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
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Trustees of the British Museum (Natural Histo
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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, 6s.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
(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)