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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 
scientific series, Botany, Entomology, Geology (incorporating Mineralogy) and Zoology, and an 
Historical series. 

Papers in the Bulletin are primarily the results of research carried out on the unique and 
ever-growing collections of the Museum, both by the scientific staff of the Museum and by 
specialists from elsewhere who make use of the Museum's resources. Many of the papers are 
works of reference that will remain indispensable for years to come. 

Parts are published at irregular intervals as they become ready, each is complete in itself, 
available separately, and individually priced. Volumes contain about 300 pages and several 
volumes may appear within a calendar year. Subscriptions may be placed for one or more of the 
series on either an Annual or Per Volume basis. Prices vary according to the contents of the 
individual parts. Orders and enquiries should be sent to : 



Publications Sales, 

British Museum (Natural History), 
Cromwell Road, 

London SW7 5BD, 
England. 



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'fusiformis 1 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 f r S 3 elsan 3 

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. [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. [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, Id 1 , 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 'J 1 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 d 1 , 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?/CSTownes 

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. 

References 

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- 19736. Revision des especes ouest-palearctiques du genre Triclistus Foerster (Hymenoptera: Ichneu- 
monidae). Mitt, schweiz. ent. Ges. 46: 219-252. 

1975. Revision des especes ouest-palearctiques du genre Chorinaeus Holmgren [Hymenoptera, Ich- 



neumonidae]. Annls Soc. ent. Fr. (N.S.) 11 : 723-744. 
Aubert, J. F. 1959. Les Ichneumonides du rivage Mediterraneen Francais (Cote d'Azur). Annls Soc. ent. Fr. 
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- 1960. Les Ichneumonides des Pyrenees-Orientales. Vie Milieu 11 : 473-493. 



THE ICHNEUMONIDAE DESCRIBED BY C. G. THOMSON 101 

- 1961. Revision des travaux concernant les Ichneumonides de France et 3me supplement au cat- 
alogue de Gaulle (80 especes nouvelles pour la faune Franpaise). Bull, metis. Soc. linn. Lyon 30: 195-200. 
1964. Les Ichneumonides du rivage Mediterraneen Fran9ais [Hym.]. 7e serie: Ichneumoninae, 



Cryptinae, Ophioninae et Mesochorinae de 1'Herault et des Bouches-du- Rhone. Bull. Soc. ent. Fr. 69: 
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1965. Les Ichneumonides du rivage Mediterraneen Fran9ais (8e Serie, Region cotiere entre La 



Ciotat et Saint-Tropez). Vie Milieu 16 (c): 549-573. 

1966. Fixations d'Ichneumonides lectotypes dans la collection C. G. Thomson conservee a Lund. 



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1968. Fixation des types, lectotypes et paratypes dans les collections d'Ichneumonides, et premiere 



liste de types perdus ou conserves. Mitt, schweiz. ent. Ges. 41 : 175-201. 

1969. Les Ichneumonides ouest-palearctiques et leurs holes. 1 (Pimplinae, Xoridinae, Acaenitinae). 



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de Gaulle (100 especes nouvelles pour la faune Fran9aise). Bull. mens. Soc. linn. Lyon 39: 269-280. 

1972. Etude commentee de nouveaux lectotypes choisis dans les collections Holmgren et Thomson 



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19766. Les Mesoleius Holmgren des collections Holmgren et Thomson (Hymenoptera: Ichneumon- 



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abdominal sclerites in the male (Hym.: Ichneum.). Trans. R. ent. Soc. Lond. 91 : 661-712. 
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102 M. G. FITTON 

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Gravenhorst, J. L. C. 1829a. Ichnewnonologia Europaea. 1, xxxi + 827 pp. Vratislaviae. 

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Hinz, R. 1963. Ober einige Typen der Gattung Dusona Cameron (Hymenoptera: Ichneumonidae). Beitr. Ent. 
13: 335-344. 

- 1975. Die Arten der Gattung Glyptorhaestus Thomson (Hymenoptera, Ichneumonidae). Z. ArbGem. 
ost.Ent.27: 39^*6. 

Holmgren, A. E. 1860. Forsok till uppstallning och beskrifning af Sveriges ichneumonider. Tredje Serien. 

Fam. Pimplariae. (Monographia Pimplariarum Sueciae). K. svenska VetenskAkad. Handl. (B) 3(10): 1-76. 
Horstmann, K. 1967a. Untersuchungen zur Systematik einiger Phygadeuon Arten aus der Verwandtschaft 

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Munch. 98: 1-22. 

- 19676. Bemerkungen zur Taxonomie der Tersilochinen (Hym. Ichneumonidae). Opusc. ent. 32: 
123-130. 

1968. Revision einiger Arten der Gattungen Mesostenus Gravenhorst, Agrothereutes Foerster und 



Ischnus Gravenhorst (Hymenoptera, Ichneumonidae). Entomophaga 13: 121-133. 

1969. Typenrevision der europaischen Arten der Gattung Diadegma Foerster (syn. Angitia Holm- 



gren) (Hymenoptera: Ichneumonidae). Beitr. Ent. 19: 413^472. 

1971a. Revision der europaischen Arten der Gattung Lathrostizus Foerster (Hymenoptera, Ichneu- 



monidae). [part]. Mitt. dt. ent. Ges. 30: 8-12. 

19716. Revision der europaischen Tersilochinen (Hymenoptera, Ichneumonidae). Teil 1. Veroff. zool. 



StSamml. Munch. 15: 45-138. 

- 1971c. Revision der europaischen Arten der Gattung Lathrostizus Foerster (Hymenoptera, Ichneu- 
monidae) [part]. Mitt. dt. ent. Ges. 30: 16-18. 

1972. Type revision of the species of Cryptinae and Campopleginae described by J. B. Bridgman 



(Hymenoptera, Ichneumonidae). Entomologist 105: 217-228. 

1973a. Revision der westpalaarktischen Arten der Gattung Nemeritis Holmgren (Hymenoptera, 



Ichneumonidae). Opusc. zool. Munch. 125: 1-14. 

19736. Revision der europaischen Arten der Gattung Dichrogaster Doumerc (Hym. Ichneumon- 



idae). Ent. scand. 4: 65-72. 

1973c. Revision der Gattung Nepiesta Foerster (mit einer Obersicht iiber die Arten der Gattung 



Leptoperilissus Schmiedeknecht (Hymenoptera Ichneumonidae). Polskie Pismo ent. 43: 729-741. 

1974a. Revision der westpalaarktischen Arten der Schlupfwespen-Gattungen Bathyplectes und Bioly- 



sia (Hymenoptera: Ichneumonidae). Entomologica germ. 1 : 58-81. 

19746. Typenrevision der von Strobl in der Gattung Hemiteles Gravenhorst s.l. beschriebenen 



Arten und Formen (Hymenoptera, Ichneumonidae). Z. ArbGem. ost. Ent. 25: 52-56. 

1974c. Typenrevision der von E. Zilahi-Kiss beschriebenen Hemitelinen mit Bemurkungen zu den 



Gattungen Hemiteles Grav. (s.str.), Gnotus Foerst. und Xiphulcus Townes (Hymenoptera, Ichneumon- 
idae). Annls hist.-nat. Mus. natn. Hung. 66: 339-346. 

1976a. Wenig bekannte oder neue europaische Hemitelinen-Gattungen (Hymenoptera, Ichneumon- 



idae, Cryptinae). NachrBl. bayer. Ent. 25: 22-31. 

19766. Nachtrag zur Revision der europaischen Dichrogaster-Arten (Hymenoptera, Ichneumon- 



idae). Z. ArbGem. ost. Ent. 28: 55-61. 

1977. Bemerkungen zur Systematik einiger Gattungen der Campopleginae II (Hymenoptera, Ich- 



neumonidae). Mitt, munch, ent. Ges. 67: 65-83. 

1978. Revision der Gattungen der Mastrina Townes (Hymenoptera, Ichneumonidae, Hemitelinae). 



Z. ArbGem. ost. Ent. 30: 65-70. 

- 1979a. A revision of the types of the Hemiteles spp. described by Thomson (Hymenoptera: Ichneu- 
monidae). Ent. scand. 10: 297-302. 

- 19796. Revision der europaischen Arten der Gattung Ceratophygadeuon Viereck (Hymenoptera, 
Ichneumonidae). Z. ArbGem. ost. Ent. 31 : 41-48. 

1981. Revision der europaischen Tersilochinae. II. (Hymenoptera, Ichneumonidae). Spixiana Sup- 



plement 4: 1-76. 

Huggert, L. 1973. Taxonomical studies on Platygastrinae (Hym. Proctotrupoidea). Ent. Tidskr. 94: 97-108. 
Idar, M. 1973. Designation of types for the species of Hadrodactylus Frst. (Hym. Ichneumonidae) described 

by C. G. Thomson, with redescriptions of H. bidentulus (Ths.) and H. villosulus (Ths.). Ent. scand. 4: 23-28. 
- 1974. Redescriptions of Hadrodactylus tarsator Ths., H. gracilipes Ths. and H. nigrifemur Ths., with 

notes on H. bidentulus Ths. and H. villosulus Ths. (Hym. Ichneumonidae). Ent. Tidskr. 95: 107-1 16. 



THE ICHNEUMONIDAE DESCRIBED BY C. G. THOMSON 103 

- 1975. Redescriptions of Hadrodactylus paludicola (Hlgr.), H.femoralis (Hlgr.) and H. insignis (Krb.) 
(Hymenoptera, Ichneumonidae). Ent. scand. 6: 182-190. 

Jussila, R. 1965. The Ichneumonidae of the Kevojoki area in Inari Lapland (Finland). Annls Univ. turku 
(A.II) 34: 1-186. 

- 1966. The males of the ichneumonid species (Hym.) Alloplasta breviventris Hellen, Mesoleius erythro- 
gaster Hlmgr. and Homotropus crassicrus Thorns. Acta ent.fenn. 32: 317-321. 

- 1967. Ichneumonological (Hym.) reports from Finland 1. Suomenhyont. Aikak. 33: 107-112. 

1979. A revision of the genus Atractodes (Hymenoptera, Ichneumonidae) in the western Palaearctic 



region. Acta ent.fenn. 34: 1-44. 
Kasparyan, D. R. 1969. Palaearctic species of the genus Tryphon Fallen (Hymenoptera, Ichneumonidae), I. 
[In Russian.] Ent. Obozr. 48: 639-662. 

- 1970. Ichneumonids of the genus Polyblastus Hartig (Hymenoptera, Ichneumonidae) of Palaearctic. 
[In Russian.] Ent. Obozr. 49: 852-868. 

1971. A revision of the Palaearctic species of the genus Cosmoconus Foerster (Hymenoptera, Ichneu- 



monidae). Trudy vses. ent. Obshch. 54: 286-307. 

1973. Hymenoptera 3 (1). Ichneumonidae, subfamily Tryphoninae, tribe Tryphonini. [In Russian.] 



Fauna SSSR (N.S.) 106: 1-320. 

1975. The species of the genus Exyston Schi^dte (Hymenoptera, Ichneumonidae) in the fauna of the 



USSR and the Mongolian People's Republic. [In Russian.] Insects of Mongolia 3: 294-305. 
Kerrich, G. J. 1938. A revision of the British species of the genus Hygrocryptus Thorns. (Hym., Ichneumon- 
idae, Cryptini). Trans. Soc. Br. Ent. 5: 169-177. 

- 1939. Systematic notes on the Oxytorina (Hym., Ichneumonidae, Mesoleptini Auctt.). Opusc. ent. 4: 
126-128. 

1942. Second review of literature concerning British Ichneumonidae (Hym.), with notes on Palaearc- 



tic species. Trans. Soc. Br. Ent. 8 : 43-77. 

1952. A review, and a revision in greater part, of the Cteniscini of the Old World (Hym., Ichneumon- 



idae). Bull. Br. Mus. nat. Hist. (Ent.) 2: 305^460. 

1953. A preliminary study of the European species of the genus Eudiaborus mihi (Hym., Ichneumon- 



idae). Opusc. ent. 18: 151-159. 

1962. Systematic notes on Tryphonine Ichneumonidae (Hym.). Opusc. ent. 27: 45-56. 



Oehlke, J. 1967. Westpalaarktische Ichneumonidae I : Ephialtinae. H ymenopterorum Catalogus (nova editio) 

2, vii + 49 pp. 's-Gravenhage. 

Perkins, J. F. 1941. A synopsis of the British Pimplini with notes on the synonymy of the European species 
(Hymenoptera, Ichneumonidae). Trans. R. ent. Soc. Lond. 91 : 637-659. 

- 1943a. Preliminary notes on the synonymy of the European species of the Ephialtes complex (Hym.), 
Ichneumonidae. Ann. Mag. nat. Hist. (1 1) 10: 249-273. 

19436. Notes on the British species of Adelognathini Roman with descriptions of two new species 



(Hym. Ichneumonidae). Trans. R. ent. Soc. Lond. 93: 95-1 14. 

1953. Notes on British Ichneumoninae with descriptions of new species (Hym., Ichneumonidae). 



Bull. Br. Mus. nat. Hist. (Ent.) 3: 103-176. 

1959. Ichneumonidae, key to subfamilies and Ichneumoninae-I. Handbk I dent. Br. Insects 7 (2ai): 



1-116. 

1960. Ichneumonidae, subfamilies Ichneumoninae II, Alomyinae, Agriotypinae and Lycorininae. 



Handbk I dent. Br. Insects 7 (2aii): 117-213. 

1962. On the type species of Foerster's genera (Hymenoptera: Ichneumonidae). Bull. Br. Mus. nat. 



Hist. (Ent.) 11: 383^83. 
Provancher, L. 1874. Les Ichneumonides de Quebec avec description de plusieurs especes nouvelles [part]. 

Naturalistecan.6: 143-151. 

Roman, A. 1939. Nordische Ichneumoniden und einige andere. Ent. Tidskr. 60: 176-205. 
Rossem, G. van. 1966. A study of the genus Trychosis Foerster in Europe (Hymenoptera, Ichneumonidae, 
Cryptinae). Zool Verh. Leiden 79: 1-40. 

- 1969a. A study of the genus Meringopus Foerster in Europe and of some related species from Asia 
(Hymenoptera, Ichneumonidae, Cryptinae). Tijdschr. Ent. 112: 165-196. 

19696. A revision of the genus Cry plus Fabricius s. str. in the western Palaearctic region, with keys 



to genera of Cryptina and species of Cryptus (Hymenoptera, Ichneumonidae). Tijdschr. Ent. 112: 299-374. 
1971. The genus Buathra Cameron in Europe (Hymenoptera, Ichneumonidae). Tijdschr. Ent. 114: 



201-207. 

Sawoniewicz, J. 1978. Zur Systematik und Faunistik der Ichneumonidae (Hymenoptera). Annls zool. Warsz. 
34:121-137. 



104 M. G. FITTON 

- 1980. Revision of European species of the genus Bathythrix Foerster (Hymenoptera, Ichneumon- 
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Schmiedeknecht, 0. 1900. Die palaarktischen Gattungen und Arten der Ichneumonidentribus der Lissonoti- 
den. Zoo/. Jb. 13(Syst.): 299-398. 

- 1903. Opuscula Ichneumonologia 1, Fasc. IV: 241-320. Blankenburg. 

Sedivy, J. 1961. Beitrag zur Kenntnis der Tryphoninen-Gattungen Phytodietus Grav. und Weisia Schmdkn. 
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- 1963. Die europaischen Arten der Gattungen Laufeia Tosq., Polysphincta Grav. und Zatypota Forst. 
(Hym., Ichneumonidae). Ada ent. Mus. natn. Prague 35: 243-261. 

1970. Westpalaarktischen Arten der Gattungen Dimophora, Pristomerus, Eucremastus und Cre- 



mastus (Hym. Ichneumonidae). Pfirodov Pr. Cesk. Akad. Ved. 4(11): 1-38. 

1971. Revision der europaischen Temelucha- Arten (Hym. Ichneumonidae). Pfirodov. Pr. Cesk. Akad. 



Fed. 5(1): 1-34. 
Stelfox, A. W. 1941. Description of six new species of bassine ichneumon-flies, with notes on some others. 

Proc. R. Ir. Acad. (B) 46: 109-1 19. 

Thomson, C. G. 1873. Opuscula Entomologica. 5: 455-530. Lund. 
1874. Opuscula Entomologica. 6: 535-612. Lund. 

- 1877. Opuscula Entomologica. 8: 732-841. Trelleborg. 

- 1883. Opuscula Entomologica. 9: 843-936. Lund. 

- 1884. Opuscula Entomologica. 10: 939-1040. Lund. 

- 1885. Notes Hymenopterologiques (Premiere partie: Cryptidae). Annls Soc. ent. Fr. (6) 5: 17-32. 

- 1886a. Notes Hymenopterologiques. Deuxieme partie. (Genre Mesochorus). Annls Soc. ent. Fr. (6) 
5: 327-344. 

- 18866. Notes Hymenopterologiques. Troisieme partie. Observations sur le genre Ichneumon et 
descriptions de nouvelles especes. No ler. Annls Soc. ent. Fr. (6)6: 1 1-24. 

1887a. Notes Hymenopterologiques. Quatrieme partie. Observations sur le genre Ichneumon et 



descriptions de nouvelles especes. No II. Annls Soc. ent. Fr. (6)1: 5-16. 

- 18876. Hymenopterologische Beitrage. Dt. ent. Z. 31 : 193-218. 

- 1887c. Opuscula Entomologica. 11: 1043-1182. Lund. 

1888a. Notes Hymenopterologiques. Cinquieme partie. Observations sur le genre Ichneumon et 



descriptions de nouvelles especes. No III. Annls Soc. ent. Fr. (6) 8: 105-126. 

- 18886. Opuscula Entomologica. 12: 1185-1318. Lund. 

- 1889. Opuscula Entomologica. 13: 1321-1438. Lund. 

- 1890. Opuscula Entomologica. 14: 1441-1534. Lund. 

- 1891. Opuscula Entomologica. 15: 1537-1656. Lund. 

- 1892a. Opuscula Entomologica. 16: 1659-1773. Lund. 

- 18926. Opuscula Entomologica. 17: 1777-1886. Lund. 

1893. Opuscula Entomologica. 18: 1889-1967. Lund. 

1894. Opuscula Entomologica. 19: 1971-2137. Lund. 

1896. Opuscula Entomologica. 21 : 2343-2404. Lund. 

1897. Opuscula Entomologica. 22 : 2407-2452. Lund. 



Tolkanit/., V. I. 1973. Ichneumon-flies of the genus Phytodietus Gravenhorst (Hymenoptera, Ichneumon- 
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Townes, H. 1959. The present condition of the Gravenhorst collection of Ichneumonidae (Hymenoptera). 
Proc. ent. Soc. Wash. 61 : 76-78. 

- 1969. The genera of Ichneumonidae. Part 1. Mem. Am. ent. Inst. 11 : 1-300. 

- 1970a. The genera of Ichneumonidae. Part 2. Mem. Am. ent. Inst. 12: 1-537. 

- 19706. The genera of Ichneumonidae. Part 3. Mem. Am. ent. Inst. 13: 1 307. 

- 1971. The genera of Ichneumonidae. Part 4. Mem. Am. ent. Inst. 17: 1-372. 

Townes, H., Momoi, S. & Townes, M. 1965. A catalogue and reclassification of the eastern Palaeartic 

Ichneumonidae. Mem. Am. ent. Inst. 5: 1-661. 
Townes, H. & Townes, M. 1959. Ichneumon-flies of America North of Mexico: 1. Subfamily Metopiinae. 

Bull. U.S. natn. Mus. 216(1): 1-318. 

- 1962. Ichneumon-flies of America North of Mexico: 3. Subfamily Gelinae, Tribe Meso- 

stenini. Bull. U.S. natn. Mus. 216(3): 1-602. 
Viktorov, G. A. & Athanasov, A. Z. 1974. Materials on the revision of Palaearctic ichneumonids of the tribe 

Theriini (Hymenoptera, Ichneumonidae). [In Russian.] Ent. Obozr. 53: 374-381. 



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) 
Chance, change & challenge 

Two multi-author volumes from one of the foremost scientific institutions in the world. 

General Editor: P. H. Greenwood 



The Evolving Earth 

Editor:L.R.M. Cocks 



The Evolving Biosphere 

Editor: P. L. Forey 



In the first volume, The Evolving Earth, twenty scientists have been asked to review the present 
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sediments and soils to continental drift and palaeogeography. 

In the companion volume, The Evolving Biosphere, museum scientists have chosen an 
evolutionary concept speciation, coevolution, biogeography etc. and related this to the group 
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In both volumes the text is supplemented by over one hundred specially-commissioned pieces of 
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These two books will be invaluable to all sixth-form and undergraduate biology and geology 
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The Evolving Earth: 276 x 219 mm, 280pp, 138 line illustrations, 42 halftones 
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Published: May 1981 

Co-published by the British Museum (Natural History), London and Cambridge University 
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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 



Typeset by Santype International Ltd., Salisbury and Printed by Henry Ling Ltd., Dorchester. 



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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ISSN 0524-6431 

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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 R 2 + 3 has been displaced towards the wing tip so that the vein 
seems to come from /? 4 , and R 2 has been reduced (Hennig, 1969: 385). R 2 + 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 R 2 + 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 20C 
(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 49N, and in Asia P. chinensis is the most 
northerly species (Perfil'ev, 1968: 89), reaching about 48N (Beklemishev & Dolmatova, 1948: 
354). In Canada sandflies are known from about 5039'N near Kamloops, from 4939'N at 
Coulee Creek in Alberta, and at 4441'N near Ottawa. The southern boundary of sandflies in the 
Old and New Worlds is about 40S (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 R 2 + 3 and R* and the 

origin of R 5 



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. 

R 2 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 R 4 and R s 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 R 4 and R 5 relatively long, at least a third of width of wing. Genital 
pump without adjacent rods. Palpal sensilla spatulate 3 

2 Vein M 1 + 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 M 1 + 2 forking beyond level of radio-median cross- vein, beyond base of R 4 . 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, R 2 /R 2 + 3 4-33, /? t overlap//? 2 0-66, R^ and R 2 
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,K 2 / 2 + 3 5-42 (4-35-6-10), 
R 1 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, 504'S 3902'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, M 1 + 2 fork 0-105 mm beyond origin of R 4 . 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, M 1 + 2 fork = +0-091 mm beyond 
origin of R 4 . 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, 3609'N, 6824'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. 
3167'N, 3533'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 (3202'N, 
3604'E, 15.vii.1978, E. K. S.); Sweilah (3202'N, 3550'E, 2.vii.l978, E. K. S.). Kuwait: Al Fantas, 2910'N, 
4806'E, Al Farwaniya area, 2916'N, 4755'E, 'Al Kazuma', Kuwait & 'Malboula', 1974, K. B.); Hussien & 
Behbehani (1976). Libya: Ashford et al. (1977: 265, Al Birkah, 3205'N, 2005'E, etc.); Nalut & Yafran area, 
3204'N, 1231'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, 4550'N, 1346'E, Ba Tays, 1320'N, 4518'E, 
Ba Zulayfah, 1350'N, 4547'E, Dali, 1342'N, 4444'E, Khodad, 1309'N, 4450'E, Mukalla area, Nigda, 
1356'N, 4555'E, Sah, 1534'N, 4851'E and Saywun area, 1556'N, 4927'E (1962, S. A. S.); 'Haski' (1939, 
cJ, P.); Sha'b Subeihi, 1305'N, 4430'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, 3630'N, 3751'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 6C 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, 3737'N, 6210'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, 4530'N, 9930'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 4250'N, 5050'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 (3202'N, 3604'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 R 2 /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, 4752'N, 017'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, 
3516'N, 2542'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, 4903'N, 007'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 4928'N, 0158'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, 731'N, 
3630'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, 3304'N, 3520'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 4912'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 16C, 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, 3439'N, 6942'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, 3146'N, 10406'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 R 4 near that of R 5 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 ofR 2 + 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 R 2 longer than R 2 + 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 R 2 shorter than R 2 + 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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204 D. J. LEWIS 

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THE GENUS PHLEBOTOMUS 



207 



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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). 
By V. O. Becker 

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Stenomine moths of the Neotropical genus 
Timocratica (Oecophoridae) 



Vitor O. Becker 



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Vol 45 No 3 26 August 1982 



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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 (CuA t and CuA 2 ) 
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 CuA l and 
CuA 2 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 CuA l and CuA 2 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 6C 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 CuA l and CuA 2 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 jR 4 and R 5 , 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 R 4 and R 5 stalked 
p. Fore wing with R 4 and R 5 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 

butx ro ' 
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]), R l from middle of cell, 
R 2 closer to R 3 than to R l} R 4 and R s very close, connate or stalked; CuA l and CuA 2 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 R 4 and R 5 have a short 
common stalk while the left shows the normal condition with R 4 and R 5 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 R 4 and/? 5 free (monotonia-group) 6 

Fore wing with R 4 and R 5 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 CuA r and CuA 2 stalked species 3 (p. 243) 

Fore wing with veins CuA l 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 K 4 and R 5 stalked venifurcata (p. 245) 

Fore wing with R 4 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 R 4 and R 5 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 R 4 and R 5 and CuA { and CuA 2 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; CuA v and CuA 2 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 CuA t and CuA 2 of the fore wing, and from parvifusca by the larger size 
and veins R 4 and R 5 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 M 3 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 




12C 



24C 



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 




12C 



-24C 



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 



24C 



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 




12C 



24C 



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 CuA l and CuA 2 , or R 4 and R 5 , 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 R 4 and R 5 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 CuA 2 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 CuA 2 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 CuA 2 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 M 3 and CuA v 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; R 4 and R s connate basally, CuA l and CuA 2 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 




12C 



24C 



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; R 4 and 
R s , and CuA r and CuA 2 , 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 
R 5 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 CuA 2 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 R 4 and R s . 

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 




- 24C 



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 




12C 



- 24C 



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 CuA 2 
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 CuA 2 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 




24C 



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 R 3 Af 3 , 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; CuA t and CuA 2 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 CuA l and CuA 2 , 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 CuA 2 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 




12C 




24C 



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 K 5 , andCuAt and 
CuA 2 , 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 R 4 and R 5 , and CuA l and CuA 2 , connate or stalked, and 
some have R 4 and R 5 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 
R 5 , CM/I! and CuA 2 stalked, M l 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 
R 4 and R 5 , and the lack of M l 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 



V. O. BECKER 




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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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 



Typeset by Santype International Ltd., Salisbury and Printed by Henry Ling Ltd., Dorchester 



/* GENERAL *\ 

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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. 14 U 
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. 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 close