53, 97

gravenhorsti (Microcryptus, Polytribax) 61,91
Gravenhorstia 17, 98
Grypocentrus 34, 72, 88
guttifer (Cremastus, Temelucha) 25, 96
guttifer (Exetastes) 30, 93

Habrocryptus 34

hadrocera (Omorga, Campoplex) 65, 95
hadrocerus (Hemiteles, Orthizema) 37, 90
Hadrodactylus 34, 35, 60, 94
haemosternus (Ctenochira) 78, 88
hamulus (Mesochorus, Astiphrommus,

Astiphromma) 53, 97
Handaoia 36, 89
hastator (Banchus) 19, 93
Hedycryptina 92
Helictes51,98
Hemichneumon 35, 99
Hemiteles 35, 36, 37, 38, 39, 40, 89, 92

INDEX

111

Hemitelina 89

Heterischnus 47, 99

heterocera (Glypta) 32, 92

heterocerus (Phaestus) 70, 93

heterocerus (Plectiscus, Proclitus) 77, 98

heterocerus (Thersilochus, Tersilochus) 84, 97

Heterocola 84, 97

heterogaster (Holocremna, Olesicampe) 41, 96

heterogaster (Phygadeuon) 72, 90

heteropus (Coleocentrus) 24, 98

heteropus (Ephialtes, Dolichomitus) 28, 87

heteropus (Leptocryptus, Bathythrix) 48, 90

heteropus (Mesoleius, Saotus, Saotis) 57, 94

heteropus (Phygadeuon, Dichrogaster) 72, 89

heteropus (Synetaeris, Pyracmon) 83, 95

hians (Lissonota) 50, 92

Hidrytal9,91

Himerta 29, 94

Himertus 29

hirticeps (Hemiteles, Zoophthorus) 37, 89

Hodostates 40, 93

Hodostatus 40

holmgreni (Mesoleptus, Mesoleptidea) 60, 94

Holocremna 40, 41

holopyga (Angitia, Diadegma) 14, 96

Homaspis 63, 93

homocerus (Hemiteles, Sulcarius) 37, 90

Homoporus 41, 42

Hoplocryptus 42, 43

hostilis (Anisobas) 12

humerella (Lissonota) 50, 93

humerellus (Mesoleius, Campodorus) 57, 94

hygrobia (Meloboris, Diadegma) 52, 96

hygrobius (Homoporus, Syrphoctonus) 42, 98

Hygrocryptus 43

Hypamblys 82

Hyperbatus 59, 94

hyperborea (Limneria, Tranosema) 49, 95

hypolius (Ichneumon) 45, 99

hypomelas (Mesoleius, Otlophorus) 57, 93

Hyposoter 16, 17,96

Ichneumon 43, 44, 45, 46, 47, 72, 99

Ichneumoninae 99

Ichneumonini 99

Idiogramma51,88

Idiogrammatini 88

Idiolispa49, 91

immarginatus (Mesoleius) 57, 94

immolator (Biolysia, Bathyplectes) 62, 95

impressifrons (Lissonota) 50, 93

incidens (Exochus) 31, 97

incidens (Mesochorus, Astiphrommus,

Astiphromma) 53, 97
incidens (Mesoleius, Campodorus) 57, 94
incidens (Thersilochus, Aneuclis) 84, 97
incisus (Homoporus, Syrphoctonus) 42, 98
incisus (Mesoleius) 57, 94
infelix (Phygadeuon) 72, 90

inferus (Euryproctus) 30, 94

inferus (Gambrus) 32, 91

inflatus (Caenocryptus, Enclisis) 20, 92

inflatus (Hemiteles, Platyrhabdus) 37, 90

inflatus Provancher (Ichneumon, Phygadeuon,

Endasys) 72

inflatus Thomson (Phygadeuon) 72, 90
inflexus (Dicoelotus, Dicaelotus) 28, 99
infumatus (Cryptus, Itamoplex) 25, 92
infumatus (Hemiteles, Gelis) 38, 89
ingratus (Trychosis) 34, 91
inimicus (Isadelphus) 39, 89
innotatus (Orthocentrus, Stenomacrus) 67, 98
inquinatus (Ichneumon) 44, 99
insignis (Hadrodactylus) 60, 94
interruptus (Metopius, Peltocarus) 60, 97
intersectus (Atractodes) 18, 91
interstitialis (Thersilochus, Phradis) 84, 97
Ipoctonus 70
irrigua (Lissonota) 50, 92
Isadelphus 36, 39, 89
ischnocera (Meloboris, Diadegma) 52, 96
ischnocerus (Hemiteles, Tricholinum) 38, 90
ischnogaster (Casinaria) 22, 95
Ischnus 34, 47, 92
Itamoplex 25, 92
Itoplectis 75, 76, 87

Javra24,61,91
jesperi (Ichneumon) 45, 99
jucundus (Smicroplectrus) 80, 88
junior (Erromenus) 29, 88

kriechbaumeri (Spudastica) 81, 95
kriechbaumeri (Trematopygus) 85, 93
Kristotomus 27, 88

lacticrus (Angitia, Diadegma) 14, 96

laetus (Kristotomus) 27, 88

laeviceps (Porizon, Barycnemis) 79, 96

laevicollis (Adelognathus) 1 1, 88

laevifrons (Pimpla, Delomerista) 75, 87

laevifrous (Pimpla, Delomerista) 75, 87

laevipectus (Mesoleius, Campodorus) 57, 94

laeviusculus (Cremastus) 25, 96

laeviusculus (Mesoleius, Barytarbus, Barytarbes)

57,93

laeviventris (Phygadeuon) 72, 90
Lagarotis 56, 93
Lagarotus 56
Lamachus 55, 93

lamina (Leptocryptus, Bathythrix) 48, 90
lancifer (Xylophrurus) 20, 92
lapponica (Pimpla) 100
lapponicum (Anomalon, Gravenhorstia,

Erigorgus)17,98

lapponicus (Goniocryptus, Trychosis) 34, 92
lapponicus (Mesochorus) 53, 97
lapponicus (Microcryptus, Aptesis) 61,91

112

INDEX

lapponicus (Phygadeuon) 72, 90

laricinus (Exenterus) 30, 88

larvatus (Eurylabus) 29, 99

lateralis (Pyracmon) 79, 95

Lathiponus 59, 93

Lathrolestes 47, 69, 93

Lathrolestus 47

Lathroplex 48

Lathrostiza 48

Lathrostizus 14, 15,96

laticarpus (Promethus, Sussaba) 79, 99

laticeps (Anomalon, Barylypa) 17, 98

laticeps (Hadrodactylus) 35, 94

laticeps (Mesochorus, Stictopisthus) 53, 97

laticrus (Caenocryptus, Enclisis) 20, 92

latipes (Phobetus, Ipoctonus, Phobetes) 70, 94

latiscapus (Mesoleius, Campodorus) 57, 94

latiscapus (Platylabus, Asthenolabus) 77, 99

latitarsis (Cryptus, Meringopus) 25, 92

latiuscula (Tranosema, Dolophron) 85, 95

lativentris (Nemeritis) 62, 95

lativentris (Platylabus) 77, 99

lativentris (Triclistus) 86, 97

latungula (Angitia, Diadegma) 14, 96

latungula (Campoplex, Dusona) 21, 95

latungula (Parabatus, Netelia) 68, 87

legator (Trychosis) 34, 92

Leipaulus 67, 98

Leptocryptoides 48, 90

Leptocryptus 48

Leptopygus 79

Lethades 85, 93

lethierryi (Trematopygus) 85, 93

leucomera (Anilasta, Hyposoter) 16, 96

leucopeltis (Ichneumon) 45, 99

leucopygus (Thrybius) 43, 91

liambus (Hemiteles) 38, 92

limbata (Oedimopsis, Oedemopsis) 63, 88

limbatus (Adelognathus) 1 1, 88

Limneria 48, 49

Limneriini 95

limnobius (Campoplex, Dusona) 21, 95

limnophilus (Ambly teles, Spilichneumon) 12, 99

linearis (Ateleute) 38, 92

linearis (Bathythrix) 48, 90

lineatus (Cremastus) 25, 96

lineifrons (Exochus) 31, 97

lineiger (Cteniscus, Eridolius) 26, 88

lineiger (Syndipnus, Hypamblys, Synodites) 82, 94

liocnemis (Ichneumon, Coelichneumon) 45, 99

Liocryptus 49

liogaster (Atractodes) 18, 91

liogaster (Omorga, Campoplex) 65, 95

liogaster (Phygadeuon) 72, 90

liopleuris (Euryproctus, Phobetus, Phobetes) 30,

94

liopleuris (Mesoleius, Saotus, Saotis) 57, 94
liopleuris (Thersilochus, Terilochus) 84, 97
liosternus (Mesoleius, Campodorus) 58, 94

liosternus (Phygadeuon) 72, 90

liosternus (Saotus) 58

liostylus (Hemiteles, Dichrogaster) 38, 89

liostylus (Ichneumon, Cratichneumon) 45, 99

Liotryphon 28, 87

Lissonota 49, 50, 51,92

Lissonotini 92

lissonotoides (Hemiteles, Ateleute) 38, 92

Listrodromini 99

litorea (Omorga, Campoplex) 65, 95

lobatus (Mesoleius, Campodorus) 58, 94

Lochetica 73, 89

longeareolatus (Ichneumon) 45, 99

longicalcar (Chorinaeus) 23, 97

longicalcar (Cymodusa) 26, 95

longicalcar (Triclistus) 86, 97

longicauda (Hemiteles, Gelis) 38, 89

longicauda (Mesochorus) 53, 97

longicauda (Microcryptus, Cubocephalus) 61, 91

longicaudatus (Hemiteles, Dichrogaster) 38, 89

longiceps (Phygadeuon, Ceratophygadeuon) 73, 90

longiceps (Pimpla) 75, 87

longicornis (Chorinaeus) 23, 97

longicornis (Exochus) 31, 97

longicornis (Thersilochus, Tersilochus) 84, 97

longigena (Amblyteles, Diphyus) 12, 99

longigena (Mesoleius, Scopesus, Neostroblia) 58,

94

longigena (Monoblastus, Rhorus) 62, 93
longigena (Ophion) 66, 97
longigena (Phygadeuon) 73, 90
longitarsis (Plectiscus, Proclitus) 77, 98
longiventris (Homoporus, Syrphoctonus) 42, 98
longiventris (Mesoleius, Lamachus) 58, 93
longiventris (Mesoleius, Saotus Saotis) 58, 94
longula (Anilasta, Hyposoter) 16, 96
longulus (Hemiteles, Xiphulcus) 38, 89
lucidulus (Acrotomus) 27, 88
Luphyroscopus 69
luteipes (Campoplex, Dusona) 21, 95
luteipes (Ephialtes, Paraperithous) 28, 87
luteipes (Olesicampa, Olesicampe) 64, 96
luteolus (Lathrolestus, Lathrolestes) 47, 93
lyrata (Omorga, Campoplex) 65, 95

macrocentrus (Polyblastus) 78, 88
macrocerus (Euryproctus, Syndipnus) 30, 94
macrocerus (Ichneumon) 45, 99
Macrochasmus 51
MacrocryptusSl

macropus (Mesoleius, Scopesus, Scopesis) 58, 94
macrostigma (Cremastus, Temelucha) 25, 96
macrostoma (Angitia, Lathrostizus) 14, 96
macroura (Sagaritis, Campoletis) 80, 95
macrourus (Goniocryptus, Trychosis) 34, 91
macrurus (Ephialtes, Dolichomitus) 28, 87
macrurus (Hemiteles, Charitopes) 38, 90
macrurus (Mesochorus) 53, 97
maculata (Diadegma) 15, 96

INDEX

113

maculipennis (Chirotica) 37, 89
magnicornis (Hemiteles, Phygadeuon) 38, 90
majalis (Diadegma) 14, 96
mandibularis (Mesochorus, Astiphrommus,

Astiphromma) 53, 97
mandibularis (Pezomachus, Gelis) 69, 89
mandibularis (Spudaeus, Campodorus) 81, 94
marginatus (Atractodes, Exolytus, Mesoleptus) 18,

91

marginatus (Cteniscus, Eridolius) 26, 88
marginatus (Delotomus, Kristotomus) 27, 88
marginatus (Lathrolestus, Lathrolestes) 47, 93
marginatus (Mesochorus) 53, 97
marginella (Nepiesta, Biolysia, Bathyplectes) 62,

95

maritimus (Thersilochus, Aneuclis) 84, 97
Mastrina 89
Mastrulus 36, 89
Mastrus 36, 37, 39, 89
medialis (Diadromus) 28, 100
Medophron71,72,74,90
megaspis (Homoporus, Syrphoctonus) 42, 98
Megastylus51,98

melampus (Omorga, Campoplex) 65, 95
melanaspis (Promethus, Promethes) 79, 98
melania (Angitia, Diadegma) 14, 96
melanocarus (Mesoleius, Otlophorus) 58, 93
melanocerus (Trematopygus) 85, 93
melanogaster (Hemiteles, Gelis) 38, 89
melanogaster (Holocremna, Olesicampe) 41, 96
melanogaster (Thersilochus, Tersilochus) 84, 97
melanopygus (Theroscopus) 40, 90
melanostoma (Gonotypa, Gonotypus) 34, 95
melanotus (Erromenus) 29, 88
melanurus (Mesoleius, Protarchus) 58, 93
melanurus (Paniscus, Netelia) 68, 87
Meloboris 52
Meringopus 25, 92 .

mesocastanus (Spudaeus, Rhinotorus) 81, 94
mesocastanus (Trychosis) 33, 92
Mesochorinae 97
Mesochorus 52, 53, 54, 97
Mesocryptus 54, 55, 100
Mesoleiini 93

Mesoleius 55, 56, 57, 58, 59, 60, 94
Mesoleptidea 56, 60, 94
Mesoleptus 18,60,91
Mesostenidea 60, 91
Mesostenina 92
Mesostenini 91
Mesostenus 60, 72, 92

mesostilpnus (Ichneumon, Barichneumon) 45, 99
mesoxanthus (Hoplocryptus, Aritranis) 43, 91
mesoxanthus (Mesoleius, Perispudus) 58, 94
messor (Dolichomitus) 28, 87
Metopiinae 97
Metopius 60, 97
microcera (Glypta) 32, 92
Microcryptus 60, 61, 62, 92, 100

Microdiaparsis 84, 96

Microleptes 62, 98

micropnygus (Ichneumon) 45, 46, 99

microstomus (Hemiteles, Zoophthorus) 38, 89

micrura (Angitia, Diadegma) 14, 96

minutorius (Ichneumon) 44, 99

minutulus (Stylocryptus, Endasys) 82, 90

Miomeris 62

monilicornis (Angitia, Lathrostizus) 1 5, 96

Monoblastus 62

monodon (Hemiteles, Platyrhabdus) 38, 90

monodon (Phygadeuon) 73, 90

monospila (Angitia, Diadegma) 15, 96

monospilus (Ichneumon) 45, 99

montanus (Phaeogenes) 70, 100

monticola (Casinaria) 22, 95

monticola (Thersilochus, Heterocola) 84, 97

mordax (Notopygus, Xenoschesis) 63, 93

mucronella (Sagaritis, Campoletis) 80, 95

mutanda (Lissonota) 49, 93

muticus (Platylabus) 77, 99

myrmecinus (Pezomachus, Gelis) 69, 89

Nanodiaparsis 83, 97

nasutus (Spilocryptus, Agrothereutes) 80, 91

neglectus (Hyposoter) 17, 96

neglectus (Trychosis) 34, 92

nemati (Mesoleius, Campodorus) 58, 94

Nemeritis 62, 95

Nemioblastus 77

Neostroblia 58, 94

Nepiesta 62, 95

nereni (Ichneumon) 45, 99

Netelia 68, 87

Neurateles 66, 67, 98

nigerrimus (Meringopus) 25, 92

nigricans (Pimpla, Scambus) 76, 87

nigricarpus (Parabatus, Netelia) 68, 87

nigriceps (Acanthocryptus, Rhembobius) 10, 89

nigriceps (Adelognathus) 1 1, 88

nigriceps (Mesochorus) 53, 97

nigricollis (Acanthocryptus, Rhembobius) 11, 89

nigricollis (Perilissus, Luphyroscopus,

Lathrolestes) 69, 93
nigricornis (Adelognathus) 11, 88
nigricornis (Centeterus) 23, 100
nigricornis (Glypta) 32, 92
nigricornis (Hemiteles, Sulcarius) 38, 90
nigricornis (Homoporus, Enizemum) 42, 98
nigricornis (Microcryptus, Oresbius) 61,91
nigricornis (Nyxeophilus, Xylophrurus) 63, 92
nigricornis (Sinarachna) 78, 87
nigricoxa (Mesoleptus, Hadrodactylus) 60, 94
nigricoxa (Olesicampa, Olesicampe) 64, 96
nigridens (Lissonota) 50, 93
nigridens (Omorga, Tranosema) 66, 95
nigridens (Spudaeus, Campodorus) 81, 94
nigrifemur (Hadrodactylus) 35, 94
nigrifrons (Diaborus, Cteniscus) 27, 88

114

INDEX

nigrifrons (Exochus) 31, 97

nigrina (Glypta) 32, 92

nigripalpis (Exochus) 31, 97

nigripalpis (Polyblastus, Ctenacmus, Ctenochira)

78,88

nigriscaposa (Pimpla, Scambus) 76, 87
nigriscuta (Mesoleius, Saotus, Saotis) 58, 94
nigritella (Limneria, Sinophorus) 49, 95
nigritulus (Microcryptus, Aptesis) 61,91
nigriventris (Glypta) 32, 92
nigriventris (Hemiteles, Isadelphus) 39, 89
nigriventris (Mesocryptus, Oresbius) 54, 91
nigriventris (Microcryptus, Mesocryptus) 100
nigriventris (Promethus, Promethes) 79, 98
nigriventris (Saotus, Saotis) 80, 94
nigriventris (Stenocryptus, Cubocephalus) 81,91
nigroplica (Glypta) 33, 92
nigroplica (Olesicampa, Olesicampe) 64, 96
nitidulus (Euryproctus) 30, 94
nitidulus (Goniocryptus, Trychosis) 34, 91
nitifrons (Chorinaeus, Trieces) 24, 97
nitifrons (Triclistus) 86, 97
nordenstromi (Ichneumon) 45, 99
notaticrus (Hemiteles, Zoophthorus) 39, 89
Notopygus 63

nubifer (Caenocryptus, Enclisis) 20, 92
nudicoxa (Ichneumon, Barichneumon) 45, 99
numidicus (Pezomachus) 69, 92
nutritor (Diaparsis) 83, 97
Nyxeophilus 63

obliquus (Hemiteles) 39, 92

obliquus (Mesoleius) 58, 94

obliquus (Plectiscus, Aperileptus) 77, 98

obliquus (Thersilochus, Tersilochus) 84, 97

obnoxius (Mesostenidea) 60, 91

obscura (Gnypetomorpha) 35, 89

obscuripes (Hemiteles, Isadelphus) 39, 89

obscurus (Phytodietus) 75, 87

ocellaris (Paniscus, Netelia) 68, 87

ochrogaster (Phygadeuon, Theroscopus) 73, 90

ochrostomus (Mesocryptus, Aptesis) 55, 91

ocularis (Phygadeuon) 73, 90

Odontocolon 63, 89

Odontomerus 63

Oecotelma 37, 90

Oedemipsis 63, 88

Oedimopsis 63

oenescens (Baeosomus, Baeosemus) 19, 100

Olesicampa 63, 64

Olesicampe 41, 63, 64, 65, 96

Omorga 65, 66

opaculus (Amblyteles, Platyabus) 12, 99

opaculus (Hemiteles, Diaglyptellana) 39, 89

opaculus (Microcryptus, Schenkia) 61, 91

opaculus (Paniscus, Netelia) 68, 87

opacus (Campoplex, Dusona) 21, 95

opacus (Cratocryptus, Javra) 24, 91

Ophion66,97

Ophioninae 97

ophthalmica (Temelucha) 25, 96

oppositus (Phygadeuon) 73, 90

orbitale (Anomalon, Gravenhorstia, Erigorgus) 17,

98

orbitalis (Casinaria) 22, 95
orbitalis (Dicoelotus, Dicaelotus) 28, 99
orbitalis (Mesoleius, Hyperbatus) 59, 94
orbitalis (Microcryptus, Aptesis) 61,91
orbitalis (Syndipnus, Synodytes, Synodites) 82, 94
orbitatorious (Habrocryptus, Ischnus) 34, 92
Oresbius 54, 61, 62, 91
orgyiae (Mesochorus) 54, 97
oriolus (Exenterus) 30, 88
ornaticeps (Microcryptus) 61, 92
ornatulus (Hemiteles, Gelis) 39, 89
ornatulus (Spilocryptus, Gambrus) 80, 91
ornatus (Phytodietus) 75, 87
Orthizema 37, 40, 90
Orthocentrinae 98
Orthocentrus 66, 67, 98
Otlophorus 57, 58, 93
ovaliformis (Phygadeuon) 73, 90
ovalis Provancher (Phygadeuon) 73
ovalis Thomson (Phygadeuon) 73, 90
ovalis (Pimpla, Itoplectis) 76, 87
ovivora (Tromatobia) 76, 87
Oxytorinae 98
Oxytorus 67, 98

Pachymerus 67

pallicarpus (Hemiteles, Eudelus) 39, 89

pallicarpus (Phygadeuon) 73, 90

pallicarpus (Thersilochus, Heterocola) 84, 97

pallicoxa (Aethecerus) 12, 100

pallicoxa (Polyblastus) 78, 88

pallida(Barylypa)17,98

pallidicarpus (Phygadeuon) 73, 90

pallidus (Pristomerus) 79, 96

pallipes (Trichomastix) 100

pallipes (Triclistus) 86, 97

pallitarsis (Diaborus, Cteniscus) 27, 88

pallitarsis (Ichneumon, Cratichneumon) 46, 99

palpalis (Lissonota) 50, 93

palustris (Hygrocryptus, Gambrus) 43, 91

Paniscus 68

Parabatus 68

parallela (Pimpla, Tromatobia) 76, 87

parallelus (Atractodes) 18, 91

parallelus (Ephialtes, Dolichomitus) 29, 87

Paraperithous 28, 87

Parmortha24, 91

parvicalcar (Syndipnus, Smicrolius) 82, 94

parvicanda (Angitia, Diadegma) 15, 96

parvicauda (Angitia, Diadegma) 1 5, 96

parvicauda (Phygadeuon, Ceratophygadeuon) 73,

90

parviceps (Amblyteles, Anisobas) 12, 99
parviceps (Syndipnus, Synodytes, Synodites) 82, 94

INDEX

115

parviceps (Thersilochus, Diaparsus,

Microdiaparsis) 84, 96

parvipennis (Phygadeuon, Arotrephes) 73, 89
parviscopa (Ichneumon, Cratichneumon) 46, 99
parvispina (Exochus) 31, 97
parvulus (Delotomus, Cycasis) 27, 88
parvulus (Euryproctus) 30, 94
parvus (Epitomus) 29, 100
patellana (Olesicampa, Olesicampe) 64, 96
pauper (Trychosis) 34, 92
pauxillus (Atractodes) 18, 91
pectinata (Anilasta, Hyposoter) 16, 96
pectinipes Bridgman (Mesochorus) 53
pectinipes Thomson (Mesochorus) 53, 97
Pectinolochus 83, 85, 96
pectoralis (Microcryptus, Aptesis) 61,91
pectoralis (Plectocryptus, Aconias) 77, 91
pectoralis (Syndipnus) 82, 94
pedatorius (Cteniscus) 27, 88
Peltocarus 60

pentagonus (Colpognathus, Dicaelotus) 24, 100
percontatoria (Zatypota) 78, 87
Perilissini 93
Perilissus 68, 69, 93
Perispuda 56, 93
Perispudus 56, 58
petiolaris (Atractodes, Exolytus, Mesoleptus) 18,

91

petiolaris (Orthocentrus) 67, 98
petiolaris (Spudastica) 81, 95
Pezolochus 69
Pezomachus 69, 92
Phaeogenes 69, 70, 100
Phaeogenini 99
Phaestus 70, 93
Phobetes 30, 70, 94
Phobetus 30, 70
Phobocampa 70
Phobocampe 70, 95
Phradis 83, 84, 85, 97
Phthorima 42, 98

Phygadeuon 10, 35, 38, 70, 71, 72, 73, 74, 75, 90
Phygadeuontina 90
Phygadeuontinae 89
Phygadeuontini 89
Phytodietini 87
Phytodietus 75, 87
Picrostigeus 67, 98

picticollis (Anilasta, Hyposoter) 16, 96
picticollis (Polysphincta, Zatypota) 78, 87
picticoxa (Mesoleius) 59, 94
picticrus (Mesochorus) 54, 97
picticrus (Omorga, Campoplex, Sesioplex) 66, 95
pictifrons (Pimpla, Dreisbachia) 76, 87
pictipes (Acropimpla) 76, 87
pictus (Goniocryptus, Trychosis) 34, 92
pilicornis (Megastylus, Helictes) 51, 98
pilosulus (Pezomachus, Pezolochus, Gelis) 69, 89
pilosus (Adelognathus, Cnemischus) 1 1, 88

Pimpla 75, 76, 87, 100

pimplarius (Allomacrus) 12,98

pimplarius (Phygadeuon, Lochetica) 73, 89

Pimplinae 87

Pimplini 87

pineti (Mesoleius, Campodorus) 59, 94

pineticola (Limneria, Sinophorus) 49, 95

pinetorum (Odontomerus, Odontocolon) 63, 89

Pionini 93

plagiatus (Mesochorus, Astiphrommus,

Astriphromma) 54, 97
planifrons (Ephialtes, Dolichomitus) 29, 87
planiscapus (Limneria, Sinophorus) 49, 95
Platylabini 99
Platylabus 12, 13,76,77,99
platylabus (Anisobas) 12
Platyrhabdus 37, 38, 90
platystylus (Amblyteles, Anisobas) 12, 99
plebejus (Erromenus) 29, 88
Plectiscidea 77, 98
Plectiscus 77
Plectocryptus 77
Pleolophus60,91

pleuralis (Cratocryptus, Parmortha) 24, 91
pleuralis (Ephialtes, Liotryphon) 29, 87
pleuralis (Goniocryptus, Trychosis) 34, 92
pleuralis (Lathrolestus, Lathrolestes) 47, 93
pleuralis (Limneria, Sinophorus) 49, 95
pleuralis (Megastylus) 51, 98
pleuralis (Mesoleius, Campodorus) 59, 94
pleuralis Cresson (Tryphon) 86
pleuralis Thomson (Tryphon) 86, 88
Pleurogyrus 36, 89

plumbeus (Hemiteles, Zoophthorus) 39, 89
polita(Dusona)21,95
Polyblastus 77, 78, 88
Polyoncus 69
Polyrhembia 18
Polyrhysius 82
Polysphincta 78
Polysphinctini 87
Polytribax61,91

polyzona (Angitia, Diadegma) 15, 96
populneus (Dolichomitus) 28, 87
Porizon 78, 79
pratorum (Exyston) 31, 88
Pristomerus 79, 96

proboscidalis (Thersilochus, Heterocola) 84, 97
Proclitus 77, 98
Promethes 79, 98
Promethus 79
Proscus 69
Protarchus 57, 58, 93
protensa (Casinaria) 22, 95
Protichneumonini 99
Pseudocryptini91

pubiventris (Caenocryptus, Enclisis) 20, 92
pubiventris (Triclistus) 86, 97
pulchella (Phobocampa, Phobocampe) 70, 95

116

INDEX

pulchella (Sussaba) 79, 99
pulchellus (Ischnus, Heterischnus) 47, 99
pulcher (Hoplocryptus, Aritranis) 43, 91
pulcherrimus (Mesoleius, Lathiponus) 59, 93
pulchrator (Polyspincta, Zatypota) 78, 87
punctata (Pimpla, Exeristes) 76, 87
punctatus (Erromenus) 29, 88
punctatus (Rhaestus, Glyptorhaestus) 79, 93
puncticeps (Pachymerus, Collyria) 67, 98
puncticollis (Adelognathus) 1 1, 88
puncticollis (Microcryptus, Aptesis) 61, 91
punctifer (Microcryptus, Oresbius) 62, 91
punctifrons (Amblyteles, Platylabus) 1 3, 99
punctigena (Phygadeuon) 74, 90
punctiger (Habrocryptus, Ischnus) 34, 92
punctipes (Angitia, Lathrostizus) 15, 96
punctipes (Cteniscus, Eridolius) 26, 88
punctipleuris (Cteniscus, Eridolius) 26, 88
punctipleuris (Mesochorus) 54, 97
punctipleuris (Phygadeuon) 74, 90
punctiscuta (Syndipnus) 82, 94
punctitarsis (Olesicampa, Olesicampe) 64, 96
punctiventris (Adelognathus) 1 1, 88
punctiventris (Deloglyptus, Dicaelotus) 26, 100
punctiventris (Hemiteles, Zoophthorus) 39, 89
punctiventris (Homoporus, Sussaba) 42, 99
punctiventris Thomson, 1877 (Lissonota) 50, 51, 93
punctiventris Thomson, 1894 (Lissonota,

Syzeucta, Syzeuctus) 51, 93
punctiventris (Phygadeuon) 74, 90
punctiventris (Pimpla, Scambus) 76, 87
punctulatus (Adelognathus) 1 1, 88
punctulatus (Glyptorhaestus) 80, 93
punctulatus (Odontomerus, Odontocolon) 63, 89
pungens (Cremastus) 25, 96
pusilla (Eusterinx) 23, 98
pusillus (Atractodes) 18, 91
Pygocryptus 72, 89
Pyracmon 79, 83, 95
pyramidatus (Anomalon, Exochilum, Therion) 17,

98

quadriannellatus (Ichneumon) 46, 99
quadriannulatus Gravenhorst (Ichneumon) 46
quadriannulatus Thomson (Ichneumon) 46, 99
quadridentata (Pimpla, Apechthis) 76, 87
quadrinotata (Anilasta, Echthronomas) 16, 96
quadrinotata (Lissonota) 49, 93
quadrinotatus (Cteniscus, Eridolius) 26, 88
quadrispinosus (Phygadeuon) 10
quadrispinus (Phygadeuon) 10
quercinus (Odontomerus, Odontocolon) 63, 89
quinquenotatus (Ichneumon) 46, 99

radialis (Cremastus) 25, 96

radialis (Orthocentrus) 67, 98

radiella (Olesicampa, Olesicampe) 64, 96

rectangulus (Miomeris, Microleptes) 62, 98

recticauda (Orthocentrus, Pictrostigeus) 67, 98

rectus (Campoplex, Dusona) 21, 95

recurvus (Phygadeuon, Medophron) 74, 90

relator (Tryphon) 86, 88

retusa (Olesicampa, Olesicampe) 64, 96

Rhaestus 79, 80, 93

Rhembobius 10, 11,89

Rhinotorus 55, 81,94

Rhorus 62, 93

rimator (Angitia, Diadegma) 15, 96

rimator (Lissonota) 51, 93

rimulosus (Ichneumon, Stenichneumon) 46, 99

ripicola (Atractodes, Exolytus, Mesoleptus) 18, 91

ripicola (Phygadeuon) 74, 90

roborator (Exeristes) 75, 76, 87

robusta (Dimophora) 27, 96

robustus (Notopygus, Homaspis) 63, 93

rostralis (Tranosema) 85, 95

rostrata (Canidia, Bathyplectes) 22, 95

rotundipennis (Phygadeuon) 74, 90

rubidus (Mesoleius) 59, 94

rubiginosus (Cycasis) 27, 88

rubricollis (Hemiteles, Gelis) 39, 89

rubricollis (Microcryptus, Schenkia) 62, 91

rubricosus (Phytodietus) 75, 87

rubripes (Hemiteles, Isadelphus) 39, 89

rubrotinctus (Hemiteles, Chirotica) 39, 89

ruficoxa (Omorga, Campoplex) 66, 95

ruficoxa (Phaeogenes, Dirophanes) 70, 100

ruficoxis (Cratocryptus, Cubocephalus) 24, 9 1

ruficrus (Anilasta, Hyposoter) 16, 96

rufifemur (Limneria, Sinophorus) 49, 95

rufipes Foerster (Asyncrita, Atractodes) 19

rufipes Foester (Atractodes) 18

rufipes Provancher (Atractodes) 18

rufipes Thomson (Atractodes, Asyncrita) 18, 91

rufipes Brischke (Glypta) 33

rufipes Spinola (Glypta) 33

rufipes Thomson (Glypta) 33, 92

rufipes (Phobetus, Ipoctonus, Phobetes) 70, 94

rufocincta (Acrolyta) 36, 89

rufonotatus (Eridolius) 26, 88

rufulus (Hemiteles, Mastrus) 39, 89

rugifer (Anomalon, Agrypon) 17, 98

rugifer (Hemiteles, Gelis) 39, 89

rugifrons (Hemiteles, Clypeoteles) 40, 89

rugipectus (Phygadeuon) 74, 90

Rugodiaparsis 83, 97

rugolosus (Leptocryptus, Bathythrix) 48, 90

Sagaritis 80
salicis (Glypta) 33, 92
salicis (Mesochorus) 54, 97
sanguinipes (Spudaeus, Arbelus) 81, 94
Saotis 55, 56, 57, 58, 59, 80, 94
Saotus 55, 56, 57, 58, 59, 80
saturator (Lissonota) 49, 93
scabra (Casinaria) 22, 95
scabriculus (Adelognathus) 11, 88

INDEX

117

scabriculus (Catoglyptus, Asthenarus, Syntactus)

23,93

scabriculus (Hemiteles, Eudelus) 40, 89
scabriculus (Trematopygus, Lethades) 85, 93
Scambus 76, 87

scansor (Plectocryptus, Giraudia) 77, 91
scaposa (Omorga, Campoplex) 66, 95
scaposus (Phygadeuon) 74, 90
Schenkia61,62,91

schoenobius (Cremastus, Temelucha) 25, 96
Scopesis 56, 58, 59, 94
Scopesus 56, 58, 59
Scopiorus 78

scutellaris (Ephialtes, Dolichomitus) 29, 87
scutellaris (Glypta) 33, 92
scutellaris (Ophion) 66, 97
scutellaris Holmgren (Polyblastus) 78
scutellaris Thomson (Polyblastus, Ctenacmus,

Ctenochira) 78, 88

scutellatus (Polyblastus, Ctenochira) 78, 88
segmentator (Hyperbatus) 59, 94
septentrionalis (Microcryptus, Oresbius) 62, 91
sericea (Cacotropa, Sphecophaga) 19, 92
serratus (Cryptus, Meringopus) 25, 92
Sesioplex 66

signatus (Syrphoctonus) 42, 98
signifer (Cteniscus, Eridolius) 26, 88
signifrons (Exochus) 31, 97
similis (Glypta) 33, 92
similis (Hemiteles) 40, 89
simillimus (Eudelus) 39, 40, 89
simplex (Erromenus) 29, 88
simplex (Exenterus), 30, 88
simplex (Mesochorus, Astiphrommus,

Astiphromma) 54, 97
simplex (Olesicampa, Olesicampe) 64, 96
simplicidens (Amblyteles, Spilichneumon) 13, 99
simulosus (Ichneumon, Stenichneumon) 46, 99
Sinarachna 78, 87
Sinophorus 49, 95

sinuata (Holocremna, Olesicampe) 41, 96
sinuatus (Mesoleius) 59, 94
Smicrolius 82, 94
Smicroplectrus 80, 88
sodalis (Pimpla) 75, 87
solutus (Hemiteles, Aclastus) 40, 89
sordidulus (Brachycryptus, Hidryta) 19, 91
sordipes (Angitia, Diadegma) 1 5, 96
Spanotecnus 68

specularis (Angitia, Diadegma) 15, 96
Sphecophaga 19,92
Sphecophagina 92
Spilichneumon 12, 13, 99
Spilocryptus 80, 81
Spilothyrateles 13,99
spiniger (Perilissus) 69, 93
spinipes (Campoplex, Dusona) 21, 95
spinula (Pezomachus, Gelis) 69, 89
spiracularis Thomson (Ichneumon) 45, 46, 99

spiracularis Tischbein (Ichneumon) 45, 46

spiracularis (Triclistus) 86, 97

spireae (Holocremna, Olesicampe) 41, 96

splendens (Campoplex, Dusona) 21, 95

sponsorius (Exyston) 31, 88

Spudaeus 55,81

Spudastica81,95

spuria (Pimpla) 76, 87

stagnalis (Hemiteles, Agasthenes) 40, 89

stagnicola (Amblyteles, Spilichneumon) 13, 99

Stenaraeus 60

Stenichneumon 46, 99

stenocarus (Campoplex, Dusona) 21, 95

stenocarus (Ichneumon, Cratichneumon) 46, 99

stenocerus (Ichneumon) 46, 99

stenocerus (Spudaeus, Campodorus) 81, 94

Stenocryptus 81

Stenomacrus 66, 67, 98

stenostigma (Anomalon, Agrypon) 17, 98

stenostigma (Canidia, Bathyplectes) 22, 95

stenostigma (Mesoleius) 59, 94

stenostigma (Pimpla, Acropimpla) 76, 87

stenura (Nemeritis) 62, 95

sternella (Olesicampa, Olesicampe) 64, 96

sternocera (Lathrostiza, Lathrostizus) 48, 96

sternocerus (Cratocryptus, Cubocephalus) 24, 91

Stibeutes36,71,90

Stictopisthus 52, 97

stigmaticus Brischke (Mesochorus) 54

stigmaticus Thomson (Mesochorus) 54, 97

Stilpnina91

stilpninus (Phygadeuon) 74, 90

Stilpnus81,91

Stiphrosomus 23

stramineipes (Diaparsis) 83, 97

strigipleuris (Pimpla) 76, 87

strigosus (Leptocryptus, Bathythrix) 48, 90

striola (Thersilochus, Pectinolochus) 85, 97

striolata (Omorga, Tranosema) 66, 95

striolatus (Caenocryptus, Enclisis) 20, 92

Stylocryptus81,82

subbuccata (Angitia, Diadegma) 15, 96

subcallosa (Olesicampa, Olesicampe) 64, 65, 96

subcircularis (Mesostenus, Mesostenidea) 60, 91

subclavata (Nepiesta) 62, 95

subdepressus (Thersilochus, Tersilochus) 85, 97

subfumata (Lissonota, Cryptopimpla) 51, 93

subglabra (Casinaria) 23, 95

subimpressus (Spudaeus, Rhinotorus) 81, 94

submuticus (Phygadeuon) 74, 90

subnasutus (Cremastus, Temelucha) 25, 96

subovalis (Mesostenus, Mesostenidea) 60, 91

subquadratus (Cryptus, Itamoplex) 25, 92

subquadratus (Ichneumon) 47, 99

subroseus (Mesoleius) 59, 94

subscaber (Syndipnus, Synodytes, Synodites) 82,

94

subteres (Plectiscus, Plectiscidea) 77, 98
subtilis (Polyblastus) 78, 88

118

INDEX

suecicus (Mesochorus) 53, 97

Sulcarius 37, 38, 90

sulcatus (Catoglyptus, Stiphrosomus, Sympherta)

23,93

superus (Gambrus) 32, 91
superus (Orthocentrus, Stenomacrus) 67, 98
Sussaba 42, 79, 99
Sympherta 23, 93
Symplecis 82, 98
Syndipnus 29, 30, 82, 94
Synetaeris 83, 95
Synocoetes 82, 93
Synodites81,82,94
Synodytes 82
Synomelix 82, 94
Syntactus 23, 93
Syrphoctonus41,42,98
Syzeucta 5 1
Syzeuctus 51,93

tarsator (Hadrodactylus) 35, 94

tarsator (Holocremna, Olesicampe) 41, 96

tarsoleuca (Buathra) 25, 92

tegularis (Glypta) 33, 92

tegularis (Limneria, Sinophorus) 49, 95

tegularis (Mesoleius, Scopesus, Scopesis) 59, 94

tegularis (Phaeogenes) 70, 100

Temelucha 25, 96

temporalis (Mesochorus) 54, 97

temporalis (Thersilochus, Phradis) 85, 97

tener (Caenocryptus, Enclisis) 20, 92

tenerrima (Lissonota) 51, 93

tenuicornis (Glypta) 33, 92

tenuicornis (Liocryptus, Idiolispa) 49, 91

tenuicornis (Mesochorus, Astiphrommus,

Astiphromma) 54, 97
tenuicosta (Anilasta, Hyposoter) 16, 96
tenuicosta (Phygadeuon) 74, 90
tenuifasciatus (Syzeuctus) 51, 93
tenuipes (Angitia, Diadegma) 1 5, 96
tenuipes (Atractodes) 19, 91
tenuipes (Stilpnus) 81,91
tenuiscapus (Mesochorus) 54, 97
tenuiscapus (Phygadeuon) 74, 90
tenuitarsis (Glypta) 33, 92, 100
tenuitarsis (Ichneumon, Coelichneumon) 47, 99
tenuitarsis (Mesoleius, Campodorus) 59, 94
tenuiventris (Glypta) 33, 100
tenuiventris (Townesia) 28, 87
terebrans (Dolichomitus) 29, 87
Tersilochinae 96
Tersilochus 83, 84, 85, 97
tetracinctorius (Adelognathus) 1 1, 88
Therion 17,98
Therionini 98

Theroscopus 37, 40, 70, 72, 73, 75, 90
Thersilochus 83, 84, 85
thomsoni Dalla Torre (Asyncrita, Atractodes) 19,

91

thomsoni Jussila (Atractodes) 19

thomsoni (Hemiteles) 37, 92

thomsoni (Lamachus) 58, 93

thomsonii (Atractodes) 19

thomsonii (Glypta) 33, 92

Thrybius43,91

Thymaridini 88

Thymaris 85, 88

Thymarus 85

tibialis (Spilocryptus, Agrothereutes) 80, 91

tiphae (Hadrodactylus) 35, 94

titillator (Meringopus) 25, 92

titubator (Itamoplex) 25, 92

t-nigrum (Cteniscus, Eridolius) 26, 88

Townesia 28, 87

Trachyarus 85, 99

Tranosema 49, 65, 66, 85, 95

transversus (Platylabus) 77, 99

Trematopygus 85, 93

triangulatorius (Exenterus) 31

triannulatus (Hemiteles, Orthizema) 40, 90

Trichocryptus 85

Tricholabus 77, 99

Tricholinum 38, 90

Trichomastrix 100

trichophthalmus (Pachymerus, Collyria) 67, 98

trichops (Catomicrus, Eusterinx) 23, 98

trichops (Phygadeuon) 75, 90

tricincta Cresson (Pimpla) 76

trincincta Thomson (Pimpla, Itoplectis) 76, 87

tricineta (Pimpla, Itoplectis) 76, 87

Triclistus 86, 97

tricolor (Mesoleius, Saotus, Saotis) 59, 94

Trieces 23, 24, 97

triplicatus (Amblyteles, Spilichneumon, Diphyus)

13,99

trispilus (Ichneumon) 47, 99
tristator (Trychosis) 34, 92
tristis (Canidia, Biolysia) 22, 95
trochantella (Canidia, Biolysia) 22, 95
trochanteralis (Hemiteles, Theroscopus) 40, 90
trochanterata (Angitia, Diadegma) 16, 96
trochanteratus Thomson 1884 (Hemiteles,

Theroscopus) 40, 90
trochanteratus Thomson 1885 (Hemiteles,

Theroscopus) 40, 90
Tromatobia 76, 87
Trophoctonus 82
Tropistes 37, 90

truncata (Angitia, Diadegma) 15, 16, 96
truncatulus (Ichneumon, Coelichneumon) 47, 99
truncicola (Amblyteles, Spilothyrateles) 13, 99
truncicola (Pyracmon) 79, 95
Trychosis 33, 34,91,92
Tryphon 86, 88
Tryphoninae 87
Tryphonini 88

tuberculatus (Dolichomitus) 29, 87
tuberculiger (Mesochorus) 54, 97

INDEX

119

ungula (Orthocentrus, Stenomacrus) 67, 98
ungularis (Hemiteles, Theroscopus) 40, 90
ungularis (Lathrolestus, Lathrolestes) 47, 93
ungularis (Phygadeuon, Theroscopus) 75, 90
unicolor (Hemiteles) 40, 89

validicornis (Ctenochira) 78, 88
validicornis (Hemiteles, Theroscopus) 40, 90
varians (Sagaritis, Campoletis) 80, 95
varicolor (Notopygus, Homaspis) 63, 93
varicorne (Anomalon, Gravenhorstia, Erigorgus)

17,98

varicornis (Phygadeuon) 75, 90
varicoxa (Anilasta, Hyposoter) 17, 96
varicoxa (Bassus, Diplazon) 19, 98
varicoxa (Glypta) 33, 92
varicoxa (Lissonota) 51, 92
varicoxa (Mesoleius) 60, 94
varicoxa (Mesoleptus, Hadrodactylus) 60, 94
varicoxa (Saotus, Saotis) 80, 94
varitarsus (Agasthenes) 40, 89
varitarsus (Polyblastus) 78, 88
versutus (Microdiaparsis) 84, 96
verticina (Ctenopelma) 26, 93

vesparum (Sphecophaga) 19, 92
viduata (Itoplectis) 76, 87
villosulus (Hadrodactylus) 35, 94
vindex (Caenocryptus) 20
vinulator (Eurylabus) 29, 99
vividus (Hyposoter) 16, 96

wesmaeli (Ichneumon, Eupalamus) 47, 99
wuestneii (Rhaestus, Glyptorhaestus) 80, 93

xanthaspis (Homoporus, Phthorima) 42, 98
xanthognatha (Glypta) 33, 92
xanthognathus (Ichneumon) 47, 99
Xenoschesis 63, 93
Xiphulcus 36, 38, 89
Xorides 87, 89
Xoridinae 89
Xylonomus 87
Xylophrurus20,51,63,92

Zatypota 78, 87

Zoophthorus 36, 37, 38, 39, 89

zygaenarum (Spilocryptus, Agrothereutes) 81, 91

British Museum (Natural History)
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Titles to be published in Volume 45

A catalogue and reclassification of the Ichneumonidae
(Hymenoptera) described by C. G. Thomson.

By M. G. Fitton

A taxonomic review of the genus Phlebotomus (Diptera: Psychodidae).

By D. J. Lewis

Stenomine moths of the Neotropical genus Timocratica (Oecophoridae).

By V. O. Becker

Afrotropical species of the myrmicine ant genera Cardiocondyla, Leptothorax,
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By Barry Bolton

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24
V -«

Bulletin of the

British Museum (Natural History)

A taxonomic review of the genus Phlebotomus
(Diptera : Psychodidae)

t). J. Lewis

Entomology series

Vol 45 No 2 24 June 1982

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Entomology series

Vol 45 No 2 pp 121-209

Issued 24 June 1982

^fAS* v

j* GENERAL '

A taxonomic review of the genus Phlebotomus I + LIBR>
(Diptera : Psychodidae)

D. J. Lewis

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

Contents

Synopsis 121

Introduction 122

General 122

Fossil Phlebotominae 124

Distribution 125

Biology 126

Relation to disease 126

Explanation of terms 127

Various 127

Names of collectors mentioned 128

Depositories, actual, probable or original 129

Keys, citations, distribution and notes 129

Genus Phlebotomus Rondani & Berte 129

Key to the subgenera of Phlebotomus 130

Tibia 3 in certain species 131

Subgenus Spelaeophlebotomus Theodor 131

Subgenus Idiophlebotomus Quate & Fairchild 133

Subgenus Australophlebotomus Theodor 135

Subgenus Phlebotomus Rondani & Berte 137

Subgenus Paraphlebotomus Theodor 142

Subgenus Synphlebotomus Theodor 148

Subgenus Larroussius Nitzulescu 150

Subgenus Adlerius Nitzulescu 163

Subgenus Euphlebotomus Theodor 168

Subgenus Anaphlebotomus Theodor 170

Subgenus Kasaulius subgen. n 172

Nomen nudum 172

Discussion 173

Leg ratios 173

Evolution of Phlebotominae 175

Aspects of leishmanial evolution in relation to that of Phlebotominae . . .177

Acknowledgements 191

References 191

Index 207

Synopsis

The 11 subgenera (one new), 96 species (one new) and 17 subspecies of Phlebotomus are reviewed and keys
are provided for their identification. Accounts are given of fossil sandflies and of the role of Phlebotomus in
the transmission of disease. Taxonomic citations are provided for each species and subspecies, and ah
annotated distribution list referring to a map. For some species further notes are given, including references
to transmission of disease. It is suggested that 'leg ratio' is worth recording as a measure of leg length in a
readily comparable form, and that it provides additional information about certain genera, subgenera,
species and infraspecific forms. Evolutionary hypotheses are put forward to explain features of the present
distribution of Phlebotominae and leishmaniasis.

Bull. Br. Mus. not. Hist. (Ent.) 45 (2): 121-209 Issued 24 June 1982

122 D. J. LEWIS

Introduction
General

Phlebotomus Rondani & Berte is one of the two Old World genera of Phlebotominae and
includes all the habitual mammal-biters and the vectors of human leishmaniasis in the Old
World. Disease of this group have recently increased in several countries and epidemics have
followed interruption of malaria control, so that renewed concern about the diseases and new
research programmes demand up to date information about the vectors. During the past 80 years
intensive study has yielded many widely scattered publications about Phlebotomus, particularly
from leishmaniasis areas (Anonymous, 1977), and reviews of the genus in three zoogeographical
regions have been published. Many species occur in all three of them (Lewis, 19786: 311), and a
general survey is required. The present work deals with some basic aspects of Phlebotomus
throughout the Old World.

The classification of Phlebotomus and the Phlebotominae has been discussed by Abonnenc
(1972), Fairchild (1955), Lewis et al (1977), Theodor (1948; 1958) and others. I recognize the
division of the living Phlebotominae into five genera, Phlebotomus and Sergentomyia Franga &
Parrot in the Old World and Warileya Hertig (Fairchild, 1955: 183; Lewis et al., 1977: 325),
Brumptomyia Franga & Parrot and Lutzomyia Franc, a in the New World. Ready et al. (1980)
have stressed the undoubted importance of subgenus Psychodopygus Mangabeira of Lutzomyia
and treated it as a genus. Lewis et al. (1977: 324) gave reasons against such a course which would
involve the elevation to generic rank of several, much more distinctive, Old World subgenera of
sandflies and could lead to a general multiplication of genera. Such questions are among the
'pitfalls of perfection' (Nelson, 1978) and are 'handicaps of the human need to compress into
linear form the three dimensional world of nature' (Campbell, 1974: 15). Taking a world- wide
perspective, I regard Psychodopygus as an important subgenus without changing its rank.

Publications (most with keys) dealing with Phlebotomus in various areas include the following.

The Old World: Artemiev (1979: 19, Euphlebotomus; 1980, Adlerius), Lewis (1973), Theodor
(1948).

The Palaearctic Region: Artemiev (1978, key with figures for Afghanistan), Croset (1978: 713,
key with figures for Tunisia), Lewis & Biittiker, 1981, Saudi Arabia), Nadim & Javadian (1976,
Iran), Perfil'ev (1968, key with figures for the U.S.S.R.), Theodor (1958, key and figures for the
region).

The Afrotropical Region: Abonnenc (1972, key with figures), Quate (1964, Sudan).

The Oriental Region: Lewis (19786).

The Australian Region : papers by Lewis and Dyce are being completed.

The taxonomic characters are easily seen in flies mounted in gum-chloral medium which may
be ringed with Glyceel (Kevan, 1955: 417, 418; Southey, 1970: 51, 53, 56; Tribe, 1972). Potash
was hardly ever used for maceration because it weakens intersegmental membranes and makes
specimens difficult to remount. It was occasionally used for treating the tip of the abdomen to
clarify the spermathecae although it may distort the ducts.

The characters used are described by Abonnenc (1972), Artemiev (1978: 1-8), Forattini (1973),
Lewis (1973; 19786: 219), Perfil'ev (1968), Theodor (1958), Young (1979: 5-8) and many others.
Lewis's (1973) account is being amplified to include recently introduced characters, some of
which are mentioned below.

Head length may be measured from the tip of the clypeus to the most posterior parts of the
head, and eye length to include the fore and hind facets. The inter-ocular suture is of some use,
but mainly for American species. The inter-arcal area lies between the cibarial chitinous arch and
the cibarial teeth. The labrum is measured to include the anterior sensilla. The antennal papillae
(Parrot, 1953) were discussed by Wirth & Navai (1978 : fig. 5, 47). The dental depth is the distance
from the tip of the maxilla to the most proximal tooth.

The relative lengths of various leg segments have been used for classification in several groups
of insects, including Lepidoptera (Imms, 1964: 555, 556), aphids (Eastop, 1972: 173), Culicidae
(Reid, 1953: 75), Ceratopogonidae (Wirth et al, 1977: 621), Chironomidae (Pinder, 1978: 11, 19;
Saether, 1976), Mycetophilidae (Hutson & Kidd, 1975: 29; Hutson et al, 1980: 42), Cecidomy-
iidae (Panelius, 1965: 5, 132), and Phoridae (Borgmeier, 1964; Schmitz, 1957: 431, couplet 8;

THE GENUS PHLEBOTOMUS 123

1958). For the Phlebotominae, Franga (1919: 125) pointed out that leg-segment lengths of each of
the species then known varied within narrow limits, and since then many authors have recorded
the actual lengths of several or a few segments, mainly in species of Lutzomyia. French writers
have measured the hind leg of many species. Raynal (1934: 350) indicated the value of the hind
tibia-femur ratio for separating two species of Phlebotomus, and Zariquiey (1937: 417) used the
lengths of basitarsus 1 (longer or shorter than femur 1) and of all tibiae of certain species of
Phlebotomus. Theodor (1958: 4) remarked that the legs were particularly short in Palaearctic
Sergentomyia, Artemiev (1978: 4) referred to various measurements of the hind leg, and Young
(1979: 7) mentioned tibia length in Lutzomyia. L. W. Quate often recorded leg measurements
regardless of sex, implying that the sexes are similar in this respect, and other publications
indicate that differences are usually small.

In recent years some authors have recorded lengths of leg segments but not always the same
ones, some have ceased to make such records, and others have never done so. It is now time to
appraise the value of leg characters and of the time spent in measuring them. In the present work,
therefore, the lengths of the long segments of each leg, of females when possible, are recorded in a
way to allow quick comparison of species. The legs were measured at x 60, with occasional use
of x 120 to locate extensions into preceding segments, which were included. Legs detached from
the body could usually be recognized as first, second or third because tibia 2 is nearly always
longer than 1, and 3 than 2. All lengths are expressed in units of which 100 are the length of femur
1 of a particular species, and the relative lengths of the nine long segments of one side, usually of
one fly, are followed by the actual length in mm of femur 1, and of the wing in some cases. Leg
diagrams, first drawn on the scale of one unit to one centimetre (examples in Figs 1 5-24) are
useful for comparing species and picking out features of individual species for additional
measurements.

The aedeagus comprises two side pieces fused at the base (Perfil'ev, 1968: 32, 42) and protects
the tips of the sperm tubes. According to Theodor (1958: 5) these tubes are the true aedeagus, and
the 'aedeagus' strictly speaking is the aedeagus sheath. Some authors have recorded the length of
the aedeagus but without indicating the basal point from which it was measured. The most
convenient point is usually the dorsal hind end near the bases of the coxites, and if other points
are used in certain cases they can be indicated.

The last abdominal segment or proctiger of male sandflies is the ninth (Just, 1973: 314, 315,
316, 332) and shows some specific differences. Isaev (1935: 98) noted three types, in P. papatasi, in
P. sergenti and a species of Sergentomyia, and in P. chinensis, characterized by the length of the
surstyles, the nature of their junction to the segment, and the ventral shape of the latter. Appreci-
able differences are shown by the six species illustrated in Figs 8 to 14. Surstyle is a convenient
name for the lateral lobes of the ninth tergite.

Keys to the subgenera and their species are provided and should be used in conjunction with
descriptions. Taxonomic citations serve as a guide to literature on the genus, subgenera, species
and subspecies. Distribution lists of all species show the sources of information for the maps.
References to disease transmission by known or possible vector species show many which are or
may be important, and indicate publications on biology as well as disease.

For some species full lists of taxonomic citations would be unduly complex and long, and early
references are confined to a few of historic interest.

Where the original or later depository of a holotype, syntypes or other type-material is not
shown by a describer, later author or other source, it is deduced (with a query) either from the
original paper or another publication which is indicated. Some syntypes have been located with
the aid of Abonnenc (1972) although he refers to them as holotypes. Information about the
depositories of some types from Afghanistan is given by Artemiev (1978: 23). Types of species
described by Professor O. Theodor were kept in the Hadassah Hebrew University Medical
School, Jerusalem, until the collection was purchased from the University by the British Museum
(Natural History) in 1981.

Distribution data, on which the maps are based, are of three kinds, viz. information about
types, publications which give detailed information and often earlier references, and previously
unpublished records indicated by collectors' initials or 'BMNH'.

124 D. J. LEWIS

Some Chinese records were not available when this work was being prepared, and are being
assembled for publication by Professor Leng Y.-j. They include the description of 'P. major wui\
for which a preliminary note is included below under P. major, and records of P. longiductus from
Xinjiang (Wu etal., 1979).

Fossil Phlebotominae

It is appropriate to consider the fossils of Phlebotominae and their ancestors because they help to
explain the relation of Phlebotomus to other genera and to the evolution of leishmaniasis. Leish-
mania probably arose from a monoxenic flagellate parasite of the ancestors of sandflies, so there is
likely to be a phylogenetic relationship between the leishmaniae and their vectors (Saf yanova,
19776:281).

The hopping flight of sandflies doubtless caused many to be trapped in resin, and some
excellent fossil specimens have survived in several of the sources of insects in amber (Hennig,
1973: 6). Their approximate ages in MYA (millions of years ago) quoted below were supplied by
Dr P. E. S. Whalley or taken from the British Museum (Natural History) (1972) time scale, Riek
(1970) or the work of Smith & Briden (1977) which was also consulted for continental move-
ments. Wings of the following species are illustrated (Figs 20-33) to give an impression of the
groups mentioned here and later: Permotipula patricia Tillyard, 1929: 779 (Rohdendorf, 1974: 6),
Phlebotomites brevifilis Hennig, 1972: 40, 62, Phlebotomus tipuliformis Meunier (Fig. 27 after
Hennig), Warileya nigrosacculus Fairchild & Hertig, P. (Spelaeophlebotomus) minteri, P. (Idio-
phlebotomus) frondifer, Lutzomyia paterna (Quate, 1963), Brumptomyia galindoi (Fairchild &
Hertig), P. (Paraphlebotomus) sergenti, Lu. (Dampfomyia) permira (Fairchild & Hertig), Sergen-
tomyia (Neophlebotomus) gombaki (Lewis & Wharton), S. (Sergentomyia) bedfordi (Newstead), S.
(Sergentomyia)fallax (Parrot) and S. (Parvidens) lesleyae (Lewis & Kirk).

370 MY A, Devonian

The earliest known insect, a wingless form, was living about this time (Riek, 1970: 168).

230 MYA, Upper Permian

The mecopteran Permotipula exemplifies a primitive wing to which that of Nemopalpus Mac-
quart, though unrelated (Rohdendorf, 1974: 6), is remarkably similar. Nemopalpus is probably
among the most primitive living Diptera, close to the basic stock of the Psychodidae and to the
Phlebotominae in the matter of venation (Fairchild, 1955: 182; Lewis et al., 1977: 323).

The original Diptera, present at this period, were probably biting flies feeding on insects or
vertebrates and contemporaneous with the beginning of the reptile age, when the theromorph
ancestors of mammals existed before the origin of birds (Downes, 1971 : 241, 261, 262).

220 MY A, Lower Triassic

The infraorder Dictyodipteromorpha of the dipterous suborder Archidiptera was probably in
existence; it flourished in the Upper Triassic and was apparently the ancestral group which gave
rise to two branches, the infraorder Tipulimorpha Rohdendorf and all other later Diptera (Roh-
dendorf, 1974: 27, 55, 129, 136, 289, 329).

760 MYA, Middle Jurassic

The Tipulimorpha were established (Rohdendorf, 1974: 3, 291, 292) and included the tipulid
family Tanydophryneidae Rohdendorf which appears to have been ancestral to 'superfamily'
Psychodidea [fossil Psychodidae] (Rohdendorf, 1974: 3, 53, 219, 228, 291-293). This ancient
group, distinguished from all other Tipulimorpha by primitive larval features, has retained a
complex wing venation but its members have become smaller and thus been able to colonize
microhabitats (Rohdendorf, 1974: 53, 58, 292). Before the end of the Jurassic the ancient group of
the Phlebotominae, among the smallest of Diptera, must have come into existence (Hennig, 1972:
38, 55, 58), in which the origin of R2 + 3 has been displaced towards the wing tip so that the vein
seems to come from /?4, and R2 has been reduced (Hennig, 1969: 385). R2 + 3 is usually branched
only in the most primitive Diptera (Colless & McAlpine, 1970: 664). Hennig's important 1972
paper was probably based largely on previous work unpublished owing to the second world war
(Schlee, 1978:382).

THE GENUS PHLEBOTOMUS 125

720 MY A, Lower Cretaceous

The first known two species of Phlebotominae existed in what is now the Lebanon and was south
of the Tethys Sea (Hennig, 1972: 38; Melville, 1967: 293). The small, evidently primitive Phleb-
otomites longifilis Hennig, 1972: 40, 62, and Phlebotomites brevifilis Hennig, 1972: 40, 62
(Stuckenberg, 1975: 459), had wings with a broad distal half and broadly rounded tip which may
have accounted for a displacement of the origin of R2 + 3 beyond that of/?5 (Hennig, 1972: 8, 27,
39, 43, 51). Although these species show few very striking differences from some recent forms they
were included in a new genus because close relationship to Phlebotomus was not indicated.
Hennig (1972: 21, 28) considered that they might belong to the ancestral group of the Phleb-
otominae or to his probably monophyletic 'Phlebotominae s. str.' which comprises Phlebotomus,
Sergentomyia, Brumptomyia and Lutzomyia. Stuckenberg drew attention to the short palpal
segment 5 of Phlebotomites brevifilis which is like that of some American sandflies. The two
Cretaceous species and the present-day Neotropical Warileya have a similar type of wing struc-
ture and may be the sole remnants of an early movement from Africa to South America or vice
versa across a south Atlantic connection in the Lower Cretaceous or earlier (Hennig, 1972: 38, 39,
44).

30 MY A, probably Upper Eocene

One poorly described species is known from Baltic amber (Rohdendorf, 1974: 275), P. (Phleb-
otomiella) tipuliformis Meunier, 19056 [as P. tipuliformis]; 1906: 103 [as Phlebotomiella tipuli-
formis']; 1912: 71 [as P. (Phlebotomiella) tipuliformis'] (Fairchild, 1955: 183-187; Hennig, 1972:
51-55; Stuckenberg, 1975) and may have lived in the amber forest and fed on thin-skinned
reptiles (Larsson, 1978: 92, 93). Hennig regarded it as a member of his Phlebotominae s. str. and
perhaps of genus Phlebotomus and of subgenus Euphlebotomus or Anaphlebotomus, which showed
that splitting of the ancient Phlebotominae was already far advanced. Stuckenberg referred to the
short palpal segment 5 and primitive wing of P. tipuliformis and considered it to be congeneric
with Phlebotomites and somewhat intermediate between it and 'Phlebotominae s. str.\

26 M YA, Miocene

Lu. paterna (Quate, 1963: 114) (Hennig, 1972: 56, 59, 62, fig. 41) is the first known phlebotomine

with a narrow wing and is related to living reptile-feeding species.

One M YA to the present day

Philaematus pungens Loew, 1845: 8 (Parrot, 1951 : 28; Duckhouse & Lewis, 1980: 99) from copal
of unknown origin, ' Phlebotomus pungens' Meunier, 1905a: 209 (Duckhouse & Lewis, 1980: 99)
from Zanzibar copal, and S. succini (Stuckenberg, 1975: 456) (Lewis et al., 1977: 326; Duckhouse
& Lewis, 1980: 105) from copal, possibly East African, may be less than one MYA and represent
living species of Sergentomyia. Several specimens from African copal examined proved to belong
to this genus, and one, treated with xylol and mounted in Euparal, clearly shows pharyngeal
teeth, antennal ascoids and palpal sensilla.

Distribution

Quate (1962: 169, 170) regarded the Phlebotominae as tropical with northern intrusions.
Sandflies occupy most of the Old World other than cold regions and oceanic islands, and they are
absent from the Seychelles (Scott, 1933: 369), and Phlebotomus from Madagascar (Brygoo, 1974).
Sandflies are considered to need at least 50 days a year with a temperature not less than 20°C
(Perfil'ev, 1968: 98). Map 1, showing the general distribution of the subfamily in the Old World, is
based on data cited by Lewis (1974) and Leger & Rodhain (1978). In western Europe P. per-
niciosus and P. mascittii occur about as far north as 49°N, and in Asia P. chinensis is the most
northerly species (Perfil'ev, 1968: 89), reaching about 48°N (Beklemishev & Dolmatova, 1948:
354). In Canada sandflies are known from about 50°39'N near Kamloops, from 49°39'N at
Coulee Creek in Alberta, and at 44°41'N near Ottawa. The southern boundary of sandflies in the
Old and New Worlds is about 40°S (Perfil'ev, 1968: 90). The maps illustrate a mainly northern
distribution of Phlebotomus, which is discussed later. It is exemplified by the northern distri-
bution of Larroussius, and therefore of kala-azar, in Tunisia (Croset et al, 1978: 744), and by the

126 D J. LEWIS

presence of five Phlebotomus species out of six Phlebotominae in France, and two out of 26 in
Zaire (Vattier & Bimangou, 1974: 92; Vattier & Trouillet, 1975: 2; 1978: 701). Some 35 species,
including P. orientalis, have marked eastern or western limits.

Biology

Numerous publications dealing with this extensive subject may be located by reference to Abon-
nenc (1972), Lewis (1973; 1974a; 1977; 1978a; 19786), Perfil'ev (1968) and others, and notes on
various species in the present work. The following brief note refers to a few aspects.

Sandfly larvae are difficult to find and many live in soil or burrows of animals. Development
from egg to adult takes weeks or months according to temperature, and larvae undergo diapause
in some northern and other areas. Many adults of both sexes feed on sugar and the females take
vertebrate blood. Adults are active at night and rest in various shelters by day. Movement varies
from short hops to flights of a few hundred metres and occasionally nearly 2 km, and is usually
stopped by moderate wind. Palaearctic species tend to have one or two generations a year, and
some tropical ones flourish in either the dry or the wet season.

The genus Phlebotomus includes all the habitual mammal-biters and therefore all the sandfly
vectors of human disease in the Old World.

Relation to disease

The following summary of relation to disease in the Old World is supplemented by notes on some
species. The leishmaniases are the main group of sandfly-borne vertebrate infections. It seems
probable that Leishmania Ross, 1903; Wenyon, 1926: 396, having arisen as an insect parasite,
came to infect reptiles and eventually mammals (Lewis, 19780: 94; Telford, 1979: 322; Wilson &
Southgate, 1979: 243), so that sandflies may be regarded as the primary hosts (Lainson & Shaw,
1979: 2). This phylogenetic priority is not only of historical interest for it is reflected in present-
day associations which have a practical significance. Lizard leishmanias now occurs in the Old
World and possibly in the New World (Lainson & Shaw, 1979: 34). No Leishmania is known in
birds (Adler, 1964:42).

Many forms of Leishmania are transmitted among mammals by species of Phlebotomus. Basi-
cally, each causes a zoonosis into which man may enter to a varying extent, so that human
involvement ranges from sporadic cases to a purely man-sandfly infection. Probably in Asia
leishmaniae caused enzootics in canids and rodents which led to certain anthroponotic forms
which spread to some other Palaearctic areas (Garnham, 1971 : 482, 488; 1977: 18; Hoogstraal &
Heyneman, 1969: 1185; Lysenko, 1971: 515-518).

The forms of Leishmania are now being classified by means of objective biochemical, serologi-
cal and other studies of their intrinsic characters (Chance, 1979; Chance et al, 1977; Garnham,
1976: 536; Lumsden, 1977: 47; de Raadt, 1977: 314; Taqi & Evans, 1978: 56; Williams & Coelho,
1978; Zuckerman & Lainson, 1977: 89), and many forms will probably be recognized.

For a long time the leishmaniae were grouped, according to their normal (Lainson & Shaw,
1971 : 21) effect on the (secondary) human host, into visceral leishmaniasis (VL) or kala-azar and
cutaneous or dermal leishmaniasis (CL) which causes oriental sore and other diseases. This
grouping is unsatisfactory (Chance et al., 1977: 53, 56) but, despite rapidly changing concepts, is
still of some practical value. It is used here, where the taxonomic names of the parasites are taken
mainly from Lumsden (1977a: 46, 49; 19776).

VL is caused by forms of the Le. donovani (Laveran & Mesnil, 19030; 19036: 958) complex
which occur largely in wild Canidae and are transmitted mainly by species of the subgenera
Larroussius and Adlerius. The anthroponotic leishmaniasis of eastern India is due to Le. d.
donovani which has no dog or other animal reservoir and is transmitted by a species of Euphlebo-
tomus. Le. d. infantum Nicole, 1908, probably spread from Asia via Transcaucasia into the
Mediterranean area where it attacks dogs and children rather than adults. VL probably spread
eastwards via the Gobi Desert to eastern China (Beklemishev & Dolmatova, 1948: 351). The east
African VL is transmitted by a species of Synphlebotomus, and may infect animals as secondary
hosts (Lysenko, 1971: 518).

CL is due largely to members of the Le. tropica (Wright, 1903) group. The wild hosts, if any, are
usually rodents, and most of the sandfly vectors belong to the subgenera Phlebotomus and

THE GENUS PHLEBOTOMUS 127

Paraphlebotomus. Le. t. major Yakimov: 1915: 501; Zuckerman & Lainson, 1977: 67 occurs
largely in central Asia (Lysenko, 1971: 518; Lysenko & Belaev, 1977: 250, map) where it infects
Rhombomys opimus Lichtenstein, 1823, and some other rodents, and causes 'moist sore' in man.
Le. t. tropica (= minor Yakimov) was possibly derived (Hoogstraal & Heyneman, 1969: 1184)
from Le. t. major, occurs from the Mediterranean area to India (Lysenko, 1971 : 58), is largely
urban, causes 'dry sore' in man and infects dogs. Le. aethiopica Bray, Ashford & Bray is a hyrax
parasite which causes disseminated CL in Ethiopia and is transmitted by a species of Larroussius.

VL and CL usually occur in different areas (Lysenko, 1971 : 518, 519; Lysenko & Beliaev, 1977:
250; Theodor, 1964: 487), largely owing to the distribution of their vectors.

The recognition of a vector is a complex process involving many subjects which include sandfly
taxonomy, distribution, host choice and other aspects of ecology, determination of flagellates
found in wild flies, development of leishmaniae ingested by flies in the laboratory, and experimen-
tal transmission. Proof that a species is a vector can seldom be obtained, and it applies only to a
particular place and time; de Raadt (1977: 314) pointed out that detailed study of epidemiology
only gives an instantaneous reflection of a process continuing over a long period. The significance
of a vector may alter (Lysenko & Beliaev, 1977ft: 263; Sergiev, 1977: 283). There are many
gradations from occasional to habitual minor and major vectors. It is therefore impossible to
draw up a simple list of vectors, but a list of vectors and suspected vectors is of some value,
especially if followed by a summary of the evidence related to each species. In the present work
this is confined to references to the literature.

Killick-Kendrick (1978: 299, 300) listed 52 taxa, 28 of them Old World form, of Phlebotomus,
known or suspected of being vectors of leishmaniasis. The Old World taxa, listed in relation to
types of the disease in man are : visceral (Synphlebotomus) celiae, martini, vansomerenae, (Larrouss-
ius) ariasi, kandelakii kandelakii, longicuspis, major s. 1., orientalis, perniciosus perniciosus, tobbi,
(Adlerius) chinensis chinensis, ch. halepensis, longiductus, simici, (Euphlebotomus) argentipes; cu-
taneous (with four marked '+ VL?' which may transmit kala-azar locally): P. (Phlebotomus)
bergeroti, duboscqi, papatasi (+ VL?), salehi, (Paraphlebotomus) alexandri, caucasicus (+ VL?),
chabaudi, mongolensis ( + VL?), sergenti sergenti, (Synphlebotomus) ansarii, (Larroussius) longipes,
pedifer, perfiliewi ( + VL?). P. rossi is a recent suspect.

In addition to known vectors some sandflies presumably transmit VL among animals in large
areas of Africa where the human disease occurs but is rare (Gigade, 1978: 239), and in part of the
Sudan (Hoogstraal & Heyneman, 1969: 1141) and elsewhere where the infection is present with
no apparent vector.

Many apsects of vectors have been discussed by Adler (1964: 48, 80), Bray (1974: 91), Hoog-
straal & Heyneman (1969), Killick-Kendrick (1978; 1979), Lewis (1971; 1974; 19780), Minter
(1972), Molyneux (1977: 43-53), Safyanova (1967), Sergiev (1967: 26; 1979) and Williams &
Coelho(1978).

Sandfly fever virus, transmitted by P. papatasi and possibly other species (PerfiFev, 1968: 128),
occurs mainly in the Mediterranean area.

Most vector species are difficult, and some impossible, to control. Domestic species were
largely controlled by house-spraying against malaria vectors but have increased where this has
been stopped, in India, for instance, and in Greece where VL and CL increased when mosquito-
spraying ceased (Leger et al., 1979: 12). Sandflies have shown little resistance to insecticides
(Killick-Kendrick, 1978: 304) till recent instances in India.

Explanation of terms

Various

Antenna 3 etc. Antennal segment 3 etc.

Chahar Mahal Part of Bakhtiar va Chahar Mahal Province, Iran

Chinese, romanization The Pinyin system, adopted in the 1980 edition of The Times Atlas,

of spelling is used here

CL Cutaneous leishmaniasis

Gamma The distance between the origin of wing- veins R2 + 3 and R* and the

origin of R5

128 D. J. LEWIS

Gruziya Georgian S.S.S.R. (Georgia)

ICZN International Code of Zoological Nomenclature (1964)

and Amendments (1973)

Inverted commas Places in distribution lists not located

Kosovo i Metohija Present name for Kosmet, Yugoslavia

Le. Leishmania

Leg segments 100 units = length of femur 1

Lu. Lutzomyia
Map symbol underlined Locality approximate

MYA Millions of years ago

P. Phlebotomus

Palp 3 etc. Palp segment 3 etc.

R2 etc. Radius branch 2 and other wing veins

S. Sergentomyia
Sperm pump and tubes Genital pump and filaments

Transcaucasia Historic name for U.S.S.R. area south of Caucasus (now Armenia,

Azerbaydzhan and Gruziya)

VL Visceral leishmaniasis

WL Wing length in mm

Names of collectors mentioned

A. E. E. A. E. Eaton

C. A. V. B. C. A. V. Barkhuus

D. J. L. D. J. Lewis

D. M. A. D. M. Ackland

D. M. M. D. M. Minter

E. K. S. E. K. Saliba
E. M. Unknown
G. B. W. G. B. White
G. S. G. Shidrawi
H. C. B. H. C. Barnett
H. W. L. H. W. Leathern
J. A. S. J. A. Sinton

J. O. C. J. Omer-Cooper

J. P. M. J. P. McMahon

J. P. T. B. J. P. T. Boorman

J. P. D. J.-P. Dedet

J. W. J. Waterston

J. Wn. Jane Wilson

K. B. K. Behbehani

K. K. K. Kertesz

K. Z. D. K. Zein el Dine

L. E. S. L. E. Stephen

L. Y.-j. Leng Yan-jia

M. A. M. Ashraf

M. A. R. M. A. Rifa'at

N. L. C. N. L. Corkill

P. A. B. P. A. Buxton

P. Petrie

R. A. B. R. A. Bolt

R. E. D. B. R. E. Drake Brockman

R. L. C. R. L. Coe

R. P. L. R. P. Lane

R. W. A. R. W. Ashford

S. A. S. Adler

S. A. S. S. A. Smith

S. J. R. S. J. Rahman

S. T. S. Taussig

V. D. V. Dhanda

Y. S. Y. Schlein

THE GENUS PHLEBOTOMUS

129

Depositories, actual, probable
ANIC, Canberra

BMNH

BPBM, Honolulu
CA, Los Banos
CFHS, Nanking
CIH, Sydney

CRI, Kasauli
EM, Montpellier
FM, Paris
IH, Scoplje
IP, Algiers
IP, Paris
IPH, Tehran
L, Bastia

LSHTM, London
NM, Vienna
MB, Corales
MC, Kweiyang
MH, Sinferopol
MI, Moscow

MR AC, Tervuren

NM, Nairobi

PIPD, Shantung

SAIMR, Johannesburg

TI, Dushanbe

TI, Tbilisi

TM

U, Moscow

U, Pavia

U, Vienna

US, Tashkent

ZSI, Calcutta
ZI, Leningrad

or original

Australian National Insect Collection, Commonwealth Scientific and

Industrial Research Organisation, Canberra.
British Museum (Natural History)
Bernice P. Bishop Museum, Honolulu
College of Agriculture, Los Banos, Philippines
Central Field Health Station, Nanking
Commonwealth Institute of Health, Sydney, Australia (till 1980 School of

Public Health and Tropical Medicine)
Central Research Institute, Kasauli
Laboratoire d'Ecologie, Universite de Montpellier, France
Laboratoire de Parasitologie, Faculte de Medicine, Paris
Institute of Hygiene, Skoplje, Yugoslavia
Institut Pasteur, Algiers
Institut Pasteur, Paris

School of Public Health and Institute of Public Health, Tehran
Lycee, Bastia, Corsica

London School of Hygiene and Tropical Medicine, London
Naturhistorisches Museum, Vienna
Musee Bocage, Colares, Portugal
Medical College, Kweiyang, China
Military Hospital, Sinferopol, U.S.S.R.
Institute of Tropical Medicine and Parasitology, Moscow

[Location of some holotypes mentioned by Artemiev, 1978: 23.]
Musee Royal de 1'Afrique Centrale, Tervuren, Belgium
National Museum of Kenya, Nairobi
Provincial Institute of Parasitic Diseases, Shandong
South African Institute of Medical Research, Johannesburg, South Africa
Tropical Institute of Tadzhiskaya S.S.S.R., Dushanbe
Tropical Institute, Tbilisi
T. Maa's collection
University, Moscow
University of Pavia, Italy
University, Vienna
Protozoology Division, Uzbekistan Sanitary and Biological Institute,

Tashkent

Zoological Survey of India, Calcutta
Zoological Institute, Academy of Sciences of the U.S.S.R., Leningrad

Keys, citations, distribution and notes

Genus PHLEBOTOMUS Rondani & Berte

Flebotomus Rondani & Berte in Rondani, 1840: 12. Type-species : Bibio papatasi Scopoli, by monotypy.

Phlebotomus Rondani & Berte; Loew, 1845: 9 [emendation; first use of this name and Phlebotomidae
mentioned by Lewis in Lewis et al., 1977: 321 incorrect; spelling fixed under suspension of rules by ICZN,
1954, Opinion 256: 199]; Summers, 1911;Theodor, 1948:96; 1958:316; 1965: 179; Fairchild, 1955: 188;
Quate, 1964: 237, 238; Lewis, 1967: 14; 1973: 162; 1978ft: 233; Perfil'ev, 1968: 218; Abonnenc, 1972: 75,
92; Lewis, Young, Fairchild & Minter, 1977: 321, 326; Abonnenc & Leger, 1977: 71, 76; Duckhouse &
Lewis, 1980:99.

Cibarium of female usually without a row of teeth but often having a group of spicules, pigment patch
usually absent. Antenna 3 usually long, three or more segments of male with two ascoids. Mesanepisternum
usually with a few antero-ventrad hairs (Abonnenc & Leger, 1977: 71, 72). Abdominal tergites 2-6 with
many erect hairs. Spermathecae usually segmented. Style with three to five spines, only one or two terminal.
Paramere often complex. Species often large and pale.

There are a few omissions from the keys because only the female is described for P. sejunctus,
teshi, tubifer (male found), pexopharynx, betisi and somaliensis, and only the male for P. buccina-

130 D. J. LEWIS

tor, papuensis, trifilis, katangensis,fantalensis, chadlii, langeroni, mariae, perfiliewi galilaeus, coma-
tus (female found) and caudatus, and because species A, B, C and D are not yet described, the
females of P. brevis brevis, P. chinensis halepensis and P. ch. kyreniae and the male of the latter are
not sufficiently described, and the descriptions of 'P. major wuf and P. (Eu.) autumnalis Artemiev
were not available in time. Suitable descriptions of the missing forms could lead to improved
keys.

Key to the subgenera of Phlebotomus

1 Distance between bases of R4 and Rs relatively short, not more than a quarter of width of wing.

A pair of rods present next to genital pump. Palpal sensilla not spatulate.
Antenna 3 very long and much longer than palp. Palp short, with segment 5 shorter than or

equal to 3. Style very long. Spermathecal ducts usually short and wide 2

Distance between bases of R4 and R5 relatively long, at least a third of width of wing. Genital
pump without adjacent rods. Palpal sensilla spatulate 3

2 Vein M1 + 2 forking at level of radio-median cross-vein, before base of/?4. Cibarium of female

unarmed. Antenna 3 = 2-3 to 2-5 times length of labrum. Palp segment 3 not enlarged at base,
with sensilla scattered on flat surface. Style with four spines and a long hair. Afrotropical

Region Subgenus SPELAEOPHLEBOTOMUS (p. 131)

Vein M1 + 2 forking beyond level of radio-median cross- vein, beyond base of R4. Cibarium of
female with teeth covering a large area. Antenna 3 about three or more times length of
labrum. Palp segment 3 enlarged at base, with sunken patch of sensilla. Style with three to five
thick spines and sometimes several thick hairs. Oriental, Palaearctic and Australian Regions

Subgenus IDIOPHLEBOTOMUS (p. 133)

3 Style with three spines.

Female with row of about five to ten cibarial teeth, few or no hypopharyngeal teeth, and
thin- walled spermathecae. Male with genital filaments short or very short, paramere simple
and beak-like, and coxite with simple hair pattern

Subgenus AUSTRALOPHLEBOTOMUS(p. 135)
Style with four or more spines 4

4 Coxite with hairy process near base. Genital filaments short, 1-3 to 2-3 times length of pump . 5
Coxite without such process. Genital filaments 3 to 1 1 times length of pump .... 7

5 Coxite 0-37 to 0-74 mm long, its process very small. Style long and cylindrical with three distal

spatulate spines and two other spines. Paramere with two upward processes. Surstyle with
distal spines. Pharyngeal armature of female comprising either a network of lines or scales.
Spermatheca with nearly equal segments and a refractive membrane near the distal one

Subgenus PHLEBOTOMUS (p. 137)

Coxite 0-20 to 0-33 mm long, its process usually large, and having a brush of long hairs. Style
not long, with four or five spines. Paramere simple, distal upper surface flat and elliptical with
short hairs. Surstyle without distal spines. Pharynx of female with teeth or scales. Sperma-
theca sometimes with differentiated rounded end-segment 6

6 Style with four long spines, two near the tip and two near the base. Pharynx of female with large

backwardly directed teeth Subgenus PARAPHLEBOTOMUS (p. 142)

Style with five long spines, two at the tip and three near the middle. Pharynx of female with
irregular scales or punctiform teeth . . . Subgenus SYNPHLEBOTOMUS (p. 148)

7 Style with four long spines, one distal, one subterminal, and two near middle.

Paramere with one or two extra lobes, with or without accessory spine. Aedeagus some-
times conical. Pharynx of female with a small group of teeth in middle, and behind it some
concentric lines. Spermatheca segmented, end-segment not enlarged

Subgenus ANAPHLEBOTOMUS (p. 170)
Style with five long spines 8

8 Paramere with one or two extra lobes, with or without accessory spine. Pharynx of female as in

Anaphlebotomus.

Spermatheca with differentiated end-segment . . Subgenus EUPHLEBOTOMUS (p. 168)
Paramere without extra lobes. Pharyngeal armature otherwise 9

9 Paramere truncated.

Antenna 3 and legs long, and wings narrow. Spermatheca moniliform. Haltere of male with
broad stalk. Paramere with adjacent rod .... Subgenus KASAULWS (p. 172)

- Paramere not truncated 10

THE GENUS PHLEBOTOMUS 131

10 Pharynx of female and male with punctiform teeth (large in wenyoni), except in mascittii which
has large irregular teeth. Spermatheca segmented, with long finger-like neck except in soma-
liensis, which has a rather long end-segment, and mascittii, which has a spermatheca with
transverse striations often in distal part, a small head, little or no narrowing, and a wide duct.
Genital filaments three to five times as long as pump . . Subgenus LARROUSSWS (p. 150)

- Pharynx of female with triangular or rounded group of medium-size teeth. Spermatheca incom-
pletely segmented. Genital filaments usually very long, 6-6 to 1 1-0 length of pump

Subgenus ADLERIUS(p. 163)

Tibia 3 in certain species

The following records of relative lengths of tibia 3 (femur 1 = 100 units, females unless males
indicated) are placed here for species about which no other taxonomic information is given.

P. aculeatus (Kenya) <$, 182 (2-26 mm); betisi, 199; gibiensistf, 212 (2-30 mm); guggisbergi, 188; kandelakii
kandelakii, 164; longicuspis (Algeria) <J, 181; major major (Nepal), 193; orientalis (Yemen), 173; pedifer
(Kenya), 1 77 ; perfiliewi perfiliewi (Italy), 177;smirrnwi(U.S.S.R..)<J, 1 47 ;tobbi (Greece), I79;angustus<3, 189;
hindustanicus (Pakistan) ^, 202; rupester <$, 163; simici (Yugoslavia)^, 164; kiangsuensistf, 185; colabaensis,
1 85 ;rodhaini (Sudan), 167; sp. D <J, 189(2-66 mm); stantoni, 176.

Subgenus SP EL A EO PHLEBOTOMUS Theodor

Phlebotomus subgenus Spelaeophlebotomus Theodor, 1948: (94), 100, 108; Quate & Fairchild, 1961: 208;

Lewis, 1973: 162; Lewis, Young, Fairchild & Minter, 1977: 325, 326. Type-species: Phlebotomus gigas

Parrot & Schwetz, 1937, by original designation.
Spelaeophlebotomus Theodor; Abonnenc & Minter, 1965: 30; Vattier-Bernard, 1970: 189; Forattini, 1971:

97 ; Abonnenc, 1 972 : 88 ; Abonnenc & Leger, 1 976 : 76.

Theodor provisionally treated this taxon as a subgenus and Abonnenc & Minter raised it to
generic rank on the strength of wing venation. After discussion with Professor Theodor (1972;
1974, in litt.). I consider that wing features of this group, and possibly the American Warileya, do
not justify generic treatment. For this reason, and in the interests of stability (Lewis et a/., 1977),
Spelaeophlebotomus is here treated as a subgenus.

Key to the species of subgenus Spelaeophlebotomus

1 Antenna 3 of female 2-3-2-5 length of labrum. Paramere of male with slight basal swelling gigas (p. 131)
Antenna 3 of female 3-9 length of labrum. Paramere of male with long setiferous lobe . minteri (p. 131)

Phlebotomus (Spelaeophlebotomus) gigas Parrot & Schwetz
(Map 1)

Phlebotomus gigas Parrot & Schwetz, 1937: 224 [?]; Parrot & Wanson, 1938: 153 [<£]; 1946: 143; Kirk &

Lewis, 1946b: 119; 1948:327; 195 1:438; Parrot, 1953: 114.Syntypes4$, ZAIRE (MRAC, Tervuren).
Phlebotomus (Spelaeophlebotomus) gigas Parrot & Schwetz; Theodor, 1948: 94, 108.
Spelaeophlebotomus gigas Parrot & Schwetz; Vattier-Bernard, 1970: 194; Abonnenc, 1972: 89.

DISTRIBUTION. Africa: Abonnenc (1972: 261, map); Vattier-Bernard (1970: 194, 221, early stages, map).

NOTE. P. gigas is a very large species which lives in caves and bites bats and other small animals
and will attack man (Abonnenc, 1972: 90).

Phlebotomus (Spelaeophlebotomus) minteri sp. n.

(Figs 1-9, Map 1)

?. Head 0-52 mm long, eye 0-45 of its length with more than 50 facets, interocular suture complete, clypeus
with few hairs. Labrum 0-31 mm long, 0-61 length of head, 0-09 length of wing, with narrow apical part,
subapical sensilla very close together, and few cibarial sensilla. Cibarium with no chitinous arch, teeth or
pigment patch. Pharynx with marked subterminal bulge and scarcely visible spicules. Hypopharynx with
about 50 teeth on each side. Antenna 3 = 1-22 mm long, 1-17 length of 4 + 5, 3-88 length of labrum, 0-35

132

D. J. LEWIS

length of wing, ascoids not clear but probably as in P. gigas, papillae not seen. Mandible pointed, with
minute teeth. Maxilla with no lateral teeth, 50 or more ventrals, some of them in two rows, and a dental
depth of 0-17 mm; palpal formula 10, 20, 42, 20, 41 ; eight mesad sensilla on segment 3. Scutum and pleuron
pale, prosternal lobes short, mesanepisternum without hairs. Leg formula 100, 184, 201; 86, 195, 181; 97,
211, 198 (3-50, 1-59). Wing length 3-50 mm, 3-18 width, R2/R2 + 3 4-33, /?t overlap//?2 0-66, R^ and R2
markedly curved, venation as in P. gigas. Haltere long. Abdominal tergal hairs erect and rather evenly
spread. Spermathecae invisible in specimen.

cJ. Eye 0-45 length of head. Interocular suture complete. Labrum 0-20 (0-19-0-21) mm long, 0-07 (0-07-
0-07) length of wing. Cibarium and pharynx virtually unarmed. Antenna 3 = 1-23 (1-17-1-29) mm long, 1-12

y

-o

Figs 1-9 Phlebotomus minteri. (1) $, head; (2) $, labro-cibarium; (3) $, tip of labrum; (4) 9, maxilla,
with fore, middle and hind ventral teeth enlarged; (5)$, pharynx; (6)$, wing; (7)<^, terminalia; (8, 9)
c?, abdominal segment 9.

THE GENUS PHLEBOTOMUS 133

(1-10-1 -15) length of 4 + 5, 6-18 (6-09-6-25) length of labrum, 0-43 (0-43-0-44) length of wing. Ascoids
unclear in balsam mounts. Wing length 2-85 (2-66-2-95) mm long, 3-45 times width,K2/2 + 3 5-42 (4-35-6-10),
R1 overlap//? 2 0-71 (0-66-0-86). Aedeagus curving downward, with blunt end; filament 2-4 length of pump,
with nearby pair of rods. Paramere with sub-basal hairy lobe, widening near tip. Coxite with two long
ventral hairs near tip; style narrow with four spines and one seta. Segment 9 as figured.

MATERIAL EXAMINED

Holotype <J, Tanzania : Amboni Cave, 5°04'S 39°02'E, 26.X.1957 (D.M.M.) (BMNH).
Paratypes. 1 $, 4 <$, same data (all in BMNH except one $ in LSHTM, London).

COMMENTS. This species is named after Dr D. M. Minter in appreciation of his important work on
sandflies and leishmaniasis. P. minteri differs from P. gigas chiefly in having a relatively longer
labrum in both sexes and a pronounced parameral lobe in the male. The eyes are not very small.
The maxillary dentition is remarkable, presumably suited to bat-feeding, and the partly double
row of teeth may be unique to this subgenus.

The Amboni Cave has been described by Cooke (1970) and Peet (1957: 152). Dr Minter caught
the sandflies on the wall of the cave, and informs me that about the same year four species of bats,
Tritaenops persicus afer Peters, 1877, Coleura afra (Peters, 1852), Tadarida (Chaerepha) pumila
(Cretzschmar, 1826) and Minioptems minor (Peters, 1867) were caught in the caves by Mr Colley
and identified by Dr D. L. Harrison.

Subgenus 1DIOPHLEBOTOMUS Quate & Fairchild

Phlebotomus subgenus Idiophlebotomus Quate & Fairchild, 1961: 208; Theodor, 1965: 176; Lewis, 1973:
162; 1978: 250; Lewis & Lane, 1976: 53; Abonnenc & Leger, 1976: 76. Type-species: Phlebotomus
asperulus Quate & Fairchild, 1961, by original designation.

The status of this subgenus is discussed with that of Spelaeophlebotomus. Most species live in
caves and probably feed on bats.

Key to the species of subgenus Idiophlebotomus

Females

1 Cibarial armature with median rod or big tooth 2

Cibarial armature without median rod or big tooth 7

2 Cibarial median rod with large serrations asperulus (p. 134)

Cibarial median rod with minute serrations or none ......... 3

3 Cibarium without patch of teeth except a few granulose spicules .... erebicolus (p. 134)
- Cibarium with patch of teeth ............. 4

4 Median cibarial projection short and tooth-shaped teshi (p. 135)

Median cibarial projection long and rod-shaped 5

5 Cibarial teeth in radiating lines tubifer (p. 135)

Cibarial teeth not in radiating lines 6

6 Cibarial teeth very long and parallel frondifer (p. 134)

Cibarial teeth not very long and parallel pholetor (p. 134)

7 Pharynx with scale-like teeth longiforceps (p. 134)

Pharynx unarmed 8

8 Cibarial teeth all small stellae (p. 135)

Anterior cibarial teeth very long sejunctus (p. 135)

Males

1 Apical spine of style with marked basal expansion asperulus (p. 134)

Apical spine of style without such expansion 2

2 Style with three spines 3

Style with more than three spines 4

3 Coxite with the two non-distal spines at 0-39. Paramere nearly straight with bulbous apex

erebicolus (p. 134)

Coxite with the two non-distal spines at 0-5 or 0-6, and 0-7. Paramere curved upward and pointed

longiforceps (p. 134)

4 Style with four spines frondifer (p. 134)

134 D. J. LEWIS

Style with five spines .5

5 Aedeagus prominent and capitate. Paramere slender and without dorsal appendage . pholetor (p. 134)
Aedeagus small and triangular. Paramere with basal dorsal curved appendage . . stellae (p. 135)

Phlebotomus (Idiophlebotomus) asperulus Quate & Fairchild

(Map 1)

Phlebotomus (Idiophlebotomus) asperulus Quate & Fairchild, 1961: 208 [? £]; Lewis & Lane, 1976: 54;
Lewis, 1978ft: 250. Holotype & WEST MALAYSIA (BPBM, Honolulu).

DISTRIBUTION. West Malaysia: Lewis (19786: 251, map).

Phlebotomus (Idiophlebotomus) erebicolus Quate
(Map 1)

Phlebotomus (Idiophlebotomus) erebicolus Quate, 1965: 22 [$ £); Lewis & Lane, 1976: 57; Lewis, 1978ft:
251. Holotype <J, PHILIPPINES (BPBM, Honolulu).

?. Leg formula, after Quate, 10, 186, 108; 92, 186, 106; 103, 203, 106.

Phlebotomus (Idiophlebotomus) frondifer Lewis & Lane
(Map 1)

Phlebotomus (Idiophlebotomus) frondifer Lewis & Lane, 1976: 57 [$ £]; Lewis, 1978ft: 251; Holotype^,
WEST MALAYSIA (BMNH) [examined].

9 (extrafact). Leg formula 100, 149, 91 ; 99, 167, 93; 103, 216, 101 (2-16, 0-89).

Phlebotomus (Idiophlebotomus) longiforceps (Wang, Ku & Yuan)

(Map 1)

Sergentomyia longiforceps Wang, Ku & Yuan, 1974: 334 [? <£]. Holotype cj, CHINA: Rongjiang (Chung-

Kiang Hsien) (MC, Guiyang).

Phlebotomus longiforceps (Wang, Ku & Yuan) Lewis, Young, Fairchild & Minter, 1977: 326.
Idiophlebotomus longiforceps (Wang, Ku & Yuan) Xiong et a/., 1980: 322 [intra-abdominal rods].

The description is in Chinese with an English summary, and some features are as follows.

$. Cibarium and pharynx figured. Cibarium with more than 60 small triangular teeth in rows and no
pigment patch. Pharynx with many wedge-shaped teeth of different sizes, arranged irregularly. Antenna
3 = 0-526 mm long, 1-82 length of 4 + 5, 2-3 length of labrum, 0-61 length of wing width. Palp formula
1, 4, 2, 5, 3; relative lengths 1:2-5:4:2:3. Wing figured, length 2-618 mm, 3-0 width (0-861 mm),
RI overlap = +0-406 mm, M1 + 2 fork 0-105 mm beyond origin of R4. Abdominal tergites 2-6 with
many erect hairs. Spermatheca figured, rather large, carrot-shaped and unsegmented with irregular wrinkles.
<J. Cibarium and pharynx figured. Antenna 3 = 0-714 mm long, 1-76 length of 4 + 5, 2-69 length of
labrum, two ascoids on 3-15. Palp ratio 1, 4, 2, 5, 3, relative lengths 1 : 2-37 : 4-5 : 1-87 : 2-95. Wing figured,
length 2-356 mm, 3-17 width (0-742 mm), /?t overlap = +0-280 mm, M1 + 2 fork = +0-091 mm beyond
origin of R4 . Terminalia as figured (but with intra-abdominal rods) ; paramere single, and claw-shaped
coxite approximately 2-4 length of style, with long hairs on apical half; style with three spines, one of them
apical.

Phlebotomus (Idiophlebotomus) pholetor Quate & Fairchild
(Map 1)

Phlebotomus (Idiophlebotomus) pholetor Quate & Fairchild, 1961 : 210 [$ £] ; Lewis & Lane, 1976: 57; Lewis,
1978ft: 251. Holotype 3, BORNEO: Sabah (BPBM, Honolulu).

? (extrafact, Sabah, 12.xi.1972). Leg formula 100, 149, 90; 95, 159,90; 121, 187, 100(2-13,0-88).
DISTRIBUTION. Borneo (Sabah) and Philippines: Lewis (1978ft: 236, map).

THE GENUS PHLEBOTOMUS 135

Phlebotomus (Idiophlebotomus) sejunctus (Quate)
(Map 1)

Phlebotomus (Idiophlebotomus) sejunctus Quate, 1965: 22 [?]; Lewis, 19786: 251. Holotype9, PHILIPPINES
(BPBM, Honolulu).

Phlebotomus (Idiophlebotomus) stellae Quate
(Map 1)

Phlebotomus (Idiophlebotomus) stellae Quate, 1965: 20 [? <J]; Lewis & Lane, 1976: 59; Lewis, 19786: 251.
Holotype & PHILIPPINES (BPBM, Honolulu).

cJ. Leg formula after Quate, 100, 135, 88; 97, 157, 91 ; 104, 185, 101 (0-84).

Phlebotomus (Idiophlebotomus) teshi Lewis
(Map 1)

Phlebotomus (Idiophlebotomus) teshi Lewis, 19786: 252 [$]. Holotype $, NEPAL: Pokhara (BPBM, Hono-
lulu).

$. Leg formula 100, 120, 90; 92, 156, 83; 96, 182, 134.

Phlebotomus (Idiophlebotomus) tubifer Lewis & Lane
(Map 1)

Phlebotomus (Idiophlebotomus) tubifer Lewis & Lane, 1976: 59 [?]; Lewis, 19786: 252. Holotype?, INDIA
(BMNH) [examined].

DISTRIBUTION. India: Lewis (19786: 326, map); G. B. Modi, 1979, pers. comm., Sagar in Shimoga district,
?cJ.

Phlebotomus (Idiophlebotomus) sp. A

(Map 1)

This is being described by Lewis & Dyce and is placed here in accordance with the first letter of
its future name.

Subgenus A USTRALOPHLEBOTOMUSTheodor

Phlebotomus subgenus Australophlebotomus Theodor, 1946: 99; Quate & Quate, 1967: 11, 14. Type-species:

Phlebotomus brevifilis Tonnoir, 1935, by original designation.
Phlebotomus brevifilis group; Fairchild, 1952: 192, 194, 196, 198, 199.

Key to the species of subgenus Australophlebotomus

Females

1 Pharyngeal teeth short, about 5 /im.

Spermathecal ducts wrinkled, outlets touching. Cibarium with about seven to nine main
teeth and several laterals, arch strong with lateral bulges. Maxilla with more than 150 ventral

teeth. Pharynx largely brown brevifiloides (p. 136)

Pharyngeal teeth long, about 25 /im 2

2 Spermathecal ducts with sleletal lattice near furca.

Maxilla with about 150 ventral teeth. Pharynx brown in hind half . . brevifilis (p. 136)

Spermathecal ducts without such lattice pexopharynx (p. 136)

Males

1 Aedeagus very short, less than half as long as coxite 2

Aedeagus long or rather long, more than half as long as co xite 3

2 Plunger of sperm pump wide. Genital filament with thin-walled tip opening backward brevifilis (p. 136)

136 D. J. LEWIS

Plunger of sperm pump narrow. Genital filament with thick-walled tip opening somewhat down-
ward brevifiloides (p. 136)

3 Genital filament 3-6 length of pump which has plunger without expanded base. Inter-arcal area

much wider than long trifilis (p. 137)

Genital filament 1-2-1-8 length of pump which has plunger with normal expanded base. Inter-
arcal area about as wide as long 4

4 Antenna 3 = 1-6 length of labrum. Aedeagus with straight tip .... papuensis (p. 136)
Antenna 3 = 1-1 length of labrum. Aedeagus with up-turned tip .... buccinator (p. 135)

Phlebotomus (Australophlebotomus) sp. B

(Map 1)
This is being described by Lewis & Dyce.

Phlebotomus (Australophlebotomus) brevifilis Tonnoir
(Map 1)

Phlebotomus brevifilis Tonnoir, 1935: 145 [? £]; Fairchild, 1952: 192. Specimens stated by Tonnoir to be?

and c? types being studied by Lewis & Dyce, AUSTRALIA (ANIC, Canberra).
Phlebotomus (Australophlebotomus} brevifilis Tonnoir ;Theodor, 1948: 100, 108.

DISTRIBUTION. Australia : information mainly from A. L. Dyce.

Phlebotomus (Australophlebotomus) brevifiloides Fairchild

(Map 1)

Phlebotomus brevifiloides Fairchild, 1952: 194 [?]. Holotype $, AUSTRALIA (CIH, Sydney).
DISTRIBUTION. Australia : information mainly from A. L. Dyce.

Phlebotomus (Australophlebotomus) buccinator Fairchild
(Map 1)

Phlebotomus buccinator Fairchild, 1952: 194, 195, 205 [£]. Holotype <J, AUSTRALIA: Cairns (SPHTM,
Sydney).

Phlebotomus (Australophlebotomus) sp. C

(Map 1)
This is being described by Lewis & Dyce.

Phlebotomus (Australophlebotomus) papuensis Fairchild
(Map 1)

Phlebotomus papuensis Fairchild, 1952: 200 & 193, 195, 203 [£]. Holotype <J, PAPUA NEW GUINEA: Doba-

dura (CIH, Sydney).
Phlebotomus (Australophlebotomus) papuensis Fairchild ; Quate & Quate, 1967: 11.

Phlebotomus (Australophlebotomus) pexopharynx Fairchild

(Map 1)

Phlebotomus pexopharynx Fairchild, 1952: 196 [?]; Quate & Quate, 1967: 14. Holotype?, AUSTRALIA:
Cairns (CIH, Sydney).

THE GENUS PHLEBOTOMUS 137

Phlebotomus (Australophlebotomus) trifilis Quate & Quate
(Map 1)

Phlebotomus (Australophlebotomus) trifilis Quate & Quate, 1967: 1 1 [<?]. HolotypecJ, PAPUA NEW GUINEA:
Vogelkop (BPBM, Honolulu).

NOTE. This species is presumably named after the three spines on the style.

Subgenus PHLEBOTOMUS Rondani & Berte

Phlebotomus subgenus Phlebotomus Rondani & Berte; Theodor, 1948: 96; 1958: 16; Quate, 1964: 238;
PernTev, 1968: 62, 63, 65, 79, 227; Hennig, 1972: 24, 51, 53 [this subgenus, Euphlebotomus and Anaphlebo-
tomus seem to be related to the fossil P. tipuliformis and should perhaps be united under a common
name]; Lewis, 19786:231.

Key to the species of subgenus Phlebotomus
Females

1 Most pharyngeal scaly teeth arranged obliquely and pointing backward . . . bergeroti (p. 137)
Most pharyngeal scaly teeth not pointing backward duboscqi (p. 138), papatasi (p. 138), sale hi (p. 141)

Males

1 Upper process of paramere not longer than paramere 2

Upper process of paramere longer than paramere 3

2 Upper process of paramere clothed with hairs on all sides duboscqi (p. 138)

- Upper process of paramere clothed with hairs on hind part sale hi (p. 138)

3 Antenna 3 = 0-27 to 0-34 mm long. Wing length about 2-2-2-7 mm. Coxite long, 0-55 to 0-6 mm.

Style 0-37-0-41 mm long, distance between basal and middle spines less than that between
middle and distal spines. Eye appearing small because head relatively short . papatasi (p. 138)
Antenna 3 = 0-24-0-28 mm long. Wing length about 1-8-2 mm long. Coxite not long, 0-37-
0-4 mm. Style 0-27 mm long, distance between basal and middle spines greater than or equal to
that between middle and distal spines. Eye appearing large because head relatively long.

Distal spines of surstyle long and thin bergeroti (p. 137)

Phlebotomus (Phlebotomus) bergeroti Parrot
(Map 2)

Phlebotomus papatasi var. bergeroti Parrot, 1934: 383 [<?]; 19416: 237 [$]; Parrot & Bellon, 1952: 60.

Syntypes 3 J, ALGERIA (IP, Algiers).
Phlebotomus (Phlebotomus) viduus Parrot, 1936a: 34 [$]. Syntypes 3 ?, ETHIOPIA (IP, Algiers?). [Synony-

mized by Theodor, 1948: 106.]
Phlebotomus (Phlebotomus) bergeroti Parrot; Lewis & Biittiker, 1980 [synonymy including description of 9

by Bellon in 1936]; PernTev, 1968: 66; Artemiev, 1978: 16.

? (extra fact, Yemen, Taiz, 29.xi.1970). Leg formula 100, 106, 68; 103, 136, 76; 120, 175, 93 (0-78).
<J (extra fact, Saudi Arabia). Head 0-41 (0-38-0-43) mm long, 0-22 (0-22-0-23) length of wing (n = 5).

DISTRIBUTION. Africa: Abonnenc & Rioux (1961: 35, map). Africa, Mediterranean area etc.: Abonnenc
(1972: 255, map). Djibouti area: Courtois (1972). Ethiopia: Danakil ('Asaita', 4.U968, 2$ biting man,
R. W.A.); Humera (29.V.1968, 1 & R. W.A.); Kirk & Lewis (1952: 339). Iran: Theodor & Mesghali (1964:
286). Morocco: Rioux et al. (1975: 495, map). Saudi Arabia: Lewis & Biittiker (1980, map). Somalia: Burao
(1913? 1 (J, R. E. D. B.). South Yemen: Pringle (1960, 'Mazu' and Yashbum, assumed to be Wadi Yeshbum);
Ba Zulayfah (x.1962, S. A. S., see under P. papatasi). Sudan: Lewis & Kirk (1951 : 565; 1957: 632, revised list).
United Arab Emirates: Al Ain (Ain al Faidr, 1980, G. B. W., det. R. P. L.). Yemen: Lewis (1974).

NOTE. This species is thermophilic and xerophilic in Afghanistan (Artemiev, 1978: 16). It bites
man readily and has been suspected of transmitting CL in the central Sahara and sandfly-fever in
Ethiopia (Abonnenc, 1972: 95).

138 D. J. LEWIS

Phlebotomus (Phlebotomus) duboscqi Neveu-Lemaire
(Map 2)

Phlebotomus duboscqi Neveu-Lemaire, 1906: 65 [? £1; Austen, 1909: 20; Abonnenc, 1958: 61; 1959: 333;

Abonnenc & Lariviere, 1958: 260 [egg & larva]; Abonnenc & Rioux, 1961: 32; Ashford, 1974: 607.

Syntypes 6 $, 6 c?, MALI (depository unknown).

Phlebotomus duboscqi Neveu-Lemaire; Picard, 1909: 165. [Mis-spelling.]
[Phlebotomus pappatasi (Scopoli); Picard, 1909: 165. Misidentification, Abonnenc, 1958: 65.]
Phlebotomus duboscqii Neveu-Lemaire; Alcock, 191 1 : 119. [Mis-spelling.]

Phlebotomus duboscqui Neveu-Lemaire; Newstead, 1912: 367; 1913: 124; Summers, 1913: 114. [Mis-
spellings.]

[Phlebotomus papatasii (Scopoli); Newstead, 1913 : 125. Misidentification, Abonnenc, 1958 : 65.]
Phlebotomus roubaudi Newstead, 1913: 125 [conditional name]; 1914: 187; Rageau, 1951: 794. Holotype^,

MAURETANIA (IP, Paris?). [Synonymized by Larrousse, 1921 : 44; Abonnenc, 1958: 65.]
Phlebotomus (Phlebotomus) roubaudi Newstead; Parrot & Gougis, 1944: 40; Kirk & Lewis, 1946a: 42;

1946b: 120; 1947: 875; 1951 : 424; Kervran, 1946: 155; Lewis & Kirk, 1957: 634; Perfil'ev, 1968: 4.
Phlebotomus (Phlebotomus) duboscqi Neveu-Lemaire; Kirk & Lewis, 1946a: 41 ; 1951 : 427; Perfil'ev, 1968 : 4,

65, 66; Abonnenc, 1972: 95.
Phlebotomus roubaudi var. fourtoni Floch & Abonnenc, 1948: 1. Holotype <J, UPPER VOLTA (depository

unknown). [Synonymized by Abonnenc, 1958: 65.]
Phlebotomus (Phlebotomus) roubaudi var. fourtoni Floch & Abonnenc; Kirk & Lewis, 1951: 325; Lewis &

Kirk, 1954: 34.

? (extra fact, Senegal, 1977-1978). Leg formula 100, 108, 61 ; 101, 106, 60; 126, 173, 98 (0-75).

DISTRIBUTION. Africa etc.: Abonnenc (1972: 255, map; 1973: 185, northernmost point 20° near Akjoujt,
Mauretania). Ethiopia: Ashford (1974: 607); Gemetchu et al. (1976: 81). Gambia: Snow (1979). Ghana:
Nangodi (10.iv.1962, <J, D. J. L.). Sudan: Hoogstraal & Heyneman (1969: 1155); Lewis & Kirk (1951: 566,
map; 1957: 632, revised list); Qutubuddin (1962: 594). Togo: Abonnenc (1973: 190, map).

In the Senegal River valley the finding by Dedet et al. (1980) of only 30 P. duboscqi and two P.
rodhaini with 2245 Sergentomyia exemplified the paucity of Phlebotomus in the tropics.

NOTE. P. duboscqi is probably more primitive than bergeroti or papatasi; its distribution corresponds with
that of CL in West Africa; it is common in dwellings in Upper Volta, and has been suspected as a vector in
Niger (Abonnenc, 1972: 34, 99) and Senegal (Abonnenc, 1973: 185; Hoogstraal & Heyneman, 1969: 1170).
Dedet et al. (1979: 435, 436) found it infected with Le. major in Senegal.

Phlebotomus (Phlebotomus) papatasi (Scopoli)
(Map 2)

Bibio papatasi Scopoli, 1786: 55 [$]. Type(s), ITALY (U, Pavia?).

? M usca papatasi (Scopoli) Gmelin, 1 790 : 2866 ; Costa, 1 843 : 6 ; Sinton, 1928: 300. [Position uncertain.]

ICiniphes molesta Costa, 1840: 2225; 1843: 4. Syntype(s), ITALY (depository unknown). [Synonymized by

Sinton, 1928: 300; Kirk & Lewis, 1951: 422; Perfil'ev, 1968: 1, 7 [P. molestus from Asia], 228 [as

'Cyniphes molesta'].] [Position uncertain.]
Flebotomus papatasii (Scopoli) Rondani, 1840: 13; Costa, 1843: 4, 6 [Terra di Otranto] [^, short note on

terminalia; Scopoli's description of fly superficial].

Hebotomus papatasii (Scopoli) Rondani, 1843 : 265, pi. 10, recognizable figure of style).
Hebotomus molestus (Costa); Rondani, 1843: 266 [habitat Kingdom of Naples; quotes "Cyniphes molesta J.

Costa' and 'Cyniphes J. Costa'].
Phlebotomus papatasi (Scopoli) Loew, 1847: 151; Sacca, 1950: 684 [early stages]; Hemming, 1958: 16, 189

[in official list of specific names] ; Abonnenc, 1958: 63; 1959: 329; Schmidt & Schmidt, 1963: 567; Bhat &

Modi, 1976: 265; Lysenko & Beliaev, 19776: 263 [variation] ; Guevara-Benitez, Ubeda-Ontiveros &

Morillas-Marquez, 1978: 824, 832.
M usca papatasii (Scopoli); Rondani, 1856: 178.
Phlebotomus papatasii (Scopoli); Noe, 1905: 722; Grassi, 1907: 356 ['J. Costa' mentioned], 359, 384, [all

Italian Phlebotominae thought to be one species]; 1908: 681 [two species known in Rome]; Summers,

1911: 105; Handlirsch, 1925: 96 [in list of names not to be changed]; Pierantoni, 1925: 4; Sinton, 1928:

THE GENUS PHLEBOTOMUS

139

300 [synonymy]; Patton & Evans, 1929: 101 ; Shchurenkova, Demina & Pavlova, 1929: 683; Isaev, 1935:
95, 103 [cibarium, spermatheca etc.]. [Mis-spellings.]

Phlebotomus pappatasi (Scopoli); Picard, 1909: 164. [Mis-spelling.]

Phlebotomus pappatasii (Scopoli); Mansion, 1913: 639. [Mis-spelling.]

Phlebotomus (Phlebotomus) papatasi var. breviventris Ristorcelli, 1941: 369 [?]. Syntypes 2 ?, MOROCCO
(depository unknown) [treated as former subspecies according to ICZN, 1974: Article 45 (e)]. Syn. n.

Phlebotomus (Phlebotomus) papatasi (Scopoli); Theodor, 1948: 86, 106, [genus and species in "nomina con-
servanda" of Handlirsch, 1925]; 1958: 17; Kirk & Lewis, 1951 : 422 [synonymy, Handlirsch's list] ; Parrot,
1953: 115. Fairchild, 1955: 188; Lewis & Kirk, 1957: 632; Quate, 1964: 240; PernTev, 1968: 1, 5, 49
[larva], 50, 60 [egg], 62, 65, 66, 228; Croset, 1969: 360; Rioux & Golvan, 1969: 68; Bailly-Choumara,
Abonnenc & Pastre, 1971: 436; Artemiev, 1972: 300 [spiracles]; Biocca, Coluzzi & Constantini, 1977a:
158, 160 [absent from Milan area and very rare in Italy where there are seven species]; 19776: 31;
Artemiev, 1978: 15; Lewis, 19786: 233 [synonymy]; Croset, Rioux, Maistre & Bayar, 1978: 726, 732;
Lewis & Buttiker, 1980.

Phlebotomus papatasi papatasi (Scopoli); Abonnenc & Rioux, 1961 : 31 [in distinction from bergeroti].

9 (extra facts). Leg formula (Hofuf, Saudi Arabia, n = 10): 100, 103-0 (96-108), 58-4 (55-62); 100-6 (98-103),
124-6 (114-133), 69-2 (64-74); 119-7 (117-123), 164-3 (154-174), 91-2 (87-96).

cJ (extra facts). Head 0-42 (0-38-0-42) mm long, 0-208 (0-20-0-21) length of wing (n = 5, Saudi Arabia). Leg
formula (Hofuf, Saudi Arabia, n = 10): 100, 110-8 (104-117), 63-4 (57-79); 93-7 (67-103), 132-5 (124-135),
74-9 (69-78); 114-1 (110-118), 165-6 (158-175), 91-9 (80-97). Basitarsus 3/femur 3: India, 84-1 (79-86,
n = 10); Saudi Arabia, 80-7 (71-87, n = 10).

11

Figs 10-14 Abdominal segment 9 of males of Phlebotomus species in dorsal view. (10) P. papatasi; (11)
P. alexandri; (12) P. guggisbergi;(i3) P. hindustanicus; (14) P. argentipes.

140 D J LEWIS

DISTRIBUTION. Western part of Old World: Beklemishev & Dolmatova (1948: 354, map); Croset (1969: 275,
map); Dolmatova & Demina (1971 : 121, map); Hennig (1966: 59, 62, map to indicate marginal speciation);
Lewis in Tesh et al (1976: 666, map); Perfil'ev (1968: 89). Africa, Mediterranean area etc.: Abonnenc (1972:
255, map). Indian subcontinent: Lewis (19786: 324, map). Afghanistan: Artemiev (1974: 157, map; 1978: 15);
Arsenieva & Neronov (1978: 30, 32); Nadim et al. (1979: 33, Robatak, 36°09'N, 68°24'E). Algeria: Biskra
(20.V.1893, A. E. £.); Dedet & Addadi (1977: 86); Dedet et al. (1977: 256). Crete: Hadjinicolaou (1958: 974,
975); Hertig (1949a: 782, 787-789. Egypt: Abu Aweigila (Sinai, vii.1979, Y. S.); Alexandria, Aswan, Asyut,
Bahtim, Beni Suef, Giza, Imbaba, 'Kafra el Sheikh', Luxor, Maadi, Marsa Matruh, Matariya, Qaliubiya,
Rashda, Sharqiyah and Tanta (1967, M. A. R.); Birqet Qarun (ix.1945, R. L. C.); Siwa (23.V.1935, J. 0. C.);
Zein el Dine (1972: 271, Dakhla oasis). Ethiopia: Gemetchu et al. (1977: 209). France: Colas-Belcour (1958:
826); Raynal (1954: 315, map); Rioux & Golvan (1969: 51, 72, maps). Greece: Hadjinicolaou (1958: 968-
973); Hertig (1949a: 778, 779, 781-783, 786-789); Leger et al. (1979: 17); Volo (15.viii.l892,& E. M., sent to
BMNH 18.viii.1892 by A. A. Merlin, H. M. Consul, Volo). Hungary: 'Carlapago' (12.vii— , K. K., be-
queathed to BMNH in 1929 by A. E. E.). India: Modi et al. (1978: 748, map); Modi & Soman (1978: 159);
Pandya et al. (1977: 133-135). Iran: Lewis, Mesghali & Djanbakhsh (1961 : 206); Mesghali (1961 : 26; 1963:
1073); Nadim, Mesghali & Javadian (1977: 215). Iraq: Ahmad (1976: 86, 96, 97); Amara (14.vi.1918,
P. A. B.); Basra (x.1918, H. W. L.); Pringle (1953: 723, map). Israel: Adler & Theodor (1929: 271); Pazael, c.
31°67'N, 35°33'E, 12.vii.1979, Y. S.). Italy: Biocca et al. (1977a: 160, map; 19776: 20, rare, 29, map);
Corradetti et al. (1956a: 6, map); Hertig (1949a: 796); Maroli & Bettini (1977: 318). Jordan: Awjan (32°02'N,
36°04'E, 15.vii.1978, E. K. S.); Sweilah (32°02'N, 35°50'E, 2.vii.l978, E. K. S.). Kuwait: Al Fantas, 29°10'N,
48°06'E, Al Farwaniya area, 29°16'N, 47°55'E, 'Al Kazuma', Kuwait & 'Malboula', 1974, K. B.); Hussien &
Behbehani (1976). Libya: Ashford et al. (1977: 265, Al Birkah, 32°05'N, 20°05'E, etc.); Nalut & Yafran area,
32°04'N, 12°31'E, M. A.). Morocco: Bailly-Choumara et al. (1971: 453, map); Gaud (1954: 95). Oman:
Barka (x.1979, G. S.). Pakistan: Robinson & Blackham (1912). Sardinia: Hertig, 1949a: 798). Saudi Arabia:
Lewis & Buttiker (1980, map). Southern Yemen: Amwadhia, 45°50'N, 13°46'E, Ba Tays, 13°20'N, 45°18'E,
Ba Zulayfah, 13°50'N, 45°47'E, Dali, 13°42'N, 44°44'E, Khodad, 13°09'N, 44°50'E, Mukalla area, Nigda,
13°56'N, 45°55'E, Sah, 15°34'N, 48°51'E and Saywun area, 15°56'N, 49°27'E (1962, S. A. S.); 'Haski' (1939,
cJ, P.); Sha'b Subeihi, 13°05'N, 44°30'E(?) (iv.1954, N. L. C.). Spain: Gil Collado (1977: 186, map);
Guevara-Benitez et al. (1978: 814); Najera (1937: 1488); Zariquiey (1944: 18). Sudan: Akasha, Dal Island,
Ferka, Sai Island, 1980, G. S.); Hoogstraal & Heyneman (1969: 1544). Syria: Adler & Theodor (1929: 271);
Arab Hassan, 36°30'N, 37°51'E, 'Halab Halwan', 'Kanat Albu', 'Klemis', 'Khan el Sobol', 'North Kena
Halab' (1978, K. Z. D.). Tunisia: Chadli, Dancescu et al. (1970: 363); Dancescu, Romain et al. (1970: 357);
Croset (1969: 273, map); Croset et al. (1978: 727, map); Dedet (1971 : 157). Turkey: Yasarol (1980). U.S.S.R.:
Gaibov (1975a: 55, Fergan area; 1976: 49, Surkhandar'ya area); Grebelsky (1937: 200, Khiva); Karapet'yan
& Babayants (1979: 67, Bayram-Ali); Latyshev & Posyval (1937: 184, Seraks); Petrishcheva (1937: 148);
Ponyrovskyi (1971: 495, Sumbar Valley). Yemen: Buttiker & Lewis (1979: 371). Yugoslavia: Trebinje
(barracks, 24.vii.1908, S. T.);Zivkovic (1972: 27).

NOTE. It is relevant to review briefly some aspects of the taxonomic history of this, the type-
species of the genus and the first-known sandfly in the world, and an important disease vector. In
pre-Linnaean times, in 1691, in Rome Philippo Bonanni (discussed by M. Lavoipierre in a paper
cited by Theodor, 1948: 86, and Lewis, 1977: 94) published the first description of a (male)
Phlebotomus but the species is unknown and may not have been papatasi. The next description, of
a female of Bibio papatasi fron the Milan area once known as Insubria, was published in 1786 in
the city of Pavia, then in the Austrian Empire, by Johann Anton Scopoli (1723-1788), an eminent
naturalist (Ambrosi, 1889; Gilbert, 1977; Higgins, 1963; Voss, 1881). The species was named from
'pappataci', the vernacular name for a sandfly, which means a silent gorger. A collection of
Scopoli was probably destroyed by shipwreck and fire in 1766 (Horn & Kahle, 1935-1937); he
was shipwrecked on the River Inn in Bavaria and had two fires in his house in Istria (Voss, 1882:
21). However, he probably studied P. papatasi later, when professor of chemistry and botany at
the University of Pavia from 1776 to 1788 (Conci, 1975: 1013). The type is not in the NM, Vienna
and is presumed to be lost. Schiner (1856: 405) praised Scopoli's work on Diptera, which had
been disregarded until recently brought to light by English workers, but pointed out that some of
his descriptions were out of date by modern standards. His figure of P. papatasi is rather crude.

In 1840, when P. papatasi was the only known sandfly in the World, 'C. molesta\ from Terra di
Otranto, a former province around Otranto, was described by Giuseppe Costa, son of O. G.
Costa (1787-1867), professor of zoology at Naples from 1836 to 1849 (Conci, 1975: 887). In 1843,

THE GENUS PHLEBOTOMUS 141

in a paper in antiquated Italian, G. Costa linked the names molesta and papatasi but seemed to
consider the species different, pointed out that Ciniphes was an ancient name [gnat in Latin,
Bailey, 1828: 361] and that papatasio was the vulgar name for a notorious insect in parts of
Lombardy. Also in 1843 Camillo Rondani, the eminent entomologist of Parma (Gilbert, 1977:
322; Meade, 1879: 138) figured the style of a species from central Italy which is now generally
regarded as P. papatasi, gave the area of H. molestus as the Kingdom of Naples, and described
another species of sandfly (Sergentomyia minuta) from the plain of Parma. Grassi (1907: 8), who
referred to Rondani's descriptions as imperfect, recognized (Grassi, 1908) two species of Phleb-
otomus s. str. in Italy, and Biocca et al. (1977 'a) reported seven, stated that P. papatasi was very
rare, and showed no record of it from Pavia. The listing of P. papatasi among 'nomina conser-
vanda' did not settle the specific name because it was intended to decide on the generic name. It is
uncertain whether B. papatasi (Scopoli, 1786) and H. papatasii (Rondani, 1843) were the same
species but here, for practical reasons, the situation is regarded as stabilized and the synonymy as
shown above.

The spelling papatasii is a mistake which lasted for more than a century.
The description off. p. var. breviventris does not seem to justify its recognition as a taxon.
Small females (and males) are sometimes seen with a wing length much less than the 2-0 to
2-4 mm given by Theodor (1958 : 18). For example, one from Israel (Pazael, 12.vii.1979) had a WL
of 1-78 mm and width of 0-48 mm.

In the U.S.S.R. P. papatasi needs a warm summer and a mild winter, with a mean temperature
not below — 6°C and a short rainy season, and it has a limited altitude range (PernTev, 1968: 89).
In Afghanistan it is found up to 2100 m but occurs mainly in the plains where the water table is
high (Artemiev, 1978: 15). This species is largely domestic over most of its range (Lewis &
Biittiker, 1980) and, perhaps for this reason, is widespread and abundant. It is mainly domestic in
France, Tunisia and Yugoslavia (Croset et al., 1978: 726, 738) and Greece (Leger et al., 1979: 19),
and purely domestic in Spain (Gil Collado, 1977: 185). P. papatasi is much more anthropophilic
than some species (Strelkova, 1974).

It has been suspected of transmitting VL in some areas where no likely vector was found
(Lewis & Biittiker, 1980), including Iraq (Abul-hab & Azawi, 1978: 406; Adler, 1964: 79; Azawi &
Abul-hab, 1976; Sukkar, 1972: 69) but is evidently a poor potential vector (Abonnenc, 1972: 100;
Safyanova, 1967: 38; Williams & Coelho, 1978: 17), and another species may be responsible.

P. papatasi and P. sergenti are the main vectors of CL to man (Theodor, 1964: 488; Wilcocks &
Manson-Bahr, 1972: 135). P. papatasi (according to Lysenko & Beliaev, 19776: 262) and P.
caucasicus transmit CL among rodents in Central Asia (Theodor, 1964: 489), and Safyanova
(1977: 257, 259) has distinguished two types of zoonotic foci in the U.S.S.R., a dangerous one with
P. papatasi, and another with P. caucasicus and P. andrejevi as vectors under different conditions.
In some areas P. papatasi transmits zoonotic Le. t. major from animals to man (Adler, 1964: 67;
Lysenko, 1971 : 516; M6skovskij & Dukhanina, 1971 : 732; Sergiev, 1979: 199). P. papatasi is the
principal vector of zoonotic CL in Central Asia (Safyanova & Alekseev, 1977: 154) and appears
to be the vector of CL in Iran (Adler, 1964: 791; Bray, 1974: 93; Javadian et al., 1977: 203;
Nadim & Rashti, 1971 : 100), Saudi Arabia (Nadim, Rashti & Ashi, 1979), Afghanistan (Nadim,
Javadian et al., 1979) and India (Lewis, 1977: 134). This species is a troublesome biter and a well
known vector of CL in many areas (Abonnenc, 1972: 100) including Israel (Molyneux, 1977: 49;
Egoz & Michaeli, 1978) and the Mediterranean basin generally (Dedet, 1979: 72). It transmits
sandfly fever.

Phlebotomus (Phlebotomus) salehi Mesghali

(Map 2)

Phlebotomus (Phlebotomus} salehi Mesghali, 1965: 264 [£]; Mesghali & Rashti, 1968: 770 [?]; Kalra &

Lewis, 1976: 522; Lewis, 19786: 235; Artemiev, 1978: 16. Holotype^, IRAN (IPH, Tehran).
\_Phlebotomussaheli Mesghali; Killick-Kendrick, 1978: 300. Mis-spelling.]

3 (extra fact, India). Leg formula 100, 1 13, 63; 122, 172, 98; 125, 170, 98 (0-74).

142 D. J. LEWIS

DISTRIBUTION. India: Lewis (19786: 235, map). Iran: Mesghali & Rashti (1968: 768, 769, map); Nadim et al.
(1977:215).

NOTE. P. salehi probably transmits CL among rodents in India (Kalra & Lewis, 1976; Killick-
Kendrick, 1978:298).

Subgenus PARAPHLEBOTOMUS Theodor

Phlebotomus subgenus Paraphlebotomus Theodor, 1948: 97; 1958: 19; 1965: 175; Perfil'ev, 1968: 49 [larva],
63, 66, 80, 232; Abonnenc, 1972 [as sergenti group]; Lewis, 19786: 235. Type-species: Phlebotomus
sergenti Parrot, 1917, by original designation.

The inclusion of P. kazeruni necessitates the following slight change to Theodor's (1958) defini-
tion of the subgenus: 'usually the spermatheca segmented. . .; in P. kazeruni very short, unseg-
mented and expanded distally.'

Key to the species and subspecies of subgenus Paraphlebotomus

Females

1 Antenna 3 short (0-12-0-16 mm), 0-5-0-6 length of labrum 2

Antenna 3 long (0-22-0-33 mm), 0-7-1-0 length of labrum 3

2 Armature about 0-17 length of pharynx with straight hind edge; pharynx conical with hind

width 2-0-2- 5 times fore width alexandri (p. 143)

Armature about 0-25 length of pharynx with distinctly concave hind edge and smaller teeth;
pharynx bulging, with hind width 3-4 times fore width marismortui (p. 146)

3 Spermatheca not segmented, its body about as long as wide.

Mesonotum brown kazeruni (p. 145)

Spermatheca segmented, its body much longer than wide 4

4 Pharyngeal teeth weakly developed, most obliquely arranged, occupying only fifth or sixth

length of pharynx andrejevi (p. 144), caucasicus (p. 144), mongolensis (p. 147)

Pharyngeal teeth very distinct, hind ones rounded, most teeth pointing inward and posteriorly,
armature usually occupying hind 0-1 6 or 0-20 of pharynx length 5

5 Pharyngeal armature 0-1 5-0- 17 length of pharynx 6

Pharyngeal armature 0-20-0- 3 3 length of pharynx 7

6 Antenna 3 = 0-44 mm long. Africa and Saudi Arabia saevus (p. 147)

Antenna 3 -0-3 1-0-33 mm long. U.S.S.R sergenti similis (p. 148)

7 End segment of spermatheca bell-shaped on narrow stem.

Pharyngeal teeth numerous chabaudi (p. 145)

End segment of spermatheca ring-like and sessile 8

8 Spermatheca with four or five segments.

Wing index 1-2-1-9 sergenti sergenti (p. 147)

Spermatheca with seven to nine segments 9

9 Pharynx with eight or nine rows of four to five teeth. Antenna 3 about 0-23 mm long jacusieti (p. 145)
Pharynx with 10-12 rows of six to eight teeth. Antenna 3 about 0-26-0-30 mm long . nuri (p. 146)

Males

1 Basal lobe of coxite with most of hairy surface ventral, lobe very large, about 40 /zm wide

caucasicus (p. 144)
Basal lobe of coxite with all or most of hairs terminal 2

2 Antenna 3 short (0-12-0-16 mm), 0-7-0-9 length of labrum. Sperm pump small (0-12 mm) with

plunger slightly wider than barrel 3

Antenna 3 long (0-24-0-34 mm), 1-0-1-4 length of labrum. Sperm pump large (0-17-0-20 mm)

with plunger much wider than barrel 4

3 Style three times as long as thick, less than half as long as coxite. Two spines on style terminal

marismortui (p. 146)
Style four times as long as thick, more than half as long as coxite. One spine of style terminal, the

next at 0-7 alexandri (p. 143)

4 Aedeagus with a narrow tip 5

Aedeagus tip curving outward and slightly backward so that in side view it appears angular,

prow-shaped or occasionally rounded . 6

THE GENUS PHLEBOTOMUS 143

5 Lobe of coxite large, about 0-080 mm long. Tip of aedeagus pointed .... saevus (p. 147)
Lobe of coxite small, about 0-034 mm long. Tip of aedeagus narrowly rounded . chabaudi (p. 145)

6 Style less than half as long as coxite, three times as long as thick.

Basal lobe of coxite slender, distinctly longer than thick 7

Style length equal to or more than half that of coxite 8

7 Basal lobe of coxite 0-06 mm long and 0-02 mm wide with small scarcely differentiated head.

Surstyle 0-26-0-28 mm long sergenti sergenti (p. 147)

Basal lobe of coxite 0-07-0-08 mm long and 0-03 mm wide with rounded slightly differentiated
head. Surstyle 0-3 1-0-32 mm long. U.S.S.R sergenti similis (p. 148)

8 Style about half length of coxite, about four times as long as thick.

Basal lobe of coxite about 0-03 mm wide. Tip of aedeagus appearing rounded or prow-
shaped in side view kazeruni (p. 145)

Style longer than half length of coxite 9

9 Second spine of style at 0-7 10

Second spine of style at 0-8 or nearer tip 11

10 Style 3-3 times as long as wide mongolensis (p. 147)

Style 4-0 times as long as wide nuri (p. 146)

11 Style 0-35 length of coxite. Basal lobe of coxite 0-08 mm wide .... andrejevi (p. 144)
Style 0-53 length of coxite. Basal lobe of coxite 0-07 mm wide .... jacusieli (p. 145)

Phlebotomus (Paraphlebotomus) alexandri Sinton
(Map 3)

Phlebotomus sergenti var.; Newstead, 1920: 309 [£]. [Synonymized by Sinton, 1928: 308.]

Phlebotomus sergenti var. alexandri Sinton, 1928: 308 [<J]; Adler, Theodor & Lourie, 1930: 533 [?].

Lectotype <£ IRAQ: Amara, 19.ix.1918 (P. A. B.) (BMNH), here designated [examined].
Phlebotomus (Paraphlebotomus) alexandri Sinton; Perfil'ev, 1968: 63, 67, 72-74, 241; Croset, 1969: 285;

Artemiev, 1978: 17; Croset, Rioux, Maistre & Bayar, 1978: 728, 732; Lewis & Buttiker, 1980 [synonymy].
Phlebotomus (Phlebotomus) alexandri Sinton; Abonnenc, 1972: 102, 103 ['Localite type: Amara

(Mesopotamie). Holotype: Depose au B. M.'] ; Guevara-Benitez, Ubeda-Ontiveros & Morillas-Marquez,

1978:833.

? (extra fact, Saudi Arabia, Wadi Mizbil, 11. viii. 1977). Leg formula 100,94, 51; 99, 115,62; 124, 153,87(1-72,
0-55).

DISTRIBUTION. Africa, Mediterranean area etc.: Abonnenc (1972: 256, map); Croset (1969: 289, map).
Afghanistan: Artemiev (1974: 157, map); Nadim et al. (1979: 33, Robatak). Algeria: Dedet (1979, map in
litt.); Dedet & Addadi (1977: 86); Rioux et al. (1970: 877). China: Xiong et al. (1964, Xinjiang); Wang et al.
(1963, Gansu Province). Cyprus: Croset et al. (1978: 729). Djibouti area: Courtois (1972). Ethiopia: Ge-
metchu et al. (1977: 209). Greece: Hertig (1949a: 781, 784-786); Leger et al. (1979: 17). Iran: Lewis et al.
(1961: 306); Mesghali (1961: 55, map); Nadim et al. (1977: 215); Theodor & Mesghali (1964: 286). Iraq:
Ahmad (1976: 86, 98); Pringle (1953: 721). Israel: Theodor (1947: 95). Pakistan: Lewis (19786: 325, map).
Rumania: Duport et al (1971 : 387). Saudi Arabia: Lewis & Buttiker (1980, map). Spain: Rioux et al. (1974a:
121, map). Tunisia: Croset (1969: 287, map); Croset et al. (1970: 872; 1978: 727, map). Turkey: Houin et al.
(1971: 638); Yasarol (1980). United Arab Republic: Al Ain (G. B. W., 1980, Ain al Faidr, det. R. P. L.\
U.S.S.R.: Croset et al. (1978: 728, Crimea); Gaibov (1975a: 55; 19756, Fergan area); Karapet'yan & Babay-
ants (1979: 67, Bayram-Ali, 37°37'N, 62°10'E); Latyshev & Pozyvai (1937: 184); Petrishcheva (1935: 206;
1937: 148); Rasnitsyna (1974). Yemen: Lewis (19746).

NOTE. Newstead cited two males of 'sergenti var.' and also six females which prove to be another
species. All eight are in the BMNH.

In Afghanistan P. alexandri is mainly a mountain species, is thermophilic and somewhat
hydrophilic, and bites man readily (Artemiev, 1978: 17). It may play some part in VL transmis-
sion in China (Xiong et al., 19636: 610), and is considered to be an important vector of CL in the
southern U.S.S.R. (Dedet, 1979: 72; Petrishcheva, 1971: 573). Females have been found infected
with flagellates in a CL area of Iran (Javadian et al, 1977: 203, 204) and the species is suspected
of transmitting CL in Tunisia (Croset et al, 1978 : 729), owing to its relationship to P. sergenti.

144 D. J. LEWIS

Phlebotomus (Paraphlebotomus) andrejevi Shakirzyanova
(Map 3)

Phlebotomus sergenti var. andrejevi Shakirzyanova, 1953: 103 [9 £|; Gaibov, 1956: 63. Syntypes?, about
760(10 per cent of 7613), U.S.S.R. (depository unknown).

Phlebotomus (Paraphlebotomus) andrejevi Shakirzyanova; Theodor & Mesghali, 1964: 286 [till then un-
known outside the U.S.S.R.]; Perfil'ev, 1968: 67, 72, 249; Croset, Abonnenc & Rioux, 1970: 867; Lewis,
1971 : 535 [close to caucasicus~\ ; Artemiev, 1974: 159.

Phlebotomus (Paraphlebotomus) mofidii Theodor & Mesghali, 1964: 289; Croset, Abonnenc & Rioux, 1970:
867. Holotype <J, IRAN (BMNH) [Synonymized by Artemiev, 1978 : 18.]

DISTRIBUTION. Afghanistan : Artemiev (1974: 157, map; 1978: 18). Iran: Theodor & Mesghali (1964: 287, 290,
north-east, partly as mofidii). Mongolia: Artemiev (1978: 18); Neronov & Gunin (1978: 23, Bayan-
Khongorsk, 45°30'N, 99°30'E, central, eastern and southern Gobi). U.S.S.R.: Dergacheva (1974: 1668,
Karshinskaya Steppe); Dergacheva & Turzhanova (1977, Mangyshlak Peninsula); Dergacheva & Zerikhina
(1974: 425); Karapet'yan & Babayants (1979: 67, Bayram-Ali); PernTev (1968: 251, Alma Ata Province and
Kzyl-Orda); Rasnitsyna (1974); Theodor & Mesghali (1964: 287, 290).

NOTE. In the U.S.S.R. (Eliseev & Dergacheva, 1972) and in Afghanistan (Artemiev, 1978: 18) this
species occurs mainly in sandy deserts, and in south-east Turkmenia it and P. caucasicus live in
drier areas than P. papatasi, according to Safyanova (1979: 78). In the U.S.S.R. (Dubrovsky,
1976; Sergiev, 1979: 199) P. andrejevi plays a major part in disseminating Le. tropica major.

Phlebotomus (Paraphlebotomus) caucasicus Marzinowsky
(Map 3)

Phlebotomus (Hebotomus, Haemasson) grimmi Porchinski, 1876: 32 [<£]; PernTev, 1968: 11. Type(s),
U.S.S.R.: Baku (depository unknown). [A senior synonym which should be suppressed; discussed below.]

Phlebotomus caucasicus Marzinowsky, 1917: 613 [<J]; Popov, 1926: 241; 1935: 108; Marzinowsky &
Shchurenkova, 1929: 67; Adler, Theodor & Lourie, 1930: 531; Isaev, 1935: 103 [cibarium]; PerfiFev,
1935 : 98. Type(s), U.S.S.R. (U, Moscow?).

\_Phlebotomussergenti Parrot; Newstead, 1920: 307 [?]; Nasonov, 1926: 240. Misidentifications.]

Phlebotomus li Popov, 1926: 241; Khodukin, 1929: 92; Sinton, 1928: 309; Shchurenkova, 1929a: 674.
Syntypes 18 $, U.S.S.R.: Armenia (MI, Moscow?). [Synonymized by Marzinowsky & Shchurenkova,
1929:673.]

Phlebotomus grimmi Porchinsky; PernTev, 1937 [status not clear]; 1960: 271 [regarded as a senior synonym
of caucasicus] ; 1968: 7, 11, 236 [description of coxite insufficient for identification; specific independence
first discussed by Nasonov.]

Phlebotomus sergenti var. Hi Popov; Popov, 1928: 33.

Phlebotomus selectus Khodukin, 1929: 99. Syntypes ? <J, U.S.S.R.: Andizhan (US, Tashkent?). [Synony-
mized by PernTev, 1968: 236.]

Phlebotomus (Paraphlebotomus) grimmi Porchinsky; Perfil'ev, 1963: 69; 1968: 1, 7, 49 [larva], 50, 62, 67, 72,
74, 233.

Phlebotomus (Paraphlebotomus) caucasicus Marzinowsky; Theodor, 1958: 19; Theodor & Mesghali, 1964:
287 [identity of grimmi uncertain]; Perfil'ev, 1968: 11; Croset, Abonnenc & Rioux, 1970: 864; Artemiev,
1974: 158; 1978: 18; Lewis, Young, Fairchild & Minter, 1977: 326 [discussion].

$ (extra fact, Iran, Abs Forushan, 5.vii.l969). Leg formula 100, 114, 72; 100, 131, 79; 121, 175, 98 (0-96); tibia
3 = 176 in cJ.

DISTRIBUTION. Afghanistan: Artemiev (1974: 157, map); Nadim et al. (1979: 33, Robatak). China: Xiong
et al. (1964, Xinjiang). Iran: Lewis et al. (1961 : 206); Mesghali (1961 : 25, map); Theodor & Mesghali (1964:
288, species mainly in the north). U.S.S.R.: Beklemishev & Dolmatova (1948: 358, map); Dergacheva (1974:
1668, Karshinskaya steppe); Dergacheva & Zerikhina (1974: 524); Gaibov (1975a: 55, Fergan area); Gre-
belsky (1937: 200, Khiva); Karapet'yan & Babayants (1979: 67, Bayram-Ali); Khodukin (1929, Tashkent);
Latyshev & Posyvai (1937: 184, Serakhs); Marzinowsky & Shchurenkova (1929: 672); Perfil'ev (1968: 89,
236, Andizhan, Baku and Tashkent); Petrishcheva (1937: 148); Ponirovsky (1971: 495, Sumbar Valley);
Rasnitsyna (1974); Zakhar'yants (1958).

THE GENUS PHLEBOTOMUS 145

NOTE. The above citations are selected from the complex taxonomic history of this species.
Artemiev (1976, in litt.) called it caucasicus because, before PerfiTev's (1960) paper, the name
grimmi had been out of use for more than 50 years. During the 49 years from 1925 to 1973 the
Zoological Record cited caucasicus seven times and grimmi not at all. In 1975 and 1976 the Review
of applied Entomology (Series B) did not mention grimmi, but recorded citations of caucasicus by
1 1 authors in 1 1 publications, and two other citations by one of the authors. I propose to request
the ICZN to suppress the name grimmi according to Articles 23 (a-b) and 79 (1973).

This species occurs in moderately hot and dry regions (PerfiTev, 1968: 89), and was considered
to transmit VL in Central Asia and Kazakhstan (Sergiev, 1979: 208). It plays a major part in
disseminating Le. tropica major in the U.S.S.R. (Dubrovsky, 1976: 275; Hoare, 1949: 167; Lewis,
1977: 134; Lysenko, 1971: 516; Sergiev, 1979: 199, 208). Perfil'ev (1968: 139) reported that it
probably transmitted Le. tropica among rodents while the relatively anthropophilic P. papatasi
and P. sergenti passed it on to man. It has been found infected with flagellates in gerbil burrows in
Iran (Nadim & Rashti, 1971 : 100) and may transmit CL among gerbils in Afghanistan (Nadim
etal, 1979:33).

Phlebotomus (Paraphlebotomus) chabaudi Croset, Abonnenc & Rioux

(Map 3)

Phlebotomus (Paraphlebotomus) chabaudi Croset, Abonnenc & Rioux, 1970: 864 [<3]; Rioux, Croset & Guy,
1970: 877; Croset, Leger, Abonnenc & Rioux, 1974: 104[$]; Rioux, Croset & Leger, 1974: 506; Dedet &
Addadi, 1974: 309 [9]; Rioux, Croset, Leger, Benmansur & Soussi, 1975: 497; Croset, Rioux, Maistre &
Bayar, 1978 : 729, 732. Holotype & TUNISIA (IP, Paris).

Phlebotomus chabaudi Croset, Abonnenc & Rioux; Guevara-Benitez, Ubeda-Ontiveros & Morillas-
Marquez, 1978:833.

9. Leg formula after Dedet & Addadi, 100, 128, 80; 93, 143, 83; 114, 183, 108 (0-94).

DISTRIBUTION. Algeria: Dedet (1979, map in litt.); Dedet & Addadi (1974: 308, Biskra; 1977: 86); Rioux
et al. (1970: 877, Ghardaia). Morocco: Rioux et al. (1974: 99, 100, map; 1975: 495, map). Spain: Rioux et al
( 1 9746 : 505, map). Tunisia : Croset et al. ( 1 970 : 872, map ; 1 978 : 727, map, 731).

NOTE. P. chabaudi is suspected of being a vector of CL in North Africa (Croset et al., 1978: 731;
Dedet, 1979:72).

Phlebotomus (Paraphlebotomus) jacusieli Theodor
(Map 3)

Phlebotomus (Paraphlebotomus) jacusieli Theodor, 1947: 95 [£]; 1958: 20 [$]; Theodor & Mesghali, 1964:
288; Croset, Abonnenc & Rioux, 1970: 867; Artemiev, 1974: 158, 161; Artemiev & Dergacheva, 1978: 84
[old records of 'mongolensis' from Transcaucasia wrong, and refer to jacusieli which occurs at Agdam in
Azerbayzhan]. Holotype J, ISRAEL (BMNH).

DISTRIBUTION. Iran: Theodor & Mesghali (1964: 288); Nadim et al. (1977: 215). Israel: Theodor (1947: 95,
Rosh Pinna). Turkey: Yasarol (1980). UJS.S.R.: Artemiev (1974: 160); Artemiev & Dergacheva (1978 : 87).

Phlebotomus (Paraphlebotomus) kazeruni Theodor & Mesghali

(Map 3)

Phlebotomus (Paraphlebotomus) kazeruni Theodor & Mesghali, 1964: 289 [^ (9 = sergenti)] ; Nadim &
Mesghali, 1968: 239 [$]; Croset, Abonnenc & Rioux, 1970: 867; Artemiev, 1974: 158; 1978: 17; Lewis &
Buttiker, 1980. Holotype rf, IRAN (IPH, Tehran).

? (extra fact, Saudi Arabia). Leg formula 100, 117,72; 100, 135,72; 118, 175, 103.

DISTRIBUTION. Afghanistan: Artemiev (1974: 157, map). Iran: Nadim & Mesghali (1968: 240, map); Theodor
& Mesghali (1964: 289, Kazerun). Saudi Arabia: Lewis & Buttiker (1980, map).

146 D. J. LEWIS

NOTE. In Afghanistan P. kazeruni occurs mainly in deserts and low rocky mountains (Artemiev,
1978: 17). In Saudi Arabia it seems common enough to play a part in transmitting Leishmania
among rodents.

Phlebotomus (Paraphlebotomus) marismortui Theodor
(Map 4)

Phlebotomus (Paraphlebotomus) maris-mortui Theodor, 1947: 92 [9 ^]; 1958: 21. Syntypes4 9, 1 £, ISRAEL
(BMNH).

The spelling is emended according to ICZN (1964) Articles 26 (a) and 32.

Phlebotomus (Paraphlebotomus) mongolensis Sinton
(Map 4)

Phlebotomus 'C; Young & Hertig, 1926: 611. CHINA (5 9, 6 $ in BMNH). [Synonymized by Patton &

Hindle, 1926:405.]
Phlebotomus sergenti var.; Patton & Hindle, 1926: 408 [9 £J. CHINA (1 9 in BMNH). [Synonymized by

Theodor & Mesghali, 1964: 290.]
Phlebotomus sergenti var. mongolensis Sinton, 1928: 309 [<£]; Raynal, 1937: 53 [9 c?, first full description].

Type(s), CHINA: North.
Phlebotomus (Paraphlebotomus) mongolensis Sinton; Theodor, 1958: 21; Theodor & Mesghali, 1964: 290

[synonymy]; Perfil'ev, 1968: 63, 67, 72-74, 246; Croset, Abonnenc & Rioux, 1970: 864; Artemiev, 1974:

159; 1978: 18.
Phlebotomus (Paraphlebotomus) imitabilis Artemiev, 1974: 158. Syntypes 29, 2 <J, AFGHANISTAN: 16 km

north of Kabul (MI, Moscow). [Synonymized by Artemiev, 1978: 18.]

The reference to sergenti var. under P. alexandri is relevant.

9 (extrafact, China, 7.ix.l927). Leg formula 100, 114, 68; 100, 131, 77; 124, 169, 97.

DISTRIBUTION. Afghanistan : Artemiev (1974: 157, as imitabilis; 1978: 18). China: Beijing area(1916 or before,
R. A. B.); Patton & Hindle (1926: 409, Xuzhou, = Hsu-chowfu); Raynal (1937: 58, Beijing and Nanjing);
Wang et al. (1963, Gansu Province desert area); Young & Hertig (1926: 611). Iran: Mesghali (1961: 33,
map); Nadim et al. (1977: 215); Theodor & Mesghali (1964: 291). Mongolia: Artemiev (1978: 18, Bayan-
Khongorsk, central and southern Gobi, and Tsagan-Bogdo which is 42°50'N, 50°50'E). UJS.S.R.: Artemiev
(1978: 18, Kazakhstan); Dergacheva (1974: 1668, Karshinskaya steppe); Dergacheva & Turzhanova (1977,
Mangyshlak Peninsula); Dergacheva & Zerikhina (1974: 424); Sorokin (1978, lower River Emba). Many
early records are omitted owing to identification problems.

NOTE. This species seems to be a poor vector (Adler, 1964: 81 ; Garnham, 1974: 225; Molyneux,
1977: 48; Safyanova, 1967: 10) of Le. donovani, but was stated by Sergiev (1979: 208) to be a main
vector of VL in Central Asia and Kazakhstan. It plays a major part in disseminating Le. t. major
in the U.S.S.R. (Dubrovsky, 1976: 275; Lewis, 1977: 134; Sergiev, 1979: 199) where it transmits
the infection among rodents (Bray, 1972: 40; Fischer, 1978: 102). It was found infected with
flagellates in gerbil burrows in Iran (Nadim & Rashti, 1971 : 100).

Phlebotomus (Paraphlebotomus) nuri Lewis
(Map 4)

Phlebotomus (Paraphlebotomus) nuri Lewis, 1967: 15 [£]; 19786: 236; Artemiev, 1974: 160, 161; 1978: 17
[9]. Holotype J, PAKISTAN (BMNH) [examined].

cJ (extrafact, Pakistan, Said Pur, 6.vi.l959). Leg formula 100, 137, 89; 94, 162, 99; 1 11, 196, 1 19 (0-83).

DISTRIBUTION. Southern Afghanistan and southern Iran: Artemiev (1978: 17). Pakistan: Lewis (19786: 325,
map).

NOTE. In Afghanistan P. nuri occurs rarely in southern rocky mountains (Artemiev, 1978 : 18).

THE GENUS PHLEBOTOMUS 147

Phlebotomus (Paraphlebotomus) saevus Parrot
(Map 4)

Phlebotomus sergenti var. saevus Parrot & Martin, 1939: 484 [? ^]. Syntypes 5 9, 1 <$, ETHIOPIA (one in IP,

Algiers; Abonnenc, 1972: 103).
Phlebotomus (Paraphlebotomus) saevus Parrot & Martin; Biittiker & Lewis, 1980 [synonymy].

? (extra fact, Saudi Arabia, Wadi Mizbil, 4.vii.l977). Leg formula 100, 127, 84; 93, 145, 86; 110, 175, 103
(0-95).

DISTRIBUTION. Africa: Abonnenc (1972: 256, map). Ethiopia : Ashford (1974: 607). Kenya: Minter (1964: 207,
map). Saudi Arabia: Lewis & Buttiker (1980).

Phlebotomus (Paraphlebotomus) sergenti Parrot
Phlebotomus sergenti Parrot, 1917: 564.
Represented by two subspecies.

Phlebotomus (Paraphlebotomus) sergenti sergenti Parrot
(Map 4)

Phlebotomus sergenti Parrot, 1917: 564 [£|; Franca, 1918: 731 [$]; Khodukin, 1929: 92; Patton & Evans,
1929: 195, 227; Isaev, 1935: 103 [cibarium]; Abonnenc & Lariviere, 1957 [larva]; Ashford, 1964: 607
[great variation in antenna 3 may indicate two species] ; Guevara-Benitez, Ubeda-Ontiveros & Morillas-
Marquez, 1978: 825, 833. Syntypes <J, ALGERIA (one in IP, Algiers; Abonnenc, 1972: 105).

? Phlebotomus crimicus Shtefko & Minkevich, 1923: 52; Nasonov, 1926: 54 [original depository unknown,
description poor, may be P. sergenti'] ; PernTev, 1968: 253. Syntypes 1 ?, 1 <£ U.S.S.R. (MH, Simferopol?).
[Position doubtful.]

[Phlebotomus caucasicus Marzinowsky; Popov, 1925: 90. Misidentification according to Marzinowsky &
Shchurenkova, 1929: 672.]

Phlebotomus (Phlebotomus) sergenti Parrot; Abonnenc & Lariviere, 1957: 395; Abonnenc, 1972: 105 [table
of differences of$ and^1 from P. alexandri and P. saevus].

Phlebotomus (Paraphlebotomus) sergenti Parrot; PernTev, 1968: 8, 9, 49 [larva], 50, 60 [egg], 67, 72-74, 81,
236; Croset, 1961: 281; Rioux & Golvan, 1969: 72; Biocca, Coluzzi & Constantini, 19776: 162; Lewis,
19786: 236 [synonymy] ; Artemiev, 1978: 16; Croset, Rioux, Maistre & Bayar, 1978: 731, 732.

Phlebotomus (Paraphlebotomus) sergenti sergenti Parrot; Lewis & Buttiker, 1980: 263 [synonymy].

? (extra fact, U.S.S.R., Akhsunskiyi, 1966). Leg formula 100, 1 19, 76; 97, 137, 80; 1 16, 168, 103 (0-94).

DISTRIBUTION. West of Old World: Croset (1969: 283, map). Africa, Mediterranean area etc.: Abonnenc
(1972: 256, map). Orient: Lewis (19786: 325, map). Afghanistan: Arsenieva & Neronov (1978: 32); Artemiev
(1974: 157, map); Nadim et al. (1979: 33, Robatak). Algeria: Dedet (1979, map in litt.); Dedet & Addadi
(1977: 86); Dedet et al. (1977: 256). Crete: Hadjinicolaou (1958: 974, 975); Hertig (1949a: 782, 787-789).
Egypt: Sharqiya Division (1967, M. A. R.); Theodor (1947: 91, Maadi). France: Rioux & Golvan (1969: 82).
Greece: Hadjinicolaou (1958: 968, 970-973); Hertig (1949a: 779, 781); Leger et al. (1979: 17). Iran: Lewis et
al. (1961 : 206); Mesghali (1961: 33, map); Nadim, Mesghali & Javadian (1977: 215); Theodor & Mesghali
(1964: 291, rare in south). Iraq: Ahmad (1976: 86, 98); Pringle (1953: 723, map). Israel: Theodor (1947: 91).
Italy: Biocca et al. (1977a: 161, map; 19776: 20, 29, map). Jordan: Awajan (32°02'N, 36°04'E, 20.X.1978,
E. K. S.). Lebanon: Theodor (1947: 91). Libya: Ashford et al. (1977: 266, Bir Ayyad area). Mali: Ranque et
al. (1975, few near Bamako). Portugal: Franca (1918: 731). Saudi Arabia: Lewis & Buttiker (1980, map).
Somali Republic: Ranque et al. (1975: 4, 1600 m). Southern Yemen: Pringle (1960: 19). Spain: Gil Collado
(1977: 186, map); Guevara-Benitez et al. (1978: 814); Najera (1937: 1489); Zariquiey (1944: 19). Syria:
'Khan el Solol' (1970, caves, wells and pits, K. Z. D.); Theodor (1947: 91, Damascus). Tunisia: Chadli,
Dancescu et al. (1970: 364); Chadli, Romain et al. (1970: 358); Croset (1969: 281, map); Croset et al.
(1978: 735, map). Turkey: Houin et al. (1971: 635); Yasarol (1980). U.S.S.R.: Gaibov (1975a: 55; 19756,
Fergan area; 1976: 491, Surkhandar'ya area); Karapet'yan & Babayants (1979: 67, Bayram Ali). Yemen:
Lewis (1974); Buttiker & Lewis (1979: 370). Yugoslavia : Simic & Zivkovic (1956: 383, Makedonija).

NOTE. P. sergenti is less sensitive to temperature than P. papatasi and can stand colder winters
and extends further north (PernTev, 1968: 80). In Tunisia (Croset et a/., 1978: 734) it is common

148 D. J. LEWIS

both in houses and out of doors, unlike P. papatasi. In Afghanistan (Artemiev, 1978: 17) it is
moderately thermophilic and hydrophilic and bites man readily, usually indoors, and in many
areas is the main vector of Le. t. tropica in numerous villages and towns. In the U.S.S.R. it is the
main vector of anthroponotic CL (Sergiev, 1979: 206). It transmits CL in Crete (Molyneux,
1977: 49), is probably the vector of Le. t. tropica in Iran (Nadim & Rashti, 1971 : 102; Nadim et
al., 1977) and Yugoslavia (Lupascu et al., 1977: 192), and is the main vector in Iraq (Baghdad) and
India (Abonnenc, 1972: 108).

Phlebotomus (Paraphlebotomus) sergenti simitts PernTev
(Map 4)

Phlebotomus (Paraphlebotomus) sergenti similis PernTev, 1963: 75 [? <£); 1968: 239, 251, 252. Syntypes?, <$,
U.S.S.R. (ZI, Leningrad?).

The spermathecae are said to be usually larger than those of P. s. sergenti. The status of this form
needs to be reexamined in view of PernTev's (1968 : viii & 17) remarks on subspecies.

DISTRIBUTION. U.S.S.R.: Dzhavadov et al. (1978: 143, Dzhalilabad and Tazakent, in Azerbaydzhan);
PernTev (1963: Caucasus, Pyatagorsk, southern Crimea and southern Ukraine).

Subgenus SYNPHLEBOTOMUSTheodor

Phlebotomus subgenus Synphlebotomus Theodor, 1948: 97; 1958: 22; PernTev, 1968: 66; Lewis & Ledger,
1976: 406; Lewis, 19786: 236. Type-species : Phlebotomus martini Parrot, 1936, by original designation.

Key to the species of subgenus Synphlebotomus

Females

1 Ventral plates of pharynx with coarse teeth which obscure most of dorsal spiculate armature.

Ascoid on antenna 3 about 0-3 length of segment. India, Iran • eleanorae (p. 149)
Ventral plates of pharynx with armature which does not obscure most of dorsal spiculate arma-
ture • 2

2 Spermatheca with 12 segments. Iran ansarii (p. 149)

Spermatheca with six to 10 segments 3

3 Ascoid on antenna 4 about half length of segment.

Pharynx with few non-spiculate ridges rossi (p. 150)

Ascoid on antenna 4 more than 0-6 length of segment and reaching its tip . ... 4

4 Spermatheca with six segments grovei (p. 149)

Spermatheca with eight or more segments . celiae (p. 149), martini (p. 150), vansomerenae (p. 150)

Males

1 Coxite lobe with about 80 hairs. Iran ansarii (p. 149)

Coxite lobe with about 30 hairs or less

2 Coxite lobe with about 30 filiform hairs of unequal lengths which increase from base of lobe to

apex, the four apical hairs being distinctly longer than the others . . . katangensis (p. 149)
Coxite lobe with not more than 20 hairs 3

3 Coxite lobe with four or five postero-dorsal stout hairs which, in normal curved position, are 1-4

length of style; about nine postero- ventral short narrow ones present.

Paramere with long dorsal row of stout sinuous hairs rossi (p. 150)

Coxite lobe with longest hairs same length as style or less 4

4 Coxite lobe small and bearing about 10 subequal hairs. India, Iran . . . eleanorae (p. 149)
Coxite lobe bearing 1 5-22 subequal hairs 5

5 Coxite lobe with longest hairs same length as style.

Paramere with a few short nearly straight rather thick hairs near tip . . . grovei (p. 149)
Coxite lobe with longest hairs 0-8 length of style or less 6

6 Coxite lobe with seven flat spatulate hairs and some thin ones celiae (p. 149)

Coxite lobe without spatulate hairs ...... . .

7 Coxite lobe with six flat hairs and about 12 thin shorter ones martini (p. 150)

Coxite lobe with 10 flat hairs ending in filiform points, and some thin ones . vansomerenae (p. 150)

THE GENUS PHLEBOTOMUS 149

Phlebotomus (Synphlebotomus) ansarii Lewis
(Map 5)

Phlebotomus (Phlebotomus) ansarii Lewis, 1957: 689 [? £]. Holotype & IRAN (BMNH) [examined].
Phlebotomus (Synphlebotomus) ansarii Lewis; Theodor, 1958: 22; Mesghali, 1961: 37; Lewis & Ledger,
1976: 406; Artemiev, 1978: 25.

$ (extra fact). Leg formula 100, 112,66; 106, 135,76; 122, 169,93(1-67,0-62).
DISTRIBUTION. Iran: Mesghali (1961 : 56, map); Theodor & Mesghali (1964: 291).

NOTE. P. ansarii has been found infected with flagellates in gerbil burrows in Iran (Nadim &
Rashti, 1971 : 100) where it is involved in the transmission of zoonotic CL (Bray, 1972: 40).

Phlebotomus (Synphlebotomus) celiae Minter
(Map 5)

Phlebotomus (Phlebotomus) celiae Minter, 1962: 457 [$ <£]; Abonnenc & Minter, 1965: 32; Abonnenc, 1972:

1 10. Holotype <J, KENYA (BMNH) [examined].
Phlebotomus (Synphlebotomus) celiae Minter; Lewis & Ledger, 1976: 406.

? (extrafact). Leg formula 100, 106, 65; 104, 127, 76; 122, 162, 94 (1-87, 0-67).

NOTE. P. celiae transmits VL in Kenya (Lysenko, 1971 : 517; Mutinga, 1975: 340; Perfil'ev, 1968:
142). It and P. vansomerenae are indistinguishable from P. martini in the female sex, so their roles
are not fully known.

Phlebotomus (Synphlebotomus) eleanorae Sinton
(Map 5)

Phlebotomus eleanorae Sinton, 1931a: 817 [£]; 1933: 418. Holotype & INDIA (BMNH) [examined].
Phlebotomus (Synphlebotomus) eleanorae Sinton; Mesghali, 1965: 267 [$]; Lewis & Ledger, 1976: 405;
Lewis, 19786: 217 [synonymy] ; Artemiev, 1978: 25.

<J (extrafact, holotype). Leg formula 100, 106, 64; 110, 140, 78; 126, 172, 94 (1-68, 0-60).
DISTRIBUTION. India: Sinton (1931 : 817). Iran: Mesghali (1935: 269).

Phlebotomus (Synphlebotomus) grovei Downes
(Map 5)

Phlebotomus (Synphlebotomus) grovei Downes, 1971: 283; [$ <£]; Lewis & Ledger, 1976: 406. Holotype c?,
NAMIBIA (SAIMR, Johannesburg).

$ (extrafact). Leg formula 100, 105, 71, - - , 123, 167, 100 (0-69).
DISTRIBUTION. Namibia: Ledger (1977: 58, map).

Phlebotomus (Synphlebotomus) katangensis Bequaert & Walravens

(Map 5)

Phlebotomus katangensis Bequaert & Walravens, 1930: 35 [£|. Syntypes 2 $, ZAIRE: Lubumbashi ( =

Elizabethville) (MRAC, Tervuren).
Phlebotomus (Synphlebotomus) katangensis Bequaert & Walravens; Lewis & Ledger, 1976: 407 [synonymy

& citations].

P. rossi was treated as a synonym of P. katangensis by some authors for several years.

150 D. J. LEWIS

Phlebotomm (SynpMebotomus) martini Parrot
(Map 5)

Phlebotomus (Phlebotomus) martini Parrot, 1936: 35 [9 <£]. Syntypes 9 9, 2 $, ETHIOPIA (one in IP, Algiers;

Abonnenc, 1972: 117).
Phlebotomus (SynpMebotomus) martini Parrot; Lewis & Ledger, 1976: 406 [synonymy].

? (extra fact, Kenya). Leg formula 100, 116,72; 102, 135,75; 117, 157,84(0-68).

DISTRIBUTION. Africa: Abonnenc (1972: 117). Kenya: Kangondi (termite hill record from D. M. M., 1980);
Minter (1964: 207, map); Wijers & Ngoka (1974: 26). Sudan: Lewis & Kirk (1954: 35, map); Qutubuddin
(1962: 594). Uganda: Wykoff et al. (1969).

NOTE. The work of Minter and colleagues showed that P. martini transmits VL in Kenya (Adler,
1964:87; Bray, 1972:40; Diesfeld, 1978:50; Manson-Bahr, 1971:434; Perfil'ev, 1968: 142;
Southgate, 1977: 245; Wilcocks & Manson-Bahr, 1972: 122). It is uncommon in western Tharaka
but may be the vector there (Mutinga & Ngoka, 1975; Wijers & Ngoka, 1974: 29). It may
transmit the disease in south-west Ethiopia (Fuller et al., 1979: 429).

Phlebotomus (SynpMebotomus) rossi De Meillon & Lavoipierre

(Map 5)

Phlebotomus rossi De Meillon & Lavoipierre, 1944: 44 [<J]; Parrot, 1957: 49 [proposed as synonym of P.

katangensis]. Holotype <J, ZIMBABWE (SAIMR, Johannesburg).
Phlebotomus (Phlebotomus) rossi De Meillon & Lavoipierre; Minter, 1962: 459; Abonnenc, 1967: 4;

1972: 112.
Phlebotomus (Synphlebotomus) rossi De Meillon & Lavoipierre; Mesghali, 1965: 269; Downes, 1971: 284;

Lewis & Ledger, 1976: 407 [9, reinstated as species ; synonymy] ; Ledger, 1977: 578.

9 (extra fact, Namibia, Sandmodder, 18.iv.1975). Leg formula 100, 113, 66; 105, 137, 75; 120, 170, 93 (2-39,
0-28).

DISTRIBUTION. Southern Africa: Lewis & Ledger (1976: 410). Namibia: Ledger (1977: 582, map).

NOTE. In Namibia P. rossi lives in damp hyrax burrows and has been found infected with
flagellates in an area of CL (Ledger, 1977: 579).

Phlebotomus (Synphlebotomus) vansomerenae Heisch, Guggisberg & Teesdale

(Map 5)

Phlebotomus (Phlebotomus) vansomerenae Heisch, Guggisberg & Teesdale, 1956: 211 [9 <$]. Holotype 9,

KENYA (BMNH) [examined].
Phlebotomus (Synphlebotomus) vansomerenae Heisch, Guggisberg & Teesdale; Lewis & Ledger, 1976: 406

[synonymy].

DISTRIBUTION. Kenya: Minter (1964: 207); Wijers & Ngoka (1974: 26).

NOTE. This species may transmit kala-azar in western Tharaka (Wijers & Ngoka, 1974: 28).

Subgenus L^/?/?OC/55/l/5Nitzulescu

Phlebotomus subgenus Larroussius Nitzulescu, 1931: 274; Theodor, 1948: 97; 1958: 22; Perfil'ev, 1968: 63,
73, 82, 250, 252; Lewis, Minter & Ashford, 1974: 435; Biocca, Coluzzi & Constantini, 1977: 162 [trochan-
ter spines in two groups in four species]; Lewis, 19786: 237; Artemiev, 1980: 1181. Type-species: Phleb-
otomus major Annandale, 1910, by original designation.

Key to the species and subspecies of subgenus Larroussius

Females

1 Spermatheca with indistinct segmentation and no neck. Pharynx with large irregular teeth . 2
Spermatheca with distinct segmentation and neck. Pharynx with punctiform teeth ... 3

2 Palpal segments 2 and 3 each 0-1 8-0- 20 mm long mascittii mascittii (p. 158)

THE GENUS PHLEBOTOMUS 151

Palpal segments 2 and 3 each 0-22-0-24 mm long .... mascittii canaaniticus (p. 158)

3 Spermatheca with neck about as long as head somaliensis (p. 162)

Spermatheca with neck much longer than head 4

4 Spermathecal ducts bag-like for most of their length gibiensis(p. 154)

Spermathecal ducts mainly tubular 5

5 Pharyngeal teeth not very fine, in regular transverse rows, occupying nearly half the pharynx.

Spermatheca with about 15 segments. Iran and Turkestan wenyoni (p. 163)

Pharyngeal teeth very fine and punctiform 6

6 Spermatheca very long with 30^35 segments.

Pharyngeal teeth bigger in middle than at sides

kandelakii burneyi (p. 154), kandelakii kandelakti (p. 154)
Spermatheca with 8-22 segments ............ 7

7 Spermatheca with 18-22 segments 8

Spermatheca with 8- 16 segments 9

8 Spermatheca with about 22 segments, long neck and a very small head. Pharynx with scarcely

visible spicules. West Malaysia betisi (p. 153)

Spermatheca with 18-21 segments, broad at end and narrowing towards base, end process (neck
and head) relatively short, about three times as long as wide. Hind half of pharynx with fine
punctiform teeth which become scale-like anteriorly keshishiani (p. 155)

9 Outer 0-43 of Spermathecal ducts sac-like.

Pharyngeal armature occupying 0-15-0-17 length of pharynx. Antenna 3 = 0-38-0-42 mm

long ariasi (p. 153)

Spermathecal ducts not like this 10

10 Wing length usually 1-8-2-5 mm 11

Wing length usually 2-8-3-6 mm 13

11 Spermatheca with 10 (8-12) segments . longicuspis (p. 155), orientals (p. \59), perniciosus (p. 161)
Spermatheca with 12- 16 segments 12

12 Palpal formula 1, 4, 2, 3, 5 . . . perfiliem perfiliem (p. 160), perfiliem transcaucasicus (p. 161)
Palpal formula 1, 2, 4, 3, 5 tobbi (p. 162)

13 Armature occupying half length of pharynx major krimensis(p. 156)

Armature occupying about a third to a quarter length of pharynx 14

14 Armature occupying about hind third or more of pharynx, fore teeth scale-like with secondary

spicules 15

Armature occupying about hind quarter of pharynx 17

15 Armature occupying hind third of pharynx. Palpal formula 1,4, (2, 3), 5 . . major major (p. 157)
Armature of pharynx extending further forward. Palpal formula 1,4, 2, 3, 5 .... 16

16 Wing about 3-5 mm long major neglectus (p. 157)

Wing about 2 mm long major syriacus (p. 157)

17 Hind teeth of pharyngeal armature relatively large guggisbergi (p. 154)

Hind teeth of pharyngeal armature not relatively large 18

18 Most of pharyngeal armature comprising rows of fused denticles . . . . smirnovi (p. 162)
Most of pharyngeal armature comprising distinct denticles

aculeatus (p. 153), longipes (p. 156), pedifer (p. 159)

Males

1 Aedeagus with distal long slightly-curved transparent process bearing fine dorsal teeth . . 2
Aedeagus otherwise ............... 4

2 Transparent part of aedeagus narrow and nearly straight, aedeagus 0-16 mm long

perfiliem transcaucasicus (p. 161)
Transparent part of aedeagus narrow and curved, or broad 3

3 Two ascoids on antenna 3-15. Aedeagus 0-13-0-14 mm long, transparent process short and

wide with four or five teeth concentrated near tip . . . . perfiliem perfiliem (p. 160)
Two ascoids on antenna 3-15. Aedeagus 0-17-0-19 mm long, transparent process long and
narrow with teeth evenly spread perfiliewi galilaeus (p. 160)

4 Aedeagus with ventral teeth in the middle, and narrowing gradually to a point .... 5
Aedeagus with smooth sides and no ventral teeth 6

5 Two ascoids on antenna 3-5 kandelakii kandelakii (p. 154)

Two ascoids on antenna 3-7 kandelakii burneyi (p. 154)

6 Aedeagus with rounded end 7

Aedeagus with sharp tip and nearly parallel sides 18

152 D. J. LEWIS

7 Aedeagus with marked distal bulge thickest at 0-84, distinctly shorter than paramere, with

sperm tubes emerging at tip 8

Aedeagus with bulge nearer base or with none or virtually none 9

8 Coxite 0-33 mm long, 1-9 length of aedeagus, with 16-32 hairs in tuft. Aedeagus clapper-like,

with moderate subapical expansion ariasi (p. 153)

Coxite 0-48 mm long, three times length of aedeagus, with 37-78 hairs in group. Aedeagus
sword-like, with thick subapical expansion and nearly pointed tip .... chadlii(p. 153)

9 Aedeagus gradually thickening from each end till 0-6. Afrotropical • . . gibiensis (p. 154)
Aedeagus without marked local thickening 10

10 Length of aedeagus 7-8-1 1-0 middle thickness ll

Length of aedeagus 13-26 middle thickness . 14

11 Coxite about 6-9 as long as thick. Afrotropical . . . . . . . fantalensis (p. 154)

Coxite about 4-6 or less as long as thick. Palaearctic 12

12 Paramere with inconspicuous mid-ventral row of about four spines . . . smirnovi (p. 162)
Paramere with conspicuous mid- ventral row of about ten spines 13

13 Style rather longer than half length of coxite. Surstyle longer than coxite. Antenna 3 = 0-45 mm

long, 1-55 length of labrum mascittii mascittii (p. 158)

Style less than half length of coxite. Surstyle not longer than coxite. Antenna 3 = 0-37-0-40 mm
long, 1-3-1 -5 length of labrum mascittii canaaniticus (p. 158)

14 Aedeagus clearly shorter than paramere, sperm tubes emerging from tip 15

Aedeagus nearly as long as paramere, sperm tubes emerging before tip.

Two ascoids on antenna 3-8 16

15 Aedeagus narrowing towards tip. Coxite hair-group with 18-30 hairs.

Sperm tubes 6- 11 times length of pump keshishiani (p. 155)

Aedeagus with scarcely visible subapical swelling. Coxite hair-group with 25-30 hairs mariae (p. 158)

1 6 Teeth occupying nearly half length of pharynx, relatively large and in regular transverse rows

wenyoni (p. 163)
Teeth occupying only a third of length of pharynx, fine and punctiform.

Tip of aedeagus like drum-stick 17

17 Coxite 0-33-0-35 mm long, style 0-16-0- 19 mm long .... major neglectus (p. 157)
Coxite 0-40-0-45 mm long, style 0-20-0-22 mm long 18

18 Palpal formula 1, 4, (2, 3), 5 . major major (p. 157)

Palpal formula 1, 4, 2, 3, 5 - . . . . . 19

19 Coxite hair-group with more than 20 widely spaced hairs. Style less than half length of coxite

major krimensis (p. 156)
Coxite hair-group with about 30 densely packed hairs. Style half as long as coxite

major syriacus (p. 157)

20 Aedeagus bifid 21

Aedeagus not bifid 22

21 Both points of aedeagus the same size and sharp. Sperm pump 0-14-0-15 mm long perniciosus (p. 161)
Distal point of aedeagus distinctly longer than the other and rounded. Sperm pump 0-05-

0-18 mm long.

Both middle spines of style nearer basal spine than in perniciosus. Style longer than coxite.
Aedeagus narrowing more markedly from the base, and more conical . . . tobbi (p. 162)

22 Hair group of coxite on low pad ......... guggisbergi (p. 154)

Hair group of coxite not on pad 23

23 Aedeagus with one to three subterminal spicules aculeatus (p. 153)

Aedeagus without subterminal spicules ........... 24

24 Tip of aedeagus tapering gradually, far from that of paramere except in P. longicuspis . . 25
Tip of aedeagus tapering abruptly, near that of paramere 27

25 Tip of aedeagus curving down ward. North Africa longicuspis (p. 155)

Tip of aedeagus curving upward. North-east tropical Africa 26

26 Tip of aedeagus curving slightly upward ........ longipes (p. 156)

Tip of aedeagus curving sharply upward pedifer (p. 159)

27 Antenna 3-1 2 with two ascoids. Tip of aedeagus ventral and mesad . . . langeroni (p. 155)
Antenna 3-7 with two ascoids. Tip of aedeagus dorsal and laterad . . . orientalis (p. 159)

THE GENUS PHLEBOTOMUS 153

Phlebotomus (Larroussius) aculeatus Lewis, Minter & Ashford

(Map 6)

Phlebotomus species C; Ashford, 1974: 610 [9].

Phlebotomus (Larroussius) aculeatus Lewis, Minter & Ashford, 1974: 437 [9 <$\ differences between Ethio-
pian and Kenyan forms noted]. Holotype <^, KENYA (BMNH) [examined].

Phlebotomus (Larroussius) elgonensis Ngoka, Madel & Mutinga, 1975: 132. Holotype <$, KENYA (NM,
Nairobi). Syn. n.

NOTE. The descriptions of aculeatus and elgonensis tally fairly well but there are a few discrep-
ancies in the description of the latter. The palp formulae of both sexes do not agree with the
lengths of segments given; and in the paratype male, which the authors kindly placed in the
BMNH, there are two ascoids on antenna 3-7 only, and R2/2 + 3 is 2-3 and not 0-88. The two
appear to be the same species.

DISTRIBUTION. Ethiopia and Kenya: Lewis et al. (1974: 439). Kenya: Ngoka et al. (1975: 132, 136, three caves
cited for holotype of P. elgonensis).

Phlebotomus (Larroussius) ariasi Tonnoir
(Map 6)

Phlebotomus ariasi Tonnoir, 19216: 53 [^] ; Nitzulescu, 193(W: 531 ; Raynal & Le Gac, 1933: 652 [9; Parrot,
1934: 386; 1936: 48; Najera, 1936: 309; Zariquiey, 1937: 410; Rageau & Colas-Belcour, 1956: 235;
Abonnenc & Lariviere, 1957: 392 [larva]; Rioux, Abonnenc & Bauduoy, 1965: 615; Croset, 1969: 349;
Bailly-Choumara, Abonnenc & Pastre, 1971: 436; Biocca, Coluzzi & Constantini, 1977 a: 162; Croset,
Rioux, Maistre & Bayar, 1978: 734;Guevara-Benitez, Ubeda-Ontiveros & Morillas-Marquez, 1978: 817,
832. Holotype <$, SPAIN (depository unknown, believed lost).

Phlebotomus (Larroussius) ariasi Tonnoir; Theodor, 1958: 23; Juminer & Gibily, 1966: 86; Rioux & Golvan,
1969: 22, 88 [review]; Dedet & Dib, 1972: 56; Rioux, Croset, Leger & Benmansur, 1974: 96; Rioux,
Croset, Leger, Benmansur & Soussi, 1975: 499; Biocca, Coluzzi & Constantini, 1977a: 160-162; 19776:
30; Croset, Rioux, Maistre & Bayar, 1978: 734.

9. Leg formula, after Zariquiey. 100, 133, 86; 138, 149, 104; 107, 183, 116 (0-98).

DISTRIBUTION. Western Mediterranean: Croset (1969: 357, map); Rioux & Golvan (1969: 89, 99, map).
Algeria: Dedet (1979, map in litt); Dedet & Addadi (1977: 86); Dedet, Addadi & Lannuzel (1977: 256);
Parrot (1936: 48); Parrot & Clastrier (1939: 633). France: Croset (1969: 353, map); Dedet & Dib (1973: 55,
57-61, map); Houin et al. (1977: 113, map, 114, Juigne-sur-Sarthe, 47°52'N, 0°17'W); Rioux & Golvan
(1969: 51, 89, 100, map). Italy: Biocca et al. (1977a: 160, map; 19776: 20, 28, map); Rioux et al. (1964: 966).
Morocco: Bailly-Choumara et al. (1971 : 453, map). Portugal: Meira & Ferreira (1944: 274, map). Spain: Gil
Collado (1977: 186, map); Guevara-Benitez et al. (1978: 815); Najera (1937: 1488); Rioux & Golvan (1969:
100, map); Zariquiey (1944: 19). Tunisia: Croset (1969: 355, map); Croset et al. (1966: 549, map; 1978: 733,
map); Rioux et al. (1966: 88, map, rare in humid areas; 1974: 505, map).

NOTE. P. ariasi has been readily infected with the parasites of VL (Molyneux, 1977: 48) and is the
vector of Le. donovani in the south of France (Killick-Kendrick, 1978: 301; Lanotte et al., 1977:
126; Molyneux, 1977: 48; Rioux et al., 1977: 299, 303 ; Zuckerman & Lainson, 1977: 73).

Phlebotomus (Larroussius) betisi Lewis & Wharton
(Map 6)

Phlebotomus (Larroussius) betisi Lewis & Wharton, 1963: 117 [?]; Lewis, 19786: 237. Holotype 9, WEST
MALAYSIA (BMNH) [examined].

DISTRIBUTION. West Malaysia : (Lewis, 19786: 237, map).

Phlebotomus (Larroussius) chadlii Rioux, Juminer & Gibily
(Map 6)

Phlebotomus (Larroussius) chadlii Rioux, Juminer & Gibily, 1966: 83 [<£]; Croset, Rioux, Juminer & Tour,
1966: 547; Croset, 1969: 342; Rioux & Golvan, 1969: 96; Rioux, Guy, Corroller, Croset & Addadi, 1970:

154 D. J. LEWIS

101 [$ unknown] ; Rioux, Croset, Leger & Bailly-Choumara, 1974: 96, 97; Rioux, Croset, Leger, Benman-
sur & Soussi, 1975: 498; Croset, Rioux, Maistre & Bayar, 1978: 735 [? unknown]. Holotype, TUNISIA
(EM, Montpellier).

cJ. Leg formula, Tunisia, after Rioux et al. (1966: 85). 100, 142, 96; 88, 162, 100; 104, 200, 125 (1-20).

DISTRIBUTION. Algeria: Dedet (1979, in litt); Dedet & Addadi (1977: 86); Dedet et al. (1977: 256). Morocco:
Rioux et al. (1974: 99, 100; 1975: 495, map). Tunisia : Croset et al. (1978: 733, map); Dedet (1971 : 157); Rioux
et al. (1966: 83, 88, in arid areas).

Phlebotomus (Larroussius) fantalensis Lewis, Minter & Ashford

(Map 6)

Phlebotomus species B; Ashford, 1974: 610 [£j.

Phlebotomus (Larroussius) fantalensis Lewis, Minter & Ashford, 1974: 439 [£]. Holotype c?, ETHIOPIA
(BMNH) [examined].

Phlebotomus (Larroussius) gibiensis Lewis, Minter & Ashford

(Map 6)

Phlebotomus species A; Ashford, 1974: 610.

Phlebotomus (Larroussius) gibiensis Lewis, Minter & Ashford, 1974: 439 [? $]. Holotype J, ETHIOPIA
(BMNH) [examined].

Phlebotomus (Larroussius) guggisbergi Kirk & Lewis
(Map 6)

Phlebotomus (Synphlebotomus) guggisbergi Kirk & Lewis, 1952: 339, 340 [$ £]. Lectotype & KENYA

(BMNH), designated by Lewis, Minter & Ashford, 1974: 440 [examined].
Phlebotomus guggisbergi Kirk & Lewis; Lewis & Minter, 1960: 352.
Phlebotomus (Phlebotomus) guggisbergi Kirk & Lewis; Abonnenc & Minter, 1965: 31; Abonnenc, 1972 [in

'Synphlebotomus' group].
Phlebotomus (Larroussius) guggisbergi Kirk & Lewis; Lewis, Minter & Ashford, 1974: 440.

DISTRIBUTION. Kenya: Minter (1964: 207, map; 1966: 180, map). Tanzania: Minter (1964: 208). Uganda:
Kidepo Park (ix.1969, received from D. M. M. in 1980).

NOTE. This very large species is found in caves and among trees and bites man (Abonnenc, 1972:
112).

Phlebotomus (Larroussius) kandelakii Shchurenkova
Phlebotomus kandelakii Shchurenkova, 19296: 693 ; Perfil'ev, 1968: 78, 261.

Phlebotomus (Larroussius) kandelakii burneyi Lewis
(Map 6)

Phlebotomus (Larroussius) kandelakii burneyi Lewis, 1967: 17 [? £]; 19786: 238; Artemiev, 1974: 160.
Holotype <£ PAKISTAN (BMNH) [examined].

DISTRIBUTION. Pakistan: Lewis (19786: 238, map).

Phlebotomus (Larroussius) kandelakii kandelakii Shchurenkova

(Map 6)

Phlebotomus sp. n.; Shchurenkova, Demina & Pavlova, 1929: 681, 684, 686, 688.

Phlebotomus kandelakii Shchurenkova, 19296: 693 [9 £1; Adler, Theodor & Lourie, 1930: 536. Syntypes?
cJ, U.S.S.R. (TI, Tbilsi).

THE GENUS PHLEBOTOMUS 155

Phlebotomus (Larroussius) kandelakii Shchurenkova; Theodor, 1958: 23; Perfil'ev, 1968: 261; Lewis, 1978:
237; Artemiev, 1978:19.

DISTRIBUTION. Central Asia: Dolmatova (1962: 461, map); Dolmatova & Demina (1971 : 120, map). Afghani-
stan: Artemiev (1974: 157, map; 1978: 19). Iran: Mesghali (1961: 47, map); Nadim & Rashti (1978: 27,
Chahar Mahal); Theodor & Mesghali (1964: 291, only in north). Lebanon: Theodor & Mesghali (1964: 291).
Turkey: Artemiev (1978: 19); Yasarol (1980). U.S.S.R.: Gaibov (19756, Fergana area); Petrishcheva (1937:
148).

NOTE. In Afghanistan P. kandelakii is very hydrophilic and moderately thermophilic and bites
man and large animals readily (Artemiev, 1978: 19). It appears to be a vector of Le. donovani in
Georgia (Maruashvili, 1958: 595; PernTev, 1968: 142), and was considered to be a main vector of
VL in Transcaucasia (Sergiev, 1979: 208).

Phlebotomus (Larroussius) keshishiani Shchurenkova
(Map 6)

Phlebotomus keshishiani Shchurenkova, 1936: 892 [? <J]. Syntypes? <J, U.S.S.R. (TI, Dushanbe).
Phlebotomus (Larroussius) keshishiani Shchurenkova; Theodor & Mesghali, 1964: 291 [had probably often

been confused with major and wenyonf] ; Lewis, 1967: 19; 1978: 238; Perfil'ev, 1968: 274; Artemiev, 1978:

19.

<J (extra facts, Pakistan, Said Pur, 6.vi.l965). Leg formula 100, 161, 113; 87, 181, 118; 108, 214, 141 (2-69,
1-09); tibia 3 = 2-65 mm long.

DISTRIBUTION. Afghanistan: Artemiev (1974: 157, map; 1978: 15). Iran: Nadim & Rashti (1978: 271, general
map, 277); Nadim et al. (1977: 215). Pakistan: Lewis (1978ft: 238, map). U.S.S.R.: Gaibov (19756, Fergana
area; 1976: 49, Surkhandar'ya); Perfil'ev (1968: 277); Petrishcheva (1935: 206).

NOTE. In the U.S.S.R. this species is numerous between 1900 and 2300 m (Perfil'ev, 1968: 277),
and in Afghanistan ranges from 1000 to 2800 m and will bite man (Artemiev, 1978: 19).

Phlebotomus (Larroussius) langeroni Nitzulescu
(Map 7)

Phlebotomus perniciosus var. ; Nitzulescu, 1930c: 382 [<£].

Phlebotomus langeroni Nitzulescu, 1930e: 548 [<£]. Holotype <£, TUNISIA (FM, Paris?).

Phlebotomus (Phlebotomus) langeroni Nitzulescu; Parrot, 1940: 310.

Phlebotomus (Larroussius) langeroni Nitzulescu; Theodor, 1958: 24; Perfil'ev, 1968: 77; Croset, 1969: 311 [$

apparently unknown]; Bailly-Choumara, Abonnenc & Pastre, 1971 : 437; Rioux, Croset, Leger & Bailly-

Choumara, 1974: 96; Croset, Rioux, Maistre & Bayar, 1978: 735.

DISTRIBUTION. North Africa: Croset (1969: 313, 316, maps, evidently very rare). Tunisia: Chadli et al. (1970:
358; 1970: 363; 1978: 733, map).

NOTE. The unknown female (Theodor, 1958: 24) is probably difficult to distinguish from P.
longicuspis and perniciosus. The species seems to be rare in Tunisia (Croset et al., 1978: 736).

Phlebotomus (Larroussius) longicuspis Nitzulescu
(Map 7)

Phlebotomus perniciosus var. ; Nitzulescu, 1930c: 384 [in part, one^].

Phlebotomus langeroni var. longicuspis Nitzulescu, 1930e: 551 [£|; Ristorcelli, 1941: 372. Syntypes cJ,

TUNISIA (FM, Paris?).

Phlebotomus longicuspis Nitzulescu; Parrot, 1936: 138 [$] [raised to species].
Phlebotomus (Larroussius) longicuspis Nitzulescu; Perfil'ev, 1968: 77; Croset, 1969: 312; Bailly-Choumara,

Abonnenc & Pastre, 1971: 436; Rioux, Croset, Leger & Bailly-Choumara, 1974: 96; Croset, Rioux,

Maistre & Bayar, 1978: 736.

156 D. J. LEWIS

DISTRIBUTION. North Africa: Croset (1969: 321, 322, map). Algeria: Dedet (1979, map in litt.); Dedet et al.
(1975: 185; 1977: 276). Libya: Ashford et al. (1977: 265, Bir Ayyad area, south of Surman). Morocco:
Bailly-Choumara et al. (1971: 453); Rioux et al. (1974: 99). Tunisia: Chadli, Dancescu et al. (1970: 363);
Chadli, Romain et al. (1970: 358); Croset et al. (1978: 737, map).

NOTE. This species has been found infected with Le. d. infantum in Algeria (Dedet, 1979: 58;
Theodor, 1964: 480) and regarded as a VL vector in North Africa (Abonnenc, 1972: 34; Hoog-
straal & Heyneman, 1969: 1185, 1186; Wilcocks & Manson-Bahr, 1972: 121) but in Tunisia
seems too rare to be important (Croset et al., 1978 : 736).

Phlebotomus (Larroussius) longipes Parrot & Martin

(Map 7)

Phlebotomus (Phlebotomus) longipes Parrot & Martin, 1939: 143 [? £|; Kirk & Lewis, 1946b: 119; 1951:
434; Parrot, 1940: 316; 1953: 113 [papillae]; Minter, 1964: 209 [variation]; Abonnenc & Minter, 1965:
32; Abonnenc, 1972: 115. Syntypes 642 $, 896 J, ETHIOPIA (one $, labelled 'type' in Parrot's writing, in
BMNH).

Phlebotomus longipes Parrot & Martin; Kirk & Lewis, 1947: 873 ; Abonnenc & Lariviere, 1957: 399 [larva];
Ashford, 1974: 610; Gemetchu, 1974: 114.

Phlebotomus (Larroussius) longipes Parrot & Martin; Lewis, Mutinga & Ashford, 1972: 119.

$ (extra fact, Ethiopia, Addis Ababa). Leg formula 100, 129, 81; 91, 148, 89; 102, 174, 108 (3-42, 1-26); tibia
3 = 2-19 mm. The length of the legs is evidently due to the size of the insect.

DISTRIBUTION. Africa : Abonnenc (1972: 259, map). Ethiopia : Ashford (1974: 607). Kenya: Minter (1964: 209,
map).

NOTE. This species transmits CL in Ethiopia (Ashford, 1977: 236; Schaller, 1972: 102; White,
1977: 163; Wilcocks & Manson-Bahr, 1972: 135).

Phlebotomus (Larroussius) major Annandale

Phlebotomus major Annandale, 1910: 46.

Phlebotomus (Larroussius) major Annandale; Perfil'ev, 1968: vii, 7, 49 [larva], 50, 54, 62, 75, 85, 253.

Some early records of P. major refer to other species described later. After their recognition
Theodor & Mesghali (1964: 291) referred to P. major, generally found in mountainous country, as
an eastern Mediterranean species occurring from Italy to north-west India where it was found
mainly in the western Himalayas. Perfil'ev (1968: 94, 261) discussed its distribution and doubted
the truth of Central Asian records.

Theodor (1958) recognized three subspecies, major in India, neglectus in Dalmatia and Italy,
and syriacus in the Mediterranean and Caucasus. The following are records of P. major s. 1.

Afghanistan: Artemiev(1978: 19). Crete: Hadjinicolaou ( 1958: 974); Hertig(1949a: 782, 787-789). Greece:
Hadjinicolaou (1958: 968, 970, 972, 973); Hertig (1949a: 779, 781-786). India: Theodor (1958: 25). Iran:
Nadim et al. (1977: 215; 1978: 26-28, maps; Chahar Mahal and other areas without details); Theodor &
Mesghali (1964: 291, few at Ramadan, Kazerun and Yazd; some early records may refer to P. tobbi). Italy:
Biocca et al. (1977a: 162, map; 19776: 20, 28, map). Corradetti et al. (1956a: 6, map); Puccini et al. (1977: 38,
map). Rumania: Duport et al. (1971: 394). Turkey: Yasarol (1980). U.S.S.R.: Petrishcheva (1937: 148).
Yugoslavia: Simic & Zivkovic (1956: 383-385, north and south Dalmatia, Hercegovina, Istra, Kosovo i
Metohija, Makedonija, Montenegro, Serbia and Vojvodina).

Phlebotomus (Larroussius) major krimensis Perfil'ev
(Map 7)

Phlebotomus (Larroussius) major krimensis Perfil'ev, 1966: 282 [? £|; 1968: 226, 254, 256, 258, 259. Syntypes

$ cJ, U.S.S.R. (ZI, Leningrad?).
? Phlebotomus perniciosus var. tauricus Nasonov, 1927: 369 [$]. No type, U.S.S.R.: Crimea. Listed by

Perfil'ev (1968: 253) under P. major. [Position doubtful.]

THE GENUS PHLEBOTOMUS 157

DISTRIBUTION. U.S.S.R.: Perfil'ev (1968: 255, 259, 261, Crimean subspecies).

NOTE. The names tauricus and tauriae ( = longiductus) appear to have nothing to do with bulls or
the Taurus Mountains but to be derived from Tauri, the name of an ancient Crimean tribe.

This form tends to remain in houses by day (Perfil'ev, 1968: 110) but not as much as P.
papatasi.

Phlebotomus (Larroussius) major major Annandale
(Map 7)

Phlebotomus major Annandale, 1910: 46 [<J]; Summers, 1911: 110; Sinton, 1925: 107 [?]; 1928: 303 [in

part]. Lectotype 9, INDIA (ZSI, Calcutta), designated by Quate, 1962a: 157.
[Phlebotomus major var. perniciosus Newstead [in part]; Brunetti, 1912: 201. Synonymized by Perfil'ev,

1968:253.]
Phlebotomus (Larroussius) major Annandale; Theodor & Mesghali, 1964: 291; Perfil'ev, 1968: 253, 260 [in

part].
Phlebotomus (Larroussius) major major Annandale; Lewis, 1978fo: 238 [synonymy, including synonym gris-

eus of which the type seems to be lost (Quate, 1962a: 157)].

DISTRIBUTION. India, Nepal and Pakistan: Lewis (19786: 239).

Phlebotomus (Larroussius) major neglectus Tonnoir

(Map 7)

Phlebotomus neglectus Tonnoir, 1921a : 333. Syntypes 12 9, 3 <J, ALBANIA, YUGOSLAVIA, ITALY (NM, Vienna).
Phlebotomus (Larroussius) major neglectus Tonnoir; Theodor, 1958: 25; Perfil'ev, 1968: 254.

DISTRIBUTION. Albania, Italy & Yugoslavia : Tonnoir (1921 : 333). Italy & Dalmatia: Theodor (1958: 25).

NOTE. P. major is considered to be a vector of VL in the western Mediterranean (Theodor, 1964:
480) and Yugoslavia (Lupascu et al., 1977: 192).

Phlebotomus (Larroussius) major syriacus Adler & Theodor
(Map 7)

Phlebotomus major Annandale [in part]; Adler & Theodor, 1929: 275 [9 £); Adler, 1946: 501 ; Cristescu &
Dancescu, 1967: 320; Rioux & Golvan, 1969: 93; Rioux, Croset, Leger & Bailly-Choumara, 1974: 96.

Phlebotomus major var. syriacus Adler & Theodor, 1931: 467; Adler, 1946: 500 [9]. Type(s), PALESTINE,
SYRIA (depository unknown).

Phlebotomus (Larroussius) major syriacus Adler & Theodor; Theodor, 1958: 25 [cJ]; Perfil'ev, 1968: 254;
Houin, Abonnenc & Deniau, 1971 : 644; Leger et al., 1974: 20.

Phlebotomus (Larroussius) major Annandale [in part]; Biocca, Coluzzi & Constantini, 1977a: 160-162.

DISTRIBUTION. Mediterranean, Caucasus, Crete, Crimea, Greece, Israel and Syria: Theodor (1958: 25).
Greece: Leger et al. (1979: 17). Jordan: Perfil'ev (1968: 255). Turkey: Houin et al. (1971: 644). U.S.S.R.:
Izmail and Transcaucasia probably, Perfil'ev (1968: 255).

NOTE. In Greece, as in the Cevennes, Corsica and Serbia, P. major is abundant only above 300 m
(Leger et al, 1979: 20). It is evidently a good vector of VL (Molyneux, 1977: 48) and is a vector in
the eastern Mediterranean area (Hoogstraal & Heyneman, 1969: 1185; Theodor, 1964: 480;
Wilcocks & Manson-Bahr, 1972: 121). It is a main vector in Crete (Leger et al, 1979: 20) and
Greece (Adler, 1964: 80; Lupascu et al, 1977: 192; Perfil'ev, 1968: 142), and may transmit VL in
southern Italy and Sicily (Biocca et al, 1917 a: 165).

Phlebotomus (Larroussius) major wui Yang & Xiong
(Map 7, type area)

\_Phlebotomus major Annandale; Ding & He, 1962: 388. Misidentification.]

Phlebotomus major wui Yang & Xiong, 1965: 412 [9 <£]. Syntypes 9 c?, CHINA: north and south Xinjiang
(Institute of Parasitic Diseases, Chinese Academy of Medical Sciences, Shanghai).

158 D. J. LEWIS

DISTRIBUTION. China: Artux, Dunhuang, Ha-mi, Kashi, Tacheng and Yning (Yang & Xiong, 1965); desert
areas of Xinjiang and of Ejina County in Inner Mongolia (Wu et al., 1979).

NOTE. This is probably P. smirnovi according to Professor Leng Y.-j. (1981, pers. comm.), and is a
desert zoophilic form in south Xinjiang (Xiong et al., 1970). The following are among features
reported by Wu et al. (1979). In Xinjiang the form occurs from early May to late September, with
population peaks in June and August; in the Karamay Desert it rests in gerbil burrows, and in
the Tarim Basin attacks people near villages and is attracted to light. In Inner Mongolia the form
occurs from early June to late August and has one peak in July.

Phlebotomus (Larroussius) marine Rioux, Croset, Leger & Bailly-Choumara

(Map 7)

Phlebotomus (Larroussius) mariae Rioux, Croset, Leger & Bailly-Choumara, 1974: 92 [<£]. Syntypes 6<$,
MOROCCO (EM, Montpellier).

Phlebotomus (Larroussius) mascittti Grassi

Phlebotomus mascittii Grassi, 1908: 681.

Phlebotomus (Adlerius) mascittii Grassi; Perfil'ev, 1968: 8, 16.

Phlebotomus (Larroussius) mascittii canaaniticus Adler & Theodor

(Map 8)

Phlebotus Adler & Theodor, 1 93 1 b : 468 [? £] . Syntypes 2 $, 4 <J, ISRAEL (BMNH).

Phlebotomus larroussei var. canaaniticus Adler & Theodor; Adler, Theodor & Witenberg, 1938 : 501.

Phlebotomus (Larroussius) mascittii canaaniticus Adler & Theodor ; Theodor, 1958: 31 ; Perfil'ev, 1968: 95.

DISTRIBUTION. General: Perfil'ev (1968: 95, east of typical subspecies, in Israel, Jordan and Syria). Israel:
Adler & Theodor (19316: 471, Ben Shemen, Rosh Pinna, Tel Aviv); Adler & Witenberg (1938: 500, Jerusa-
lem area).

Phlebotomus (Larroussius) mascittii mascittii Grassi
(Map 8)

Phlebotomus mascittii Grassi, 1908: 681 [? £); Newstead, 1914: 182; Sinton, 1928: 310; Adler & Theodor,

1931a: 106; Hertig, 1950: 453 [discussion]; Raynal, 1954: 306; Guevara-Benttez, Ubeda Ontiveros &

Morillas Marquez, 1978: 832. Lectotype <J, ITALY (BMNH), designated by Hertig, 1950: 457 [examined].
Phlebotomus larroussei Langeron & Nitzulescu, 1931: 73; Raynal & Le Gac, 1932: 504; Adler, Theodor &

Witenberg, 1938: 498; Hertig, 1950: 455 [probably mascittii]. Syntypes ?, FRANCE (depository unknown).

[Synonymized by Sacca, 1948a: 226; Raynal, 1954: 307.]
Phlebotomus vesuvianus Adler & Theodor, 1931: 108. Syntypes 13 $, ITALY (BMNH). [Synonymized by

Raynal, 1954: 307; Theodor, 1958: 29.]
Phlebotomus perniciosus var. nitzulescui Simic, 1932: 432. Syntypes 3 $, YUGOSLAVIA (depository unknown).

[Synonymized by Raynal, 1954: 307; Theodor, 1958: 29.]
Phlebotomus (Phlebotomus) larroussei Langeron & Nitzulescu; Parrot, 1941 : 45.
Phlebotomus (Adlerius) larroussei Langeron & Nitzulescu; Theodor, 1948: 108.
Phlebotomus (Adlerius) mascittii mascittii Grassi; Sacca, 1949ft: 552; Theodor, 1958: 29; Croset, 1969: 300;

Houin, Abonnenc & Deniau, 1971 : 642.
Phlebotomus (Adlerius) mascittii Grassi; Perfil'ev, 1968: 16 [discussion], 95; Rioux & Golvan, 1969: 27, 51,

73; Croset, 1969: 300; Houin, Abonnenc & Deniau, 1971 : 642; Biocca, Coluzzi & Constantini, 1977ft: 31.
Phlebotomus mascittii Grassi ; Artemiev, 1980: 1181 [male fits Larroussius], 1 185.

9. Leg formula (after Raynal & Le Gac, 1932) 100, 135, 83; 135, 158, 89; 1 14, 187, 109 (0-86).

DISTRIBUTION. General: Perfil'ev (1968: 95, western Europe as far east as Greece); Theodor (1958: 31,
Corsica, Crete, Cyprus, France, Italy and Yugoslavia). Europe: Croset (1969: 305, map). Crete: Elounda,
35°16'N, 25°42'E (13.V.1979, D. M. A., biting man indoors). Cyprus: Adler (1946: 503). France: Colas-
Belcour & Rageau (1956, map); Croset (1969: 303, map); Houin et al. (1977: 113, map, 114); Rioux &

THE GENUS PHLEBOTOMUS 159

Golvan (1969: 75, 76, map, Calvados, Savignies etc.). Italy: Corradetti et al. (1956a: 6, map); Biocca et al.
(1977 a: 160, map; 1911 b: 20, rare, 29, map); Maroli & Bettini (1977: 318); Puccini et al. (1977: 38, map).
Switzerland :Gaschen(1956a: 225; I956b: 228); Perfil'ev( 1968: 1 6). Turkey : Houinet al. (197 '1: 642).

NOTE. P. mascittii is here placed provisionally in Larroussius.

Professor J. A. Rioux (1979, in litt.) told me the history of the Calvados (France) record, one of
the most northerly for a sandfly, which was recounted to him by the late J. Colas Belcour. 'Dans
sa proprietee de Normandie [at Conde-sur-Ife, 49°03'N, 0°07'W] sejournait a cetteepoque, Mme
Colas Belcour. En plain jour, elle est attaquee par un Phlebotome. Elle a le reflexe de couvrir
1'insecte avec un verre a boire et done de le capturer vivant. Elle alerte immediatement son mari
qui travaillait a 1'Institut Pasteur, dans de service de M. Roubaud. Colas Belcour recoupere le
Phlebotome, 1'identifie et le public avec son epouse [Colas Belcour & Colas Belcour, 1929]
comme Phlebotomus perniciosus'. Langeron & Nitzulescu (1932: 293) thought it was probably P.
larroussei (= mascittii), and Colas Belcour & Tisseuil (1936: 121, footnote) and Raynal (1954:
309) agreed.

Savignies is 49°28'N, 01°58'E, and Rioux & Golvan (1969: 75) remarked that the species
doubtless existed in Belgium, Germany and Luxembourg.

This species bites man but its ecology is little known (Croset, 1969: 309).

Phlebotomus (Larroussius) orientalis Parrot
(Map 8)

Phlebotomus (Phlebotomus) langeroni var. orientalis Parrot, 1936: 30 [9 ^] ; Kirk & Lewis, 1946a: 39; 19466:

120; 1948 : 326; 1955: 235. Syntypes 24 9, 32 & ETHIOPIA (IP, Algiers).
[Phlebotomus perniciosus Newstead; Archibald & Mansour, 1937: 395; Sinton, 1937: 404; Kirk, 1939: 541.

Misidentifications according to Kirk & Lewis, 1940: 627.]
[Phlebotomus langeroni Nitzulescu; Theodor, 1938: 165. Misidentification according to Kirk & Lewis, 1940:

627.]
Phlebotomus (Phlebotomus) orientalis Parrot; Parrot & Clastrier, 1946: 64; Kirk & Lewis, 1951 : 432; 1952:

340; Heisch & Guggisberg, 1952: 427; Parrot, 1953: 113; Abonnenc, Dyemkouma & Hamon, 1964: 160;

Abonnenc & Minter, 1965: 72; Hoogstraal & Heyneman, 1969: 1156; Abonnenc, 1972: 118.
Phlebotomus (Larroussius) langeroni orientalis Parrot; Theodor, 1958: 24; Qutubuddin, 1962: 597; Lewis &

Hitchcock, 1968: 1 18; Perfil'ev, 1968: 92; Lewis, Minter & Ashford, 1974: 440.
Phlebotomus (Phlebotomus) langeroni orientalis Parrot; Quate, 1964: 238.
Phlebotomus orientalis Parrot; Davis, 1967: 52; Ashford, 1974: 610; Hoogstraal & Heyneman, 1969: 1156

[synonymy].

DISTRIBUTION. Africa: Abonnenc (1972: 259). Ethiopia: Ashford (1974: 608; 1977: 236, wide altitude range);
Diredawa (1936, C. A. V. B.); Fuller et al. (1979: 419, map); Gemetchu & Fuller (1976: 82); Gemetchu et al.
(1975: 45; 1977: 209). Kenya: Minter (1964: 207); Wajir (1943, J. P. M.). Saudi Arabia: Lewis & Buttiker,
1980: 263). Southern Yemen: 'Wadi Ayaraq' (1962, S. A. S.); Whittingham (1937, as P. perniciosus). Sudan:
Hoogstraal & Heyneman (1969: 1155). Yemen: Buttiker & Lewis (1979: 370); Hoogstraal & Heyneman
(1969: 11 57); Lewis (19746: 188).

NOTE. In the Sudan and Ethiopia this species occurs in Acacia seyal-Balanites forest which is
associated with deeply cracking dark clay soils (Fuller et al., 1979: 429). It was suspected of being
the vector of VL in the Sudan by Kirk & Lewis (Adler, 1964: 78, 90) and shown to be so by the
extensive work of Hoogstraal & Heyneman (1969: 1185, 1186, 1194) (Abonnenc, 1972: 34, 120;
Wilcocks & Manson-Bahr, 1972: 122; Williams & Coelho, 1978: 17). It may transmit the disease
in Ethiopia (Ashford et al., 1973: 263; Fuller et al., 1979: 429; Gemetchu et al, 1977: 209; White,
1977: 163).

Phlebotomus (Larroussius) pedifer Lewis, Mutinga & Ashford

(Map 8)

Phlebotomus (Larroussius) pedifer Lewis, Mutinga & Ashford, 1972: 12 [9 c?]; Mutinga, 1975: 347. Holotype

cJ, KENYA (BMNH) [examined].
Phlebotomus pedifer Lewis, Mutinga & Ashford ; Ashford, 1974: 610.

160 D. J. LEWIS

DISTRIBUTION. Ethiopia : Ashford (1974: 610, Boleta Forest, Ochollo); Lewis et al. (1972: 132, Shabe, 7°31'N,
36°30'E). Kenya: Lewis et al. (1972: 132, Mount Elgon area). Sudan: Lewis et al. (1972: 132, Gilo and
Katire).

NOTE. P. pedifer bites man readily out of doors in the Mount Elgon area and transmits CL
(Mutinga, 1975: 346) and is a vector of CL in Ethiopia (Ashford, 1977: 236; Bray, 1974: 92;
Peters et al., 1977: 502; White, 1977: 163).

Phlebotomus (Larroussius) perfiliem Parrot

Phlebotomus perfiliewi Parrot, 1930: 383 ; Corradetti, Sacca & Neri, 19566: 105; 1957: 226.
Phlebotomus (Larroussius) perfiliewi Parrot; Perfil'ev, 1968: 48 [larva], 50, 62, 75, 76, 83.

Phlebotomus (Larroussius) perfiliewi galilaeus Theodor
(Map 8)

Phlebotomus (Larroussius) perfiliewi galilaeus Theodor, 1958: 26 [<$; distinction from subsp. transcaucasicus
uncertain] ; Theodor & Mesghali, 1964: 292; Perfil'ev, 1968: 267. Syntypes^, CYPRUS, ISRAEL (BMNH).

DISTRIBUTION. Cyprus and Israel: Theodor (1958: 26). Cyprus: Liopetri, Panagera, 33°04'N, 35°20'E, Sa-
lamis,Sotira area (x. 1971, J. P. T. B.). Turkey :Yasarol( 1980).

Phlebotomus (Larroussius) perfiliem perfiliem Parrot
(Map 8)

Phlebotomus perfiliewi Parrot, 1930: 383 [<J]; Hertig, 19496: 286; Sacca, 1950: 681 [early stages]; Corra-
detti, Sacca & Neri, 19566: 105 [figures in pi. 2 transposed]; 1957: 226; Corradetti, Neri, Verolimi,
Palmieri & Proietti, 1961 : 102. Syntypes 4^, U.S.S.R. (IP, Algiers?).
Phlebotomus macedonicus Adler & Theodor, 1931: 468 [? £]. Syntypes 4 ?, 19 £, GREECE (BMNH).

[Synonymized by Adler, 1946: 500.]

Phlebotomus perniciosus var. ; Simic, 1932 : 432. [Synonymized by Simic & Zivkovic, 1956: 384.]
Phlebotomus sp. n.; Simic & Zivkovic, 1947: 195. [Synonymized by Simic & Zivkovic, 1956: 384.]
Phlebotomus (Larroussius) perfiliewi perfiliewi Parrot; Theodor, 1958: 25; Perfil'ev, 1968: 263.
Phlebotomus (Larroussius) perfiliewi Parrot; Perfil'ev, 1968: 263 etc.; Croset, 1969: 327; Biocca, Coluzzi &
Constantini, 1977a: 160-162; 19776: 30; Rioux, Croset, Leger & Rosin, 1977: 378; Croset, Rioux, Maistre
&Bayar, 1978:738.

DISTRIBUTION. Balkans, Italy, Malta, north-west Africa and U.S.S.R. (Crimea): Dolmatova (1962: 459);
Theodor (1958: 26). Mediterranean area: Croset (1969: 333, map); Dedet et al. (1977: 256); Rioux et al.
(1977: 379, map). Algeria: Dedet (1979, map in litt.). Greece: Hadjinicolaou (1958: 968); Leger et al. (1979:
20); Macedonia (viii.1918, J. W.). Italy: Biocca et al. (1977a: 161, map; 19776: 28, map); Hertig (1949a: 796,
797); Maroli & Bettini (1977: 318). Morocco: Rioux et al. (1977: 377). Rumania: Duport et al. (1971).
Sardinia: Hertig (1949a: 798). Tunisia: Chadli, Dancescu et al. (1970: 361); Chadli, Remain et al. (1970:
358); Croset (1969: 331, map); Croset et al. (1978: 737, map); Dedet (1971: 157). Turkey: Yasarol (1980).
U.S.S.R.: Parrot (1930: 383); Perfil'ev (1968: 266, Moldavia, north-east Caucasus). Yugoslavia: Simic &
Zivkovic (1956: 383-385, Kosovo i Metohija, Makedonija, Serbia, Sibenik, Split and Vojvodina).

NOTE. P. perfiliewi is a main vector of Le. donovani in Greece (Perfil'ev, 1968: 142), transmits VL
in Serbia and possibly Greece, and canine leishmaniasis in Tunisia (Leger et al, 1979: 20). It may
have transmitted VL in Emilia-Romagna in 1971 under unusual weather conditions (Killick-
Kendrick et al. (1977: 169, 173), and is probably a vector of VL in Rumania (Adler, 1964: 79). Its
secondary role in canine kala-azar in Tunisia is discussed by Maroli & Bettini (1977: 320), and its
relation to VL in Yugoslavia by them and Lupascu et al. (1977 : 192).

This species transmits CL in Italy (Biocca et al., 1977 'b: 20; Corradetti, 1977: 194; Rivosecchi
et al., 1977: 135), is a possible vector in Italy (Killick-Kendrick et al., 1977: 169, 170, 173;
Lupascu et al., 1977: 192), and is the probable vector in the Abruzzi (Croset et al., 1978: 739). It is
strongly suspected of transmitting CL in part of Grosseto Province of Italy and of being the
vector in the Abruzzi and Emilia-Romagna (Maroli & Bettini, 1977: 315, 320).

THE GENUS PHLEBOTOMUS 161

Phlebotomm (Larroussius) perfiliewi transcaucasicus Perfil'ev

(Map 8)

Phlebotomus transcaucasicus Perfil'ev, 1937: 108 [£]• Type(s), U.S.S.R.: Transcaucasia, Divichi (near Baku)

and Nakhichevan (ZI, Leningrad?).
Phlebotomus (Larroussius) perfiliewi transcaucasicus Perfil'ev; Theodor, 1958: 26 [status undecided, possibly

a synonym of galilaeusj ; Theodor & Mesghali, 1964: 291; Perfil'ev, 1968: 267 [9]; Ahmad, 1976: 43, 152,

156.

DISTRIBUTION. Iran: Theodor & Mesghali (1964: 292, one at Kazvin). Iraq: Ahmad (1976: 99). U.S.S.R.:
Perfil'ev (1968: 267) and Theodor & Mesghali (1964, Azerbaydzhan, Baku, Nakhichevan, north-east Cau-
casus and Transcaucasia); Dzhavadov et al. (1978 : 143, Astanly, Dzhalilabad, Kanaga and Tazakent).

NOTE. This form appears to be a vector of Le. donovani in Azerbaydzhan (PerfiFev, 1968 : 142).

Phlebotomus (Larroussius) perniciosus Newstead
(Map 9)

Phlebotomus nigerrimus Newstead, 191 la: 68 [$]; Summers, 1913: 106; Perfil'ev, 1968: 9, 253. Syntypes2 9,

MALTA. [Synonymized with P. perniciosus by Raynal, 1954: 301.]
Phlebotomus perniciosus Newstead, 191 la: 70 [? <J]; Gaschen, 1945: 140 [9 £|; 1956: 226; Hertig, 1950:

453; Sacca, 1950: 684 [early stages] ; Raynal, 1954: 306; Corradetti, Sacca & Neri, 19566: 105; 1957: 226;

Abonnenc & Lariviere, 1957: 401 [larva]; Guevara-Benitez, Ubeda-Ontiveros & Morillas-Marquez,

1978: 821, 831. Syntypes 9 3, MALTA (depository unknown).
Phlebotomus legeri Mansion, 1913: 639; 1914: 588. Syntypes 9 & CORSICA (L, Bastia?; Mansion, 1914: 590).

[Synonymized by Larrousse, 1921 : 40; Raynal, 1954: 301 ; Perfil'ev, 1968: 16; Theodor, 1958: 26.]
Phlebotomus perniciosus var. nigerrimus Newstead [?]; Newstead, 1914: 184; Perfil'ev, 1968: 9, 161, 253.

[Synonymized by Larrousse, 1921 : 40; Raynal, 1954: 301 ; Perfil'ev, 1968 : 16.]
Phlebotomus lusitanicus Franca, 1918: 732. Type(s), PORTUGAL (MB, Colares?; Franca, 1918: 731; 1922: 9,

18). [Synonymized by Larrousse, 1921 : 37; Raynal, 1954: 301.]
Phlebotomus grassii Pierantoni, 1925: 5. Type(s), ITALY: near Naples (MZ, Turin?; Pierantoni, 1925: 1, 8).

[Synonymized by Adler & Theodor, 193 la: 106.]

Phlebotomus major Annandale var. perniciosus Newstead; Sinton, 1928: 303.
Phlebotomus (Larroussius) perniciosus Newstead; Theodor, 1948: 107; 1958: 26; Sacca, 19496: 552; Nicoli,

1956: 112 [hairs]; Perfil'ev, 1968: 9; Rioux & Golvan, 1969: 25, 78; Croset, 1969: 397; Rioux, Croset,

Leger & Bailly-Choumara, 1974: 96; Biocca, Coluzzi & Constantini, 1977a: 162; 19776: 30; Croset,

Rioux, Maistre & Bayar, 1978: 740.
Phlebotomus perniciosus legeri Mansion; Nicoli, 1955: 33; 1956: 110 [hairs].

9. Leg formula, Spain, after Zariquiey, 1937: 411, 100, 110, 66; 101, 133, 77; 117, 177, 101 (0-78)(? &<£ by
ToumanofT& Chassignet, 1954: 680).

DISTRIBUTION. Western Europe and Africa: Croset (1969: 407). Algeria: Dedet (1979, map in litt.); Dedet &
Addadi (1977: 86); Dedet et al. (1977: 254, map). France: Colas-Belcour (1958: 826, map); Colas-Belcour &
Rageau (1956, map); Croset (1969: 403, map); Houin et al. (1977: 113, map, 114); Rioux & Golvan (1969: 51,
83, map). Italy: Biocca et al. (1977a: 161, map; 19776: 20, common and widespread, 28, map); Corradetti et
al. (1956a: 6, map); Maroli & Bettini (1977: 318); Puccini et al. (1977: 38, map). Jersey: [?] Marett (1923a;
19236, St. Helier which is 49°12'N). Libya: Ashford et al. (1977: 265). Morocco: Bailly-Choumara et al.
(1971: 453). Portugal: Azavedo (1954: 247); Franca (1918: 731). Sardinia: Hertig (1949a: 798). Spain:
'Fulgencio' and 'Jalavara' (1931, through J. A. S.); Gil Collado (1977: 186); Najera (1937: 1488); Zariquiey
(1944: 18). Switzerland: Gaschen (19566: 228). Tunisia: Chadli, Dancescu et al. (1970: 363); Chadli, Romain
et al. (1970: 358); Croset (1969: 405); Croset et al. (1978: 737, map); Dedet (1971: 157). Turkey: Yasarol
(1980). Yugoslavia: Simic & Zivkovic 1956: 383-385, north Dalmatia and Istra).

NOTE. 'P. nigerrimus'' was described from females, and Newstead (191 la) hoped that P. J. Marret
would find males. Newstead (1914) examined some and considered them a dark variant of
perniciosus, but a male labelled 'Malta. 1910. Capt. Marett. Phlebotomus perniciosus var. ni-
gerrimus, Newst. Type $. Pres. R. Newstead B. M. 1947-141.' is P. perfiliewi. It seems likely that
the original nigerrimus were perniciosus, but could be confused with dark forms of at least one
other species.

162 D. J. LEWIS

The name nigerrimus has page preference over perniciosus but became a junior synonym of it
following the action of Raynal, the first reviser (ICZN, 1964: Article 24 (a)).

In Jersey on 1 September 1923 Marett (1923a, b) was surprised to find a phlebotomine and
wrote to the press as follows. 'Sir — I have the honour to request you to insert this letter in your
paper. On the night of the 1st instant, I captured a Phlebotomus fly, and should be glad to know
if any scientist may be breeding the fly in the Island. I have the honour to be, Sir, Your obedient
servant, P. Jauvin Marett Lt. Col., M.O.H., States, Jersey. Royal Square 2/9/23.' The fly, a male,
was identified by Newstead as P. perniciosus, but, in view of Hertig's (1953: 453) remarks on the
aedeagus, it may conceivably have been P. mascittii. Dr W. J. Le Quesne (1970, 1971, in litt.) told
me of the letter and informed me that Marett lived successively in two houses on the outskirts of
St Helier. Le Quesne and other entomologists have sought this species without result. It may
have been in Jersey since the island was attached to the mainland (Ragge, 1965: 256) and have
disappeared with housing developments. It probably exists at the same latitude in France (Rioux
&Gol van, 1969: 83).

P. perniciosus bites man indoors in Tunisia where its two annual peaks correspond with the
transmission periods of VL (Croset et al, 1978: 741, 743). It has been readily infected with the
parasites of VL (Molyneux, 1977: 48) and is a vector in the western Mediterranean basin (Hoog-
straal & Heyneman, 1969: 1185, 1186; Wilcocks & Manson-Bahr, 1972: 121), the main vector in
north Africa (Dedet, 1976: 422; PernTev, 1968: 142; Theodor, 1964: 480), and very probably the
habitual vector in Tunisia (Croset et a/., 1978: 744). It transmits canine leishmaniasis near Tours
in France (Houin et al., 1977: 114), is probably the main vector of VL in Provence (Ranque et al.,
1977: 286, 292), has been proved to be the vector of VL in southern Italy (Biocca et al., 19776;
Rivosecchi, 1977: 135), Sicily and Malta (Biocca et al., 1911 a: 165), and is probably the vector of
VL in the Monte Argentario area of the Italian Province of Grosseto (Maroli & Bettini, 1977:
315, 320). Its relation to VL in general was discussed by Saf'yanova (1967: 36) and in Italy by
Killick-Kendrick et al. (1977: 170, 173). It has been found to harbour a sandfly fever virus in Italy
(Rivosecchi, 1977: 136).

Phlebotomus (Larroussius) smirnovi PerfiPev
(Map 9)

Phlebotomus smirnovi Perfil'ev, 1941: 279 [$ <J]; Shakirzyanova, 1950: 26. Syntypes & <J, U.S.S.R. (ZI,

Leningrad?).
Phlebotomus (Larroussius) smirnovi Perfil'ev; Theodor, 1958: 27; Perfil'ev, 1968: 277.

DISTRIBUTION. U.S.S.R.: Dergacheva et al. (1978, Kzyl-Orda area); Perfil'ev (1968: 95, 279, 281, Osh area in
Kirgiziya).

NOTE. P. smirnovi probably transmits VL in the Kzyl-Orda region (Dergacheva et al., 1978).

Phlebotomus (Larroussius) somaliensis Abonnenc, Adam & Bailly-Choumara

(Map 9)

Phlebotomus somaliensis Abonnenc, Adam & Bailly-Choumara, 1959: 588 [$]; Abonnenc & Minter, 1965:
38; Abonnenc, 1972: 120. Holotype ?, SOMALI REPUBLIC: cave at 'Shamah Aleh' near the Asseh Hills (IP,
Algiers).

Phlebotomus (Larroussius) tobbi Adler & Theodor
(Map 9)

Phlebotomus perniciosus var. tobbi Adler & Theodor in Adler, Theodor & Lourie, 1930: 536 [$ £]\ Ni-
tzulescu, 19316: 267. Syntypes 27 $, 40 J, IRAN; ISRAEL (BMNH).

Phlebotomus tobbi Adler & Theodor; Parrot, 1934: 80.

Phlebotomus pirumovi Burakova & Mirzayan, 1934: 89 [short description in a footnote & reference to full
description which was apparently not published]; Perfil'ev, 1968: 271, 273. Type(s), U.S.S.R. (ZI, Len-
ingrad?). [Synonymized by Perfil'ev, 1941 : 273, 281.]

THE GENUS PHLEBOTOMVS 163

Phlebotomus perniciosus Newstead [in part]; Adler & Theodor, 1957: 215 [proboscis].

Phlebotomus (Larroussius) perniciosus tobbi Adler & Theodor; Theodor, 1958: 27; Rioux & Golvan, 1969:

81, 82; Ahmad, 1976: 43, 144; Guevara-Bemtez, Ubeda-Ontiveros & Morillas-Marquez, 1978: 832.
Phlebotomus (Larroussius) tobbi Adler & Theodor; Perfil'ev, 1968: 271; Croset, 1969: 341; Houin et al.,

1971:635.

DISTRIBUTION. Eastern Mediterranean area: Perfil'ev (1968: 273, Cyprus, Greece, Israel, Jordan and Syria);
Theodor & Mesghali (1964: 292, widely distributed from Yugoslavia to north-west Iran). Europe and North
Africa: Croset (1969: 342, probably not in western Europe; 1967: 177, Tunisian records probably refer to P.
longicuspis or P. perfiliewi); Dolmatova (1962: 460, map, evidently including P. perniciosus); Houin (1977:
167, not west of Yugoslavia; old records from France and Spain must refer to P. perniciosus). Albania:
Perfil'ev (1968: 8, possibly P. perniciosus). Greece: Hadjinicolaou (1958: 968, 972); Hertig (1949a: 779,
781-783); 'Kerken', Struma Valley (26.vi.1935, P. A. B.); Leger et al. (1979: 17). Iran: Adler et al. (1930: 537,
Resht); Lewis et al. (1961: 206); Nadim & Rashti (1978: 27, areas indicated); Nadim et al. (1977: 215);
Theodor & Mesghali (1964: 291, 292, in north-west; probably some early records refer to P. major). Iraq:
Ahmad (1976: 99). Israel: Adler et al. (1930: 537, Ajaleth, Rosh Pinna); Pazael (7.V.1979, Y. S.). Lebanon:
Mechref (1964, L. E. S.). Sicily: Catania (<$ presented to BMNH in 1931 by S. A. as P. perniciosus; 'var.
tobbf added in Theodor's writing). Turkey: Houin et al. (1971 : 635). U.S.S.R.: Dzhavadov et al. (1978: 143,
Astanty, Dzhalilabad and Tazakent); Perfil'ev (1968: 273, Armenia, Azerbaydzhan and Gruziya; absent
from Turkestan); Theodor & Mesghali (1964: 292). Yugoslavia: Nitzulescu (19316: 267, Skoplje); Simic &
Zivkovic (1956: 383-385, north and south Dalmatia, Hercegovinia, Istra, Kosovo i Metohija, Makedonija,
Montenegro and Serbia).

NOTE. P. tobbi is a vector of VL in the eastern Mediterranean area and probably in Transcaucasia
(Theodor, 1964: 480, 485) and is probably a vector in Azerbaydzhan (Perfil'ev, 1968: 142) and
Cyprus (Adler, 1946: 510). It is too rare to be significant in Greece(Leger et al., 1979: 23).

Phlebotomus (Larroussius) wenyoni Adler & Theodor
(Map 9)

Phlebotomus wenyoni Adler & Theodor in Adler, Theodor & Lourie, 1930: 353 [9 £]. Syntypes 25 ?, 53 <J,

IRAN (BMNH).
Phlebotomus (Larroussius) wenyoni Adler & Theodor; Theodor, 1958: 27; Theodor & Mesghali, 1964: 291;

Ahmad, 1976:49, 162.

DISTRIBUTION. General: Theodor & Mesghali (1964: 292, apparently very restricted). Iran: Nadim et al.
(1978: 27, Chahar Mahal and other areas); Theodor & Mesghali (1964: 292, Hamadan, Kermanshah,
Malayer and Tehran ; seemed restricted to north-west). Iraq: Ahmad (1976: 99); Theodor & Mesghali (1964:
292, Salahuddin). Turkey: Yasarol (1980). U.S.S.R.: Perfil'ev (1968: 270, Ashkabad and Karakala areas in
Turkmeniya); Petrishcheva (1935: 20).

According to Theodor & Mesghali (1964) some old records may refer to P. keshishiani or P.
major.

Subgenus ADLERWS Nitzulescu

Phlebotomus subgenus Adlerius Nitzulescu, 1931: 271; Theodor, 1948: 98; 1958: 27; Theodor & Mesghali,
1964: 292; Perfil'ev, 1968: 73, 81, 280 [also vii, 8, 48 (larva), 51, 60 (egg), 75, 77, 280 on P. chinensis s. 1.];
Lewis, 1978ft: 239; Artemiev, 1978: 19, 20, 75; 1980: 1171, 1180. Type-species: Phlebotomus chinensis
Newstead, 1926, by original designation.

Professor Leng Yan-jia informed me in 1980 that several taxa in China await study, therefore
most 'P. chinensis' in Map 10 are marked with a query.

Key to the species of subgenus Adlerius (after Artemiev, 1980).

Females

Artemiev recommends that males should be identified first, and accompanying females compared with data
in his table. Some characters are indefinite, and descriptions should be consulted.

164

D. J. LEWIS

Males

1

10

11

12

13

14

15

16

17

18

Two ascoids on antennal segments 3-1 5 2

One ascoid on antennal segments 9-15 3

Subterminal tubercle of aedeagus far (30-35 /mi) from tip

Subterminal tubercle of aedeagus near (6-8 um) tip

Antenna 8 with two ascoids

Antenna 8 with one ascoid

Coxite with 14-27 hairs in group, rarely 29

Coxite with 29-1 15 hairs in group, rarely 27

Whole hair-group on basal half of coxite. Tubercle of aedeagus 6-16 um from tip
Part of hair-group on distal half of coxite. Tubercle of aedeagus 19-28 um from tip
All 69- 1 1 4 hairs of group on basal half of coxite ; coxite wide ....

Some of 27-85 hairs of group on distal half of coxite

Aedeagus with rectangular subterminal notch

Aedeagus with normal obtuse-angled subterminal notch

Coxite with 27-50 group-hairs. Sandfly dark

Coxite with 50-85 group-hairs. Sandfly of normal colour or pale

Sperm tubes long (1200-1 700 /im)

Sperm tubes of average length (900-1 100 /zm)

Coxite with 50-60 group-hairs. Sandfly large (antenna 3 = 400-^430 um) .

Coxite with 65-75 group-hairs. Sandfly of normal size (antenna 3 = 340-375 /mi). Iran

'Sp. 1' of doubtful status (related to arabicus)

Antenna 6 and 7 with two ascoids of same length 12

Antenna 6 and 7 with one ascoid or with one long and one short 14

Coxite with 35-60 group-hairs. Sperm tubes 740-1000 /zm long. Sandfly small . . davidi (p. 166)

Coxite with 69-1 10 group-hairs 13

Coxite with 69-94 group-hairs hindustanicus (p. 166)

Coxite with 99-1 11 group-hairs. Afghanistan . . . Sp. 2 (possibly a subsp. of hindustanicus)

chinensis (p. 165)

simici (p. 168)

4

11

5

6

turanicus (p. 168)

brevis (p. 165)

. rupester (p. 167)

7

halepensis (p. 166)
8

kabulensis (p. 167)
9

longiductus (p. 167)
10
arabicus (p. 164)

Coxite with 90-220 group-hairs. Ventral process of style long (about 20 um)
Coxite with 30-85 group-hairs. Ventral process of style long or short •
Coxite very wide, whole group of 125-200 hairs on its basal half

Coxite narrow, part of hair-group on its distal half

Ventral process of style long

Ventral process of style short

Whole hair-group (40-85) of coxite on its basal half

Part of hair-group on distal half of coxite

Antenna 3 = 1-20-1-55 times length of labrum. Coxite with 35-70 group-hairs
Antenna 3 = 1 -05-1-20 times length of labrum. Coxite with 30-50 group-hairs

Phlebotomus (Adlerius) angustus Artemiev

15
16

comatus (p. 166)

balcanicus (p. 165)

zulfagarensis (p. 168)

17

salangensis (p. 168)
18

. angustus (p. 164)
kyreniae (p. 167)

(Map 10)

[Phlebotomus (Adlerius) chinensis longiductus Parrot; Lewis, 1967: 21 [in part]. Misidentification.]
[Phlebotomus (Adlerius) longiductus Parrot; Artemiev, 1974: 163 [in part]; Lewis, 19786: 240 [in part].

Misidentifications.]
Phlebotomus (Adlerius) angustus Artemiev, 1978: 22 [? (J]; 1980: 1189. Holotype £, AFGHANISTAN (MI,

Moscow).

DISTRIBUTION. Afghanistan: Artemiev (1978: 22, in north and centre in high rocky mountains). Pakistan:
Lewis (1967: 23, Gilgit area, i.e. Gilgit, Gol, Gwari, Keris and Parkuta; 84 $ of Adlerius from this area
examined in 1979 found to be a mixture of P. angustus and salangensis). U.S.S.R.: Artemiev (1978: 22,
Tadjikistan and Uzbekistan).

In Afghanistan P. angustus occurs in high rocky mountains (Artemiev, 1978: 22).

Phlebotomus (Adlerius) arabicus Theodor
(Map 10)

Phlebotomus (Adlerius) chinensis arabicus Theodor, 1953: 120 [$ ^]; Artemiev, 1974: 163. Syntypes2 9, 1 c?,
YEMEN (BMNH).

THE GENUS PHLEBOTOMUS

165

Phlebotomus chinensis arabicus Theodor; Abonnenc & Minter, 1965: 32; Abonnenc, 1972: 111.
Phlebotomus (Adlerius) arabicus Theodor; Artemiev, 1980: 1 190.

DISTRIBUTION. Ethiopia (?): Ashford (1974: 610). Saudi Arabia : Biittiker & Lewis (1980). Yemen : Buttiker &
Lewis (1978 : 371); Theodor (1953 : 120).

Phlebotomus (Adlerius) balcanicm Theodor
(Map 10)

[Phlebotomus chinensis Newstead; Nitzulescu, 1930a: 367 [9 <£]• Misidentification.]

Phlebotomus (Adlerius) chinensis balcanicus Theodor, 1958: 28 [differs in some respects from Nitzulescu's
form but has similar hair-group on coxite]; Theodor & Mesghali, 1964: 292; Dancescu, 1967: 426; 1968:
189; Perfil'ev, 1968: 290; Leger, Saratsiotis, Pesson & Leger, 1979: 23, 24 [found variants which cast
doubt on status of taxon]. Holotype <£, GREECE (BMNH).
Phlebotomus (Adlerius) balcanicus Theodor; Artemiev, 1980: 1 188 [raised to species].

DISTRIBUTION. South-east Europe: Theodor (1958: 29). Crete: (as P. chinensis) Hadjinicolaou (1958: 974);
Hertig (1949a: 788, 789). Greece: (as P. chinensis) Hadjinicolaou (1958: 968, 970, 972); Hertig (1949a:
781-786); Leger et al. (1979: 17); Theodor (1958: 28, Yannitsa). Iran: Theodor & Mesghali (1964: 293).
Rumania: Duport et al. (1971: 388, 389). Turkey: Yasarol (1980). U.S.S.R.: Dergacheva (1977: 1572, Azer-
baydzhan). Yugoslavia : Zivkovic (1974: 4, map).

NOTE. This species may transmit VL in Greece (Leger et al, 1979: 23).

Phlebotomus (Adlerius) brevis Theodor & Mesghali
(Map 10)

Phlebotomus (Adlerius) chinensis brevis Theodor & Mesghali, 1964: 293 [? <£]. Holotype $, IRAN (IPH,

Tehran).
Phlebotomus (Adlerius) chinensis ismailicus Perfil'ev, 1966: 314 [<J]; 1968: 288 [$]. Type(s), U.S.S.R.: Ismail

(ZI, Leningrad?). [Synonymized by Artemiev, 1980: 1183.]

Phlebotomus (Adlerius) brevis ismailicus Perfil'ev; Artemiev & Dergacheva, 1977: 1574.
Phlebotomus (Adlerius) brevis Theodor & Mesghali; Artemiev & Dergacheva, 1977: 1572 ['chinensis' from

Agdam area of Azerbaydzhan sympatric with balcanicus and halepensis and distinct from Chinese form] ;

Artemiev, 1980: 1183.

DISTRIBUTION. Iran: Nadim et al. (1978: 27, 28); Theodor & Mesghali (1964: 293). Turkey: Yasarol (1980).
U.S.S.R.: Artemiev & Dergacheva (1977: 1574); Dzhavadov et al. (1978: 143, Astanty, Dzhalilabad, Kha-
naga and Tazakent).

Phlebotomus (Adlerius) chinensis Newstead
(Map 10)

Phlebotomus major var. chinensis Newstead, 1916: 191 [$ cj]; Foo-Hai, 1934: 498. Lectotype <$, CHINA

(BMNH), designated by Lewis, 1978ft: 239 [examined].
Phlebotomus chinensis Newstead; Sinton, 1928: 306 [in part, synonymy]; 1932: 59; 1933: 418; Patton &

Evans, 1929: 29, 83, 137, 151, 162, 166, 215, 219, 223, 227; Yao & Wu, 1941: 78; Guan et al., 1980: 25

[variation].
Phlebotomus (Adlerius) chinensis chinensis Newstead; Theodor, 1958; Theodor & Mesghali, 1964: 292;

Perfil'ev, 1968: 281 ; Lewis, 1978ft: 239.
Phlebotomus (Adlerius) chinensis Newstead; Artemiev, 1980: 1183.

cJ (extra fact, China, Wo Fu Su Temple, l-6.vii.1914, R. A. B.). Leg formula 100, 131, 83; 98, 160, 95; 111,
188, 108 (0-81). Legs also examined by Nitzulescu (1930a: 365, 369) and Patton & Hindle (1926: 406).

RECORDED DISTRIBUTION. See note under subgenus. China: Guan et al. (1980, Gansu, Shanxi and Sichuan
Provinces); He K'ai-zeng et al. (1959); Leng (1978: 7, no. 5 on map); Leng (1980, pers. comm., Qianshan);
Leng & Chang ( 1 964 : 208) ; Lewis ( 1 978ft : 239, 325) ; Patton & Hindle (1928: 546, Chi-nan, Tsinan).

NOTE. P. chinensis is the vector of VL in China (Dedet, 1976: 421; Hoogstraal & Heyneman,
1969: 1185; Perfil'ev, 1968: 140; Theodor, 1964:484; Wilcocks& Manson-Bahr, 1972: 121).

166 D. J. LEWIS

Phlebotomus (Adlerius) comatus Artemiev
(Map 10)

Phlebotomus (Adlerius) comatus Artemiev, 1978: 21 [£]; 1980: 1188 [?]. Holotype & AFGHANISTAN (MI,
Moscow).

The female is known but not described (Artemiev, 1979, in litt.).

DISTRIBUTION. Afghanistan: Artemiev (1978: 21). Nepal (?):Chobhar (1976, J. W. JV.,$ only examined).

NOTE. In Afghanistan P. comatus is a rare species of rocky mountains between 1000 and 2600 m
(Artemiev, 1978: 21).

Phlebotomus (Adlerius) davidi Artemiev
(Map 10)

[Phlebotomus (Adlerius} chinensis arabicus Theodor; Lewis, 1974: 189; Lewis & Buttiker, 1980: 263. Mis-
identifications.]

[? Phlebotomus chinensis Newstead; Ashford, 1974: 610 [very like arabicus']. Misidentification.]
Phlebotomus (Adlerius) davidi Artemiev, 1980: 1 191 [$ d~\. Holotype ^, YEMEN : Ta'izz (MI, Moscow).

DISTRIBUTION. Ethiopia : Artemiev (1980, probably this species). Yemen: Ta'izz.

Phlebotomus (Adlerius) halepensis Theodor
(Map 10)

Phlebotomus (Adlerius} chinensis halepensis Theodor, 1958: 29 [§ £]; Mesghali, 1963: 1075; Theodor &

Mesghali, 1964: 293; Perfil'ev, 1968: 291. Syntypes, IRAN, SYRIA (BMNH).
^Phlebotomus chinensis monticola Tarvit-Gontar, 1956: 158; Perfil'ev, 1968: 295 [position doubtful].

Type(s), U.S.S.R. (depository unknown). [Junior primary homonym of Phlebotomus monticolus Lima,

1932.]
Phlebotomus (Adlerius) halepensis Theodor; Artemiev, 1980: 1190.

DISTRIBUTION. Iran, north Syria and U.S.S.R.: Theodor (1958: 29). Iran: Nadim et al. (1977: 215; 1978: 27,
28); Theodor & Mesghali (1964: 293, Tehran etc.). Israel: Dishon(= Deishum, 28.vii.1942, det. 0. T}. Syria:
Theodor & Mesghali (1964: 293, Aleppo). Turkey: Yasarol (1980). U.S.S.R.: Artemiev & Dergacheva (1977:
1572); Theodor & Mesghali (1964: 293, Tbilisi).

NOTE. P. monticola might be a synonym of P. halepensis but the name must be permanently
rejected as a junior primary homonym of P. monticolus Lima, 1932: 50 (ICZN, 1964: Articles 53,
57 and 57(b)).

In the U.S.S.R. (Perfil'ev, 1935 : 96, map; 1968 : 89) 'P. chinensis' occurs as far north as 47° in the
southern Ukraine, in a part of East Kazakhstan with a mean winter temperature of — 16°C, and
up to 2800 m.

'P. chinensis'' appears to be a vector of Le. donovani in Gruziya (Perfil'ev, 1968: 142) and
possibly in Turkestan (Theodor, 1964: 485) and was considered to be a main vector in Trans-
caucasia, Central Asia and Kazakhstan (Sergiev, 1979: 208).

Phlebotomus (Adlerius) hindustanicus Theodor
(Map 10)

[Phlebotomus chinensis Newstead; Sinton, 1928: 306 [in part]. Misidentification.]

Phlebotomus (Adlerius} chinensis hindustanicus Theodor, 1958: 29, 30 [$ £]. Syntypes <3, NORTH- WEST OF

INDIAN SUBCONTINENT (BMNH).

[Phlebotomus (Adlerius} chinensis longiductus Parrot; Lewis, 1967: 21 [in part]. Misidentification.]
Phlebotomus (Adlerius) hindustanicus Theodor; Artemiev, 1978: 23 [and a possible variant classed as Species

1]; 1980: 1191.

THE GENUS PHLEBOTOMUS 167

DISTRIBUTION. Afghanistan: Artemiev (1978: 23, Gorband valley, Mahi-Par, 34°39'N, 69°42'E and Sarobi; in
low rocky mountains). India: Bin Sar in Uttar Pradesh (BMNH); Kasauli (J. A. S.); Tapoban village in
Chamoli district (13.vi.1969, V. £>.). Pakistan: Rawalpindi (1959, H. C. R; all 11 ^ Adlerius examined in
1979 were this species). Nepal: Chobhar (1976, J. Wn.).

NOTE. In Afghanistan P. hindustanicus occurs in low rocky mountains (Artemiev, 1978: 23).

Phlebotomus (Adlerius) kabulensis Artemiev
(Map 10)

Phlebotomus (Adlerius} kabulensis Artemiev, 1978: 21 [9 £]; 1980: 1188. Holotype <J, AFGHANISTAN (MI,
Moscow).

DISTRIBUTION. Afghanistan: 'Gorband', Kabul, Kandahar and Pangshir (Artemiev, 1978).
NOTE. This species is found in dwellings and is rather thermophilic and hydrophilic.

Phlebotomus (Adlerius) kyreniae Theodor
(Map 11)

[Phlebotomus chinensis Newstead; Adler, 1946: 498 [9 £]; Theodor, 1953: 120. Misidentifications.]
Phlebotomus (Adlerius) chinensis kyreniae Theodor, 1958: 29 [9 <£]. Syntypes 9 cJ, CYPRUS (BMNH).
Phlebotomus (Adlerius) kyreniae Theodor ; Artemiev, 1980: 1185.

DISTRIBUTION. Cyprus: Theodor (1978). Turkey: Yasarol (1980).

NOTE. This form probably transmits canine kala-azar in Cyprus (Adler, 1946: 510; Lupascu et al.,
1977:192).

Phlebotomus (Adlerius) longiductus Parrot
(Map 11)

Phlebotomus major var. longiductus Parrot, 1928: 29 [<J]; 1940: 310 [in part; ascoid formula of 'longiductus'

variable]; 1946: 68; Nitzulescu, 1929: 430. Syntypes 2 <J, U.S.S.R.: Shakrisyabz near Samarkand (de-

pository unknown).

[Phlebotomus chinensis Newstead [in part] ; Nitzulescu, 1930a: 373 [?]. Misidentification.]
Phlebotomus chinensis var. longiductus Parrot; Nitzulescu, 193 la: 264 [9].
Phlebotomus (Adlerius) chinensis var. longiductus Parrot ; Theodor, 1948: 107.
Phlebotomus (Adlerius) chinensis longiductus Parrot; Theodor, 1958: 29 [in part]; PernTev, 1968: 285;

Dancescu, Cristescu, Costin et al, 1970: 57.
Phlebotomus (Adlerius) chinensis tauriae PernTev, 1966: 286; 1968: 62, 286. Type(s), U.S.S.R.: Crimea (MI,

Moscow?) [Synonymized by Artemiev, 1978: 20.]

Phlebotomus chinensis tanriae PernTev; Saladze, 1972: 617. [Mis-spelling.]
Phlebotomus (Adlerius) longiductus Parrot ; Artemiev, 1974: 163 [in part]; 1978: 20; 1980: 1 190.

DISTRIBUTION. Afghanistan: Artemiev (1978: 20, north and centre). Rumania : Artemiev (1978: 20); Dancesco
et al. (1970: 60, map); Duport et al. (1971). U.S.S.R.: Artemiev (1978: 20, Central Asia, Crimea, Kazakhstan,
northern Caucasus and southern Ukraine); Gaibov (19756, rare in Fergana area); Saladze (1972: 617,
Mtshketa district).

NOTE. In Afghanistan P. longiductus occurs in houses in plains between 1000 and 2000 m and is
very anthropophilic; in the mountains of Kazakhstan it is the main vector of VL (Artemiev, 1978:
20).

Phlebotomus (Adlerius) rupester Artemiev

(Map 11)

Phlebotomus (Adlerius) rupester Artemiev, 1978: 21 [9 <£]; 1980: 1188. Holotype <?, AFGHANISTAN (MI,
Moscow).

NOTE. In Afghanistan P. rupester occurs in very high rocky mountains up to 3300 m (Artemiev,
1978:21)

168 D. J. LEWIS

Phlebotomus (Adlerius) salangensis Artemiev
(Map 11)

Phlebotomus (Adlerius) salangensis Artemiev, 1978: 22 [? <J]; 1980: 1189. Holotypec?, AFGHANISTAN (MI,
Moscow).

Phlebotomus (Adlerius) simici Nitzulescu
(Map 11)

Phlebotomus chinensis var. simici Nitzulescu, 193 la: 264 [$ cJ]. Syntypes? <J, YUGOSLAVIA (IH, Skoplje?).

Phlebotomus (Phlebotomus) chinensis var. simici Nitzulescu; Parrot, 1941 a: 45.

Phlebotomus (Adlerius) simici Nitzulescu; Theodor, 1958: 3 1 ; Perfil'ev, 1968: 282, 293; Artemiev, 1974: 164;

1980: 1185; Leger, Saratsiotis, Pesson & Leger, 1979: 24.
Phlebotomus (Adlerius) chinensis simici Nitzulescu; Houin, Abonnenc & Deniau, 1971 : 642.

DISTRIBUTION. Balkans, Iran, Syria and Turkey: Theodor (1958: 32). Crete: Hertig (1949a: 787). Greece:

Leger et al. (1979: 17). Turkey: Houin et al. (1971: 642); Yasarol (1980). U.S.S.R.: Gaibov (1975ft, Fergana
area); Perfil'ev (1968: 285, central Asia and Transcaucasia). Yugoslavia: Nitzulescu (1931a: 265, Skoplje);
Simic & Zivkovic (1956: 383-385, Hercegovina, Kosovo i Metohija, Makedonija and Serbia).

NOTE. P. simici is a vector of VL in the eastern Mediterranean area (Theodor, 1964: 480) and is
considered to be one in Yugoslavia (Lupascu et al. 1977: 192).

Phlebotomus (Adlerius) turanicus Artemiev
(Map 11)

Phlebotomus (Adlerius) simici turanicus Artemiev, 1974: 163 [9 cJ]. Types 6 $, 4 <J, AFGHANISTAN (MI,

Moscow).
Phlebotomus (Adlerius) turanicus Artemiev, 1978: 20; 1980: 1185.

DISTRIBUTION. Afghanistan: Artemiev (1978: 20, Aliabad etc. in north, probably also in Iran). UJSJS.R.:
Artemiev (1978 : 20, southern Tadzhikistan, southern Turkmeniya and southern Uzbekistan).

NOTE. In Afghanistan P. turanicus seems to be thermophilic and xerophilic and can stand a cold
winter, is found mainly in rodent and bird burrows and sometimes in houses, and will attack man
(Artemiev, 1978:20).

Phlebotomus (Adlerius) zulfagarensis Artemiev
(Map 11)

Phlebotomus (Adlerius) zulfagarensis Artemiev, 1978: 22 [? £|; 1980: 1189. Holotype <J, U.S.S.R. (MI,
Moscow).

DISTRIBUTION. Iran: Artemiev (1978: 22). UJSJS.R.: Artemiev (1978: 22, Turkmeniya).

Subgenus EUPHLEBOTOMUS Theodor

Phlebotomus subgenus Euphlebotomus Theodor, 1948: 98; 1958: 32; Perfil'ev, 1968: 33 [parameres], 34, 78,
83; Hennig, 1972: 53; Lewis, 1978ft: 240; Artemiev, 1979: 19 [key to J including one species not
described]. Type-species : Phlebotomus argentipes Annandale & Brunetti, 1908, by original designation.

Key to the species and subspecies of subgenus Euphlebotomus

Females

1 Spermatheca bulbous with faint striations near duct, with head on a distinct narrow neck.

Pharyngeal armature comprising five or six anterior rows of grouped spicules and many

minutely spiculate transverse ridges tumenensis (p. 170)

Spermatheca not bulbous, without narrow neck. Pharyngeal armature comprising distinct teeth
or ridges 2

2 Spermatheca with faint transverse striations or indistinct segments 3

Spermatheca distinctly segmented 4

THE GENUS PHLEBOTOMUS 169

3 Spermatheca with large apical segment and about 15 indistinct segments. Pharynx with many

uniform teeth mesghalii (p. 170)

Spermatheca with small apical segment and about 30 transverse striations. Pharynx with a
median group of small teeth and, behind them, some close-set concentric lines kiangsuensis (p. 1 70)

4 Spermathecal common duct with rather thin walls. Antenna 5 without papilla . argentipes (p. 169)
Spermathecal common duct with thick walls. Antenna 5 with papilla 5

5 Antenna 3/labrum 1-0 philippinensis philippinensis (p. 170)

Antenna 3/labrum 1-4 philippinensis gouldi (p. 1 70)

Males

1 Paramere with two lobes mesghalii (p. 170)

Paramere with three lobes • 2

2 Middle lobe of paramere nearly rectangular caudatus (p. 169)

Middle lobe of paramere with rounded end 3

3 Middle lobe of paramere thicker than main (upper) lobe kiangsuensis (p. 170)

Middle lobe of paramere thinner than main lobe 4

4 Main lobe of paramere much more than twice length of middle lobe, lower lobe narrow, depth of

paramere about 0-29 of its length (measured to junction with coxite) . . argentipes (p. 169)
Main lobe of paramere about twice length of middle lobe 5

5 Aedeagus without accessory spines tumenensis (p. 1 70)

- Aedeagus with accessory spines.

Lower lobe of paramere appearing narrow but extending mesally, depth of paramere about
0-35 of its length • 6

6 Antenna 3/labrum 1-7. Style 0-54 length of coxite, and more than four times as long as thick

philippinensis philippinensis (p. 170)
Antenna 3/labrum 2-0. Style 0-61 length of coxite, and about three times as long as thick

philippinensis gouldi (p. 170)

Phlebotomus (Euphlebotomus) argentipes Annandale & Brunetti

(Map 12)

Phlebotomus argentipes Annandale & Brunetti in Annandale, 1908: 101 [$ $]; Summers, 1911: 108; Patton
& Evans, 1929: 228. Lectotype & INDIA (ZSI, Calcutta), designated by Quate, 1962a: 157.

Phlebotomus (Euphlebotomus) argentipes Annandale & Brunetti; PernTev, 1968: 61 [egg]; Lewis, 19786: 240
[synonymy including annandalei, marginatus (apparently lost, according to Quate, 1962a: 157) and
zeylanicus <$ as synonyms] ; Artemiev, 1978: 24; Killick-Kendrick, 1978: 309 [variation].

? (extra facts). Leg formula (two from West Malaysia, Lamir) 100, 126, 79; 97, 154, 95; 107, 182, 135 (2-23,
0-85); 100, 129, 76; 98, 148,94; 107, 165, 104; (India, Ranighat) 100, 125, 75; 95, 151,93; 106, 180, 108.
cJ. Shaft of haltere 0-15 length of wing.

DISTRIBUTION. The Orient: Lewis (19786: 325, map). India: Modi et al. (1978: 748, Maharashtra, map).
Pandya et al. (1977: 133). The record from Iran by Javadian (1975: 207) was omitted by Artemiev (1978) and
appears to be incorrect.

NOTE. Lewis (19786: 323, 330, 331) summarized aspects of this species and reported geographical
variation which was partly associated with differences in feeding habits. Spiculate ascoids were
noted in India by S. Das (1973, in litt.). S. argentipes is an important vector of VL in India (Adler,
1964: 69; Bray, 1974: 93; Hoogstraal & Heyneman, 1969: 1186; PernTev, 1968: 141; Shanmu-
gham et al, 1977: 796; Theodor, 1964: 482; Wilcocks & Manson-Bahr, 1972: 122). Houses were
sprayed against malaria vectors in the early 1950s but P. argentipes seemed likely to flourish in
neighbouring vegetation (Anonymous, 1955). Kala-azar, greatly reduced during the anti-malarial
campaign, broke out on a large scale in 1977 (Killick-Kendrick, 1978: 306) when both spraying
and treatment seemed necessary to cope with it. By 1978 control was not complete (Anonymous,
1978).

Phlebotomus (Euphlebotomus) caudatus Artemiev

(Map 12)

Phlebotomus (Euphlebotomus) caudatus Artemiev, 1978: 25; 1979: 17 [<£). Holotype <$, AFGHANISTAN:
Farah-Rud area (BMNH) [examined].

NOTE. In Afghanistan P. caudatus occurs in low desert mountains (Artemiev, 1978: 25).

170 D. J. LEWIS

Phlebotomus (Euphlebotomus) kiangsuensis Yao & Wu
(Map 12)

Phlebotomus kiangsuensis Yao & Wu, 1938: 527 [9 <J]. Holotype^, CHINA (CFHS, Nanking?).
Phlebotomus (Euphlebotomus) kiangsuensis Yao & Wu; Lewis, 1978ft: 244 [synonymy].

DISTRIBUTION. China, Taiwan and West Malaysia: Lewis (1978: 245). China: Leng(1978: 7, no. 10 on map).

Phlebotomus (Euphlebotomus) mesghalii Rashti & Nadim
(Map 12)

Phlebotomus (Euphlebotomus) mesghalii Rashti & Nadim, 1970: 145 [9 <?]; Artemiev, 1978: 24. Holotype^,
IRAN (IPH, Tehran).

In the original description the names mesghali and mesghalii were both used, obviously for the
same taxonomic unit. As first reviser, within the meaning of ICZN (1964), Article 24 (a) (i), I here
select mesghalii as the name which will ensure stability of nomenclature.
A possible variant in Afghanistan is provisionally classed as species 2 by Artemiev (1978: 25).

Phlebotomus (Euphlebotomus) philippinensis Manalang

Phlebotomus philippinensis Manalang, 1930: 175.

Phlebotomus (Euphlebotomus) philippinensis gouldi Lewis
(Map 12)

Phlebotomus (Euphlebotomus) philippinensis gouldi Lewis, 19786: 245 [9 ^]. Holotype?, THAILAND (BMNH)
[examined].

Phlebotomus (Euphlebotomus) philippinensis philippinensis Manalang

(Map 12)

Phlebotomus philippinensis Manalang, 1930: 175 [9 <J]. Syntypes 9 <J, PHILIPPINES (CA, Los Banos (?) but

destroyed according to Quate & Rosario, 1962: 787, 789, 791).
Phlebotomus (Euphlebotomus) philippinensis philippinensis Manalang; Lewis, 1978ft: 245 [synonymy].

9 (extrafact). Leg formula 100, 133, 89; 111, 155, 100; 111, 178, 111.
DISTRIBUTION. Philippines: Lewis (1978ft: 325, map).

Phlebotomus (Euphlebotomus) tumenensis Wang & Chang
(Map 12)

Phlebotomus tumenensis Wang & Chang, 1963: 511 [9 £|. Syntypes? <J, CHINA: Tumen, 31°46'N, 104°06'E

(PIPD, Shandong).
Phlebotomus (Euphlebotomus) tumenensis Wang & Chang; Artemiev, 1979: 19.

Subgenus ANAPHLEBOTOMUSTheodor

Phlebotomus subgenus Anaphlebotomus Theodor, 1948: 99; Perfil'ev, 1968: 67, 78, 83; Hennig, 1972: 53;
Lewis, 1978ft: 247. Type-species : Phlebotomus stantoni Newstead, 1914, by original designation.

Key to the species of subgenus Anaphlebotomus

Females

1 Spermatheca long (about eight times as long as wide) and tubular with very long duct. Afrotropi-

cal • rodhaini(p. 170)

Spermatheca not long and tubular. Oriental 2

THE GENUS PHLEBOTOMUS 171

2 Spermatheca slightly carrot-shaped with small end-segment, individual duct about lour (possibly

more) times length of spermatheca.

Sternal tubercle broad colabaensis (p. 171)

Spermatheca spindle-shaped with very narrow cylindrical apical segment, duct short but
common duct very long.
Ascoids long. Palp 3 with peg sensilla grouped around middle 3

3 Pharyngeal armature with antero-median numerous long pointed teeth which blend laterally

with ridges. Individual ducts longer than spermathecae stantoni (p. 172)

- Pharyngeal armature with several antero-median rows of small short teeth, and antero-laterally a

number of backward-pointing teeth. Individual ducts shorter than spermathecae . hoepplii (p. 171)

Males

1 Paramere bilobed colabaensis (p. 171)

Paramere trilobed 2

2 Plunger of sperm pump much wider than body of barrel. Afrotropical .... rodhaini (p. 171)
Plunger of sperm pump narrower than body of barrel. Oriental 3

3 Spine near aedeagus not longer than it. Pharynx with a series of oblique ridges radiating from

mid-line and ending in loops laterally stantoni (p. 172)

Spine near aedeagus much longer than it. Pharynx with a series of posterior ridges and, antero-
laterally, a number of teeth projecting medio-posteriorly hoepplii (p. 171)

Phlebotomus (Anaphlebotomus) colabaensis Young & Chalam

(Map 13)

Phlebotomus colabaensis Young & Chalam, 1927: 859 [£]. Holotype^, INDIA (CSI Kasauli, now in NICD,

Delhi?).
Phlebotomus (Anaphlebotomus) colabaensis Young & Chalam; Lewis, 19786: 247 [references including one

to description of $ by Sinton in 1933].
Phlebotomus (Anaphlebotomus) colobaensis Young & Chalam ; Artemiev, 1978: 24. [Mis-spelling.]

DISTRIBUTION. India & Pakistan: Lewis (19786: 326, map). India: Delhi (1979, S. J. R.); Modi et al. (1977: 3;
1978: 748, map of Maharashtra).

Phlebotomus (Anaphlebotomus) hoepplii Tang & Maa
(Map 13)

Phlebotomus hoepplii Tang & Maa, 1945 : 25 [$ £]. Holotype & CHINA (TM).
Phlebotomus (Anaphlebotomus) hoepplii Tang & Maa; Lewis, 19786: 247 [references].

DISTRIBUTION. China: Leng (1978: 7, no. 9 on map); Lewis (19786: 326, map).

Phlebotomus (Anaphlebotomus) rodhaini Parrot
(Map 13)

Phlebotomus rodhaini Parrot, 1930a: 187 [<J]; 19306: 103; Kirk & Lewis, 1947: 875; Heisch & Guggisberg,
1952: 428. Holotype <J, ZAIRE (MRAC, Tervuren).

Phlebotomus (Phlebotomus) rodhaini Parrot; Parrot, 1948: 127 [?]; Kirk & Lewis, 19466: 120; 1951: 437;
Minter, 1963: 490; Quate, 1964: 245; Abonnenc & Minter, 1965: 32; Abonnenc, 1967: 70; 1972: 108.

Phlebotomus grenieri Rageau, 1951 : 796 [$]. Syntypes 2 $, CAMEROUN (IP, Paris). [Synonymized by Abon-
nenc, 1967:70.]

Phlebotomus (Anaphlebotomus) rodhaini Parrot; Qutubuddin, 1962: 597.

DISTRIBUTION. Africa: Abonnenc (1972: 251, map). Congo: Vattier-Bernard & Bimangou (1974: 105). Benin
Republic and Togo: Abonnenc (1973: 190, map). Ethiopia: Ashford (1974: 610); Gemetchu et al (1977: 209).
Gambia: Snow (1979: 245). Guinea: Abonnenc & Clastrier (1974: 61). Kenya: Minter (1964: 407, map);
Kiboko (D. M. M., record of 1980); Wijers & Ngoka (1974: 26). Senegal: (1977, J. P. D). Sudan: Hoogstraal
& Heyneman (1969: 1155); Lewis & Kirk (1954: 35, map); Qutubuddin (1962: 597, Gedaref). Uganda:
Wykoff etal. (1969: 206).

172 D. J. LEWIS

NOTE. This species apparently feeds mainly on rodents (Abonnenc, 1972: 110) and has been
known to bite man (Ashford, 1974: 610; Ashford et at, 1973: 261; Hoogstraal & Heyneman,
1969:1156,1170).

Phlebotomus (Anaphlebotomus) sp. D

(Map 13)
This species is being described by Dr I. H. Davidson.

Phlebotomus (Anaphlebotomus) stantoni Newstead
(Map 13)

Phlebotomus stantoni Newstead, 1914: 190 [?]; Leng, Liu, Huang & Liu, 1979: 189. Holotype ?, WEST
MALAYSIA (BMNH) [examined].

Phlebotomus (Anaphlebotomus) stantoni Newstead; Lewis, 1978ft: 248 [synonymy including maynei as syn-
onym and description of $ P. stantoni by Raynal in 1934].

DISTRIBUTION. China and Orient: Lewis (19786: 326, map). China: Leng (1978: 7, no. 8 on map); Leng et al.
(1979: 189, Ch'ang-chiang ( = Zhanjiang)).

Subgenus KASA ULIUS subgen. n.

Type-species : Phlebotomus newsteadi Sinton.

DIAGNOSIS. Pharynx of $ with well-developed teeth pointing backward. Antenna 3 long, 0-39 mm or more in
9 and 0-52 or more in <J; two ascoids on segments 3-15 in 9, and 3-10 in^. Palp formula 1 (2, 4), 3, 5 in$
and 1, (4, 2), 3, 5 in £. Mesanepisternum with two lower hairs. Legs very long, in <$ tibia 3 = 2-18, and
basitarsus 3 = 1-36, as long as femur 1. Wing narrow, in 9 about 4-3, and in $ 4-7, as long as wide; index
1-94-2-14 in 9 and 1-5-2-3 in £. Haltere of $ with broad stalk, hind half marked off by a furrow and having a
projecting group of large sensilla. Spermatheca moniliform with about 25 segments, the end one large.
Genital filaments about twice length of pump. Aedeagus narrow, with two accompanying pointed rods.
Paramere with truncated end having four small ventral serrations, a few hairs on a small projection, and an
accessory spine. Coxite without group of hairs. Style with five spines.

DISCUSSION. Theodor (1948 : 108) placed the single species provisionally in the subgenus Euphlebo-
tomus although the description of the paramere did not indicate this, and Artemiev (1979: 19)
considered that P. newsteadi was closest to Euphlebotomus in view of the lateral spine of the
aedeagus and the shape of the paramere, coxite and style. The unique nature of this species is
emphasized by its narrow wings and by comparison of its leg-segments with those of other
species, and a new subgenus is therefore proposed for it. It is given a territorial name because P.
newsteadi, though discovered 60 years ago, is known only from the one place.

Phlebotomus (Kasaulius) newsteadi Sinton
(Map 13)

Phlebotomus newsteadi Sinton, 1926: 559 [<£]; Lewis, 19786: 250 [synonymy including description of 9 by
Sinton in 1928]; Artemiev, 1978: 24; 1979: 19 [related to Euphlebotomus^. Lectotype^, INDIA (BMNH),
designated by Lewis, 19786: 250 [examined].

$ (extra facts). Labrum 0-28 mm long, 0-59 length of head, 0-09 length of wing. Antenna 4 and 5 with papilla.
Two lower mesanepisternal hairs present. Leg formula 100, 176, 119; 93, 194, 125; 106, 218, 136 (3-03, 0-20).
Wing length 2-93 (2-84-3-03) mm long, 4-7 times width. Haltere shaft 0-19 length of wing.

MATERIAL EXAMINED
India: 1 <J, Kasauli, 5.viii.l928, garden house (J. A. S.) (RSTMH, London).

NOTE. The rarity of this species and the relative lengths of the hind tibia and the basitarsi suggest
that it may be a cave form seen rarely outside caves.

Nomen nudum

Phlebotomus algeriensis Jenkins, 1964: 31.

THE GENUS PHLEBOTOMUS

173

Discussion

Leg ratios (Figs 1 5-24)

The relative length of tibia 3 of a species was taken as an indicator of leg length, and species with
a tibia 3 length of 165 units or less were classed as short-leg species, and the rest as long-leg
species. For estimating the mean lengths in subgenera, figures for species were placed in working
groups of 121-130, 131-140 and so on.

Phlebotomites brevifilis Hennig (120 MYA)

The leg formula appears to be 100, 132, 53; 143, 143, 64; 117, 168, 68 (0-41), with the three tibiae

and the basitarsi approaching equality and tibia 1 and femur 2 very long.

Genus Warileya

The leg formula of the male of W. nigrosacculus is 100, 132, 70; 100, 136, 72; 106, 138, 74 (0-47),
each leg being like the others, and tibia 3 very short. The remarkable degree of uniformity of this
species may be related to that shown by the fossil forms.

Genus Phlebotomus

Out of 55 species studied the length of tibia 3 ranged from 155 to 218, and there were eight (14 per

cent) short-leg species and 47 (86 per cent) long-leg species.

P'tes Warileya P.

brevifilis nigrosac- tipuli-

culus formis

15

P.

minteri

h6

'17

200

0.150

100

.t: 50

Figs 15-24 The relative lengths of long segments of each leg of 10 species of Phlebotominae.

174 D. J. LEWIS

Phlebotomus tipuliformis (Meunier) (30 MYA). The leg formula (Hennig, 1972: 52) appears to
be 100, 137, 112; 128, 178, 133; 123, 220, 133 (0-49). All tibiae are long and tibia 3 very long, and
the basitarsi longer than the femora and about equal to each other. If the measurements of flies in
amber are reliable the two fossil species show a tendency to uniformity.

Subgenus Spelaeophlebotomus. One species was measured. Tibia 3 = 211. All tibiae and basi-
tarsi are very long, probably as an adaptation to caves.

Subgenus Idiophlebotomus. Six species were measured. Tibia 3 = 197 (182-216). The tibiae and
basitarsi show a tendency to uniformity, probably a primitive feature. Femur 1 is always longer
than femur 2, and this and the long tibia 3 may represent an adaptation to life in caves.

Subgenus Australophlebotomus. Two species were measured. Tibia 3 = 155 (155-156).

Subgenus Phlebotomus. Four species were measured. Tibia 3 = 171 (163-175).

Subgenus Paraphlebotomus. Eight species were measured. Tibia 3 = 172 (153-194). In P. alex-
andri each tibia is shorter than in other species, and basitarsus 1 is shorter than femur 1 (longer in
all other species). All the tibiae of P. nuri are longer than those of other species studied. Measure-
ments of P. sergenti from Corsica and Morocco (Croset et a/., 1974: 107) and the U.S.S.R.
indicate local variation.

Subgenus Synphlebotomus. Six species were measured. Tibia 3 = 165 (157-172) and there is
little specific variation.

Subgenus Larroussius. Seventeen species were measured. Tibia 3 = 181 (147-212), one being
below 150 and two above 200.

Subgenus Adlerius. Six species were measured. Tibia 3 = 182 (163-202).

Subgenus Euphlebotomus. Three species were measured. Tibia 3 = 180 (176-185).

Subgenus Anaphlebotomus. Four species were measured. Tibia 3 = 179 (167-189).

Subgenus Kasaulius. In the one species tibia 3 = 218.

Other genera

Records of 27 species of Sergentomyia show a variation of 124-229 in the length of tibia 3, even in
this small sample, with 18 short-leg and nine long-leg species. Published records of three species
of Brumptomyia show an average length of 184 (178-190) for tibia 3. Published records of 100
species, of 30 subgeneric groups, of Lutzomyia give a tibia 3 length of 136 to 230 units, with 42
short-leg and 58 long-leg species. Among groups represented by five or more species, the number
of species considered and the mean length of tibia 3 were: Psychodopygus Mangabeira, 8, 169;
Pressatia Mangabeira, 5, 161; Helcocyrtomyia Barretto, 9, 172; Psathyromyia Barretto, 7, 196;
verrucarum-group, 10, 155.

Conclusion

The records for P. papatasi and P. caucasicus suggest that leg measurements differ little between
the sexes, and the former shows little variation in a particular area. Extinct and some primitive
species (Phlebotomites, Warileya, P. tipuliformis, Spelaeophlebotomus and Idiophlebotomus) show
a trend towards equality of segment 1, 2 or 3 in successive legs. Most species of Phlebotomus show
a pattern in which each femur, tibia and basitarsus is longer than the preceding one, and each
tibia is somewhat longer, and each basitarsus much shorter, than its femur.

The length of tibia 3 in Phlebotomus tends to be greater than in Sergentomyia and Brumpto-
myia, and varies more than in Brumptomyia and less than in Sergentomyia. In all three genera the
length of tibia 3 apparently constitutes a more or less clear-cut feature of several subgeneric
groups, and in the case of Kasaulius it was a factor in deciding to treat this group as a subgenus.
Certain species can be distinguished from others by the length of tibia 3. Some species with a
similar tibia 3 length differ in the form of another leg, which may be useful if the others are
missing. It is therefore worth while to examine all three legs, and to follow up differences
suggested by leg diagrams. Infra-specific local variation seems to occur in at least one species (P.
sergenti). These findings suggest that the measurement of all long leg segments of at least one
female of a sandfly species yields enough information to be worth doing. Further work could
provide averages instead of measurements of individual flies, indicate the length of tibia 3 in
additional species, and evaluate any apparently distinctive specific characters.

THE GENUS PHLEBOTOMUS 175

Evolution of Phlebotominae

It is now possible to speculate on the history of Phlebotomus and some other sandflies on the
basis of fossils, palaeogeography, morphology and distribution. The following remarks refer to
the Old World unless the New is mentioned.

Old and New World sandfly faunas

Before 120 MYA Phlebotominae had probably lived for a long time in Pangaea, and possibly fed
on insects before taking to vertebrate blood (Rohdendorf, 1974: 58). After the opening of the
south Atlantic about that time the two sandfly faunas of Africa and South America probably
evolved separately. It is not known whether this separation continued to the present day, because
the north Atlantic did not open (Papavero, 1977: 212) till perhaps 60 MYA or later, there were
probably land connections in the Bering area and elsewhere from time to time, and the effects of
past polar and climatic changes on sandfly distribution are unknown. After the development of
Phlebotomus its species probably did not extend far enough into the cold north to cross the
Bering land bridge (PerfiTev, 1968: 90). The present differences between Old and New World
sandflies, though somewhat indefinite, are enough to suggest that isolation has prevailed for the
last 120 MYA.

The proboscis in relation to reptile and mammalian hosts

According to present evidence nearly all living species of sandflies are either reptile or mammal
feeders. In general the reptile feeders are distinguished morphologically as ridge-tip species,
according to the shape of the maxilla and other stylets, and have a short labrum, not more than
0-23 mm long (Lewis, 1975: 507, 520, 525). Mammal feeders are hook-tip species, usually with a
labrum 0-24 or more in length. Species of Sergentomyia typically have a labrum about 0-11
(0-09-0-14) as long as the wing, and Phlebotomus about 0-14 (0-11-0-22) as long, and the differ-
ence is usually accentuated by the greater size of species of Phlebotomus. Current studies on the
mouth parts of early fossil-piercing psychodids, living in the age of reptiles and presumably
ridge-tips, had a short labrum. It is likely that in reptile feeders the labrum has remained short,
and that in mammal feeders it has progressively lengthened during the rise of mammals, so that
now the two groups have a somewhat midge-like and mosquito-like facies respectively.

Wing shape

Wings vary from being very broad with round ends to being very narrow with almost pointed
tips. There is no clear distinction between narrowly rounded and bluntly pointed tips, but there is
generally little difficulty in dividing species as follows among four groups designated by typical
values of wing length divided by width, although in a doubtful case a subjective view of shape
may be better than exact measurement: very wide (3-2, Figs 26-30), wide (3-5, Figs 32-35),
narrow (3-9 Fig. 36) and very narrow (5-5, Figs 37, 38).

Very wide wings are seen in Phlebotomites species and Phlebotomus tipuliformis, in the primi-
tive Warileya, in the subgenera Spelaeophlebotomus and Idiophlebotomus, and in some species of
subgenus Australophlebotomus. Except in this subgenus the wings are of a primitive type with the
origin of R4 near that of R5 so that gamma is short (Abonnenc, 1972: 75). The closely related
Spelaeophlebotomus and Idiophlebotomus share an archaic wing structure (Abonnenc, 1972: 74,
75). An inter ocular suture (Young, 1979: 5), presumably a primitive character, occurs (Lewis et
al, 1977: 326) in Warileya and Spelaeophlebotomus. Idiophlebotomus and most species (Young &
Chaniotis, 1973; Lewis et al, 1977: 325) of Warileya share the unusual feature of rods near the
sperm pump.

In all other phlebotomines the origin ofR2 + 3 has become distally displaced, the wing less wide,
and its tip less rounded, possibly in relation to the development of a hairy fringe (Hennig, 1972: 8,
27). The first known narrowing occurred, in America, at least 26 MYA.

The wing tends to be wide, with R2 longer than R2 + 3 (Theodor, 1958: 24, 33, 48; PernTev,
1968: 26, more or less oval) in Phlebotomus, in subgenus Neophlebotomus Franca & Parrot and
some ungrouped and other species of Sergentomyia, and in Brumptomyia and Lutzomyia. In-
cluded in Phlebotomus are the pair of closely related subgenera Euphlebotomus and Anaphlebo-
tomus which somewhat resemble (Hennig, 1972: 53-55) the ancient P. tipuliformis, and their type

176

D. J. LEWIS

32

33

120 MYA

P. (Paraphlebotomus) $

27

Phlebotomus tipuliformis

29

P. (Spelaeophlebotomus)

30

P. (Idiophlebotomus) ?

34

35

Sergentomyia (Neophlebotomus)

S. (Sergentomyia)

S. (Sergentomyia) ?

26 MYA

38

S. (Parvidens) ?

Figs 25-38 Wings of Permotipula and of examples of 13 groups of Phlebotominae to illustrate evol-
utionary trends.

THE GENUS PHLEBOTOMUS 111

of spermathecal duct suggested to Perfil'ev (1968: 83) that they are transitional between Sergento-
myia and Phlebotomus. Their long parameral spines and those of Kasaulius may (Artemiev, 1979:
18) be homologous with the intra-abdominal rods of the primitive Warileya and Idiophlebotomus.
In three subgenera of the wide-wing group, Synphlebotomus, Euphlebotomus and Anaphlebotomus,
the mean length of the labrum is less than in Phlebotomus, Paraphlebotomus, Larroussius and
Adlerius, and may indicate an earlier stage in evolution.

The wing is often narrow, lanceolate and nearly pointed, with R2 shorter than R2 + 3 (Fairchild,
1955: 183; Theodor, 1958: 4; Perfil'ev, 1968: 26, 27, 295) in subgenera Sintonius Nitzulescu,
Grassomyia Theodor, Sergentomyia, Parrotomyia Theodor and some ungrouped species of genus
Sergentomyia.

Very narrow wings are seen in a few species of genus Sergentomyia.

It appears that the phlebotomine wing has gradually narrowed, particularly in Sergentomyia
and mainly in its savanna species, so that now in the Old World reptile and mammal feeders
usually have narrow and wide wings respectively.

Widespread and northern groups of Phlebotomus

If distribution is considered in relation to the above discussions on labrum length and wing shape
it is possible to postulate two evolutionary groups of Phlebotomus, the widespread and northern
groups. The former includes all except subgenus Phlebotomus and virtually all Paraphlebotomus,
Adlerius and Larroussius, and the latter group comprises most species of these subgenera.

The widespread group shows a number of primitive or apparently primitive features, and in
general is widely distributed (Maps 1, 5, 12, 13). The northern group seems to exhibit the ultimate
stage in lengthening of the labrum, and has an extraordinarily north-western and restricted
distribution in the Old World (Lewis, 1974: 364; 1978a: 97; Maps 2-4, 6-11) in relation to other
Phlebotominae (the widespread group and Sergentomyia, Map 1), with little specific variation in
distribution. It corresponds roughly with the distribution of moles, hedgehogs and jerboas (Bar-
tholomew et al, 1911) and also (Ellerman, 1950) of the gerbil genera Meriones, Psammomys and
Rhombomys with which some sandflies are associated. Gerbils appeared late in geological time
(Chaline, 1977) like many rodents (Petrishcheva, 1971: 569). The large northern group includes
most of the Old World vectors of mammalian leishmaniasis.

It is suggested that the northern group increased in numbers of individuals, and in species
except in subgenus Phlebotomus, in late geological times in the north-west, and that during the
pluvial periods (Banister & Clarke, 1977: 147, 151; Roberts, 1975: 276) a few species extended
into Ethiopia, Yemen and China. P. orientalis, closely related to the Mediterranean P. langeroni,
may be a product of this process, and its present extension into the Sudan, but not far into West
Africa (Map 8), could be a stage in its progress. In China species of Phlebotomus (Maps 3, 4, 10)
may now be cut off from the west unless there is a connection along a corridor south of Mongolia
(PernTev, 1968: 96). Further spread of the northern cold- winter group may have been prevented
by climatic factors and by natural barriers like the Sahara which has existed for a long time
(Corbet, 1967: 334). The distribution of P. papatasi and P. duboscqi differs somewhat from that of
the others, their subgenus is unusually small, and (Hennig, 1972: 53) they may be related to
Euphlebotomus, so their history may possibly be different.

Aspects of leishmanial evolution in relation to that of Phlebotominae

The availability of a few sandfly fossils and knowledge of the present distribution of leishmaniasis
make it possible to speculate, as follows, on the history of Leishmania in a way that may have
some bearing on the classification of the genus.

Before 120 MY A

In the ancient continent of Pangaea Phlebotominae may have acted as primary hosts of leish-

maniae of reptiles for a long period.

120 to 20 MY A

In the New World some 120 MYA leishmaniae probably began to evolve in isolation from the
Old Word, and this separation may have been continuous, or nearly so, till historic times and
accounted for the considerable difference (Chance et al., 1977: 59) between the parasites of Old

178

D. J. LEWIS

THE GENUS PHLEBOTOMUS

179

o.

55

180

D. J. LEWIS

THE GENUS PHLEBOTOMUS

181

182

D. J. LEWIS

THE GENUS PHLEBOTOMUS

183

D. J. LEWIS

OS

5

THE GENUS PHLEBOTOMUS

185

90

O.

186

D. J. LEWIS

a.

08

THE GENUS PHLEBOTOMUS

187

i

188

D. J. LEWIS

ca

5

THE GENUS PHLEBOTOMUS

189

a,

S3

190

D. J. LEWIS

THE GENUS PHLEBOTOMUS 191

and New World CL at the present day. Mammalian leishmaniae may have developed separately
in the Old and New Worlds during this period. Leishmaniae may have been absent from Aus-
tralia some 120 MYA when, with Antarctica, it separated from the rest of the Old World.

In the Old World reptile leishmaniae presumably continued to exist and, with the appearance
of the genus Phlebotomus, mammalian leishmaniae may have developed and become widespread,
with no special relation to Central Asia (Bray, 1974: 95).

20 M YA to the present day

The increase, postulated above, of the subgenera Phlebotomus, Paraphlebotomus, Larroussius and
Adlerius in the north of the Old World may have aided the development of mammalian leish-
maniae, and thus account for the fact that much of the Old World human leishmaniasis is in
northern latitudes, where at least some forms of VL and CL are thought to have arisen. The VL
of Kenya and eastern India may have been secondary developments (Garnham, 1977). The vector
in India, the only habitual man-biter in the widespread group of subgenera, is the western form of
P. argentipes, which has adopted peridomestic habits (centred on cattle) in an area of dense
human population and consequent destruction of many natural resting sites and wild animals.

Man had become numerous in the Old World about one MYA, long before his arrival in the
New World, and in due course a few sandfly species became more or less domestic and therefore
more effective vectors of leishmaniae.

The above hypotheses on sandfly and leishmanial evolution, based on circumstantial evidence
and conjecture, and postulating an early split between Old and New World sandflies, and a late
expansion of Phlebotomus in the north of the Old World, offer an explanation of some features of
present-day distribution of leishmaniae.

Acknowledgements

I am very grateful to the Medical Research Council for grant support; the Keeper of En-
tomology, British Museum (Natural History), for providing facilities; Professor A. Corradetti, Dr
W. J. Le Quesne, Dr Ruth Lichtenberg, Professor J. A. Rioux and Dr C. W. Sabrosky for
information about some early records and nomenclature; Dr O. Escola and Dr J. Gil Collado for
information about the location of types; Dr M. M. Artemiev, Dr J. P. Dedet, Mr J. Lane and Dr
D. M. Minter for a number of specimens; Dr I. H. Davidson, Mr A. L. Dyce and Professor Leng
Y.-j. for advice about species D, Australian species and T. major wuf ; Miss Carolyn Couch for
technical assistance; Dr P. E. S. Whalley for advice about fossils; and Mr P. M. Hammond and
Miss Ann Lum for translating Chinese.

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Index

Invalid names are in italics; principal references are in bold.

A sp. 130, 135

aculeatus 131, 151, 152, 153

Adlerius 122, 126, 127, 131, 163, 174, 177, 191

alexandri 127, 139, 142, 143, 146, 147, 174

algeriensis 172

Anaphlebotomus 125, 130, 137, 170, 174, 175, 177

andrejevi 141, 142, 143, 144

angustus 131, 164

annandalei 169

ansarii 127, 148, 149

arabicus 164, 166

argentipes 127, 139, 168, 169, 173, 191

ariasi 127, 151, 152, 153

asperulus 133, 134

Australophlebotomus 130, 135, 174, 175

autumnalis 130

avellari 173

B sp. 130, 136
balcanicus 164, 165
bedfordi 124
bergeroti 127, 137
betisi 129, 131, 151, 153
Bibio 129, 138, 140

brevifilis, Phlebotomites 124, 125, 173

brevifilis, Phlebotomus 135, 136

brevifiloides 135, 136

brevis 130, 164, 165

breviventris 139

Brumptomyia 122, 124, 125, 173, 175, 176

buccinator 129, 136

burneyi 151, 154

C sp. 130, 136

canaaniticus 151, 152, 158

caucasicus 127, 141, 142, 144, 145, 174

caudatus 130, 169

celiae 127, 148, 149

chabaudi 127, 142, 143, 145

chadlii 130, 152, 153

chinensis 123, 125, 127, 130, 163, 164, 165

Ciniphes 138, 141

clydei 173

colabaensis 131, 171

colobaensis 131

comatus 130, 164, 166

crimicus 147

Cyniphes 138

208

D. J. LEWIS

D sp. 130, 131, 172
Dampfomyia 124, 176
davidi 164, 166
duboscqi 127, 137, 138, 177
duboscqui 138
duboscquii 138
dubosqi 138

eleanorae 148, 149
elgonensis 153
erebicolus 133, 134

Euphlebotomus 122, 125, 126, 127, 130, 137, 168,
172, 174, 175, 177

fallax 124

fantalensis 130, 152, 154

Flebotomus 129

fourtoni 138

frondifer 124, 133, 134, 173

galilaeus 130, 151, 160, 161

galindoi 124

gibiensis 131, 151, 152, 154

gigas 131, 132, 133

gombaki 124

gouldi 169, 170

grassii 161

Grassomyia 177

grenieri 171

grimmi 144, 145

griseus 157

grovei 148, 149

guggisbergi 131, 139, 151, 152, 154

Haemasson 144
halepensis 127, 130, 164, 166
Hebotomus 138
Helcocyrtomyia 174
hindustanicus 131, 139, 164, 166
hoepplii 171

Idiophlebotomus 124, 130, 133, 174, 175, 176, 177
imitabilis 146
ismailicus 165

jacusieli 142, 143, 145

kabulensis 164, 167
kandelakii 127, 131, 151, 154, 155
Kasaulius 130, 172, 174, 177
katangensis 130, 148, 149, 150
kazeruni 142, 143, 145
keshishiani 151, 152, 155
kiangsuensis 131, 169, 170
krimensis 151, 152, 156
kyreniae 130, 164, 167

langeroni 130, 152, 155, 177
larroussei 158

Larroussius 125, 126, 127, 131, 150, 174, 177, 191

legeri 161

lesleyae 124

li 144

Hi 144

longicuspis 127, 131, 151, 152, 155

longiductus 124, 127, 164, 167

longifilis 125

longiforceps 133, 134

longipes 127, 151, 152, 156

lusitanicus 161

Lutzomyia 122, 123, 124, 125, 174, 175

macedonicus 160

major 124, 127, 130, 131, 141, 150, 151, 152, 155,

156, 157
marginatus 169
mariae 130, 152, 158
marts mortui 146
marismortui 142, 146
martini 127, 148, 149, 150
mascittii 125, 131, 150, 151, 152, 158, 159, 162
maynei 172
mesghali 170
mesghalii 169, 170
minteri 124, 131, 132, 133, 173
minuta 141
mofidii 144
molesta 138, 140, 141
molestus 138

mongolensis 127, 142, 143, 146
monticola 166
monticolus 166
Musca 138

neglectus 151, 152, 157
Nemopalpus 124
Neophlebotomus 124, 175, 176
newsteadi 172, 173
nigerrimus 161, 162
nigrosacculus 124, 173
nitzulescui 158
nuri 142, 143, 146, 174

orientalis 126, 127, 131, 151, 152, 159, 177

papatasi 123, 127, 129, 137, 138, 139, 141, 144, 147,

148, 174

papatasii 138, 141
pa^patasi 139
pappatasii 139
papuensis 130, 136
Paraphlebotomus 124, 127, 130, 142, 174, 176, 177,

191

Parrotomyia 177
Parvidens 124, 176
paterna 124, 125, 176
pedifer 127, 131, 151, 152, 159
perfiliewi 127, 130, 131, 151, 160

THE GENUS PHLEBOTOMUS

209

permira 124

perniciosus 125, 127, 151, 152, 159, 161, 163

pexopharynx 135, 136

Philaematus 125

philippinensis 169, 170

Phlebotomiella 125

Phlebotominae 122

Phlebotomites 124, 125, 174, 176

Phlebotomus 122, 123, 124, 125, 126, 127, 129, 130,

134, 137, 174, 175, 177, 191
pholetor 133, 134
pirumovi 162
Pressatia 174
Psathyromyia 174
Psychodopygus 122, 174
pungens 125

rodhaini 131, 138, 170, 171
rossi 127, 148, 149, 150, 173
roubaudi 138
rupester 131, 164, 167

saevus 142, 143, 147

saheli 141

salangensis 164, 168

salehi 127, 137, 141

sejunctus 129, 133, 135

selectus 144

sergenti 123, 124, 127, 141, 142, 143, 145, 146, 147,

148, 174
Sergentomyia 122, 123, 124, 125, 134, 138, 173,

175, 176, 177
simici 127, 131, 164, 168
similis 142, 143, 148

Sintonius 177

smirnovi 131, 151, 152, 158, 162

somaliensis 129, 131, 151, 162

Spelaeophlebotomus 124, 130, 131, 174, 175, 176

stantoni 131, 170, 171, 172

stellae 133, 134, 135

succini 125

Synphlebotomus 126, 127, 130, 148, 174, 177

syriacus 151, 152, 157

tanriae 167

tauriae 157, 167

tauricus 156

teshi 129, 133, 135

tipuliformis 124, 125, 137, 173, 174, 175, 176

tobbi 127, 131, 151, 152, 162

transcaucasicus 151, 161

trifilis 130, 136, 137

tubifer 129, 133, 135

tumenensis 168, 169, 170

turanicus 164, 168

vansomerenae 127, 148, 149, 150
verrucarum 174
vesuvianus 158
viduus 137

Warileya 122, 124, 125, 173, 174, 175, 176, 177
wenyoni 131, 151, 152, 155, 163
wui 124, 130, 157

zeylanicus 169
zulfagarensis 164, 168

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Stenomine moths of the Neotropical genus Timocratica (Oecophoridae).
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Stenomine moths of the Neotropical genus
Timocratica (Oecophoridae)

Vitor O. Becker

Entomology series

Vol 45 No 3 26 August 1982

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ISSN 0524-6431

British Museum (Natural History) 2 6AUG T

Entomology series
Vol45No 3 pp 2 11-306

Issued 26 August 1982

Stenomine moths of the Neotropical genus

T" ^« /r\ L «J \ t

Timocratica (Oecophoridae)

Vitor O. Becker

Centre de Pesquisa Agropecuaria dos Cerrados, Caixa postal, 70-0023, 73300-Planaltina, DF,
Brazil

Contents

Synopsis ............... 211

Nomenclatural summary ............ 211

Introduction ............... 213

Nomenclatural history ............. 213

Material and methods ............. 213

Abbreviations of depositories ........... 216

Colour-pattern and defence ............ 216

Geographical and ecological distribution ......... 217

Classification of Timocratica ............ 217

Cladistic analysis ............. 217

Phenetic analysis ............. 221

Timocratica Meyrick ............. 225

Key to species and subspecies ........... 228

Division of Timocratica into species-groups ... ..... 230

The monotonia-group ............ 230

The leucocapna-group ............ 238

The albella-group ............. 240

Species transferred from Timocratica .......... 271

Acknowledgements ............. 272

References ............... 273

Index ......... ....... 305

Synopsis

The genus Timocratica Meyrick is revised and a key to the 46 species is provided together with distribution
maps, illustrations of the male and female genitalia, and cladistic and phenetic analyses. Biological data and
descriptions of the larvae and pupae are given for T. palpalis (Zeller) and T. melanocosta sp. n. Seventeen
new species and one new subspecies are described, and 1 1 specific synonyms are newly established. Five
species previously included in Timocratica are provisionally transferred to Stenoma Zeller; S. butyrota
Meyrick (as comb, n.) and Lychnocrates leucocapna Meyrick (as comb, rev.) are transferred to Timocratica.
The genus is restricted to the Neotropical Region and ranges from the gulf area of Mexico to northern
Argentina. The species occur mainly in three Life Zones: Tropical Moist Forest, Tropical Premontane
Moist Forest, and Tropical Premontane Wet Forest. The larvae of palpalis and melanocosta are injurious to
various species of trees and bore into the trunks, feeding on the bark surrounding the entrance holes.

Nomenclatural summary

TIMOCRATICA Meyrick, 1912
Lychnocrates Meyrick, 1926
albella-group

albella (Zeller, 1839) Surinam

albitogata sp. n. Brazil

amst'li Duckworth, 1962 sp. rev. Venezuela

albella Amsel, 1956 (nom. preocc.)

Bull. Br. Mus. not. Hist. (Ent.) 45 (3): 211-306 Issued 26 August 1982

212

V. O. BECKER

anelaea (Meyrick, 1932)
argonais (Meyrick, 1932)

argonias Clarke, 1955 (misspelling)
bicornuta sp. n.
hut yr ota (Meyrick, 1929) comb. n.

syndicastis (Meyrick, 1929) syn. n.
constrictivalva sp. n.
fuscipalpalis sp. n.
grandis (Perty, [1833])
guarani sp. n.
isarga (Meyrick, 1925)
leucorectis (Meyrick, 1925)
macroleuca (Meyrick, 1932)
mature scens (Meyrick, 1925)
megaleuca (Meyrick, 1912)
melanocosta sp. n.
melanostriga sp. n.
nivea sp. n.
palpalis (Zeller, 1877)

auxoleuca (Meyrick, 1925)

haywardi Busck, 1939 syn. n.
parvifusca sp. n.
parvileuca sp. n.
philomela (Meyrick, 1925)
spinignatha sp. n.
subovalis (Meyrick, 1932)

stomatocosma (Meyrick, 1932) syn. n.
titanoleuca sp. n.
venifurcata sp. n.
xanthosoma xanthosoma (Dognin, 1913)

sacra (Meyrick, 1918)
xanthosoma leucocephala subsp. n.
xanthotarsa sp. n.
species 3
species 4
species 5
species 6
species 7

leucocapna-group
effluxa (Meyrick, 1930)
leucocapna (Meyrick, 1926) comb. rev.
species 2
monotonia-group
agramma sp. n.
fraternella (Busck, 1910)
longidlia sp. n.
loxotoma (Busck, 1909)
major (Busck, 1911)
meridionals sp. n.
monotonia (Strand, 1911)

isographa Meyrick, 1912 syn. n.

claudescens Meyrick, 1925 syn. n.

crassa Meyrick, 1925 syn. n.
pompeiana Meyrick, 1925
species 1

Brazil

Brazil, Guyana, French Guiana

Brazil

Colombia, Costa Rica, Panama, Peru

Ecuador

Venezuela

Brazil, French Guiana, Panama

Argentina, Paraguay

Bolivia

Bolivia, Brazil, French Guiana, Colombia, Peru

Bolivia

Colombia, French Guiana, Venezuela

Colombia

Brazil

Brazil

Brazil

Argentina, Bolivia, Brazil

Costa Rica

Brazil

Peru

Peru

Brazil

Peru

Brazil

French Guiana

Colombia, Panama

Panama

Peru

French Guiana

Brazil

Brazil

Colombia

Bolivia

Colombia, Costa Rica, Peru, Venezuela

Peru

Brazil

Costa Rica

Colombia

Costa Rica, Guatemala, Mexico

Brazil, Peru

Brazil, Paraguay, Bolivia

Brazil, Guyana, Colombia, Ecuador

Peru
Costa Rica

NEOTROPICAL GENUS TIMOCRATICA 213

Introduction

The genus Timocratica includes the largest known species of Oecophoridae in the world. The
females of some species, such as leucorectis, have a fore wing length of up to 32 mm, equivalent to
about 80 mm wing-span. According to the male and female genitalia, it is a very homogeneous
group, but externally the species show great variation, mainly in shape, venation and colour-
pattern of the fore wings.

Timocratica is confined to the Neotropical Region, ranging from the Gulf of Mexico in the
north, to the northern part of Argentina in the south. In South America it is almost restricted to
the eastern side of the Andes, with most of the species represented in the Amazonian Basin.

Despite its wide geographical distribution, the genus is confined ecologically to a few Life
Zones, chiefly to three: Tropical Moist Forest, Tropical Premontane Moist Forest, and Tropical
Premontane Wet Forest (Fig. 1).

During the last 10 years I have reared bark-feeding larvae from several different host-plants in
a number of localities, and found the white ' bark-feeder ' Timocratica albella sensu auctorum to
be a species complex. The vexed question of whether these white species and those described in
Lychnocrates were related to the fuscous species of Timocratica, as suggested by Busck, was still
unresolved. It was clear that a detailed revision of these groups was needed to solve this question
and to provide accurate descriptions and definitions of the bark-feeding species.

The species with dull fuscous fore wings belonging to the monotonia- and leucocapna-gioups
are presumably cryptic, while the white species of the albella-group are considered to be mimetic.
Possible models are the white species of the arctiid genus Agylla Walker, which are very abun-
dant and are sympatric with those of Timocratica. This hypothesis is supported by field tests in
which species of Agylla were rejected by birds. As Timocratica species are presumably not
distasteful to predators, they may form a Batesian mimetic group of Agylla.

Nomenclatural history

The genus Timocratica was proposed by Meyrick (1912) to accommodate his fuscous species
isographa; tristrigata Zeller and major Busck were included provisionally and later Meyrick
(1925) added three new species to the genus. In 1926 he described Lychnocrates for another
fuscous species, leucocapna, and added effluxa in 1930.

Although Meyrick described most of the white Timocratica species, he never considered them
to be congeneric with isographa since, according to him, they had veins 2 and 3 (CuAt and CuA2)
of the fore wings free, not stalked as is usual in the fuscous species. As pointed out by Busck
(1938: 283), the venation of the fore wings in this group (Timocratica sensu Busck, i.e. Timo-
cratica + Lychnocrates + the white species) is highly variable, particularly in the white species; it
varies not only between but also within species. Some of the white species also have CuAl and
CuA2 of the fore wings stalked as in isographa and related fuscous species.

Busck (1935) was the first to unite into one genus the fuscous species included here in the
monotonia-group, the white species and the fuscous Lychnocrates species, his decision having
been based on the similarity of the male and female genitalia. Clarke (1955) removed Lychno-
crates from synonymy with Timocratica, on account of the free CuAl and CuA2 of the fore wings,
but retained effluxa (described by Meyrick in Lychnocrates and undoubtedly congeneric with
leucocapna) in Timocratica.

Although colour-pattern and wing venation can be used to define three clearly distinct sub-
groups, these features are not sufficient to treat them as separate genera. In the Stenominae wing
venation seems to have little taxonomic value, and the structure of the male genitalia is the main
basis for generic division. As the genitalia of all the species discussed above are so similar, there is
little doubt that they constitute a monophyletic group and they are here regarded as a single
genus.

Material and methods

About 500 adult specimens of Timocratica were examined, representing 46 species. Of these
about 250 are from my collection, 200 are from the BMNH collection, and 50 are from the

214 V. O. BECKER

NMNH and other institutions indicated in the text. Although my collection contained half of the
specimens, these represented only one-third of the species, six of them new. In the BMNH
collection four-fifths of the species were represented, including 17 primary types of previously
known species and seven of the new species described here. I reared about 100 specimens of two
species, palpalis and melanocosta, and eight larvae and four pupae of these have been studied.

Of the specimens studied, about 300 belonged to only four species: argonais, butyrota, melano-
costa and palpalis. In contrast, 15 species were represented only by single specimens; nine of these
by females, including five of the seven described but unnamed species.

The classification and descriptions of the Timocratica species were based on characters of dry
adult specimens. Thirty-four characters were selected, as discussed in the section on classification,
and used in the cladistic analysis. A selection of these characters was used again in the phenetic
analysis as two-state characters.

In the cluster analysis three methods were used to assess the overall similarity or dissimilarity
of all species: (a) the product-moment correlation coefficient; (b) the taxonomic distance coef-
ficient (Sneath & Sokal, 1973: 124); (c) Gower's coefficient (Gower, 1971). The data were used
either in their untransformed state or after standardization by characters to zero means and unit
standard deviations. Clustering of the taxa from the between-taxon similarity/dissimilarity
matrix was accomplished by the weighted pair-group method of Sokal & Sneath (1963). All these
computations were carried out by a program of Davis (1973), as amended and extended by
Dr R. G. Davies (unpublished). The program also constructed and drew on the line-printer, the
dendrogram expressing the results of the cluster analysis.

The measurements at the beginning of each description are those of the fore wing length of the
smallest and largest specimen, measured in millimetres from the base to the apex of the wing. In
several instances the number of available specimens was limited and variation in size of those
species may be greater than is indicated by the recorded measurements.

Dissections and slide preparations followed the method described by Robinson (1976). The
number of genitalia preparations varied with the relative similarity of species and material
available, and is detailed for each species under ' Material examined '. Head preparations were
made and illustrated only for representative species of each of the three species-groups. The wing
venation is illustrated for representatives of each species-group and for species which differ from
the pattern in the group. Drawings of the genitalia are based on individual specimens and are not
composite. The size of the illustrations depended on the size of the specimens. Large specimens
were drawn to a smaller scale and in some instances figures of different scale appear on the same
page. The photographs of the moths show the right-hand wings; where these were unsuitable for
photography the left-hand wings were taken and the image reversed. All the drawings and
half-tone illustrations except one were made by myself.

The geographical distribution of each species is based on specimen labels, completed and/or
corrected, when necessary, according to the 1968 edition of the Times Atlas of the World.
Localities not traced in this atlas were corrected and completed following Brown (1979). Altitude,
when given in feet on the specimen label, was converted into metres, for example: " 1000 m
('3100ft')".

The ecological distribution of the species of Timocratica is expressed according to Holdridge's
system of ' Classification of World Life Zones' (Holdridge, 1967; 1978; Holdridge et al, 1971).
This system has the advantages of being simple and easily used by any biologist, not only by
ecologists, and of being objective, since it is based mainly upon meteorological data, viz., annual
average temperature and total annual precipitation. Another advantage is that most of the
Central and South American countries have already been mapped following this system.

Data on temperature and precipitation were taken from Wernsted (1972). However, as there
are no meteorological stations at the localities of many of the species, the data used were those
provided by the nearest station at the same altitude. In a few cases, no nearby meteorological
station at a similar altitude was found in Wernsted, and the nearest station was selected and the
temperature corrected assuming a decrease of about 6°C per 1000 m of elevation (Holdridge,
1971: 13). Following these procedures a list of all the localities taken from specimen-labels was
organized, including geographic coordinates, altitude, mean annual temperature, total annual

NEOTROPICAL GENUS TIMOCRATICA

215

Mean Annual Biotemperature in Degrees Centigrade

3
•a

o

o
N

oo
12
-5
"o
DC

216 V. O. BECKER

precipitation and the respective Life Zone. This list was submitted to Dr Holdridge who checked
it and made corrections and comments.

It must be remembered that label data are sometimes vague or wrong, particularly on speci-
mens from old collections. Therefore, especially in mountainous areas where the climate changes
over relatively short distances, many specimens could have been collected in Life Zones different
from those calculated from the available label data. Another problem resulting from lack of data
and particularly of specimens is that some specimens were probably collected in associations
atypical of the Life Zone ('Climatic Association' of Holdridge, 1971: 16). A good example of
non-climatic association and of specimens presumably mislabelled is that of T. major.

Despite the problems pointed out in the last paragraph, Holdridge's Life Zone system provides
valuable information about the ecological adaptations of the species and also gives a good
indication of where a species could be expected to occur.

The host-plants collected by the author were identified by Dra M. Brandao Ferreira, Empressa
de Pesquisa Agropecuaria de Minas Gerais, Belo Horizonte, and are marked in the Table with an
asterisk (*). Other hosts are quoted from Araujo et al. (1968: 290) and from Hayward (1969: 72).
Brazilian vernacular names for the Myrtaceae are taken from Legrand & Klein (1967-1978).
English vernacular names are quoted from. Bailey (1900-1902) and Adams (1972). The ichneu-
monid parasite ofpalpalis was identified by Dr M. G. Fitton, BMNH.

Abbreviations of depositories

BMNH British Museum (Natural History), London, England

ESALQ Escola Superior de Agricultura " Luiz de Queiroz ", Piracicaba, Sao Paulo, Brazil

IP Institut fur Pflanzenschutzforschung, Eberswalde, East Germany

LN Landessammlungen fur Naturkunde, Karlsruhe, West Germany

MNHU Museum fur Naturkunde der Humboldt-Universitat, Berlin, East Germany

MN Museu Nacional, Rio de Janeiro, Brazil

NM Naturhistorisches Museum, Vienna, Austria

NMNH National Museum of Natural History, Smithsonian Institution, Washington, D.C., U.S.A.

UCV Universidad Central de Venezuela, Maracay, Venezuela

VB V. O. Becker collection, Centre de Pesquisa Agropecuaria dos Cerrados, Planaltina, Brazil

ZSBS Zoologische Sammlung des Bayerischen Staates, Munich, West Germany

Colour-pattern and defence

Species of Timocratica show two basic colour-patterns. Those belonging to the monotonia and
leucocapna species-groups, as well as parvifusca, which belongs to the albella-group, have dull
fuscous fore wings and bright golden-ochreous hind wings and abdomen. The species of the
albella-group, except for parvifusca, have white fore wings and white or golden-yellow hind wings
and abdomen.

Concerning protection against predation, the first group is probably cryptic, or possibly cryp-
tic only when at rest as the raised golden-ochreous hind wings are possibly aposematic. The
second group, the whites, are probably mimetic. Adults of many species of arctiids are known to
be toxic (Rothschild et al., 1979), and field tests carried out by Collins & Watson (1981) in
Venezuela showed that white species of lithosiids (Arctiidae) were rejected by birds. These mostly
white lithosiids form a very large and common group, now included in the genus Agylla, and are
sympatric with the species of Timocratica. If the species of Timocratica prove not to be toxic, they
would form a Batesian mimetic group of the species of Agylla.

This complex of white mimics may also include other moths, such as species of Rupela Walker
(Schoenobiinae), a large genus of Neotropical pyralids.

The cryptic group of Timocratica includes 12 species, the supposed mimic group 34 species.
Thus, if other factors that might affect the success of a group of living organisms are excluded, it
seems that a mimetic habitus provides more effective protection than crypsis in Timocratica.

NEOTROPICAL GENUS TIMOCRATICA 217

Geographical and ecological distribution

The monotonia-group, despite its relatively small number of species, has the widest geographical
and ecological distribution. It ranges from Mexico and Central America, where it is represented
by two species, possibly three, to the Warm Temperate Moist Forest in the southern part of
Brazil, where it is represented by meridionalis; one species, longicilia, occurs in the Tropical
Montane Rain Forest in the mountains of Colombia.

The leucocapna-group appears to be a montane group as it has been collected only in the
mountains of Peru, Colombia and Costa Rica, being restricted to the Tropical Premontane
Moist Forest and the Tropical Premontane Wet Forest Life Zones.

The albella-group, the largest of the species-groups, is mainly South American and only two
species, xanthotarsa and parvifusca, are known to occur in Central America (Panama and Costa
Rica). Ecologically the group is almost confined to the Tropical Moist Forest and the Tropical
Premontane Moist Forest and Wet Forest, but it is represented by three species in the Warm
Temperate and Subtropical Moist Forest, and by one, guarani, in the Warm Temperate Dry
Forest.

Classification of Timocratica

Like many other lepidopterous groups of the Neotropical Region, particularly Microlepidoptera,
the Stenominae have been very little studied or even collected. Therefore it is very difficult to
appreciate the degree of variation in the group, or the relationship between the different groups in
the subfamily.

An attempt was made to work out the phyletic relationship of the species based on a cladistic
analysis of the genus. However, as discussed below, many difficulties were found and only a basic
division of the genus into three species-groups, as arranged in the cladogram (Fig. 2), seems to be
sound; because of this, the species of each of the three species-groups are arranged phenetically,
following numerical methods.

Cladistic analysis

Although the species now included in the genus Timocratica show great external differences
between species-groups, mainly in colour-pattern, they apparently constitute a monophyletic
group. According to the characters discussed below, the genus is composed of three species-
groups, each of them also apparently monophyletic.

This basic division of the genus into three species-groups seems consistent and presumably
reflects very well the first steps of its evolution. However, above this level it was impossible to
understand the relationship between the species within each of the three groups. The main
difficulty is lack of data. Obviously many of the apomorphies are not expressed morphologically,
but reflected in behaviour, host-preferences, ecological adaptations, and other biological and
physiological features. As the biology of Timocratica is insufficiently known, none of this infor-
mation can be included in the analysis. Another difficulty is related to the method itself, as
conceived by Hennig (1966). It seems that cladistic analysis may work very well at generic level
and for higher classification, but not at specific level, except perhaps with relatively small and
well-known groups. At the specific level it is often very difficult to decide whether a particular
state of a character is primitive (plesiomorphic) or derived (apomorphic).

Although most of the important genera of the subfamily are still poorly known, it seems very
likely that the sister-group of Timocratica is the genus Loxotoma Zeller, which includes only two
described species. They have similarly broad valvae, with sacculus and ampulla not differentiated,
and a strong uncus that is basally broad and bent ventrad. Loxotoma also constitutes a mono-
phyletic group, whose species share at least the apomorphic state of character 1. The monophyly
of Timocratica is defined by characters 2-4 (Fig. 3). In the following list the headings denote the
apomorphic state.

218

V. O. BECKER

(1) Basal third of fore wing costa concave

In most Stenominae, including Timocratica, the fore wing costa is straight to convex. In
Loxotoma the costa is concave, as are the subcostal and first radial veins to some extent (see
figure in Duckworth, 1967: 7).

(2) Gnathos undivided medially

Timocratica has an undivided gnathos expanded medially to form a strongly sclerotized
projection which is bent ventrad, the ' apex '. In Loxotoma as well as in other presumably related
groups, such as Falculina Zeller, the gnathos is divided in the middle, often forming a pair of
apically dentate arms.

LOXOTOMA

T / MOCR AT/CA

a

3

o

i_
en

c

o

•*-

o

c
o
E

a.

o

i—

D)
I
O

a
o

u
O
u

a

D
O

k-

O)

I
o

"aJ

-Q

2-4

Fig. 2 Cladogram representing the relationship between Timocratica and Loxotoma, and the primary
division of Timocratica into species-groups. Open squares denote plesiomorphy, filled represent apo-
morphy. Numbers refer to characters discussed in the text.

NEOTROPICAL GENUS TIMOCRATICA 219

T/MOCRAT/CA

monoronia-group 'eucocapna a/be/la-group

-group

P °° o E ~

O 'u C QJ -2 '~ O ^ C O 03

X

2 E! o al I J &J? -2 £t

an ••••••••• nnn nnnn DD nn nan an nan na ana anna nan an a n n a

7n nnnonnonn ••• DDDD an an nnn an a an an an a n nan ana an cm n a

6D 100000000 ana ••••••••••••••••••••••••••••••••••

so ooooooooo mm® ••••••••••••••••••••••••••••••••• o

40 ••••••••• ••• ••••••••••••••••••••••••••••••••••

30 ••••••••• ••• ••••••••••••••••••••••••••••••••••

20 ••••••••• ••• ••••••••••••••••••••••••••••••••••

IB onaaoaoao oaa 0000000000000000000000000000000000

Fig. 3 Character matrix of the genus Loxotoma and the species of Timocratica. Open squares denote
plesiomorphy, filled squares apomorphy, shaded squares the intermediate state of an apomorphic trans-
formation series. Numbers refer to characters discussed in the text.

(3) Juxta with two simple, almost straight, lateral processes

In Timocratica the two lateral processes of the juxta are simple and almost straight, whereas in
Loxotoma each process is divided in two as in Falculina where these processes are developed into
very complex structures.

(4) Vesica armed with cornuti

In Timocratica the vesica is armed with a strong cornutus or covered with many spines, or with
both. These features are not developed in Loxotoma or other related genera such as Falculina.

(5) Fore wings with white scales

The species of the monotonia-group have the ground-colour of the fore wings plain fuscous, a
state which is also found in the related genus Loxotoma. The species of the albella-group have a
plain white ground-colour, and in the leucocapna-group the ground-colour is fuscous but with
areas of white scales on the upper surface of the fore wing, mainly beyond the cell. The presence of
such white scales in the leucocapna-group and the white fore wings in the albella-group could
therefore be considered an apomorphic transformation series. This makes these two species-
groups the sister-clade of the monotonia-group.

(6) Digitate processes of juxta long

The monotonia- and /eucocapna-groups have the digitate processes of the juxta short, not
reaching the anterior side of the gnathos (except in major where they are long, as in the albella-
group). In the albella-group these processes are often very long, overlapping the anterior side of
the gnathos. Short processes are here considered plesiomorphic, as the homologous hairy pro-
cesses in the related genera Loxotoma and Falculina are also short.

(7) Third segment of labial palpus small

In the leucocapna-group the third segment of the labial palpus is much reduced (Figs 5, 8), less
than one-third the length of the second ; in the other species-groups, as well as in Loxotoma, it is
longer, about two-thirds the length of the second (Figs 4, 6, 7, 9).

(8) Mesonotum with long scales

In the monotonia-group the scales along the middle of the mesonotum are very long and raised
to form a crest (Fig. 9); in the remaining groups these scales are normal as in Loxotoma and
Falculina. Therefore, this development is an autapomorphic state for the monotonia-group.

220

V. O. BECKER

(9) Fore wings with cubital veins stalked

The species of the monotonia-group have the fore wings with CuAl and CuA2 stalked (Fig. 12); in
the leucocapna-group all the veins are free (Fig. 13); and in the albella-group the veins are free in
most species but in some the cubitals are stalked (Figs 13, 16, 24). However, the stalking in these
few species seems to be linked to the stalking of veins jR4 and R5, a character not found in the
other two species-groups. Considering that Loxotoma also has all the veins free, the stalking of
the cubital veins in the monotonia-group can be considered apomorphic.

Figs 4-6 Timocratica species, frontal view of denuded heads. 4, T. palpalis (Zeller). 5, T. leucocapna
(Meyrick). 6, T. monotonia (Strand).

NEOTROPICAL GENUS T1MOCRATICA

221

Figs 7-9 Timocratica species, lateral view of heads with dorsal outline of thorax. 7, T. palpalis (Zeller). 8,
T. leucocapna (Meyrick). 9, T. monotonia (Strand).

Phenetic analysis

A satisfactory arrangement based on a cladistic analysis of the species-groups, mainly those
within the albella-group, could not be produced. This failure was due mainly to the mosaic
pattern of evolution shown by the species. An assessment of phenetic similarity was therefore
made, following numerical methods, as shown in Figs 10, 1 1.

222
4

.5

.6

V. O. BECKER

.7 .8

.9

major

/ong/c»/»a
monofonia
agramma
pompe/ana

me rid/ on a//s

loxofoma

fraferne/la

Fig. 10 Phenogram of the species of the monotonia-group of Timocratica calculated by weighted pair-
group method of average linkage from matrix of between OTU Gower's coefficient. Numbers at top
denote magnitude of correlation coefficient.

In addition to the nine characters used in the cladistic analysis a further twenty-five were
selected and used in the phenetic analysis (Table 1).

The arrangements given by the three phenetic methods showed few differences from one
another, and in a few cases they were even identical. However, the most generally satisfactory
results were provided by Gower's Coefficient.

The clustering of the monotonia-group (Fig. 10) was based on characters 6 and 10 to 16. The
basic division of this group into three subgroups, as shown in the phenogram, reflects very well
what was expected from the overall similarity of the species. T. major is really a very distinctive
species in the group, and its separation from the others seems plausible. The division of the other
species into two groups, as shown in the same phenogram, also seems correct, although the
arrangement of the species inside each of these subgroups does not reflect very well what might
have been expected from their overall resemblance. As discussed in the taxonomic section,
monotonia and pompeiana would have been expected to form a pair of very closely related species,
or even to appear as forms of the same species. In the other subgroup, for the same reasons,
fraternella and loxotoma should form a pair of closely related species, as indicated by the absence
of coremata on the male abdomens and by their geographic distribution. The species meridionalis
and loxotoma came together in the phenogram because the free coremata (character 12), as
present in meridionalis, and the absence of coremata, as shown in loxotoma and fraternella, were
both considered as apomorphic,Tvhile the presence of coremata bound into pockets, as shown by
the other species of the group, is considered plesiomorphic. As meridionalis and loxotoma are
almost identical in respect of the remaining characters, they were therefore clustered together,
while fraternella, which shares the absence of coremata with loxotoma, was separated from both
by the apomorphic fuscous hind wings.

NEOTROPICAL GENUS TIMOCRATICA 223

Table 1 Characters used in the phenetic analysis as two-state variables, a = presumed apomorphies,
p = presumed plesiomorphies, ? = assessment of apomorphy/plesiomorphy not established.

Character

no. Description

10 a. Fore wing with apex pointed
p. Fore wing with apex rounded

11 a. Fore wing with fasciae linked posteriorly
p. Fore wing with fasciae free

12 a. Coremata of second abdominal segment free or absent

p. Coremata of second abdominal segment located in a pocket

13 a. Hind wing fuscous

p. Hind wing golden-ochreous or golden-yellow

14 a. Apex of gnathos folded

p. Apex of gnathos not folded

15 a. Fore wing with transverse fasciae absent
p. Fore wing with transverse fasciae present

16 a. Valva with base broader than distal part

p. Valva with dorsal and ventral margins almost parallel or narrowed basally

17 a. Hind wing bordered with fuscous
p. Hind wing plain golden-yellow

18 a. Hind wing white or tinged with yellow
p. Hind wing plain golden-yellow

19 a. Abdomen above white or tinged with yellow
p. Abdomen above plain golden-ochreous

20 a. Fore wing underside with yellow colour absent
p. Fore wing underside with yellow colour present

21 a. Hind tarsus white

p. Hind tarsus golden-ochreous

22 a. Fore tarsus white
p. Fore tarsus fuscous

23 a. Fore tarsus golden-ochreous
p. Fore tarsus fuscous

24 a. Frons plain white

p. Frons edged with fuscous

25 a. Labial palpus with ochreous colour absent
p. Labial palpus with ochreous colour present

26 a. Labial palpus with fuscous colour absent
p. Labial palpus with fuscous colour present

27 a. Gnathos with lateral arms modified

p. Gnathos with lateral arms not modified

28 a. Vesica with small spines missing
p. Vesica with small spines present

29 a. Fore wing with R4 and R5 stalked
p. Fore wing with R4 and R5 free

30 a. Fore wing with base of costa tinged with grey
p. Fore wing with base of costa white

31 a. Veins marked with dark fuscous

p. Veins not marked with dark fuscous

32 a. Ductus bursae and corpus bursae not differentiated
p. Ductus bursae and corpus bursae differentiated

33 ?. Gnathos with pointed apex
?. Gnathos with rounded apex

34 ?. Margin of ostium bursae convex

?. Margin of ostium bursae concave or straight

224

V. O. BECKER

.7

grondis
xonfhoforso
consfr/cfiva/va
species 3

bicornufa

am sell

subova/is

ane/aea

isargo

tifano/euco

parvi/euca

macro/euco

species 5
species 7
pa/pa/is
mega/euca
maturescens

leucorecf/s
species 6
spinignatha
argonais

a/be/la
guarani

ph/lomela
melanosfriga

venifurcafa

nivea

species 4

fuscipa/pa//s

xanfhosoma

a/b/togata

me/anocosfo

butxro'°
parvifusco

Fig. 11 Phonogram of the species of the albella-group of Timocratica calculated by weighted pair-group
method of average linkage from matrix of between OTU Gower's coefficient. Numbers at top denote
magnitude of correlation coefficient.

NEOTROPICAL GENUS TIMOCRATICA 225

The arrangement of the species of the albella-group was based on the similarity of characters 5,
9 and 19-34. The resulting grouping (Fig. 1 1) also looks reasonable, except that the species with a
golden-ochreous abdomen and white hind wings, related to xanthosoma, were mixed up with
those related to albella which have a white abdomen and white hind wings. If this character (the
colour of the abdomen in the albella-group) is considered as more important than most of the
others analysed, then amseli, subovalis, venifurcata, xanthosoma and fuse ipalpalis should form a
group of closely related species. A few other species, viz., anelaea + isarga and titanoleuca + par-
vileuca, also seem to be wrongly associated. As discussed in the taxonomic section, isarga appears
to be related to palpalis, and parvileuca to butyrota. These apparent inaccuracies in the phenetic
cluster analysis probably resulted from the lack of representation of males or females in about
two-thirds of the species, and from the equal weight given to all characters.

TIMOCRATICA Meyrick

Timocratica Meyrick, 1912; 706; Busck, 1935: 16 [catalogue]; Clarke, 1955: 384 [adult, genitalia]. Type-
species: Timocratica isographa Meyrick, 1912 [ = Cryptolechia monotonia Strand, 1911], by original
designation and monotypy.

Lychnocrates Meyrick, 1926: 226; Clarke, 1955: 224 [adult, genitalia]. Type-species: Lychnocrates leuco-
capna Meyrick, 1926, by monotypy. [Synonymized by Busck, 1935: 16.]

Vertex densely covered with long, narrow scales. Haustellum slighly shorter than second segment of labial
palpus, covered basally with long scales, distal half with sensory papillae. Maxillary palpus four-segmented,
about length of first segment of labial palpus. Labial palpus ascending, reaching to vertex or beyond; first
segment very short, one-quarter length of second; second segment about twice length eye diameter, slightly
curved upwards, with long rough scales below; third segment one-third to same size as second, smooth-
scaled, thick to slender, slightly curved upwards, or straight. Antenna three-quarters length of fore wing,
ciliation one-half to twice diameter of flagellum. Thorax with or without dorsal crest, densely or sparsely
covered with long hair-like scales below. Metascutum with pair of long, hair-like, posteriorly directed
groups of scales. Fore tarsus thickened by long scales; hind tibia covered with long, rough, hair-like scales.
Fore wing subrectangular to suboval; apex rounded, pointed in few species; fuscous, often with three
oblique fasciae, or plain white; 12 veins (11 in holotype of syndicast is [ = butyrota]), Rl from middle of cell,
R2 closer to R3 than to Rl} R4 and Rs very close, connate or stalked; CuAl and CuA2 very close, connate or
stalked (Figs 12-24). Hind wing sometimes golden-ochreous to plain white, rarely fuscous. Abdomen long,
robust, weakly sclerotized, reaching tornus in resting position, densely covered with narrow scales; male
often with pair of cor^mata on second sternite (Fig. 25); apodemes on second abdominal sternite short in
female, modified in male to accommodate coremata; sternites two to seven with some small sparsely
distributed setae; eight with several longer ones; first tergite as well as genitalia covered with long narrow
scales; female with dorsal membrane between eighth and ninth segments expanded as a wide inwardly
directed sac.

GENITALIA $. Symmetrical. Uncus very broad basally, nearly triangular or with lateral margins nearly
parallel, long, strong, bent ventrad, naked. Gnathos often belt-like, modified medially into strong, scler-
otized, often pointed process. Juxta a transverse plate with two long, usually symmetrical, posteriorly
directed lobes covered distally with long setae. Vinculum complete, often rounded. Valva long, broad, lateral
margins nearly parallel or constricted basally, inner surface of distal half covered with many modified,
strong, apically divided setae; ampulla and sacculus slightly or not differentiated, covered with very long
setae. Aedeagus cylindrical, straight or bent ventrad; inception of bulbus ejaculatorius basal, often on dorsal
side of aedeagus ; vesica often with one strong cornutus and many smaller, acutely pointed spines.

GENITALIA 9- Papillae anales slightly to strongly sclerotized, sparsely covered with very long setae. Eighth
segment strongly sclerotized, tergite with irregular row of long setae on posterior margin, sternite covered
with very long setae. Apophyses anteriores and posteriores of same length or latter slightly longer. Ostium
bursae narrow to broad; antrum cylindrical or conical; ductus bursae straight or bent posteriorly; corpus
bursae nearly globular ; signa present as plates with inwardly directed spines ; inception of ductus seminalis
near ostium; walls of ductus and corpus bursae often scobinate.

REMARKS. In the right fore wing of the holotype of isographa, veins R4 and R5 have a short
common stalk while the left shows the normal condition with R4 and R5 separate. This was noted
by Meyrick in the original description; however, Clarke (1955: pi. 192, fig. la) illustrated the
anomalous wing which is not representative of the venation of the monotonia-group.

226

V. O. BECKER

14

Figs 12-15 Wing venation of Timocratica <$. 12, T. monotonia (Strand). 13, T. palpalis (Zeller). 14, T. major
(Busck). 15, T. leucocapna (Meyrick).

The non-white Timocratica species show some external resemblance to Thioscelis Meyrick and
Loxotoma and probably form the sister-group of the latter. They can easily be distinguished from
these genera as Thioscelis has unusually long hind legs and Loxotoma has conspicuous shades of
pink on the hind wings. These two genera also have coremata in pockets on the second abdomi-
nal sternite, and the entire valva has strong, apically bifurcated setae. However, in Loxotoma the
gnathos is divided in the middle and each of the digitate processes of the juxta are split into two
branches. Thioscelis has an entire gnathos but the valva has a well-developed ampulla, a charac-
ter absent in Timocratica.

PUPA. Pupae of only palpalis and melanocosta were available for this study, and these show
generalized gelechioid characters. However, like other Stenominae, they are slightly flattened
dorso-ventrally and have the abdomen sharply curved ventrad, with the fifth, sixth and seventh
segments free. They also have strong dorsal incisions on these three segments, allowing dorso-
ventral movements only.

Two specializations are not shared by the other known pupae: the two long cremaster pro-
cesses, called 'anal legs' by Powell (1973: 26), and the peculiar projection of the pronotum

NEOTROPICAL GENUS TIMOCRATICA

227

17

18

19

21

20

22

23

24

Figs 16-24 Wing venation of Timocratica <$. 16-19, T. butyrota (Meyrick). 20, T. parvifusca sp. n. 21, T.
venifurcata sp. n. 22-24, T. nivea sp. n.

(Figs 46-48). The ' anal legs ', which anchor the pupa to the gallery walls, are also known in other
gelechioid groups, such as Ethmia Hiibner (Ethmiidae) (Powell, 1973: 26) and Agonoxena Mey-
rick (Agonoxenidae) (Bradley, 1966: 468), and it seems likely that they might occur in other
Stenominae. For the time being, the expansion of the pronotum can be used to distinguish
palpalis and melanocosta from the other known stenomine pupae.

LARVA. Larvae of only palpalis and melanocosta were available for this study, both species
belonging to the albella-group. As this does not include monotonia, the type-species of the genus,
it is not certain that the characters in these two species are representative of the whole genus.

Apart from these, the larvae (and pupae) of only five other stenomine species have been
described: Antaeotricha dissimilis (Kearfott) (Becker, 1970), A. schlaegeri (Zeller) (Mackay, 1972),
Stenoma crambina Busck (Dampf, 1929), S. decora (Zeller) (Silva & Heinrich, 1946) and S. ybyra-
juba Becker (Becker, 1971). The available information is therefore inadequate for generalizations
to be made about the taxonomic value of characters, and the relationship of the Stenominae with
other gelechioid groups, although some characters are very similar in the larvae of all seven
species.

Like other gelechioids, stenomine larvae have three prespiracular setae, LI, L2, L3, on the
prothorax, and LI and L2 on the same pinaculum on abdominal segments 1-8. In Timocratica
and the five species mentioned above, the adfrontal area of the head does not extend to the
vertical angle and the distance between setae P2 is the same as or less than the distance between
setae PI (Fig. 39). These characters are also found in the Xyloryctinae, whereas in all other

228

V. O. BECKER

25

27

Figs 25-27 Timocratica species, abdominal segments 1-2. 25, T. monotonia (Strand). 26, T. meridionalis
sp. n. 27, T. loxotoma (Busck).

gelechioids the distance between setae P2 is greater than that of setae PI, and the adfrontal area
reaches the vertical angle of the head, except in some Ethmiidae. According to Powell (1973) the
adfrontal area in Ethmia often reaches the vertical angle, but in a few species it does not.
However, as in other gelechioids, except for Stenominae and Xyloryctinae, the distance between
setae P2 is always greater than the distance between setae PI.

In an attempt to trace relationships among the gelechioids I examined the larvae of four
Australian Xyloryctinae in the BMNH: Cryptophasa hyalinopa Lower, C. balteata Meyrick,
Echiomima mythica Meyrick and Perixestis eucephala (Turner). These species also have the
adfrontal area not reaching the vertical angle of the head and the distance between setae P2 is
almost the same as that between setae PI. Therefore it seems that the combination of both
characters of the head, i.e., adfrontal area not reaching the vertical angle and the distance
between setae P2 about the same as between setae PI, is a good diagnostic feature for dis-
tinguishing the larvae of Stenominae and Xyloryctinae from those of other Gelechioidea.

The larvae of the two species of Timocratica described here can be distinguished easily from
the other five, as the former have setae D and setae SD on the same pinaculum on both the meso-
and metathorax, and a series of extra sclerotized areas, ' pinacula without setae ', on the meso-
and metathorax and on segments 1-7 of the abdomen.

Key to species and subspecies

Note. The males of albella, isarga, megaleuca, melanostriga, species 1, and species 3 to species 7, and the
females of anelaea, constrictivalva, fuscipalpalis, guarani, macroleuca, maturescens, parvifusca, parvileuca,
philomela, spinignatha, subovalis, titanoleuca, venifurcata, xanthotarsa, effluxa, agramma, fraternella, lon-
gicilia, pompeiana and species 2 are unknown.

NEOTROPICAL GENUS TIMOCRATICA 229

1 Ground-colour of fore wing white (albella-group) 14

Groundcolour of fore wing not white. .......... 2

2 (1) Fore wing with all veins free (leucocapna-group) 4

Fore wing with not all veins free ............ 3

3 (2) Fore wing with R4 and/? 5 free (monotonia-group) 6

Fore wing with R4 and R5 stalked parvifusca (p. 270)

4 (2) Hind wing edged with fuscous effluxa (p. 240)

Hind wing not edged with fuscous 5

5 (4) Fore wing of male less than 23 mm leucocapna(p.23S)

Fore wing of male more than 25 mm species 2 (p. 239)

6 (3) Apex of fore wing pointed major (p. 231)

Apex of fore wing rounded 7

7 (6) Hind wing golden-ochreous ............ 9

Hind wing yellowish fuscous or dark fuscous 8

8 (7) Abdomen golden-ochreous species 1 (p. 237)

Abdomen golden-ochreous crossed with fuscous bands .... fraternella (p. 237)

9 (7) Fore wing without distinctive fasciae agramma (p. 232)

Fore wing with three distinctive fasciae 10

10 (9) Antenna with ciliation clearly longer than diameter of flagellum . . . longicilia (p. 232)

Antenna with ciliation about diameter of flagellum 11

11 (10) Second abdominal sternite without coremata ...... loxotoma (p. 236)

Second abdominal sternite with coremata 12

12 (11) Coremata on second abdominal sternite in inverted pockets 13

Coremata on second abdominal sternite free, attached to the sternite surface meridionalis (p. 235)

13 (12) Male genitalia with margins of valva nearly parallel monotonia (p. 234)

Male genitalia with margins of valva not parallel, converging towards apex pompeiana (p. 233)

14 (1) Fore wing plain white 15

Fore wing marked with dark fuscous melanostriga (p. 266)

15 (14) Hind wing plain golden-yellow 16

Hind wing white or tinged with yellow 20

16 (15) Fore tarsus fuscous 19

Fore tarsus golden-ochreous 17

17(16) Fore wing with veins CuAr and CuA 2 stalked species 3 (p. 243)

Fore wing with veins CuAl and CuA 2 not stalked 18

18(17) Gnathos basally with long, digitate processes constrictivalva (p. 243)

Gnathos basally without long, digitate processes xanthotarsa (p. 242)

19(16) Mid tarsus fuscous above grandis (p. 240)

Mid tarsus golden-ochreous bicornuta(p. 241)

20 (15) Abdomen plain golden-ochreous above

Abdomen white, or tinged with yellow, or crossed with white bands above ....

21 (20) Hind wing plain white 22

Hind wing tinged with yellow species 4 (p. 244)

22(21) Fore wing with K4 and R 5 stalked venifurcata (p. 245)

Fore wing with R4 and R 5 not stalked

23 (22) Fore wing with base of costa fuscous 24

Fore wing with base of costa not fuscous ....
24(23) Head with vertex golden-yellow xanthosoma xanthosoma (p. 247)

Head with vertex white xanthosoma leucocephala (p. 247)

25(23) Second segment of labial palpus almost fuscous externally . . . . fuscipalpalis (p. 246)

Second segment of labial palpus almost ochreous externally . . 26

26 (25) Vesica with strong cornutus and many smaller spines .... subovatts (p. 243)

Vesica with strong cornutus only • amsett (p. 244)

27(20) Abdomen golden-ochreous crossed with white bands above . . . anelaea (p. 248)

Abdomen white or tinged with yellow above

28 (27) Hind tarsus plain white

Hind tarsus golden-yellow, or tinged with yellow

29 (28) Fore tarsus white or mixed with white scales above ... 30

Fore tarsus fuscous above • isarga (p. 266)

30 (29) Abdomen tinged with yellow above .

Abdomen plain white above ....... 32

230

V. O. BECKER

31 (30) Frons white edged with fuscous guarani (p. 268)

Frons plain white albella (p. 267)

32(30) Fore wing with costa dark grey basally melanocosta(p. 261)

Fore wing with costa white basally nivea (p. 262)

33 (28) Fore tarsus white above 34

Fore tarsus dark fuscous above 36

34(33) Abdomen and underside of wings plain white albitogata (p. 264)

Abdomen and underside of wings tinged with yellow 35

35(34) Second segment of labial palpus golden-yellow below macroleuca(p.24S)

Second segment of labial palpus dark fuscous below titanoleuca (p. 248)

36 (33) Hind wing tinged with yellow above

Hind wing white above 41

37(36) Second segment of labial palpus golden-yellow below 39

Second segment of labial palpus dark fuscous below 38

38 (37) Fore wing with R4 and R5 stalked butyrota (p. 269)

Fore wing with all veins free parvileuca (p. 269)

39 (37) Abdomen white above 40

Abdomen tinged with yellow above spinignatha (p. 250)

40 (39) Fore wing more than 20 mm species 6 (p. 265)

Fore wing less than 15mm philomela (p. 268)

41 (36) Second segment of labial palpus white internally 42

Second segment of labial palpus almost go Iden-ochreous .... maturescens (p. 252)

42 (41) Fore femur dark fuscous above 43

Fore femur yellow above 44

43 (42) Fore wing white below along apex and termen . . . palpalis (p. 253), megaleuca (p. 253)

Fore wing dark fuscous below along apex and termen .... argonais (p. 251)

44 (42) Mid femur yellow above species 7 (p. 265)

Mid femur white above 45

45 (44) Fore wing more than 25 mm leucorectis (p. 249)

Fore wing less than 20 mm species 5 (p. 250)

Division of Timocratica into species-groups

Except for parvifusca, a small species of uncertain position, all species can easily be clustered into
three well-defined natural groups: 1) the albella-group which includes species with white fore
wings (Figs 68-78); 2) the monotonia-group whose species have fuscous fore wings with three
oblique, nearly parallel fasciae (Figs 55-63); 3) the leucocapna-group which includes species with
dark fuscous fore wings, without fasciae, but with a diffuse white area beyond the cell and an area
of yellow scales on the basal half of the costa (Figs 64-66). T. parvifusca (Fig. 67) is a small, dark
fuscous species in which the fore wing has R4 and R5 and CuA{ and CuA2 stalked. The wing-
shape, genitalia, wing-venation, and distribution put it very close to butyrota, a small white
species in the albella-group. The fuscous colour of parvifusca is presumably due to a secondary
loss of the advanced state.

These three species-groups are defined by characters 5-9 (Figs 2, 3), discussed as follows.
Character 5 is the apomorphy of the clade comprising the albella-group + leucocapna-group, 6 is
the apomorphy of the albella-group, 1 of the leucocapna-group, and 8-9 the apomorphies which
define the monotonia-group (see cladogram, Fig. 2).

The monotonia-group

cJ, ?, 14-30 mm. Head, thorax above, and fore wing fuscous. Third segment of labial palpus about two-thirds
as long as second, ascending vertex very close to head. Thorax with crest of long, narrow scales (Fig. 9). Fore
wing with apex rounded, acute in major; CuAv and CuA2 stalked at basal quarter; three oblique, nearly
parallel fasciae crossing wing, except in agramma. Hind wing often pale to golden-ochreous, fuscous in
fraternella. Abdomen ochreous, crossed with fuscous bands in fraternella; coremata absent in loxotoma and
fraternella, on surface of sternite in meridionalis (Fig. 26).

GENITALIA cJ. Digitate processes of juxta not reaching anterior margin of gnathos apex, except in major.

NEOTROPICAL GENUS TIMOCRATICA 231

REMARKS. The species of this group can easily be distinguished from others by the crest of long
scales on the thorax; from the albella-group also by their fuscous colour, from the leucocapna-
group by the stalked veins CuAt and CuA2 of the fore wing, and from parvifusca by the larger size
and veins R4 and R5 free on the fore wing.

Timocratica major (Busck)
(Figs 14, 28, 63, 79, 80, 154)

Stenoma major Busck, 1911: 212, pi. 8, fig. 8. Holotype <J, PERU: Lima, Callao (Pusey) (NMNH) [not

examined].
Timocratica major (Busck) Meyrick, 1912 : 707 [list] ; Busck, 1935 : 17 [catalogue].

cJ 22-25 mm. Head pale yellow, ochreous towards clypeus; vertex and crown with grey and ochreous-tipped
scales. Second segment of labial palpus ochreous, whitish internally above; third segment whitish. Antenna
pale yellow, scape with greyish-tipped scales; flagellum progressively fuscous from base to apex, ciliation
half diameter of flagellum. Thorax pale yellow with greyish and ochreous-tipped scales. Fore wing with apex
pointed, pale yellow; margins, oblique fasciae, and fold ochreous; underside golden-ochreous. Hind wing
pale golden-yellow, cilia golden. Legs ochreous, paler above; fore tarsus dark ochreous on outer side.
Abdomen ochreous.

9 26 mm. Slightly darker than male. Fore wing irrorate with ferruginous scales; margins, fasciae, fold and
cilia ferruginous.

GENITALIA $ (Figs 79, 80). Uncus slightly narrowed at middle; apex strongly concave, nearly bifurcate. Apex
of gnathos blunt. Digitate processes of juxta narrow and long, overlapping proximal side of gnathos, curved
inwards, covered with setae towards apex. Anterior margin of vinculum nearly straight. Valvae progress-
ively broadening distad; sacculus slightly expanded; apex evenly rounded. Aedeagus bent ventrad at base,
slightly dilated medially; vesica with single strong cornutus.

GENITALIA $ (Fig. 1 54). Ostium bursae narrow, margin straight. Antrum short. Ductus bursae about twice as
long as corpus bursae, nearly cylindrical, walls slightly wrinkled. Corpus bursae globular, walls smooth.
Signum an elongate plate, slightly constricted at middle, weakly sclerotized along and across middle.

REMARKS. T. major is easily separated by its pointed fore wings; these are rounded in all other
species in the group.

Busck (1911) stated that the median and post-median fasciae reach the dorsum, but they
merely reach the fold, and on M3 the post-median forms an acute angle with a fascia that follows
the fold, parallel to the tornus. Busck also stated that the alar expanse was 50-60 mm, but it was
impossible to find specimens larger than 55 mm amongst the material studied. The presence of a
thoracic crest in this species is not clear. All the specimens examined have the thorax more or less
rubbed, except one which has some long, loose scales around the pin, which may indicate the
presence of this character. The shape of the juxta and valva is somewhat unusual for this group.
No other species has the digitate processes of the juxta overlapping the proximal side of the
gnathos, nor the characteristic expansion of the sacculus. This is also the only species in the genus
with such a long ductus bursae.

DISTRIBUTION (Fig. 28). Brazil (Amazonian Basin and Central Plateau), Peru (Pacific coast [prob-
ably erroneous, see below]). This species is presumably associated with Tropical Moist Forest as
indicated by the specimens collected in Borba and Fonte Boa, Amazonas (dots in lowest hexagon
of Fig. 28). The specimens from 'Callao, Peru' are probably mislabelled as this locality is in a
desert area. The specimens from Mato Grosso and Goias come from an area covered predomi-
nantly by two associations. The savanna-type vegetation, called 'cerrado' in Brazil, covers most
of the area and is the result of the monsoon-type of rainfall, corresponding to an 'atmospheric
association' in Holdridge's system. The other association is represented by gallery-forests along
the river banks. These gallery forests represent the climatic association of the area. Thus, the two
dots which represent the two localities in Goias and Mato Grosso (Fig. 28) could be moved
further to the right, probably over the 2000-precipitation line, and falling very close to the other
two dots which represent the most likely ecological association of major.

232

V. O. BECKER

MATERIAL EXAMINED

8 f?, 1 $ (4 cJ, 1 9 genitalia preparations).

Peru: 1 <$ (paratype), Lima, Callao (Pusey) (BMNH). Brazil: 5 £, Mato Grosso, Rio Brilhante, 22.X.1970
(Becker) (VB; BMNH; NMNH); 1 9, Amazonas, Fonte Boa, vii.1906 (Klages) (BMNH); 1 <J, Amazonas,
Borba, Rio Madeira, x.1943 (Pohl) (NMNH); 1 <?, Goias, Leopoldo Bulhoes, x.1935 (Spitz) (BMNH).

Timocratica agramma sp. n.

(Figs 28, 55, 8 1,82)

c? 30 mm. Head fuscous. Labial palpus dark ochreous; second segment above and third segment fuscous.
Antenna fuscous. Thorax fuscous, crest dark brown. Fore wing fuscous, costa ferruginous-ochreous, dorsum
ferruginous, oblique fasciae indistinct; underside golden-yellow, deep golden-yellow along margins. Hind
wing golden-ochreous, cilia and dorsum deep golden-ochreous. Legs deep golden-ochreous, fore tarsus dark
fuscous above; third to fifth segments of mid tarsus fuscous brown.

GENITALIA $ (Figs 81, 82). Uncus narrow, lateral margins nearly parallel; apex medially concave. Apex of
gnathos sharply pointed, strongly sclerotized. Digitate processes of juxta straight, apex with several setae.
Valva with margins nearly parallel, basal third of ventral margin slightly sinuous. Aedeagus slightly bent
ventrad, vesica with a single strong, pointed cornutus.

REMARKS. T. agramma is easily separated from others in this group by the absence of distinctive
oblique fasciae on the fore wings.

DISTRIBUTION (Fig. 28). Brazil (Atlantic coast). The data 'Espirito Santo' on the label of the only
specimen known are not precise enough. However, it can be assumed that the specimen was
collected in the lowlands around the capital, Vitoria, and the species may belong to a transitional
association between the Tropical Dry Forest and the Subtropical Moist Forest.

MATERIAL EXAMINED
Holotype <J, Brazil: Espirito Santo (Johnson) (NMNH).

• longicifia

• major

A agramma

12°C

24°C

Fig. 28 Geographical and ecological distribution of the monot oma-group of Timocratica.

Timocratica longicilia sp. n.

(Figs 28, 56, 83-85)

cJ 28-30 mm. Head fuscous, frons whitish edged with ochreous and fuscous scales, vertex and crown
brownish fuscous. Second segment of labial palpus dark ochreous, brownish fuscous above; third segment
brownish fuscous. Antenna pale yellow, scape fuscous, distal half of flagellum progressively darker towards
apex, ciliation one and a half times diameter of flagellum. Thorax fuscous, dark brown along middle, apex of

NEOTROPICAL GENUS TIMOCRATICA 233

crest scales dark brownish fuscous. Fore wing light fuscous; costa ochreous to ferruginous; termen, dorsum,
oblique fasciae and cilia fuscous; golden-yellow below. Hind wing golden-yellow, cilia deep golden-yellow.
Legs deep golden-yellow; fore tarsus brownish fuscous above, darker outwardly; mid tarsus slightly tinged
with fuscous above. Abdomen deep ochreous, fourth to seventh tergites tinged with brownish fuscous.

GENITALIA $ (Figs 83-85). Uncus broad, lateral margins converging slightly towards apex, apex slightly
concave. Apex of gnathos folded, not strongly sclerotized as in other species. Digitate processes of juxta
short, dorsal side and apex with several setae. Valva very broad, costal margin nearly straight, ventral
margin evenly rounded. Aedeagus bent ventrad, vesica with a single, broad-based, sharply pointed cornutus.

REMARKS. T. longicilia can easily be distinguished from other species in this group by its relatively
long antennal ciliation and by the brownish fuscous third segment of the labial palpus. It is the
only species in the group which has the base of the valva narrower than the distal part. Like major
and agramma, from which it is very distinct externally, it has a single strong cornutus in the
vesica. The specimen from Antioquia, Mesopotamia has the distal half of the valva slightly
narrower than the typical form. Possibly it is a lowland form of the species.

DISTRIBUTION (Fig. 28). Colombia (Oriental Cordillera and Mesopotamia). This species repre-
sents the genus in the high mountains of Colombia, in the Tropical Montane and Tropical Lower
Montane Wet Forest. No other species in the genus is known to occur in these two Life Zones.

MATERIAL EXAMINED

3 cJ (2 c? genitalia preparations).

Holotype <J, Colombia : Tolima, Mt Tolima, 3200 m (Fassl) (BMNH).

Paratypes. Colombia: 1 & Tolima, Mt del Eden, Ibague, 2700 m, xii.1909 (Fassl) (BMNH); 1 £, Anti-
oquia, Mesopotamia, 1500 m('5000 ft')(NMNH).

Timocratica pompeiana (Meyrick)
(Figs 29, 57, 86, 87)

Timocratica pompeiana Meyrick, 1925: 176; Busck, 1935: 17 [catalogue]; Clarke, 1955: 391, pi. 195, figs
4~4b [adult, genitalia]. Holotype & PERU (BMNH) [examined].

cJ 27-30 mm. Head whitish, frons edged with fuscous, vertex and crown with fuscous scales. Second segment
of labial palpus with basal half ochreous, dark grey above, outer half above and near articulations whitish;
third segment whitish, fuscous below. Antenna fuscous; scape whitish, basal half above with fuscous scales.
Thorax fuscous, dark fuscous along middle; crest dark brown, patagia pale yellow, tegulae fuscous. Fore
wing fuscous, costa deep ochreous to ferruginous brown, termen, dorsum, oblique fasciae and fold ferrugi-
nous brown, cilia fuscous, dark ochreous below. Hind wing golden-ochreous, cilia and dorsum deep golden-
ochreous. Legs deep golden-ochreous, fore tarsus fuscous above, dark brown on claws ; mid tarsus and third
to fifth segments of hind tarsus fuscous brown above. Abdomen deep golden-ochreous.

GENITALIA <$ (Figs 86, 87). Uncus nearly triangular, base broad, tapering strongly towards apex. Apex of
gnathos flat and rounded. Digitate processes of juxta straight, dorsal side of apex with several setae. Valva
very broad basally, ventral margin strongly curved near middle. Aedeagus bent ventrad at basal third, vesica
with strong, bent cornutus and several smaller spines opposite.

REMARKS. T. pompeiana is a little larger than monotonia but otherwise very similar externally. The
only difference is in the shape of the valva; in pompeiana it is nearly triangular with a very broad
base, while in monotonia the margins are almost parallel. T. pompeiana could be a local form of
monotonia, as variation of genitalia has been found in different populations of the latter. This view
is supported by the ecological distribution of both forms as discussed below. However, pompeiana
is retained as a distinct species until further material and information is available.

DISTRIBUTION (Fig. 29). Peru (eastern side of the Andes). All specimens were collected in the same
place, the locality being Tropical Premontane Wet Forest. This is also the Life Zone of monotonia
and suggests that pompeiana could be a local form of that species.

MATERIAL EXAMINED

5 c? (3 c? genitalia preparations).

Peru: holotype J, Carabaya, La Oroya (BMNH); 4 & Puno, Carabaya, La Oroya, R. Inambari, iii.1905
(Ockenden) (BMNH).

234

V. O. BECKER

Timocratica monotonia (Strand)
(Figs 6, 9, 12, 25, 29, 58, 88-93, 155)

Cryptolechia monotonia Strand, 1911: 151; 1914: 58, pi. 11, fig. 18 [redescr., adult]. Holotype9, ECUADOR:

Macas (colln Niepelt) [not traced].
Timocratica isographa Meyrick, 1912: 707; 1925: 176 [addition to descript.]; Busck, 1935: 17 [catalogue];

Clarke, 1955: 384, pi. 192, figs 1-ld [adult, wing venation, genitalia]. HolotypecJ, VENEZUELA (BMNH)

[examined]. Syn. n.
Timocratica claudescens Meyrick, 1925: 177; Busck, 1935: 16 [catalogue]; Clarke, 1955: 387, pi. 193, figs

4-4a [adult, genitalia]. Lectotype $, PERU (BMNH), designated by Clarke (1955 : 387) [examined]. Syn. n.
Timocratica crassa Meyrick, 1925: 177; Busck, 1935: 16 [catalogue]; Clarke, 1955: 388, pi. 194, figs 1-la

[adult, genitalia]. Lectotype <$, BRAZIL (BMNH), designated by Clarke (1955: 388) [examined]. Syn. n.
Timocratica monotonia (Strand) Busck, 1935: 17 [catalogue].

<J 20-24 mm, 9 26-28 mm. Head whitish fuscous, frons edged with dark ochreous and fuscous scales; vertex
and crown with fuscous scales, darker along middle. Second segment of labial palpus deep ochreous, dark
grey above, whitish near outer articulation; third segment whitish with dark fuscous scales towards apex,
slightly tinged with ochre above. Scape whitish, with fuscous-tipped scales above; flagellum fuscous, diffus-
ely ringed with whitish scales on articulations; ciliation about diameter of flagellum. Thorax fuscous, dark
brown along middle, scales whitish basally; tegula slightly edged with ochreous scales; crest dark brown
apically. Fore wing fuscous, scales whitish basally; costa deep ochreous to ferruginous brown; termen,
dorsum, oblique fasciae and fold ferruginous brown; cilia fuscous, pale ochreous basally; underside dark
golden-ochreous. Hind wing golden-ochreous, dorsum and cilia deep golden-ochreous. Legs golden-
ochreous; fore tarsus whitish fuscous above, progressively darker distally. Abdomen deep golden-ochreous.

GENITALIA $ (Figs 88-93). Lateral margins of uncus parallel or narrowing slightly towards apex. Apex of
gnathos variable, gradually to abruptly tapered. Juxta with digitate processes straight or with internal
margins slightly sinuate. Margins of valva nearly parallel or valva broader at base and strongly angled at
one-third ; apex evenly rounded. Aedeagus bent ventrad ; vesica with several cornuti on dorsal side, several
smaller acute spines ventrally.

GENITALIA 9 (Fig. 155). Ostium bursae wide, margin shallowly convex. Antrum conical, anterior part
constricted, wrinkled longitudinally. Ductus bursae widening gradually towards corpus bursae. Corpus
bursae pear-shaped, walls smooth as in ductus bursae. Signum a single elongate plate.

REMARKS. T. monotonia is very similar externally to pompeiana and meridionalis. The former is
larger and has the basal half of the valva very broad. The latter has a distinctive lighter area on
the basal half of the fore wing and the coremata on the abdomen attached to the sternite surface.

Although there are no good external features to distinguish specimens from different localities,
Meyrick described this species three times, giving no evidence as to why he believed they were

• monotonia
V pompeiana

• meridiona/is

12°C

-24°C

Fig. 29 Geographical and ecological distribution of the monotonia-group of Timocratica.

NEOTROPICAL GENUS TIMOCRATICA 235

different. The male genitalia exhibit slight differences between specimens from different places,
but seem to be constant in those from the same locality. This is regarded as geographic variation
as specimens from places between the type-localities have intermediate genitalia.

The type of monotonia has not been traced. According to Horn & Kahle (1936: 191, 270),
Niepelt's collection was sold, the Strand types being deposited in the MNHU and IP. Dr. H.-J.
Hannemann and Dr. R. Gaedike of these respective institutions were unable to find the type
(pers. comm.), nor could it be traced at the BMNH where part of the Niepelt collection is now
deposited.

According to Strand's figure monotonia can be associated with only three species: isographa,
longicilia, and pompeiana. It is unlikely that it represents either longicilia or pompeiana, as the
former is known only from the high mountains of Colombia, and the latter (which could be a
local form of isographa) from the type-locality in southern Peru. The third species, isographa
(with its synonyms crassa and claudescens), is widely distributed in the north of South America
including Ecuador; in my opinion monotonia belongs to the same population and is the senior
name for this species.

DISTRIBUTION (Fig. 29). Brazil (Amazonian Basin), Guyana, Colombia, Ecuador, Peru, Ven-
ezuela.

Despite its wide geographic distribution, this species appears to be associated mainly with only
two Life Zones, Tropical Moist Forest and Tropical Premontane Wet Forest. The specimen from
Palma Sola, Venezuela, which is in the Tropical Dry Forest, presumably is associated with
gallery forests in this savanna area.
MATERIAL EXAMINED
21 c£, 3 $ (1 1 c?, 2 $ genitalia preparations).

Brazil: lectotype £ of T. crassa, Para, Belem, vii.1919 (Parish) (BMNH); 8 6*, Para, Belem ['Para'] (Moss)
(BMNH); 1 cJ, 1 ?, Amazonas, Fonte Boa (Klages) (BMNH). Guyana: 1 <J, Essequibo, Potaro, v.1908
(Klages) (BMNH). Colombia: 1 £, Cundinamarca, Medina (Fassl) (BMNH). Ecuador: 1 $, Pichincha, Santo
Domingo de los Colorados, 14.ii.1959 (Hodges) (NMNH). Venezuela: holotype 3 of T. isographa, Falcon,
Palma Sola (BMNH); 1 <J, Aragua, Rancho Grande, 4.vi.l968 (Feige) (VB); 1 & 1 $, same data, 1100 m,
lO.iv, lO.v.1967 (Salcedo & Rodriguez) (VB); 1 & Falcon, Palma Sola (BMNH); 2 <$, Las Quinguas, near San
Esteban (Klages) (BMNH); 1 & San Esteban (Klages) (BMNH). Peru: lectotype $ of T. claudescens, Puno,
San Gaban, iv.1913 (BMNH); 1 $ (paralectotype of T. claudescens Meyrick), Puno, San Gaban [river], 760
m ('2500 ft'), iv.1913 (NMNH).

Timocratica meridionalis sp. n.

(Figs 26, 29, 59, 94, 95, 156)
[Timocratica claudescens Meyrick ; Biezanko, \96lb: 6. Misidentification.]

cJ 23-28 mm. Head whitish, frons edged with ferruginous, vertex and crown dark fuscous along middle.
Second segment of labial palpus dark ochreous, whitish with fuscous scales above and around distal
articulations; third segment whitish with fuscous scales below. Scape whitish fuscous above; flagellum
whitish at base, progressively fuscous towards apex. Thorax light fuscous, ferruginous brown along middle ;
crest dark brown apically; tegula edged with ochreous scales. Fore wing light fuscous; basal half between R
and costa whitish; costa ochreous to ferruginous; apex, termen, tornus, and oblique fasciae ferruginous
brown; fold with ochreous scales; cilia fuscous; underside dark ochreous. Hind wing golden-ochreous, cilia
and dorsum deep golden-ochreous. Legs deep golden-ochreous; fore tarsus whitish with grey scales above,
progressively dark brown outwards; mid tarsus dark fuscous brown above; third to fifth segments of hind
tarsus fuscous brown. Abdomen deep golden ochreous.

9 26-30 mm. Lighter than male; second segment of labial palpus without grey scales above.

GENITALIA <$ (Figs 94, 95). Uncus narrow, lateral margins progressively convergent towards apex; apex
rounded. Apex of gnathos short, blunt. Digitate processes of juxta nearly straight, distal half dorsally and
apex with several long setae. Valva strongly curved near basal third, then with margins almost parallel ; apex
evenly rounded. Aedeagus bent ventrad, slightly narrower at middle; vesica with short, bent cornutus and
few sharply pointed, smaller spines.

GENITALIA $ (Fig. 156). Margin of ostium bursae straight. Antrum with lateral margins almost parallel.
Ductus bursae broadening progressively towards corpus bursae. Corpus bursae pear-shaped. Signum a
single plate, weakly sclerotized along middle.

236

V. O. BECKER

REMARKS. T. meridionalis, the southern species in the group, is very similar to monotonia but can
easily be distinguished by the lighter area on the basal half of the fore wing between R and the
costa, and by the genitalia. Although this species also has coremata on the second abdominal
sternite, these are not located in pockets but are attached to the sternite surface (Fig. 26). This is
probably an intermediate development between loxotoma and fraternella which lack coremata,
and the remaining species of the genus which have them located in pockets (Fig. 25).

It was impossible to examine the material studied by Biezanko (196 Ib: 6), but it certainly
belongs to meridionalis and not to claudescens which is a synonym of monotonia, a species
occurring in the tropical areas of northern South America.

BIOLOGY. Like palpalis, this species emerges earlier (October) in the northern and warmer areas of
its range, and later (February to March) in the southern areas. This seems to indicate that it is
univoltine in southern, colder regions, but further collecting from the northern and warmer
localities may show that it is bivoltine in these areas.

DISTRIBUTION (Fig. 29). Southern Brazil, Paraguay and Bolivia. This species is the only represen-
tative of the monotonia-group in the Warm Temperate and Subtropical regions of South America.
It is restricted to two Life Zones, Warm Temperate Moist Forest and Subtropical Lower Mon-
tane Moist Forest.

MATERIAL EXAMINED

21 cJ, 8 9 (7 (J, 2 9 genitalia preparations).

Holotype £, Brazil: Parana, Curitiba, 920 m, 12.iii.1975 (Becker) (MN).

Paratypes. Brazil: !<?,!$, Minas Gerais, Sete Lagoas, 720 m 18-20.X.1969 (Becker) (VB); 1 & Sao Paulo,
Ipiranga, iii.1926 (Spitz) (BMNH); 1 $, Sao Paulo, Sao Bernardo, iii.1926 (Spitz) (BMNH); 2<$, 19, Sao
Paulo, Piracicaba, 14-19.U966 (ESALQ); 9 & 4 $, Parana, Curitiba, 920 m, 15.ii-20.iii.1975 (Becker) (VB;
BMNH; NMNH; MNHU; NM); 1 J, 1 9, Parana, Igua?u, 20.ii-5.iii.1922 (BMNH); 1 <J, Rio Grande do
Sul, Elsenau, 1905 (Martin) (BMNH); 1 & Rio Grande do Sul, Santa Maria, 25.iii.1971 (Link) (VB); 1<J, Rio
Grande do Sul, Pelotas, 14.ii.1961 (Biezanko) (VB). Bolivia: 1 9, Santa Cruz, Prov. del Sara, 450 m (Stein-
bach) (BMNH). Paraguay: 1 $, Ibapa, 20.X.1924 (BMNH); 1 <?, Sapucay, 20.U905 (Forster) (BMNH); 1 rf,
Villa Rica (Jorgensen)) (NMNH).

Excluded from type-series. [South Africa:] 1 <J, Natal, Stellenbosch (C.K.B.) (BMNH) [mislabelled].

Timocratica loxotoma (Busck)
(Figs 27, 30, 60, 96, 97, 157)

Stenoma loxotoma Busck, 1910: 212; Walsingham, 1913: 179 [list]. Holotype $, MEXICO: Vera Cruz,

Orizaba, vi (Muller) (NMNH) [not examined].
Timocratica loxotoma (Busck) Busck, 1935: 17 [catalogue].

$ 20-22 mm. Head whitish, frons edged with fuscous, vertex and crown with fuscous scales along middle.
Second segment of labial palpus deep ochreous below, dark fuscous above, pale distally, third segment pale
fuscous. Antenna whitish fuscous. Thorax fuscous above, dark fuscous along middle, apical half of crest
scales dark fuscous brown. Fore wing light fuscous; costa, apex, termen, dorsum, oblique fasciae and fold
ferruginous brown ; cilia fuscous. Hind wing and underside of fore wing golfen-ochreous. Legs deep golden-
ochreous; fore tarsus light fuscous above, dark fuscous on claws, mid tarsus progressively dark fuscous
towards claws. Abdomen deep golden-ochreous above, paler below.

9 22-28 mm. Slightly paler than male. Second segment of labial palpus without fuscous tinge above.

GENITALIA $ (Figs 96, 97). Margins of uncus nearly parallel, apex slightly concabe. Digitive expansions of
juxta nearly straight, with few setae apically. Margins of valva nearly parallel. Aedeagus bent ventrad, vesica
with several long spines dorsally.

GENITALIA 9 (Fig. 157). Margin of ostium bursae expanded posteriorly, concave medially. Antrum slightly
constricted at middle, with longitudinal wrinkles anteriorly. Ductus bursae broadening progressively to-
wards corpus bursae. Corpus bursae globular, walls smooth. Signum an elongate plate weakly sclerotized at
middle.

REMARKS. Although this Central American species is very similar to monotonia externally, it can
easily be distinguished in the male as it lacks coremata on the second abdominal sternite
(Fig. 27). The female can be distinguished by the shape of the margin of the ostium bursae and by
the wide, globular corpus bursae.

NEOTROPICAL GENUS TIMOCRATICA

237

DISTRIBUTION (Fig. 30). Mexico (Gulf of Mexico and Yucatan Peninsula), Guatemala and Costa
Rica. This species is associated with a wide range of Life zones and has been collected from
Tropical Dry Forest and Tropical Moist Forest, up to Tropical Premontane Wet Forest.

MATERIAL EXAMINED

13 cJ, 5 9 (3 cJ, 2 9 genitalia preparations).

Mexico: 1 <$ paratype, Veracruz, Veracruz (Schwarz) (BMNH); 3 <J, Veracruz, Huatuxco (BMNH); 3 cJ,
Campeche, Escarcega, 30.ix.1973 (Becker) (VB); 1 & San Luiz Potosi, Palitla, 5.viii.l966 (Flint) (NMNH); 1
cJ, San Luiz Potosi, Tamazunchale, 26.vi.1965 (Flint) (NMNH). Guatemala: 1 & Chejel, vi (Schaus)
(NMNH); 1 (J, Peten, Tikal, 19-22: ix.1973 (Becker) (VB). Costa Rica: 3 $, Cartago, Turrialba, 10.ix.1971,
lO.x.1971, 10.xii.1971 (Becker)(VE); 1 9, Cartago, Turrialba, 10.vi.l972(Becfcer)(BMNH).

Timocratica species 1

(Figs 30, 62, 158)
9 23-24 mm. Similar to loxotoma and monotonia. Hind wing yellowish-fuscous. Abdomen golden-ochreous.

GENITALIA 9 (Fig. 158). Margin of ostium bursae rounded, expanded posteriorly. Antrum slightly constric-
ted posteriorly, anterior half wrinkled longitudinally. Ductus bursae constricted posteriorly, widening pro-
gressively towards corpus bursae, wrinkled longitudinally. Corpus bursae globular. Signum a nearly circu-
lar, diffuse plate.

REMARKS. The two females considered here are very similar to loxotoma and monotonia but their
hind wings are yellowish-fuscous, not golden-ochreous. The genitalia are very close to those of
loxotoma but the margin of the ostium bursae is evenly rounded, whereas it is concave in
loxotoma. Their yellowish-fuscous hind wings suggest some relationship with fraternella, and they
could well represent the female of that species, but as they are very much lighter and in poor
condition it seems better not to name them until more material is available for study.

DISTRIBUTION (Fig. 30). Costa Rica (known only from Turrialba). This species occurs v/ithfrater-
nella in the same Life Zone, viz., Tropical Premontane Wet Forest.

• (oxofoma

• frafernella
T species J

r

ire

24°C

Fig. 30 Geographical and ecological distribution of the monotonia-group of Timocratica.

MATERIAL EXAMINED

2 9 (2 9 genitalia preparations)

Costa Rica: 2 9, Cartago, Turrialba, 600 m, IS.vii, 20.x. 1972 (Becker) (VB; BMNH).

Timocratica fraternella (Busck)
(Figs 30, 6 1,98, 99)

Stenoma fraternella Busck, 1910: 80; Walsingham, 1913: 179 [list]. Holotype cJ, COSTA RICA: Cartago, Juan

Vinas (Schaus) (NMNH) [not examined].
Timocratica fraternella (Busck) Busck, 1935: 16 [catalogue].

cJ 14-19 mm. Head whitish, frons edged with fuscous; crown with fuscous scales along middle. Labial palpus
bright fuscous, second segment dark fuscous above, third segment progressively dark fuscous towards apex.

238 V. O. BECKER

Antenna fuscous, scape whitish. Thorax fuscous, apex of crest scales dark brownish-fuscous, metascutum
and first abdominal tergite yellowish-fuscous. Fore wing dark shiny fuscous, costa edged with deep ochre;
termen, dorsum, oblique fasciae and fold dark brownish-fuscous. Hind wing dark shiny fuscous. Legs deep
golden-ochreous; fore tarsus light fuscous above, dark fuscous distally; mid tarsus dark fuscous above; hind
claws dark fuscous. Abdomen deep golden-ochreous; tergites crossed with dark fuscous bands near articu-
lations.

GENITALIA <$ (Figs 98, 99). Uncus slightly dilated medially. Apex of gnathos short, blunt. Digitate processes
of juxta straight, gently narrowed at middle; dorsal side of apex with several short setae. Valva with margins
nearly parallel, ventral margin evenly rounded at basal third. Aedeagus with basal third strongly curved
ventrad ; vesica with several spines on dorsal side, progressively longer distally.

REMARKS. T. fraternella is easily separated from other species in this group by its dark fuscous
hind wings, and from parvifusca by its oblique fasciae on the fore wings. The male genitalia are
very similar to those of loxotoma and, as in that species, lack the pair of coremata on the second
abdominal sternite.

DISTRIBUTION (Fig. 30). Costa Rica. Known only from the type-locality, Juan Vinas, and Tur-
rialba, about 20 km distant. Both localities are in Tropical Premontane Wet Forest.

MATERIAL EXAMINED

5 <J (2 cJ genitalia preparations).

Costa Rica: 1 <J, 1910 (Lankaster) (BMNH); 1 $ paratype, Cartago, Juan Vinas (Schaus) (BMNH); 2(J,
Cartago, Turrialba, 10.xi.1971 (Becker) (VB); 1 & Cartago, Turrialba, lO.v.1973 (Becker) (VB).

The leucocapna-group

cJ, $, 18-26 mm. Head whitish, vertex with fuscous or brownish scales. Third segment of labial palpus
one-third length of second, nearly straight. Thorax without crest of long scales, covered with fuscous and
whitish scales. Fore wing elongate; basal half of costa nearly straight, distal half evenly rounded, apex
acutely angled; veins free; dark fuscous, basal half with yellow scales between Rs and costa, diffuse whitish
area crossed with dark fuscous veins beyond cell. Hind wing golden-yellow. Abdomen golden-yellow with
dark fuscous scales on tergites in some specimens ; coremata located in pockets.

GENITALIA <$. Digitate processes of juxta not reaching anterior margin of gnathos apex.

REMARKS. This group can be easily distinguished from the monotonia-group and parvifusca by
the free veins of the fore wing, and from the albella-group by the dark fuscous fore wings.

Timocratica leucocapna (Meyrick) comb. rev.
(Figs 5, 8, 15, 31, 66, 100, 101, 159)

Lychnocrates leucocapna Meyrick, 1926: 227; Clarke, 1955: 224, pi. 112, figs 1-ld [adult, wing venation,

genitalia]. Holotype $, COLOMBIA (BMNH) [examined].
Timocratica leucocapna (Meyrick) Busck, 1935: 17 [catalogue].

$ 18-22 mm, 9 26 mm. Head whitish, frons light ochreous, progressively deep ochreous towards clypeus,
vertex suffused with fuscous. Haustellum covered with white scales at base. Second segment of labial palpus
ochreous, slightly tinged with fuscous below near distal articulations, white dorsally; third segment white,
dark fuscous below. Antenna with scape whitish, slightly tinged with ochreous, flagellum white basally, basal
half ochreous suffused with fuscous, then progressively ochreous towards apex. Thorax whitish, suffused
with fuscous ; patagium whitish ; tegula light fuscous. Fore wing dark fuscous, basal half between Rs and
costa ochreous with white scales ; a large, diffuse, whitish area beyond cell, crossed with dark fuscous veins ;
cilia dark fuscous with white dots near veins; underside ochreous. Hind wing light to golden-ochreous. Legs
ochreous, fore tibia and tarsus dark fuscous with white scales on articulations, mid and hind tarsi with dark
fuscous scales above. Abdomen ochreous, some specimens with third to sixth tergites dark fuscous.

GENITALIA £ (Figs 100, 101). Uncus with lateral margins nearly parallel, posterior margin concave, slightly
broadened in some specimens. Apex of gnathos pointed, bent posteriorly. Digitate processes of juxta straight
or bent gently inwards near apex, apex with several setae. Valva with margins nearly parallel or somewhat
broadened at middle. Aedeagus bent ventrad; vesica with two cornuti, a strong one and another smaller, on
the opposite side.

NEOTROPICAL GENUS TIMOCRATICA

239

GENITALIA $ (Fig. 159). Ventral margin of ostium bursae expanded posteriorly, falcate at middle. Antrum
slightly constricted anteriorly. Ductus bursae twisted posteriorly, nearly cylindrical, broadening progress-
ively towards corpus bursae. Corpus bursae nearly globular. Signum a single irregular plate.

REMARKS. T. leucocapna is easily distinguished from effluxa by the absence of fuscous on the hind
wing margins. The specimens from Turrialba, Costa Rica, have the abdomen plain golden-
ochreous ; some have the hind wings deep golden-ochreous.

DISTRIBUTION (Fig. 31). Costa Rica, Colombia, Peru and Venezuela. Despite its wide geogra-
phical distribution this species appears to be confined to a single Life Zone, Tropical Premontane
Wet Forest.

MATERIAL EXAMINED

16 cJ, 1 $ (5 cJ, 1 $ genitalia preparations).

Costa Rica: 1 <J, 1 $, Cartago, Turrialba, 600 m, 17-22. ii.1965 (Duckworth) (NMNH); 5 <J, Cartago,
Turrialba, IS.vii, 10.ix.1971, 10.iv.1972 (Becker) (VB; BMNH). Colombia: holotype & Cundinamarca,
Medina, 500 m (' 1650 ft') (BMNH); 2 <$, Cundinamarca, Medina (Fassl) (BMNH). Peru: 3cJ, La Oroya, R.
Inambari, 1000 m ('3100 ft'), iii, xi-xii.1906 (Ockenden) (BMNH). Venezuela: 2 £, Barinas, La Chimenea,
5 km south La Soledad, 1500 m, 28-29.V.1975 (Dietz) (VB; UCV); 1 & Lara, Anzoategui, Quebrada Guaco,
1440 m, 13-16.vi.1972 (Salcedo & Zambrano) (UCV).

•leucocapna
® species 2

A effluxa

12°C

24°C

Fig. 31 Geographical and ecological distribution of the leucocapna-group of Timocratica.

Timocratica species 2
(Figs 31, 65)

<$ 26 mm. Similar to leucocapna. Fore wing with the ochreous area on the basal half less pronounced ; area
beyond cell darker.

GENITALIA ^. Similar to those ofleucocapna.

REMARKS. The only specimen representing this form was collected in the same locality as three
specimens of typical leucocapna. It is larger than any male of the series representing leucocapna
and quite distinctive, but its genitalia are almost identical. As in effluxa, more material is necess-
ary to clarify this form.

DISTRIBUTION (Fig. 31). Peru (eastern slopes of the Andes), in Tropical Premontane Wet Forest,
a Life Zone where leucocapna also occurs.

MATERIAL EXAMINED
Peru: 1 <J, Puno, La Oroya, Rio Inambari, 1000 m (' 3100 ft '), iii. 1905 (Ockenden) (BMNH).

240 V. O. BECKER

Timocratica effluxa (Meyrick)
(Figs 3 1,64, 102, 103)

Lychnocrates effluxa Meyrick, 1930; 19. Holotype <J, BOLIVIA (BMNH) [examined].

Timocratica effluxa (Meyrick) Busck, 1935: 16 [catalogue]; Clarke, 1955: 388, pi. 194, figs 2-2b [adult,
genitalia].

<£ 20 mm. Head whitish, tinged with light ochreous; vertex with long brownish-fuscous scales. Second
segment of labial palpus ochreous, dark fuscous above; third segment white basally, with dark fuscous scales
towards apex. Antenna fuscous, scape with white scales, distal half of flagellum with ochreous scales. Thorax
fuscous, with white scales. Fore wing dark fuscous, veins darker than ground colour, sparsely mixed with
white scales; white streak poorly defined at base, between Rs and Sc; a diffuse white area crossed with
fuscous veins beyond cell, not reaching margins; cilia dark fuscous with white spots near veins; underside
ochreous, distal quarter fuscous. Hind wing ochreous, termen and dorsum fuscous; cilia fuscous. Legs
ochreous, fore tarsus dark fuscous, mid and hind tarsi with dark fuscous scales.

GENITALIA £ (Figs 102, 103). Similar to leucocapna.

REMARKS. T. effluxa is easily recognized by the fuscous borders of its hind wings. Although its
genitalia are almost identical to those of leucocapna it seems to be a distinct species. The colour
pattern of the wings in leucocapna shows little variation, the hind wings are plain golden-
ochreous and the fore wings have a distinctive ochreous area on the basal half between Rs and
the costa. All these features are absent in effluxa.

DISTRIBUTION (Fig. 31). Bolivia. The type-locality is in Subtropical Moist Forest.

MATERIAL EXAMINED
Bolivia: holotype <J, La Paz, Rio Songo, 750 m (Fassl) (BMNH).

The albella-group

cJ, $, 9-32 mm. Head, thorax and ground-colour of fore wing white. Third segment of labial palpus half to
same length as second. Thorax without crest of scales. Fore wing elongate, subrectangular or suboval; veins
free, or CuAl and CuA2, or R4 and R5, or both, stalked; plain white above except for melanostriga and
parvifusca; white below, tinged with golden-yellow, and/or fuscous along apex and termen. Hind wing white,
golden-yellow or tinged with yellow. Abdomen golden-ochreous, tinged with golden-yellow or white above,
white below; coremata located in pockets.

GENITALIA <$. Digitate processes of juxta often reaching anterior margin of gnathos apex.

REMARKS. The species of this group can easily be recognized by the white ground-colour of the
fore wings. T. parvifusca is the only fuscous species in this group but is easily separated* from
other fuscous species by the stalked R4 and R5 of the fore wing.

Timocratica grandis (Perty)
(Figs 32, 69, 104, 105, 160)

Yponomeuta grandis Perty, [1833]: 163, pi. 32, fig. [12] [legend of figure transposed with Pyralis bahiensis

Perty]. Holotype [<3 ?], BRAZIL: Piaui (Spix & Martius) (lost).
Cryptolechia grandis (Perty) Zeller, 1854: 378 [transcription]; Felder & Rogenhofer, 1875: pi. 139, fig. 56

[adult] jZeller, 1877: 260 [list].
[Cryptolechia bahiensis (Perty); Walker, 1864: 712 [catalogue; name quoted from figure legend in Perty,

[1833]: pi. 32, fig. 12].]

Stenoma grandis (Perty) Walsingham, 1913: 185 [catalogue].
Timocratica grandis (Perty) Busck, 1935 : 16 [catalogue].

cJ 22-26 mm. Frons yellowish-fuscous. Second segment of labial palpus dark fuscous, outer half above and
internally white, with yellowish-fuscous scales below ; third segment white, progressively dark grey internally
towards apex. Antenna with scape and base of flagellum white, progressively fuscous to apex. Legs golden-
ochreous, fore tibia and tarsus greyish-fuscous above. Fore wing with apex, termen and tornus evenly
rounded; all veins free; underside golden-yellow, slightly tinged with fuscous along apex. Hind wing golden-
yellow. Abdomen golden-ochreous, first tergite and all sternites white.
$ 28-30 mm. Fore wing broader than in male. Hind wing deep golden-yellow.

NEOTROPICAL GENUS TIMOCRATICA 241

GENITALIA <J (Figs 104, 105). Uncus slightly narrowed at base, apex strongly concave. Apex of gnathos
pointed. Digitate processes of juxta very long, reaching middle of gnathos, distal half of dorsal side with
sparse setae. Margins of valva nearly parallel. Aedeagus bent ventrad; vesica with cornutus undeveloped,
represented as a sclerotized area, and with many small acutely pointed spines.

GENITALIA $ (Fig. 160). Margin of ostium bursae expanded posteriorly, slightly concave at middle. Antrum
cylindrical. Ductus bursae straight, nearly cylindrical, posterior quarter sclerotized, with few longitudinal
wrinkles. Corpus bursae oblong, walls, as in ductus bursae, densely scobinated. Signum an elongate plate
constricted at middle, concave at both extremities.

REMARKS. T. grandis has golden-yellow hind wings and is thus very similar to bicornuta, constric-
tivalva and xanthotarsa. However, it can be easily distinguished from bicornuta by its fuscous fore
tibiae, and from the other two species by its fuscous fore tarsi.

One female from French Guiana has veins CuA± and CuA2 of the fore wing stalked.

In pi. 32 of Perty's work two species of Stenominae are illustrated. Fig. 12 represents a large
species with white fore wings and yellow hind wings, named Pyralis bahiensis; fig. 13 represents a
smaller species with pale wings and black markings on the fore wings, named Yponomeuta
grandis. However, as may easily be recognized from the descriptions, there is no doubt that the
legends were transposed and the large white species represents grandis, while the smaller rep-
resents bahiensis. With the exception of Walker (1864), all subsequent authors (Zeller, 1854, 1877,
Felder & Rogenhofer, 1875, Walsingham, 1913 and Busck, 1935), recognized fig. 12 as represen-
ting grandis.

In the BMNH and NMNH there were series totalling 25 specimens with white fore wings and
golden-yellow hind wings; these were identified as grandis and agreed with Perty's fig. 12. Upon
closer examination it was found that they represent four distinct species. In the absence of other
evidence it seems reasonable to apply the name grandis to the only species with golden-yellow
hind wings of this complex known to occur in the Amazon Basin of Brazil (type-locality of
grandis). According to Horn & Kahle (1936: 206), Perty's types were deposited in the ZSBS. Dr
Dierl informed me (pers. comm.) that the types cannot be found in that Museum and are believed
to have been destroyed during World War II.

DISTRIBUTION (Fig. 32). Brazil (Amazon Basin), French Guiana, Panama. Despite its wide distri-
bution, this species appears to be restricted to a single Life Zone, Tropical Moist Forest. It is
interesting that the other species with the hind wings and abdomen golden-ochreous, except for
bicornuta, were also collected in this Life Zone.

MATERIAL EXAMINED

1 1 c?, 6 $ (3 cJ, 1 $ genitalia preparations).

Brazil: 1 <J, Amazonas, Sao Paulo de Oliven9a (Staudinger) (MNHU); 1 <$, Para, Belem (' Para ') ([Bates])
(BMNH); 1 (J, 3 $, Para, Belem ('Para') (Moss) (BMNH). French Guiana: 1 & Guyanne, Cayenne (Felder)
(BMNH); 2 & 1 9, Guyanne, Cayenne (Deyrolle) (BMNH); 1 <J, Guyanne, Cayenne (BMNH); 2^, Guy-
anne, St Jean, R. Maroni (Le Moult) (BMNH); 2 9, Guyanne, R. Maroni (Bar) (BMNH). Panama: 2 <J,
Canal Zone, Barro Colorado Island, 10-1 7.v. 1964 (Duckworth) (NMNH).

Timocratica bicornuta sp. n.

(Figs 32, 110, 111, 161)

cJ ? 18-20 mm. Frons white, edged with fuscous. Second segment of labial palpus with proximal half tinged
with ochreous below, distal half fuscous, dark fuscous above, except near articulations ; third segment white,
dark grey internally. Antenna white; flagellum somewhat yellow towards apex. Legs golden-yellow, fore
tarsus dark fuscous above. Fore wing with apex rounded or somewhat angled, all veins free; underside
golden-yellow, slightly tinged with fuscous along apex and termen. Hind wing golden-yellow. Abdomen
golden-yellow, first tergite and all sternites white.

GENITALIA J (Figs 110, 111). Uncus with lateral margins parallel, apex concave. Apex of gnathos blunt.
Digitate processes of juxta bent outwards, diverging progressively from each other towards apex, with long
setae apically. Valva long, narrow, lateral margins nearly parallel. Aedeagus bent ventrad at basal third,
vesica with two strong bent cornuti opposite each other.

242

V. O. BECKER

GENITALIA ? (Fig. 161). Margin of ostium bursae with two small posteriorly directed lobes. Antrum very
broad medially. Ductus bursae somewhat broadened towards corpus bursae. Corpus bursae nearly globu-
lar, walls, as in ductus bursae, densely scobinate. Signum a rectangular plate.

REMARKS. T. bicornuta is easily separated from xanthotarsa and constrictivalva by its fuscous fore
tarsi, and from grandis by its yellow fore tibiae. It is also the only species in the group with two
strong cornuti in the vesica.

The only known female agrees in every detail with the males but is doubtfully associated with
this species.

DISTRIBUTION (Fig. 162). Brazil (south-eastern coast), French Guiana. The holotype of this
species, the only Brazilian specimen bearing detailed data, was collected in Subtropical Lower
Montane Wet Forest. The female from French Guiana came from Tropical Moist Forest, like the
others of the grandis complex. This difference in ecological adaptation may indicate that the
female belongs to a different species.

MATERIAL EXAMINED

3 J, 1 $ (3 cJ, 1 $ genitalia preparations).

Holotype & Brazil: Rio de Janeiro, Pico do Itatiaia, 28.iii-l.iv. 1958 (Kettlewell) (BMNH).

Paratypes. Brazil: 2 3 (Ragonot) (BMNH).

Excluded from types-series. French Guiana: 1 $, Guyanne, St Jean, R. Maroni (Le Moult) (BMNH).

• grandis

"" A xanthotarsa

• constrictiva/va
(•I species 3

A bicornuta

ire

24X

Fig. 32 Geographical and ecological distribution of the albella-group of Timocratica.

Timocratica xanthotarsa sp. n.

(Figs 32, 70, 106, 107)

cJ 21-22 mm. Frons white, edged with fuscous. Second segment of labial palpus ochreous below, basal
two-thirds black above, distal third and internal side white; basal third of third segment white, apical
two-thirds black. Antenna white, somewhat tinged with yellow towards apex. Legs golden-yellow above,
white below. Fore wing with apex, termen and tornus evenly rounded; all veins free; underside golden-
ochreous, slightly tinged with fuscous along apex. Hind wing golden-yellow. Abdomen golden-ochreous,
first tergite and sternites white.

GENITALIA <$ (Figs 106, 107). Uncus with lateral margins nearly parallel, apical margin concave. Apex of
gnathos long, strongly sclerotized. Digitate processes of juxta tapered towards apex, apex pointed, dorsal
side with sparse setae apically. Valva with basal third narrow, distal two-thirds wide, margins evenly
rounded; apex acutely rounded. Aedeagus slightly bent ventrad, vesica with strong curved cornutus.

REMARKS. T. xanthotarsa is easily distinguished from grandis and bicornuta by its golden-yellow
and white legs, and from constrictivalva by the unmodified gnathos.

NEOTROPICAL GENUS TIMOCRATICA 243

DISTRIBUTION (Fig. 32). Panama. The type-series was collected in Tropical Moist Forest.

MATERIAL EXAMINED

2 cJ (1 cJ genitalia preparation).

Holotype <J, Panama: Barro Colorado Island, 1-9. v. 1964 (Duckworth) (NMNH).

Paratype. Panama: 1 <$, Barro Colorado Island, l-9.v. 1964 (Duckworth) (NMNH).

Timocratica constrictivalva sp. n.

(Figs 32, 108, 109)

c? 21 mm. Frons white. Antenna with scape and basal half of flagellum white. Legs golden-yellow above,
white below. Fore wing with apex, termen and tornus evenly rounded; veins free; underside golden-yellow,
slightly tinged with fuscous along apex. Hind wing golden-yellow. Abdomen golden-ochreous, first tergite
and all sternites white.

GENITALIA J (Figs 108, 109). Basal two-thirds of uncus narrow, apical third broadened, apex concave.
Gnathos with two long, digitate, ventrally directed processes basally; apex triangular, dorsoventrally com-
pressed. Digitate processes of juxta long and narrow, widely separated, with long setae at middle. Basal third
of valva strongly constricted, distal two-thirds abruptly rounded. Aedeagus nearly straight, vesica with long,
strong, curved cornutus and few smaller spines opposite.

REMARKS. T. constrictivalva is very similar externally to xanthotarsa, but can be easily dis-
tinguished by its constricted valvae; from grandis and bicornuta it can be separated by the
absence of fuscous scales on the legs. Timocratica species 3 is possibly the female of this species
(see below).

The only specimen representing constrictivalva is not in very good condition. It lacks the palpi,
the right and half of the left antenna, one of each of the mid and hind legs, as well as most of the
scales on the thorax and remaining legs. Nevertheless, as the genitalia are so peculiar and
distinctive, it cannot be confused with any other species in the genus. It therefore seems justified
to name and describe it.

DISTRIBUTION (Fig. 32.). Ecuador (eastern side of the Andes). The only specimen was collected in
Tropical Moist Forest.

MATERIAL EXAMINED
Holotype <J, Ecuador: Pastaza, Sarayacu (Buckley) (BMNH).

Timocratica species 3

(Figs 32, 162)
$18 mm. Externally very similar to constrictivalva. Veins CuA^ and CuA2 stalked on fore wing.

GENITALIA 9 (Fig. 162). Margin of ostium bursae expanded posteriorly as two lobes. Antrum long, some-
what broadened at middle, with few strong longitudinal wrinkles. Ductus bursae constricted posteriorly,
broadening progressively towards corpus bursae. Corpus bursae wide, globular, walls, as in ductus bursae,
densely scobinated. Signum a single subrectangular plate weakly sclerotized along middle.

REMARKS. The specimen considered here is externally very similar to constrictivalva and xan-
thotarsa, and may well represent the female of the former species, but it differs by the stalked veins
CuA{ and CuA2 of the fore wing.

DISTRIBUTION (Fig. 32). Peru (eastern side of the Andes). The single specimen was collected in
Tropical Moist Forest.

MATERIAL EXAMINED
Peru: 1 $, Loreto, Iquitos (Strecker) (NMNH).

Timocratica subovalis (Meyrick)
(Figs 33, 112, 113)

Stenoma subovalis Meyrick, 1932: 304; Busck, 1935: 58 [catalogue]. Holotype J, BRAZIL (NMNH)
[examined].

244 V. O. BECKER

Stenoma stomatocosma Meyrick, 1932: 304. Holotype <$, BRAZIL: (NMNH) [examined]. Syn. n.
Timocratica stomatocosma (Meyrick) Busck, 1935: 17 [catalogue].
Timocratica subovalis (Meyrick) Duckworth, 1962: 113.

cJ 16-17 mm. Frons white, edged with fuscous. Second segment of labial palpus golden-ochreous below,
basal half dark fuscous above, white internally and at distal articulation ; third segment white with fuscous
scales towards apex. Antenna ochreous with fuscous scales, scape white. Fore coxa golden-yellow below,
tibia and tarsus dark fuscous ; mid and hind tarsi golden-yellow. Fore wing with costa evenly rounded, apex
rounded, somewhat angled, termen and tornus evenly rounded; veins free; underside of both wings golden-
yellow above cell. Hind wing white above. Abdomen golden-ochreous, first tergite, anal tuft and sternites
white.

GENITALIA <$ (Figs 112, 113). Uncus slightly broadened basally or with lateral margins nearly parallel; apex
strongly concave, nearly bifurcate. Apex of gnathos short, pointed. Digitate processes of juxta well separated
basally; distal half progressively pointed, covered with long setae dorsally. Valva wide, dorsal margin
straight, ventral margin evenly rounded. Aedeagus bent ventrad, vesica with long, strong, bent cornutus and
many smaller, pointed spines opposite.

REMARKS. T. subovalis is the only species with a golden-ochreous abdomen and white hind wings
that has the mid and hind tarsi golden-yellow.

The holotype of stomatocosma is an anomalous specimen in which M3 and CuAv are shortly
stalked in the right fore wing and stalked from the middle in the left. Meyrick considered it a
distinct species, probably because of this feature; since both holotypes agree in all other details,
including genitalia, they are considered here to be conspecific.

DISTRIBUTION (Fig. 33). Brazil (Amazon Basin). The two type-localities belong to the same Life
Zone, Tropical Moist Forest.

MATERIAL EXAMINED

Brazil: holotype <$ of S. subovalis, Amazonas, Ponte Nova, Rio Xingu (NMNH); holotype <$ of S.
stomatocosma, Tefe, ix (Fassal) (NMNH).

Timocratica species 4
(Figs 33, 177)

9 17 mm. Head white. Second segment of labial palpus white, basal half dark fuscous above; third segment
white, progressively fuscous towards apex. Legs white; fore coxa and femur golden-yellow above, tibia and
tarsus dark fuscous below ; mid and hind tarsus golden-yellow below. Fore wing with costa evenly rounded,
apex angled, termen and tornus obliquely rounded; R4 and Rs connate basally, CuAl and CuA2 stalked at
basal fourth; underside golden-yellow, slightly tinged with fuscous along apex. Hind wing slightly tinged
with golden-yellow above, deeper towards apex, underside golden-yellow above cell and along termen.
Abdomen golden-ochreous, first tergite, anal tuft and sternites white.

GENITALIA $ (Fig. 177). Margin of ostium bursae slightly concave at middle. Antrum straight, nearly
cylindrical. Ductus bursae nearly cylindrical, posterior third thickened, wrinkled, strongly scobinate; an-
terior two-thirds wrinkled longitudinally. Corpus bursae globular. Signum an irregular, strongly concave
plate.

REMARKS. The specimen described here is the only one that combines an ochreous abdomen and
golden-yellow hind wings with fore tarsi which are white above.

DISTRIBUTION (Fig. 33). Brazil (Amazon Basin): in Tropical Moist Forest.

MATERIAL EXAMINED
Brazil: 1 $, Para, Belem (' Para ') (Moss) (BMNH).

Timocratica amseli Duckworth sp. rev.
(Figs 33, 116, 117, 179)

Timocratica! albella Amsel, 1956: 306, pi. 63, fig. 6, pi. 107, fig. 8. Holotype <$, VENEZUELA (ZSBS) [exam-
ined]. [Junior secondary homonym of Depressor ia ( Volucra) albella Zeller, 1839.]

Timocratica amseli Duckworth, 1962: 113. [Objective replacement name for Timocratical albella Amsel,
1956.]

[Timocraticaxanthosoma(Dognm); Duckworth, 1966: 197(partim). Misidentification.]

NEOTROPICAL GENUS TIMOCRATICA

245

cJ 13 mm, $ 17 mm. Head white. Basal two-thirds of labial palpus ochreous externally, basal half dark
fuscous above, distal third and internally white; third segment white basally, progressively fuscous towards
apex. Fore coxa golden-yellow below, femur and basal two-thirds of tibia golden-ochreous above, distal
third of tibia and tarsus dark fuscous; mid tarsus tinged with ochreous below. Fore wing with basal third of
costa gently arched, distal two-thirds nearly straight; apex rounded, somewhat angled; termen and torn us
obliquely rounded; veins free; basal third of costa tinged with golden-yellow below. Hind wing white.
Abdomen golden-ochreous above, first tergite and sternites white.

GENITALIA $ (Figs 116, 117). Uncus somewhat broadened at middle. Apex of gnathos narrow, nearly
pointed. Digitate processes of juxta very long, compressed laterally at base, distal half with several long setae
dorsally. Margins of valva parallel, evenly rounded. Aedeagus strongly bent ventrad at basal third, vesica
with strong cornutus.

GENITALIA 9 (Fig. 179). Margin of ostium bursae slightly concave at middle. Antrum wide, anterior half
narrowing progressively towards ductus bursae, strongly wrinkled. Ductus bursae widening progressively
towards corpus bursae. Corpus bursae pear-shaped. Signum an irregular, sclerotized plate, concave across
middle.

REMARKS. T. amseli is easily distinguished from xanthosoma by its white fore wing costa, and from
fuscipalpalis by the ochreous tinge on the second segment of the labial palpus (almost dark
fuscous in fuscipalpalis).

Duckworth (1966: 197) synonymized this species with xanthosoma, but my examination of the
types of both species has shown them to be distinct; this is supported by their different ecological
distribution.

DISTRIBUTION (Fig. 33). Northern Venezuela, in Tropical Dry Forest.

MATERIAL EXAMINED

1 (J, 1 $ (1 J, 1 9 genitalia preparation).

Venezuela : holotype cJ, Distrito Federal, Caracas, Los Venados, vi-viii. 1937 (Vogl) (ZSBS); 1$ paratype;
Distrito Federal, Caracas, Berg Avila, 1000 m, vi-vii (Vogl) (ZSBS).

A amseli

+ species 4
20 A fuscipalpa/is

• x. leucocephala

®x. xanthosoma
-"> • subova/is

H ven/furcata

•7 - Ik

12°C

24°C

Fig. 33 Geographical and ecological distribution of the albella-group of Timocratica.

Timocratica venifurcata sp. n.

(Figs 21, 33, 78, 120, 121)

c? 16-17 mm. Head white. Second segment of labial palpus white, basal two-thirds dark grey externally;
third segment white, with grey scales near apex. Legs white, distal half of fore tibia, and tarsi dark fuscous
below. Fore wing with costa gently arched, apex rounded, termen and tornus obliquely rounded; R4 and
Rs, and CuAr and CuA2, stalked; basal half of costa with fuscous and golden-yellow scales below. Hind
wing white. Abdomen golden-ochreous, first tergite, anal tuft and sternites white.

246 V. O. BECKER

GENITALIA <£ (Figs 120, 121). Uncus wide, lateral margins nearly parallel, apex bifurcate. Apex of gnathos
narrow, pointed. Digitate processes of juxta flat, triangular, distal half with long sparse setae. Valva with
basal half wide, narrowing progressively towards apex. Aedeagus bent ventrad, vesica with strong, short
cornutus and many smaller spines opposite.

REMARKS. T. venifurcata is the only white species which has an ochreous abdomen and #4 and
R5 stalked on the fore wings (Fig. 21). It can be easily distinguished from all others with an
ochreous abdomen by the lack of ochreous coloration on its palpi and legs.

DISTRIBUTION (Fig. 33). Brazil (Central Plateau), in Tropical Premontane Moist Forest.

MATERIAL EXAMINED

2 <$ (1 <$ genitalia preparation).

Holotype & Brazil: Distrito Federal, Planaltina, 1000 m, 9.xi.l977 (Becker) (MN).

Paratype. Brazil: 1 <J, Distrito Federal, Planaltina, 1000 m, ll.xi. 1976 (Becker) (BMNH).

Timocraticafuscipalpalis sp. n.

(Figs 33, 114, 115)

cJ 12 mm. Frons white, edged with fuscous. Second segment of labial palpus dark fuscous, tinged with
ochreous basally below, white internally and at distal articulation; third segment white, fuscous internally.
Antenna light ochreous, scape white. Fore coxa tinged with golden-yellow below; mid and hind legs white.
Fore wing with basal third of costa gently arched, apex rounded, termen straight, tornus rounded; veins
free; costa golden-yellow below. Hind wing white. Abdomen golden-ochreous above, first tergite and
sternites white.

GENITALIA 3 (Figs 114, 115). Lateral margins of uncus nearly parallel, apex strongly concave, nearly
bifurcate. Apex of gnathos broad, rounded. Digitate processes of juxta very long, external margins nearly
straight, distal half of internal margins sinuate, distal two-thirds with sparse, irregular row of setae. Dorsal
margin of valva straight, ventral margin with basal third sinuate, middle third parallel to dorsal margin,
distal third converging progressively towards apex; apex acute. Aedeagus bent ventrad, vesica with single,
strong, bent cornutus.

REMARKS. T. fuscipalpalis is very close to amseli but is easily distinguished by the almost dark
fuscous second segment of the labial palpi. It can also easily be distinguished from venifurcata by
the free veins on the fore wing, from subovalis by the plain white hind wings, and from xan-
thosoma by the white costa.

DISTRIBUTION (Fig. 33). Southern Venezuela, in Tropical Premontane Moist Forest, a Life Zone
not shared by its closest relatives, amseli and xanthosoma.

MATERIAL EXAMINED

Holotype £, Venezuela: Bolivar, Guayaraca, Auyan Tepui, 1100m, 14.iv.1956 (Fernandez & Rosales)

(NMNH).

Timocratica xanthosoma (Dognin)

Stenoma xanthosoma Dognin, 1913: 416.

Timocratica xanthosoma (Dognin) Duckworth, 1966: 197 (partim) [synonymy].

c? 10-17 mm, $ 13-18 mm. Frons white, edged with golden-yellow; vertex golden-yellow or white. Second
segment of labial palpus white, basal third dark grey above; third segment white with dark grey scales.
Antenna white with dark fuscous scales. Legs white; fore coxa golden-yellow below; fore tibia, fore and mid
tarsi dark fuscous with white scales. Fore wing with base of costa arched, distal two-thirds nearly straight,
apex rounded, termen and tornus obliquely rounded; veins free or CuA^ and CuA2 stalked; basal third of
costa dark fuscous or black; white below. Hind wing white. Abdomen golden-ochreous, first tergite and
sternites white.

GENITALIA <J (Figs 118, 119). Uncus narrow, lateral margins nearly parallel, apex concave. Apex of gnathos
long, narrow. Digitate processes of juxta with lateral margins nearly parallel, apex with few short setae
dorsally. Valva narrow, somewhat broadened basally or with margins nearly parallel. Aedeagus slightly
bent ventrad, ventral side of apex with two lateral, pointed projections; vesica with single, long, curved
cornutus.

NEOTROPICAL GENUS TIMOCRATICA 247

GENITALIA 9 (Fig. 1 76). Margin of ostium bursae expanded posteriorly, strongly concave at middle. Antrum
wide, wrinkled anteriorly. Ductus bursae cylindrical, expanded progressively towards corpus bursae.
Corpus bursae reniform. Signum a transverse, irregular plate, slightly constricted at middle.

REMARKS. T. xanthosoma can be easily distinguished from other species with an ochreous abdo-
men and white hind wings by the dark fuscous or black basal half of its fore wing costa, and from
venifurcata by the free veins R4 and Rs .

Duckworth (1966: 197) synonymized amseli with xanthosoma and added Guatemala, Panama
and Colombia to the distribution. After examining his material it was found that amseli is a good
species and the specimens from Panama represented a subspecies of xanthosoma, described below
as leucocephala. The material from Guatemala was not available for study.

DISTRIBUTION (Fig. 33). Colombia, French Guiana, Panama. Both subspecies appear to be re-
stricted to Tropical Moist Forest and Tropical Wet Forest.

Timocratica xanthosoma xanthosoma (Dognin)
(Figs 33, 176)

Stenoma xanthosoma Dognin, 1913: 416; Meyrick, 1925: 192 [synonymy]; Busck, 1935: 60 [catalogue].

Holotype <J, FRENCH GUIANA (NMNH) [examined].
Stenoma sacra Meyrick, 1918: 209. Holotype 9, FRENCH GUIANA (BMNH) [examined]. [Synonymized by

Meyrick, 1925: 192.]
Timocratica xanthosoma (Dognin) Clarke, 1955: 392, pi. 196, figs 3-3c [adult, genitalia of holotype of sacra

Meyrick].

cJ 10-14 mm, 9 13-16 mm. Vertex golden yellow.

REMARKS. T. xanthosoma xanthosoma has the vertex golden-yellow, whereas in x. leucocephala it
is white. The nominate subspecies is also smaller on average than the latter, and the white scales
on the palpi, tarsi and antennae predominate over the black, making these appendages look
lighter.

DISTRIBUTION (Fig. 33). French Guiana.

MATERIAL STUDIED

5 c?, 2 9 (4 (J, 2 9 genitalia preparations).

French Guiana: holotype $ of S. xanthosoma, St Laurent du Maroni (NMNH); holotype 9 of S. sacra, R.
Maroni, 1916 (Le Moult) (BMNH); 4 <J, 1 9, St Jean du Maroni (Le Moult) (BMNH).

Timocratica xanthosoma leucocephala subsp. n.

(Figs 33, 77, 118, 119)
Timocratica xanthosoma (Dognin); Duckworth, 1966: 197(partim) [synonymy].

c? 14-17 mm, 9 18 mm. Vertex white. Antenna white mixed with black scales. Fore femur, fore and mid
tibiae above, and tarsi black, with scattered white scales mainly on the mid and hind tarsi.

REMARKS. T. xanthosoma leucocephala is distinguished from the nominate subspecies by the
white vertex. It is also larger on average and the black scales on the palpi, antennae and tarsi
predominate over the white so that these appendages look darker.

DISTRIBUTION (Fig. 33). Colombia and Panama.

MATERIAL EXAMINED

6 (J, 1 9 (3 ^, 1 9 genitalia preparations).

Holotype <$, Panama : Canal Zone, Barro Colorado Island, 5 -10.iv.1965 (Duckworth) (NMNH).

Paratypes. Colombia: 1 J [Bogota?] (Nolcken) (BMNH); 1 J, Choco, Juntas, Rio San Juan, 100m
('400 ft'), ii.1909 (Palmer) (BMNH). Panama: 3 & 1 9, Canal Zone, Barro Colorado Island, 10-28.iv.1964,
5-10.iv.1965 (Duckworth) (NMNH).

248 V. O. BECKER

Timocratica anelaea (Meyrick)
(Figs 34, 7 1,1 22, 123)

Stenoma anelaea Meyrick, 1932: 305. Holotype <$, BRAZIL (NMNH) [examined].
Timocratica anelaea (Meyrick) Busck, 1935: 16 [catalogue].

<J 25-26 mm. Frons white, edged with golden-yellow. Second segment of labial palpus ochreous, basal half
dark grey above; third segment black. Antenna white. Fore coxa and tibia with dark grey scales above, fore
and mid tarsi black. Fore wing elongate, costa gently arched, apex angled, termen and tornus obliquely
rounded; veins free; white below. Hind wing white. Abdomen golden-ochreous with white transverse bands
on articulations, first tergite, anal tuft and sternites white.

GENITALIA <$ (Figs 122, 123). Uncus with lateral margins nearly parallel, apex strongly concave. Apex of
gnathos broad, triangular. Digitate processes of juxta long, broad basally, narrowing progressively towards
apex ; apex pointed, distal half with long setae above, few setae on ventral side. Margins of valva evenly
rounded, nearly parallel, somewhat convergent from distal third to apex. Aedeagus bent ventrad at base,
vesica with strong elongate, bent cornutus and many minute spines.

REMARKS. T. anelaea is the only species in the group with an ochreous abdomen transversely
banded with white on the articulations.

DISTRIBUTION (Fig. 34). Brazil (Amazon Basin). Both localities are in Tropical Moist Forest.

MATERIAL EXAMINED

3 cJ (2 c? genitalia preparation).

Brazil: holotype cJ, Amazonas, Ponte Nova, Rio Xingu (NMNH); 2 <J, Para, Belem ('Para') (Moss)
(BMNH).

Timocratica titanoleuca sp. n.

(Figs 34, 73, 124, 125)

cJ 27-28 mm. Frons white, edged with fuscous. Second segment of labial palpus dark fuscous below and
externally, white internally and near distal articulation; third segment white basally, dark fuscous towards
apex. Antenna white. Fore coxa above, mid and hind tarsi golden-ochreous; fore tibia and tarsus white
above, dark fuscous below. Wings below golden-ochreous above cell and on veins, fore wing with costa
gently arched; apex angled, somewhat pointed; termen straight, oblique; tornus rounded; veins free. Abdo-
men white, somewhat tinged with cream above.

GENITALIA <$ (Figs 124, 125). Uncus with lateral margins nearly parallel, basal third slightly broadened, apex
strongly concave. Apex of gnathos narrow, pointed, strongly sclerotized. Digitate processes of juxta
broadened basally, narrowing progressively towards apex, distal half with row of long setae dorsally.
Ventral margin of valva evenly rounded, dorsal margin nearly straight. Aedeagus slightly bent ventrad at
middle, vesica with strong bent cornutus and many spines of different sizes opposite.

REMARKS. T. titanoleuca is very similar externally to macroleuca and leucorectis, but can be easily
distinguished by the absence of ochreous colouring on the labial palpi.

DISTRIBUTION (Fig. 34). Peru (eastern side of Andes). Both localities are in Tropical Premontane
Wet Forest.

MATERIAL EXAMINED

2 c? (2 3 genitalia preparations).

Holotype & Peru: Puno, La Oroya, R. Inambari, Carabaya, 1000m ('3100ft'), ix.1905 (Ockenden)
(BMNH).

Paratype. Peru: 1 <J, Huanuco, Tingo Maria, 2.xi.l949 (Allard) (NMNH).

Timocratica macroleuca (Meyrick)
(Figs 34, 72, 130, 131)

Stenoma macroleuca Meyrick, 1932: 304. Holotype J, BOLIVIA (NMNH) [examined].
Timocratica macroleuca (Meyrick) Busck, 1935: 17 [catalogue].

NEOTROPICAL GENUS TIMOCRATICA

249

<J 27-30 mm. Frons white, edged with fuscous. Second segment of labial palpus white, basal half ochreous
below, dark grey above ; third segment white, with few black scales on apex. Antenna white. Fore coxa and
tibia ochreous above; fore tarsus white above, dark fuscous below; mid and hind tarsi golden-ochreous.
Fore wing with costa gently arched; apex angled, somewhat pointed; termen straight, oblique; torn us
rounded; veins free; underside golden-yellow, deep golden-yellow along costa and veins. Hind wing below
golden-yellow along costa and on veins. Abdomen tinged with golden-yellow above, becoming progressively
white towards base, white below.

GENITALIA <$ (Figs 130, 131). Uncus with lateral margins nearly parallel, somewhat broadened at basal
two-thirds, apex concave. Apex of gnathos short, strongly sclerotized, rounded. Digitate processes of juxta
very long, narrow, distal two-thirds with sparse setae. Margins of valva evenly rounded, nearly parallel.
Aedeagus nearly straight, vesica with strong bent cornutus and many spines of different sizes.

REMARKS. T, macroleuca is very similar externally to leucorectis and titanoleuca, but can be easily
distinguished from titanoleuca by the ochreous tinge of the second segment of its labial palpi and
from leucorectis by the fore tarsi, which are white above.

DISTRIBUTION (Fig. 34). Bolivia (eastern slopes of the Andes). The type-locality is in Subtropical
Moist Forest, transitional to Tropical Premontane Moist Forest.

MATERIAL EXAMINED

2 c? (2 J genitalia preparation).

Bolivia: holotype <J, La Paz, Rio Songo (Fassl) (NMNH); 1 £, La Paz, Rio Songo, 750 m (Fassl) (BMNH).

• /eucorec
•"' • species 5

^titanoleuca

®anelaea
30 ® macroleuca

A spinignafha

- 24°C

Fig. 34 Geographical and ecological distribution of the albella-group of Timocratica.

Timocratica leucorectis (Meyrick)
(Figs 34, 126, 127, 164)

Stenoma leucorectis Meyrick, 1925: 223. Lectotype <J, BRAZIL (BMNH), designated by Clarke (1955: 388)

[examined].
Timocratica leucorectis (Meyrick) Busck, 1935: 17 [catalogue]; Clarke, 1955: 388, pi. 194, figs 4, 4b [adult,

genitalia].

3 28-30 mm, $ 30-32 mm. Frons white, edged with fuscous in some specimens. Second segment of labial
palpus ochreous below, basal half dark grey above, distal half white above and internally; third segment
progressively black from basal third to apex. Antenna white. Distal half of fore tibia above, and fore tarsus
dark fuscous, mid and hind tarsi golden-yellow. Fore wing with costa gently arched, apex angled, somewhat
pointed; termen straight, oblique; tornus rounded; veins free; underside of wings above cell golden-yellow,
some specimens tinged with fuscous along apex and termen. Abdomen white.

GENITALIA <J (Figs 126, 127). Uncus with lateral margins nearly parallel, slightly constricted at basal third,
apex strongly concave. Gnathos very long, lateral arms nearly parallel from basal third towards apex, two

250 V. O. BECKER

digitate posteriorly directed processes at basal third ; apex rounded, strongly sclerotized. Digitate processes
of juxta very long; distal half somewhat bent outwards, with long setae. Margins of valva nearly parallel,
distal half of ventral margin evenly rounded. Aedeagus curved ventrad, vesica with strong cornutus and
several spines.

GENITALIA ? (Fig. 164). Lamella antevaginalis as two broad, triangular lobes. Ostium bursae wide, margin
nearly straight. Antrum bent at middle, anterior half with a few longitudinal wrinkles. Ductus bursae twisted
posteriorly, wrinkled longitudinally. Corpus bursae nearly globular, walls, as in ductus bursae, densely
scobinate. Signum a nearly circular, diffuse plate.

REMARKS. T. leucorectis is one of the largest species in the group, and very similar externally to
titanoleuca and macroleuca. It can be easily distinguished from the former by the ochreous
underside of the labial palpi and from the latter by the fuscous fore tarsi. The base of the gnathos
arms is modified as in spinignatha and constrictivalva but in leucorectis the digitate processes are
directed posteriorly.

DISTRIBUTION (Fig. 34). Brazil, Bolivia, Colombia, French Guiana and Peru. Despite its wide
geographic distribution this species appears to be confined to only two Life Zones, Subtropical
Moist Forest and Tropical Moist Forest.

MATERIAL EXAMINED

7 (J, 2 9 (4 cJ, 1 $ genitalia preparations).

Bolivia: 2 <$, La Paz, Rio Songo (Fassl) (BMNH). Brazil: lectotype 3, Minas Gerais, Leopoldina, 1924
(BMNH); 1 (J paralectotype, Para, Belem ('Para'), vii.1919 (Parish) (NMNH). Colombia: 1 <J, Putumayo,
Mocoa (Hopp) (MNHU). French Guiana: 1 <$, Guyanne, Maroni River, 60 m, viii.1904 (Schaus) (NMNH).
Peru: 1 & San Martin, Moyobamba (de Mathan) (BMNH); 1 $, Loreto, Pebas (de Mathan) (BMNH); 1 ?,
San Martin, Huallaga, Chambirayacu (de Mathan) (BMNH).

Timocratica spinignatha sp. n.

(Figs 34, 128, 129)

cJ 19-23 mm. Frons white, edged with fuscous. Second segment of labial palpus ochreous below, dark
fuscous above except distal quarter, distal quarter white above; third segment white, apical third dark
fuscous. Fore coxa golden-yellow, femora ochreous, basal half dark fuscous above; tarsus and distal half of
tibia dark fuscous; mid femur and tibia tinged externally with golden-ochreous, mid and hind tarsi
golden-ochreous. Fore wing with costa gently arched, apex angled, termen and tornus rounded; veins free;
underside golden-ochreous above cell and on veins, tinged with fuscous along apex and termen. Hind wing
golden-ochreous below above cell and on veins. Abdomen tinged with cream above, white below.

GENITALIA <$ (Figs 128, 129). Uncus constricted at middle, apex bifurcate. Gnathos arms expanded ventrad
at base, densely covered with minute spines, apex short, rounded. Digitate processes of juxta long, slender,
lateral margins nearly parallel, with sparse long setae, mainly along dorsal side. Valva with dorsal margin
straight except at base, ventral margin evenly rounded. Aedeagus bent ventrad at basal third, vesica with
bent cornutus and many spines of different sizes.

REMARKS. T. spinignatha is similar externally to leucorectis, macroleuca and titanoleuca but is
smaller. It differs externally from macroleuca and titanoleuca by the dark fuscous fore tarsi and
from leucorectis by the dark fuscous tinge on the fore femora. The modified gnathos makes the
male genitalia very distinctive.

DISTRIBUTION (Fig. 34). Peru. The type-series was collected in Tropical Premontane Wet Forest.

MATERIAL EXAMINED

4 3 (2 J genitalia preparations).

Holotype <J, Peru: Puno, La Oroya, R. Inambari, 1000 m ('3100 ft'), iii.1905 (Ockenden) (BMNH).

Paratypes."Peru: 3(J, Puno, La Oroya, R. Inambari, 1000 m ('3100 ft'), iii.1905 (Ockenden) (BMNH).

Timocratica species 5
(Figs 34, 163)

9 17 mm. Frons white, edged with fuscous. Second segment of labial palpus golden-yellow, basal half tinged
with dark fuscous above; third segment fuscous internally, white externally. Fore femur golden-yellow;

NEOTROPICAL GENUS TIMOCRATICA 251

basal half of fore tibia ochreous above, distal half, and tarsus dark fuscous; mid leg and hind tarsus tinged
with golden-yellow. Fore wing with costa evenly arched; apex, termen and tornus rounded; veins free;
underside golden-yellow above cell, tinged with fuscous along apex and termen. Hind wing white, underside
golden-yellow above cell. Abdomen white.

GENITALIA 9 (Fig. 163). Lamella antevaginalis expanded posteriorly as two lobes. Antrum very broad
posteriorly, anterior half funnel-shaped, wrinkled longitudinally. Ductus bursae broadening progressively
towards corpus bursae. Corpus bursae pear-shaped. Signum an irregular plate, strongly constricted at
middle.

REMARKS. This species is very similar externally to spinignatha, but is smaller and lacks the
fuscous tinge on the fore femora.

DISTRIBUTION (Fig. 34). Peru (eastern side of the Andes), in Tropical Moist Forest.

MATERIAL EXAMINED
Peru: 1 9, San Martin, Tarapoto (de Mathan) (BMNH).

Timocratica argonais (Meyrick)
(Figs 35, 132, 133, 170)

Stenoma argonais Meyrick, 1925: 224. Holotype $, BRAZIL (BMNH) [examined].

[Stenoma maturescens Meyrick, 1925: 223 (partim). Misidentification.]

Timocratica argonais (Meyrick) Busck, 1935: 16 [catalogue].

Timocratica argonias: Clarke, 1955: 387, pi. 193, figs 1-lc. [Incorrect subsequent spelling.]

c£ 20-23 mm, $ 20-25 mm. Head white, frons edged with fuscous. Second segment of labial palpus ochreous
below, basal two-thirds dark grey above, distal third above and internally white; distal half of third segment
progressively fuscous towards apex. Fore coxa and femur above, and tarsus dark fuscous; mid coxa and
femur tinged with golden-ochreous externally, tarsus ochreous; hind tibia tinged with golden-yellow exter-
nally, tarsus golden-yellow. Fore wing with costa gently arched, apex rounded, somewhat angled; termen
and tornus obliquely rounded; veins free; golden-ochreous below, slightly tinged with fuscous along apex
and termen. Hind wing golden-yellow above cell. Abdomen white.

GENITALIA $ (Figs 132, 133). Uncus with lateral margins nearly parallel, apex strongly concave. Apex of
gnathos long and narrow. Digitate processes of juxta broadened basally, narrowing progressively towards
apex, distal half with sparse setae above. Ventral margin of valva evenly rounded, dorsal margin straight
except at base. Aedeagus somewhat curved ventrad, vesica with strong, curved cornutus and many acutely
pointed spines of different sizes.

GENITALIA $ (Fig. 170). Margin of ostium bursae straight. Antrum long with some longitudinal wrinkles,
strongly bent at connection with ductus bursae. Ductus bursae broadening progressively towards corpus
bursae, walls slightly wrinkled. Corpus bursae oblong, walls plain and smooth. Signum an elongate plate
weakly sclerotized and smooth along middle.

REMARKS. T. argonais is very similar externally to maturescens, megaleuca and palpalis. However,
it can be easily distinguished externally from the first by the white inner surface of the labial
palpus, and from the others by the fuscous tinge along the apex and termen on the underside of
the fore wing. The large series from French Guiana and Guyana agrees well with the lectotype of
argonais but their conspecificity should be confirmed by males from the type-locality.

DISTRIBUTION (Fig. 35). Brazil, French Guiana and Guyana. All the known localities of this
species are in Tropical Moist Forest.

MATERIAL EXAMINED

37 J, 15 9 (2 c£, 4 $ genitalia preparations).

Brazil: holotype $, Amazonas, Fonte Boa, ii.1920 (Parish) (BMNH); 1 $, Amazonas, Fonte Boa, vii.1906
(Klages) (BMNH). French Guiana: 6 $, 1 $ (paralectotypes of S. maturescens), R. Maroni (Le Moult)
(BMNH; NMNH); 29 & 10 $, Nouveaux Chantier, i-x (Le Moult] (BMNH); 1 & St Jean du Maroni (Le
Moult) (BMNH). Guiana: 1 ?, Berbice, New River, 250 m ('750 ft'), 20.i-23.iii.1938 (Hudson) (BMNH).

252

V. O. BECKER

Timocratica maturescens (Meyrick)
(Figs 35, 134, 135)

Stenoma maturescens Meyrick, 1925: 223. Lectotype ^, FRENCH GUIANA (BMNH), designated by Clarke

(1955: 391) [examined].
Timocratica maturescens (Meyrick) Busck, 1935: 17 [catalogue] ; Clarke, 1955: 391, pi. 195, figs 1-lb [adult,

genitalia].

c? 9 20-22 mm. Head white, frons edged with fuscous. Second segment of labial palpus ochreous, basal third
tinged with dark grey above; third segment white with some fuscous scales on apex. Fore coxa and femur
above, and tarsus dark fuscous; fore coxa and femur below, and mid and hind tarsi ochreous. Fore wing
with costa gently arched, apex angled, termen and tornus rounded; veins free; underside ochreous above
cell, white or tinged with fuscous along apex and termen. Hind wing underside ochreous above cell.
Abdomen white.

GENITALIA £ (Figs 134, 135). Uncus with lateral margins nearly parallel, slightly broader basally, apex
strongly concave. Apex of gnathos broad, rounded. Digitate processes of juxta long, lateral margins nearly
parallel, with sparse long setae mainly along dorsal distal two-thirds. Ventral margin of valva evenly
rounded, dorsal margin nearly straight. Aedeagus nearly straight, vesica with strong, curved cornutus and
many acutely pointed spines of different sizes.

REMARKS. T. maturescens is very similar externally to argonais and palpalis but can be easily
distinguished by the lack of white on the second segment of the labial palpus. Meyrick described
this species from a series of 10 specimens, of which only nine were traced, eight in the BMNH and
one in the NMNH. Only one paralectotype is conspecific with the lectotype of maturescens; the
other seven belong to argonais.

DISTRIBUTION (Fig. 35). French Guiana, Colombia and Venezuela, in Tropical Moist Forest and
Tropical Premontane Wet Forest.

MATERIAL EXAMINED

7 cJ» 1 ? (3 (J, 1 ? genitalia preparations).

French Guiana: lectotype $, R. Maroni, 1916 (Le Moult) (BMNH); 1 <$ paralectotype, R. Maroni (Le
Moult) (BMNH); 1 & St Laurent, R. Maroni, 16.xi.1906 (Le Moult) (BMNH). Colombia: 2 J, Cundina-
marca, Medina (Fassl) (BMNH); 1 <J, Guainia, Rio Negro ('Ost Colomb.'), 800 m (Fassl) (BMNH). Ven-
ezuela: 1 9, Amazonas, San Carlos de Rio Negro, 125 m, 1 9-3 l.viii. 1976 (Salcedo & Fernandez) (UCV); 1 $,
Bolivar, Rio Guaniamo, 160 m, 25-28.V.1979 (Clavijo, Chacon & Fernandez) (UCV).

® argonais

• maturescens

• mega/euca
A palpalis

12°C

- 24°C

Fig. 35 Geographical and ecological distribution of the albella-group of Timocratica.

NEOTROPICAL GENUS TIMOCRATICA 253

Timocratica megaleuca (Meyrick)
(Figs 35, 169)

Stenoma megaleuca Meyrick, 1912: 711; 1925: 224 [remarks]. Holotype9, COLOMBIA (BMNH) [examined].
Timocratica megaleuca (Meyrick) Busck, 1935: 17 [catalogue]; Clarke, 1955: 391, pi. 195, figs 2-2b, 4a
[adult, genitalia].

$ 25 mm. Frons white, edged with fuscous. Second segment of labial palpus ochreous externally, basal half
dark grey above, white internally; third segment white, distal half black. Thorax light fuscous above, except
tegulae. Fore coxa above, distal half of tibia and fore tarsus dark fuscous; mid and hind tarsi ochreous. Fore
wing with costa gently arched, apex angled, termen and tornus obliquely rounded; veins free; ochreous
below, except along apex, termen and below cell. Hind wing white.

GENITALIA 9 (Fig. 169). Margin of ostium bursae slightly expanded posteriorly as two small lobes. Antrum
bent dorsally, posterior third cylindrical, anterior two-thirds narrowing progressively towards ductus
bursae. Ductus bursae widening progressively towards corpus bursae. Corpus bursae pear-shaped, walls
slightly wrinkled longitudinally as in ductus bursae. Signum an elliptical plate without spines along middle.

REMARKS. T. megaleuca is extremely similar to palpalis and possibly synonymous, as suggested by
their ecological distribution. The genitalia of the only known specimen, the female holotype,
differ very slightly from those of palpalis. However, as no males are available and, as the geo-
graphic distribution does not agree with that of palpalis, it seems preferable to retain it as a
distinct species.

Meyrick (1925: 224), commenting upon his original description stated, "Thorax and abdomen
of original type (still unique) described as white (I supposed them to be discoloured); actually the
thorax is tinged greyish-ochreous except patagia, abdomen suffused pale greyish-ochreous be-
coming greyer posteriorly; I now think this colouring may be natural, but am not certain; there is
nothing of the sort in any of the allied species. Otherwise the species is nearest auxoleuca
[palpalis]."

DISTRIBUTION (Fig. 35). Colombia (known only from the type-locality), in Tropical Lower Mon-
tane Moist Forest.

MATERIAL EXAMINED
Colombia: holotype 9, Cauca, Popayan, 1906 (L.) (BMNH).

Timocratica palpalis (Zeller)
(Figs 4, 7, 13, 35^5, 136, 137, 168)

Cryptolechia (Cryptolechia) palpalis Zeller, 1877: 275. Holotype <^, BRAZIL (MNHU) [examined].

Stenoma auxoleuca Meyrick, 1925: 223. Lectotype ^, BRAZIL (BMNH), designated by Clarke, 1955: 387
[examined]. [Synonymized by Meyrick, 1926: 239.]

Timocratica haywardi Busck, 1938 : 280, figs 1-2. Holotype £, ARGENTINA : Entre Rios, Concordia (Hayward)
(NMNH) [not examined]. Syn. n.

Stenoma palpalis (Zeller) Meyrick, 1926: 239 [synonymy, distribution].

{Timocratica palpalis (Zeller);] Hempel, 1909: 68 [host, damage]; Ihering, 1909a: 228 [host, damage];
19096: 525 [host, damage].

[Stenoma albella (Zeller); Bondar, 1912: 15, figs 1-6, pi. 1 [host, damage, description]; Bondar, 1913: 24,
figs 17-20 [host, damage, description]; Lima, 1928: 161 [host]; Andrade, 1928 [host, damage]; Lima,
1930 [cat.]; Santos, 1934: 36 [host, damage]; Barbosa, 1933: 288, fig. 113 [host, damage]; Ronna,
1933: 332 [host, damage]; Fonseca, 1934: 228 [host]; Monte, 1934: 176, figs 161-162 [host, damage];
Ronna, 1934a [host, damage]; 19346 [host, damage]; Pyenson, 1938 [host, damage]; Carvalho &
Carvalho, 1939: 47 [hosts]; Lima, 1950: 1 [damage, control]; Silva & Heinrich, 1950: 9 [hosts]; Bertels,
1954: 61 [hosts]. Misidentifications.]

[Timocratica albella (Zeller); Lima, 1936: 277 [hosts] ; Araujo, 1937: 310 [host, control] ; Caldeira & Vieira,
1938 [host]; Biezanko & Freitas, 1938: 27 [catalogue, hosts]; Biezanko & Seta, 1939 [hosts]; Costa,
1942 : 248 [host, damage] ; Lima, 1945 : 269 [hosts, damage, description, genitalia] ; Lepage & Figueiredo,
1946 [hosts]; Duarte, 1947: 192 [host, damage, control]; Biezanko, Bertoldi & Baucke, 1949 [hosts];
Lofti, 1949: 20 [host, damage, control]; Robbs, 1953: 80 [host]; Costa, 1958: 139 [host, damage];
Robbs, 1960: 91 [host, damage]; Biezanko, 196 la: 12 [hosts]; 19616: 6 [host]; Mariconi & Soubihe,

254

V. O. BECKER

1961: 35 [host, damage]; Maranhao, 1962: 9 [host]; Pinheiro, 1962: 248 [host]; Mariconi, 1963: 389,

figs 178C-D [hosts, damage, description, control]; Sefer, 1963: 42 [host]; Silva et alii, 1968 [hosts];

Gallo et alii, 1970: 570 [hosts, damage]. Misidentifications.]
Timocratica palpalis (Zeller) Busck, 1935: 17 [catalogue].
Timocratica auxoleuca (Meyrick) Clarke, 1955: 387, pi. 193, figs 2-2b [adult, genitalia]; Hayward, 1969: 72

[hosts].

<$ 14-24 mm, ? 19-25 mm. Frons white, edged with fuscous. Second segment of labial palpus ochreous
below except near distal articulation, basal half dark grey above, white internally and near distal articu-
lation; third segment white, distal half becoming progressively dark grey towards apex. Fore coxa, femur
and basal half of tibia ochreous above, femur tinged with dark grey above, distal half of tibia, and tarsus
dark greyish-fuscous ; mid and hind tarsi ochreous. Fore wing with costa gently arched, apex angled, termen
and tornus obliquely rounded; veins free; underside above cell and veins golden-ochreous, except along
apex and termen. Hind wing and abdomen white.

GENITALIA <$ (Figs 136, 137). Uncus with lateral margins parallel or narrowing slightly towards apex, apex
concave. Apex of gnathos wide, rounded. Digitate processes of juxta wide basally, narrowing towards apex,
distal two-thirds with long setae dorsally. Valva with margins nearly parallel or somewhat broadened at
distal third, sacculus slightly pronounced. Aedeagus somewhat bent ventrad, vesica with strong bent cor-
nutus and several smaller spines.

37

Figs 36, 37 Timocratica palpalis (Zeller), last instar larva, Brazil, Santa Catarina, ex Psidium guajava. 36,
dorsal view; 37, lateral view.

NEOTROPICAL GENUS TIMOCRATICA 255

GENITALIA $ (Fig. 168). Margin of ostium bursae slightly concave at middle. Antrum long, posterior third
nearly cylindrical, anterior two-thirds narrowing progressively towards ductus bursae, with few longitudinal
wrinkles. Ductus bursae widening progressively towards corpus bursae. Corpus bursae pear-shaped, walls
strongly wrinkled as in ductus bursae. Signum a long elliptical plate, slightly constricted at middle, without
spines in central area.

PUPA. J, 9, length 15-19 mm, maximum diameter 4-5-5-5 mm. Indistinguishable from that of melanocost a.

LARVA (Figs 36-42). Maximum length 35 mm; cylindrical, dark pinkish violet; pinacula large, well defined,
slightly prominent, dark brown. Anal comb absent. Meso- and metathorax with extra sclerotized area,
'pinacula without setae', between setae L and setae SV1 ; abdominal segments 1-2 with two extra sclerotized
areas, 'pinacula', on each segment, one behind spiracle, between setae SD and setae L, the other behind
LI + L2, above L3; segments 3-7 with three extra sclerotized areas, two as in segments 1-2 and a third in
front of L3, above setae SV. Abdominal prolegs with 1 12-1 16 crochets in triordinal circle, anal prolegs with
58-62 crochets arranged triordinally in anal penelipse. Head hypognathous, nearly spherical, with irregular,
hexagonally sculptured surface, dark brown; adfrontal area not reaching to vertical angle; only primary
setae present; mandible with two small blunt teeth; ocellus V below antenna; adfrontal area slightly
prominent near clypeus; setae P2 closer to each other than setae PI. Pro thoracic plate prominent, strongly
sclerotized, dark brown, with irregularly sculptured area behind setae SD2; Dl equidistant to XD1 and D2,
below level of former, posterior to second; SD2 between XD2 and SD1, slightly posterior to both; MXD1,
MD1 and MSD1 present; LI, L2, L3 on same pinaculum; SV1 and SV2 on same pinaculum; spiracle
vertically elongated. Meso- and metathorax with Dl, D2, SD1 and SD2 on same pinaculum; Dl slightly
posterior to D2; pinaculum LI + L2 slightly connected with L3; SV1 below L3. Abdomen with spiracle on
segment 1 twice the size of others; setae Dl on segments 1-2 slightly closer to each other than setae D2,
further apart on 3-7; SD2 on segments 1-8 present but greatly reduced; SV3 absent on segments 1 and 7-9.

REMARKS. T. palpalis is easily distinguished from any other species from southern South
America, except isarga, by the dark fuscous fore tarsi and dark fuscous distal half of fore tibiae. T.
isarga has similarly coloured tarsi and tibiae but has the fore wings white below, CuA± and CuA2
stalked, and the hind tarsi white. T. megaleuca from Colombia is probably also a synonym of
palpalis, as discussed on p. 253.

As a large number of specimens were available for study, either collected at light or reared on
different hosts and from different places, some variation in size and genitalia was found. Speci-
mens from warmer areas such as the east, the Central Plateau and the coast of Brazil, south to the
lowlands of Santa Catarina, are on average larger than those from the Parana Plateau, south
Brazil and Argentina. Variation of the male genitalia occurs mainly in the shape of the valva and
digitate processes of the juxta. The valva may be slightly narrowed with the margins nearly
parallel, or have the distal half somewhat broadened and the sacculus slightly pronounced. In
some specimens the digitate processes of the juxta are broad, nearly triangular, with the margins
converging progressively towards the apex, whereas in others the distal half is very narrow.

This species, referred to as albella (Zeller) in the Brazilian economic literature, is the most
common species in the south of South America, mainly in southern Brazil and northern Argen-
tina, and is a pest of ornamental, fruit and timber trees. As the adults are almost white, it was
originally identified by Bondar (1912: 15) as Stenoma albella (Zeller) and this name has been used
by all subsequent authors ; at that time albella appeared to be the only available name for a large
white stenomine, since grandis has golden-yellow hind wings, and palpalis was still considered an
Indian species.

Although the holotype bears the label ' Bengal ', there is no doubt that it represents this South
American species, as pointed out by Meyrick (1926: 239).

BIOLOGY. The larvae of T. palpalis are polyphagous bark-feeders and considered to be pests of
ornamental, fruit and timber trees in Brazil and Argentina. They tunnel into the trunk and larger
branches of the host-plant, feeding on the bark surrounding the holes (Fig. 43). No branches of
less than 2 cm diameter were found infested. The tunnel is shallow, only 5-8 cm long and about
0.5 cm wide when the larva is ready to pupate. The tunnel begins more or less at right angles to
the axis of the wood and then follows the pith (Fig. 45). Larvae collected in Brusque, Santa
Catarina, were tunnelling the trunk upwards, whereas those collected in Sete Lagoas, Minas
Gerais, were tunnelling down towards the base of the tree. In the former locality it rains through-
out the year, making it necessary to tunnel upwards to avoid flooding of the hole, whereas in Sete

256

V. O. BECKER

Lagoas, in the Central Plateau of Brazil, the larvae develop during a well-defined dry season,
when there is no such danger. The larvae cover the eaten areas of the bark with frass (Fig. 44),
and so remain hidden when feeding outside the tunnel. No larvae were found feeding during the
day.

38

41

42

Figs 3&-42 Timocratica palpalis (Zeller), last instar larva. 38, setal map. 39, frontal view of head. 40, lateral
view of head. 41, dorsal view of last three abdominal segments. 42, inner surface of mandible.

NEOTROPICAL GENUS TIMOCRATICA

257

Fig. 43 Damage caused by larvae of Timocratica palpalis (Zeller) on the trunk of Tibouchina candolleiana.

258

V. O. BECKER

Pupation takes place inside the tunnel; the pupa is attached by the cremaster to a few strands
of silk on the tunnel wall.

Field observations and label data show that there are differences in the flight period of adults,
and specimens from warmer areas emerge earlier in the season than those from cooler places.
Most specimens from Minas Gerais were collected from early October to late February, those
from the lowlands of Santa Catarina, further south, emerged at the beginning of December,
specimens from Rio Grande do Sul were collected in January, while those bred by Hayward in
Argentina emerged in March. Possibly the species has two generations in warmer localities, but is
univoltine further south where the average temperature is lower.

T. palpalis is usually a minor pest, but when infestation is high it may seriously damage the
trees (Fig. 43). When the trunk or branch is ring-barked, the tree dies beyond that point. In some
Myrtaceae, such as guava (Psidium guajava L.), the bark never recovers and the trunk becomes
deformed where it was damaged. In Brasilia, strong infestation was found in a number of
Tibouchina candolleiana (Melastomataceae), an ornamental tree with attractive pink and violet
flowers. Some of the trees contained more than 100 larvae. Considering that one larva can
seriously damage or even kill a whole branch, such an infestation is very serious.

It seems that the preferred host-plants are Myrtaceae, mainly species of Psidium, and it is very
easy to find the larvae feeding on guava, a common fruit tree in South America. Table 2 gives a
list of the host plants of palpalis based on my own observations and the literature.

Table 2 Food plants of T. palpalis

Scientific names

English
vernacular names

Brazilian
vernacular names

ACERACEAE

Acer saccharinum
A. platanoides

Silver maple
Norway maple

CASUARINACEAE

Casuarina equisetifolia

Casuarina, Willow, Whistling pine

Casuarina

CUNONIACEAE

Belangera tpmentosa

Cangalheiro

EBENACEAE

Diospyros kaki

Kaki

Caqui

FAGACEAE

Castanea saliva

Sweet chestnut

Castanheira

Quercus robur

British oak

Carvalho-ingles

LAURACEAE

Persea america

Avocado pear

Abacateiro

MELASTOMATACEAE

* Tibouchina candolleiana
*T. urvilleana

Quaresmeira
Quaresmeira

MYRTACEAE

Calycorectes pohlianus
Campomanesia acida
Eucalyptus alba
E. camaldulensis
(= E. rostrata)
E. citriodora
E. propinqua
*E. saligna
E. tereticornis

Timor white gum
Murray red gum

Lemon scented spotted gum

Sydney blue gum, Saligna gum
Forest red gum

Cambucazeiro
Ara^a-do-Para
Eucalipto
Eucalipto

Eucalipto
Eucalipto
Eucalipto
Eucalipto

* Author's observations; others were quoted from Araujo et al. (1968) and Hayward (1969), and from label data.

English vernacular names follow Adams (1972) and Bailey (1900-02); Brazilian vernacular names of the
Myrtaceae follow Legrand & Klein (1967-78).

NEOTROPICAL GENUS TIMOCRATICA

259

Table 2 (continued)

Scientific names

English vernacular
names

Brazilian
vernacular names

Eugenia brasiliensis
E. uniflora
*E. involucrata

Hexachlamys edulis

Marlierea tomentosa
Myrcia fenzliana
Myrciaria trunciflora
*Psidium guajava
P. guineense
P. humile
Syzygium jambos
S. malaccense

PLATANACEAE

Platanus orientalis

PROTEACEAE

Macadamia ternifolia

PUNICACEAE

Punica granatum

ROSACEAE

Cydonia vulgaris
Eriobotrya japonica
*Malus domestica
M. sylvestris
Prunus amygdalus
P. armeniaca
P. domestica
P. persica
*Pyrus communis
*P. sinensis

RUBIACEAE

Coffea arabica

SALICACEAE

Salix viminalis

TILIACEAE

Luehea divaricata

ULMACEAE

Ulmus americana

Surinam cherry

Guava
Guiana guava

Rose apple
Otaheite apple

Oriental plane
Queensland nut
Pomegranate

Quince

Loquat

Apple

Crab apple

Almond

Common apricot

Common garden plum

Peach

Common pear

Sand pear, Japanese pear, Chinese pear

Arabian coffee
Osier willow

White elm

Grumixameira

Pitangueira

Cerejeira-de-

folha-miuda
Cereja-do-

Rio Grande
Garapuruna
Guamirim-ara9a
Jaboticabeira
Goiabeira
Ara9a-azedo
Ara9a-vermelho
Jambeiro
Jambeiro-vermelho

Platano-oriental

Macadamia

Romanzeira

Marmeleiro

Ameixeira-do-Japao

Macieira

Macieira-silvestre

Amendoeira

Abrico

Ameixeira

Pessego

Pereira

Pereira-do-Japao

Cafeeiro
Vimeiro
A9oita-cavalo

Sometimes the larvae are heavily parasitized. About 80 per cent of the larvae collected in
Brasilia on Tibouchina candolleiana were parasitized by an apparently undescribed species of
Eudeleboea Blanchard (Ichneumonidae). They may also be preyed on by birds. A wild guava
(Psidium sp.), found near Planaltina, Distrito Federal, at the beginning of September 1978 had
branches attacked by three larvae. However, it was found that each larval gallery had a fresh hole
near the middle, made by an unidentified species of woodpecker, through which the bird had
removed the larva. It is interesting to note that Psidium species have a very hard wood and the
bird had to make a hole 2 cm deep to reach the larvae.

DISTRIBUTION (Fig. 35). Northern Argentina, Bolivia, Brazil. This species has not only a wide
range of food-plant preference and geographical distribution, but also a wide range of ecological

260

V. O. BECKER

45

Figs 44-45 Damage caused by larvae of Timocratica palpalis (Zeller). 44, branch of Pyrus communis with
areas partially covered by frass (O. Mielke photo). 45, split branch of Psidium guajava showing larva inside
gallery.

range of food-plant preference and geographical distribution, but also a wide range of ecological
distribution. It has been collected from Warm Temperate Dry Forest, in the North Argentina
'Chaco' area, and Warm Temperate Moist Forest of southern Brazil and Argentina, crossing the
Subtropical Moist Forest of southern Brazil, up to the Tropical Moist Forest of the north-east
Brazilian coast. The high concentration of localities in Warm Temperate and Subtropical Moist
Forest Life Zones does not indicate that this species is chiefly associated with these Life Zones,
but probably means that the species has been more intensely collected there.

MATERIAL EXAMINED

71 (J, 31 9, 8 larvae, 4 pupae (9 $, 5 9 genitalia preparations).

Argentina: 5 <$ paratypes of T. haywardi, Entre Rios, Concordia, ex guava~[Psi#mm guajava L.], Hi. 1937
(Hayward) (BMNH; NMNH); 1 $ paratype of T. haywardi, Entre Rios, Concordia, ex pomegranate [Punica
granatum L.], iii.1938 (Hayward} (NMNH): 1 & Santa Fe, Villa Ana, iii.1924 (Hayward) (BMNH). Bolivia: 1
cJ, Nuflo de Chaves, Esperanza (BMNH). Brazil: holotype J of C. palpalis, ' Bengal ' (MNHU); lectotype <$
of S. auxoleuca, Espirito Santo, Leopoldina, 1924 (BMNH); 1 <£ no further data (BMNH); 1 <J, 2 9, Alagoas,
Maceio ['Maceo'] (BMNH); 1 <J, 2 9, Bahia, Salvador (Fruhstorfer) (BMNH); 3 <$, Distrito Federal,
Brasilia, lO.x.1979 (Gomes) (VB); 10 <J, 2 9, Distrito Federal, Planaltina, 5-25.xi.1975, 12.ii.1976, 6-
15.xi.1977, 20.ii.1978, 21.xi.1978 (Becker) (VB; BMNH; MNHU; NM; NMNH); 3 £, 8 larvae, 4 pupae,
Distrito Federal, Planaltina, 1000m, ex Tibouchina candolleiana, 9-16.xi.1978 (Becker) (VB; BMNH;

NEOTROPICAL GENUS TIMOCRATICA 261

NMNH); 1 9, Espirito Santo, 1910 (Fruhstorfer) (BMNH); 1 <$, Minas Gerais, Leopoldina (Staudinger)
(MNHU); 1 9, Maranhao (BMNH); 1 & Agua Suja, x.1906 (Baer) (BMNH); 2 & Minas Gerais, Cordis-
burgo, ex Psidium guajava, 5, 16.xi.1974 (Becker) (VB); 3 J, 2 9, Minas Gerais, Cordisburgo, ex Eugenia
involucrata, 3.x-23.xi.l974 (Becker) (VB; BMNH); 3 <J, Minas Gerais, Sete Lagoas, 720 m, 20.i, 10, 18.ii.1969
(Becker; Biezanko) (VB; LN); 4 <$, 2 9, Minas Gerais, Sete Lagoas, 720 m, ex Psidium guajava, 2.x-
14.xii.1974, 8.U.1975 (Becker) (VB; BMNH; NMNH); 7 & Parana, Castro, 1896-1898 (Jones) (BMNH); 1 &
1 9, Parana, Curitiba, 920m, ll.xii.1971, 2.ii.l974 (Becker) (VB); 2 <J, Parana, Mandirituba, 13.xii.1969
(Becker) (VB); 1 9, Pernambuco, Serra do Comunati (Gounelle) (BMNH); 1 <J, 1 9, Rio de Janeiro (BMNH);
1 (J, Rio Grande do Sul, Guarani, 711954 (Biezanko) (BMNH); 1 $, Rio Grande do Sul, Pelotas, 2911960
(Biezanko) (BMNH); 1 <J, Rio Grande do Sul, Pelotas, 2211965 (Guerra) (VB); 2 9, Rio Grande do Sul,
Santa Maria, ex Mains domestica, 16.ii.1979 (Link); 1 <$, 2 9, Rio Grande do Sul, Santa Maria, ex Pyrus
communis, 16.ii.1979 (Link); 1 <$, 3 9, Rio Grande do Sul, Santa Maria, ex Psidium guajava, 17.ii.1979 (Link);
1 (J, 1 9, Rio Grande do Sul, Santa Maria, ex Luehea divaricata, 28.ii.1979 (Link) (all VB); 1 cJ, 4 ?, Santa
Catarina, Brusque, ex Psidium guajava, 6-29.xii.1970 (Becker) (VB; BMNH; NMNH); 1 9, Santa Catarina,
Corupa, xii.1955 (Mailer) (NMNH); 1 & Santa Catarina, Rio Vermelho, vii.1954 (Mailer) (NMNH); 2 &
Sao Paulo, Sao Paulo, 1889 (Jones) (BMNH); 1 & 1 ?, Sao Paulo, 1910 (Ihering) (BMNH); 4 rf, Sao Paulo,
Piracicaba, 540 m, xii.1965-i.1966 (ESALQ).

Timocratica melanocosta sp. n.

(Figs 46-53, 138, 139, 173)

cJ 14-18 mm, 9 16-19 mm. Frons white. Second segment of labial palpus golden-ochreous, white internally
and around distal articulations; third segment black with white scales towards base. Fore coxa below, femur
and basal two-thirds of tibia golden-ochreous above; distal third of tibia, and tarsus dark fuscous below,
mixed with white scales above; distal joint of mid femur, and tibia golden-ochreous externally, mid tarsus
dark fuscous below, or white, proximal articulations of hind tibia slightly tinged with golden-ochreous. Fore
wing with costa strongly arched at base, then gently arched, apex angled, termen slightly obliquely rounded,
tornus rounded ; veins free; basal third of costa tinged with dark grey; white below. Hind wing and abdomen
white.

GENITALIA $ (Figs 138, 139). Lateral margins of uncus nearly parallel, apex strongly concave, nearly
bifurcate. Apex of gnathos nearly triangular. Digitate processes of juxta broadened basally, narrowing
progressively to basal third, then straight; distal half with sparse setae dorsally. Valva with margins nearly
parallel, sacculus slightly pronounced. Aedeagus nearly straight, vesica with strong cornutus and many
acutely pointed spines opposite.

GENITALIA 9 (Fig. 173). Margin of ostium bursae rounded. Antrum funnel-shaped, bent, with strong longi-
tudinal wrinkles. Ductus bursae broadening progressively towards corpus bursae. Corpus bursae pear-
shaped, walls slightly wrinkled as in ductus bursae. Signum a round or elliptical plate without spines at
middle.

PUPA (Figs 45-47). <$, length 16-17 mm, maximum diameter 4.5-5.0 mm. Brown, darker towards head.
Setae minute. Pronotum strongly expanded forwards, displacing pronotum-head suture to a ventral posi-
tion, longitudinally wrinkled. Meso- and metanotum irregularly wrinkled. Cremaster with pair of long
processes, directed cephalad, reaching sixth segment, apices with many small, hook-like setae.

LARVA. Maximum length 30 mm. Very similar to that of palpalis and almost indistinguishable from it,
except for the following characters : extra sclerotized area, 'pinaculum without setae ', behind spiracle greatly
reduced on abdominal segments 3-4, absent on segments 5-7.

REMARKS. T. melanocosta is nearest to palpalis, but easily distinguishable from it, and others in
the group, by the grey tinge along the basal third of the fore wing costa.

Specimens from Bananal Island, reared from Byrsonima sp., have the mid tarsus almost white
whereas most specimens, such as those from Minas Gerais and Distrito Federal, have tarsi which
are dark fuscous below. However, in all other respects, including genitalia, the specimens agree
very well.

BIOLOGY. The behaviour and feeding habits of the larvae of this species are almost the same as in
palpalis (Figs 50-52). Most of the specimens studied were obtained from larvae feeding on
Erythroxylum suberosum (Erythroxylaceae) and Byrsonima sp. (Malpighiaceae).

262

V. O. BECKER

Figs 46-48 Timocratica melanocosta sp. n., pupa. 46, ventral view. 47, lateral view. 48, dorsal view.

DISTRIBUTION (Fig. 53). Brazil (Central Plateau and dry areas of the southern border of the
Amazon Basin). Specimens were collected in Tropical Premontane Moist Forest and Tropical
Moist Forest. However, it is important to point out that its host plants are not part of a climatic
association, but are one of the components of the atmospheric monsoon-type of association
called 'cerrado' in Brazil (Fig. 49) (see discussion of this association under major, p. 231).

MATERIAL EXAMINED

34 cJ, 27 9, 8 larvae, 4 pupae (6 cJ, 5 9 genitalia preparations).

Holotype <J, Brazil: Distrito Federal, Planaltina, 1000 m, 9.xi.l977 (Becker) (MN).

Paratypes. Brazil: 4 ^, 3 $, Minas Gerais, Sete Lagoas, 720 m, ex Erythroxylum suberosum, 13.x-
18.xi.1974, 26.x. 1978 (Becker) (VB; BMNH); 2 rf, 1 ?, Distrito Federal, Brasilia, 1000 m, 10.x. 1979 (Gomes)
(VB); 8 (J, 2 ?, Distrito Federal, Planaltina, 1000m, 5.xi, 23.xi, 16.xii.1975, ll.xi.1977, 26.x- lO.xi. 1978
(Becker) (VB; BMNH; MNHU; NMNH); 19 <£, 21 9, 8 larvae, 4 pupae, Goias, Bananal I., ex Byrsonima sp.,
24.x-15.xi.1977 (fleeter) (VB; BMNH; LN; MNHU; NM;NMNH; ZSBS).

Timocratica nivea sp. n.

(Figs 22-24, 54, 140, 141, 171)

cJ 15-17 mm, 9 19-20 mm. Frons white. Second segment of labial palpus white with black scales basally
above; third segment white below, black above. Legs white; fore coxa tinged with golden-yellow, fore femur
above, tibia and tarsus dark grey below. Fore wing with basal third of costa arched, distal two-thirds nearly
straight; apex angled, termen and tornus obliquely rounded; veins free or CuA^ and CuA2 connate or
stalked; white below. Hind wing and abdomen white.

GENITALIA $ (Figs 140, 141). Lateral margins of uncus nearly parallel; apex strongly concave, nearly
bifurcate. Apex of gnathos pointed. Digitate processes of juxta with lateral margins parallel, narrowing

NEOTROPICAL GENUS TIMOCRATICA

263

progressively near apex, with sparse setae on distal half dorsally. Valva with margins nearly parallel or
somewhat broadened at middle, bent dorsally. Aedeagus bent ventrad at basal third, vesica with strong bent
cornutus and many smaller spines of different sizes opposite.

GENITALIA $ (Fig. 143). Margin of ostium bursae rounded. Antrum wide, somewhat narrowed towards
anterior end, with a few strong longitudinal wrinkles. Ductus bursae broadening progressively towards

Figs 49-52 Habitat and food-plant (Erythroxylum suberosum) of Timocratica melanocosta sp. n. 49, habi-
tat and food-plant. 50-52, branches of the food-plant showing (50) eaten areas of bark covered with frass,
(51) eaten area exposed with entrance hole of larval gallery, (52) split branch with gallery and pupa.

264 V. O. BECKER

corpus bursae, scobinate. Corpus bursae pear-shaped, finely scobinate. Signum a round plate without spines
at middle.

REMARKS. T. nivea is similar to melanocosta, albella and isarga but is easily distinguished by the
absence of ochreous coloration on its labial palpi. The male and female genitalia are very close to
those of melanocosta, except for the narrow base of the digitate processes of the juxta in the male.
The arrangement of the CuA veins is very variable. Of 1 1 specimens examined, two females and
four males have these veins free, two have them connate, two shortly stalked, and in one specimen
they are connate on the left wing and shortly stalked on the right (Figs 22-24).

DISTRIBUTION (Fig. 53). Brazil (Central Plateau). This species is sympatric with melanocosta (see
discussion on ecology under that species, p. 262).

MATERIAL EXAMINED

10 <J, 3 9 (3 c?, 2 $ genitalia preparations).

Holotype J, Brazil: Distrito Federal, Planaltina, 1000 m, 15.xi.1975 (Becker) (MN).

Paratypes. Brazil: 1 & Minas Gerais, Sete Lagoas, 720 m, 18.x. 1969 (Becker) (VB); 8 <$, 3 $, Distrito
Federal, Planaltina, 1000 m, 5-15.xi.1975, ll.xi.1976, 10.xi.1978, lO.x.1979 (Becker, Gomes) (VB; BMNH;
NMNH;MNHU).

Timocratica albitogata sp. n.

(Figs 53, 144, 145, 167)

c? 19-25 mm, $ 25-28 mm. Frons white. Second segment of labial palpus dark grey above on basal half,
tinged with ochreous basally below, white internally; third segment white, becoming progressively black
from basal third to apex. Fore coxa and femur golden-ochreous below, fore tibia and tarsus dark greyish-
fuscous below; distal half of tibia, and tarsus white above; mid and hind tarsus golden-ochreous. Fore wing
with costa gently, evenly rounded; apex nearly rounded, somewhat angled; termen and tornus obliquely
rounded ; veins free; white below. Hind wing and abdomen white.

GENITALIA <$ (Figs 144, 145). Uncus with lateral margins nearly parallel, apex strongly concave. Apex of
gnathos narrow. Digitate processes of juxta very long, lateral margins nearly parallel, distal half with long
setae above. Valva long, margins evenly rounded, nearly parallel, narrowing progressively from apical third
to apex. Aedeagus nearly straight, vesica with strong bent cornutus and several smaller spines.

GENITALIA $ (Fig. 167). Margin of ostium bursae nearly straight. Antrum very wide posteriorly, narrowing
progressively towards ductus bursae, with few longitudinal wrinkles. Ductus bursae somewhat constricted
near antrum, widening progressively towards corpus bursae. Corpus bursae pear-shaped, walls wrinkled
longitudinally as in ductus bursae. Signum an elongate elliptical plate without spines at centre.

REMARKS. T. albitogata is very close to palpalis, but usually larger. Like melanocosta, it can easily
be distinguished from palpalis by the white underside of the fore wings and the white dorsal side
of the fore tarsi. It differs from melanocosta by its larger size, white fore wing costa and ochreous
hind tarsi.

DISTRIBUTION (Fig. 53). South-east Brazil. This species appears to be restricted to the Subtropi-
cal Region of South America. The localities where the type-series was collected are in Subtropical
Moist Forest, Subtropical Lower Montane Moist Forest and Subtropical Lower Montane Wet
Forest. It can be expected to occur further south, in the Warm Temperate Moist Forest of Rio
Grande do Sul and possibly in Uruguay and Argentina.

MATERIAL EXAMINED

21 <J, 4 9 (3 (J, 2 $ genitalia preparations).

Holotype & Brazil: Parana, Curitiba, 920 m, 10.ii.1975 (Becker) (MN).

Paratypes. Brazil: 17 <J, 4 $, Parana, Curitiba, 920 m, xii.1974-ii.1975 (Becker) (VB; BMNH; NMNH;
NM; MNHU; ZSBS); 2 & Mato Grosso, Rio Brilhante, 25.U971 (Becker) (VB); 1 & Rio de Janeiro, [?
Itatiaia] (Zifcfln)(NMNH).

NEOTROPICAL GENUS TIMOCRATICA 265

Timocratica species 6
(Figs 53, 165)

9 23-27 mm. Frons white, edged with fuscous. Second segment of labial palpus ochreous below, basal half
dark fuscous above, distal half white above and internally; third segment white, fuscous towards apex.
Antenna fuscous, except scape and base of flagellum white. Fore femur above, distal half of tibia above and
fore tarsus dark fuscous ; fore coxa below, mid and hind tarsi golden-yellow. Fore wing with costa gently
arched, apex angled, termen and tornus rounded; veins free; underside golden-yellow, fuscous along apex
and termen. Hind wing tinged with golden-yellow along costa and termen. Abdomen white.

H species
20 A species 7

® me/anocosta

A isarga
30 •a/bitogafa

• melanostrii

24°C

Fig. 53 Geographical and ecological distribution of the albella-group of Timocratica.

GENITALIA 9 (Fig. 165). Margin of ostium bursae slightly concave. Antrum long, posterior third funnel-
shaped, anterior two-thirds strongly wrinkled longitudinally, bent at connection with ductus seminalis.
Ductus bursae broadening progressively towards corpus bursae. Corpus bursae oblong, walls slightly
wrinkled longitudinally as in ductus bursae. Signum an elongate plate without spines along middle.

REMARKS. This species is very similar to argonais but can easily be distinguished by the hind
wings which are tinged with yellow above along the costa and termen, and by the narrow antrum
of the female genitalia.

Although the ecology of the Bolivian specimen differs from that of specimens from Brazil, their
genitalia are similar; because of the poor condition of the Bolivian specimen they cannot be
separated on superficial characters.

DISTRIBUTION (Fig. 53). Bolivia, Brazil. Both Brazilian localities are in Tropical Moist Forest;
the Bolivian locality is in a transitional zone between Subtropical Moist Forest and Tropical
Premontane Dry Forest.

MATERIAL EXAMINED

3 $ (2 genitalia preparations).

Bolivia: 1 $, Santa Cruz, Santa Cruz de la Sierra (Steinbach) (BMNH). Brazil: 1 9, Amazonas, Benjamin
Constant, xi.l942(P0W)(NMNH); 1 9, Para, Belem (Moss) (BMNH).

Timocratica species 7
(Figs 53, 166)

9 22 mm. Frons white, edged with light fuscous. Second segment of labial palpus white, basal half ochreous
below and dark fuscous externally above; third segment dark fuscous, few white scales basally. Antenna
white, slightly fuscous towards apex. Fore coxa below, fore and mid femur above, basal half of fore tibia

266 V. O. BECKER

above, mid and hind tarsi golden-yellow; distal half of fore tibia above and fore tarsus dark fuscous. Fore
wing with costa gently arched, apex right-angled, termen and tornus rounded; golden-ochreous below
above cell, dark fuscous along apex and termen. Hind wing upperside white, underside golden-yellow along
costa. Abdomen white.

GENITALIA $ (Fig. 166). Margin of ostium bursae slightly concave. Antrum long, posterior third nearly
cylindrical, anterior two-thirds curved, strongly wrinkled longitudinally. Ductus bursae nearly cylindrical.
Corpus bursae oblong, walls plain and smooth. Signum an elongate plate without spines along middle.

REMARKS. This species is very similar externally to species 6 and to argonais; however, it can
easily be distinguished from the former by the white upperside of the hind wings, and from the
latter by the narrower antrum and undivided signum. It almost certainly represents a distinct
species, but as no males are known it has not been named here.

DISTRIBUTION (Fig. 53). Colombia. The locality where the only specimen was collected is presum-
ably in Tropical Premontane Wet Forest.

MATERIAL EXAMINED
Colombia: 1 9, Cundinamarca, Medina (Fassl) (BMNH).

Timocratica melanostriga sp. n.

(Figs 53, 68, 174)

$ 21 mm. Frons white. Second segment of labial palpus golden-ochreous except distal half above and
around distal articulation ; third segment dark grey with a few white scales basally. Antenna dark fuscous,
scape and base of flagellum white. Fore coxa, femur and tibia golden-ochreous, distal third of tibia except
articulations dark greyish fuscous; tarsus dark greyish-fuscous, ochreous on articulations; distal half of mid
femur ochreous below, tinged with grey externally at middle ; mid tibia fuscous with ochreous scales above,
tarsus fuscous with ochreous scales; third tarsus ochreous below with fuscous scales. Thorax and abdomen
white with a dark grey mediodorsal line. Fore wing with costa gently arched, apex rounded, termen and
tornus obliquely rounded; veins free; basal half of veins R3 — Af3, I A + 2 A, fold and middle of cell dashed
with dark greyish-fuscous; underside white, same veins marked with fuscous. Hind wing with veins and
dorsal half of cell fuscous.

GENITALIA $ (Fig. 174). Margin of ostium bursae slightly expanded posteriorly, concave at middle. Antrum
slightly constricted after insertion of ductus seminalis, anterior two-thirds narrowing slightly towards ductus
bursae, walls scobinate and wrinkled longitudinally. Ductus bursae widening progressively towards corpus
bursae. Corpus bursae pear-shaped, walls as in ductus bursae, finely scobinate. Signum a small, irregular
plate.

REMARKS. T. melanostriga is the only white Timocratica species with dark markings on the
wings. The female genitalia, mainly the shape of the ostium, put this species close to palpalis and
its allies; however, its correct position cannot be ascertained until males are known.

DISTRIBUTION (Fig. 53). Brazil. The only specimen was collected in the lowlands on the coast of
Santa Catarina, which is in Subtropical Moist Forest, transitional to Warm Temperate Moist
Forest.

MATERIAL EXAMINED

Holotype $, Brazil: Santa Catarina (Hoffmann) (NMNH).

Timocratica isarga (Meyrick)
(Figs 53, 172)

Stenoma isarga Meyrick, 1925 : 224. Holotype $, BOLIVIA (BMNH) [examined].

Timocratica isarga (Meyrick) Busck, 1935: 17 [catalogue]; Clarke, 1955: 388, pi. 194, figs 3-3c [adult,
genitalia].

$ 19 mm. Frons white. Basal half of second segment of labial palpus tinged with ochreous externally and
with dark fuscous above, white internally; third segment dark fuscous with white scales basally. Fore coxa
below and basal half of tibia above ochreous; fore femur above, distal half of tibia, and tarsus dark fuscous

NEOTROPICAL GENUS TIMOCRATICA 267

with few white scales on articulations ; articulations of mid femur, near tibia, tinged with golden-yellow. Fore
wing with basal third of costa arched, distal two-thirds nearly straight; apex rounded, slightly angled,
termen and tornus obliquely rounded; CuAt and CuA2 shortly stalked; white below. Hind wing and
abdomen white.

GENITALIA 9 (Fig. 172). Margin of ostium bursae evenly concave. Antrum widened posteriorly, abruptly
narrowed at middle, anterior half with margins nearly parallel, wrinkled longitudinally. Ductus bursae
broadening progressively towards corpus bursae. Corpus bursae pear-shaped; walls, as in ductus bursae,
plain. Signum an elongate plate without spines along middle.

REMARKS. T. isarga is very similar to palpalis, melanocosta and nivea, but is easily distinguished
from the first by the stalked CuAl and CuA2, the plain white underside of the fore wings and
white hind tarsi, and from the last two by the fuscous fore tarsi. The genitalia are very similar to
those of melanocosta from which, however, it differs by the white costa of the fore wings. T. nivea
has no yellow on the palpi and legs. Clarification of the relationship of isarga within the group
depends upon the discovery of males.

DISTRIBUTION (Fig. 53). Bolivia (eastern side of the Andes); in Warm Temperate Moist Forest.
According to Dr Holdridge (in litt.) this area is affected periodically by frost, resulting from cold
air masses which come from Antarctica, and is colder than would be suggested by the mean
annual temperature. According to the meteorological data it should be classified as Tropical
Premontane Dry Forest, transitional to Subtropical Moist Forest.

MATERIAL EXAMINED
Bolivia: holotype 9, Santa Cruz, Prov. del Sara, 450 m, xi (Steinbach) (BMNH).

Timocratica albella (Zeller)
(Figs 54, 75, 175)

Depressaria (Volucra) albella Zeller, 1839: 197. Holotype 9, SURINAM (BMNH) [examined].
Cryptolechia albella (Zeller) Zeller, 1854: 377 [redescription] ; Walker, 1864: 713 [catalogue].
Timocratica albella (Zeller) Busck, 1935: 16 [catalogue].

9 17 mm. Frons white. Second segment of labial palpus tinged with golden-yellow below, basal half tinged
with fuscous above, white internally; third segment white externally, dark fuscous internally. Fore tarsus
white above; fore coxa below, femur and basal half of tibia above golden-yellow, distal half of tibia, and
tarsus dark fuscous below; mid and hind legs white. Fore wing with costa gently arched, apex right-angled,
termen and tornus obliquely rounded; veins free; costa golden-yellow on underside. Hind wing white.
Abdomen with second to fourth tergites tinged golden-yellow.

GENITALIA 9 (Fig. 175). Ostium bursae wide, expanded posteriorly, concave at middle. Antrum very wide,
funnel-shaped, anterior half with a few wrinkles. Ductus bursae broadening progressively towards corpus
bursae, walls finely scobinate. Corpus bursae pear-shaped. Signum a long, transverse plate, concave across
middle.

REMARKS. T. albella is very similar externally to nivea and guarani, but is easily distinguished
from the former by the yellow tinge on its abdomen, fore legs, palpi and underside of the fore
wing costa; from guarani it can be separated by the white head. The female genitalia are distinct
from others in the group by the very broad antrum and the expansion of the margin of the ostium
bursae. However, the relationship of this species within the group must remain unknown until
males are discovered.

Although albella is known only from the holotype there are at least 50 references to it in the
Brazilian literature; these are based on misidentifications, probably through following Bondar
(19 13) and Lima (1928; 1936; 1945), and most apply to palpalis.

DISTRIBUTION (Fig. 54). Surinam; probably from Paramaribo which is in Tropical Moist Forest.

MATERIAL EXAMINED
Surinam : holotype 9, no further data (BMNH).

268 V. O. BECKER

Timocratica guarani sp. n.

(Figs 54, 76, 142, 143)

cJ 12-14 mm. Frons white, edged with fuscous and few ochreous scales. Basal two-thirds of second segment
of labial palpus ochreous externally, dark grey above, distal third and internally white; third segment dark
fuscous with white scales basally. Fore coxa, femur and tibia ochreous below ; distal half of tibia, and tarsus
white above, dark fuscous below; distal articulations of mid and hind femora tinged with golden-ochreous.
Fore wing with costa gently arched, apex somewhat angled, termen and tornus obliquely rounded; veins
free, CuA± and CuA2 very close basally in some specimens; underside golden-ochreous above cell. Hind
wing white. Abdomen slightly tinged with cream-yellow dorsally.

GENITALIA J (Figs 142, 143). Uncus with lateral margins nearly parallel, apical third with long sparse setae
in some specimens, apex strongly concave. Apex of gnathos broadly rounded. Digitate processes of juxta
with basal half broad, distal half narrow, covered with long sparse setae. Basal half of valva with margins
nearly parallel, then narrowing progressively towards apex, sacculus slightly expanded. Aedeagus bent
basally, vesica with strong bent cornutus and many acute spines of different sizes opposite.

REMARKS. T. guarani is externally very similar to albella, but their relationship cannot be clar-
ified until males of the latter are discovered. It differs from other white species from southern
South America by the white fore tarsi and yellow-tinged abdomen, and from albella by having the
frons edged with fuscous.

DISTRIBUTION (Fig. 54). Northern Argentina and Paraguay. It appears that this is the only
species restricted to Warm Temperate Dry Forest. The only other species recorded from the same
region is palpalis (collected by Hay ward in Villa Ana) which, however, is not restricted to it.

MATERIAL EXAMINED

3 cJ (2 genitalia preparations).

Holotype <J, Argentina : Santa Fe, Villa Guillermina, 20. ii.1925 (Hayward) (BMNH).

Paratypes. Argentina: 1 <J, Santa Fe, Villa Ana, ii.1924 (Hayward) (BMNH). Paraguay: 1 cJ, Paraguay
Central, 1885 (Germain) (BMNH).

Timocratica philomela (Meyrick)
(Figs 54, 146, 147)

Stenoma philomela Meyrick, 1925: 224. Holotype <$, PERU (BMNH) [examined].

Timocratica philomela (Meyrick) Busck, 1935: 17 [catalogue]; Clarke, 1955: 391, pi. 195, figs 3-3b [adult,
genitalia].

<J 10 mm. Head white, edged with fuscous. Second segment of labial palpus ochreous below, basal two-
thirds dark fuscous above, distal third above and internally white; third segment white, fuscous towards
apex. Antenna fuscous, scape white. Fore coxa below, femur and basal half of tibia above, and tarsus dark
fuscous. Hind tarsus golden-ochreous. Fore wing with costa gently arched, apex, termen and tornus evenly
rounded; veins free; underside golden-yellow, slightly tinged with fuscous along apex. Hind wing tinged with
cream-yellow. Abdomen white.

GENITALIA <$ (Figs 146, 147). Uncus wide, basal half slightly constricted; apex strongly concave. Apex of
gnathos wide, triangular. Digitate processes of juxta long, broadened basally, narrowing progressively
towards apex, few setae at distal half dorsally. Valva slightly constricted basally, dorsal margin nearly
straight, ventral margin evenly rounded. Aedeagus bent ventrad, vesica with strong bent cornutus and many
spines opposite.

REMARKS. T. philomela is a small species similar to butyrota and parvileuca but is easily dis-
tinguished by the ochreous colour of the labial palpi.

DISTRIBUTION (Fig. 54). Peru (Amazonian side of the Andes), in Tropical Moist Forest.

MATERIAL EXAMINED
Peru: holotype cJ, Loreto, Yurimaguas, iii.1920 (Parish) (BMNH).

NEOTROPICAL GENUS TIMOCRATICA

269

Amvea
• afbefta
i0 ^ parvifusca

bufyrofa
A phi/ome/a
® parvileuca
® guarani

12°C

24°C

Fig. 54 Geographical and ecological distribution of the albella-group of Timocratica.

Timocratica parvileuca sp. n.

(Figs 54, 150, 151)

cJ 9-10 mm. Frons white, edged with fuscous. Second segment of labial palpus fuscous, white internally;
third segment white, few fuscous scales internally towards apex. Antenna cream-yellow, scape white. Thorax
cream-yellow. Fore coxa tinged with fuscous below; femur and tibia above, and tarsus dark fuscous; mid
and hind legs light golden-yellow. For wings sub-oval, costa gently arched, apex, termen and tornus evenly
rounded ; veins free; golden-yellow below. Hind wing and abdomen cream-yellow.

GENITALIA <$ (Figs 150, 151). Uncus narrow, basal half slightly broadened. Apex of gnathos long, narrow.
Digitate processes of juxta long, narrow, distal half with several setae dorsally. Valva with margins nearly
parallel or somewhat broadened at middle, dorsal margin nearly straight or somewhat bent basally, ventral
margin evenly rounded. Aedeagus strongly bent ventrad, vesica with strong short cornutus and many small
spines.

REMARKS. T. parvileuca is one of the smallest species in the genus; it is very similar to butyrota
and philomela, but is easily distinguished from the former by the numerous spines of the vesica,
and from the latter by the lack of ochreous coloration of the palpi. Although it is closest to
butyrota it seems to vary less in wing venation.

DISTRIBUTION (Fig. 54). Brazil (Plateau of Parana and Sao Paulo), in Subtropical Lower Mon-
tane Forest, a different Life Zone from that of its nearest related species, butyrota.

MATERIAL EXAMINED

4 $ (2 genitalia preparations).

Holotype <J, Brazil: Sao Paulo, Sao Paulo (Jones) (BMNH).

Paratypes. Brazil: 2 J, Sao Paulo, Sao Paulo (Jones) (BMNH); 1 & Parana, Castro (Jones) (BMNH).

Timocratica butyrota (Meyrick) comb. n.
(Figs 16-19, 54, 74, 148, 149, 178)

Stenoma butyrota Meyrick, 1929: 516; Busck, 1935: 35 [catalogue]; Clarke, 1955: 276, pi. 138, figs 1-lc

[adult, genitalia]. Holotype £, COLOMBIA (BMNH) [examined].
Stenoma syndicastis Meyrick, 1929: 516; Busck, 1935: 58 [catalogue]. Holotype <$, COLOMBIA (BMNH)

[examined]. Syn. n.
Timocratica syndicastis (Meyrick) Clarke, 1955: 392, pi. 196, figs 1-lb [adult, genitalia].

270

V. O. BECKER

<$ 9-13 mm, 9 14-17 mm. Frons white, edged with fuscous. Second segment of labial palpus dark fuscous,
white internally and at distal articulation; third segment white in males, dark fuscous in females. Antenna
fuscous, scape white. Fore coxa fuscous below, femora and basal half of tibia golden-ochreous above, distal
half of tibia, and tarsus dark fuscous ; mid leg golden-yellow above ; hind tarsus golden-yellow. Fore wing
sub-oval, costa rounded, apex, termen and tornus evenly rounded; veins free or/?4 and K5 , andCuAt and
CuA2, stalked or connate; underside golden-yellow, fuscous or white along apex and termen. Hind wing
tinged with cream-yellow. Second to sixth abdominal tergites cream-yellow.

GENITALIA $ (Figs 148, 149). Lateral margins of uncus nearly parallel or slightly convergent towards apex;
apex slightly concave to slightly convex. Apex of gnathos narrow. Digitate processes of juxta short, laterally
compressed, with short setae dorsally. Valva with margins nearly parallel, evenly rounded, or broadened at
middle; ampulla slightly pronounced. Aedeagus bent ventrad, vesica with strong, short cornutus.

GENITALIA 9 (Fig. 178). Margin of ostium bursae straight or slightly convex. Antrum wide, short, with few
wrinkles anteriorly. Ductus and corpus bursae not differentiated, forming a long, wide sac. Signum a long,
irregular plate.

REMARKS. T. butyrota is very similar to philomela and parvileuca, but is easily distinguished from
the former by the lack of ochreous coloration on the palpi, and from the latter by the absence of
spines in the vesica.

The venation of the fore wing is very variable in this species, as it is in nivea. Some specimens
have all the veins free, others have R4 and R5, and CuAl and CuA2, connate or stalked, and
some have R4 and R5 stalked very close to the apex (Figs 16-19). These two veins can be
completely fused, and this is the case in the holotype of syndicastis, which has only eleven veins in
the fore wing. Variation occurs even within the same locality, and in a large series from Turrialba,
Costa Rica, specimens with all the above combinations were found.

The absence of a differentiated ductus bursae in this species is unique within the genus.

DISTRIBUTION (Fig. 54). Costa Rica, Colombia and Peru. No climatic data could be obtained for
Gorgona I., the type-locality, to establish the Life Zone. However, as suggested by other localities
it is probably Tropical Moist Forest or Tropical Premontane Wet Forest.

MATERIAL EXAMINED

71 cJ, 10 9 (8 (J, 2 9 genitalia preparations).

Colombia: holotype <J of S. butyrota, Cauca, Gorgona I., 60 m ('200 ft'), 7.x. 1924 (Collenette) (BMNH);
holotype ^ of S. syndicastis, Cauca, Gorgona I., 60 m ('200ft'), 7.vii.l924 (Collenette) (BMNH). Costa Rica:
61 (J, 109, Cartago, Turrialba, 600 m, 6-15.vii, 10.xi.1971; lO.ii, 4-20.V.1972; 10-20.iv.1973 (Becker) (VB;
BMNH; LN; MNHU; NM; NMNH; ZS3S). Panama: 5 <J, Canal Zone, Barro Colorado I., 10.x-
12.xii.1934 (Bates) (NMNH). Peru: 1 & Puno, Carabaya, La Oroya, Rio Inambari, 1000m ('3100ft'),
iii.1905 (Ockenden) (BMNH).

Timocratica parvifusca sp. n.

(Figs 20, 54, 67, 152, 153)

<J 9 mm. Head fuscous. Second segment of labial palpus dark fuscous with dark grey scales externally,
whitish internally; third segment light fuscous, whitish basally. Antenna dark fuscous. Thorax dark fuscous.
Fore coxa and femora light fuscous, fore tibia and tarsus dark fuscous; mid tibia and tarsus ochreous; tarsi
tinged with fuscous distally. Fore wing nearly oval; apex, termen and tornus evenly rounded; veins /?4 and
R5, CM/I! and CuA2 stalked, Ml missing; dark fuscous, costa ochreous below. Abdomen fuscous above,
whitish below.

GENITALIA $ (Figs 152, 153). Uncus broad, lateral margins nearly parallel, slightly broadened at middle.
Apex of gnathos long, narrow, pointed. Digitate processes of juxta slightly broadened at middle, gently
curved dorsad. Aedeagus strongly curved ventrad, vesica with long curved cornutus.

REMARKS. T. parvifusca is similar externally to fraternella, but differs by its smaller size, stalked
R4 and R5 , and the lack of Ml and the oblique fasciae of the fore wings.

Despite its colour pattern, which does not agree with that of any species of the albella-group,
there is no doubt that it belongs here. The shape of the genitalia and wings, and the venation, as
well as its geographical and ecological distribution, indicate that it is very close to if not a melanic
form of butyrota.

NEOTROPICAL GENUS TIMOCRATICA 271

DISTRIBUTION (Fig. 54). Costa Rica, in Tropical Premontane Wet Forest.

MATERIAL EXAMINED
Holotype <J, Costa Rica : Cartago, Turrialba, 600 m, 5.vii.l971 (Becker) (BMNH).

Species transferred from Timocratica

The following species have been included in Timocratica but are here transferred provisionally to
Stenoma, although none of them seem to be congeneric with lit ura Zeller, the type-species, which
is represented only by the holotype $ in the BMNH. Since the characters of female Stenominae
are inadequate for generic divisions, the relationship of litura with other species in the subfamily
cannot be established. It is probably congeneric with griseana Fabricius, the type-species of
Antaeotricha Zeller. If this proves true, most of the species now in Antaeotricha should be referred
to Stenoma, and most of the species now in Stenoma should be transferred to other genera, most
of these still to be defined. This would involve more than 1000 new combinations. The decision to
synonymize Antaeotricha with Stenoma, and the erection of new genera to accommodate species
now in this genus, should be taken only after further research, and possibly not before the male of
litura is known.

Stenoma completella (Walker)

Cryptolechia completella Walker, 1864: 718. LECTOTYPE $, BRAZIL: Amazonas, Tefe ['Ega'] (Bates)

(BMNH), here designated [examined].
Timocratica completella (Walker) Busck, 1935: 16.
Stenoma completella (Walker) Duckworth, 1962: 113.

As the abdomen of the male syntype is lost, the female syntype is here selected as the lectotype;
the male is labelled paralectotype. This species is very similar externally to Antaeotricha rhipi-
daula (Meyrick).

The larvae of completella, unlike those of Timocratica, skeletonize leaves tied together with
silk; I have reared a specimen from a larva feeding on Brosimum costaricanum (Moraceae) in
Costa Rica.

Stenoma convexicostata (Zeller) comb. n.

Cryptolechia convexicostata Zeller, 1877: 272. Holotype $ [not $ as stated by Zeller], BRAZIL: Rio de
Janeiro, Nova Friburgo (MNHU) [examined].

Stenoma liniella Busck, 1910: 80. Holotype $, COSTA RICA: Sixaola River (Schaus) (NMNH) [not examined].
Syn. n.

Stenoma cantatrix Meyrick, 1925: 221. Holotype £, BOLIVIA: Santa Cruz, Prov. del Sara, 450 m, x (Stein-
bach) (BMNH) [examined]. Syn. n.

Timocratica cantatrix (Meyrick) Clarke, 1955: 387.

Timocratica liniella (Busck) Duckworth, 1962: 113.

The holotypes of convexicostata and cantatrix, and a paratype of liniella in the BMNH, have
been examined and there is little doubt that they represent the same species, which is distributed
from the gulf area of Mexico to south Brazil. The specimens examined show some variation in
colour, some being darker than others, and in the female genitalia. However, the male genitalia
are almost identical in specimens from different localities and of different colour-pattern.

Duckworth (1962: 113) pointed out that the genitalia of liniella and cantatrix are atypical for
Timocratica and that both might require a new genus.

Stenoma grandaeva (Zeller) comb. n.

Cryptolechia grandaeva Zeller, 1854: 381. Holotype $, BRAZIL [no further data] (MNHU) [examined].
Stenoma chrysogastra Meyrick, 1915: 476. Holotype cJ, FRENCH GUIANA: St Jean du Maroni, 1915 (Le

Moult) (BMNH) [examined]. Syn. n.
Timocratica grandaeva (Zeller) Busck, 1935: 16.

272 V. O. BECKER

Although the holotype of grandaeva is in quite poor condition, with the abdomen, part of the
antennae and most of the legs missing, there is no doubt that it is conspecific with the holotype of
chrysogastra.

Stenoma sexmaculata (Dognin) comb. n.

Cryptolechia sexmaculata Dognin, 1904: 133. Holotype $, ECUADOR: San Francisco, near Loja (NMNH)

[examined].
Timocratica sexmaculata (Dognin) Busck, 1935: 17.

Although this species has the ground colour of the fore wing white, the metallic blue-green dots,
as well as its genitalia, are atypical of Timocratica.

Stenoma staudingerana (Maassen) comb. n.

Tortrix staudingerana Maassen, 1890: 25, pi. 9, fig. 29. Holotype [^?], COLOMBIA: Villavicencio [not exam-
ined].

Stenoma contophora Meyrick, 1915: 472. Lectotype $, FRENCH GUIANA: Godebert, R. Maroni, 1915 (Le
Moult] (BMNH) [examined]. Syn. n.

Stenoma heterosema Meyrick, 1930: 244. Holotype £, BRAZIL: Para, Taperinha, 21-30. vi.1927 (Zerny) (NM)
[examined]. Syn. n.

Timocratica staudingerana (Maassen) Busck, 1935: 17.

According to Horn & Kahle (1935: 162, 272) the Stiibel collection, including the Maassen types,
was deposited in the MNHU. Dr H.-J. Hannemann was unable to find the holotype of staud-
ingerana there (pers. comm.), neither is it in the IP (Dr Gaedike, pers. comm.). However, from the
excellent illustration which accompanies the description, there is no doubt that staudingerana
represents the species described later by Meyrick as contophora and heterosema. It is widely
distributed in South America; in the BMNH there are specimens from Bolivia, Brazil, Peru,
French Guiana and Surinam.
The genitalia of staudingerana exclude this species from Timocratica.

Stenoma tristrigata (Zeller) comb. n.

Cryptolechia tristrigata Zeller, 1854: 382, pi. 3, fig. 21. Holotype 9, BRAZIL [no further data] (MNHU)

[examined].
Stenoma aphanodesma Meyrick, 1915: 478. Holotype <$, FRENCH GUIANA: Godebert, R. Maroni, 1915 (Le

Moult) (BMNH) [examined]. [Synonymi/ed by Busck, 1935: 17.]
ITimocratica tristrigata (Zeller) Meyrick, 1912: 706; Busck, 1935: 17; Clarke, 1955: 392, pi. 196, figs 2, 2a.

Although the holotypes are of different sex there is no doubt that they belong to the same species.
The male genitalia of tristrigata are similar to those of staudingerana, and exclude both species
from Timocratica.

Acknowledgements

This study was undertaken as part of a Ph.D degree project at the Department of Zoology and
Applied Entomology, Imperial College of Science and Technology, University of London. The
work was supported by Empresa Brasileira de Pesquisa Agropecuaria (EMBRAPA), and par-
tially financed by the Institute of International Education, HE Fellowship no. 15783257. The
success of the application to carry out this study should be credited to Dr D. Gifford, University
of Brasilia, Brazil, who gave the project special support and to whom I express my particular
gratitude.

I am most grateful to my supervisor Dr R. G. Davies, Imperial College, London, for guidance
during this study and for making available his computer programs. I am also grateful to the
Keeper of Entomology, BMNH, and many of his staff, particularly Dr K. Sattler, for facilities and
advice provided during three years' work in that department. Much help was also received from
Dr J. D. Bradley, Commonwealth Institute of Entomology, London.

NEOTROPICAL GENUS TIMOCRATICA 273

Gratitude is specially acknowledged to the following people, who provided assistance with
material and information: Dra M. Brandao Ferreira, Empresa de Pesquisa Agropecuaria de
Minas Gerais, Belo Horizonte; Dr W. Died, ZSBS; Dr W. D. Duckworth, NMNH; Mr R. Feige,
Caracas, Venezuela; Dr F. Fernandez, UCV; Dr J. Furlan Jr, Centro de Pesquisa Agropecuaria
do Tropico Umido, Belem; Dr R. Gaedike, IP; Mr A. L. de Lima Gomes, Centro de Pesquisa
Agropecuaria dos Cerrados, Planaltina, DF, Brazil; Dr H.-J. Hannemann, MNHU; Dr J. B.
Heppner, NMNH; Dr L. R. Holdridge, Tropical Science Center, San Jose, Costa Rica; Dr F.
Kasy, NM; Dr D. Link, University of Santa Maria, Rio Grande do Sul; Dr O. H. H. Mielke,
Federal University of Parana, Curitiba; Dr R.-U. Roessler, LN; and Dr S. Silveira Neto, ESALQ,
Piracicaba, Brazil.

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276

V. O. BECKER

Figs 55-60 Wings of Timocratica species. 55, T. agramma sp. n., holotype c£, Brazil. 56, T. longicilia sp. n.,
holotype $, Colombia. 57, T. pompeiana Meyrick,^, Peru. 58, T. monotonia (Strand), cJ, Venezuela. 59, T.
meridionalis sp. n., holotype <$, Brazil. 60, T. loxotoma (Busck), <$, Mexico.

NEOTROPICAL GENUS TIMOCRATICA

211

Figs 61-66 Wings of Timocratica species. 61, T.fraternella (Busck), <J, Costa Rica. 62, T. species 1, ?,
Costa Rica. 63, T. major (Busck), $, Brazil. 64, T. effluxa (Meyrick), holotype rf, Bolivia. 65, T. species 2, <J,
Peru. 66, T. leucocapna (Meyrick), holotype <^, Colombia.

278

V. O. BECKER

Figs 67-72 Wings of Timocratica species. 67, T. parvifusca sp. n., holotype (J, Costa Rica. 68, T. melano-
striga sp. n., holotype 9, Brazil. 69, T. grandis (Perty), $, French Guiana. 70, T. xanthotarsa sp. n.,
holotype c?, Panama. 71, T. anelaea(Meyrick),($, Brazil. 72, T. macroleuca (Meyrick), cJ, Bolivia.

NEOTROPICAL GENUS TIMOCRATICA

279

Figs 73-78 Wings of Timocratica species. 73, T. titanoleuca sp. n., holotype <$, Peru. 74, T. butyrota
(Meyrick), cJ, Costa Rica. 75, T. albella (Zeller), holotype $, Surinam. 76, T. guarani sp. n., holotype (J,
Argentina. 77, T. xanthosoma leucocephala subsp. n., holotype J, Panama. 78, T. venifurcata sp. n.,
holotype <$, Brazil.

280

V. O. BECKER

79

80

81

Figs 79-82 Genitalia of Timocratica £. 79, 80, T. major (Busck), Brazil. 8 1, 82, T. agramma sp. n., holotype,
Brazil.

NEOTROPICAL GENUS TIMOCRATICA

281

85

86

Figs 83-87 Genitalia of Timocratica <J. 83-85, T. longicilia sp. n. (83, 84) paratype, Colombia. (85) right
valva, paratype, Colombia. 86, 87, T. pompeiana Meyrick, Peru.

282

V. O. BECKER

88

89

90

Figs 88-91 Genitalia of Timocratica monotonia (Strand), J. 88, 89, Colombia. 90, 9 1 , Brazil.

NEOTROPICAL GENUS TIMOCRATICA

283

93

94

95

Figs 92-95 Genitalia of Timocratica J. 92, 93, T. monotonia (Strand), Venezuela. 94, 95, T. meridionalis
sp. n., paratype, Brazil.

284

V. O. BECKER

98

99

Figs 96-99 Genitalia of Timocratica £. 96, 97, T. loxotoma (Busck), Mexico. 98, 99, T '. fraternella (Busck),
Costa Rica.

NEOTROPICAL GENUS TIMOCRATICA

285

100

102

103

Figs 100-103 Genitalia of Timocratica rf. 100, 101, T. leucocapna (Meyrick), Costa Rica. 102, 103, T.
effluxa (Meyrick), holotype, Bolivia.

286

V. O. BECKER

104

105

106

107

Figs 104-107 Genitalia of Timocratica <J. 104, 105, T. grandis (Perty), Panama. 106, 107, T. xanthotarsa
sp. n., paratype, Panama.

NEOTROPICAL GENUS TIMOCRATICA

287

109

111

Figs 108-111 Genitalia of Timocratica <$. 108, 109, T. constrictivalva sp. n., holotype, Ecuador. 110, 111,
T. bicornuta sp. n., paratype, Brazil.

288

V. O. BECKER

113

Figs 112-115 Genitalia of Timocratica <$. 112, 113, T. subovalis (Meyrick), holotype, Brazil. 114, 115, T.
fuscipalpalis sp. n., holotype, Venezuela.

NEOTROPICAL GENUS TIMOCRATICA

289

120

Figs 116-121 Genitalia of Timocratica <J. 116, 117, T. amseli Duckworth (holotype of Timocratica albella
Amsel), Venezuela. 1 18, 1 19, T. xanthosoma leucocephala subsp. n., paratype, Panama. 120, 121, T. venifur-
cata sp. n., paratype, Brazil.

290

V. O. BECKER

123

124

125

Figs 122-125 Genitalia of Timocratica <J. 122, 123, T. anelaea (Meyrick), Brazil. 124, 125, T. titanoleuca
sp. n., paratype, Peru.

NEOTROPICAL GENUS TIMOCRATICA

291

127

Figs 126-129 Genitalia of Timocratica <$. 126, 127, T. leucorectis (Meyrick), Peru. 128, 129, T. spinignatha
sp. n., holotype, Peru.

292

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Figs 130-133 Genitalia of Timocratica £. 130, 131, T. macroleuca (Meyrick), Bolivia. 132, 133, T. argonais
(Meyrick), French Guiana.

NEOTROPICAL GENUS TIMOCRATICA

293

137

Figs 134-137 Genitalia of Timocratica $. 134, 135, T. maturescens (Meyrick), Colombia. 136, 137, T.
palpalis (Zeller), Brazil.

294

V. O. BECKER

Figs 138-141 Genitalia of Timocratica <J. 138, 139, T. melanocosta sp. n., paratype, Brazil. 140, 141, T.
nivea sp. n., paratype, Brazil.

NEOTROPICAL GENUS TIMOCRATICA

295

144

145

Figs 142-145 Genitalia of Timocratica <$. 142, 143, T. guarani sp. n., holotype, Argentina. 144, 145, T.
albitogata sp. n., paratype, Brazil.

296

V. O. BECKER

147

146

148

149

Figs 146-149 Genitalia of Timocratica <$. 146, 147, T. philomela (Meyrick), holotype, Peru. 148, 149, T.
butyrota (Meyrick), Costa Rica.

NEOTROPICAL GENUS TIMOCRATICA

297

153

Figs 150-153 Genitalia of Timocratica £. 150, 151, T. parvileuca sp. n., paratype, Brazil. 152, 153, T.
parvifusca sp. n., holotype, Costa Rica.

298

V. O. BECKER

/

r/ "^p

155

Figs 154, 155 Genitalia of Timocratica $. 154, T. mo/or (Busck), Brazil. 155, T. monotonia (Strand), Brazil.

NEOTROPICAL GENUS TIMOCRATICA

299

159

Figs 156-159 Genitalia of Timocratica $. 156, T. meridionalis sp. n., paratype, Brazil. 157, T. loxotoma
(Busck), Mexico. 158, T. species 1, Costa Rica. 159, T. leucocapna (Meyrick), Costa Rica.

300

V. O. BECKER

160

Figs 160-163 Genitalia of Timocratica $. 160, T. grandis (Perty), French Guiana. 161, T. bicornuta sp. n.
French Guiana. 162, T. species 3, Peru. 163, T. species 5, Peru.

NEOTROPICAL GENUS TIMOCRATICA

301

164

166

Figs 164-166 Genitalia of Timocratica 9. 164, T. leucorectis (Meyrick), Peru. 165, T. species 6, Brazil. 166,
T. species 7, Colombia.

302

V. O. BECKER

168

Figs 167-169 Genitalia of Timocratica $. 167, T. albitogata sp. n., paratype, Brazil. 168, T. palpalis
(Zeller), Brazil. 169, T. megaleuca (Meyrick), holotype, Colombia.

NEOTROPICAL GENUS TIMOCRATICA

303

172

Figs 170-173 Genitalia of Timocratica $. 170, T. argonais (Meyrick), holotype, Brazil. 171, T. nivea sp. n.
paratype, Brazil. 172, T. isarga (Meyrick), holotype, Bolivia. 173, T. melanocosta sp. n., paratype, Brazil.

304

V. O. BECKER

176

177

178

179

Figs 174-179 Genitalia of Timocratica ?. 174, T. melanostriga sp. n., holotype, Brazil. 175, T.
(Teller), holotype, Surinam. 176, T. xanthosoma xanthosoma (Dognin) (holotype of Stenoma sacra
Meyrick), French Guiana. 177, T. species 4, Brazil. 178, T. butyrota (Meyrick), Costa Rica. 179,
T. amseli Duckworth (paratype of Timocratica albella Amsel).

305

Index

Synonyms and unavailable names are in italics; principal references are in bold.

Agonoxena Meyrick 227

Agonoxenidae 227

agramma sp. n. 212, 228-230, 232, 233

Agylla Walker 213, 216

albella Amsel 21 1,244, 247

albella Zeller 211, 213, 225, 228, 230, 244, 253,

255, 267, 268

albitogata sp. n. 211, 230, 264
amseli Duckworth 211, 225, 229, 244, 245, 246
anelaea Meyrick 212, 225, 228, 229, 248
Antaeotricha Zeller 227, 271
aphanodesma Meyrick 272
Arctiidae 216
argonais Meyrick 212, 214, 230, 251, 252, 265,

266

argonias Clarke 212, 251
auxoleuca Meyrick 212, 253

bahiensis Perty 241
balteata Meyrick 228
bicornuta sp. n. 212, 229, 241, 242, 243
butyrota Meyrick 212, 214, 225, 230, 268, 269,
270

cantatrix Meyrick 271

chrysogastra Meyrick 271, 272

claudescens Meyrick 212, 234, 235, 236

completella Walker 271

constrictivalva sp. n. 212, 228, 229, 241, 242,

243, 250

contophora Meyrick 272
convexicostata Zeller 271
crambina Busck 227
crassa Meyrick 212, 234, 235
Cryptophasa Lewin 228

decora Zeller 227
dissimilis Kearfott 227

Echiomima Meyrick 228

effluxa Meyrick 213, 228, 229, 239, 240

Ethmia Hubner 227, 228

Ethmiidae 227, 228

eucephala Turner 228

Eudeleboea Blanchard 259

Falculina Zeller 218, 219

fraternella Busck 212, 222, 228-230, 236, 237,

238, 270
fuscipalpalis sp. n. 212, 225, 228, 229, 245, 246

Gelechioidea 228

grandaeva Zeller 271, 272

grandis Perty 212, 229, 240, 241, 255

griseana Fabricius 271

guarani sp. n. 212, 217, 228, 230, 267, 268

haywardi Busck 212, 253
heterosema Meyrick 272
hyalinopa Lower 228

Ichneumonidae 259

isarga Meyrick 212, 225, 228, 229, 253, 255, 264,

266, 267
isographa Meyrick 212, 213, 225, 234, 235

leucocapna Meyrick 212, 213, 216, 220, 221, 229,

238, 239, 240

leucocephala subsp. n. 212, 229, 247
leucorectis Meyrick 212, 213, 230, 248, 249, 250
liniella Busck 271
litura Zeller 271

longicilia sp. n. 212, 217, 228, 229, 232, 233, 235
loxotoma Busck 212, 222, 229, 230, 236, 237, 238
Loxotoma Zeller 217, 218-220, 226
Lychnocrates Meyrick 211, 213, 225

macroleuca Meyrick 212, 228, 230, 248, 249, 250
major Busck 212, 213, 216, 219, 222, 229, 230,

231, 233

maturescens Meyrick 212, 228, 230, 251, 252
megaleuca Meyrick 212, 228, 230, 251, 253, 255
melanocosta sp. n. 212, 214, 226, 227, 230, 261,

264, 267

melanostriga sp. n. 212, 228, 229, 240, 266
meridionalis sp. n. 212, 217, 222, 229, 230, 234,

235, 236
monotonia Strand 212, 216, 220-222, 227, 229,

233, 234, 235-237
mythica Meyrick 228

nivea sp. n. 212, 230, 262, 264, 267
Oecophoridae 213

palpalis Zeller 212, 214, 216, 220, 221, 225-227,
230, 251, 252, 253, 255, 258, 261, 264, 267, 268

parvifusca sp. n. 212, 216, 217, 228-231, 238,
240, 270

parvileuca sp. n. 212, 225, 228, 268, 269, 270

Perixestis Meyrick 228

Philomela Meyrick 212, 228, 230, 268, 269, 270

pompeiana Meyrick 212, 222, 228, 229, 233, 234,
235

rhipidaula Meyrick 271
Rupela Walker 216

sacra Meyrick 212, 247
schlaegeri Zeller 227
Schoenobiinae 216

306 V. O. BECKER

sexmaculata Dognin 272 titanoleuca sp. n. 212, 225, 228, 230, 248, 249,

spinignatha sp. n. 212, 228, 230, 250, 251 250

staudingerana Maassen 272 tristrigata Zeller 213, 272

Stenoma Zeller 227, 255, 271

Stenominae 217, 218, 226, 227, 228 venifurcata sp. n. 212, 225, 228, 229, 245, 246,

subovalis Meyrick 212, 225, 228, 229, 243 247

stomatocosma Meyrick 212, 244

syndicastis Meyrick 212, 225, 269, 270 xanthosoma Dognin 212, 225, 229, 245, 246, 247

xanthotarsa sp. n. 212, 217, 228, 241, 242, 243

Thioscelis Meyrick 226 Xyloryctinae 227, 228
Timocratica Meyrick 211, 213, 216-219, 225,

226-230, 271, 272 ybyrajuba Becker 227

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Afrotropical species of the myrmicine ant genera Cardiocondyla, Leptothorax,
Melissotarsus, Messor and Cataulacus (Formicidae).

By Barry Bolton

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Afrotropical species of the myrmicine ant
genera Cardiocondyla, Leptothorax,
Melissotarsus, Messor and Cataulacus
(Formicidae)

Barry Bolton

Entomology series

Vol 45 No 4 30 September 198^

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World List abbreviation : Bull. Br. Mus. not. Hist. (Ent.)

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ISSN 0524-6431 Entomology series

Vol 45 No 4 pp 307-370
British Museum (Natural History)
Cromwell Road
London SW7 5BD Issued 30 September 1982

Afrotropical species of the myrmicine ant genera
Cardiocondyla, Leptothorax, Melissotarsus, Mes;
and Cataulacus (Formicidae)

Barry Bolton

1 V* LIBRARY ^

Department of Entomology, British Museum (Natural History), Cromwell RoaM ^Londoa^>
SW7 5BD

Contents

Synopsis 307

Introduction 307

Measurements and indices 308

Abbreviations of museums . 309

Cardiocondyla Emery ........ 309

Synonymic list of Afrotropical Cardiocondyla species 311

Key to species (workers) .312

Leptothorax Mayr ......... 319

Synonymic list of Afrotropical Leptothorax species 323

Key to species (workers) 323

Melissotarsus Emery 333

Synonymic list of Afrotropical Melissotarsus species 335

Key to species (workers) 335

Messor Forel ........... . 338

Synonymic list of Afrotropical Messor species 342

Key to species (medium to large workers) 342

Cataulacus F. Smith 354

Key to species (workers) 354

Appendix 354

Acknowledgements 365

References 365

Index ... 369

Synopsis

The Afrotropical species of the myrmicine ant genera Cardiocondyla Emery, Leptothorax Mayr,
Melissotarsus Emery and Messor Forel are revised and keyed, and a revised key to Cataulacus F. Smith is
presented. At genus-level Loncyda Santschi, Dyclona Santschi and Prosopidris Wheeler are newly
synonymized with Cardiocondyla; Nesomyrmex Wheeler and Tetramyrma Forel with Leptothorax; and
Veromessor with Messor. The current synonymy of Aphaenogaster Mayr, a genus very close to Messor, is
listed with the inclusion of Brunella Forel as a new synonym. At species-level nine Cardiocondyla (four new),
1 1 Leptothorax (one new), three Melissotarsus and 12 Messor (one new) are recognised in the regional fauna.
New species-level synonymy includes 10 names in Cardiocondyla, three in Leptothorax, four in Melissotarsus
and 14 in Messor, most of the last being of former infraspecific names. Five former infraspecific names in
Messor are given new status here as valid species. In Cataulacus six new species are described and four
previously synonymized names are reinstated as valid species.

Introduction

This paper is presented as a further contribution towards a revision of the subfamily Myrmicinae
in the Afrotropical region which, for the purpose of this study, excludes the fauna of the Malagasy
region. Previously issued parts of this series include studies of the genera Epitritus Emery (Bolton,
1972), Cataulacus F. Smith (Bolton, 1974), Decamorium Forel, Rhoptromyrmex Mayr and
Triglyphothrix Forel (Bolton, 1976), Tetramorium Mayr (Bolton, 1980), Meranoplus F. Smith,

Bull. Br. Mus. nat. Hist. (Ent.) 45 (4): 307-370

Issued 30 September 1982

308 B. BOLTON

Dicroaspis Emery and Calyptomyrmex Emery (Bolton, 198 la), Ankylomyrma Bolton,
Atopomyrmex Andre, Baracidris Bolton, Cyphoidris Weber, Ocymyrmex Emery, Pristomyrmex
Mayr and Terataner Emery (Bolton, 19816).

With the inclusion of the four genera treated in this paper a total of 20 of the region's 43
presently recognized myrmicine genera have been revised in the present series. The Afrotropical
fauna of some myrmicine genera has been studied by Brown who, beside revising Rhoptromyrmex
(Brown, 1964), has also analysed the genera of the myrmicine tribe Dacetini and revised its main
genera on a world-wide basis. In the case of sub-Saharan Africa this included the genera
Serrastruma Brown (Brown, 1952), Smithistruma Brown (Brown, 1953) and Strwnigenys F. Smith
(Brown, 1954).

Prior to these studies very little synthesising work had been carried out on the Afrotropical
myrmicines, the only notable contributions being the series of papers produced by Arnold
between 1916 and 1926 on the fauna of South Africa, and a catalogue of species by Wheeler
(1922) who also included a key to world genera. This key is now very much out of date, is difficult
to use and cannot be trusted. Similarly Arnold's (1916) key to the South African myrmicine
genera has, through subsequent synonymies and descriptions of new genera, become unusable.
More recently Bolton (1973) presented a subfamilial and generic key for the Afrotropical region
but again detailed investigation of the individual genera mentioned above has already rendered
this partially obsolete. A key to the 19 myrmicine genera in which the antennal club is restricted
to two segments has been constructed by Bolton (19816) and a key to the remaining genera is
presently being built up.

The four genera newly revised in this paper, which are discussed in more detail under their
individual sections, constitute a relatively minor proportion of the regional fauna in terms of
number of species. Melissotarsus and Cardiocondyla are arbitrarily regarded as small genera, with
three and nine species respectively in the region, whilst Leptothorax with 1 1 and Messor with 12
species are of moderate size. Apart from Melissotarsus, which is restricted to sub-Saharan Africa
and Madagascar, most species of the other three genera are primarily distributed elsewhere, the
Afrotropical fauna merely representing the few species which have successfully invaded the region
from the north.

Measurements and indices

Total Length (TL). The total outstretched length of the individual, from mandibular apex to
gastral apex.

Head Length (HL). The length of the head proper, excluding the mandibles, measured in a
straight line from the anteriormost point of the median clypeal margin to the mid-point of the
occipital margin, in full-face view. (In species with strongly concave occipital margin the head
length is measured to the mid-point of a line connecting the posterolateral corners.)

Head Width (HW). The maximum width of the head in full-face view, measured behind the eyes.

HW x 100
Cephalic Index (CI). —

HL

Eye Length (EL). In Cataulacus; the maximum length of the eye in full-face view.

EL x 100

Ocular Index (OI). In Cataulacus;

HW

Scape Length (SL). The maximum straight-line length of the antennal scape excluding the basal
constriction or neck. (In Cataulacus the SL usually measured in profile view with the scape in its
scrobe, as it is usually in this position in mounted specimens.)

c _ SL x 100

Scape Index (SI).

Pronotal Width (PW). The maximum width of the pronotum in dorsal view.

AFROTROPICAL MYRMICINE ANT GENERA 309

Alitrunk Length (AL). The diagonal length of the alitrunk in profile from the point at which the
pronotum meets the cervical shield to the posterior base of the metapleural lobes or teeth. (In
Melissotarsus measured to posteroventral corner of alitrunk as metapleural lobes absent.)

Abbreviations of museums

AMNH, New York American Museum of Natural History, New York, U.S.A.

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

IE, Bologna Istituto di Entomologia del'Universita, Bologna, Italy.

MCSN, Genoa Museo Civico di Storia Naturale 'Giacomo Doria', Genoa, Italy.

MCZ, Cambridge Museum of Comparative Zoology, Cambridge, Massachusetts, U.S.A.

MHN, Geneva Museum d'Histoire Naturelle, Geneva, Switzerland.

MNHN, Paris Museum National d'Histoire Naturelle, Paris, France.

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

MR AC, Tervuren Musee Royal de 1'Afrique Centrale, Tervuren, Belgium.

NM, Basle Naturhistorisches Museum, Basle, Switzerland.

NM, Bulawayo National Museum, Bulawayo, Zimbabwe. (Hymenoptera from this museum are

now deposited in SAM, Cape Town.)

NM, Vienna Naturhistorisches Museum, Vienna, Austria.

SAM, Cape Town South African Museum, Cape Town, South Africa.

USNM, Washington United States National Museum, Washington, D.C., U.S.A.

ZM, Kiev Zoological Museum, Institute of Zoology, Academy of Sciences of Ukrainian

S.S.R., Kiev, U.S.S.R.

CARDIOCONDYLA Emery
(Figs 1-7)

Cardiocondyla Emery, 1869: 20. Type-species: Cardiocondyla elegans Emery, 1869: 21, by monotypy.
Emeryia Forel, 1890: ex. Type-species: Emeryia wroughtonii Forel, 1890: cxi, by monotypy. [Synonymy by

Forel, 1892: 313.]
Xenometra Emery, 1917:96. Type-species: Xenometra monilicornis Emery, 1917:96. ( = Cardiocondyla

emeryi Forel), by monotypy. [Synonymy by Urbani, 1973: 199.]
Loncyda Santschi, 1930: 70 [as subgenus of Cardiocondyla]. Type-species: Cardiocondyla (Loncyda)

monardi Santschi, 1930: 70, by monotypy. Syn. n.
Dyclona Santschi, 1930: 70 [as subgenus of Cardiocondyla]. Type-species: Monomorium cristatum Santschi,

1912: 163, by original designation. Syn. n.
Prosopidris Wheeler, 1935: 40 [as subgenus of Cardiocondyla]. Type-species: Cardiocondyla (Prosopidris)

sima Wheeler, 1935: 41, by original designation. Syn. n.
Prosopidris Wheeler; Reiskind, 1965: 80. [Raised to genus.]

DIAGNOSIS OF WORKER. Small to minute monomorphic myrmicine ants. Mandibles with 5 teeth which
decrease in size from apical to basal. Palp formula 5, 3 (16 species examined). Clypeus with flattened and
prominent projecting lateral portions which are fused to the raised projecting median portion to form a shelf
which projects forward over the mandibles (Fig. 2). Sometimes the lateral portions of the clypeus extend
further forward than the median so that the anterior margin of the projecting shelf is concave medially.
Median portion of clypeus posteriorly broadly inserted between small narrow frontal lobes. Frontal carinae
and antennal scrobes absent. Eyes present, generally large and conspicuous, situated in front of the
midlength of the sides. Antennae with 1 1-12 segments, usually with a distinct 3-segmented club but the first
club segment may be relatively small. Promesonotal dorsum flattened to evenly convex in profile, the dorsal
alitrunk without sutures but the metanotal groove commonly (but by no means universally) impressed.
Pronotal corners in dorsal view broadly rounded to bluntly angular and projecting. Propodeal spiracle
small, situated approximately at the midlength, often low down on the side but not shifted back towards the
margin of the declivity. Propodeum unarmed to strongly bispinose. Metapleural lobes low and rounded.
Petiole nodiform with a moderate to long, usually slender, anterior peduncle. Postpetiole dorsoventrally
flattened in profile, in dorsal view very broad, much broader than the petiole node. Sting large and strongly
developed, knife blade-like and broad in profile, without lamelliform appendages. Dorsal surfaces of body
usually hairless.

310

B. BOLTON

The genus Cardiocondyla contains about 40 species, mostly distributed in the Old World.
Discounting tramp species only two have been described from the New World (ectopia Snelling
and venmtula Wheeler) but it is quite possible that both represent introductions, although to the
present no conspecific forms have been found among Old World material of the genus.

Cardiocondyla contains several very successful tramp species which are easily and apparently
frequently spread by human commerce. Such tramps include the cosmopolitan emeryi,
tropicopolitan wroughtonii and the Pacific island-hopping nuda (Mayr), which sometimes reaches

Figs 1-7 Cardiocondyla workers. 1, profile of shuckardi. 2, head of shuckardi. 3-7, alitrunk and pedicel
segments of (3) monardi, (4) wroughtonii, (5) emeryi, (6) weserka, (7) neferka.

AFROTROPICAL MYRMICINE ANT GENERA 311

the Pacific coast of North America. The fauna of the Afrotropical region includes 9
Cardiocondyla species. Of these six are found only in this region, two are the common tramps
emeryi and wroughtonii, and one also occurs on Madagascar (shuckardi). One of the six endemic
species, zoserka, described from a series of females, is suspected of being the first inquiline to be
found in this genus.

The majority of species of the world are known only from workers; a few queens are known
and these are quite normal apart from having the wing venation much reduced. The peculiarity of
Cardiocondyla lies in the males, which are known to be dimorphic in several species. Ordinary
alate males are known for a fair number of species but in some (emeryi, wroughtonii, elegans,
batesii Forel) dealate, highly ergatoid males are also produced; such peculiar males were
responsible for two of the generic names in the synonymy above, Emeryia and Xenometra. In a
further species, papuana (Reiskind), the only known male is an ergatoid. The problem is that the
extent of ergatoid male production among the species, and the reasons for the production of such
males, is unknown. It may well be that all species of Cardiocondyla are capable of developing
both normal and ergatoid males, given the right conditions, but it may be that some species only
have normal alate males, some only have ergatoid males, and some have both. It is certainly an
intriguing problem and deserves further investigation.

Recent studies of Cardiocondyla include the works of Wilson & Taylor (1967) on the Pacific
species, and of Bernard (1956) on the Palaearctic fauna; the species of sub-Saharan Africa have
not been dealt with previously.

To the present Cardiocondyla has occupied its own tribe, the Cardiocondylini, characterized
primarily by its prominent clypeus and broad postpetiole in the worker, and the reduced
venation in the female. Other features noted by Emery (1922a) and Wheeler (1922) have been
eroded away by subsequent discoveries of species not then known. Nevertheless, the tribal status
has remained as such since 1922 although Urbani (1977) has recently pointed out the similarity
between C. monardi and Leptothorax. He interpreted this as convergence but I consider that a
real relationship exists between Leptothorax and Cardiocondyla and that the latter belongs in
tribe Leptothoracini. Comparing the two genera there is broad agreement in head shape,
dentition, high palp formula, position of eyes, antennal segmentation, size and shape of frontal
lobes, broad insertion of the posterior clypeus between the frontal lobes, lack of scrobes and
frontal carinae, size and position of propodeal spiracle, and form of the metapleural lobes. The
presence of all these characters together in both genera argues strongly that they are genuinely
closely related and I propose the dissolution of Cardiocondylini and the incorporation of its sole
genus in the Leptothoracini. Within the tribe Cardiocondyla is still separated from Leptothorax
and its close relatives (as discussed under that genus) by the characters devised by Emery and
Wheeler, namely the specialized form of the anterior clypeus (although this is hinted at in some
Leptothorax), the characteristic form of the postpetiole and the reduced wing venation of the
females. A further character distinguishing the two is the specialized blade-like sting of
Cardiocondyla, not seen in Leptothorax.

Synonymic list of Afrotropical Cardiocondyla species

emeryi Forel

emeryi var. rasalamae Forel syn. n.

emeryi subsp. mahdii Karavaiev syn. n.

monilicornis Emery

nuda subsp. nereis Wheeler

mauritia Donisthorpe syn. n.
monardi Santschi
neferka sp. n.
nilotica Weber
sekhemka sp. n.
shuckardi Forel

globinodis Stitz syn. n.

badonei Arnold syn. n.

wassmanni [sic] Santschi syn. n.

312 B. BOLTON

wasmanni var. sculptior Santschi syn. n.

brevispinosa Weber syn. n.

fusca Weber syn. n.
weserka sp. n.
wroughtonii (Forel)

wroughtonii var. hawaiensis Forel

emeryi subsp. chlorotica Menozzi syn. n.
zoserka sp. n.

Key to species (workers)

Note. C. zoserka, described from suspected inquiline females, is omitted from the key.

1 With alitrunk in profile the dorsum without trace of a metanotal groove or impression (Fig. 3).

Propodeum unarmed. Postpetiole in dorsal view distinctly longer than broad. (Angola)

mortar di (p. 314)

With alitrunk in profile the dorsum with a distinct metanotal groove or impression (Figs 1, 4-7).
Propodeum sharply angulate to bispinose. Postpetiole in dorsal view as broad as to markedly
broader than long 2

2 With the head in full-face view the scapes, when laid back, distinctly exceeding the occipital

corners. (Sudan) nilotica (p. 315)

With the head in full-face view the scapes when laid back, either failing to reach or just reaching
the occipital corners, never exceeding them 3

3 Dorsal surfaces of head and alitrunk smooth and glossy, unsculptured everywhere except for

widely separated minute punctulae on the head. Head relatively broad and scapes short, CI 86,

SI 74. (Ghana) . sekhemka (p. 315)

Dorsal surfaces of head, alitrunk or both finely and densely sculptured, the sculpture usually
conspicuous. Scapes longer, SI > 80. Head with CI usually < 80, rarely otherwise ... 4

4 Propodeum in absolute profile bluntly angulate to bidenticulate (Fig. 1), never with a pair of

strong teeth or spines which are longer than their basal width in profile and which are as long
as half the distance separating their bases in dorsal view. Scapes relatively long, Si in range
93- 100. (Widespread in sub-Saharan Africa; Madagascar) .... shuckardt (p. 316)
Propodeum in absolute profile strongly bidentate to bispinose (Figs 4-7), the teeth or spines
longer than their basal width in profile and at least as long as half the distance separating their
bases in dorsal view. Scapes relatively short, SI in range 8 1-94 5

5 With alitrunk in profile the propodeal dorsum approximately flat behind the metanotal groove

and more or less level with the promesonotal dorsum, the propodeal dorsum not showing a

long gradual slope down to the spines (Fig. 6). (Cameroun) weserka (p. 317)

With alitrunk in profile the propodeal dorsum convex behind the metanotal groove and then
showing a long gradual slope down to the spines (Figs 4, 5, 7) 6

6 With alitrunk in profile the mesonotal dorsum abruptly changing slope posteriorly and

descending steeply to the metanotal groove (Fig. 4). Petiole node in dorsal view subglobular,
usually slightly broader than long. Head relatively broad, CI in range 79-86. (Pantropical

tramp species) wroughtonii (p. 317)

With alitrunk in profile the mesonotal dorsum curving evenly into the meianotal groove, without
an abrupt change of slope posteriorly (Figs 5, 7). Petiole node in dorsal view not subglobular,
usually quite distinctly longer than broad. CI in range 72-79 7

7 Pronotal corners bluntly but conspicuously angular in dorsal view. Propodeal spines relatively

long and slender (Fig. 7). (Ghana, Cameroun) neferka (p. 314)

Pronotal corners rounded in dorsal view. Propodeal spines relatively short and stout (Fig. 5).
(Cosmopolitan tramp species, very common) emeryi (p. 312)

Cardiocondyla emeryi Forel
(Fig. 5)

Cardiocondyla emeryi Forel, 1881: 5. Syntype workers, VIRGIN Is.: St Thomas I.. 1878 (MHN, Geneva)

[examined].
Cardiacondyla emeryi var. rasalamae Forel, 1891: 161. Syntype workers, MADAGASCAR: Imerina (P.

Camboue) (MHN, Geneva) [examined]. Syn. n.

AFROTROPICAL MYRMICINE ANT GENERA 313

Cardiocondyla emeryi subsp. mahdii Karavaiev, 1911:8. Syntype workers, SUDAN: Khartoum, Sirdargarten,

no. 1900 ( V. Karavaiev) (ZM, Kiev) [examined]. Syn. n.
Xenometra monilicornis Emery, 1917: 96. Holotype ergatoid male [not female], VIRGIN Is.: St Thomas I.

(MCSN, Genoa). [Synonymy by Urbani, 1973: 200.]
Cardiocondyla nuda subsp. nereis Wheeler, 1927: 140. Syntype workers, females, NORFOLK I.: 1915 (A. M.

Lea) (MCZ, Cambridge). [Synonymy by Wilson & Taylor, 1967: 53.]
Cardiocondyla mauritia Donisthorpe, 1946: 776. Holotype and paratype workers, MAURITIUS: 1941-45, no.

102 (R. Mamet) (BMNH) [examined]. Syn. n.

WORKER. TL 1.7-2.1, HL 0.45-0.52, HW 0.34-0.38, CI 72-78, SL 0.30-0.36, SI 86-94, PW 0.22-0.28, AL
0.48-0.58 (40 measured).

Antennal scapes of moderate length (SI, above), when laid back on the head usually failing to reach the
occipital corners but in a few samples just reaching them; never distinctly exceeding the occipital corners.
Maximum diameter of eye 0.10-0.12, about 0.28-0.32 x HW and with 8-10 ommatidia in the longest row.
Head always conspicuously longer than broad in full-face view, CI > 80 in all samples examined. With the
alitrunk in dorsal view the pronotal corners narrowly but evenly rounded, not produced into angular
shoulders. In profile the alitrunk with the promesonotal dorsum forming an even shallow convexity from
front to back, the slope of the dorsum not changing radically just in front of the metanotal groove.
Metanotal groove sharply and conspicuously impressed, the propodeal dorsum convex behind the groove,
then entering a long slope down to the propodeal spines. In profile the propodeal spines short and stoutly
constructed but longer than their basal width. In dorsal view each spine longer than half the distance
separating their bases. Petiole and postpetiole shaped as in Fig. 5, the petiole node showing some variation
in shape but in dorsal view always at least as long as broad and usually distinctly longer than broad.
Peduncle of petiole moderately long, the sternite of the postpetiole showing a blunt anteroventral
prominence or bulge. Postpetiole in dorsal view much broader than long, with a shallowly concave anterior
margin and evenly convex sides. Dorsal surfaces of head and alitrunk usually with scattered fine punctures,
the surface between them finely and densely shagreened or granular. In some specimens the punctures are
very small or widely scattered and inconspicuous, in which case the entire surface appears shagreened to
granular. Occasionally the granular ground-sculpture is reduced leaving the fine punctures on a more or less
smooth surface. Sculpture on the dorsal head is frequently stronger and better defined than on the dorsal
alitrunk. Hairs absent except on mouthparts and around gastral apex but a fine appressed pubescence is
present all over the body, being more conspicuous on the darkly coloured gaster than elsewhere. Head and
alitrunk yellow to light brown, sometimes orange-brown; gaster much darker, blackish brown to black and
contrasting strongly with the head and alitrunk.

A well known highly successful tramp-species, emeryi has been spread widely over the earth's
surface, mainly by human commerce. In the tropics and subtropics it survives outside, but in the
temperate zones it is more or less restricted to constantly heated buildings and greenhouses. The
presence of two very closely related species in West Africa, neferka and weserka, implies that the
Afrotropical region is most probably the place of origin of emeryi.

Like a few other species emeryi is known to have dimorphic males (see discussion, p. 311). The
species usually produces normal winged males but sometimes also develops highly ergatoid
males which may be found wandering alone, far from any nest.

MATERIAL EXAMINED

Afrotropical region. Ghana: Polcoase (W. Bellfield); Kibi (D. Leston). Nigeria: Gambari (B. Bolton); Bussa
(J. T. Medler). Cameroun: Nkoemvon (D. Jackson). Angola: Luanda (G. R. Gradwell & D. Snow). Sudan:
Khartoum (V. Karavaiev). Uganda: Ruwenzori, Semliki Forest (D. S. Fletcher). Kenya: Embu, Ishiara (V.
Mahnert & J.-L. Ferret). Tanzania: Lindi (D. V. Fitzgerald); Manyara Nat. Park (M. E. Irwin & E. S. Ross);
Zanzibar (L. F. Brown). Zimbabwe: Bembesi (G. Arnold). Botswana: Shorobe (A. Russell-Smith). South
Africa: Durban (C. B. Cooper); Nelspruit (M. Samways).

Other regions. Madagascar: Joffreville (J. M. Betsch); Imerina (P. Camboue). Seychelles: Little Sister I. (U.
M'iiller). Aldabra: South I. (B. Cogan & A. M. Hutson). Chagos Archipelago: Diego Garcia (A. M. Hutson).
Ascension I. (E. A. G. Duffey). Egypt: Gizeh (F. Morey); Siwa (J. Omer-Cooper); Zegawa (J. Omer-Cooper).
Madeira: Funchal (N. L. H. Krauss). Cape Verde Is.: Fogo (Lindberg); Fogo (Groh); S. Vincente (Lindberg);
S. Tiago (Lindberg); Nicolau (Lindberg); St Helena (Wollaston). Virgin Is.: St Vincent I. (H. H. Smith); St
Thomas I. West Indies: Anguilla (A. G. Parker). Puerto Rico: Mayaquez (M. R. Smith). Norfolk I. (A. M.
Lea). Mauritius (R. Mamet).

For Pacific distribution see Wilson & Taylor (1967); for Neotropical distribution see Kempf (1972).

314 B. BOLTON

Cardiocondyla monardi Santschi
(Fig. 3)

Cardiocondyla (Loncyda) monardi Santschi, 1930: 70, fig. 5. Syntype workers, ANGOLA: Rio Mbale,
ix.1928-i.1929 (A. Monard) (NM, Basle) [examined].

WORKER. TL 2.7, HL 0.58, HW 0.46, CI 79, SL 0.49, SI 107, PW 0.33, AL 0.68.

Antennal scapes relatively long, SI > 100; when laid back on the head exceeding the occipital corners.
Maximum diameter of eye 0.14, about 0.30 x HW and with approximately 14 ommatidia in the longest row.
Pronotal corners in dorsal view broadly and evenly rounded. Alitrunk in profile with the dorsum forming a
single uninterrupted surface, without trace of a metanotal groove or impression. Propodeum unarmed, the
dorsum rounding broadly, smoothly and evenly into the declivity. Petiole in profile with a very long anterior
peduncle and a long low feebly convex node. Petiole node in dorsal view subglobular, only very slightly
longer than broad. Postpetiole in dorsal view somewhat longer than broad, narrow (c. 0.13) at its junction
with the petiole, then rapidly broadening posteriorly to a maximum width of c. 0.26 at about its midlength,
and behind this narrowing again to a posteriormost width of c. 0.20. Dorsal length of postpetiole about 0.30,
of petiole peduncle plus node about 0.40. All dorsal surfaces of head, alitrunk, petiole, postpetiole and first
gastral tergite reticulate-punctate. Whole of body dorsally with glinting silvery pubescence which is mostly
set within the punctures. Colour yellow with glinting silvery highlights due to the pubescence.

This very distinctive species should not be confused with any other African form. It is quickly
separated from all its congeners in the Afrotropical region by its long scapes, lack of a metanotal
groove or impression, absolutely unarmed propodeum, elongate pedicel segments and glinting
silvery pubescence on a yellow background.

MATERIAL EXAMINED
Angola: Rio Mbale (A. Monard).

Cardiocondyla neferka sp. n.

(Fig. 7)

HOLOTYPE WORKER. TL 1.8, HL 0.48, HW 0.36, CI 75, SL 0.32, SI 89, PW 0.26, AL 0.49.

Antennal scapes of moderate length (SI 87-91 in type-series), when laid back on the head not reaching the
occipital corners in full-face view. Maximum diameter of eye 0.11, about 0.31 x HW and with 9-10
ommatidia in the longest row. Head conspicuously longer than broad, CI < 80. Pronotum in dorsal view
with narrowly rounded, somewhat prominent corners, giving the ant a conspicuously square-shouldered
appearance. With the alitrunk in profile the promesonotum forming an even shallow convexity from front to
back which grades into the metanotal groove without passing through an abrupt change of slope.
Metanotal groove shallowly impressed, the propodeal dorsum shallowly convex behind the groove, then
sloping downwards posteriorly towards the spines. Propodeal spines elongate and narrow, in profile much
longer than their basal width; in dorsal view the spines slightly incurved and each as long as the distance
separating their bases. Shape of pedicel segments as in Fig. 7. In dorsal view the petiole node longer than
broad, the postpetiole distinctly broader than long and broadest at its midlength. Dorsum of head
shagreened-granular, the sculpture very fine and dense, blanketing the surface. Promesonotal dorsum very
finely and densely superficially shagreened and mat, but the propodeal dorsum with only vestigial sculpture
and glossy, much less densely sculptured than the promesonotum. Dorsal surfaces of petiole, postpetiole and
gaster unsculptured except for a faint and patchy superficial patterning. Hairs absent except on mouthparts
but a fine appressed pubescence is present which is most apparent on the gaster. Colour uniform light
brownish yellow, the dorsum of the head slightly darker than the sides; sides of the first gastral tergite a rich
darker brown.

PARATYPE WORKERS. TL 1.80-1.81, HL 0.46-0.48, HW 0.35-0.37, CI 76-79, SL 0.32-0.33, SI 87-91, PW
0.24-0.27, AL 0.48-0.51 (3 measured).

Maximum diameter of eye 0.10-0.11, about 0.27-0.31 x HW and with 9-10 ommatidia in the longest
row. As holotype but in a couple the darker colour of the sides of the first gastral tergite extends onto the
dorsum.

Holotype worker, Ghana: Mampong, 10.ii.1970 (P. Room) (BMNH).

Paratypes. 3 workers with same data as holotype (BMNH; NM, Basle; MCZ, Cambridge).

Non-paratypic material examined. Cameroun: Nkoemvon (D. Jackson).

AFROTROPICAL MYRMICINE ANT GENERA 315

The Cameroun material differs from the type-series only in colour as here the dorsum of the head
is conspicuously much darker than the sides and the gaster is uniformly dark brown. This is
merely an intensification of the condition seen in the type-series and has no significance at
species-level.

C. neferka is closest related to emeryi but is quickly separable by its elongate narrow propodeal
spines and conspicuously square-shouldered appearance when the pronotum is seen in dorsal
view.

Cardiocondyla nilotica Weber

Cardiocondyla nilotica Weber, 1952: 8, fig. 13. Holotype worker, SUDAN: White Nile R., Ed Dueim, lat. 14U
00' N., 2.vii.l939, no. 1234 (N. A. Weber) (not in AMNH, New York ; presumed lost).

The only known representative of this species cannot be found in AMNH, New York and must
be presumed lost. However, Weber's original description contains enough information to give a
reasonable picture of this species, and it appears distinct from all other species of the Afrotropical
region. The following diagnostic characters are taken from Weber's description.

WORKER. TL 2.5. Antennal scapes when laid back distinctly exceeding the occipital corners. Metanotal
groove broad and rounded-concave. Propodeum armed with a pair of short triangular tubercles. Peduncle
of petiole slender. Petiole node in dorsal view broader than long, the postpetiole slightly broader than long
(taken from Weber's fig. 13, where the postpetiole appears subglobular in dorsal view). Densely and finely
punctate on head and alitrunk, gaster smooth and shining. Colour bright ferruginous, the head with a dark
area dorsally; appendages pale and gaster dark brown.

The overall picture which emerges is of a relatively large species closely related to shuckardi but
with decidedly longer scapes, narrower postpetiole and lighter colour, although a few pale
coloured individuals of shuckardi are known.

Cardiocondyla sekhemka sp. n.

HOLOTYPE WORKER. TL 1.8, HL 0.44, HW 0.38, CI 86, SL 0.28, SI 74, PW 0.12, AL 0.32.

Head relatively short and broad, scapes relatively short (CI and SI, above). When laid back on the head
the scapes failing to reach the occipital corners in full-face view. Projecting median portion of clypeus and
flattened prominent lateral parts of clypeus closely fused and forming a more or less evenly semicircular
projecting lobe which hides most of the mandibles in full-face view (only the two apicalmost teeth of the
right mandible can be seen in the holotype). Eyes relatively large, maximum diameter 0.12, about
0.32 x HW and with 10-11 ommatidia in the longest row. Shape of eye irregular in profile, narrowed and
drawn out anteroventrally, rounding the lower curve of the sides and onto the margins of the ventral surface
of the head. Pronotal corners rounded in dorsal view. With alitrunk in profile the promesonotum evenly
convex from front to back, sloping posteriorly to the feebly impressed metanotal groove. Propodeal dorsum
more shallowly convex than promesonotum and on a much lower level so that there is a distinct step-down
from the promesonotum to the propodeum. Posteriorly the propodeal dorsum sloping down to a pair of
broad blunt and very low tubercles which are much shorter than the metapleural lobes and which are
shorter than their basal widths. In dorsal view the tubercles distinctly shorter than half the distance
separating their bases. Petiole in profile with a short peduncle and rounded node. In dorsal view the petiole
node subglobular, slightly broader than long. Postpetiole in dorsal view much broader than long, with a
shallowly concave anterior margin and evenly convex sides. Dorsum of head sculptured with widely
scattered superficial minute punctulae, the surface between the punctulae smooth and shining. Remainder of
body unsculptured, smooth and shining. Hairs absent except on mouthparts and gastral apex. Colour
uniform glossy blackish brown, the legs and antennae lighter.

Holotype worker, Ghana: Tumu, 24.xii. 1969 (P. Room) (BMNH).

This small, virtually unsculptured darkly coloured species is easily recognised by its relatively
short scapes, broad head, characteristically shaped eyes, lack of developed propodeal spines and
feebly impressed metanotal groove followed by a depressed propodeum. In the Afrotropical
region only wroughtonii approaches the CI value of sekhemka, but in that species the propodeal
spines are long and strongly developed. Only shuckardi has the propodeal armament as feebly
developed as in sekhemka but here the head and body are usually strongly sculptured, the eye is
not drawn out anteroventrally, and the dimensions are very different.

316 B. BOLTON

Cardiocondyla shuckardi Forel
(Figs 1,2)

Cardiocondyla shuckardi Forel, 1891: 161. Syntype workers, MADAGASCAR: Imerina, Antananarivo
(Cambou'e) (MHN, Geneva) [examined].

Cardiocondyla globinodis Stitz, 1923: 154. Syntype workers, SOUTH WEST AFRICA: Omaruru, 22.vi.191 1 (W.
Michaelsen) (MNHU, Berlin) [examined]. Syn. n.

Cardiocondyla badonei Arnold, 1926: 225, fig. 64. Syntype workers, MOZAMBIQUE: Amatongas Forest,
ii.1917 (G. Arnold) (BMNH; MCZ, Cambridge) [examined]. Syn. n.

Cardiocondyla wassmanni [sic] Santschi, 1926: 241. Holotype worker, CAMEROUN: Gr. Batanga (R. P. E.
Wasmann) (NM, Basle) [examined]. Syn. n.

Cardiocondyla wasmanni var. sculptior Santschi, 1926: 241. Holotype worker, GABON: Samkita (F. Faure)
(NM, Basle) [missing from mount]. Syn. n.

Cardiocondyla brevispinosa Weber, 1952: 6. Holotype worker, ZAIRE: Beni, lat. 0° 24' N., long. 29° 24' E.,
24.ii.1948, no. 2116 (N. A. Weber) (not in AMNH, New York; presumed lost). [Junior secondary homo-
nym of Pheidole brevispinosa Donisthorpe 1947: 593 (= Cardiocondyla paradoxa Emery); synonymy by
M. R. Smith, 1955: 305.] Syn. n.

Cardiocondyla fusca Weber, 1952: 7. Holotype worker, UGANDA: Jinja, 15.viii.1939, no. 1495 (N. A. Weber)
(not in AMNH, New York ; presumed lost). Syn. n.

WORKER. TL 2.0-2.6, HL 0.50-0.60, HW 0.38-0.46, CI 75-79, SL 0.36-0.45, SI 93-100, PW 0.27-0.35, AL
0.54-0.69 (35 measured).

Antennal scapes when laid back on the head in full-face view either just failing to reach or just reaching
the occipital corners, never distinctly surpassing them; the scapes moderately long, SI > 90. Maximum
diameter of eye 0.1 1-0.14, about 0.26-0.30 x HW and with 9-12 ommatidia in the longest row. Head always
obviously longer than broad, CI < 80 in material examined. Pronotal corners in dorsal view broadly and
evenly rounded. With the alitrunk in profile the promesonotal dorsum forming an even shallow convexity
from front to back, sloping evenly into the metanotal groove. Metanotal groove impressed but the depth of
the impression varying between samples. To some extent the apparent variation in depth is caused by the
convexity of the propodeum behind the groove as in some cases it rises more steeply and is more convex
than in others. Propodeal dorsum behind the convex portion sloping downwards posteriorly to the junction
with the declivity. Propodeal armament very reduced, at best represented only by a pair of minute triangular
denticles which may be acute or blunted, or by a pair of tubercles, or merely bluntly angular; never with
developed teeth or spines (Fig. 1). In dorsal view the propodeal armament scarcely visible, the length of each
component constituting only a fraction of the distance separating their bases. Petiole node in dorsal view
subglobular, usually broader than long but in some only about as broad as long. Postpetiole distinctly
broader than long. In profile the petiole and postpetiole as in Fig. 1, the petiolar dorsum convex and
somewhat variable in length. Sculpture of dorsal head and alitrunk usually of fine, very dense blanketing
shagreening or granulation, but this may be reduced on the alitrunk or even on the head, though less
frequently on the latter than on the former. In extreme cases the dorsal alitrunk may be almost smooth.
Hairs absent except on mouthparts and gastral apex. Colour varying from medium brown to blackish
brown, sometimes black.

The commonest and most widespread endemic species in the Afrotropical region, shuckardi is
recognised by its dimensions and extremely reduced propodeal armament. Other species in the
region with reduced propodeal armament include monardi, sekhemka and nilotica. In the first of
these the metanotal groove is absent and the pedicel segments are very elongate (Figs 1, 3). C.
sekhemka is a much smaller species with shorter scapes and a broader head, and nilotica has
longer scapes than shuckardi and a narrower postpetiole.

MATERIAL EXAMINED

Ghana: Kibi (D. Lesion); Mampong (P. Room); Mole G. R. (J. C. Greig). Nigeria: Ibadan (K. Whitney};
Ibadan (B. Critchley). Cameroun: Nkoemvon (D. Jackson); Batanga (Wasmann). Zimbabwe: Umtali (G.
Arnold). Botswana: Shorobe (A. Russell-Smith). South West Africa: Okahanja (P. Hammond); Omaruru (W.
Michaelsen). South Africa: Transvaal, Plaston (M. Samways); Nelspruit (M. Samways); Natal, Ubombo (W.
L. & D. E. Brown); Illovo (P. Atkinson). Mozambique: Amatongas Forest (G. Arnold). Madagascar: Mont
d'Ambre (J. M. Betsch); Antananarivo (Camboue).

AFROTROPICAL MYRMICINE ANT GENERA 317

Cardiocondyla weserka sp. n.

(Fig. 6)

HOLOTYPE WORKER. TL 1.9, HL 0.46, HW 0.35, CI 76, SL 0.32, SI 91, PW 0.25, AL 0.48.

Antennal scapes moderately long but when laid back on the head failing to reach the occipital corners in
full-face view. Maximum diameter of eye 0.12, about 0.34 x HW and with 9-10 ommatidia in the longest
row. Pronotum in dorsal view with the corners narrowly rounded but not prominent. With the alitrunk in
profile the promesonotum with its dorsum almost flat, rounding broadly into its anterior declivity but
running into the metanotal groove almost in a straight line, with only the feeblest of curves. Metanotal
groove narrowly but quite distinctly impressed. Behind the metanotal groove the propodeal dorsum more
or less flat and on a slightly higher level than the posterior part of the promesonotum; the propodeal
convexity behind the metanotal groove followed by a long slope down to the spines, which is characteristic
of most species of the region, is absent here. Propodeal spines elongate and narrow, much longer than their
basal width in profile; in dorsal view the spines somewhat incurved, each spine easily as long as the distance
separating their bases. Shape of pedicel segments in profile as in Fig. 6. In dorsal view the petiole node
conspicuously longer than broad, its dorsal surface narrow. Postpetiole much broader than long, its anterior
face slightly concave, its sides convex. Dorsum of head blanketed by a fine dense granular sculpture or
shagreening. Dorsal promesonotum more lightly shagreened than head, the sculpture here being extremely
fine and very dense indeed. Propodeal dorsum with same sculpture as promesonotum but somewhat weaker
and appearing shiny in places. Petiole and postpetiole very finely and superficially shagreened. Hairs absent
except on mouthparts but a fine appressed pubescence is present, most easily visible on the first gastral
tergite. Alitrunk medium brown, the appendages slightly lighter. Head dorsally and gaster blackish brown
to black. Pedicel segments intermediate in shade between alitrunk and gaster.

Holotype worker, Cameroun: Nkoemvon, 1980, no. M35 (D. Jackson) (BMNH).

Among the species of the region in which the metanotal groove is impressed, weserka is
immediately distinguished by the shape of the propodeal dorsum. In general the propodeal
dorsum is convex behind the groove and then enters a long slope down to the tubercles, spines or
teeth (Figs 1, 4, 5, 7), but in weserka the dorsum is almost flat and does not conform to this usual
shape (Fig. 6).

Cardiocondyla wroughtonii (Forel)
(Fig. 4)

Emeryia wroughtonii Forel, 1890: cxi. Holotype male [ergatoid, not worker], INDIA: Poona (Wroughtori)

(MHN, Geneva) [examined].
Cardiocondyla wroughtonii (Forel) Forel, 1892: 313.
Cardiocondyla wroughtonii var. hawaiensis Forel, 1899: 119. Syntype workers, HAWAII: Molokai (MHN,

Geneva). [Synonymy by Wilson & Taylor, 1967: 56.]
Cardiocondyla emeryi subsp. chlorotica Menozzi, 1930: 84. Syntype workers, female, SOMALI REPUBLIC:

Duca Abruzzi, x.1926 (G. Paoli & A. Chiaromonte) (IE, Bologna) [examined]. Syn. n.

WORKER. TL 1.6-1.9, HL 0.42-0.50, HW 0.34-0.40, CI 79-86, SL 0.30-0.36, SI 81-89, PW 0.24-O.28, AL
0.46-0.55 (25 measured).

Small species with relatively broad head and short scapes, CI and SI above. When laid back on the head
the scapes failing to reach the occipital corners in full-face view. Maximum diameter of eye 0.09-0.11, about
0.26-0.30 x HW and with 9-11 ommatidia in the longest row. Pronotal corners rounded in dorsal view.
With the alitrunk in profile the promesonotum forming a shallow convexity from front to back but the slope
changing sharply posteriorly and becoming quite steep where it slopes down to the strongly impressed
metanotal groove; this change in slope very conspicuous in absolute profile. Propodeal dorsum behind the
metanotal groove convex in profile, then entering a long downward slope to the propodeal spines.
Propodeal spines enlongate and narrow in profile, longer than their basal width; in dorsal view each spine as
long as the distance separating their bases. Petiole node in dorsal view subglobular, as broad as or slightly
broader than long. Postpetiole distinctly broader than long. Dorsal surfaces of head and alitrunk blanketed
by fine shagreening or punctulate shagreening. Petiole and postpetiole finely superficially shagreened. Hairs
absent except on mouthparts and gastral apex but a sparse appressed pubescence is present, easiest seen on
the first gastral tergite. Head, alitrunk and appendages yellow to yellowish brown, colour of gaster variable.
Frequently the gaster is the same colour as the head and alitrunk but in some the sides of the tergite are

318 B. BOLTON

darker than the dorsum. In others the darker colour has also extended across the posterior portion of the
first tergite and in some the gaster is uniformly dark.

A tramp species probably originating in South East Asia, wroughtonii is now widespread in the
tropics and subtropics. Amongst the Afrotropical region species wroughtonii is recognizable by its
small size, relatively short scapes and broad head, subglobular petiole node in dorsal view, and
the characteristic shape of the promesonotum in profile. In terms of CI it is approached only by
sekhemka, but this species is uniformly dark in colour, has much shorter scapes (SI 74), and has a
differently shaped alitrunk.

MATERIAL EXAMINED

Afrotropical Region. Somali Republic: Duca Abruzzi (Paoli & Chiaromonte). Tanzania: Dar es Salaam (A.
J. Halstead); Zanzibar (M. J. Way).

Other regions. West Malaysia: Alor Star (G. H. Lowe); Gombak (B. Bolton). Australia: Qld, Mackay
(R. E. Turner). Japan: Chichi-jima, Ogasahara (M. Tanaka). Hawaii: Molokai (R. C. L. Perkins). Sri Lanka:
Peradeniya (A. Rutherford); Nawalapitiya. India: Poona (Wrought on); Pusa (S. D. Agarwala). Thailand.
U.S.A.: Fla, Dade Co., Tamiami Trail (W. F. Bur en).

Cardiocondyla zoserka sp. n.

HOLOTYPE FEMALE. TL 3.3, HL 0.68, HW 0.55, CI 81, SL 0.46, SI 84, PW 0.47, AL 1.04.

With the head in full-face view the outer margins of the mandibles conspicuously sinuate, passing through
a right-angle apically and forming a flat transverse anterior margin along to the apical tooth. Masticatory
margin of mandible with the usual five teeth but the apical tooth considerably enlarged, the three basalmost
teeth very small. Form of clypeus more Leptothorax-like than is usual in the genus, with a broadly and
evenly convex anterior lobe which projects over the base of the mandibles and with an impressed area
between the frontal lobes behind the posterior margin of the clypeus. Funicular segments of antennae with
bizarre modification and highly characteristic. In dorsal view funicular segment 1 slightly longer than broad,
2 slightly broader than long, but thereafter segments 3-10 short and very broad, becoming even broader
apically and with segments 8-10 extremely broad. The apical funicular segment swollen-conical in dorsal
view. In ventral view the funiculus even more bizarre. Segments 1-5 appearing the same as in dorsal view,
segments 6-7 flattened dorsoventrally, segment 8 slightly transversely concave, the very broad segment 9
strongly transversely concave and segment 10 so concave that the strongly arched ventral surface appears
almost to touch the dorsal at the point of maximum concavity. Apical segment invaginated and forming a
cup-shaped hollow which extends deep into the segment. Ocelli distinct. Maximum diameter of eye 0.24,
about 0.44 x HW. With alitrunk in dorsal view the mesoscutum slightly broader than long, the rounded
pronotal corners visible anteriorly. In profile the propodeal dorsum sloping down posteriorly to a pair of
small acute denticles. Petiole and postpetiole nodes both distinctly broader than long in dorsal view. Dorsal
surfaces of head, mesoscutum and scutellum granular to shagreened, with scattered punctures, the
mesoscutum also with very faint striate vestiges longitudinally. Dorsal propodeum with ground-sculpture
vestigial to absent, with a few feeble transverse rugulae. Petiole, postpetiole and gaster with scattered minute
punctulae dorsally. Hairs absent except on mouthparts but the body with a fairly dense and quite
conspicuous appressed pubescence which is most easily visible on the first gastral tergite. Colour dark
brown to blackish brown, the appendages lighter.

PARATYPE FEMALES. TL 2.9-3.3, HL 0.62-0.67, HW 0.51-0.55, CI 82-84, SL 0.42-0.46, SI 82-85, PW
0.42-0.46, AL 0.90-1.00 (4 measured).

As holotype but may be slightly lighter in colour. Sculpture reduced in some, the propodeal dorsum
almost smooth and the dorsal alitrunk less intensely sculptured. Maximum diameter of eye 0.21-0.24, about
0.41-0.44 x HW.

Holotype female, Nigeria: nr Abuja, Gurara Falls, 20.iii.1972 (E. Classey) (BMNH).
Paratypes. 4 females with same data as holotype (BMNH; NM, Basle; MCZ, Cambridge).

Although it is not usual practice to describe ant species from isolated females I make an exception
in this case for two reasons. Firstly, the modification of the mandibles, clypeal structure and
antennal funiculi lead me to suspect that this species is an inquiline. Secondly, the bizarre
modification of the funiculi renders the species immediately recognizable. To the best of my
knowledge no other ant has funiculi even remotely resembling this one, and certainly they cannot
be confused with any other member of Cardiocondyla. Assuming that I am correct in my

AFROTROPICAL MYRMICINE ANT GENERA 319

supposition that zoserka is an inquiline species (which makes it the first one known in the genus),
it is interesting to speculate what its host might be. Apart from the modifications of the head and
its appendages the overall appearance of zoserka is very like that of shuckardi females. The two
are definitely closely related and it may be that shuckardi represents the host of zoserka.

LEPTOTHORAX Mayr

(Figs 8-22)

Leptothorax Mayr, 1855: 431. Type-species: Formica acervorum F., 1793: 358, by subsequent designation of

Bingham, 1903:214.
Temnothorax Mayr, 1861: 68. Type-species: Myrmica (Leptothorax) recedens Nylander, 1856: 94, by

monotypy. [Synonymy by Forel, 1890<a: Ixxii.]
Dichothorax Emery, 18956: 323 [as subgenus of Leptothorax}. Type-species: Leptothorax (Dichothorax)

pergandei Emery, 18956: 323, by subsequent designation of Wheeler, 1911: 161. [Synonymy by Brown,

1973:180.]
Goniothorax Emery, 1896: 58 [as subgenus of Leptothorax]. Type-species: Leptothorax vicinus Mayr, 1887:

620, by subsequent designation of Wheeler, 1911: 164. [Junior homonym of Goniothorax Milne-Edwards,

1879: 103 (Crustacea).]
Mychothorax Ruzsky, 1904: 288 [as subgenus of Leptothorax]. Type-species: Formica acervorum F., 1793:

358, by original designation. [Synonymy by M. R. Smith, 1950: 29.]
Nesomyrmex Wheeler, 1910: 259. Type-species: Nesomyrmex clavipilis Wheeler, 1910: 259, by monotypy.

[As subgenus of Leptothorax and first available replacement name for Goniothorax Emery; M. R. Smith,

1950: 30.]Syn.n.
Tetramyrma Forel, 1912: 766 [as subgenus of Dilobocondyla Santschi]. Type-species: Dilobocondyla

(Tetramyrma) braunsi Forel, 1912: 767, by monotypy. [Raised to genus; Forel, 19136: 122. See also

Bolton, 1976:291.]Syn. n.
Caulomyrma Forel, 1914: 233 [as subgenus of Leptothorax]. Type-species: Leptothorax echinatinodis Forel,

1886a: xlviii, by original designation. [Synonymized with Nesomyrmex by Forel, 1915: 364.]
M yrmammophilus Menozzi, 1924: 29 [as subgenus of Leptothorax]. Type-species: Leptothorax

(M yrmammophilus) finzii Menozzi, 1924: 29, by monotypy. [Synonymy by Brown, 1973: 182.]
Limnomyrmex Arnold, 1948: 222. Type-species: Limnomyrmex stramineus Arnold, 1948: 223, by original

designation. [Synonymized with Nesomyrmex by Brown, 1971 : 4.]
Myrafant M. R. Smith, 1950: 29 [as subgenus of Leptothorax]. Type-species: Leptothorax curvispinosus

Mayr, 1866: 508, by original designation. [Synonymy by Brown, 1973: 182.]
Icothorax Hamann & Klemm, 1967: 415 [as subgenus of Leptothorax]. Type-species: Leptothorax

(Icothorax) megalops Hamann & Klemm, 1967: 417, by monotypy. [Synonymized with Myrafant by

Urbani, 1978: 556.]

DIAGNOSIS OF WORKER. Monomorphic myrmicine ants. Mandibles usually with five teeth (very rarely with 6)
which decrease in size from apex to base. Palp formula 5, 3 (60 species examined by dissection or in situ
count). Median portion of clypeus unmodified, broad and broadly inserted between the frontal lobes.
Anterior margin of median portion of clypeus evenly arched-convex to strongly lobate, the lobe often
prominent and concealing the basal border of the mandible or the basal tooth. Lateral portions of clypeus
unmodified, not forming a raised narrow ridge or shield-wall in front of the antennal insertions. Frontal
carinae usually absent but very rarely represented by a pair of faint narrow lines which run back from the
ends of the narrow frontal lobes. Antennal scrobes absent. Antennae with 11-12 segments, with a
conspicuous 3-segmented apical club. Eyes present, moderate to large in size and situated at or slightly in
front of the midlength of the sides. Propodeal spiracle circular and frequently very small, situated usually at
about the midlength of the segment and generally quite high up on the sides; never shifted back and down to
a position close to the bases of the propodeal spines. Pronotal corners dentate to evenly rounded.
Metapleural lobes rounded, usually small. Metanotal groove varying from absent to deeply impressed.
Propodeum commonly bidentate or bispinose, only very rarely unarmed. Petiole nodiform, variable in
shape, the anterior peduncle very variable in length and often with a denticulate process on each side
dorsally where peduncle meets node. Sting strong and acute, without apical or apicodorsal lamelliform
appendages, roughly cylindrical in section, not knife blade-like. Pilosity usually of short stout blunt hairs
but sometimes hairs absent and sometimes elongate.

Leptothorax is a large genus with a worldwide distribution although the majority of species are
Holarctic. Over 200 species have been described to date of which just 1 1 occur in sub-Saharan

320

B. BOLTON

Africa. The reason for this paucity of species in the Afrotropical region may well be the result of
direct competition from the extremely varied and enormously successful tetramoriine fauna of
the region (Bolton, 1976; 1980). In the past some members of Leptothorax and Tetramorium have
been confused because of an overall convergent similarity of appearance between a few members
of each genus. The following table will separate the workers of the two genera.

Figs 8-16 Leptothorax workers. 8, profile of angulatus. 9-13, heads of (9) angulatus, (10) braunsi, (11)
cenatus, (12) humerosus, (13) megalops. 14-16, alitrunk and pedicel segments of (14) megalops, (15) cenatus,
(16) humerosus. Pilosity omitted in 9-13.

AFROTROPICAL MYRMICINE ANT GENERA

321

Leptothorax
Sting simple, without an apical or apicodorsal

lamelliform appendage.
Maxillary palp with 5 segments.
Lateral portions of clypeus not raised into a

narrow ridge or shield-wall in front of the

antennal insertions.
Mandibles with 5 (rarely 6) teeth, decreasing in size

from apex to base.

Propodeal spiracle set high on side of segment and
about at its midlength; the spiracle usually in
the anterodorsal quadrant of the side of the
propodeum.

Tetramorium

Sting with an apical or apicodorsal lamelliform
appendage.

Maxillary palp with 4 (or rarely 3) segments.

Lateral portions of clypeus raised into a narrow
ridge or shield-wall in front of the antennal
insertions.

Mandibles usually with 7 teeth arranged as three
enlarged teeth followed by 4 denticles. [One or
two species with only 6 teeth but several with
> 7 by increase of the denticle series.]

Propodeal spiracle shifted back and down, set
behind the midlength; the spiracle usually in the
posteroventral quadrant of the side of the
propodeum.

Apart from the few African species revised below the taxonomy of most of the Old World
fauna of Leptothorax is in a poor condition. Only the faunas of North America (Creighton, 1950;
Brown, 1955) and of the Neotropical region (Kempf, 1959; Urbani, 1978) have been studied in
any detail. The west European fauna is mostly covered by Bernard (1968), Collingwood (1978;
1979) and Kutter (1977) but the remainder of the Old World remains unstudied by modern
methods.

Most of the generic synonymy noted above is straightforward and needs no further comment
here; a few, however, require further explanatory notes, as follows.

Temnothorax, synonymized long ago by Forel (1890a) on the grounds that it graded into
Leptothorax, has frequently been resurrected by European authors and treated either as a
subgenus of Leptothorax or even as a separate genus (most recently by Bernard, 1968). The
reason for this is not hard to find for among the west European species recedens, the type-species
of Temnothorax, stands out as an oddity as it does not belong to any of the usual west European
species-groups. However, when the extensive North African fauna is considered recedens is seen
as a fairly unexceptional Leptothorax species, and when the world fauna is taken into
consideration it seems decidedly mundane. The truth of the matter appears to be that recedens,
along with a few other species, really belongs to the North African fauna but has managed to
establish itself north of the Mediterranean. Urbani (1971) has discussed the validity of
Temnothorax and concluded that Forel's approach was the only logical one. I agree completely
and thus the original synonymy of Forel stands.

Tetramyrma, originally described as a subgenus of Dilobocondyla and later transferred into the
Tetramoriini, was recognized by Bolton (1976) to be only dubiously separable from Leptothorax.
On closer study it has not proved possible to find any genus-level characters to keep the name
separate. The type-species of Tetramyrma, braunsi, seems odd at first sight because of its domed
petiole and rounded, unarmed propodeum, but these developments are foreshadowed in maximus
Santschi and its allies. L. simoni, the only other species ever placed in Tetramyrma, provides a
good link back into the main mass of Leptothorax species, showing as it does a pair of propodeal
teeth whilst otherwise resembling braunsi very closely.

Nesomyrmex, with its own set of earlier synonyms (Caulomyrma, Goniothorax, Limnomyrmex),
is here formally synonymized with Leptothorax for the first time. Brown (1973) placed it as a
possible synonym in his world list of genera. Some members of this predominantly tropical group
appear very odd as a number of them have the petiole node denticulate, others have dentate
pronotal corners and many have very prominent clypeal lobes. However, there do not appear to
be any characters, either alone or in combination, which can serve to keep the former
Nesomyrmex species separate from the mass of Leptothorax. The largest representation of this
group occurs in South America and has been revised by Kempf (1959). His definition does not
separate Nesomyrmex from Leptothorax and one of his stated characters, the 5,3 palp formula,
seems universal in the genus. Species formerly placed in Nesomyrmex show considerable
variation in form and grade into more ordinary Leptothorax in all their specialized characters.

322 B. BOLTON

In my opinion all the earlier synonymy quoted above is valid and none of the included names
is deserving of further recognition as none of the characters invoked to separate them is
consistent or particularly functional. In fact, the similarities so enormously outweigh the
supposed differences, and the assumed diagnostic characters are so variable both within and
between the supposed subgenera, that the subgeneric system used in Leptothorax was at best
artificial, at worst misleading.

The only remaining subgeneric name in Leptothorax is Macromischa Roger ( = Antillaemyrmex
Mann, = Croesomyrmex Mann). Until recently this was treated as a good genus but Urbani
(1978), in his revision of the group, showed that the more exotic species (formerly in
Macromischa) graded into the more ordinary Leptothorax groups without it being possible to
draw any meaningful dividing line. However, instead of sinking Macromischa he chose to treat it
as a subgenus, though with considerable apprehension as some of the characters used are also
demonstrable, as Urbani says, elsewhere in Leptothorax, whilst others are not consistent through
Macromischa itself. The implication is that Macromischa is best regarded as a synonym of
Leptothorax.

The closest relatives of Leptothorax include many small inquiline or dulotic genera, all of
which are derived directly from Leptothorax. These genera are Chalepoxenus Menozzi,
Harpagoxenus Forel, Epimyrma Emery, Leonomyrma Arnoldi, Myrmoxenus Ruzsky,
Doronomyrmex Kutter, Formicoxenus Mayr, Myrmetaerus Soudek, and Symmyrmica Wheeler. Of
these Epimyrma is characterized by a reduced palp formula of 4,2 or 3,2 and usually a reduced
dentition; the genus may be valid. Harpagoxenus and Chalepoxenus both have strong frontal
carinae and short scrobes. The two are basically very similar and retain the standard
leptothoracine palp formula count of 5,3. The difference of antennae 11 -segmented versus
12-segmented which is used to separate them is not convincing as both antennomere counts
occur in Leptothorax (and several other myrmicine genera). The relationship of these two needs
further study for, although Chalepoxenus was revised quite recently (Kutter, 1973) its standing
with relation to Harpagoxenus was not discussed. The older separation based on mandibular
dentition, with Chalepoxenus having dentate and Harpagoxenus edentate mandibles works for
Europe, but the North American Harpagoxenus species have teeth.

Doronomyrmex, with its two parasitic species pads Kutter and pocahontas Buschinger, seems
indefensible as a genus. Its specialized features all result from inquiline syndrome characters
common to numerous parasitic but otherwise unrelated ants. The same appears to be true of
Myrmetaerus and Myrmoxenus, although further study of all these is needed. More information is
also required of Leonomyrma and Symmyrmica as both genera have short but fairly prominent
frontal carinae. The former also has the eyes shifted back on the head and the latter has 6-dentate
mandibles although this is not unknown in Leptothorax.

Finally Formicoxenus. Because of their very specialized inquiline lifeways in the nests of much
larger formicine ants Formicoxenus species have always presented a problem. Until recently the
genus only contained the two Palaearctic species nitidulus (Nylander) and orientalis Dlussky, and
was separated from Leptothorax by its possession of a strongly dentate subpostpetiolar process.
This postpetiolar development is a common feature in many unrelated inquilines from all parts of
the Myrmicinae and is a recognized character of the inquiline syndrome. It should not, by itself,
be regarded as being of generic significance. Dissection of nitidulus has, however, shown that the
mandibles are apparently consistently 6-dentate and the palp formula is reduced to 4,3. These
characters, coupled with the 11 -segmented antennae (again not a strong character when taken
alone) combine to form a reasonable case for maintaining Formicoxenus as a genus. An
observation in support of this comes from the decision of Buschinger (1979) to transfer the
American species hirticornis Emery and diversipilosus M. R. Smith from Leptothorax to
Formicoxenus on the grounds that their social organization is the same as in the European
nitidulus, and despite the fact that they lack a strong subpostpetiolar process. Dissection of
hirticornis has shown a 4,3 palp formula and 6-dentate mandibles as in nitidulus. I have not been
able to dissect any diversipilosus but a similar dentition and palp formula there would reinforce
the case for maintaining Formicoxenus as a genus separate from Leptothorax.

AFROTROPICAL MYRMICINE ANT GENERA 323

Synonymic list of Afrotropical Leptothorax species

angular us Mayr

angulatus st. ilgii Forel syn. n.

latinodis Mayr syn. n. (provisional)

angulatus var. concolor Santschi syn. n.
braunsi (Forel) comb. n.
cenatus sp. n.
denticulatus Mayr
evelynae Forel
grisoni Forel
humerosus Emery
innocens (Forel)
megalops Hamann & Klemm
simoni (Emery) comb. n.
stramineus (Arnold)

Key to species (workers)

1 With the alitrunk in absolute profile the dorsum forming a single uninterrupted surface which is

evenly flat or slightly convex, without trace of a metanotal impression and not having the

propodeum depressed (Fig. 8) 2

With the alitrunk in absolute profile the dorsum with the metanotal groove impressed even if
only feebly so, or the propodeum depressed below the level of the promesonotum, or both
(Figs 14-22) ... 3

2 Head and body uniform blackish brown to black. (Ghana, Zaire) .... grisoni (p. 329)
Head and body uniform yellow. (Extremely widespread) angulatus (p. 324)

3 First gastral tergite everywhere with blunt standing hairs 4

First gastral tergite either without standing hairs at all or at most with a single transverse row at

theapexofthesclerite 9

4 Petiole node narrow in profile, not denticulate (Figs 14-16). Antennal scapes longer, SI 85-1 10.

Eyes larger, maximum diameter 0.30-0.38 x HW 5

Petiole node broad in profile, denticulate (Figs 18-20). Antennal scapes shorter, SI 68-74. Eyes
smaller, maximum diameter 0.24-0.29 x HW 7

5 Anterior pronotal angles projecting as a pair of acute teeth in dorsal view; sides of pronotum

sharply marginate. Petiole node sharply triangular in profile (Fig. 16). Scapes relatively shorter

and head broader (Fig. 12), SI 85, CI 83. (' East Africa ') humerosus (p. 329)

Anterior pronotal angles evenly bluntly rounded in dorsal view; sides of pronotum not
marginate. Petiole node not sharply triangular in profile (Figs 14, 15). Scapes relatively longer
and head narrower (Figs 1 1, 13), SI 107-1 10, CI 70-78 .... 6

6 Eyes larger, maximum diameter 0.38 x HW. Petiole node in profile without a strongly

differentiated posterodorsal angle (Fig. 14). Mandibles almost smooth, with only vestiges of

sculpture. (Sudan) megalops (p. 331)

Eyes smaller, maximum diameter 0.30-0.31 x HW. Petiole node in profile with a strongly
differentiated posterodorsal angle (Fig. 15). Mandibles with strong but fine longitudinal
rugular sculpture. (Kenya) cenatus (p. 327)

7 Subpetiolar process a tooth anteriorly followed by a long cuticular flange which runs back to

the postpetiolar junction (Fig. 19). Eyes with 10-11 ommatidia in the longest row. Larger
species, HW 0.62-0.68, PW 0.46-0.52. (South Africa) . . . denticulatus (p. 328)

Subpetiolar process an anteriorly situated simple tooth or denticle (Figs 18, 20). Eyes with 7-8
ommatidia in the longest row. Smaller species, HW 0.49-0.53, PW 0.35-0.38 . 8

8 Propodeal spines short and broad, in profile about as long as their basal width, the declivity

between the spines and the metapleural lobes concave (Fig. 20). Dorsum of head densely and
sharply reticulate-punctate, with traces of fine rugulae. (Zaire) . . . innocens (p. 330)

Propodeal spines long and narrow, in profile distinctly longer than their basal width and
slightly downcurved, the declivity between the spines and the metapleural lobes straight
(Fig. 18). Dorsum of head weakly superficially reticulate-punctate, without trace of rugulae.
(South Africa) stramineus (p. 332)

9 Propodeum unarmed (Fig. 22). (South Africa) . . braunsi (p. 325)
- Propodeum armed with a pair of spines or teeth (Figs 17, 21) 10

324 B. BOLTON

10 Eye with only 7-8 ommatidia in the longest row. Alitrunk shaped as in Fig. 17. Small yellow

species with longer scapes, HW < 0.60, SI > 90. (Ghana, Zaire) .... evelynae (p. 328)
Eye with 15-16 ommatidia in the longest row. Alitrunk shaped as in Fig. 21. Large reddish
species with darker gaster and shorter scapes, HW > 0.85, SI < 85. (South Africa) . simoni (p. 331)

The few species constituting the Afrotropical fauna of Leptothorax apparently represent outliers
derived from a number of different species-groups of extralimital origin, one or two species from
each of which have managed to enter the region and to survive there. Because of the
unsatisfactory state of the taxonomy of Leptothorax the species-group limits have not been
worked out, but the 1 1 species occurring in sub-Saharan Africa aggregate as follows.

L. angulatus and grisoni. Metanotal groove absent. SI > 85. Eyes large, with 15 or more ommatidia in the
longest row. Pronotal corners acute. Petiole node large, with a short anterior peduncle; the node sculptured
but not denticulate. Frontal carinae absent. Median clypeal lobe more or less evenly convex.

L. denticulatus, innocens and stramineus. Metanotal groove present. SI < 75. Eyes relatively small, with
7-10 ommatidia in the longest row. Pronotal corners blunt. Petiole node large and denticulate, with a
moderately long anterior peduncle. Frontal carinae absent and the median clypeal lobe more or less evenly
convex.

L. braunsi and simoni. Metanotal groove present and the propodeum somewhat depressed below the level
of the promesonotum. SI in intermediate range, 78-83. Eyes large, with 16-18 ommatidia in the longest row.
Pronotal corners rounded. Petiole node massive and domed, not denticulate and with a moderately long
narrow peduncle. Clypeal lobe conspicuously produced; frontal carinae absent.

L. evelynae, cenatus and megalops. Metanotal groove present but shallow, sometimes very shallow.
SI > 90. Eyes relatively small to moderate, with 7-12 ommatidia in the longest row. Pronotal corners
bluntly angular to evenly rounded. Petiole node small, without denticles and with a moderately long
peduncle. Frontal carinae very feeble to absent and the median clypeal lobe more or less evenly rounded.

L. humerosus. Metanotal groove present. SI 85. Eyes large, with 14-15 ommatidia in the longest row.
Pronotal corners sharply dentate, the sides of the pronotum sharply marginate. Petiole node acutely
triangular, not denticulate, with a short peduncle. Feeble frontal carinae present and the median clypeal lobe
conspicuously produced.

Leptothorax angulatus Mayr
(Figs 8, 9)

Leptothorax angulatus Mayr, 1862: 739. LECTOTYPE worker, EGYPT: 'auf der sinaitischen Halbinsel' (R.

v. Frauenfeld) (NM, Vienna), here designated [examined].
Leptothorax angulatus st. ilgii Forel, 1894: 82. Holotype worker, ETHIOPIA: ' Sudabessinien ' (A. Ilg) (MHN,

Geneva) [examined]. Syn. n.
Leptothorax latinodis Mayr, 1895: 130. Holotype worker, MOZAMBIQUE: Delagoa Bay (H. Brauns) (not

found, presumed lost). Syn. n. (provisional).
Leptothorax angulatus var. concolor Santschi, 1914a: 107, fig. 15. Syntype workers, KENYA: M6mbasa, st.

no. 3, x.1911 (Alluaud & Jeannel) (NM, Basle) [examined]. Syn. n. [Data labels on syntypes read L.

(Goniothorax) angulatus var. concolor. ,]

WORKER. TL 3.1-3.8, HL 0.70-0.90, HW 0.56-0.74, CI 75-85, SL 0.50-0.66, SI 88-97, PW 0.40-0.56, AL
0.82- 1.08 (65 measured).

Mandibles delicately but densely longitudinally striate, the striation usually distinct but sometimes
superficial. Median clypeal lobe extensive, broad, covering the bases of the mandibles and having its anterior
margin conspicuously arched-convex. Median clypeal carina fine, not strongly developed but usually
discernible, only rarely the carina partially or wholly effaced. Antennal scrobes absent. Frontal carinae
absent but in some the frontal lobe followed on one or both sides by a weak rugular line which runs back on
the head. Maximum diameter of eyes 0.17-0.22, about 0.27-0.33 x HW and with 13 or more ommatidia in
the longest row. With the head in full-face view the sides narrower in front of the eyes than behind, slightly
convergent anteriorly. Sides of head behind eyes shallowly convex, slightly convergent posteriorly and
meeting the occipital margin in a blunted angle. Occipital margin transverse to very shallowly concave, with
a slightly projecting rim above the occipital foramen which is visible in full-face view. With the alitrunk in
profile the dorsum forming a single shallowly convex to almost flat surface, without trace of a metanotal
impression. Propodeum armed with a pair of triangular teeth or short broad spines of variable size. In
general the teeth are about as long as their basal width and slightly upcurved, but individuals with spines

AFROTROPICAL MYRMICINE ANT GENERA 325

longer than their basal width are fairly common. Specimens with the propodeal armament reduced to short
broad teeth, where they are shorter than the basal width, are less common. Metapleural lobes low and
rounded. In dorsal view the alitrunk with angulate to weakly dentate pronotal corners. Mesonotum
narrower than pronotum and the sides of the propodeum diverging to the level of the spiracle and then
converging to the bases of the propodeal teeth. Petiole in profile shaped as in Fig. 8, with a short anterior
peduncle which has a triangular dentiform anteroventral process. Dorsal surface of peduncle with a
denticulate process in front of the level of the spiracle on each side. Anterodorsal angle of node quite sharply
defined, the posterodorsal angle much broader and bluntly rounded. Petiole node in dorsal view variable in
shape and size. Usually the node about as broad as long, rarely slightly longer than broad but quite
commonly obviously broader than long, in some cases approaching the postpetiole in width. Dorsum of
head covered with a fine dense reticulate-punctulate ground-sculpture which in some samples is superficial
and granular in appearance. Superimposed on this are very fine irregular rugulae which frequently form a
narrow reticulum occipitally and sometimes also on the sides of the head. Dorsal surfaces of alitrunk, petiole
and postpetiole with fine granular or punctulate ground-sculpture and with disorganized fine rugulae. The
rugular sculpture is usually distinctive but in some individuals may be partially effaced. Base of first gastral
tergite generally with a superficial reticular pattern but sometimes almost completely smooth. All dorsal
surfaces of head and body with numerous short stout blunt hairs ; such hairs absent from the appendages.
Colour yellow, frequently with the antennal club darker.

L. angulatus is the most widely distributed and commonest species of this genus in sub-Saharan
Africa. It is easily identified by its yellow colour and lack of any trace of a metanotal groove or
impression. Only one other species in the region lacks a metanotal groove, grisoni, but in this
species the full adult colour is uniform blackish brown or black.

Arnold (1916: 259) noted that he only found angulatus on the trunks of trees but personal
observation has shown that it also occurs in leaf litter samples and log mould. However, the
species does seem to prefer to nest clear of the ground when possible, as colonies are often found
in West Africa in cocoa pods which are still attached to the tree, and the sample from Malawi
noted below was collected in Swartzia pods.

MATERIAL EXAMINED

Egypt: Sinai (Frauenfeld). Ghana: Legon (D. Lesion); Tafo (B. Bolton); Tafo (C. A. Collingwood); Adeiso
(P. Room); Adeiso (D. Lest on). Nigeria: Gambari (B. Taylor). Ethiopia: ' Sudabessinien ' (A. Ilg). Sudan:
Equatoria (N. A. Weber); Port Sudan (N. A. Weber); Nile above Khartoum (N. A. Weber). Kenya: Nairobe
(Patrizi); Mombasa (Alluaud & Jeannel); Tana Riv., Wema (V. Mahnert & J.-L. Ferret). Tanzania: Dar es
Salaam (N. L. H. Krauss). Malawi: nr Salima (B. J. S.). Zimbabwe: Victoria Falls (G. Arnold); Melsetter (G.
Arnold); Khami Riv. (G. Arnold). Botswana: Maxwee (A. Russell-Smith). South Africa: Natal, St Lucia (J. C.
Faure).

Leptothorax braunsi (Forel) comb. n.
(Figs 10, 22)

Dilobocondyla (Tetramyrma) braunsi Forel, 1912: 767. Holotype worker, SOUTH AFRICA: Cape Colony,

Willowmore (H. Brauns) (BMNH) [examined].
Tetramyrma braunsi (Forel) Forel, 19136: 122. [See also Bolton, 1976: 291.]

WORKER. TL 5.2-5.9, HL 1.20-1.36, HW 1.00-1.16, CI 83-86, SL 0.82-0.94, SI 79-83, PW 0.78-0.96, AL
1.44- 1.62 (9 measured).

Mandibles finely longitudinally striate, the spaces between striae finely punctulate or shagreened; the
striate sculpture sometimes inconspicuous. Median lobe of clypeus prominent (Fig. 10), its anterior margin
shallowly and evenly convex. Frontal carinae and antennal scrobes absent, the scapes of moderate length (SI
above). Maximum diameter of eye 0.28-0.31, about 0.26-0.29 x HW and with 16-18 ommatidia in the
longest row. In full-face view the head shaped as in Fig. 10. Alitrunk and pedicel segments in profile as in
Fig. 22, the promesonotum evenly convex, the metanotal groove not or only slightly impressed but the
propodeal dorsum distinctly depressed below the level of the promesonotum. Propodeum absolutely
unarmed, the dorsum rounding evenly into the declivity. In dorsal view the pronotal corners rounded, the
promesonotum narrowing posteriorly. Metapleural lobes rounded. Node of petiole in profile massive, with a
relatively narrow anterior peduncle which has a dentiform anteroventral process. In dorsal view the petiole
node subglobular, slightly broader than long; postpetiole broader than long and broader than the petiole.
Dorsum of head longitudinally rugulose with a few cross-meshes, occipitally a weak reticulum may be

326

B. BOLTON

17

19

22

Figs 17-22 Leptothorax workers. Alitrunk and pedicel segments of (17) evelynae, (18) stramineus, (19)
denticulatus, (20) innocens, (21) simoni, (22) braunsi.

AFROTROPICAL MYRMICINE ANT GENERA 327

formed. Sides of head above and behind eyes generally more obviously reticulate than the dorsum. Dorsal
alitrunk irregularly rugose, the sculpture quite strong, usually forming a reticulum on the propodeum and
anterior pronotum. Petiole and postpetiole irregularly reticulate-rugose. First gastral tergite densely
punctulate or shagreened, the sculpture generally strongest basally and usually traces of very fine
longitudinal costulae may be seen. A few short inconspicuous erect hairs present on dorsum of head but the
dorsal alitrunk, petiole and postpetiole hairless. First gastral tergite without standing hairs but with a short
fine appressed sparse pubescence. Appendages without standing hairs. Head and gaster dark brown tinged
with red to reddish black; alitrunk and pedicel segments red, the two colours strongly contrasting in fresh
specimens.

This large and conspicuous South African species is easily recognized by its large size, unarmed
propodeum, lack of hairs on alitrunk and first gastral tergite and depressed propodeal dorsum.
The closest related species in sub-Saharan Africa is simoni, but here the propodeum is distinctly
bidentate.

MATERIAL EXAMINED
South Africa: Cape Prov., Willowmore (//. Brauns).

Leptothorax cenatus sp. n.

(Figs 11, 15)

HOLOTYPE WORKER. TL 3.6, HL 0.78, HW 0.60, CI 77, SL 0.64, SI 107, PW 0.47, AL 0.98.

Mandibles finely but strongly longitudinally rugulose. Anterior clypeal margin convex and concealing the
basal tooth of the mandibles. Median clypeal carina feebly developed, weaker than the more laterally
situated clypeal carinae, which converge anteriorly. The anteriormost clypeal carina runs across the clypeus
in an unbroken transverse arc just behind the anterior margin, terminating at the antennal fossa on each
side. Frontal carinae represented by a pair of feeble meandering rugula-like ridges which run back from the
narrow frontal lobes to a point behind the level of the posterior margins of the eyes; these carinae scarcely
stronger than the rugular sculpture of the head and merging with that sculpture posteriorly. Antennal
scrobes absent, the scapes relatively long, SI > 100. Eyes quite large, maximum diameter 0.18, about
0.30 x HW and with 1 1 ommatidia in the longest row. With the head in full-face view the occipital margin
shallowly transversely convex, the occipital corners rounded. With alitrunk in dorsal view the pronotal
corners rounded. With alitrunk in profile the promesonotum shallowly evenly convex, the metanotal area
broadly but shallowly impressed and the propodeum armed with a pair of acute narrow spines. Metapleural
lobes very low, rounded. Petiole in profile with a moderately long anterior peduncle, the dorsal surface of
which is confluent with the anterior face of the node, the two not separated by an angle. Node with well
developed antero- and posterodorsal angles, the dorsum between them more or less flat. In dorsal view the
dorsum of the petiole node broader than long, and the petiole narrower than the postpetiole. Dorsum of
head with fine, widely spaced, irregular rugulae which are predominantly longitudinal. Occipitally the
rugulae are more sharply defined and have a few cross-meshes, although no reticulation is developed. Spaces
between rugulae smooth or at most with only vestiges of ground-sculpture. Sides of head above eyes
sculptured as dorsum but both in front of and behind the eyes the rugulae are more crowded and tend to
form a loose reticulum. Promesonotal dorsum finely and predominantly longitudinally rugulose, with very
sparse cross-meshes. The rugulae widely spaced and with a ground-sculpture of extremely fine superficial
punctulae between them, which in places is almost effaced. Rugulae present on propodeal dorsum but
weaker than on promesonotum. Petiole and postpetiole with fine superficial shagreening and a few weak
inconscpicuous rugulae. First gastral tergite unsculptured except for hair-pits. All dorsal surfaces of head
and body with numerous stout blunt mainly straight hairs which are shorter and more erect on the head
than on the first gastral tergite. Legs and scapes without such hairs. Dorsum of head brown, remainder of
body dull yellow with a brown tint, especially on the petiole and postpetiole which are somewhat darker
than the alitrunk and gaster but not as dark as the head.

PARATYPE WORKER. TL 3.2, HL 0.74, HW 0.58, CI 78, SL 0.62, SI 107, PW 0.42, AL 0.90.

As holotype but slightly 'smaller, maximum diameter of eye 0.18, about 0.31 x HW and with 10
ommatidia in the longest row.

Holotype worker, Kenya: Lake Nakuru, Nat. Park, 6.xi.l974, leaf litter (V. Mahnert) (MHN, Geneva).
Paratype. 1 worker, Kenya: Nakuru, Lake Elmenteita, 7.xi.l977, 1800 m (V. Mahnert & J.-L. Ferret)
(BMNH).

328 B. BOLTON

L. cenatus is closest related to megalops but differs in having smaller eyes and a differently shaped
petiole node (compare Figs 11,13 and 14, 15).

Leptothorax denticulatus Mayr
(Fig. 19)

Leptothorax denticulatus Mayr, 1901: 5. Syntype workers, female, SOUTH AFRICA: Cape Prov., Port
Elizabeth (H. Brauns) (NM, Vienna) [examined].

WORKER. TL 3.1-3.5, HL 0.74-0.84, HW 0.62-0.68, CI 81-85, SL 0.46-0.48, SI 71-74, PW 0.46-0.52, AL
0.82-0.94 (8 measured).

Mandibles finely shagreened to virtually smooth. Anterior margin of median lobe of clypeus evenly
arched-convex; median clypeal carina present and usually quite distinct. Frontal carinae absent; antennal
scrobes absent. Maximum diameter of eye 0.16-0.19, about 0.26-0.29 x HW and with 10-11 ommatidia in
the longest row; the eye in profile only very slightly longer than high. With the head in full-face view the
occipital margin straight to feebly convex, rounding evenly into the sides; the latter slightly narrower in
front of the eyes than behind and feebly convergent anteriorly. With the alitrunk in profile the
promesonotum shallowly convex dorsally, the metanotal groove impressed and the propodeal dorsum
convex. Propodeum armed with a pair of strong spines which are longer than their basal width. Metapleural
lobes rounded. In dorsal view the alitrunk with the pronotal corners bluntly angular to narrowly rounded.
Petiole node in profile large and blocky (Fig. 19), the upper sides and dorsum with numerous peaks or
denticles from which hairs arise. Peduncle of petiole short and broad, subtended by an extensive ventral
process which takes the form of a triangular denticle or tooth anteriorly, followed by a long cuticular ridge
which runs back to the junction with the postpetiole. In ventral view the subpetiolar ridge is seen to fork at
about its midlength, forming an inverted Y-shape. With the pedicel segments in dorsal view the denticles
conspicuous on the sides of both the petiole and postpetiole; both segments broader than long, the latter
somewhat broader than the former. Dorsum of head covered with a blanket of fine dense punctulate
ground-sculpture which is overlaid everywhere by dense and very fine rugular sculpture. On the dorsum the
rugulae are close and longitudinal but on the sides, above the eyes and occipitally there is a tendency for a
narrow reticulum to be formed. Dorsal alitrunk reticulate-punctate and with fine rugulae which are
predominantly longitudinal; on the promesonotum a reticulum may be formed anteriorly and in some the
rugulae are quite strongly developed. Sculpture of petiole and postpetiole dorsally predominantly
reticulate-punctate but a few fine rugulae may be present. Base of first gastral tergite superficially reticulate
to almost smooth. All dorsal surfaces of head and body densely and evenly clothed with short blunt hairs;
the appendages without such hairs. Colour uniform yellow, sometimes the posterior half of the gaster darker
than the anterior half.

Among the species in which the metanotal groove is impressed three, denticulatus, innocens, and
stramineus, have the petiole node bearing denticles from which hairs arise. Of the three
denticulatus is recognized by its strongly developed subpetiolar process, dense pilosity, larger eyes
and larger size.

MATERIAL EXAMINED
South Africa: Cape Prov., Barrydale (H. V. Daly); Port Elizabeth (H. Brauns).

Leptothorax evelynae Forel
(Fig. 17)

Leptothorax (Goniothorax) evelynae Forel, 1916: 423. Syntype workers, female, ZAIRE: St Gabriel (Kohl)
(MHN, Geneva) [examined].

WORKER. TL 2.5-2.9, HL 0.58-0.70, HW 0.47-0.54, CI 77-81, SL 0.44-0.52, SI 92-98, PW 0.35-0.44, AL
0.65-0.82 (7 measured).

Mandibles finely shagreened. Median clypeal lobe evenly arched-convex. Median clypeal canna present
but fine, incomplete in a few specimens. Frontal carinae and antennal scrobes absent. Antennal scapes
relatively long, SI > 90. Maximum diameter of eye 0.12-0.16, about 0.26-0.29 x HW and with 7-8
ommatidia in the longest row. With the alitrunk in profile the metanotal groove shallowly but
conspicuously impressed, the promesonotum evenly shallowly convex and the propodeal dorsum almost flat
to shallowly convex. Propodeal spines straight, distinctly longer than their basal width. Metapleural lobes

AFROTROPICAL MYRMICINE ANT GENERA 329

low and rounded. In dorsal view the alitrunk with the pronotal corners angulate and the sides ot the
promesonotum bluntly marginate. The dorsal surface gradually narrows from front to back but the sides of
the mesonotum are slightly convex and the sides of the propodeum diverge from the metanotal groove to
the level of the spiracle and then converge to the bases of the spines. Petiole in profile with a high narrow
node (Fig. 17) which is not equipped with denticles. Ventral process of peduncle a simple small tooth,
anteriorly situated. In dorsal view the petiole node broader than long. Dorsum of head with fine superficial
reticulate-punctate ground-sculpture which is overlaid by a very fine narrow reticulate-rugulose net
everywhere except in the area immediately behind the frontal lobes. Dorsal alitrunk with superficial
punctulate ground-sculpture overlaid by fine rugulae. In specimens from Zaire this rugular sculpture is faint
and weakly developed, forming a reticulum only on the anterior pronotum, but in material from Ghana the
rugulae are more strongly developed everywhere and reticular meshes are frequent. Petiole and postpetiole
finely punctulate or granular dorsally, sometimes with one or two fine rugulae. Base of first gastral tergite
very lightly shagreened or with a superficial reticular pattern. Dorsum of head with scattered short stout
blunt hairs. Similar hairs are present on the pronotum (several pairs), mesonotum (1-3 pairs), petiole and
postpetiole (1-2 pairs each) but are absent from the propodeum and absent from the first gastral tergite
except for a transverse row at the extreme apex. Gastral segments behind the first also with a transverse
apical row each. Colour yellow.

The characteristic distribution of the body hairs renders evelynae quickly recognizable among the
African Leptothorax species. The lack of hairs on the propodeum and first gastral tergite is
paralleled only in simoni and braunsi, but these are both much larger (HW > 0.85), darker
coloured species with much more massively developed petiole nodes and shorter antennal scapes
(SI < 85). The closest related species appears to be megalops, but here the eyes are larger and the
propodeum and first gastral tergite both have hairs present.

MATERIAL EXAMINED
Ghana: Tafo (B. Bolton). Zaire: St Gabriel (Kohl).

Leptothorax grisoni Forel

Leptothorax (Goniothorax) grisoni Forel, 1916: 425. Syntype workers, male, ZAIRE: St Gabriel (Kohl) (MHN,
Geneva; MCZ, Cambridge) [examined].

WORKER. TL 2.9-3.4, HL 0.76-0.86, HW 0.60-0.66, CI 77-81, SL 0.56-0.63, SI 94-98, PW 0.46-0.53, AL
0.85-1. 02 (9 measured).

Answering to the description of angulatus, differing only in colour and intensity of sculpture. In grisoni the
full adult colour is uniform blackish brown to black, as opposed to the uniform yellow found in angulatus.
The dorsum of the head and alitrunk in grisoni is blanketed by a dense reticulate-punctate ground-sculpture
which is overlaid by conspicuous fine rugulae which form a distinct reticulum on the alitrunk and on much
of the head. Basically this sculpture is the same as that seen in angulatus, but here it is more intensely and
sharply developed.

Although these seem relatively minor differences I am prepared to accept them as valid for the
present. The reason for this is that the two colour forms have ranges which only partially overlap.
The yellow angulatus is known from most of the continent from Ethiopia and Sudan to South
Africa and Botswana, but the dark grisoni has only been found in Ghana and Zaire. L. angulatus
also occurs in Ghana in the same areas where grisoni has been discovered, but even here the two
maintain their distinctive colours, no intermediates being known.

MATERIAL EXAMINED
Ghana: Tafo (B. Bolton); Kade (J. Majer); Asamankese (P. Room). Zaire: St Gabriel (Kohl).

Leptothorax humerosus Emery
(Figs 12, 16)

Leptothorax humerosus Emery, 1896: 62. Holotype worker, 'AFRICA ORIENTALS': no loc. (Staudinger &

Bang-Haas) (MCSN, Genoa) [examined].
WORKER. TL 3.7, HL 0.90, HW 0.75, CI 83, SL 0.64, SI 85, PW 0.52, AL 0.98.

330 B. BOLTON

Mandibles almost smooth, with only vestigial traces of superficial sculpture. Median lobe of clypeus
strongly produced, roughly rectangular, its anterior margin transverse and flat; the anterior margin meeting
the sides of the lobe in a distinct angle. Median clypeal carina absent. Narrow weakly differentiated frontal
carinae present which are scarcely stronger than the remaining cephalic sculpture but which reach back well
beyond the level of the posterior margins of the eyes. Antennal scrobes absent. Antennal scapes moderately
long, SI 85. Maximum diameter of eye 0.24, about 0.32 x HW and with 14-15 ommatidia in the longest row.
With the head in full-face view the sides narrower in front of the eyes than behind and somewhat convergent
anteriorly. Sides behind eyes rounding evenly into the occipital margin, the latter shallowly and evenly
transversely convex. Pronotum sharply marginate laterally, the anterior pronotal corners dentate. With the
alitrunk in profile the promesonotum convex, the metanotal groove deeply impressed. Propodeum broadly
and evenly convex in profile, sloping down posteriorly to the long spines; the latter blunt apically and with
their dorsal margins angled (Fig. 16). Metapleural lobes low and rounded. Petiole in profile with the node
triangular, rising to an acute peak above; anterior peduncle of petiole short. In dorsal view the sides of the
petiole roughly parallel. Postpetiole much broader than petiole. Basal face of first gastral tergite transverse
except for a median concavity where it articulates with the postpetiole. Dorsum of head everywhere
sculptured with fine longitudinal rugulae and with a fine granular to punctulate superficial
ground-sculpture. Dorsal alitrunk with ground-sculpture similar to head. Pronotum also with 7-8 broad,
coarse longitudinal rugae which are almost sulcate in appearance and are most strongly developed
anteriorly. These longitudinal rugae are continuous over the length of the promesonotum and also traverse
the base of the metanotal groove, but they either fade out or become very weakly defined on the propodeum
where a punctulate ground-sculpture predominates. Petiole and postpetiole finely and densely
reticulate-punctulate, the first gastral tergite very densely finely shagreened and opaque. All dorsal surfaces
of head and body with distinctive short stout blunt hairs. Body colour more or less uniform medium brown
but the mandibles, clypeal lobe and antennae yellow. Propodeal spines yellowish, lighter in colour than the
propodeum itself.

This very distinctive species is easily separable from all other known African forms by its
flat-margined clypeal lobe, deep metanotal groove, large eyes, sharply marginate pronotum with
dentate corners and sharply triangular petiole node. It is not obviously related to any other
species of Leptothorax in the region and, as far as is known, is only represented in collections by
the holotype.

MATERIAL EXAMINED
' East Africa ' : no loc. (Staudinger & Bang-Haas).

Leptothorax innocens (Forel)
(Fig. 20)

Tetramorium (Leptothoraxl) innocens Forel, 191 3a: 317. Holotype worker, ZAIRE: Katanga, Elizabeth ville

( = Lubumbashi) (Bequaert) (MHN, Geneva) [examined].
Leptothorax innocens (Forel) Forel, 1916: 425.

WORKER. TL 2.3-2.5, HL 0.60-0.62, HW 0.49-0.50, CI 81-82, SL 0.34-0.35, SI 69-70, PW 0.35-0.36, AL
0.66-0.68 (2 measured).

Mandibles extremely finely and delicately superficially shagreened, almost smooth. Median portion of
clypeus with anterior margin evenly arcuate-convex, with a narrow cuticular apron. Median clypeal carina
vestigial to absent. Frontal carinae and antennal scrobes absent, the scapes short. Maximum diameter of eye
0.12-0.13, about 0.24-0.26 x HW and with 7-8 ommatidia in the longest row. With the head in full-face
view the sides narrower in front of the eyes than behind and somewhat convergent anteriorly. Behind the
eyes the sides very slightly convex and rounding into the occipital corners, the occipital margin itself very
feebly impressed medially to almost straight. With the alitrunk in profile the promesonotum shallowly
convex, the metanotal groove impressed and the propodeum convex, more strongly so than the
promesonotum. Propodeat spines broadly triangular and stout, about as long as their basal width.
Propodeal declivity concave between the spines and the rounded low metapleural lobes. In dorsal view the
alitrunk with the pronotal shoulders obtusely and bluntly rounded, the promesonotum narrowing to the
metanotal groove. Behind this the sides of the propodeum diverge to about the level of the spiracle, pass
through an obtuse angle and then converge again to the bases of the spines. Petiole in dorsal view with a
tubercle at either side of the node and the posterior margin of the node with a much smaller and
inconspicuous pair of tubercles. Petiole in profile with a short anterior peduncle which has a small

AFROTROPICAL MYRMICINE ANT GENERA 331

triangular process ventrally. Dorsal surfaces of head, alitrunk, petiole and postpetiole finely densely sharply
reticulate-punctate, the promesonotum and head also with traces of fine rugular sculpture. First gastral
tergite basally with very delicate superficial shagreening. All dorsal surfaces of head and body with
numerous very short blunt hairs; legs and scapes without standing hairs. Colour uniform yellow.

This small species is closest related to stramineus but the two are separated by the characters
given in the key and noted under the discussion of stramineus.

MATERIAL EXAMINED
Zaire: Katanga, Elizabeth ville (Bequaert).

Leptothorax megalops Hamann & Klemm
(Figs 13, 14)

Leptothorax (Icothorax) megalops Hamann & Klemm, 1967: 417, fig. 1. Holotype worker, and paratype
female, SUDAN : Wadi Haifa, 2811962 (H. Hamann & W. Klemm) (NM, Vienna) [examined].

WORKER. TL 2.5, HL 0.67, HW 0.47, CI 70, SL 0.52, SI 1 10, PW 0.35, AL 0.72.

Mandibles almost smooth, with faint vestiges of superficial sculpture. Median lobe of clypeus prominent,
its anterior margin evenly but shallowly convex. Median clypeal carina present, fine but distinct. Frontal
carinae and antennal scrobes absent. Head relatively longer and narrower and scapes relatively longer than
any other known species of the region (CI and SI above). Eyes relatively large, maximum diameter 0.18,
about 0.38 x HW and with 1 1-12 ommatidia in the longest row. With the head in full-face view the occipital
margin feebly indented medially, rounding broadly and evenly into the sides; the latter approximately
parallel but converging slightly anteriorly. Alitrunk in profile with the metanotal groove shallowly
impressed, the propodeal spines short, stout and straight. Metapleural lobes low and rounded, the declivity
between the spines and the metapleural lobes more or less straight. In dorsal view the pronotal angles
broadly and evenly rounded. Petiole in profile with the node bluntly triangular (Fig. 14), the anteroventral
process small and triangular. Dorsum of head with fine longitudinal rugulae the spaces between which are
smooth or at most only faintly superficially punctulate. Dorsal alitrunk with more conspicuous punctulate
ground-sculpture which is overlaid by fine, predominantly longitudinal rugulae, although these are irregular
in places. Petiole and postpetiole finely punctulate with traces of fine rugulae. First gastral tergite
unsculptured. All dorsal surfaces of head and body with quite long blunt hairs which are, however, shorter
on the head than on the alitrunk; the appendages without such hairs. Colour uniform yellow.

This very conspicuous species is easily recognized by its long narrow head, long scapes, large
eyes, rounded pronotal corners and lack of denticles on the petiole node, coupled with its
impressed metanotal groove and possession of hairs on the first gastral tergite. The closest
relatives of megalops in the Afrotropical region are evelynae and cenatus. Both are distinguished
from megalops quite easily as the former lacks pilosity on the first gastral tergite except for a
sparse apical row, and the latter has strongly sculptured mandibles, smaller eyes and a broader
head, and has the petiole node differently shaped (Figs 14, 15).

MATERIAL EXAMINED
Sudan: Wadi Haifa (Hamann & Klemm).

Leptothorax simoni (Emery) comb. n.
(Fig. 21)

Tetramorium simoni Emery, 1895ft: 35, pi. 2, fig. 22. Lectotype worker, SOUTH AFRICA: Transvaal, Makapan

(E. Simon) (MCSN, Genoa), designated by Bolton, 1976: 292 [examined].
Tetramyrma simoni (Emery) Emery, 1922a: 291. [See also Bolton, 1976: 291.]

WORKER. TL 4.7, HL 1.10, HW 0.92, CI 84, SL 0.72, SI 78, PW 0.74, AL 1.32.

Mandibles with faint longitudinal sculpture and scattered pits. Median lobe of clypeus prominent, its
anterior margin shallowly convex. Median clypeal carina distinct. Frontal carinae and antennal scrobes
absent, the scapes of moderate length. Maximum diameter of eye 0.29, about 0.32 x HW and with 17-18
ommatidia in the longest row. With the head in full-face view the occipital margin shallowly convex
centrally, more steeply convex laterally where it rounds into the sides. With the alitrunk in profile the
promesonotum evenly convex, sloping down posteriorly to the shallow metanotal groove. Propodeal

332 B. BOLTON

dorsum more shallowly convex than the promesonotum and strongly depressed below the level of the
promesonotum (Fig. 21). Propodeum armed with a pair of teeth which are slightly shorter than the rounded
metapleural lobes. Petiole node massive, domed in profile, the anterior peduncle short and narrow, equipped
with a dentiform anteroventral process. In dorsal view both petiole and postpetiole inflated, broader than
long. Dorsum of head finely and predominantly longitudinally rugulose, with scattered cross-meshes. On
the sides of the head and occipitally a loose reticulum is present. Ground-sculpture between the rugulae a
fine dense punctulation, superficial in places. Dorsal surfaces of alitrunk, petiole and postpetiole
reticulate-rugose, the meshes narrower and usually more sharply defined on the propodeum and pedicel
segments than on the promesonotum. Fine punctulate ground-sculpture present everywhere but stronger on
the pedicel segments than on the promesonotum. Base of first gastral tergite finely and very densely
reticulate-punctulate, the sculpture weakening posteriorly on the sclerite. Extreme base of first tergite, just
behind the postpetiolar articulation, with very short but strongly developed costulae. Short erect hairs very
sparse on dorsum of head, absent from all other surfaces except the petiole where a single pair is present.
Hairs absent from appendages. Sparse short decumbent to appressed pubescence present on alitrunk and
first gastral tergite; hairs present on tergites of gaster behind the first. Head and gaster dark brown with a
dull reddish tinge, alitrunk and pedicel segments dull red. Mandibles yellow.

A distinctive species characterized by its large size, depressed propodeum, lack of standing hairs
on the first gastral tergite, large eyes and short propodeal teeth. The closest relative of simoni is
braunsi, but in the latter the propodeum is unarmed (Figs 21, 22).

MATERIAL EXAMINED
South Africa : Transvaal, Makapan (E. Simon).

Leptothorax stramineus (Arnold)
(Fig. 18)

Limnomyrmex stramineus Arnold, 1948: 223, figs 10, lOa, lOb. Holotype worker, SOUTH AFRICA: Natal,

Zululand, St Lucia Lake (J. C. Faure) (NM, Bulawayo) [examined].
Leptothorax (Nesomyrmex) stramineus (Arnold) Brown, 1971 : 4.

WORKER. TL 2.8, HL 0.63, HW 0.53, CI 84, SL 0.36, SI 68, PW 0.38, AL 0.68.

Mandibles very delicately shagreened, almost smooth. Median portion of clypeus with anterior margin
evenly arcuate-convex, with a very narrow cuticular apron. Median clypeal carina vestigial. Frontal carinae
and antennal scrobes absent, the antennal scapes short. Maximum diameter of eye 0.13, about 0.25 x HW
and with 7-8 ommatidia in the longest row. Sides of head behind eyes very feebly convex, the sides
narrowing in front of the eyes so that the width immediately in front of the eyes is 0.47 and across the
clypeus at its widest is 0.38 (compare with HW 0.53). Occipital margin approximately straight and the
occipital corners evenly rounded. Alitrunk in profile with promesonotum evenly and shallowly convex, the
propodeal dorsum on the same level but shorter and more strongly convex, the two convexities separated by
the conspicuously impressed metanotal groove. Propodeal spines narrow and somewhat downcurved along
their length, longer than their basal widths. Slope of the declivity between the spines and the rounded
metapleural lobes straight. In dorsal view the alitrunk with the pronotal shoulders obtusely and bluntly
angled, the promesonotum narrowing posteriorly to the metanotal groove. Behind this the propodeum
broadening to the level of the spiracle then narrowing again to the bases of the spines; the latter divergent
and in the holotype with the right spine slightly longer than the left. Petiole node in dorsal view with a
strong lateral tubercle on each side, a pair of smaller tubercles on the posterior margin and a very feeble pair
anterodorsally which are almost effaced. Postpetiole with a low but broad lateral tubercle on each side. In
profile the petiole peduncle short, with a small triangular anteroventral process and with a very small
sub-denticulate process dorsally. The node itself higher than long, the lateral and posterior tubercles
distinct. Sides of first gastral tergite in dorsal view curving evenly away from the articulation with the
postpetiole, without a sharp, flattened appearance. Head, alitrunk, petiole and pcstpetiole very finely and
very densely superficially reticulate-punctulate, without rugulose sculpture. First gastral tergite with only
the faintest vestiges of superficial sculpture basally. Dorsal surfaces of head and body with scattered very
short blunt stout hairs; such hairs absent from appendages and sides of head but one or two may project
from the curved part of the occipital corner on each side. Colour uniform pale yellow.

Among the species which have the metanotal groove impressed stramineus is most closely related
to innocens, the two species sharing the characters of relatively small eyes and having short hairs
on the first gastral tergite (as opposed to the first gastral tergite being hairless). In fact, stramineus

AFROTROPICAL MYRMICINE ANT GENERA 333

and innocens form an extremely close species-pair and may eventually prove to be expressions of
a single species. For the present the two may be separated as the sculpture in innocens, although
punctulate as in stramineus, is much more strongly developed and sharply defined, with traces of
rugular sculpture also present at least on the head. Besides this the petiolar tubercles are not as
strongly developed in innocens as they are in stramineus, and the propodeal spines are shorter and
broader (Figs 18,20).

MATERIAL EXAMINED
South Africa: Natal, St Lucia Lake (J. C. Faure).

MEL1SSOTARSUS Emery

(Figs 23, 24)
Melissotarsus Emery, 1877: 378. Type-species: Melissotarsus beccarii Emery, 1877: 379, fig., by monotypy.

DIAGNOSIS OF WORKER. Myrmicine ants with moderate to conspicuous size variation in most nest samples,
living under bark and in wood of live trees; general appearance as in Figs 23, 24. Mandibles short, when
unworn armed with a long finger-like apical tooth followed by two much smaller teeth and sometimes also
by a minute basal denticle. With wear these gradually become an undifterentiated blunt margin. Palp
formula 0,1 (weissi). Median portion of clypeus bluntly triangular in shape and somewhat raised, not
projecting back between the frontal lobes. Lateral portions of clypeus simple and unmodified. Frontal lobes
narrow, confluent centrally and separated only by a narrow impressed line; the anteriormost parts of the
frontal lobes abut the posterior clypeal margin. Antennal scrobes absent. Frontal carinae absent. Antennae
with six segments, the scapes very short (SI 39-47), the two apical segments forming a strong club. Eyes
present, distinctly longer than broad and set in front of the midlength of the sides. Alitrunk short, fusiform
and box-like, without dorsal sutures or impressions except in the very largest individuals where rarely a
metanotal impression is shallowly present. Propodeum unarmed and rounded. Metapleural lobes absent.
Lateral portions of pronotum reduced to a narrow V-shaped wedge below the level of the conspicuous
mesothoracic spiracle. Anterior coxae small, much smaller than the massively developed middle and hind
coxae. Propodeal spiracle round, situated low on the side of the propodeum and just behind its midlength.
Metapleural gland system easily visible through the cuticle. Basitarsal segment of each leg greatly swollen,
as wide as the preceding tibia, terminating apically in a circlet of small teeth on the anterior (leading) edge on
the middle and hind basitarsi. Petiole with an anterior peduncle and a small low posteriorly situated node
which is broadly attached to the postpetiole; the latter broadly attached to the gaster. Dorsal alitrunk finely
longitudinally costulate throughout. Elongate fine hairs present dorsally on head and body, and also present
on the upper surfaces of the scapes and outer surfaces of the tibiae.

This small genus, of which only four uncommon species are presently recognized, is restricted to
the Malagasy region (1 species) where it is rare, and the Afrotropical region (3 species) where it is,
however, very widespread. The species nest in the healthy wood of living trees, apparently
tunnelling their own galleries below the surface. For this reason most collections of Melissotarsus
are made more by luck than by intent as their presence in the wood is usually not detectable on
the surface. Delage-Darchen (1972) has shown that the method of walking in these ants is very
strange; they progress on their front and hind legs with the middle pair projecting upwards, and
presumably in contact with the gallery roof. She also noted the presence of coccids inside the
galleries, also discussed by Ben-Dov (1978). It seems probable that coccid secretions form a
major, if not the main, item in the diet of Melissotarsus species.

The genus most closely related to Melissotarsus is Rhopalomastix Forel, represented by three
or four poorly defined species distributed throughout the Oriental and Indo- Australian
zoogeographical regions and utilizing the same lifeway as Melissotarsus. Since Emery (1922a) and
Wheeler (1922) produced their classifications these two small genera have always been placed
together in a tribe of their own (Melissotarsini) and it is fairly certain that they represent two
stages on a single adaptive line. Rhopalomastix is the more generalized of the two, Melissotarsus
decidedly the more specialized, but the modifications seen in the latter are foreshadowed in the
former genus. It is the accentuation of these adaptive specializations which separates the genera,
as follows.

334

B. BOLTON

23

29

Figs 23-29 23, 24, profile and head of Melissotarsus weissi. 25-29, Messor workers. 25, profile of angularis.
26-29, heads of (26) angularis, (27) striatifrons, (28) decipiens, (29) denticornis. Pilosity omitted from 24,
26-29.

AFROTROPICAL MYRMICINE ANT GENERA

335

Rhopalomastix
Antennae 10-segmented.

Lateral portion of pronotum extensive, distinctly
larger than the mesopleuron.

First coxa as large as or larger than the second and

third coxae.
Petiole sub-sessile, with a strong ventral process.

Free posterior face of petiole node long, its
articulation with the postpetiole narrow.

Basitarsal segment of each leg not swollen, without
apical circlets of teeth.

Sting long and strong.

Melissotarsus
Antennae 6-segmented.
Lateral portion of pronotum very reduced,

forming a V-shaped narrow wedge which is

smaller than the mesopleuron.
First coxa much smaller than the swollen second

and third coxae.
Petiole short-pedunculate, with feeble or no

ventral process.
Free posterior face of petiole node very short, its

articulation with the postpetiole very broad.
Basitarsal segment of each leg strongly swollen,

with apical circlets of teeth.
Sting very reduced and probably non-functional.

So little material of Melissotarsus is available at present that this survey must be regarded as
strictly preliminary. Three species are now recognized in the Afrotropical region but it is possible
that each may be compounded of more than one different sibling-species. Conversely it is by no
means impossible that further collections will bridge what appear here as species tor the
differences between them, though consistent in the few samples to hand, are relatively minor and
may well be anulled by further collecting.

For the present I define weissi as having a dark brown to black strongly sclerotized male, and a
similarly coloured female in which the postpetiole in dorsal view is quite narrow
(1.20-1.40 x broader than long) and has a rounded or even hemispherical anterior margin. The
worker of weissi has the alitrunk medium to dark reddish brown, the anterior margin of the
pronotum in dorsal view sharply defined and angular where it meets the anterior declivity, and
the sides of the alitrunk meeting the dorsum in a fairly well-defined angle.

M. emeryi and beccarii, on the other hand, have pale yellow feebly sclerotized males, and have
females in which the postpetiole in dorsal view is quite broad (1.90-2.20 x broader than long)
and lacking a rounded anterior margin, the margin instead being more or less straight or even
slightly concave. The workers are yellow to light yellowish brown and have the sides of the
alitrunk rounding bluntly into the dorsum when seen in dorsal view. Females of emeryi differ
from those of beccarii as in the former the mesoscutum is broader than long in dorsal view; it is
longer than broad in the latter. Workers of emeryi have the anterior margin of the pronotum
sharply defined and angular where it meets the anterior declivity, whereas in beccarii there is no
such sharp differentiation between dorsum and anterior declivity, instead the one surface rounds
bluntly into the other.

The shape of the alitrunk in dorsal view shows subtle but perhaps significant differences
between separate series of workers presently grouped as single species, but discovering whether
these differences are meaningful, or even consistent, will have to await the amassing of
considerably more samples than are presently available.

Synonymic list of Afrotropical Melissotarsus species

beccarii Emery

titubans Delage-Darchen syn. n.
emeryi Forel

emeryi var. pilipes Santschi syn. n.

compressus Weber syn. n.
weissi Santschi

major Santschi syn. n.

Key to species (workers)

1 With the alitrunk in dorsal view the anterior margin of the pronotum rounding evenly into the
anterior declivity, the two not meeting in a sharp angle or edge. (Ethiopia, Tanzania, South
Africa, Ivory Coast) beccarii(p. 336)

336 B. BOLTON

With the alitrunk in dorsal view the anterior margin of the pronotum separated from the anterior

declivity by a sharp angle or edge 2

2 Sides of alitrunk meeting dorsum in a fairly well-defined angle. Alitrunk colour medium to dark

reddish brown. (Ghana, Congo, Zaire) weissi(p. 337)

Sides of alitrunk rounding bluntly into the dorsum. Alitrunk colour yellow to light yellowish
brown. (Ethiopia, Sudan, Kenya, Tanzania, Zaire, Central African Republic, South Africa,
Ivory Coast, Ghana) emeryi (p. 337)

The three presently recognised species are basically so similar that to present a full description for
each would be redundant so, for the purposes of identification, a description of the type-species
beccarii is given and the other two are compared to it.

Metissotarsus beccarii Emery

Melissotarsus beccarii Emery, 1877: 379, fig. Syntype workers, ETHIOPIA: Keren (Beccari) (MCSN, Genoa;

MHN, Geneva) [examined].
Melissotarsus titubans Delage-Darchen, 1972: 216, figs 1-10. Syntype workers, females, males, IVORY COAST:

Lamto (Delage-Darchen) (probably in collection of Delage-Darchen). Syn. n.

WORKER. TL 2.3-3.3, HL 0.56-0.82, HW 0.56-0.80, CI 97-105, SL 0.24-0.34, SI 39^17, PW 0.34-0.55, AL
0.58-0.80(15 measured).

With the head in full-face view the occipital margin concave, sometimes deeply so medially, and with the
sides convex and weakly to distinctly convergent in front of the eyes. Mandibles with a long finger-like
apical tooth, worn down to nothing in some specimens; the mandibles unsculptured. Eyes much longer than
broad, strip-like in many, the maximum diameter 0.12-0.16, about 0.18-0.22 x HW. Median portion of
clypeus raised above the level of the lateral portions, not extending back between the frontal lobes; the latter
contiguous and separated only by an impressed line. Scapes very short, SI < 50. Alitrunk in dorsal view with
anterior pronotal margin rounding into the declivity, the two surfaces not separated by a sharp edge or
angle. Dorsum of alitrunk roughly rectangular longitudinally, somewhat narrower behind than in front but
not strongly so, and with the dorsum rounding into the sides. In profile the promesonotal dorsum and
anterior propodeum are more or less flat but the posterior part of the propodeum rounds very broadly and
evenly into the declivity, without trace of armament. Fore coxae small, about half the size of the strongly
swollen middle and hind coxae. Peduncle of petiole short and grading into the relatively high narrow node,
the node with a short posterior free face, broadly attached to the postpetiole. In dorsal view the petiole node
much broader than long. Postpetiole in dorsal view much broader than long, slightly broader than the
petiole and very broadly attached to the first gastral tergite without a posterior constriction. Gaster only
feebly sclerotized, crumpled in most mounted specimens. Dorsum of head with a silky superficial
ground-sculpture upon which scattered small pits are usually superimposed. The ground-sculpture may
cover the whole head but frequently it fades out occipitally. Median portion of clypeus more densely and
strongly sculptured than dorsum of head capsule. Dorsal alitrunk finely longitudinally costulate
throughout, the costulae fading out where the propodeal dorsum rounds into the declivity. Dorsal surfaces
of head, scapes, pronotum, mesonotum, pedicel segments and gastral tergites with scattered sparse long fine
hairs. Propodeal dorsum usually with one or two shorter hairs but these are frequently missing. Dorsal
(outer) surfaces of tibiae with sparse long hairs similar to those on alitrunk. Head and alitrunk dull yellowish
brown to dark yellow, the gaster lighter, usually pale dull yellow.

The key character given to separate beccarii and emeryi workers is quite weak. In most
individuals there is a reasonable visible difference between the two, with the anterior pronotal
margin rounding bluntly into the declivity in beccarii, and with the anterior pronotal margin
separated from the declivity by an angle or edge in emeryi. Having said that, however, it should
be pointed out that the difference is not so well marked in some individuals, which in
consequence are difficult to place. Both species have a pale yellow feebly sclerotized male.
Females of both species have the postpetiole in dorsal view conspicuously broader than long but
it seems that two species are present as in some the mesoscutum is longer than broad (beccarii)
but in others broader than long (emeryi). It should be admitted that very few worker-associated
females are known and further collections may annul this apparent difference. To sum up, for the
present I recognise these two as separate on the strength of the differently shaped mesoscutum in
females and the form of the anterior pronotal margin in workers, but harbour a suspicion that
only a single real species may in fact be represented here.

AFROTROPICAL MYRMICINE ANT GENERA 337

M. beccarii differs from weissi fairly consistently in all castes. The workers of weissi are darker
in colour than those of beccarii and have both the anterior pronotal margin and the sides of the
alitrunk relatively strongly marginate. The male is dark brown to black and strongly sclerotized,
and in the female the postpetiole is relatively narrow in dorsal view with an arched-convex
anterior margin which is quite different in shape from the strongly transverse form seen in
beccarii. Measurements of the postpetiolar widths of the various forms are given under the
discussion of the genus.

MATERIAL EXAMINED

Ivory Coast: Lamto Field Station (W. L. Brown); nr Abidjan (W. L. Brown). Ethiopia: Keren (Beccari).
Tanzania: Lulanguru (G. D. H. Carpenter). South Africa: Natal, Durban (C. B. Cooper); Durban (H. B.
Mar ley).

Melissotarsus emeryi Forel

Melissotarsus emeryi Forel, 1907: 133. Syntype workers, ETHIOPIA: Colba, 1905 (M. de Rothschild) (MHN,

Geneva) [examined].
Melissotarsus emeryi var. pilipes Santschi, 1914a: 71. Syntype workers, KENYA: Taveta, 750 m, st. no. 65,

iii. 1912; and TANZANIA: Kilimanjaro, Bismarckhiigel, 2740 m, st. no. 70, iii.!912(C. Alluaud& R.Jeannel)

(NM, Basle) [examined]. Syn. n.
Melissotarsus compressus Weber, 1952: 1, figs 28, 29. Holotype female, CENTRAL AFRICAN REPUBLIC:

Ubangi-Shari, Haut Mbomu, lat. 5° 30' N., long. 25° 15' E., iii. 1948, no. 2184 (N. A. Weber) (AMNH,

New York) [examined]. Syn. n.

WORKER. TL 2.5-3.4, HL 0.66-0.88, HW 0.70-0.90, CI 100-105, SL 0.30-0.38, SI 39^3, PW 0.37-0.57, AL
0.62-0.88 (13 measured).

Answering to the description of beccarii but with the anterior pronotal margin in dorsal view separated
from the anterior declivity by a well defined angle or edge.

As pointed out under beccarii a few individuals seem intermediate between those of emeryi and
those of beccarii, and in consequence are difficult to place. My suspicions are that these two
names may represent a single species but I feel unsure enough to avoid synonymizing them whilst
the apparent difference between the females remains unresolved. So, until the taxonomic value of
the shape of the mesoscutum in females can be assessed, the two must remain as separate species.

MATERIAL EXAMINED

Ethiopia: Colba (Rothschild). Sudan: Darfur, Jebel Murra (M. Steele). Kenya: Muguga (K. Njukiine);
Taveta (Alluaud & Jeannel). Zaire: Popokabaka (E. S. Ross & R. E. Leech). Ghana: Tafo (C. A.
Collingwood). South Africa: Cape Prov., Clanwilliam (Y. Ben-Dov). Central African Republic: Haut Mbomu
(N. A. Weber).

Melissotarsus weissi Santschi

(Figs 23, 24)

Melissotarsus weissi Santschi, 1910: 356, fig. 3. Holotype female, CONGO: Brazzaville (A. Weiss) (NM, Basle).

[Only gaster and one forewing remaining on mount.]
Melissotarsus major Santschi, 1919: 85. Syntype workers, ZAIRE: Penghe, 13.ii., no. 125 (Bequaert) (NM,

Basle; MR AC, Tervuren) [examined]. Syn. n.

WORKER. TL 2.3-3.0, HL 0.58-0.74, HW 0.60-0.78, CI 98-104, SL 0.27-0.34, SI 41-47, PW 0.36-0.50, AL
0.56-0.84 (14 measured).

Answering to the description of beccarii but darker in colour, the alitrunk medium to dark reddish brown;
with the anterior pronotal margin meeting the anterior declivity in a well-defined angle or edge, and with the
sides of the alitrunk meeting the dorsum in a fairly well-marked angle.

Lighter coloured workers may sometimes be difficult to separate from emeryi, but in general the
sharper marginations of the sides of the alitrunk in weissi are fairly distinct. The sexual forms of
weissi are both easily separated from those of emeryi as the male of the former is dark brown to
black (pale yellow and feebly sclerotized in the latter), and the female of weissi has the postpetiole
relatively narrow in dorsal view with an arched-convex anterior margin, as opposed to a very
broad and distinctly transverse postpetiole in emeryi.

338 B. BOLTON

MATERIAL EXAMINED

Ghana: Tafo (B. Bolton). Zaire: Kamaiembi (H. Schouteden); Penghe (Bequaeri).

MESSOR Forel

(Figs 25-32, 35-43)

Messor Forel, 1890a: Ixviii [as subgenus of Aphaenogaster Mayr]. Type-species: Formica barbara L., 1767:

962, by subsequent designation of Bingham, 1903 : 277.
Messor Forel; Bingham, 1903: 277. [Raised to genus.]
Cratomyrmex Emery, 1891: 572. Type-species: Cratomyrmex regalis Emery, 1891: 572, by monotypy.

[Synonymy by Emery, 1922a: 357.]
Veromessor Forel, 1917: 235 [as subgenus of Novomessor Emery]. Type-species: Aphaenogaster andrei

Mayr, 1886: 448, by subsequent designation of Emery, 1921 : 67. Syn. n.
Veromessor Forel; Wheeler, 1922: 680. [Raised to genus.]
Lobognathus Enzmann, 1947: 152 [as subgenus of Veromessor]. [Erroneous entry for Veromessor

lobognathus (Andrews); see Brown, 1949: 49.]

DIAGNOSIS OF WORKER. Granivorous myrmicine ants, mostly strongly polymorphic but a few monomorphic
or only weakly polymorphic. Head massively constructed in larger workers. Mandibles large and powerful,
multidentate in smaller workers (up to 15 teeth) but this number usually decreasing with increased body size
until in largest workers only a few massive teeth or an edentate crushing edge remains. Sometimes also in
small workers the teeth are worn down to an edentate margin. Palp formula predominantly 4,3 but in
largest workers usually 5,3 (30 species dissected). Median portion of clypeus broad and shield-like, broadly
inserted between the widely separated frontal lobes; both median and lateral portions of clypeus unmodified
except for a central impression of the anterior margin in some species. Frontal lobes short but conspicuous,
at least partially concealing the antennal insertions. Frontal carinae absent. Antennal scrobes absent.
Antennae 12-segmented, either filiform and without an apical club (in which case the flagellar segments
gradually increase in size apically), or with a feebly defined incipient club where the apical 3-4 segments are
slightly enlarged. Eyes present, moderate to large in size, situated at or just behind the midlength of the sides
in full-face view. Ventral surface of head with elongate ammochaete hairs which usually form a
psammophore. This may be reduced and non-functional in some species but the hairs are still conspicuous
and generally longer than those found elsewhere on the body; in a few species the psammophore is better
developed in smaller than in larger workers. With the alitrunk in profile the promesonotum swollen and
convex, frequently dome-like and sloping down steeply behind to the metanotal groove which is weakly to
distinctly impressed. Propodeum rounded to strongly bispinose posteriorly and on a much lower level than
the convex promesonotum. Promesonotal suture fused and inflexible but its track represented by a distinct
arched impression across the dorsum. Mesonotum bounded by impressions on all sides, its boundary easily
discernible except in the smallest workers of a few species. Metapleural lobes absent or at most represented
by a pair of low broadly rounded ridges. Propodeal spiracle large and conspicuous, circular to subcircular
and situated approximately at the midlength of the propodeum or sometimes slightly behind the midlength,
but never shifted conspicuously back towards the declivity. Basal posterior portion of mesopleuron just
above the middle coxa with a few hairs projecting downwards and backwards. (Whether these are
guard-hairs indicating the exit site of a gland is not known, but the hairs remain even in species where other
body pilosity is very reduced or absent.) Spurs on posterior tibiae varying from very feebly pectinate through
partially barbate and minutely barbulate to simple. Alitrunk ventrally with a strong metasternal process
which is usually large to very large (reduced but still conspicuous only in rufotestaceus (Foerster) and
vaucheri Emery out of 45 species dissected). Petiole with a long anterior peduncle, the spiracle situated at
about the midlength of the peduncle, well in front of the node. Petiole node in profile narrow and often
bluntly triangular to conical in shape, but frequently a sloping differentiated dorsal surface is present where
the anterodorsal angle is generally the highest point.

Messor is a moderately sized genus of granivorous ants occurring in grassland and savannah, and
in arid to desert situations. The main base of the genus is in the Palaearctic region where about
70-80 species occupy a broad strip of territory reaching across the whole width of North Africa
and the southern European countries, across the Near and Middle East and thence eastwards
through the U.S.S.R. to China and Japan. Compared to this the faunas of other zoogeographical
regions are relatively minor. The Afrotropical region has 12 species and Madagascar has 1; the
Oriental region has 3-4 species and the Nearctic has 8, all distributed on the western side of the
continent and formerly occupying a genus of their own, Veromessor, now synonymized. Species

AFROTROPICAL MYRMICINE ANT GENERA 339

of Messor are absent from the Neotropical region, the Indo-Australian region and Australasia,
nor do they occur on any of the Pacific island systems.

Recent studies of Messor include those of Arnoldi (1977) on the fauna of the U.S.S.R., and
Collingwood (1978) on the species of the Iberian Peninsula. The only previous synthesis of
sub-Saharan African species is that of Arnold (1920), for the then-recognized South African
forms, but no key was given in that revue. Creighton (1950) has keyed the North American
species formerly in Veromessor. Knowledge of the detailed biology of the species is sparse, but
good basic work has been done on some African species by Levieux & Diomande (1978), and
Levieux (1979).

The closest relatives of Messor are the genera Aphaenogaster and Pheidole Westwood.
Members of the latter genus are easily separated from Messor as the palp formula is reduced to
2,2, its species are dimorphic, and the antennal funiculus ends in a strongly defined 3-segmented
club. Aphaenogaster, which is absent from sub-Saharan Africa, is more difficult to differentiate as
its species, apart from being uniformly monomorphic, are very close to Messor and share most of
its diagnostic characters, including the filiform to feebly clavate funiculi and high palp formula
(PF) count. Of 55 species of Aphaenogaster dissected 31 had PF 5,3, and 24 had PF 4,3. For some
reason, although species with the higher PF apparently outnumber those with the lower count,
the zoogeographical distribution of the latter is much wider than that of the former.
Aphaenogaster species with PF 5,3 are found in the Nearctic, Palaearctic and Oriental regions;
species with PF 4,3 are also found in these three regions and in the Neotropical, Malagasy,
Indo-Australian and Australasian regions as well.

After a study of Aphaenogaster for genus-level characters, primarily a search for strong
characters to separate it from Messor, it became apparent that Brown (1973) was correct in
relegating the former subgenera of Aphaenogaster to the synonymy. These former subgenera
(Attomyrma Emery, Deromyrma Forel, N ystalomyrma Wheeler and Planimyrma Viehmeyer) have
no significance as they are founded upon minor, inconsistent or gradient character-states.
Further, it is now clear that Brown (1974) was also correct in assigning Novomessor to the
synonymy of Aphaenogaster. The only real character separating the two was the fore-wing
venation, there being one closed cubital cell in the former and supposedly two in the latter. The
same character was invoked to separate Veromessor from Messor, again the former having one,
the latter two closed cubital cells. A survey of the venation of Aphaenogaster and Messor shows
that in both genera the same finely graded series of changes in wing venation occurs (Figs 35-43),
which obviates these supposed differences in number of closed cubital cells; it is instructive to
consider both genera together.

The most complete, and therefore most primitive, venation pattern (Fig. 35, M. galld) shows
two closed cubital cells and has Rs + M dividing well in front of the level of cross- vein m — cu, so
that m — cu arises from M itself and there is a short free section of M between the point of
division of Rs + M into its constituent parts and the point where m — cu meets M.

The free section of M then contracts (Fig. 36; M. tropicorum, angularis, nigriceps Santschi; A.
geei Wheeler, schurri (Forel)) as the fusion of Rs + M lengthens outwards along the wing until the
condition shown in Fig. 37 is seen (M. rugosus (Andre); A. schurri) where there is no free portion
of M between Rs + M and the point of origin of m — cu, the veins Rs, M and m — cu all
appearing to arise from a point at the apex of Rs + M.

Next, the fusion of Rs + M advances further out along the wing so that Rs and M now
separate a short distance beyond the point of origin of m — cu, which now arises direct from
Rs + M (Fig. 38; M. intermedius Forel, angularis, himalayanus Forel, aciculatus (Smith), structor
(Latreille), regalis; A. rudis Emery, treatae Forel). Following this the fusion of Rs + M advances
further out along the wing, drawing closer to cross-veins 2r and r — m, as shown in Figs 39, 40,
this stage constituting what may be considered as the normal pheidoline venation (M. barbarus
(L.), capitatus (Latreille), structor, galla, denticornis, capensis, leubberti, muticus (Nylander),
aegyptiacus (Emery), nigriceps Santschi, semirufus (Andre), instabilis (Smith), meridionalis (Andre);
A. geei, rudis, lamellidens Mayr,famelica (Smith), fulva Roger, japonica Forel, pallida (Nylander),
huachucana Creighton, splendida (Roger), megommatus Smith, subterranea (Latreille), occidentalis
Emery, crocea Andre, gemella (Roger), senilis Mayr).

340 B. BOLTON

As the fusion of Rs + M progresses still further along the wing a critical point is reached at
which cross-vein r — m vanishes. This occurs whilst the advancing fusion is still some little
distance away from 2r. A male of A. spinosa Emery in BMNH shows the critical point as the
specimen has r — m present on the left wing but it has vanished from the right. The dissapearance
of r — m leaves the venation as in Fig. 41, which is present in a wide range of species (M.
pergandei (Mayr), lobognathus, formerly of Veromessor; A. albisetosus Mayr, formerly of
Novomessor; A. dromedarius (Emery), longiceps (Smith), pythia Forel, phalangium Emery, beccarii
Emery, araneoides Emery, sagei Forel).

Eventually the stage seen in Fig. 42 is reached where Rs and M are fused to the point of
intersection of 2r (M. andrei (Mayr); A. cockerelli Andre), and finally in A. ensifera Forel the
fusion of Rs + M has extended beyond the level of 2r so that this cross-vein now arises from
Rs + M (Fig. 43).

It should be pointed out that there is considerable variation present along this sequence within
single species and that it is by no means rare to find specimens with different venation patterns on
the left and right forewings, representing different stages in the sequence, and thus showing it to
be a dynamic rather than a static system. Also, adventitious vein-stubs frequently arise at random
from all the main veins, and from the cross-veins too on occasion.

Thus the loss of r — m cross- vein, reducing the two cubital cells to only one, rather than being
the concise taxonomic character it was thought to be in the past, can now be seen as just one step
in a long gradual sequence of venation development in both Aphaenogaster and Messor, and of
no significance in genus-level discrimination among these ants. To draw a line at any point in the
sequence and claim that it is more significant than a line drawn at any other point is thus purely
arbitrary, and as a direct consequence of the establishment of this sequence the synonymy of
Novomessor with Aphaenogaster is confirmed and the name Veromessor falls into the synonymy
of Messor, there being no other consistent character to separate them.

A side development in the history of Novomessor, following Brown's (1974) synonymy, was the
suggestion of Holldobler, Stanton & Engel (1976) that the name might be resurrected for two of
its former members (albisetosus and cockerelli) because of the presence of an exocrine gastral
glandular system which was absent from other Aphaenogaster species examined, and incidentally
absent also from the third former Novomessor species (ensifera), which was to be retained in
Aphaenogaster despite the fact that it is otherwise very close to the first two. The obvious
inference was that the presence of such a gland system merited genus-level consideration. This is
reasonable logic as far as it goes, though many would argue (myself included) that basing genera
on such features is grossly over-weighting a relatively weak single character. The discussion
would probably have rested there but Kugler (1978) published a paper indicating that such
glands occur widely in the Myrmicinae in a range of genera, including a member of
Aphaenogaster (phalangium) whose placement in that genus has never been doubted, but which is
not closely related to any of the three mentioned above. This gives rise to three possibilities.

Firstly, that the presence of such glands is highly significant and that, following the model of
Aphaenogaster-Novomessor, every species showing such structures must be assigned to a genus
separate from the parent genus, irrespective of any basic similarities they may otherwise show.
The idea is ludicrous of course, and obviously not at all what Holldobler et al intended; the
plethora of pointless generic names thus produced would be incredible and no more sensible than
selecting genera from groups of closely related species on grounds of, say, presence or absence of
hairs on the first gastral tergite. Holldobler et al. in their study found a gastral exocrine system in
one species of Ocymyrmex Emery but not in two others; they did not suggest the creation of a
separate genus here.

Secondly, we can reassign such forms with gastral exocrine glands (or indeed any other
individual specialization) when it suits us to do so, and ignore it otherwise. Thus we can utilize
such a character to prop up an otherwise poorly defined or undefinable genus which looks like
falling irrevocably into the synonymy. This idea does not hold much merit as it again leads
unerringly to the creation of swarms of peripheral genera, each with only one or two species,
which cannot be adequately separated from their closest relatives remaining embedded in the
central mass of species.

AFROTROPICAL MYRMICINE ANT GENERA 341

Finally, we can consider that the development of such gland systems in some species of a genus
but not in others, whilst uniformly stable genus-level characters span the entire range of species,
reflects a specialization in the lifeway of the ants involved and is significant at species or
species-group level but not beyond that, providing always that other genus-level characters
remain uniform throughout. This is decidedly the alternative which I favour as in the long run it
will produce strong, well-defined genera, and realistic species-groups within those genera.

To conclude the observations on the genus-level synonymy of Aphaenogaster, it is now
apparent that the monotypic genus Brunella Forel sinks as a synonym. This Malagasy species has
had a chequered career since its original description as Aphoenogaster [sic] belli Forel, 1895; 248.
(Syntype workers, MADGASCAR: Imerina, Moramanga (M. Sikora) (MHN, Geneva) [examined].)
It was later shifted by Forel (1917) out of Aphaenogaster to form the type-species of his genus
Brunella. Emery (1922a: 242) disagreed with this and synonymized Brunella under Atopula
Emery, which for him was a catch-all genus to which a number of obscure species were relegated.
During my study of the tetramoriine genera (Bolton, 1976) it transpired that the type-species of
Atopula was in fact a Tetramorium, so that the name Atopula fell into synonymy and the
remaining former occupants of Atopula were transferred to other genera. At that time I had not
examined the type-series of belli and so referred the species back to Forel's temporarily
resurrected Brunella. Now, having at last examined the types of belli, it turns out to be a fairly
unexceptional Aphaenogaster which seems to belong to the Oriental sagei-group as it has a broad
occipital margin, relatively short antennal scapes, a moderately well-developed antennal club and
distinct propodeal spines.

A summary of the current genus-level synonymy of Aphaenogaster is given in the appendix,
p. 364.

Of the two names synonymized with Messor above, Cratomyrmex was recognized as a
synonym by Emery as long ago as 1922. The separation of the two was based on the presence of
pectinate hind tibial spurs in the latter and their supposed absence in the former. This was
quickly spotted as a feeble and variable character and the status of the genus challenged
(Santschi, 1920). The form of the hind tibial spurs is in fact very variable in Messor, showing all
stages from feebly pectinate, through barbate and minutely barbulate to simple. Even in the same
series there is sometimes variation in spur form between different-sized workers.

Veromessor, which began its existence as a subgenus of Novomessor, was given generic status
by Wheeler (1922) who separated it from Messor on the venation character discussed above and
now known to be spurious. The discussion in Wheeler & Creighton (1934: 356-360) indicated
that Messor and Veromessor were extremely closely related, but no means of separating them was
given. Presumably only the venation character invoked previously by Wheeler could be found.
The present investigation has shown the two to be synonymous for, leaving aside the venation, all
characters of Messor are duplicated in Veromessor, except for the species relictus Wheeler &
Mann. This last was originally described as a member of Aphaenogaster but was transferred to
Veromessor by Wheeler & Creighton (1934), for no apparent reason. In my opinion it is an
ordinary member of Aphaenogaster, fitting the diagnostic characters of that genus and having all
the criteria required to separate it from Messor which are tabulated below; it is herewith returned
to Aphaenogaster. Finally, the fossil species sculpturatus Carpenter, originally described in
Messor (where it is a junior secondary homonym of sculpturatus Stitz), later included in
Veromessor but suggested as a possible Pogonomyrmex species by Wheeler & Creighton, is
impossible to place at present and requires further study. The living North American species now
included in Messor are andrei (Mayr) comb, n., chamberlini Wheeler, julianus (Pergande) comb, n.,
lariversi (Smith) comb, n., lobognathus Andrews, pergandei (Mayr) comb, n., smithi (Cole)
comb, n., stoddardi (Emery) comb. n.

Aphaenogaster and Messor are very closely related and certainly derive from a single parent
stock. The characters tabulated below will separate them even though a few species show
exceptions to one or another of the characters.

342

B. BOLTON

Aphaenogaster
Entirely monomorphic.

Mostly without ammochaete hairs (present in a

very few species).
Head usually slender, CI 90 at maximum,

generally much less (range 49-90 in 75 species

measured).
Metasternal process small to absent, approaching

size seen in Messor only in A. subterranea (55

species dissected).
Outer margins of mandibles not strongly curved

towards midline, the mandibles triangular in

shape and not massive.

Messor
Mostly polymorphic species (a very few feebly

polymorphic and monomorphic species known).
Mostly with ammochaete hairs present (reduced in

a few species).
Head massive and broad, in medium to large

workers CI > 90 (range 95-125 in 64 species

measured).
Metasternal process large to very large, always

very conspicuous (45 species dissected).

Outer margins of mandibles strongly curved
towards midline, the mandibles massive and
heavy.

Synonymic list of Afrotropical Messor species

angularis Santschi stat. n.
capensis (Mayr)

pseudoaegyptiaca Emery syn. n.

barbarus subsp. capensis var. schencki Forel (unavailable)

braunsi Forel syn. n.

donisthorpei Santschi syn. n.
cephalotes (Emery)

plinii Santschi syn. n.
collingwoodi sp. n.
decipiens Santschi stat. n.

barbarum r. capense var. decipiens Forel (unavailable)

barbarus subsp. capensis var. proba Forel (unavailable)

arcistriatus Santschi syn. n.
denticornis Forel

denticornis var. parvidens Forel syn. n.

denticornis var. brunni Forel syn. n.
galla (Mayr)

barbarum subsp. caduca var. galla Emery (unavailable)

barbarus subsp. semirufus var. rufa Forel (unavailable)

barbarus st. galla var. triempressa Santschi (unavailable)

barbarus st. latinodis Santschi syn. n.

barbarus r. semirufus var. rufula Forel (unavailable)

barbarus subsp. galla var. armata Emery (unavailable)

cjalla st. nobilis Santschi syn. n.

galla var. airensis Bernard syn. n.
incisus Stitz (nomen dubium)
luebherti Forel stat. n.
piceus Stitz
regalis (Emery)

regalis var. rubea Santschi syn. n.

sculpturatus Stitz syn. n.
ruginodis Stitz stat. n. (nomen dubium)
striatifrons Stitz stat. n.
tropicorum Wheeler stat. n.

denticornis var. laevifrons Stitz syn. n.

braunsi var. nigriventris Stitz syn. n.

Key to species (medium to large workers)

Note. The nomina dubia incisus Stitz and ruginodis Stitz are omitted from the key.
1 Hairs absent from first gastral tergite or at most with a single sparse transverse row at the

extreme apex of the sclerite

Hairs present on first gastral tergite, more or less evenly distributed over the whole surface of
the sclerite . ...

AFROTROPICAL MYRMICINE ANT GENERA 343

2 Dorsumofpropodeum with one or more pairs ol standing hairs 3

Dorsum of propodeum without standing hairs 4

3 Dorsum of head coarsely and densely reticulate-punctate everywhere, the mid-dorsal strip also

rugulose. (Niger, Mali) collingwoodi (p. 346)

Dorsum of head smooth everywhere except for the rugulose mid-dorsal strip; without coarse
dense reticulate-punctate sculpture. (Throughout Sahelian zone and northern East Africa,
also occurring coastally in West Africa) galla (p. 349)

4 Head sculptured everywhere with close-packed longitudinal rugulae between which is

reticulate-punctate ground-sculpture. Eyes slightly smaller, 0.15-0.18 x HW in HW range of
2.00-3.12. (Tanzania, Zimbabwe, Angola, Botswana, South West Africa, South Africa)

leubberti(p. 351)

Head smooth except usually for a short central rugular area behind the frontal lobes. Eyes
slightly larger, 0.18-0.21 x HW, in HW range of 2.00-2.76. (Kenya) . . angularis (p. 344)

5 Basal third or more of first gastral tergite strongly and conspicuously sculptured with rugulae,

costulae, coarse reticulate-puncturation, or a combination of these 6

Basal third of first gastral tergite unsculptured except for hair pits and very faint superficial
patterning. In some very large workers a few short basigastral costulae may develop but these
are restricted to the area immediately behind the postpetiole 7

6 With the head in full-face view the sides with projecting hairs. Petiole and postpetiole coarsely

closely and deeply rugose. (Nigeria, Benin Republic, Congo) regalis (p. 352)

With the head in full-face view the sides without projecting hairs. Petiole and postpetiole finely
sculptured with feeble rugulae, dense puncturation or a combination of both. (Ethiopia,
Kenya, Tanzania) cephalotes (p. 346)

7 Posterior half of clypeus between frontal lobes with a distinct, strongly raised central step or

welt. (Angola, South West Africa) tropicorum (p. 354)

Posterior half of clypeus between frontal lobes without a raised central step or welt, usually
more or less flat or even slightly concave .......... 8

8 Eyes relatively large, the maximum eye diameter 0.21-0.25 x HW, in HW range of

2.50- > 4.00. (Botswana, South West Africa, South Africa) .... denticornis (p. 349)
Eyes smaller, the maximum eye diameter 0.14-0.19 x HW, in HW range of 2.50- > 4.00 . . 9

9 In HW range 2.80- > 4.00 the sides of the head conspicuously evenly convex in full-face view

(Fig. 27). Propodeum in profile relatively long and low (Fig. 32). (South West Africa, South

Africa) stria tifrons (p. 353)

In HW range 2.80- > 4.00 the sides of the head approximately straight in full-face view, the
sides parallel or divergent anteriorly (Fig. 28). Propodeum in profile relatively short and high
(Fig. 31) '10

10 Body pilosity very dark in colour, deep red-brown to blackish. (Botswana, South Africa)

piceus (p. 352)
Body pilosity pale, white or silvery to yellowish 11

1 1 Head red in majo'r workers, contrasting in colour with the much darker alitrunk and gaster.

(Zimbabwe, Botswana, Lesotho, South Africa) decipiens(p. 348)

Head brown to black in major worker, about the same colour as the alitrunk and gaster. (South
West Africa, Botswana, South Africa) capensis (p. 345)

Among strongly polymorphic species such as these, where there is an enormous worker
size-range, the standard measurements which I have otherwise used consistently for the
Myrmicinae become meaningless and cannot be utilized. A few standard ratios have, however,
proved to be of value in some cases and these are included in the relevant descriptions. The keys
and descriptions are based on medium to large workers as these show the best discriminating
characters, the minor workers of closely related species being sometimes indistinguishable. Size
ranges covered by the descriptions are given for each species in terms of H W range.

The presence or absence of propodeal teeth or spines, which appears to be a functional
diagnostic character in other parts of the range of Messor, is not of much use in the Ethiopian
region species for, although some always have the propodeum armed (regularis, collingwoodi) and
some always have it unarmed and rounded (angularis, luebberti), the rest show a disconcerting
variability in this character, sometimes differing even in individuals from the same nest sample.

The 12 recognizable species are distributed roughly as follows in the Afrotropical region.

344

B. BOLTON

Northern (Sahelian) species: collingwoodi, galla, cephalotes (in extreme east).

Western species : regalis, galla (coastally).

Eastern species : cephalotes, angularis, leubberti (in south), galla (in north).

Southern species : denticornis, luebberti, striatifrons, tropicorum, piceus, decipiens, capensis.

The species fall into two groups in terms of pilosity. The first group, characterized generally by
reduced pilosity and virtual absence of hairs on the first gastral tergite, contains the species
angularis, collingwoodi, galla, and luebberti. In the second group pilosity is generally dense and is
evenly distributed over the first gastral tergite. Included here are the remaining eight species
noted above. Of them regalis is very conspicuous and not obviously close to any of the others. Of
the remainder the southern complex of piceus, decipiens and capensis may represent a single
species, and denticornis, striatifrons and tropicorum are closely related.

Messor angularis Santschi stat. n.
(Figs 25, 26)

Messor barbarus st. semirufus var. angularis Santschi, 1914a: 75 [unavailable name]; Santschi, 1928: 202
[galla var. angularis, first available use of name]. Syntype workers, KENYA: Naivasha, 1900 m, st. no. 14,
xii.191 1 (C. Alluaud & R. Jeannel) (NM, Basle) [examined].

MEDIUM TO LARGE WORKER. HW 2.00- > 2.75.

Anterior clypeal margin flattened to weakly and quite broadly indented medially. With the head in
full-face view the occipital margin indented medially, the indentation becoming more distinct with increased
size. In HW range 2.00-2.80 the maximum diameter of the eye is 0.42-0.52, about 0.18-0.21 x HW, and the
CI range is 104-113. Propodeum unarmed, rounded to right-angled where dorsum meets declivity and
sometimes with a reinforcing ridge or flange following the curve, especially in largest workers. Dorsum of
head with sculpture very reduced, sometimes without sculpture. Usually with a few very feeble low
longitudinal rugulae between the frontal lobes which may extend for a short distance behind them. On each
side of this median area, moving outwards towards the eyes and occipital corners, the head is unsculptured
except for a very feeble superficial reticular pattern and a few scattered faint punctulae. Pronotum and
mesonotum dorsally unsculptured to feebly densely punctulate, generally with some weak transverse
rugulae immediately behind the cervical shield. Rarely these are absent but in some they extend further back
on the pronotum than is usual. Propodeal dorsum transversely rugose, conspicuously more strongly
sculptured than the pronotum or mesonotum. First gastral tergite unsculptured and smooth, usually with a
faint superficial reticular pattern visible. Head dorsally with very reduced pilosity; apart from the strong
mouthpart hairs and those around the frontal lobes the dorsum with only 2-3 pairs, spanning the midline of
the head. With the head in full-face view the sides both in front of and behind the eyes, the occipital corners
and the occipital margin without projecting hairs except mid-occipitally where the posteriormost dorsal pair
may project on each side of the occipital impression. Psammophore strong, the J-shaped hairs very long and
conspicuous. Pronotum dorsally with 0-3 pairs of hairs, when present situated posteriorly, close to the
promesonotal junction. Mesonotum with 0-5 pairs of hairs. Some of this variation may be due to abrasion,
the mesonotal hairs in particular seem easily lost. Propodeum always hairless dorsally. Petiole with 0-1,
postpetiole with 0-2 pairs of hairs respectively. First gastral tergite hairless or at most with a sparse
transverse row at the extreme apex of the sclerite. Colour variable, usually with reddish head and alitrunk
and black gaster, but the alitrunk often with some black, the amount of which varies from sample to sample.
In extreme cases the entire body black but even here the head with a reddish tint showing through.

One of the four sub-Saharan African species which lacks hairs on most or all of the first gastral
tergite, angularis is at present known only from Kenya. The most closely related species is the
extremely widespread galla which also occurs in Kenya. The two are separated as follows.

angularis galla

Propodeal dorsum without hairs. Propodeal dorsum with one or more pairs of hairs.

Occipital margin on each side of the median Occipital margin on each side of the median

impression without projecting hairs impression with one or more pairs of projecting

hairs.

Ventral surface of hind femora without freely Ventral surface of hind femora with numerous

projecting hairs or at most with 1-2 close to the freely projecting hairs which usually occur over

trochanter. the length of the shaft but which are often

densest proximally.

AFROTROPICAL MYRMICINE ANT GENERA 345

Ventral surface of postpetiole in profile without an Ventral surface of postpetiole in profile with a

anterior prominence or at most with a feeble sharp dentiform or angular prominence

angle, the surface immediately behind this anteriorly, the surface immediately behind this

smoothly concave. irregular, not smoothly concave.

Median strip of head dorsally unsculptured or at Median strip of head dorsally usually

most with very feeble rugulae anteriorly. conspicuously rugulose, only very rarely

reduced.

MATERIAL EXAMINED

Kenya: Tana Riv. (J. L. Clark); Olikoriti (M. G. Lepage); Kajiado (J. Darlington); Bissel (J. Darlington);
Kajiado (G. Nyamasyo); Kajiado (W. Sands); Isiolo (E. S. Ross & R. E. Leech); Naivasha (Alluaud &

Messor capensis (Mayr)
(Fig. 31)

Atta capensis Mayr, 1862: 743. LECTOTYPE worker, SOUTH AFRICA: Cape of Good Hope, Novara Expd.

" D ". (NM, Vienna), here designated [examined].
Aphaenogaster pseudoaegyptiaca Emery, 1884: 384. Syntype workers, SOUTH AFRICA: Cape of Good Hope

(MCSN, Genoa) [examined]. Syn. n.
Messor barbarus subsp. capensis var. schencki Forel, 1910a: 15. Holotype worker, SOUTH WEST AFRICA:

Bethanien (Schenck) (not found in MHN, Geneva, presumed lost). [Unavailable name.]
Messor braunsi Forel, 19136: 138. Syntype workers, SOUTH AFRICA: Cape Prov., Willowmore (H. Brauns)

(MHN, Geneva) [examined]. Syn. n.
Messor donisthorpei Santschi, 1937: 51. Syntype workers, females, SOUTH WEST AFRICA: West of Maltahohe,

1500 m, 12.xii.1934 (K. Jordan) (BMNH; MCZ, Cambridge; USNM, Washington) [examined]. Syn. n.

MEDIUM TO LARGE WORKER, HW 2.35- > 3.40.

Anterior clypeal margin varying from shallowly convex to transverse, only very rarely with the faintest
vestige of a median indentation. With the head in fullface view the sides more or less straight and
approximately parallel, never evenly convex nor obviously diverging anteriorly. Occipital margin broadly
and shallowly concave to indented medially. In HW range 2.35-3.44 the maximum diameter of the eye is
0.40-0.58, about 0.15-0.19 x HW, and the CI range is 103-119. Propodeum in profile with the dorsum
rounding narrowly into the declivity in most cases; in some more broadly rounded and in a few right-angled,
but only rarely with dentiform prominences and here usually only in the largest workers. Usual sculpture of
entire dorsum of head of fine, densely packed parallel longitudinal rugulae, most commonly with fine
punctulation between them. Variation in the sculpture consists of a reduction, in density or intensity, or one
or both of these components. Sometimes the rugulae are more widely spaced and fainter than is usual, in
which case the punctulate ground-sculpture is much more obvious and may appear as the dominant
component in places. On the other hand the punctulate sculpture may fade out, leaving the rugulae sharply
defined; the rugulae may then also become less intense and leave the head only feebly sculptured. Dorsal
alitrunk usually rugose or rugulose everywhere but, as on the head, this sculpture may be reduced until it is
very faint or even absent. When distinctly present the direction of sculpture on the pronotum shows
variation. Commonly it is longitudinal but forms with the sculpture diagonal, transverse, irregular or
varying on different parts of the surface are fairly frequent. First gastral tergite unsculptured or at most with
a very faint superficial patterning. All dorsal surfaces of head and body with numerous conspicuous standing
hairs. Colour black to dark reddish brown, the head and alitrunk always the same colour, the gaster
sometimes darker.

The taxa capensis, decipiens and piceus, treated here as separate species, may in fact represent
only a single variable species. The differences invoked to distinguish the three are minor (see key)
and may eventually prove to be gradient.

Among the species in which the first gastral tergite is uniformly hairy the three taxa mentioned
above are characterized together by their relatively small eyes, lack of strong gastral sculpture,
relatively straight-sided head and short propodeum, and lack of a median prominence on the
posterior half of the clypeus.

346 B. BOLTON

MATERIAL EXAMINED

Botswana: Kuke Pan (Vernay-Lang); Gomodimo Pan (Vernay-Lang); Gomodimo (G. U. Son). South
Africa: Cape Prov., Willowmore (G. Arnold); Willowmore (H. Brauns); Grahamstown (W. L. Brown);
Grahamstown (F. Jacot-Guillarmod); Cape Town (E. Simon); Cape Town (J. C. Bridwell); Cape Town (R. E.
Turner); Addo (M. Samways); Balfour (E. S. Ross & R. E. Leech); Fish River Valley (G. Arnold); Fort
Beaufort (J. W. G.); Oudtshorn (B. Brunhuber); Port Elizabeth (B. Brunhuber). South West Africa: W. of
Malahohe (K. Jordan).

Messor cephalotes (Emery)

Stenamma (Messor) barbarum subsp. cephalotes Emery, 1895a: 179. Syntype workers, ETHIOPIA: Arussi
Galla, Ganale Gudda, 3.V.1893 (V. Bottego) (MCSN, Genoa; MHN, Geneva) [examined].

Messor cephalotes (Emery) Emery, 1908 : 443. [Raised to species.]

Messor plinii Santschi, 1912: 165. Syntype workers, KENYA: Nakuru, 1904 (C. Alluaud) (NM, Basle)
[examined]. Syn. n.

MEDIUM TO LARGE WORKER, HW 3.20- > 5.00.

Median portion of clypeus with anterior margin broadly but shallowly indented-concave. With the head
in full-face view the occipital margin more or less transverse, very shallowly impressed medially to virtually
straight, only very rarely evenly shallowly convex. Head broad and massive, very strongly transversely
convex between the eyes, CI 109-123 in HW range 3.28-5.52. Eyes fairly small, their maximum diameter
0.54-0.72, about 0.13^0.17 x HW within the above-stated HW range; the relatively smaller eyes occurring
in larger individuals. Psammophore generally more strongly developed in smaller than in larger workers,
the characteristic hooked or J-shaped hairs sparse or absent in very large workers. Propodeal dorsum
varying from rounding bluntly and evenly into the declivity to meeting the declivity in a sharp right-angle.
In either case a low reinforcing lip or flange may be present which follows the curve, but prominent blunt
teeth or lamellae are only very rarely known to develop. Dorsum of head blanketed everywhere with
extremely fine, very densely and tightly packed, parallel longitudinal costulae; the head with a silky
appearance under low magnification. The direction of the costulae is variable but usually they run straight
back from clypeus to occiput centrally on the dorsum, and tend to diverge towards the occipital corners
away from this central strip. In very large workers there is a tendency for the direction of the sculpture to be
less regular, and even loops or whorls may occur. Dorsal alitrunk densely rugulose everywhere, the
sculpture usually transverse but sometimes irregular on the propodeum. Sculpture on propodeal dorsum
generally coarser and more widely spaced than on pronotum, and always coarser on pronotum than on
dorsum of head. Petiole and postpetiole finely and densely sculptured with feeble rugulae, dense
puncturation, or a combination of both. Base of first gastral tergite extensively sculptured with exceedingly
fine close-packed scratch-like costulae, or sometimes with dense granular puncturation, or with a
combination of both. The extent of this sculpture is variable but always at least the basal third of the first
tergite is covered. Pilosity quite dense, all dorsal surfaces of head and body with standing hairs. With the
head in full-face view the sides in front of and behind the eyes, and the curved side portions of the occipital
corners, without projecting hairs; the occipital margin itself usually with conspicuous projecting hairs.
Colour red to reddish dark brown, often with the gaster somewhat darker than the head and alitrunk.

A very distinctive East African species, cephalotes is one of the only two known African forms in
which the gaster is strongly sculptured. The other, regalis, has much coarser sculpture, as noted in
the key, and also differs by having the propodeum always bidentate or bispinose, a feature only
very rarely developed in cephalotes. Beside this the anterior clypeal margin, always concave in
cephalotes, is shallowly convex and irregular in regalis, and the sides of the head, hairless in
cephalotes, have distinct standing hairs in regalis, at least behind the eyes.

MATERIAL EXAMINED

Ethiopia: Ganale Gudda (V. Bottego). Kenya: Nakuru, (E. Pinhey); Nakuru (T. J. Anderson); Nakuru (C.
Alluaud); Lake Ngunga (Allen & Brooks); Kericho (F. W. Dry); Athi Riv. (C. S. Betton); Olikoriti (M. G.
Lepage); Kajiado (J. Darlington); Kajiado (G. Nyamasyo). Tanzania: Dodoma (W. M. Mann); Umbulu
(W. M. Mann); Arusha (C. F. D.).

Messor coltingwoodi sp. n.

(Fig. 30)

HOLOTYPE WORKER, HW 2.56.

Anterior clypeal margin broadly but shallowly indented medially. With the head in full-face view the sides
more or less straight, slightly convergent anteriorly and rounding broadly and evenly into the occipital

AFROTROPICAL MYRMICINE ANT GENERA

347

margin behind. Occipital margin sharply indented medially. Maximum diameter of eye 0.52, about
0.20 x HW, and the CI 107. Propodeum armed with a pair of short but well-developed triangular spines
which are somewhat downcurved along their length. Dorsum of head sculptured everywhere. Mid-dorsal
strip of head longitudinally rugulose to level of posterior margins of eyes; behind this the rugulae rapidly
weakening. Everywhere dorsum of head finely and very densely reticulate-punctate, with superimposed very
feeble rugulae away from the more strongly sculptured median strip. Pronotum and mesonotum dorsally

31

32

Figs 30-34 30-32, Messor workers. Alitrunk of (30) collingwoodi, (31) capensis, (32) striatifrons. 33, 34,
Cataulacus workers. Profile of (33) centrums, (34) moloch.

348 B. BOLTON

transversely rugulose, the propodeum more strongly transversely rugose. Sides of pronotum less strongly
rugulose than the pleurae. First gastral tergite unsculptured except for the usual fine superficial reticular
patterning. Dorsum of head sparsely hairy. Discounting the strong pilosity on the mouthparts and around
the frontal lobes the dorsum with only a few pairs of hairs spanning the mid-dorsal strip. With the head in
full-face view the sides both in front of and behind the eyes lacking projecting hairs. Projecting hairs also
absent from occipital corners but a single hair projecting from the occipital margin on each side of the
median indentation. Psammophore strongly developed, the J-shaped hairs conspicuous. Dorsal alitrunk
without hairs on pronotum, with 4 pairs on mesonotum and one pair on the propodeum. Petiole with one
pair, postpetiole and first gastral tergite hairless. Colour uniform very dark blackish brown.

MEDIUM TO LARGE PARATYPE WORKERS, HW 2.16-2.72. As holotype but in some individuals the mid-dorsal
rugulae of the head more sharply defined. Variation in pilosity throughout the type-series shows the dorsal
head with 2-5 pairs, pronotum with 0-1 pair, occipital margin with 0-2 pairs, mesonotum with 4-6 pairs,
propodeum with 1-3 pairs, petiole with 0-3 pairs, postpetiole with 0-3 pairs of hairs. First gastral tergite
consistently hairless. Eyes fairly large, within the HW range given above the maximum eye diameter is
0.46-0.58, about 0.20-0.22 x HW. CI range is 103-1 10.

Holotype worker, Niger: Azanyares, iii.1979 (J. Newby) (BMNH).

Paratypes. 12 workers with same data as holotype (BMNH; MCZ, Cambridge; NM, Basle; MHN,
Geneva).
Non-paratypic material examined. Mali: Tessalit (P. Room).

Among the species with hairless or near hairless first gastral tergite collingwoodi is distinguished
by having propodeal hairs present, having an extensively sculptured head, and having persistent
propodeal spines. M. luebberti, which also has the head sculptured everywhere, is reddish in
colour and lacks propodeal hairs and spines. Also, the rugular cephalic sculpture is more
extensively developed than in collingwoodi. M. angularis also lacks propodeal hairs and spines
and has the head weakly or not sculptured. M. galla, which frequently develops propodeal lobes
or teeth and which also has propodeal hairs present, lacks the characteristic cephalic sculpture of
collingwoodi.

The closest relatives of collingwoodi are, however, not to be found among the other
sub-Saharan African species but among the members of the aegyptiacus-group, of which
collingwoodi seems to be the only Afrotropical species.

Messor decipiens Santschi stat. n.
(Fig. 28)

Stenamma (Messor) barbarum r. capense var. decipiens Forel, 1905: 177 [unavailable name]; Santschi, 1917:

94 [Messor capense st. decipiens, first available use of name]. Syntype workers, females, SOUTH AFRICA:

Natal (Wroughton) (MHN, Geneva) [examined].
Messor barbarus subsp. capensis var. proba Forel, 191 la: 266. Holotype worker, SOUTH AFRICA: Orange

Free State, Bothaville (H. Brauns) (MHN, Geneva) [examined]. [Unavailable name.]
Messor arcistriatus Santschi, 1928: 202. Holotype worker, SOUTH AFRICA: Natal (Wroughton) (NM, Basle)

[examined]. Syn. n.

MEDIUM TO LARGE WORKER, HW 2.64- > 4.20.

Answering to the description of capensis in most particulars. In the HW range quoted above the
maximum diameter of the eye is 0.44-0.66, about 0.14-0.18 x HW, and the CI range is 107-121, the largest
workers known for decipiens thus being somewhat larger and broader headed than those known for
capensis. Propodeum in profile with the dorsum usually meeting the declivity in a right-angle, which may
project into a broad but quite short lobe or tooth of variable shape and size; rarely the propodeum merely
narrowly rounded. In contrast the propodeum of capensis is generally rounded, only seldom with dentiform
prominences. Sculpture of head basically the same as in capensis but here the rugae tending to be more
sharply developed and more widely separated, although there is some variation. Spaces between the rugae
usually smooth, frequently glossy, much less commonly with traces of punctulate ground-sculpture. Head
usually obviously red, contrasting in colour with the alitrunk and gaster which are darker. In smaller
workers this distinction in colour is not nearly so obvious and at the lower limit of the size range considered
here (and smaller) the ant may be unicoloured.

AFROTROPICAL MYRMICINE ANT GENERA 349

Very closely related to capensis and piceus, decipiens is separated from the former only on the
weak characters mentioned above. It is even closer to the latter, being distinguished only by the
colour of the hairs as noted in the key, and the fact that piceus does not have the head distinctly
different in colour from the alitrunk in large workers. It seems very probable that more extensive
collecting of this complex will reveal that these forms represent but a single species.

MATERIAL EXAMINED

Zimbabwe: Bulawayo (G. Arnold). Botswana: Ghazi (J. Maurice). Lesotho: Mafeteng (R. Crawshay). South
Africa: Natal, Weenen (G. Arnold); Natal (Wroughton); Durban (G. Arnold); Drakensberg, Van Reenen
(R. E. Turner); Mkuzi Reserve (C. P. Peelers); no loc. (ex coll. F. Smith); Transvaal, Brakfontein (Lingnau);
Vryburg (G. Arnold); Shiluvane (Junod); Orange Free State, Bothaville (H. Brauns).

Messor denticornis Forel
(Fig. 29)

Messor denticornis Forel, 1910a: 14. Syntype workers, female, male, SOUTH WEST AFRICA: Liideritzbucht,

1903 (L. Schultze) MHN, Geneva; BMNH) [examined].
Messor denticornis var. parvidens Forel, 19100: 15. Syntype workers, SOUTH WEST AFRICA: Kubub (L.

Schultze) (MHN, Geneva) [examined]. Syn. n.
Messor denticornis var. brunni Forel, 19106: 444. Syntype workers, SOUTH WEST AFRICA: no loc. (Brunn);

and SOUTH AFRICA: Cape Prov., Steckstown (Wartmann) (MHN, Geneva) [examined]. Syn. n.

MEDIUM TO LARGE WORKER, HW 2.48- > 3.10.

Anterior clypeal margin usually evenly convex medially, only rarely with the faintest trace of a central
indentation. With the head in full-face view the sides more or less straight and diverging anteriorly, but
sometimes the sides more nearly parallel. Occipital margin broadly but shallowly concave, this feature
fading out in smaller workers where the margin is approximately transverse. In HW range 2.48-3.16 the
maximum diameter of the eye is 0.56-0.70, about 0.21-0.25 x HW, and the CI range is 100-106. Propodeum
in profile relatively long and low, resembling that of striatifrons (Fig. 32). Propodeal armament very
variable, the junction of dorsum and declivity being rounded, acutely angled or distinctly bidentate. These
variants are commonly seen in the same series and are in fact shown by the type-series of denticornis itself.
Basic sculpture of the head finely densely packed parallel longitudinal rugulae with punctulate
ground-sculpture between them. Frequently the sculpture much reduced, either by suppression of the
ground-sculpture so that the rugulae stand out from a smooth surface or by reduction of the rugulae in
number and intensity so that the head is mostly or wholly punctulate. In smaller individuals the surface may
be almost smooth. Dorsal alitrunk rugulose to rugose, the sculpture sometimes partially or totally effaced
from the pronotum. First gastral tergite smooth and shining or at most with faint superficial pattering. All
dorsal surfaces of head and body with numerous standing hairs. Colour mid-brown to black, sometimes
with the gaster darker than the head and alitrunk.

A distinctive species amongst those with uniformly distributed pilosity on the first gastral tergite,
denticornis is immediately isolated by its relatively large eyes. Only a few workers of tropicorum
approach even the lower end of its eye size range but in the latter species the clypeus has a
conspicuous posteromedian tumulus or welt and the propodeum is shorter and higher in profile
than is the case in denticornis.

MATERIAL EXAMINED

Botswana: Ghanzi (E. S. Ross & A. R. Stephen). South West Africa: Okaukuejo (E. S. Ross & R. E. Leech);
Spitzekopfe (E. S. Ross & K. Lorenzen); Ababis (R. W. Tucker); Berseba (L. O. Sordahl); Liideritzbucht (L.
Schultze); Kubub (L. Schultze). South Africa: Cape Prov., Oudtshorn (B. Brunhuber); Strydenburg (M.
Patterson); Steckstown (Wartmann).

Messor galla (Mayr)

Stenamma (Messor) barbarum subsp. caduca var. galla Emery, 1895a: 179 [unavailable name]; Mayr, 1904:
5 [Stenamma (Messor) barbarum var. galla, first available use of name]; Santschi, 1928: 201 [galla raised
to species]. Holotype worker, ETHIOPIA: Alto Duau, Boran Galla, v. 1893 (V. Bottego) (MCSN, Genoa)
[examined].

Messor barbarus subsp. semirufus var. rufa Forel, 1910c: 250. Syntype workers, ETHIOPIA: Nefassit (K.
Escherich) (MHN, Geneva) [examined]. [Unavailable name.]

350 B. BOLTON

Messor barbarus st. galla var. triempressa Santschi, 1917: 92. Syntype workers, CHAD: Baguirmi, Techeckna;

ETHIOPIA: no loc.; SENEGAL : Casamance (Clavaux). (NM, Basle) [examined]. [Unavailable name.]
Messor barbarus st. latinoda Santschi, 1917: 93, fig. 2. Syntype workers, 'EAST AFRICA': no loc.

(Reichensperger) (NM, Basle) [examined]. Syn. n.
Messor barbarus r. semirufus var. rufula Forel, 1918: 156. [Unnecessary replacement name for rufa Forel,

1910c: 250, above.] [Unavailable name.]
Messor barbarus subsp. galla var. armata Emery, 19226: 98. Syntype workers, GHANA: no loc. (MCSN,

Genoa) [examined]. [Unavailable name.]
Messor galla st. nobilis Santschi, 1928: 201. Syntype workers, female, ETHIOPIA: Bisa Tint, 1200m

(Reichensperger) (NM, Basle) [examined]. Syn. n.
Messor galla var. airensis Bernard, 1950: 286, Syntype workers, NIGER: Air Dist., Dabaga, 600 m; Mt

Baguezans, 1500 m; Agadez, 525 m (Chopard & Villiers). [Not found in MNHN, Paris, presumed lost.]

Syn. n.

MEDIUM TO LARGE WORKER, HW 2.40- > 3.70.

Median portion of clypeus with anterior margin broadly but shallowly concave to more or less entire.
With the head in full-face view the sides very shallowly convex to roughly straight, usually slightly
convergent in front of the eyes. Occipital margin broadly indented medially. In HW range 2.40-3.76 the
maximum diameter of the eye 0.44-0.68, about 0.17-0.20 x HW, and the CI range 102-114. Propodeum
showing great variation; frequently with the dorsum rounding into the declivity but sometimes with a pair
of broad teeth or lamellae. Between these two extremes is a range of intermediates including forms with a
narrow to broad rim or flange following the curve of the surface, forms with a small to large salient angle
and forms with the angle or flange projecting to various degrees. Dorsum of head smooth and shining, away
from the median strip sculptured only with very widely scattered small pits or a faint superficial patterning.
Median strip of head behind clypeus with longitudinal rugular sculpture which usually extends back at least
as far as the level of the posterior margins of the eyes, and often distinctly further back than this; only very
rarely is the rugular strip shorter. Intensity of rugulae on the median strip very variable and the width of the
strip not usually exceeding the width across the frontal lobes and often narrower, only rarely slightly wider.
Pronotum dorsally with weak transverse rugulae which may sometimes be very feeble or even partially
effaced. Mesonotum varying from almost smooth to faintly rugulose. Propodeal dorsum generally sharply
transversely rugose but in some samples the rugae diagonal, irregular or interrupted. First gastral tergite
unsculptured but often showing a faint superficial patterning. With the head in full-face view the sides
without projecting hairs, the occipital margin with 0-4 hairs on each side of the median impression.
Generally hairs are present occipitally, specimens with zero count are very few and may be the result of
abrasion. Dorsum of head sparsely hairy, the psammophore conspicuous ventrally. Parts of dorsal alitrunk
with pilosity as follows; pronotum with 0-4 pairs, mesonotum with 4-10 pairs, metanotal groove with 1-2
pairs at least in large workers, propodeum with 1-5 pairs. Petiole with 1-3, postpetiole with 3-6 pairs of
hairs. First gastral tergite without hairs or with a sparse transverse row at the extreme apex of the sclerite.
Ventral surfaces of hind femora usually with hairs all along the shaft but in some they are denser proximally
than distally. Colour reddish brown to blackish brown, usually with the gaster darker than the head and
alitrunk. In some samples the head slightly more reddish than the alitrunk.

Without doubt the commonest, most successful and most widely distributed Messor species in
the northern half of sub-Saharan Africa, galla ranges throughout the Sahelian zone across the
entire width of the continent. On the eastern side it is found as far south as Kenya, and in the west
it occurs coastally as well as in the drier northern parts of the West African states. Of the four
species in the region which lack dense gastral pilosity galla is separated from collingwoodi and
luebberti by the extensive cephalic sculpturing of the last two. The separation of galla from its
closest African relative, angularis of Kenya, is tabulated under that name. Some aspects of the
biology of galla have been investigated by Levieux & Diomande (1978) and Levieux (1979).

MATERIAL EXAMINED

Ethiopia: Addis Ababa, Entoto Hills (K. Guichard); Addis Ababa (V. O. De Mass/); Boran Galla (V.
Bottego); Lake Zwai, Sucsuci (J. 0. Cooper); Wachacha Ravine (H. Scott); Bisa Tint (Reichensperger); Mt
Monagasha (Cloudsley-Thompson); Gondar (Cloudsley-Thompson); Tisisat Falls (Cloudsley-Thompson);
Holetta; Dessie (E. S. Ross); Nefassit (K. Escherich); Barentu (Muller); Tessenei (Muller); Amba Derho
(Muller); Om Agar (Muller); Ghinda (K. Escherich); no loc. (G. McCreagh). Somali Republic: Alabla Balleh
(P. E. Glover). Kenya: Nakuru (N. A. Weber); Marsubit (Rift Valley Expd.); Tsavo East (J. Darlington);
Maralal (M. E. Irwin & E. S. Ross). Sudan: Kadugli (C. Sweeny); Khartoum (N. A. Weber); Khartoum (R.
Cottom); Khartoum (H. H. King); Kulme (H. Lynes); Lake Kellek (C. Sweeny); Dilling-El Obeid Rd. (C.

AFROTROPICAL MYRMICINE ANT GENERA 351

Sweeny); Sennar (B. Hocking); Imatong Mts (N. A. Weber); Equatoria (N. A. Weber). Niger: Niamey (P.
Room); Niamey (J. Levieux); Ayorou (P. Room); Assode (J. Levieux). Mali: Gao (B. Malkin); Anefis (P.
Room). Upper Volta : Ougadougou (P. Room); Banfora (Betbeder). Senegal: Dakar (W. L. Brown); Dakar (N.
L. H. Krauss); Casamance (Clavaux). Ivory Coast: Korhogo (R. Lucius); Ferkessedougou (J. Levieux).
Ghana: Lawra (W. Cook); Bolgatanga (P. Room); Tamale (Anipare); Tumu (P. Room); Navrongo (C. A.
Collingwood); Dawhwenya (D. Lesion); Dawhwenya (C. A. Collingwood); Nyankpala; Prampram (W.
Belfield); Achimota (W. Belfield); Nungua (W. Belfield); Accra (C. A. Buckman). Nigeria: Kalkala (F. D.
Golding); Illela (Lelean); Katsina (J. T. Medler); Zaria (A. S. Ahman); Maiduguri (E. R. Ross & K.
Lorenzen).

Messor incisus Stitz nomen dubium

Messor incisus Stitz, 1923: 149. Holotype female, SOUTH WEST AFRICA: Okosongomingo Farm, vii-viii.1912
(H. Thomsen) [not found in MNHU, Berlin, presumed lost].

Described from a single female which has since been lost, the identity of incisus cannot be
ascertained accurately at present. In his original description of incisus Stitz compares it to the
female of denticornis. So few females of denticornis are known that it is possible for incisus to fall
within the range of variation of that species. On the other hand incisus may be the female of
striatifrons or indeed be a separate species. Considerably more samples of Messor females will be
necessary before any attempt at placing incisus can be made.

Messor luebberti Forel stat. n.

Messor barbarus subsp. lubberti Forel, 1910a: 13. Syntype workers, SOUTH WEST AFRICA: Okahandja
(Peters), and no loc. (Lubbert) (MHN, Geneva) [examined].

MEDIUM TO LARGE WORKER, HW 2.00- > 3.00.

Anterior clypeal margin flattened to slightly indented medially. With the head in full-face view the sides
more or less straight, roughly parallel or weakly convergent anteriorly. Occipital margin distinctly indented
medially in large workers but the indentation becoming obliterated with reduced size. In HW range
2.00-3.12 the maximum diameter of the eyes 0.38-0.50, about 0.15-0.18 x HW, and the CI is 100-112. With
the propodeum in profile the dorsum rounding narrowly into the declivity to meeting the declivity in a
right-angle; propodeal armament never developed. Dorsum of head everywhere finely and densely
longitudinally rugulose, the rugulae approximately parallel and becoming finer away from the mid-dorsal
strip. Ground-sculpture of minute punctulation is present between the rugulae but this is less conspicuous in
some samples than in others. Pronotal dorsum weakly and faintly to quite strongly transversely rugulose,
but always with a fairly distinct punctulate component between the rugulae. Mesonotum smooth with only
vestigial traces of sculpture to irregularly granular, only rarely with a rugular component. Propodeal
dorsum transversely rugulose to rugose, with punctures between the rugulae. First gastral tergite
unsculptured except for the fine superficial reticular patterning which is usual in the genus. With the head in
full-face view the sides and occipital margin lacking projecting hairs. Projecting hairs very sparse to absent
on dorsum of head but present on mouthparts and between frontal lobes. Psammophore strongly
developed. On dorsal alitrunk the pronotum with 0-4 pairs of hairs, the mesonotum with 2-6 pairs; the
propodeum, petiole and postpetiole lacking hairs. First gastral tergite without hairs or at most with 2-3 at
the extreme apical margin of the sclerite. Colour usually red with a blackish gaster but in some the gaster the
same shade of red as the head and alitrunk. Shade of red of head and alitrunk varying from bright, almost
orange, to very dull.

This very distinctive species is extremely widespread in the southern half of the African continent.
It is immediately recognizable by its strongly sculptured head and very reduced pilosity. Of the
sparsely hairy species of Africa only collingwoodi from Mali and Niger has the head anywhere
near as strongly sculptured as luebberti, but in that species the propodeum has hairs and the
junction of propodeal dorsum and declivity is armed with a pair of short spines.

MATERIAL EXAMINED

Tanzania: Dodoma (A. Loveridge). Zimbabwe: Bulawayo (G. Arnold); Springvale (G. Arnold). Botswana:
Damara Pan (G. U. Son); Kuke Pan (G. U. Son); Gomodimo (G. U. Son); Xani Pan (A. Russell-Smith).
Angola: Cahama (E. S. Ross & R. E. Leech). South West Africa: Gemsbok Pan (G. U. Son); Okahandja
(Peters); no loc. (Lubbert); Windhoek (Ross & Stephen). South Africa: Transvaal, Shiluvane (Junod);
Malagieskraal (Lingnau); Pretoria; Pietersburg (E. S. Ross & R. E. Leech).

352 B. BOLTON

Messorpiceus Stitz

Messor piceus Stitz, 1923: 150. Syntype workers, female, SOUTH AFRICA: Transvaal (Ulrich) [not found in
MNHU, Berlin, presumed lost].

MEDIUM TO LARGE WORKER, HW 3.28- > 4.20.

Answering to the description of capensis, but differing mainly in the colour of the body pilosity which is
white to yellowish in capensis but very deep red-brown to blackish in piceus. Apart from this the anterior
clypeal margin is indented medially in piceus; the propodeum varies from narrowly rounded through
right-angled to broadly and bluntly dentate, and the largest known workers are larger than those of
capensis. In the HW range 3.28-4.20 the maximum diameter of the eye is 0.54-0.64, about 0.15-0.17 x HW,
and the CI range is 106-1 19. The maximum known for capensis is HW 3.44 but this may not be the largest
worker of the species, merely the largest available for study at present. Relative size of eye and CI fall within
the range of capensis.

Unfortunately the type-series of piceus appears to be lost, but three short series from Transvaal
match the original description tolerably well and show the dark pilosity noted by Stitz. I am
therefore applying the name piceus to these specimens and to two other short series, from Natal
and Botswana, noted under material examined.

M. piceus is a very closely related to capensis and decipiens; these three names may ultimately
prove to represent only a single species.

MATERIAL EXAMINED

Botswana: Gomodimo (Vernay-Lang). South Africa: Natal, Pietermaritzburg (Akerman); Transvaal,
Sabie; Kimberley (G. Arnold); Oliphants River, Grootdraai (H. Lang).

Messor regalis (Emery)

Cratomyrmex regalis Emery, 1891: 572, pi. 15, fig. 16. LECTOTYPE female, NIGERIA: Benue (Staudinger)

(MCSN, Genoa), here designated [examined].
Cratomyrmex regalis var. rubea Santschi, 1913: 308. Holotype worker, BENIN REPUBLIC: no loc. (Le Moult)

(NM, Basle) [examined]. Syn. n.
Cratomyrmex sculpturatus Stitz, 1916: 377, fig. 2. Syntype workers, CONGO: Fort Possel-Fort Crampel,

xi.1910 (Schubotz); and Chutes de la Nana, 'bei Fort Crampel', 7.xi.l910 (Haberer) (MNHU, Berlin)

[examined]. Syn. n.
Messor regalis (Emery) Emery, 1922a: 357.

MEDIUM TO LARGE WORKER, HW 3.00- > 4.50.

Median portion of clypeus with anterior margin shallowly convex to somewhat flattened, irregular
because of strong sculpture but not strongly impressed-concave. In HW range 3.00-4.40 the maximum
diameter of the eye is 0.48-0.70, about 0.16-0.17 x HW, and the CI range is 109-115. With the head in
full-face view the sides in front of the eyes more or less straight, roughly parallel or slightly convergent
anteriorly. Behind the eyes the sides rounding very broadly and evenly into the occipital margin ; the latter
usually shallowly indented medially. Propodeum armed with a pair of short triangular spines. Dorsum of
head densely sculptured everywhere with coarse parallel longitudinal rugulae. On the median strip behind
the frontal lobes the rugulae tend to run straight back on the head; on each side of this strip they diverge
towards the occipital corners. Pronotal dorsum coarsely sharply and irregularly rugose, frequently
reticulate-rugose in places and generally with a strip of strong transverse rugae immediately behind the
cervical shield. Remainder of dorsum and also sides of alitrunk strongly and generally sharply rugose
everywhere, the sculpture stronger than on the dorsum of the head. Tergal portions of petiole and
postpetiole very closely and coarsely irregularly rugose, the surfaces with a crumpled and wrinkled
appearance. First gastral tergite rugulose to sharply costulate basally, the sculpture extending at least over
the basal third of the sclerite and becoming finer posteriorly. All dorsal surfaces of head and body with
numerous standing hairs, pilosity also dense on legs. With the head in full-face view projecting hairs are
present on the sides behind the eyes, on the broad curve of the occipital corners and on the occipital margin
itself. One or two hairs usually also project from the sides in front of the eyes. Psammophore conspicuously
developed. Colour dull red to reddish brown, the gaster sometimes with an orange tint.

A species of West and Central Africa regalis is easily characterized by its blanketing coarse
rugose sculpture. No other species in the region has sculpture approaching that found in regalis.
This feature coupled with the dense pilosity and persistent propodeal spines renders the species

AFROTROPICAL MYRMICINE ANT GENERA 353

quickly recognizable. Only cephalotes and regalis have extensive sculpture on the first gastral
tergite; characters separating the two are given under cephalotes.

Some aspects of the biology of regalis have recently been investigated by Levieux & Diomande
( 1 978) and Levieux( 1979).

MATERIAL EXAMINED

Nigeria: K. State, N. Bussa (J. T. Medler); Mokwa (C. Longhurst); Olokemeji (Bridwell); Benue
(Staudinger). Benin Republic: no loc. (Le Moult). Congo: Fort Crampel (Schubotz).

Messor ruginodis Stitz stat. n., nomen dubium

Messor barbarus st. ruginodis Stitz, 1916: 374, fig. 1. Syntype workers, CONGO: Fort Crampel,
xi.l910-6.i.!91 1 (Schubotz) [not found in MNHU, Berlin, presumed lost].

Apart from the very distinctive regalis this is the only other species of Messor recorded from the
Congo. It is possible to decide from Stitz's description that ruginodis is related to capensis and its
allies, but further placement cannot be attempted without the types as the description alone is not
good enough. It must suffice for the present to state that, apart from regalis, no Messor species is
known to extend its range into the Congo, so ruginodis remains an enigma.

As the species, whatever it really is, is definitely not closely related to barbarus, I have raised it
to species-level here.

Messor striatifrons Stitz stat. n.

(Figs 27, 32)

Messor denticornis var. striatifrons Stitz, 1923: 149. Syntype workers, SOUTH WEST AFRICA: no loc. (Scheben)
(MNHU, Berlin) [examined].

MEDIUM TO LARGE WORKER, HW 2.84- > 3.75.

Anterior clypeal margin usually shallowly convex medially but sometimes a weak central indentation of
the margin is present. With the head in full-face view the sides convex. Generally the convexity is distinct
(Fig. 27) in larger workers but tends to be less marked in smaller individuals; infrequently the reverse is true
and medium sized workers show the convexity more strongly than larger specimens. Occipital margin
shallowly indented medially, the indentation best developed in large workers and slowly disappearing with
decrease in size. Within the HW range 2.84-3.76 the maximum diameter of the eye is 0.52-0.68, about
0.16-0.18 x HW, and the CI range is 104-114. Propodeum in profile relatively long and low (Fig. 32),
usually rounded at the junction of dorsum and declivity but quite frequently right-angled or projecting into
a broad short tooth which is really no more than a projection of the right-angle. Dorsum of head sculptured
with extremely fine dense longitudinal rugulae which in the strongest sculptured individuals are very close
packed. Spaces between the rugulae with fairly conspicuous ground-sculpture of fine punctures. In medium
sized workers, and quite frequently in maximum sized workers also, the sculpture on the dorsal head is
modified by a weakening of the rugular component and an intensification of the punctures, so that in some
the rugular component is supressed and the head appears reticulate-punctate everywhere or almost
everywhere. Dorsal alitrunk rugulose, the direction of the sculpture variable but usually stronger on the
propodeum than elsewhere. First gastral tergite unsculptured or at most with the faint superficial patterning
so commonly seen in this genus. All dorsal surfaces of head and body with numerous standing hairs; evenly
distributed hairs conspicuous on first gastral tergite. Colour medium to dark brown, commonly uniform but
often with the gaster darker, blackish brown.

A fairly distinctive member of the group of species centring on capensis, striatifrons is
characterized by its relatively long low propodeum and convex head sides. The shape of the head
is not duplicated in other African species but denticornis has a similarly proportioned
propodeum. However, in this last-named species the eyes are larger, with a range of
0.21-02.5 x HW.

MATERIAL EXAMINED

South Africa: Cape Prov., Victoria West (G. Arnold); Steinkop (G. Arnold); Springbok (E. S. Ross & R. E.
Leech); Picketberg (E. S. Ross & R. E. Leech); Citrusdal (E. S. Ross & R. E. Leech); Papendrop (E. S. Ross &
K. Lorenzen); Clanwilliam (£. 5. Ross & R. E. Leech). South West Africa: no loc. (Scheben).

354 B. BOLTON

Messor tropicorum Wheeler stat. n.

Messor barbarus subsp. capensis var. tropicorum Forel, 1910fo: 444 [unavailable name]; Wheeler, 1922: 805

{capensis var. tropicorum, first available use of name]. Syntype workers, ANGOLA: Mossamedes (Baum &

Van der Kellen) (MHN, Geneva) [examined].
Messor denticornis var. laevifrons Stitz, 1923: 148. Syntype workers, SOUTH WEST AFRICA: Usakos,

iv.-vi.1911; and Grootfontein, 7-ll.vi.1911 (W. Michaelsen} |(MNHU, Berlin) [examined]. Syn. n.
Messor braunsi var. nigriventris Stitz, 1923: 150. Syntype workers, SOUTH WEST AFRICA: Grootfontein,

7-1 l.vi.191 1 (W. Michaelsen) (MNHU, Berlin) [examined]. Syn. n.

MEDIUM TO LARGE WORKER, HW 3.00- > 3.80.

Anterior clypeal margin entire or at most with a feeble median indentation. With the head in full-face view
the sides approximately straight, more or less parallel or feebly diverging anteriorly. Occipital margin
usually broadly and shallowly concave but this becomes less apparent with decreased size. Centre of
posterior half of clypeus, between the frontal lobes, with a conspicuously raised tumulus or welt in large
workers, this feature decreasing in intensity with reduced size and not present in smaller workers. In the HW
range 3.00-3.84 the maximum diameter of the eye is 0.64-0.72, about 0.19-0.21 x HW, and the CI range is
102-1 1 1. Propodeum in profile relatively short and high, like that of capensis (Fig. 31). Propodeal dorsum
either rounding into declivity, or meeting it in a right-angle, or armed with a pair of short triangular teeth ;
variation occurs within series. Dorsum of head sculptured with narrow fine longitudinal rugulae. In
strongest sculptured individuals the rugulae are dense and conspicuous, but often they are much reduced or
partially to entirely effaced away from the central strip. Between the rugulae the ground-sculpture is of a fine
superficial punctulation, often completely effaced. Dorsal alitrunk rugulose to rugose, the sculpture
frequently weak on the pronotum or even absent in places. First gastral tergite unsculptured or at most with
a faint superficial reticular pattering. All dorsal surfaces of head and body with numerous conspicuous
standing hairs. Head and alitrunk reddish brown, the gaster darker.

Larger workers of tropicorum are quickly isolated by their possession of a strong prominent welt
or tumulus posteromedially on the clypeus, but this character fades with reduced worker size.
The eyes are quite large, approaching the lower limit of the range seen in denticornis, but in the
latter the propodeum is longer and lower in profile.

MATERIAL EXAMINED

South West Africa: Kabiras (R. W. E. Tucker); Usakos (W. Michaelsen), Grootfontein (W. Michaelsen).
Angola: Mossamedes (Baum & Van der Kellen).

CATAULACUS F. Smith

Cataulacus F. Smith, 1853: 225. Type-species: Cataulacus taprobanae F. Smith, 1853: 225, by subsequent
designation of Bingham, 1903: 120.

For diagnosis of genus, current synonymy and generic revision see Bolton (1974). For some time I
have been unhappy about the treatment which I gave to some species in the C. tenuis-group of
Africa (Bolton, 1974). It has become apparent, with the acquisition of more material and with
further experience of the group, that I was wrong to synonymize some of the names. The
opportunity to rectify these mistakes now presents itself and the changes from the previous
system are summarized below. Following this six new species are described and a revised key to
the Afrotropical species is provided which reflects these additions and changes, and which
includes also the two African species recently described by Snelling (1979). The new key only
deals with the Afrotropical fauna; it excludes the Malagasy species which were incorporated in
the former (1974) key. For identification of such species the reader is referred back to the earlier
study.

Key to species (workers)

1 Dorsal alitrunk without standing hairs of any description or at most with only 1-2 very short
hairs at the highest point of the pronotum. Generally hairs absent from alitrunk but rarely

sparse strongly appressed hairs may be present

Dorsal alitrunk with numerous standing hairs which are usually conspicuous. If the standing
hairs are very short they are more or less evenly distributed over the dorsum and are not
restricted to the highest point of the pronotum 10

AFROTROPICAL MYRMICINE ANT GENERA 355

2 Propodeum completely unarmed, without trace of spines or teeth. (Zaire) . . . inermis Santschi
Propodeum armed with a pair of spines or teeth .3

3 Dorsal alitrunk strongly sulcate throughout. Appressed hairs present on the dorsal alitrunk.

(Ghana) . adpressus Bolton

Dorsal alitrunk reticulate-punctate to reticulate, usually also with fine rugulae or a

rugoreticulum present ; never sulcate. Appressed hairs absent from dorsal alitrunk . . 4

4 Petiole dorsally strongly transversely rugose or sulcate everywhere 5

Petiole dorsally variously sculptured but never transversely strongly rugose or sulcate . . 7

5 First gastral sternite laterobasally with a longitudinal margination or carina which parallels the

laterobasal margination of the first tergite. Femora of hind legs not excessively
anteroposteriorly compressed ........... . 6

First gastral sternite laterobasally without a longitudinal margination or carina which parallels
the laterobasal margination of the first tergite. Femora of hind legs strikingly
anteroposteriorly compressed, narrow and very deep. (Sierra Leone, Cameroun, Equatorial
Guina, Congo, Zaire, Uganda) kohli Mayr

6 Sides of head behind eyes irregular, either denticulate, crenulate or otherwise jagged. Relatively

broader-headed species, CI > 125, the head strongly broadened behind the eyes. Laterally
projecting hairs on sides of head behind eyes long and conspicuous. (Sierra Leone, Ghana,

Nigeria, Cameroun, Uganda, Zaire, Zambia) huberi Andre

Sides of head behind eyes regular, smooth, neither denticulate nor crenulate. Relatively
narrower-headed species, CI 120 or less, the head not strongly broadened behind the eyes.
Laterally projecting hairs on sides of head behind eyes minute and inconspicuous or absent.
(Ghana, Nigeria, Cameroun, Uganda, Congo, Zaire) egenus Santschi

7 Petiole and postpetiole in dorsal view strongly longitudinally sulcate. Postpetiole dorsally

divided into two projecting lobes by a deep median longitudinal cleft. (Cameroun, Zaire)

lobatus Mayr

Petiole and postpetiole in dorsal view not strongly longitudinally sulcate. Postpetiole dorsally
not divided into two projecting lobes by a deep median longitudinal cleft .... 8

8 Lateral pronotal margination with 2 teeth. Dorsal and lateral surfaces of petiole and postpetiole

with numerous tubercles and small angular prominences, presenting a multi-peaked and

irregular surface. (Cameroun, Congo, Zaire, Kenya) pullus Santschi

Lateral pronotal margination with 0-1 teeth. Dorsal and lateral surfaces of petiole and

postpetiole not equipped with tubercles and small angular prominences .... 9

9 With the head in full-face view the lateral margins behind the eyes without a row of short

projecting hairs. Lateral pronotal margination without teeth. (Ghana, Cameroun, Guinea,

Zaire) tar dux Santschi

With the head in full-face view the lateral margins behind the eyes with a row of short projecting
hairs. Lateral pronotal margination with a single tooth on each side, close to the anterior
pronotal corner. (Zaire) theobromicolus Santschi

10 Petiole and postpetiole strongly transverse, much flattened dorsoventrally and without nodes,

both very broadly thickly V-shaped in dorsal view. Propodeum armed only with a pair of
small teeth or tubercles which are inconspicuous. (Sierra Leone, Liberia, Ghana, Nigeria,

Cameroun, Zaire) . • mocquerysl Andre

Petiole and postpetiole nodiform, not strongly transverse nor flattened, not broadly V-shaped

in dorsal view. Propodeal spines well developed and conspicuous .... . 11

1 1 Hairs on clypeus and usually also on remainder of cephalic dorsum bizarre, strongly clavate or

stalked-suborbicular. In most the apex of each hair is very strongly swollen whilst the stem is
narrow; sometimes the stem may be short or very short . . . . . . . 12

Hairs on clypeus and remainder of cephalic dorsum simple, usually stout cylindrical and blunt
but sometimes very short and stubble-like, sometimes elongate and fine and occasionally
gradually increased in thickness from base to apex, but not strongly clavate or
stalked-suborbicular ............ 20

1 2 With the alitrunk in dorsal view the pronotal margin on each side without an unbroken series of

denticles which project laterally between the pronotal corner and the site of the promesonotal

junction . 13

With the alitrunk in dorsal view the pronotal margin on each side with an unbroken series of
denticles which project laterally between the pronotal corner and the site of the promesortotal
junction ................ 15

13 First gastral tergite regularly longitudinally sulcate throughout. (Cameroun) . . jacksoni (p. 360)

356 B. BOLTON

First gastral tergite reticulate-punctate or with tine rugulae overlying reticulate-punctate

ground-sculpture, never longitudinally sulcate 14

14 Propodeal dorsum longitudinally rugulose. (Nigeria, Cameroun) . . . . vorticus Bolton
Propodeal dorsum transversely rugose. (Nigeria) ....... boltoni Snelling

15 Bizarre hairs on dorsum of head behind clypeus with a very short basal stem, appearing

stud-like, the swollen apices set very close to the cephalic surface 16

Bizarre hairs on dorsum of head behind clypeus with an elongate basal stem, never short and
stud-like, the swollen apices conspicuously raised well clear of the cephalic surface . . . 17

16 Larger species, HW 0.80 or more. (Tanzania, Zimbabwe, Angola, South Africa)

brevisetosus Forel
Smaller species, HW < 0.80. (Ivory Coast, Ghana, Cameroun, Uganda, Kenya, Tanzania,

Angola) jeannett(p. 358)

17 Dorsal alitrunk with weak rugulose sculpture and a blanketing dense reticulate-punctate

ground-sculpture which is very conspicuous between the rugulae, the surface matt and dull . 1 8
Dorsal alitrunk with strong dense rugose sculpture the spaces between which are unsculptured

or at most contain some feeble superficial ground-sculpture, the surface glossy . . . 19

18 Denticles on lateral pronotal margins minute and inconspicuous in dorsal view, much smaller

than the tooth at the pronotal corner. (Cameroun) satrap(p. 363)

Denticles on lateral pronotal margins large and conspicuous, at least as large as the tooth at the
pronotal corner, sometimes larger. (Ghana, Nigeria, Cameroun, Zaire) . . . lujae(p. 358)

19 Smaller species, HW 0.80 or less. Body hairs relatively short (Fig. 34). Basal quarter of first

gastral tergite without strong rugulae, either punctate or with feeble rugulae caused by
alignment of punctures. Propodeal spines in profile evenly feebly curved. (Ghana, Nigeria)

moloch(p. 361)

Larger species, HW > 0.90. Body hairs relatively long (Fig. 33). Basal quarter of first gastral
tergite with strong longitudinal rugulae which are independent of the underlying
puncturation. Propodeal spines in profile with basal third elevated and apical two-thirds
recurved. (Cameroun) centrurus (p. 359)

20 Erect hairs on dorsal surfaces of head, alitrunk and gaster abundant, dense, very long narrow

and fine, curved or even sinuate, the entire ant with a softly pilose appearance rather than the

bristly appearance usually associated with this genus 21

Erect hairs on dorsal surfaces of head, alitrunk and gaster relatively sparse, short broad and
blunt, coarse and usually straight, the entire ant with a bristly or stubbly appearance . . 22

21 Propodeal dorsum longitudinally rugulose or rugose. Larger species, HL > 0.90, HW > 0.85.

(Ghana, Cameroun, Angola) elongatus Santschi

Propodeal dorsum transversely rugulose. Smaller species, HL < 0.90, HW < 0.85. (Zaire)

pilosus Santschi

22 Head relatively broad or very broad, the eyes small, CI > 112, OI < 30. In dorsal view the

posterolateral portion of the pronotal margin produced into a large spine or triangular
prominence. Propodeal spines long and very strong, not dorsoventrally flattened ... 23
Head relatively narrow and eyes larger, CI 1 10 or less, OI > 32. In dorsal view the
posterolateral portion of the pronotal margin usually armed with a short tooth or a denticle.
When a short tooth is present in this position it is usually comparable in size with others on
the pronotal margin. Propodeal spines usually dorsoventrally flattened, only rarely otherwise
. . . . 24

23 Sculpture of dorsal alitrunk a very distinct rugoreticulum with strongly reticulate-punctate

interspaces. Lateral margins of mesonotum usually with one or more denticles. (Liberia,
Ghana, Cameroun, Equatorial Guina, Gabon, Congo, Zaire). . . . erinaceus Stitz
Sculpture of dorsal alitrunk variable in intensity but consisting essentially of a longitudinal
rugation or sulcation which may be irregular or sinuate. Lateral margins of mesonotum
usually without denticles. (Ivory Coast, Liberia, Ghana, Nigeria, Cameroun, Equatorial
Guinae, Zaire, Uganda) guineensis F. Smith

24 Posterior one-quarter of first gastral tergite coarsely longitudinally sulcate, rugose or striate,

this sculpture always very distinct and usually extending to the apex of the tergite ... 25
Posterior one-quarter of first gastral tergite reticulate-punctate or finely superficially sculptured
and shining; a few fine scattered longitudinal rugulae formed by the fusion of the margins of
aligned punctures may sometimes be present 27

25 Smaller species, HW < 0.90, with relatively large eyes, OI 50 or more. (Zaire, Kenya)

striativentris Santschi

AFROTROPICAL MYRMICINE ANT GENERA 357

Larger species, HW > 0.95, with relatively smaller eyes, OI in range 34-48 .... 26

26 Dorsal surfaces of head and alitrunk with numerous conspicuous relatively long stout hairs.

Eyes slightly larger, OI range 43-48. (Kenya, Mozambique, South Africa) . . wissmanni Forel
Dorsal surfaces of head and alitrunk with relatively few inconspicuous very short stubbly hairs.
Eyes slightly smaller, OI range 34—40. (Ethiopia, Somali Republic, Kenya, Tanzania, Zambia,
Malawi, Zimbabwe, Mozambique, Angola, South West Africa, South Africa)

intrudens (F. Smith) (part)

27 Occiput with a distinct deeply incised transverse groove above the foramen. Below this the

remaining strip of the occiput juts out as a shield over the dorsal rim of the foramen itself.

(Uganda) impressus Bolton

Occiput without a deeply incised transverse groove above the foramen 28

28 Subpetiolar process complex, anteroventrally with a prominent broadly rounded angle and

posteroventrally with an extended heel or spur; the surface between these two usually
concave. Postpetiole with a strongly developed simple long digitiform ventral process . . 29
Either the subpetiolar process simple, a rectangular or subrectangular lobe without the above
configuration or with a feebly prominent acute angle or small tooth posteroventrally; if the
latter then the postpetiole with a short blunt or short tooth-like ventral process ... 35

29 Eyes relatively small, OI < 50 30

Eyes relatively large, OI 50 or more 32

30 Propodeal spines long, 0.40 or more in profile (in HW range 1.10-1.26), strongly divergent and

markedly elevated; in profile the spines distinctly longer than the maximum length of the

petiole. (Cameroun, Zaire) greggi Bolton

Propodeal spines short, < 0.25 in profile (in HW range 0.90-1.04), not strongly divergent nor
markedly elevated; in profile the spines distinctly shorter than the maximum length of the
petiole 31

31 Stout hairs on cephalic dorsum extremely dense, appearing as a bristly pelt in profile. A line

across the dorsum at the midlength of the eyes with many more than 10 hairs. Hairs on
dorsum of head more or less cylindrical, not spatulate; the hairs truncated apically, their sides

more or less straight and parallel. (Zaire) cestus (p. 360)

Stout hairs on cephalic dorsum sparse, not giving the appearance of a bristly pelt in profile. A
line across the dorsum at the midlength of the eyes with at most 10 hairs. Hairs on dorsum of
head conspicuously spatulate, broadly convex apically, their sides shallowly convex and
convergent basally. (Kenya) kenyensis(p. 358)

32 Most or all of stout hairs' on clypeus and dorsum of head increasing markedly in thickness from

base to apex, frequently 2-3 times broader at apex than at base. (Sierra Leone, Ghana,

Cameroun, Chad, Zaire) pygmaeus Andre

Most or all of stout hairs on clypeus and dorsum of head cylindrical or nearly so, not increasing
markedly in thickness from base to apex ; in some cases the hairs may broaden approximately
to their midlength and then continue at that width to their apices 33

33 Mesonotal and propodeal dorsa with very fine superficial low irregular weak wandering

rugulae, feeble or faded out in places but never evenly spaced nor regularly longitudinal.
Spaces between these fine rugulae densely strongly reticulate-puntate and dull. (Ghana,

Congo, Zaire) weissi (p. 358)

Mesonotal and propodeal dorsa with conspicuous strong broad longitudinal rugae which may
be parallel but which are never faded out in places. Spaces between the rugae weakly
superficially sculptured or unsculptured, the surfaces shining 34

34 Entire body exceptionally highly polished and very shiny. Longitudinal rugae on posterior half

of mesonotum and on propodeum very broad, subsulcate and parallel, without anastomoses

on the prodpodeum. (Nigeria) taylori(p. 364)

Dully shining, not obviously highly polished. Longitudinal rugae on posterior half of
mesonotum and on propodeum not subsulcate, not parallel, tending instead to diverge and
converge slightly along their lengths or to be weakly wavy; on the propodeum with
anastomoses. (Benin Republic) diffidlis Santschi

35 Hairs on dorsum of head exceptionally short, forming only a minute stubble on the surface.

Dorsum of head usually meeting occipital surface in a marked angle or edge, the one not

rounding evenly into the other 36

Hairs on dorsum of head conspicuous and quite dense, not represented only by a minute

stubble on the surface. Dorsum of head rounding into occipital surface 37

358 B. BOLTON

36 Tooth on mesokatepisternum large, long and acute, projecting anteriorly and usually clearly

visible in dorsal view, projecting beyond the margins of the mesonotum. (South Africa)

mi cans Mayr

Tooth on mesokatepisternum small and short, usually a mere denticle or acute angle,
sometimes not even as strong as this ; not visible in dorsal view. (Ethiopia, Somali Republic,
Kenya, Tanzania, Malawi, Zimbabwe, Mozambique, Angola, South West Africa, South
Africa) intrudens (F. Smith)

37 Larger species, HW > 1.10, PW > 0.90. (Zaire) bequaerti Forel

Smaller species, HW< 1.10, PW< 0.90 rife* . . 38

38 Pronotum laterally with a number of irregular rounded tuberculiform projections, without a

regular series of denticles although some of the projections appear to consist of 2 or more

denticles fused together. (South Africa) fricatidorsus Santschi

Pronotum laterally with a more or less regularly spaced series of denticles • 39

39 Dorsal surfaces of mesonotum and propodeum extremely finely and very densely more or less

evenly longitudinally rugulose, the rugulae so close together that the spaces between them are

wide enough for only 1-2 rows of punctures. (Cameroun) mckeyi Snelling

Dorsal surfaces of mesonotum and propodeum coarsely rugose, the rugae predominantly
longitudinal but with some strong cross-meshes, breaks or irregularities. The rugae widely
spaced so that the spaces between most of them accomodate many more than 2 rows of
punctures. (Ghana, Nigeria, Cameroun, Zaire, Sudan, Uganda, Tanzania, South Africa)

traegaordhi(p. 358)

Cataulacus lujae Forel sp. rev.

Cataulacus lujae Forel, 191 Ib: 311. Syntype workers, ZAIRE: Kasai, Kondue (Luja) (MHN, Geneva)
[examined]. [Wrongly synonymized with brevisetosus Forel by Bolton, 1974: 31.]

C. lujae var. gilviventris Forel should be included as a synonym oflujae, not of brevisetosus.

Cataulacus jeanneli Santschi sp. rev.

Cataulacus jeanneli Santschi, 1914a: 108, fig. 16. Holotype worker, KENYA: Gazi, 20 km S. of Mombasa, st.
no. 6, xi. 1911 (C. Alluaud & R. Jeannel) (NM, Basle) [examined]. [Wrongly synonymized with
brevisetosus by Bolton, 1974: 31.]

The names pygmaeus st. degener Santschi and janneli [sic] var. loveridgei Santschi should be
included in the synonymy of jeanneli, not of brevisetosus. The types of loveridgei still have not
been found; the holotype of brevisetosus has now been located in MHN, Geneva.

Cataulacus weissi Santschi

Cataulacus weissi Santschi, 1913: 310. Holotype worker, CONGO: Brazzaville, 1907 (A. Weiss) (NM, Basle)

[examined].
Cataulacus jeanneli var. aethiops Santschi, 1924: 220. Syntype workers, ZAIRE: Kidada-Kitobola,

14-25.ii.1922 (H. Schouteden) (MRAC, Tervuren) [examined]. Syn. n.

Cataulacus kenyensis Santschi stat. n.

Cataulacus jeanneli st. kenyensis Santschi, 1935: 272, figs 6a-c. Syntype workers, KENYA: Nairobi, st. 2,
1660 m, 1932-33 (C. Arambourg, P. Chappuis & R. Jeannel) (NM, Basle) [examined]. [Wrongly
synonymized with weissi by Bolton, 1974: 39.]

Cataulacus traegaordhi Santschi sp. rev.

Cataulacus traegaordhi Santschi, 19146: 24, fig. 3. Syntype workers, female, male, SOUTH AFRICA: Natal,
Zululand, Dukudu, 27.vii.1905 (/. Tragardh) (NM, Basle) [examined]. [Wrongly synonymized with
pygmaeus Andre by Bolton, 1974: 48.]

AFROTROPICAL MYRMICINE ANT GENERA 359

Of those names formerly included as synonyms under pygmaeus, the forms C. tragardhi [sic] var.
ugandensis Santschi, C. marleyi Forel (types in MHN, Geneva, not previously seen), and C.
pygmaeus subsp. suddensis Weber should now be included in the synonymy of traegaordhi, not of
pygmaeus.

Cataulacus centrums sp. n.

(Fig. 33)

HOLOTYPE WORKER. TL 3.9, HL 1.00, HW 0.92, CI 92, EL 0.47, OI 51, SL 0.48, SI 52, PW 0.70, AL 1.10.

With the head in full-face view the lateral margins of the head behind the eyes denticulate, terminating
posteriorly in a short tooth at the occipital corner. Occipital crest absent, the dorsum of the head rounding
evenly but narrowly into the occipital surface; the occipital margin itself unarmed except for a small tooth
situated close to the tooth at the corner. Eyes relatively large, OI > 50. Alitrunk with promesonotum both
longitudinally and transversely convex. In profile the highest point at about the midlength of the pronotum,
the remainder sloping evenly downwards posteriorly to the base of the propodeal spines. Anterior strongly
curved portion of pronotal dorsum with a number of minute peaks or tubercles from which hairs arise; such
peaks absent elsewhere on alitrunk. Tooth at base of mesokatepisternum developed. Propodeal spines in
profile with the basal third elevated at an angle of about 45°, the apical two-thirds back-curved. Metapleural
lobes low and rounded. With the alitrunk in dorsal view the pronotal corners denticulate and the lateral
margins of the pronotum armed with a series of 6-7 regularly spaced triangular denticles. Lateral margins of
mesonotum with a pair of small denticles whose bases are fused, situated at approximately the midlength.
Following the metanotal identation of the margin the sides of the propodeum are equipped with 2-3 small
tubercles. Propodeal spines in dorsal view broad and evenly divergent. Petiole in profile rising to a sharp
peak dorsally, behind which the surface slopes evenly downwards to the postpetiolar junction. Subpetiolar
process with a bluntly rounded anterior lobe and a weakly developed posteroventral tooth. Postpetiole in
profile with its dorsal and posterior surfaces tuberculate and its ventral process simple, short digitiform.
First gastral tergite not marginate laterally, conspicuously longer than broad. Dorsum of head finely and
evenly reticulate-rugulose, the recticular meshes of irregular size and the rugulae low and rounded.
Ground-sculpture in the meshes reduced to an inconspicuous vestigial superficial shagreening, without
punctulae. Pronotal dorsum similarly but somewhat more strongly sculptured, the reticulum breaking down
on the mesonotum so that the longitudinal component predominates and the cross-meshes are reduced or
incomplete. Propodeal dorsum more strongly and predominantly longitudinally rugose, irregular centrally.
Transverse rugae are present between the bases of the propodeal spines. Ground-sculpture of alitrunk
mostly as head but the mesonotum with some minute and virtually effaced punctulae. Petiole in dorsal view
longitudinally rugose, the sculpture converging posteriorly. Postpetiole dorsum irregularly rugulose. First
gastral tergite blanketed by fine dense reticulate-punctate sculpture, the basal quarter also with widely
spaced fine longitudinal costulae. Behind this level the tergite with scattered short longitudinal rugulae
which are very fine and irregular and formed by the alignment of the margins of adjacent punctures. First
gastral sternite reticulate-punctate. Sides of pronotum obliquely sulcate, the mesopleuron transversely
sulcate and the sides of the propodeum more or less vertically so behind the level of the spiracle. Sides of
petiole and postpetiole longitudinally sulcate-rugose. Discounting the long simple hairs which arise round
the eyes the entire dorsum of the head thickly clothed with stalked-suborbicular hairs, the stems of the hairs
long and fine and holding the suborbicular distal portions well clear of the surface of the head. Occipital
surface with a number of elongate narrowly clavate hairs. All remaining dorsal surfaces of body densely
clothed with moderately long stout cylindrical simple hairs which are truncated apically; those on the
alitrunk and petiole straight, those on the postpetiole and first gastral tergite weakly back-curved. Colour
uniform black, glossy ; the scapes, tibiae and tarsal segments dull yellow.

Holotype worker, Cameroun: Nkoemvon, 1979 (D. Jackson) (BMNH).

As indicated by the stalked-suborbicular cephalic hairs centrums belongs to the complex of
species centring on brevisetosus, and is most closely related to the smaller moloch. In the latter
species the simple pilosity of the alitrunk and gaster is very short and stubble-like, whereas in
centrums it is long and conspicuous (Figs 33, 34). The specialized cephalic hairs of moloch are
sparser than in centrums, have the basal stems of the hairs shorter and the apices less strongly
expanded. With the head in profile the specialized hairs immediately in front of the eye have the
basal stem longer than the swollen apex in centrums, shorter than the swollen apex in moloch. In
profile the propodeal spines of centrums have the basal third elevated and the apical two-thirds

360 B. BOLTON

recurved, a feature not seen in moloch where the spines are exceedingly feebly but evenly curved
along their length. Finally, the shape of the subpetiolar process differs in the two species, that of
moloch having the posteroventral angle more salient and the ventral surface more concave than
in centrums.

Cataulacus cestus sp. n.

HOLOTYPE WORKER. TL 4.0, HL 1.00, HW 0.99, CI 99, EL, 0.45, OI 45, SL 0.48, SI 48, PW 0.76, AL 1.10.

Sides of head behind eyes denticulate, terminating in a larger denticle at the occipital corner. Occipital
crest absent but the occipital surface shallowly concave above the foramen and meeting the dorsum in an
angle, the two surfaces not evenly rounded together. Occipital margin unarmed except for a denticle or short
tooth close to the one at the corner. Eyes relatively small, OI < 50. With the alitrunk in profile the dorsum
evenly shallowly convex between the more steeply sloped anterior portion of the pronotum and the base of
the propodeal spines. Pronotal and propodeal surfaces beset with small peaks or tubercles in profile, the
mesonotal dorsum also having such peaks but they are here more scattered and much lower, having the
appearance of minute irregularities in the outline. Mesokatepisternal tooth small. Metapleural lobes
rounded. Propodeal spines in profile short, more or less straight, only very slightly elevated. Alitrunk in
dorsal view with the pronotal corners denticulate, the lateral marginations of the pronotum behind the
corners with 6-7 sharp triangular denticles projecting laterally. Sides of mesonotum with 1-2 small denticles
and sides of propodeum also with 1-2, occurring on the convexity over the spiracle. Propodeal spines short
and broad, widely divergent. Petiole node in profile rising to an acute peak dorsally. The subpetiolar process
with a rounded and slightly prominent anteroventral lobe and a triangular projecting posteroventral tooth
or heel; the ventral surface between the two angles feebly concave. Postpetiole in profile high, its dorsal
surface with a number of conspicuous peaks or tubercles and its ventral process short-digitiform. Dorsum of
head irregularly reticulate-rugose, the meshes of varying size and the rugae low and rounded. Many of the
reticular meshes incomplete or with their walls broken. Ground-sculpture within the meshes a very fine
superficial shagreening or granular roughening of the surface, not reticulate-punctate. Dorsal alitrunk
irregularly reticulate-rugose everywhere, many of the rugular meshes incomplete or broken and very
irregular in shape. Ground-sculpture finely reticulate-punctate to densely shagreened. Petiole node in dorsal
view strongly longitudinally rugose, the rugae converging posteriorly. Postpetiole irregularly rugulose and
finely densely punctulate. First gastral tergite coarsely and densely reticulate-punctate everywhere, the
whole surface also loosely covered with anastomosing fine irregular superficial rugulae which are strongest
basally and fade out apically on the sclerite. First gastral sternite similarly sculptured. Entire dorsum of head
covered with a dense pelt of short straight erect bristly blunt hairs which are cylindrical to subcylindrical in
shape. All remaining dorsal surfaces of body with similar dense bristly pilosity. Colour uniform black; the
scapes, tibiae and tarsi dull yellow.

PARATYPE WORKERS. TL 4.0-^.1, HL 0.98-1.02, HW 0.98-1.02, CI 98-100, EL 0.45-0.48, OI 46-^7, SL
0.48-0.50, SI 49-51, PW 0.76-0.86, AL 1.08-1.16 (4 measured).

As holotype but in some the gastral rugulae are less strongly developed and in one the gastral rugulae are
effaced. The ventral surface of the subpetiolar process may be more strongly concave than is the case with
the holotype.

Holotype worker, Zaire (B. Congo on data label): Ituri For., Beni-Irumu, ii.1948, no. 2122 (N. A. Weber)
(MCZ, Cambridge).

Paratypes. 1 worker with same data as holotype; 1 worker with same data as holotype but no. 2120; 2
workers with same data as holotype but no. 21 19. (MCZ, Cambridge; BMNH).

Cataulacus jacksoni sp. n.

HOLOTYPE WORKER. TL 3.5, HL 0.98, HW 0.94, CI 96, EL 0.46, OI 50, SL 0.49, SI 52, PW 0.68, AL 0.98
(cephalic measurements approximate as head crushed).

With the head in full-face view the sides behind the eyes minutely denticulate. Occipital crest absent, the
dorsum rounding into the occipital margin. Head of holotype crushed behind level of eyes and the surface
fractured; the fracture also running forward on the head along the inner margin of the right eye to the
clypeus. With the alitrunk in profile the dorsal outline rising steeply to about the midlength of the
pronotum. Behind this the remainder of the dorsum evenly shallowly convex to the bases of the propodeal
spines, the outline not interrupted by superficial peaks or tubercles. Mesokatepisternal tooth small and
broadly triangular. Propodeal spines in profile strongly downcurved along their length. Metapleural lobes

AFROTROPICAL MYRMICINE ANT GENERA 361

very small. With the alitrunk in dorsal view the pronotal corners angular, the angle slightly projecting. Sides
of pronotum behind this not marginate, without a regular series of laterally projecting denticles. Instead the
sides with only a blunt tubercle at the point of junction of the pronotum and mesonotum and with one or
two minute irregularities, too low, small and blunt to be called tubercles or denticles, situated behind the
corner. Sides of mesonotum and propodeum unarmed and immarginate, the latter with a low salient welt at
the site of the spiracle. Propodeal spines in dorsal view curved, bowed outwards along their length. Petiole
in profile blunt above, not rising to a sharp peak. Subpetiolar process with the anteroventral angle rounded,
the posteroventral angle acute and slightly projecting. Postpetiole in profile very high and narrow, with a
flat anterior face and a long simple ventral process. In dorsal view the postpetiole with the sides converging
dorsally so that the node narrows from base to apex. Dorsum of head to level of posterior margins of eyes
finely longitudinally rugose, behind this level the head with very heavy broad strong sulci. Ventral surface of
head longitudinally sulcate. Dorsal alitrunk regularly strongly longitudinally sulcate except for the area
between the bases of the propodeal spines where the sulci are arched-transverse. Propodeal declivity
transversely sulcate. Coxae, femora and tibiae of legs all longitudinally sulcate. Anterior face of petiole node
transversely sulcate, the dorsum with U-shaped sulci. Upper half of anterior face of postpetiole vertically
sulcate. Sides of alitrunk diagonally sulcate from anteroventral to posterodorsal on each sclerite except on
the mesokatepisternum where they run from posteroventral to anterodorsal. First gastral tergite and first
sternite covered with strong parallel longitudinal sulci throughout. Dorsum of head with abundant
stalked-suborbicular hairs which have slender basal stems. Remainder of dorsal surfaces of body with sparse
fine curved hairs which are very feebly clavate apically. Colour uniform black but scapes, anterior tibiae and
tarsi, and tarsi of middle and hind legs dull yellow.

Holotype worker, Cameroun: Nkoemvon, 1980 (D. Jackson) (BMNH).

The characteristic strong sulcate sculpture of this species, coupled with its possession of
immarginate and unarmed lateral pronotal margins, stalked-suborbicular cephalic hairs, and
propodeal spines which are bowed outwards in dorsal view and downcurved in profile, make
jacksoni very easily recognisable.

Cataulacus moloch sp. n.

(Fig. 34)

HOLOTYPE WORKER. TL 3.4, HL 0.90, HW 0.80, CI 89, EL 0.43, OI 54, SL 0.42, SI 53, PW 0.60, AL 0.94.

With the head in full-face view the sides behind the eyes denticulate, ending in a low triangular tooth at
the occipital corner. Occipital margin with a small tooth close to the one at the corner but otherwise
unarmed; the occipital crest absent, the dorsum rounding evenly into the occipital surface. Eyes relatively
large, OI > 50. Alitrunk with promesonotum both longitudinally and transversely convex. In profile the
alitrunk with its highest point at about the midlength of the pronotum, behind which the dorsum is evenly
shallowly convex to the base of the propodeal spines. The steeply sloping anterior portion of the pronotal
dorsum with a number of minute peaks or tubercles from which hairs arise, such peaks absent elsewhere on
the dorsum. Tooth on mesokatepisternum moderately developed, distinct. Propodeal spines in profile
scarcely elevated and almost straight, only very feeble downcurved along their length; not having the basal
portions elevated and the distal portions recurved. With the alitrunk in dorsal view the pronotal corners
with prominent acute dentiform angles. Pronotal margin behind the corners with 5-6 triangular, laterally
projecting denticles which are quite evenly spaced. Sides of mesonotum with two small denticles, the sides of
the propodeum convex over the site of the spiracles, with one or two minute tubercles. Propodeal spines
broad in dorsal view and evenly divergent. Petiole in profile rising to an acute peak dorsally. Subpetiolar
process complex, with a blunt and strongly prominent anteroventral angle and a tooth-like projecting
posteroventral angle, the two separated by a conspicuously concave ventral margin. Postpetiole in profile
with the dorsal and posterior surfaces distinctly denticulate, the subpostpetiolar process elongate-digitiform.
Dorsum of head irregularly reticulate-rugulose, the reticular meshes of varying size and the rugulae
themselves low and rounded. Ground-sculpture within the meshes reduced to an inconspicuous vetigial
shagreening, without punctures. Pronotal dorsum similarly but more strongly sculptured, with a few low
broad transverse rugae anteriorly but with the longitudinal component predominating behind this. On the
mesonotum and propodeum the longitudinal component of the sculpture predominates, the rugae being
broader and more strongly developed ; many of the cross-meshes are feeble or incomplete. Rugae between
bases of propodeal spines transverse. Petiole in dorsal view regularly longitudinally rugose, the rugae
converging posteriorly; the postpetiole irregularly rugose. Ground-sculpture of alitrunk as on head. First
gastral tergite blanketed by dense reticulate-punctation, without strong basigastral rugulae but here and

362

B. BOLTON

r- m

-m

r-m

r-m

r-m

2r
Rs*m J Rs

J Rs

VM

Figs 35-43 Semi-diagrammatic representation of principal venation development on forewing of Messor
and Aphaenogaster. For explanation see text, pp. 339-340.

AFROTROPICAL MYRMICINE ANT GENERA 363

there with feeble short rugulae formed by the alignment of margins of adjacent punctures. First gastral
sternite reticulate-punctate. Sides of pronotum transversely sulcate. Discounting the long hairs which arise
around the eyes the dorsum of the head with numerous stalked-suborbicular hairs, the basal stems of which
are quite short. Occipital surface with longer hairs which increase in thickness from base to apex. All
remaining dorsal surfaces of body with numerous short stout blunt hairs. On the alitrunk some of these
hairs are slightly thicker apically than basally, these hairs straight everywhere except on the base of the first
gastral tergite where they are slighly back-curved. Colour uniform black; the scapes, tibiae and tarsi dull
yellow.

PARATYPE WORKERS. TL 2.8-3.2, HL 0.74-0.86, HW 0.68-0.72, CI 86-92, EL 0.39-0.42, OI 54-57, SL
0.38-0.40, SI 53-56, PW 0.50-0.60, AL 0.78-0.92 (3 measured).
As holotype but averaging slightly smaller.

Holotype worker, Ghana: Pankese, 24.ix.1968 (C. A. Collingwood) (BMNH).

Paratype. Ghana: 1 worker with same data as holotype. Nigeria: 2 workers, Onipe, CRIN, 1 l.vi.1975, tree
47-16 (A63.1), black pod project (B. Taylor] (BMNH).

C. moloch is closest related to centrums, the differences between them are noted under the latter
name.

Cataulacus satrap sp. n.

HOLOTYPE WORKER. TL 3.5, HL 0.87, HW 0.82, CI 94, EL 0.44, OI 53, SL 0.40, SI 49, PW 0.56, AL 0.96.

With the head in full-face view the sides behind the eyes minutely denticulate, the denticles partially
concealed by the thickened short hairs which project above them; the row of denticles ends in a small tooth
at the occipital corner. Occipital crest absent, the dorsum of the head rounding into the occipital surface.
Occipital margin unarmed except for a small tooth close to the one at the corner. Eyes relatively large,
OI > 50. In profile the anterior outline of the pronotal dorsum sloping steeply, the surface equipped with a
number of low peaks or tubercles. Behind this the remainder of the alitrunk shallowly but evenly convex,
sloping down posteriorly to the base of the propodeal spines. Mesokatepisternal tooth prominent,
moderately well developed. Metapleural lobes low and rounded. Propodeal spines in profile straight, only
slightly elevated. With the alitrunk in dorsal view the pronotal corners angular and projecting. Sides of
pronotum behind the corners only weakly marginate and with a series of 4-5 projecting denticles, all of
which are small and widely spaced. In the holotype the right pronotal margin with 5, the left with 4 denticles.
On both sides the posteriormost denticle the largest, the anteriormost distinctly smaller; the 2-3 between
them minute and inconspicuous. Sides of mesonotum and propodeum without differentiated denticles.
Propodeal spines in dorsal view broad and feebly divergent. Petiole in profile rising to an acute peak above.
Subpetiolar process simple, with a bluntly rounded anteroventral angle and an acute, weakly projecting
posteroventral angle, the two separated by a flat ventral surface. Postpetiole dome-like and high in profile,
with two feebly developed peaks dorsally; the subpostpetiolar process short-digitiform and blunt. Dorsum
of head irregularly reticulate-rugulose, the reticular meshes of uneven size and irregular shape, the rugulae
low and rounded. Ground-sculpture of the rugular meshes a fine dense reticulate-puncturation. Dorsal
alitrunk densely covered in fine rugulae which are low and rounded, reticulate in places but predominantly
longitudinal behind the pronotum. Entire dorsum of alitrunk also blanketed by a fine dense and very
conspicuous reticulate-punctate ground-sculpture. Petiole and postpetiole with dense reticulate-punctate
sculpture, the former also with longitudinal rugae in dorsal view, the latter only with a few vestigial irregular
rugulae. First gastral tergite strongly and densely reticulate-punctate everywhere. Dorsum of head with
numerous distinctive stalked-suborbicular hairs, those situated anteriorly on the dorsum more strongly
expanded apically than those situated behind the level of the eyes. All remaining dorsal surfaces of body with
many very short thick blunt hairs. Colour uniform black, dull; the scapes, tibiae and tarsi dull yellowish
brown.

PARATYPE WORKER. TL 3.4, HL 0.88, HW 0.80, CI 91, EL 0.43, OI 54, SL 0.40, SI 50, PW 0.57, AL 0.96.

As holotype but propodeal spines slightly less divergent and the subpetiolar process with the
anteroventral and posteroventral angles separated by a feebly concave ventral surface. On the pronotal
margins the anteriormost denticle behind the corner is no larger than those following it (except for the last in
the row, which is the largest); and the left side of the pronotum with 5 denticles, the right side with 4.

Holotype worker, Cameroun: Nkoemvon, 1970, M12 (D. Jackson) (BMNH).

Paratype. 1 worker with same data as holotype (BMNH).

Related to vorticus which it resembles closely, satrap is immediately separated by its possession of
denticles on the lateral pronotal margins.

364 B. BOLTON

Cataulacus taylori sp. n.

HOLOTYPE WORKER. TL 3.2, HL 0.82, HW 0.76, CI 93, EL 0.42, OI 55, SL 0.44, SI 58, PW 0.60, AL 0.90.

With the head in full-face view the sides behind the eyes denticulate, ending in a tooth at the occipital
corner. Occipital crest absent, the dorsum rounding evenly into the occiput; the occipital margin unarmed
except for a smaller tooth close to the one at the corner. Eyes relatively large, OI > 50. With the alitrunk in
profile the highest point of the dorsum at about the midlength of the pronotum. In front of this the dorsum
slopes down to the cervical shield and a few scattered minute peaks occur on the outline. Behind the highest
point the dorsum is shallowly convex and slopes evenly downwards towards the bases of the propodeal
spines. Mesokatepisternal tooth developed. Metapleural lobes low and rounded. Propodeal spines in profile
narrow, slightly downcurved along their length. Alitrunk in dorsal view with the pronotal corners
denticulate, the lateral margins of the pronotum with 6-7 projecting triangular denticles. Sides of
mesonotum and propodeum each with a single projecting denticle, the latter also with the sides convex at
the site of the spiracle. Propodeal spines narrow and evenly divergent in dorsal view. Petiole in profile rising
to a sharp peak above. Subpetiolar process complex, with a narrow rounded projecting blunt anteroventral
angle and a spur-like posteroventral angle, the ventral surface between the two angles strongly concave.
Postpetiole node with dorsal surface denticulate, the ventral process narrow and digitiform. Dorsum of head
feebly reticulate-rugulose, the rugulae very weak, fine, low and rounded, the reticular meshes mostly
incomplete and irregular in shape and size. Ground-sculpture in the meshes almost completely effaced, the
surface glossy. Dorsal alitrunk predominantly longitudinally rugose, with some anastomoses on the
pronotum but behind this the rugae straight and parallel, quite broad and without cross-meshes. Spaces
between the rugae glossy and almost smooth, with only the faintest vestiges of ground-sculpture. Rugae on
declivity between bases of spines transverse. Petiole and postpetiole longitudinally rugose, the rugae
converging posteriorly. First gastral tergite shiny, with superficial fine reticulate-puntulate sculpture
everywhere and with a weak pattern of very fine longitudinal irregular rugulae. Stronger longitudinal
rugulae present on the basal one-fifth of the tergite. First gastral sternite similarly but even more delicately
sculptured. Dorsum of head with numerous short stout straight cylindrical hairs which are blunt apically.
All remaining dorsal surfaces of body with similar pilosity, the longest hairs occurring on the base of the first
gastral tergite where they are slightly recurved. Colour uniform glossy jet black; the scapes, tibiae and tarsi
dull yellow.

PARATYPE WORKER. TL 3.5, HL 0.88, HW 0.81, CI 92, EL 0.45, OI 56, SL 0.46, SI 57, PW 0.67, AL 0.96.

As holotype but slightly larger, its subpostpetiolar process shorter and broader than in the holotype. The
rugae on the dorsal alitrunk not running straight back as in the holotype but slightly skewed to the left
posteriorly.

Holotype worker, Nigeria: Gambari, CRIN, 24. v. 1976, black pod project (B. Taylor) (BMNH).
Paratype. Nigeria: 1 worker, Onipe, CRIN, 25.vii.1975, black pod project (B. Taylor) (BMNH).

Appendix

The current genus-level synonymy of Aphaenogaster is as follows.

APHAENOGASTER Mayr

Aphaenogaster Mayr, 1853: 107. Type-species: Aphaenogaster sardoa Mayr, 1853: 107, by subsequent

designation of Bingham, 1903: 270.
Deromyrma Forel, 1913c: 49 [as subgenus of Ischnomyrmex Mayr]. Type-species: Aphaenogaster

(Ischnomyrmex) swammerdami Forel, 18866: cvi, by monotypy. [Synonymy by Brown, 1973: 180.]
Planimyrma Viehmeyer, 1914: 604 [as subgenus of Aphaenogaster}. Type-species : Stenamma (Ischnomyrmex)

loriai Emery, 1897: 563, by original designation. [Synonymy by Brown, 1973: 184.]
Attomyrma Emery, 1915: 70 [as subgenus of Aphaenogaster}. Type-species: Formica subterranea Latreille,

1 798 : 49, by original designation. [Synonymy by Brown, 1 973 : 1 78.]
Novomessor Emery, 1915: 73. Type-species: Aphaenogaster (Ischnomyrmex) cockerelli Andre, 1893: 150, by

original designation. [Synonymy by Brown, 1974: 47.]
Nystalomyrma Wheeler, 1916: 215 [as subgenus of Aphaenogaster}. Type-species: Myrmica longiceps F.

Smith, 1858: 128, by original designation. [Synonymy by Brown, 1973: 183.]
Brunella Forel, 1917: 234. Type-species: Aphoenogaster [sic] belli Forel, 1895: 248, by original designation.

Syn. n.

AFROTROPICAL MYRMICINE ANT GENERA 365

Acknowledgements

My sincere thanks go to the following for the loan of types and other material during the course
of this study.

Dr Claude Besuchet (MHN, Geneva); Mrs Cathy A. Car (NM, Bulawayo); Dr Jean Decelle
(MRAC, Tervuren); Mrs Marjorie Favreau (AMNH, New York); Dr Max Fischer (NM,
Vienna); Dr F. Koch (MNHU, Berlin); Prof Egidio Mellini (IE, Bologna); Mr Alfred Newton Jr
(MCZ, Cambridge); Dr Roberto Poggi (MCSN, Genoa); Dr A. G. Radchenko (ZM, Kiev); Miss
Helen Rae and Dr A. J. Prins (SAM, Cape Town); Dr David R. Smith (USNM, Washington); Dr
Cesare Baroni Urbani (NM, Basle); Mme Janine C. Weulersse (MNHN, Paris).

Finally my thanks to Ms Dorothy Jackson of Oxford University for collecting Cataulacus
material for me at my request, and to Mr David Morgan for lettering Figs 35-43.

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366 B. BOLTON

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— 1890a. Fourmis de Tunisie et de 1'Algerie orientale. Cr. Soc. ent. Belg. 34: Ixi-lxxvi.

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AFROTROPICAL MYRMICINE ANT GENERA 367

— 1905. Miscellanea myrmecologiques 2. Annls Soc. ent. Fr. 49: 155-185.

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Sudamerika und Afrika. Dt. ent. Z. 1911 : 285-312.

- 1912. Descriptions provisoires de genres, sous-genres et especes de formicides des Indes orientales.
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— 191 3c. Wissenschaftliche Ergebnisse einer Forschungsreise nach Ostindien, ausgefiihrt im Auftrage der
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— 1916. Fourmis du Congo et d'autres provenances recoltees par MM Hermann Kohl, Luja, Mayne etc.
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- 1917. Cadre synoptique actuel de la faune universelle des fourmis. Bull. Soc. vaud. Sci. nat. 51 : 229-253.
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419^64.
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368 B. BOLTON

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— 1912. Fourmis d'Afrique et de Madagascar. Annls Soc. ent. Belg. 56: 150-167, figs.

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— 1926. Description de nouveaux formicides Ethiopiens. Revue Zool. Bot. Afr. 13: 207-267, 6 figs.

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AFROTROPICAL MYRMICINE ANT GENERA

369

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28: 259-265, 3 figs.

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aethiops 358
airensis 350
angularis 344
angulatus 324
Aphaenogaster 364
arcistriatus 348
armata 350
Attomyrma 364

badonei 316

beccarii 336

braunsi (Leptothorax) 325

braunsi (Messor) 345

brevispinosa 316

Brunella 364

brunni 349

capensis 345
Cardiocondyla 309
Cataulacus 354
Caulomyrma 319
cenatus 327
centrums 359
cephalotes 346
cestus 360
chlorotica 317
collingwoodi 346
compressus 337
concolor 324
Cratomyrmex 338

decipiens 348
denticornis 349
denticulatus 328
Deromyrma 364
Dichothorax 319
donisthorpei 345
Dyclona 309

emeryi (Cardiocondyla) 312
emeryi (Melissotarsus) 337
Emery ia 309
evelynae 328

Index

Synonyms are in italics,
fusca 316

galla 349
globinodis 316
Goniothorax 319
grisoni 329

hawaiensis 317
humerosus 329

Icothorax 319
ilgii 324
incisus 351
innocens 330

jacksoni 360
jeanneli 358

kenyensis 358

laevifrons 354
latinoda 350
latinodis 324
Leptothorax 319
Limnomyrmex 319
Lobognathus 338
Loncyda 309
luebberti 351
lujae 358

mahdii 313
major 337
mauritia 313
megalops 331
Melissotarsus 333
Messor 338
moloch 361
monardi 314
monilicornis 313
Mychothorax 319
Myrafant 319
Myrmammophilus 319

370

B. BOLTON

neferka 314
nereis 313
Nesomyrmex 319
nigriventris 354
nilotica 315
nobilis 350
Novomessor 364
Nystalomyrma 364

parvidens 349
piceus 352
pilipes 337
Planimyrma 364
p/iw'i 346
profta 348
Prosopidris 309
pseudoaegyptiaca 345

rasalamae 312
regalis 352
rwftea 352
ru/oi 349
ru/w/a 350
ruginodis 353

satrap 363
schencki 345

sculptior 316
sculpturatus 352
sekhemka 315
shuckardi 316
simoni 331
stramineus 332
striatifrons 353

taylori 364
Temnothorax 319
Tetramyrma 319
titubans 336
traegaordhi 358
triempressa 350
tropicorum 354

Veromessor 338

316

weissi (Cataulacus) 358
weissi (Melissotarsus) 337
weserka 317
wroughtonii 317

Xenometra 309
zoserka 318

British Museum (Natural History)
Chance, change & challenge

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Titles to be published in Volume 45

A catalogue and reclassification of the Ichneumonidae (Hymenoptera) described by C. G.
Thomson.

By M. G. Fitton

A taxonomic review of the genus Phlebotomus (Diptera : Psychodidae).
By D. J. Lewis

Stenomine moths of the Neotropical genus Timocratica (Oecophoridae).

By V. O. Becker

Afrotropical species of the myrmicine ant genera Cardiocondyla, Leptothorax,
Melissotarsus, Messor and Cataulacus (Formicidae).
By Barry Bolton

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