contributions
of
American
the
Entomological
Institute
Volume 24, Number 1, 1988
THE MOSQUITOES OF THE SUBGENUS CULEX IN
SOUTHWESTERN
ASIA AND EGYPT (DIPTERA: CULICIDAE)
bY
Ralph E. Harbach
Form Approved
OMB No. 0704-0188
Report Documentation Page
Public reporting burden for the collection of information is estimated to average 1 hour per response, including the time for reviewing instructions, searching existing data sources, gathering and
maintaining the data needed, and completing and reviewing the collection of information. Send comments regarding this burden estimate or any other aspect of this collection of information,
including suggestions for reducing this burden, to Washington Headquarters Services, Directorate for Information Operations and Reports, 1215 Jefferson Davis Highway, Suite 1204, Arlington
VA 22202-4302. Respondents should be aware that notwithstanding any other provision of law, no person shall be subject to a penalty for failing to comply with a collection of information if it
does not display a currently valid OMB control number.
1. REPORT DATE
3. DATES COVERED
2. REPORT TYPE
1988
00-00-1988 to 00-00-1988
4. TITLE AND SUBTITLE
5a. CONTRACT NUMBER
The Mosquitoes of the Subgenus Culex in Southwestern Asia and Egypt
(Diptera: Culicidae)
5b. GRANT NUMBER
5c. PROGRAM ELEMENT NUMBER
6. AUTHOR(S)
5d. PROJECT NUMBER
5e. TASK NUMBER
5f. WORK UNIT NUMBER
7. PERFORMING ORGANIZATION NAME(S) AND ADDRESS(ES)
Walter Reed Army Institute of Research,Department of
Entomology,Walter Reed Biosystematics Unit,Washington,DC,20307
9. SPONSORING/MONITORING AGENCY NAME(S) AND ADDRESS(ES)
8. PERFORMING ORGANIZATION
REPORT NUMBER
10. SPONSOR/MONITOR’S ACRONYM(S)
11. SPONSOR/MONITOR’S REPORT
NUMBER(S)
12. DISTRIBUTION/AVAILABILITY STATEMENT
Approved for public release; distribution unlimited
13. SUPPLEMENTARY NOTES
14. ABSTRACT
see report
15. SUBJECT TERMS
16. SECURITY CLASSIFICATION OF:
a. REPORT
b. ABSTRACT
c. THIS PAGE
unclassified
unclassified
unclassified
17. LIMITATION OF
ABSTRACT
18. NUMBER
OF PAGES
Same as
Report (SAR)
246
19a. NAME OF
RESPONSIBLE PERSON
Standard Form 298 (Rev. 8-98)
Prescribed by ANSI Std Z39-18
Dedicated to
Kenneth L. Knight,
my much admired mentor, colleague and
friend, whose collections made in
Egypt and the Yemen Arab Republic
considerably enhanced this work.
...
III
PREFACE
The mosquito fauna of southwestern Asia is limited in taxonomic diversity. Only
98 species are recorded from the region, but at least 25 of these are reportedly involved
in the transmission of pathogens that cause malaria, filariasis, and arboviral diseases of
importance to man. Epidemiological studies and the control of the associated vectors
are extremely limited due to difficulties in making precise species determinations. These
difficulties are largely attributed to the absence of up-to-date taxonomic treatments for
the various species. This work is a contribution toward defining part of the mosquito
fauna of the region. It is a preliminary treatment of the species belonging to the
subgenus Culex. Its primary objective is to provide modern keys, descriptions, and
illustrations for recognizing these species in the adult, pupal, and larval stages.
I consider this work to be preliminary because too few specimens were available
to determine the actual distributions and the total range of morphological variation for
most of the species. Most of the nomenclatural problems have been solved, but many
biosystematic questions require further investigation. The taxonomic status of some
forms remains uncertain. Perhaps the greatest contribution of taxonomic study is not in
filling gaps in our knowledge about species, but in showing how many gaps still exist. If
this manual allows the user to make accurate species determinations, then its primary
purpose will be fulfilled. It is my hope that this work will also serve as a basis for further
research on the included species.
This report is based on data and observations accumulated at the Walter Reed
Biosystematics Unit (WRBU) (housed in the National Museum of Natural History,
Smithsonian Institution) between July 1980 and December 1984. The work began as a
study of Culex pipiens Linnaeus. It was broadened in January 1982 to include all
members of the subgenus Culex in southwestern Asia and Egypt.
I am indebted to many persons for the successful completion of this manual, and I
am most happy to acknowledge their help. I owe much to the stimulating discussions
about systematic problems and taxonomic procedures I had with Bruce A. Harrison, E.L.
Peyton, Curtis W. Sabrosky, F. Christian Thompson, Michael E. Faran, and Thomas J.
Zavortink while we were working together at the Smithsonian Institution. Many
individuals have been most generous in loaning specimens or otherwise assisting me
with the study of collections under their care: Peter S. Cranston, British Museum
(Natural History), London; John Lane, London School of Hygiene and Tropical
Medicine; Francois Rodhain, lnstitut Pasteur, Paris; Jean Clastrier, Museum National
d’t-listoire Naturelle, Paris; A. Rickenbach, Office de la Recherche Scientifique et
Technique Outre-Mer, Bondy; Paul H. Arnaud, California Academy of Sciences, San
Francisco; J. Callot, lnstitut de Parasitologie, Strasbourg; Amnon Freidberg, Tel Aviv
University; Joel Margalit, Ben-Gurion University of the Negev, Beer Sheva; M.W.
Service, Liverpool School of Tropical Medicine; Thomas Gaffigan and James Pecor,
WRBU, National Museum of Natural History, Smithsonian Institution; and staff members
of the Dokki Institute and the Entomological Society of Egypt, Cairo. Sherif el Said, Ain
Shams University, Cairo, and Joel Margalit (affiliation given above) are especially
acknowledged for their special effort in assisting Bruce A. Harrison and myself in
collecting a considerable amount of material in Egypt and Israel, respectively, that was
examined during the study. All of the illustrations were meticulously and skillfully
prepared by Taina Litwak, WRBU. They are a most valuable asset to this work. The
entire manuscript was read critically by Ronald A. Ward. His comments and suggestions
have resulted in many important improvements. Finally, I warmly acknowledge the con-
iv
stant love and support of my wife Roberta, son Paul, and daughter Andrea who patiently
tolerated my use of hundreds of hours of family time to complete this manual.
Bangkok,Thailand
October, 1987
RalphE. Harbach
CONTENTS
Preface ..................................................
Abstract.. ...............................................
Introduction. .............................................
Taxonomy .............................................
Material.. ...........................................
Morphology........................................
Taxa ............................................
Keys ............................................
Speciestreatments .....................................
Type depositories......................................
Illustrations. ..........................................
Keys to genera of Culicidae.....................................
Adu~s...........................................
Pupae ...........................................
La~ae ...........................................
Keysto subgeneraof Culex ....................................
Adults...........................................
Pupae ...........................................
La~ae ...........................................
GenusCulex.............................................lO
SubgenusCulex ........................................
Linear arrangementof includedtaxa ...........................
Distributionallistof the species ..............................
Zoogeographicaffinities .................................
Keysto speciesof Culex (Culex) .................................
Adu~s.............................................13
Pupae.............................................17
Larvae.............................................lg
The speciestreatments. ......................................
Culex (Culex)pipiensLinnaeus .............................
Culex (Culex) quinquefasciatus
Say ..........................
Culex (Culex) vagansWiedemann ...........................
Martini .............................
Culex (Culex) torrentium
Culex(Culex)decensTheobald .............................
Culex(Cdex) antennatus(Becker) ...........................
Culex(Culex)univitfatusTheobald ...........................
Culex (Culex)perexiguusTheobald ..........................
Culex(Culex)thei/eriTheobald .............................
Culex(Culex)laticinctusEdwards ............................
Culex(Culex)mattinglyiKnight ..............................
Culex(Culex)simpsoniTheobald ............................
Culex(Culex)sinaiticusKirkpatrick ...........................
Culex(Culex)&ttoniTheobald .............................
Culex(Culex)sitiensWiedemann ............................
Culex(Culex)poicdipes(Theobald) ...........................
Culex(Culex)mimeticusNoe ...............................
Culex (Culex)bitaeniorhynchus
Giles .........................
i
.
.2
...2
.
...3
...3
...3
4
4
4
5
...5
...5
...6
7
...7
...8
...8
..10
11
11
.12
13
23
23
.3 1
.36
40
44
48
52
57
61
67
72
75
78
82
87
92
97
10 1
Culex (Culex) tritaeniorhynchus
Giles . . . . . . . . . . . . . . . . . . . . . . . . . 106
Culex (Culex)pseudovishnuiColless . . . . . . . . . . . . . . . . . . . . . . . . .111
Referencescited. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 116
Figures . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ...142
Chaetotaxytables . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 186
Appendices . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 227
A. Speciesof Culicidaeotherthan Culex(Culex)known
to occur in southwesternAsia and Egypt . . . . . . . . . . . . . . . . . . .227
B. Complete synonymy of Culex (Culex) pipiens Linnaeus . . . . . . . . . . . 229
C. Complete synonymy of Culex (Culex) quinquefasciatusSay . . . . . . . . .232
D. Complete synonymyof Culex (Culex) sifiensWiedemann . . . . . . . . . . 235
Conspectusof taxonomicchanges . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .236
SystematicIndex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .237
THE MOSQUITOES OF THE SUBGENUS CULEXIN SOUTHWESTERN
ASIA AND EGYPT (DIPTERA: CULICIDAE)
bY
Ralph E. Harbach1f2
ABSTRACT
Twenty species of Culex of the subgenus Culex are recognized in southwestern
Asia and Egypt: pipiens Linnaeus, quinquefasciatus Say, vagans Wiedemann, torrentium Martini, decens Theobald, antennatus (Becker), univiftatus Theobald, perexiguus Theobald, theileri Theobald, laticinctus Edwards, maftinglyi Knight, simpsoni
Theobald, sinaiticus Kirkpatrick, duttoni Theobald, sitiens Wiedemann, poicilipes
(Theobald), mimeficus No&, bitaeniorhynchus Giles, tritaeniorhynchus Giles, and
pseudovishnui Colless. Keys, descriptions, and illustrations are provided for the
identification of the adult, pupal, and larval stages of each species. Keys are also
included for the genera of mosquitoes and the subgenera of Culex occurring in the
region. The individual treatment of each species also includes a complete synonymy, a
summary of previous literature, a systematic discussion, information on bionomics and
distribution, and a synopsis of material examined. Lectotype specimens are designated
for decens, simpsoni, duttoni, and 16 junior synonyms. A neotype specimen is designated for bitaeniorhynchus. Six new junior synonyms are recognized: alpha Seguy
(= theileri); calloti Rioux and Pech, erectus Iglisch, and torridus lglisch (= pipiens);
ethiopicus Edwards (= bitaeniorhynchus); and mauritanicus Callot (= simpsoni). The
recognized species are arranged into three groups and 10 subgroups. The duttoni
group and the decens and simpsoni subgroups are recognized for the first time.
The
‘
opinions or assertions contained herein are the private views of the author and are not to be
construed as official or as reflecting the views of the Department of the Army or the Department
of Defense.
2Walter Reed Biosystematics Unit, Department of Entomology, Walter Reed Army Institute of
Present address: U.S. Army Medical Component,
Research, Washington, DC 20307-5100.
Armed Forces Research Institute of Medical Sciences, APO San Francisco 963465000.
2
Contrib.Amer. Ent. Inst.,vol. 24, no. 1, 1988
INTRODUCTION
The mosquito fauna of southwestern Asia is very poorly known. No modern,
comprehensive treatment exists for any group in any part of the region and most of the
species descriptions and keys are widely scattered through the literature. There are a
number of major works on the mosquitoes of certain countries and specific parts of the
region (Edwards, 1921, 1926; Seguy, 1924; Kirkpatrick, 1925; Monchadskii, 1951;
Mattingly and Knight, 1956; Senevet and Andarelli, 1959; Gutsevitch et al. 1974), but all
of these are limited in scope and treatment, recognize junior synonyms, or include
misidentifications or confused species concepts. The majority of the species known to
occur in the region were originally described from elsewhere.
This work began in July 1980 as a study of Culex pipiens Linnaeus, 1758 in the
Middle East. The study was prompted by the alleged involvement of pipiens in the 1977
and 1978 epizootics of Rift Valley Fever in Egypt. At that time the concept of pipiens in
Egypt was confused by the uncertain status of molestus Forskal, 1775. It was unknown
whether one or more forms identifiable as pipiens may have played a role in
transmission. In January 1982 a decision was made to expand the study to include all
members of the subgenus Culex in southwestern Asia. This decision was made in
response to the need for up-to-date reliable means of identifying medically important
species in this rapidly developing part of the world. The subgenus Culex in this region
includes a number of actual and potential vector species which could not be identified
accurately with existing keys and taxonomic analyses. The decision to expand the study
was made at a time when it was becoming apparent that pipiens in Egypt was no more
than a single behaviorally and physiologicallyvariable species.
Southwestern Asia is defined here as the tract of land south of the USSR
between the Mediterranean Sea and the lndus River of Pakistan (Fig. 1). The area
includes all of Asian Turkey and the Arabian Peninsula. The eastern part of Pakistan
lying along the arid plain of the lndus River is excluded because its mosquito fauna is
almost entirely Oriental. Egypt is included in its entirety although only the Sinai
Peninsula is actually a part of Asia. All of the species of the subgenus Culex found in
Egypt west of the Gulf of Suez also occur in parts of southwestern Asia.
The mosquito fauna of the region consists of only 98 species representing seven
genera and 18 subgenera (Appendix A). The genus Culex is represented by 34
species assigned to seven subgenera. The subgenus Culex includes 20 species.
Most of southwestern Asia lies within the Mediterranean Subregion of the Palaearctic.
The southern part of the Arabian Peninsula belongs to the Afrotropical Region. The
majority of the species of the subgenus Culex have Palaearctic and Afrotropical
affinities. Seven species are clearly Afrotropical, seven Palaearctic, and three Oriental.
Three species are broadly distributed in both Asia and Africa.
TAXONOMY
The composition and organization of this section is unusual. It includes all
information of taxonomic relevance. Explanatory information is given followed by keys to
the genera of mosquitoes and the subgenera of Culex in the region. These are
followed by a brief taxonomic treatment of the subgenus Culex, keys to the species of
the subgenus, and the individual species treatments.
Material. The material examined during this study largely came from the collec-
Harbach: SubgenusCulexin SouthwesternAsia
3
tions of the British Museum (Natural History), the London School of Hygiene and
Tropical Medicine, and the National Museum of Natural History, Smithsonian Institution.
A large proportion of the material in the National Museum of Natural History came from
collections made in Egypt and the Yemen Arab Republic by Kenneth L. Knight
(194951) and in Egypt by Bruce A. Harrison and myself (1983). Important additional
material was examined from the collections of the lnstitut Pasteur (Paris); Office de la
Recherche Scientifique et Technique Outre-Mer (Bondy, France); California Academy
of Sciences; Museum National d’Histoire Naturelle (Paris); lnstitut de Parasitologie,
Faculte de Medicine, Universite de Strasbourg (Strasbourg, France); Dokki Institute
(Cairo); Entomological Society of Egypt (Cairo); and the personal collections of Amon
Freidberg, Tel Aviv University, and Joel Margalit, Ben-Gurion University of the Negev,
Beer Sheva. Some 18,600 specimens (5,129 females, 3,703 males, 1,245 male
genitalia, 4,220 pupal exuviae, 2,576 larval exuviae, and 1,727 fourth-instar larvae) were
examined, including the type specimens of 50 nominal forms. A total of 14,305
specimens (3,571 females, 2,741 males, 874 male genitalia, 3,641 pupal exuviae,
2,123 larval exuviae, and 1,355 fourth-instar larvae) were examined from southwestern
Asia. The remainder (1,558 females, 962 males, 371 male genitalia, 579 pupal exuviae,
453 larval exuviae, and 372 fourth-instar larvae) were examined from neighboring areas,
mainly in Africa.
Morphology.
The morphological terminology recommended by Harbach and
Knight (1980) is used throughout this work. No new terminology is introduced. Terms
for some details of the male genitalia are taken from Harbach (1986 [1987]) and Harbach
et al. (1983 [1984]). The system of lettering the setae borne on the subapical lobe of
the male gonocoxite is adopted from Edwards (1941) and Belkin (1962). Siphon indices
were calculated using the basal width of the larval siphon. Trumpet indices were
determined using the greatest width of the pupal trumpet.
Taxa. The species is the taxonomic unit of primary importance in this study. The
species are arranged into groups and subgroups largely for convenience.
No
subspecies are recognized in this work. Subspecies are largely meaningless biological
concepts which have little recognition in modern culicid taxonomy. Most recent workers
have either synonymized subspecific names with specific epithets or recognized them
for distinct biological species. Users will find that I have followed Sirivanakarn (1976) in
considering Culex quinquefasciatus Say, 1823 as a species distinct from pipiens, even
though these forms apparently interbreed in some areas of southwestern Asia.
Subspecies are not included among the taxa listed in Appendix A because I believe that
descrete populations currently regarded as subspecies in southwestern Asia will
eventually prove to be differentiated species.
Keys. Keys are provided for recognizing the adult, pupal, and larval stages of
the seven genera of mosquitoes, seven subgenera of Culex, and the 20 species of the
subgenus Culex currently known to occur in southwestern Asia. The keys were
prepared especially for the limited fauna of the region (Appendix A) and some species of
the included genera of mosquitoes and subgenera of Culexfrom other areas will not key
out properly. The keys were prepared for specimens in reasonably good condition.
Badly rubbed or damaged specimens will be difficult to identify. The pupal keys for the
subgenera of Culex and the species of Culex (Culex ) were difficult to prepare and are
not as reliable as they appear. Identifications made in the pupal stage should be
confirmed in the adult and larval stages when possible. The keys for adults were
prepared for both males and females. Keys are not included for the male genitalia. Such
keys are normally difficult to use, especially for Culex, and more often than not genitalia
are identified by comparing dissections with published figures. Detailed descriptions
4
Contrib.Amer. Ent. Inst.,vol. 24, no. 1, 1988
and illustrations are provided in lieu of cumbersome keys for the male genitalia.
Species treatments.
The format of the species treatments is traditional. A
complete synonymy is given for each species followed by a summary of previous work,
detailed descriptions of the adult, pupal, and larval stages, a systematic discussion,
information on bionomics and distribution,and a synopsis of material examined. A cross
(+) preceding type data in the synonymies indicates that the type specimen or
specimens of the associated nominal taxon were examined.
Exceptionally long
synonymies are included as appendices. References to previous taxonomic works are
exhaustive for the region. Some ecological papers and most major taxonomic analyses
for the included species in neighboring geographic areas are also included. The
references are listed in chronological order under the biomen, triomen, or other
combination used by the authors. Most of the abbreviations used in the synonymies
and literature summaries are self-evident. The letters A, M, F, P, L, and E indicate adult,
male, female, pupa, larva, and egg, respectively. An asterisk (‘) after one of these letters
indicates that at least part of the life stage was illustrated in the publication cited.
Abbreviations used for type depositories follow the end of this paragraph. The species
descriptions include a brief diagnosis for each life stage. All measurements and counts
are based on at least 10 specimens from southwestern Asia, when available. Diagnostic
characters were checked in most of the positively determined specimens on hand. The
chaetotaxy of the immatures was studied using a differential interference contrast
microscope. Tables are included which summarize the complete chaetotaxy of the pupa
and fourth-instar larva of each species. The names of localities given in the material
examined sections were taken from the labels accompanying the specimens. For the
most part, no attempt was made to determine the modern spellings of the names.
Type depositories. BC - Bigot Collection (in the collection of J.E. Collin, New
Market, Great Britain); BM - British Museum (Natural History), London, Great Britain; DI Dokki Insitute, Cairo, Egypt; ESE - Entomolgical Society of Egypt, Cairo, Egypt; HNM Magyar Nemzeti Muzeum, Budapest, Hungary; IPS - lnstitut de Parasitology, Faculte de
Medicine, Universite de Strasbourg, Strasbourg, France; MM - Macleay Museum of
Natural History, University of Sydney, Sydney, Australia; MNHP - Museum National
d’Histoire Naturelle, Paris, France; NMM - National Museum of Victoria, Melbourne,
Australia; NMNH - National Museum of Natural History, Smithsonian Institution,
Washington, DC, USA; NMW - NaturhistorischesMuseum Vienna, Austria; TMRI - Taiwan
Provincial Malaria Research Institute, Nankang, Taipei County, Taiwan; TMS - Tomsk
Malaria Station, Tomsk, Siberia, USSR; US - School of Public Health and Tropical
Medicine, University of Sydney, Sydney, Australia; ZM - Zoologisches Museum der
Humboldt Universitat, Berlin, East Germany; ZMC - Universitetets Zoologiske Museum,
Copenhagen, Denmark.
Illustrations. The illustrations are original. Drawings are included for the male
genitalia, pupa, and larva of each species. Special features of the adult ornamentation of
some species are illustrated on a single plate. Structures of the male genitalia are
illustrated in a manner consistent with the standard arrangement and positioning
adopted in my earlier works (Harbach, 1985b, 1986 (1987); Harbach et al., 1984, 1985).
This arrangement includes the mesal aspect of the gonocoxopodite, the dorsal aspect
of the intact phallosome, lateral and mesal aspects of the fused lateral plate and
aedeagal sclerite, and dorsal aspects of the proctiger and tergum IX. The pupal and
larval stages are illustrated in the traditional manner.
Harbach: SubgenusCulex in SouthwesternAsia
KEYS TO GENERA
OF CULICIDAE
(Not treated further except for Culex)
ADULTS
1.
Maxillary palpi of female about as long as proboscis;scutellum evenly
rounded; abdominal scales largely or completely absent . . . . Anopheles
Maxillary palpi of female much shorter than proboscis; scutellum
distinctlytrilobed; abdomen completely covered with scales . . . . . . . . . 2
2(l).
Cell R2 of wing shorter than vein R2+3; wing membrane without
distinctmicrotrichia . . . . . . . . . . . . . . . . . . . . . . . . . . Uranotaenia
Cell R2 at least as long as vein R2+3; wing with distinctmicrotrichia . . . . . . . .3
3(2)*
Spiracular setae present; wing with setae present ventrally at base
of subcosta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Culiseta
Spiracular setae absent; subcosta without setae present ventrally . . . . . . . . 4
4(3)*
Tarsomere 1 of fore- and midlegs distinctly longer than tarsomeres
2-5 combined; tarsomere 4 shorter than tarsomere 5 . . . Orthopodomyla
Tarsomere 1 no longer than combined length of tarsmeres 2-5,
usually shorter;tarsomere4 longerthan tarsomere5 . . . . . . . . . . . . . . 5
5(4).
Postspiracular setae present; foreungues toothed. . . . . . . . . . . . . . Aedes
Postspiracularsetae absent;ungues simple . . . . . . . . . . . . . . . . . . . . . 6
6(5).
Hindungues large and conspicuous; pulvilli absent . . . . . . . . Coqulllettldia
Hindungues small and inconspicuous;pulvilli present . . . . . . . . Culex (p. 7)
PUPAE
1.
Seta 9-IV-VII spiniform, arising at or near extreme posterolateral
corner of tergum; trumpet short, flared, meatus with broad
slit extending nearly to base . . . . . . . . . . . . . . . . . . . . . . Anopheles
Seta 9-IV-VII weakly developed, arising distinctly before posterolateral corner of tergum; trumpet otherwise, meatus at most with
narrow slit extending a shortdistancefrom pinna . . . . . . . . . . . . . . . . 2
2(l)*
Trumpet without pinna, apex modified for piercing plant
tissues . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Coquillettldla
Trumpetwith distinctopen pinna . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3(2).
Inner part of paddle deeply excavated toward base, distinctly
widened near middle . . . . . . . . . . . . . . . . . . . . . . . . . Uranotaenia
Inner part of paddle little if at all excavated toward base, slightly
widened (usuallyin distalthird) . . . . . . . . . . . . . . . . . . . . . . . . . . . .4
6
Contrib.Amer. Ent. Inst.,vol. 24, no. 1, 1988
4(3)*
Tracheoid area of trumpet well developed, extending a considerable distance from base; seta 2-P usually present . . . . . . . . . Culex (p. 8)
Tracheoid area absent or at most weakly developed at extreme
base; seta 2-P absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
WL
Seta 9-VIII strongly developed, about 0.5 length of paddle; paddle
somewhat rectangular with buttress distinctly ending beyond
middle of outer margin; seta 1-P branched . . . . . . . . . Orthopodomyia
Seta 9-VIII not as well developed; paddle more or less ovoid, buttress ending before middle of outer margin; seta 1-P simple . . . . . . . . . . 6
Setae 8- and 9-CT in line perpendicularto median keel .........
Seta 8-CT distinctlymore anteriorthan 9-CT .................
Culiseta
Aedes
LARVAE
1.
Siphon absent ................................
Siphon present. ......................................
Anopheles
.2
2(l)*
Spiracular apparatus at apex of siphon modified for piercing plant
tissues, with anterior toothed plate. . . . . . . . . . . . . . . . Coquillettidia
Spiracular apparatus unmodified, comprised of 5 lobes, without
anteriortoothedplate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3(2).
Siphon without pecten ........................
Siphonwith pecten. ....................................
4(3)*
Head longer than wide; comb scales arising from a sclerotized
plate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Uranotaenfa
Head at least as wide as long; comb plate absent, scales arising
from unsclerotizedintegument . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
WV*
Siphon with at least 3 pairs of seta I-S, usually more .........
Siphonwithasinglepairof setal-S ..........................
6(5).
Seta 1-S arising near base of siphon. ....................
Seta 1-S arising near last pecten spine in middle part of siphon ......
Otthopodomyia
4
Culex (p. 8)
.6
Culiseta
Aedes
Harbach: SubgenusCulexin SouthwesternAsia
KEYS TO SUBGENERA
7
OF CULEX
(Not treated further except Culex)
ADULTS
1.
No lower mesepimeral setae; probosciswith pale ring in
middle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Culex (in part,p. 13)
One or more lower mesepimeral setae; proboscis either
entirelydark or pale beneath . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2(f)
Three or more lower mesepimeral setae and prealar scales present;
occurring in southern part of Arabian Peninsula . . . . . . . . . . . . . Lutzia
One to 4 lower mesepimeral setae present; prealar scales present
or absent; normally no prealar scales if 3 or 4 lower mesepimeral
setae are present. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3w
Proboscis shorter than forefemur; hindtarsomere 1 short, not more
than 0.85 length of hindtibia . . . . . . . . . . . . . . . . . . . . . . Barraudius
Proboscis longer than forefemur; hindtarsomere 1 usually as long
or only slightly shorter than hindtibia, not less than 0.85 length
of hindtibia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4(3)
Abdominal terga with basal pale bands or basolateral pale
spots (lateral pale stripes on terga VI and VII in
antenna&s). . . . . . . . . . . . . . . . . . . . . . . . . . . Culex (in part, p. 13)
Abdominal terga with apical pale bands, apicolateral pale
spotsor entirelydark . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
S(4)
Decumbent scales of vertex mainly narrow and dark, those on ocular
line broad and white; scutal scales entirely dark and very fine;
occurring in southern part of Arabian Peninsula . . . . . . . . . Culiciomyia
All decumbent scales of vertex narrow and pale; scutal scales
predominently pale and rather coarse (except in
Neowlex martjnjo. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .6
W2
Prealar, and usually postspiracular,scales present . . . . . . . . . . . . Maillotia
Prealar and postspiracularscales absent . . . . . . . . . . . . . . . . . . . . . . . .7
WI
Small yellowish-brown species; apical markingsof abdominal terga
yellowish and indistinct against pale brown background;
maxillary palpi of male shorterthan proboscis . . . . . . . . . . Lasiosiphon
Larger, darker species; apical markings of abdominal terga white
and well contrasted against black or nearly black background;
maxillary palpi of male longer than proboscis . . . . . . . . . . . . . Neoculex
Contrib.Amer. Ent. Inst.,vol. 24, no. 1, 1966
PUPAE
2
.5
1.
Seta 6-III-VI usuallyall single................................
Seta 6-III-VI usually all branched (except sometimesduttoni) ...........
2(l)*
All or most of setae l-g-CT single; seta 9-VIII well removed cephalad
from caudolateral angle of tergum, about 0.7 from base . . . . . . . . . Lutzia
All or most of setae l-g-CT branched; seta 9-VIII immediately
adjacentto caudolateralangle of tergum . . . . . . . . . . . . . . . . . . . . . . 3
W).
Seta 5-CT very long, longer than trumpet; paddle distinctly
notchedat tip . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Culiciomyia
Seta 5-CT short; paddle evenly rounded or only slightly
notchedat tip . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4(3).
Trumpet short and broad, length less than 0.6 mm, index about
3.0; setae 1-IX and 2-P absent . . . . . . . . . . . . . . . . . . . Lasiosiphon
Trumpet longer, length normally more than 0.6 mm, index more
than 3.0; setae l-IX and 2-P usually present . . . . . . . Culex (in part, p. 17)
5(l).
Seta 5-V,VI at least nearly as long as 2 terga following, usually
as tong,sometimeslonger. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .6
Seta 5-V,VI 1.5 or less length of tergum following (except in
&Ron?, usuallyless . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
6(5).
Trumpet long and slender, index more than 7.0; seta 1-P not
much stronger than 2-P, about twice as long . . . . . . . . . . . . . Neoculex
Trumpet not as long and slender, index less than 7.0; seta 1-P
clearly strongerthan 2-P, about four times as long . . . . . . . . . . Maillotia
W).
Trumpet short and broad, length less than 0.6 mm, index
about 4.0 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Barraudius
Trumpet variable, length and index usually more than indicated,
if length less than 0.6 mm then index more than 4.0 . . Culex (in part, p. 17)
LARVAE
1.
Lateral palatal brushes developed for predation; saddle longer than
siphon; pecten extending to apex of siphon . . . . . . . . . . . . . . . Lutzia
Lateral palatal brushes developed for filter-feeding; saddle shorter
than siphon; pecten not extending to apex of siphon. . . . . . . . . . . . . . 2
w 1.
Ventral brush (seta 4-X) with 4 pairs of setae; antenna shorter than
0.5 length of head; siphon with lateral pair of transverselyaligned setae adjacent to pecten . . . . . . . . . . . . . . . . . . . Culiciomyia
Ventral brush with 5 or more (usually more) pairs of setae; antenna
longer than 0.5 length of head; siphon without lateral pair of
transversely-alignedsetae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
Harbach: SubgenusCulex in SouthwesternAsia
9
3(2).
Setae 5,6-C very short and inconspicuous, about same thickness
and length as seta 4-C . . . . . . . . . . . . . . . . . . . . . . . . Lasiosiphon
Setae 5,6-C long and conspicuous, much thicker and longer than
seta4-C..........................................4
4(3).
Ventral brush with 1 or more setae borne anteriorto grid . . . . . . . . . . . . . . 5
Ventral brushwith all setae borne on grid . . . . . . . . . . . . . . . . . . . . . . . 6
5(4).
Seta 3-P nearly length of setae 1,2-P; siphon with 2 or more
(usually more) anterolateral setae on each side; seta 1-S
unpaired, occurring in single median posteriorrow. . . . . . . . . . Maillotia
Seta 3-P much shorter than 1,2-P, about 0.5 as long; siphon
without anterolateral setae; seta 1-S more or less paired,
occurringin 2 posterolateralrows . . . . . . . . . . . . . . . . . . . . Neoculex
6(4).
Seta 1-S in single row, with all elements arranged in more or less
straightline . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Bamudius
Seta 1-S in 1 or 2 rows, with l-3 elements distinctly out of line with
the others (inserted laterally) . . . . . . . . . . . . . . . . . . . . . Culex (p. 19)
10
Contrib.Amer. Ent. inst.,vol. 24, no. 1, 1988
Genus Culex Linnaeus
Subgenus Culex Linnaeus
Culex Linnaeus, 1758: 602. Type species: pipiens Linnaeus, 1758: 602
(subsequent designation, Latreille, 1810: 442).
Heteronycha
Lynch Arribalzaga, 189la: 373; 189lb: 155. Type
species: dolosa Lynch Arribalzaga (monotypy).
Lasioconops
Theobald, 1903a: 236. Type species: poicilipes
Theobald (monotypy).
Heptaphlebomyia
Theobald, 1903a: 336. Type species: simplex
Theo bald (monotypy).
Pseudoheptaphlebomyla
Ventrillon, 1905b:427.
Type species:
madagascariensis Ventrillon (monotypy).
Trichopronomyia
Theobald, 1905b: 98. Type species: annulata
Theobald (monotypy).
Aporoculex Theobald, 1907: 316. Type species: punctipes Theobald
(monotypy).
Leucomyia
Theobald, 1907: 372. Type species: gelidus Theobald
(original designation).
Oculeomyia Theobald, 1907: 515. Type species: sarawaki Theobald
(monotYPY)*
Theobaldiomyla
Brunetti, 1912: 462. Type species: Culex gelidus
Theobald, 190la: 20 (fixed type of a replacement name).
Phalangomyia Dyar and Knab, 1914: 58. Type species: debilis Dyar
and Knab (monotypy).
Laiomyia Izquierdo, 1916: 65. Type species: Culex stigmatosoma Dyar,
1907: 123 (subsequent designation, Edwards, 1932a: 200).
Culex subgenus Transculicia
Dyar, 1917( 1918): 184. Type species:
eleuthera Dyar (monotypy).
Culex subgenus Cacoculex Dyar, 1918: 100. Type species: habilitator
Dyar and Knab, 1906: 212 (original designation).
The subgenus Culex is a large and diverse group with more than 200 species
which are distributed throughout the world. The concept of the subgenus defined by
Edwards (1932a, 1941), Belkin (1962)) Bram (1967a), Sirivanakarn (1976), and Tanaka
et al. (1979) is followed here although this concept will probably change once the
phylogenetic relationships of the species are understood. These authors have
adequately diagnosed the subgenus and it does not need to be characterized here.
The subgenus Culex in southwestern Asia is represented by 20 nominal
species which fall into three groups and 10 subgroups. The groups, subgroups, and
species are arranged and treated in the order indicated in the chart below. The species
and subgroups, with the exception of the trifilatus subgroup,
are arranged
chronologically within their respective higher categories. The species are placed in
subgroups based on presumptive phylogenetic relationships. I have made no special
effort to determine natural affinities and little is said or conjectured about evolutionary
relationships between species or groups of species. My primary aim has been to
provide a strong descriptive base for the identification and future study of each species.
The supraspecific groups are not described. They are adequately defined by the
shared characters noted for the included species.
Harbach: SubgenusCulex in SouthwesternAsia
Linear
Arrangement
of Included
Taxa
pipiens group (Edwards, 1932a: 201; excluding duttoni
Theobald)
pipiens subgroup (Sirivanakarn, 1976: 29)
1) pipiens Linnaeus, 1758 (p. 23)
2) quinquefasciatusSay, 1823 (p. 31)
trifilatussubgroup (Mattingly and Rageau, 1958: 241)
3) vagans Wiedemann, 1828 (p. 36)
4) torrentiumMartini, 1925 (p. 40)
decens subgroup (New Subgroup)
5) decens Theobald, 1901c (p. 44)
6) antennatus (Becker, 1903) (p. 48)
univittatus subgroup (Sirivanakarn, 1976: 47)
7) univittatusTheobald, 1901 b (p. 52)
8) perexiguus Theobald, 1903a (p. 57)
theileri subgroup (Sirivanakarn, 1976: 43)
9) thei/eriTheobald, 1903a (p. 61)
10) laticinctusEdwards, 1913a (p. 67)
11) matting/Vi Knight, 1953b (p. 72)
simpsoni subgroup (New Subgroup)
12) simpsoni Theobald, 1905c (p. 75)
13) sinaiticus Kirkpatrick, 1924( 1925) (p. 78)
duttoni group (New Group)
14) duttoni Theobald, 1901 c (p. 82)
sitiens group (Edwards, 1932a: 201)
sitiens subgroup (sitiens series of Edwards, 1932a: 201;
including poicilipes (Theobald))
15) sitiens Wiedemann, 1828 (p. 87)
16) poicilipes (Theobald, 1903b) (p. 92)
mimeticus subgroup (mimeticus series of Edwards,
1932a: 201)
17) mimeticus Noe, 1899 (p. 97)
bitaeniorhynchus subgroup (bitaeniorhynchus series of
Edwards, 1932a: 201, excluding poicilipes
(Theobald))
18) bitaeniorhynchus Giles, 1901 a (p. 101)
vishnui subgroup (Bram, 1967a)
19) tritaeniorhynchusGiles, 1901 a (p. 106)
20) pseudovishnui Colless, 1957 (p. 111)
Distributional
List of the Species
Too few collections have been made in most areas of southwestern Asia to plot
the distributions of the species accurately and completely. Members of the subgenus
Culex known to occur in the region are listed by country below. The list is compiled from
material examined and literature records. Species mistakenly or doubtfully recorded in
the literature are not included.
12
Contrib.Amer. Ent. Inst.,vol. 24, no. 1, 1988
AFGHANISTAN:
Culex perexiguus, pipiens, quinquefasciatus, theileri, and
vagans.
BAHRAIN : Culex quinquefasciatus.
EGYPT:
Culex antennatus, laticinctus, mimeticus, perexiguus, pipiens,
poicilipes, sinaiticus, theileri, and tritaeniorhynchus.
IRAN: Culex antennatus, bitaeniorhynchus, laticinctus, mimeticus, perexiguus,
pipiens, pseudovishnui, quinquefasciatus, sinaiticus, Miens, theileri, torrentium,
tritaeniorhynchus, and vagans.
IRAQ: Culex antennatus, laticinctus, mimeticus, perexiguus, pipiens,
pseudovishnui, quinquefasciatus, theileri, torrentium, and tritaeniorhynchus.
ISRAEL: Culex antennatus, laticinctus, mimeticus, perexiguus, pipiens,
poicilipes, sinaiticus, theileri, and tritaeniorhynchus.
JORDAN: Culex antennatus, laticinctus, mimeticus, perexiguus, pipiens,
sinaiticus, theileri, and tritaeniorhynchus.
KUWAIT: Culex quinquefasciatus.
LEBANON: Culex laticinctus, mimeticus, perexiguus, pipiens, theileri, and
tritaeniorhynchus.
OMAN: Culex laticinctus, perexiguus, quinquefasciatus, sinaiticus, sitiens, and
tritaeniorhynchus.
PAKISTAN (west of lndus River): Culex bitaeniorhynchus, mimeticus,
perexiguus, pseudovishnui, quinquefasciatus, sitiens, theileri, tritaeniorhynchus, and
vagans.
P. D. R. Y EM EN : Culex bitaeniorhynchus, decens, duttoni, laticinctus, mattinglyi,
pipiens, quinquefasciatus, sinaiticus, sitiens, theileri, tritaeniorhynchus, and univittatus.
QATAR : Culex quinquefasciatus.
Culex laticinctus, mattinglyi, perexiguus,
pipiens,
SAUDI ARABIA:
quinquefasciatus, sinaiticus, sitiens, theileri, and tritaeniorhynchus.
SYRIA:
Culex laticinctus, mimeticus, perexiguus, pipiens, theileri, and
tritaeniorhynchus.
TURKEY: Culex antennatus, laticinctus, mimeticus, perexiguus, pipiens, theileri,
torrentium, tritaeniorhynchus, and vagans.
UN ITED ARAB EM IRATES: Culex pipiens, quinquefasciatus, and sitiens.
YEMEN ARAB REPUBLIC:
Culex bitaeniorhynchus, decens, laticinctus,
mattinglyi, pipiens, quinquefasciatus,
simpsoni, sinaiticus, sitiens, theileri,
tritaeniorhynchus, and univittatus.
Zoogeographic
Affinities
The Yemen Arab Republic, the P.D.R. Yemen, and the southwestern part of
Saudi Arabia belong to the Afrotropical Region. The rest of southwestern Asia and
Egypt lie within the southern Palaearctic. The known distributions of the included
species can be examined to determine their zoogeographic affinities. Culex decens,
univittatus, mattinglyi, simpsoni, and duttoniare strictly Afrotropical species. All of these
species except mattinglyi are widely distributed south of the Sahara and occur in the
Afrotropical part of the Arabian Peninsula. Culex mattinglyi is endemic to southwestern
Asia. Culex antennatus and poicilipes are also widely distributed in sub-Saharan Africa,
but they do not occur in the Afrotropical part of Arabia. Culex poicilipes extends
northward along the Nile River into Egypt and Israel. Culex antennatus is common in
Harbach: Subgenus Culexin SouthwesternAsia
13
lands around the eastern end of the Mediterranean. Seven Afrotropical species are
present in the region.
Culex theileri appears to be a southern Palaearctic species which has spread
southward through the temperate highlands of eastern Africa and eastward through
India. Both laticinctus and sinaiticus are mainly Palaearctic with limited distributionsin the
Afrotropical Region. Culex perexiguus is very widely distributed across northern Africa
and southwestern Asia and extends eastward into the Oriental Region. This species is
apparently replaced by univittatus in southwestern Arabia. Culex mimeticus occurs in a
relatively narrow zone which extends from the shores of the Mediterranean, through the
Middle East, and beyond. Populations of this species in eastern Asia occur in both the
Palaearctic and Oriental regions. Culex vagans is widespread in the northeastern
Palaearctic. Its known range includes southern China and India in the Oriental Region
and northern countries of southwestern Asia. Culex torrentium is the only species of
the subgenus in southwestern Asia which is exclusively Palaearctic. Its range lies mainly
north of southwestern Asia. A total of seven Palaearctic species occur in the region.
Oriental species in southwestern Asia include sitiens, trifaeniorhynchus, and
pseudovishnui. Culex bitaeniorhynchus perhaps should be included among these
species, but its wide distribution in the Oriental Region is offset by its nearly equally
broad distribution in the Afrotropical Region. Culex tritaeniorhynchus is very widely
distributed in southwestern Asia, but the main part of its range lies within the Oriental
Region. Isolated populations occur in Egypt and it is known from certain coastal areas of
eastern and western Africa where it was probably introduced by man. Both sitiens and
pseudovishnui have limited distributions in the region. The former is a coastal species
which extends from the eastern coast of Africa to the South Pacific.
Culex
pseudovishnui is an indigenous member of the Oriental fauna which has spread into the
eastern part of the region.
Culex pipiens and quinquefasciatus are common widespread sister species
whose distributions undoubtedly have been influenced by man. Their natural ranges
cannot be known for certain. Culex pipiens is very widely distributed in the Palaearctic
and eastern areas of the Afrotropical Region. This species may have evolved in Africa
and dispersed to more northern latitudes. It is replaced at lower elevations in warmer
areas by quinquefasciatus. Culex quinquefasciatus is cosmotropical.
KEYS
TO SPECIES
OF
CULEX
(CULEX)
ADULTS
1.
One or more lower mesepimeral setae present (Fig. 2A) (note
presence of alveoli if setae are missing); proboscis and tarsi
withoutpale rings(except dutiond. . . . . . . . . . . . . . . . . . . . . . . . .2
No lower mesepimeral setae (note absence of alveoli); proboscis
with median pale ring; tarsi with narrow pale rings (sitiens group) . . . . . 18
2U )*
Tarsi with narrow pale rings (duttoni group); midtibia with anterior
pale stripe (Fig. 2H); postspiracularscales present . . . . . duttoni (p. 82)
Tarsi without pale rings (pipiens group); midtibia with or without
anterior pale stripe; postspiracularscales present or absent . . . . . . . . .3
14
Contrib.Amer. Ent. Inst.,vol. 24, no. 1, 1988
pipiens group
W).
Postspiracularand prealar scales present (Fig. 2A). ...............
No postspiracularscales. ................................
.4
8
All tibiae with anterior pale stripes (Fig. 2G); prealar and upper
and lower mesokatepisternal scale-patches confluent
(Fig. 2B); basal pale bands of abdominal terga produced
posteriorlyin middle . . . . . . . . . . . . . . . . . . . . . . . . theileri (p. 61)
Fore- and midtibiae normally without anterior pale stripes
(univittatus and perexiguus usually have an anterodorsal
pale stripe on the midtibia); prealar and upper and lower
mesokatepisternal scale-patches separate; basal pale
bands of abdominal terga not produced posteriorlyin middle . . . . . . . . 5
W).
Hindtibia with anterior pale stripe (Fig. 2K); wing with short line
ofpalescalesatbaseofcosta . . . . . . . . . . . . . . . . . . . . . . . . . . .6
Hindtibiawithout anterior pale stripe;wing entirely dark-scaled . . . . . . . . . 7
w-
Anterior surface of midfemur with complete narrow pale stripe
(Fig. 21);wing usually with row of scales on vein 2A ( an
unique character among mosquitoes) (Fig. 2D); occurring
in southern part of Arabian Peninsula . . . . . . . . . . univittatus (p. 52)
Anterior surface of midfemur entirely dark-scaled or with
incomplete narrow pale stripe; wing seldom with few
scales on vein 2A; occurring throughout southwestern
Asia and Egypt . . . . . . . . . . . . . . . . . . . . . . . perexiguus (p. 57)
7(5).
Hindfemur with complete or nearly complele anterodorsal
dark stripe (Fig. 2L); females: abdominal sterna with
apical dark bands; forecoxa with some dark scales;
males: abdominal sterna mainly dark-scaled, with
basolateral pale spots: maxillary palpus normal . . . . . . simpsoni (p. 75)
Hindfemur with anterodorsal dark stripe on distal 0.5 or less
(Fig. 2M); females: abdominal sterna usually entirely
pale-scaled; scales of forecoxa usually all pale; males:
abdominal sterna mainly pale-scaled, posterior sterna
usually with dark scales posteriorly;maxillary palpus
sparselysetose . . . . . . . . . . . . . . . . . . . . . . . . sinaiticus (p. 78)
8~3).
Prealarscalespresent(Fig. 2C) ............................
Prealarscalesabsent ..................................
WV
Thorax pale; wing with short line of pale scales at base of costa;
scales of forecoxa all pale; frequently with 2-4 lower
mesepimeralsetae . . . . . . . . . . . . . . . . . . . . . . laticinctus (p. 67)
Thorax darker; wing entirely dark-scaled; forecoxa with some
dark scales; normally with only one lower mesepimeral
seta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . tonwtium (p. 40)
9
10
Harbach: SubgenusCulex in SouthwesternAsia
15
1O(8).
Fore- and midfemora and all tibiae with anterior pale stripes
(Fig. 2G). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . vagans(p.36)
These with anteriorsurfacesentirelydark-scaled . . . . . . . . . . . . . . . . .l 1
ll(l0).
Abdominal sterna with apical dark bands; scutal scales with
distinct reddish tint; occurring in southern part of Arabian
Peninsula. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . decens (p. 44)
Abdominal sterna unbanded, dark scales confined to midline
and/or posterolateral corners when present; color of scutal
scalesvariable . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .12
12(11).
Abdominal terga unbanded, terga VI and VII with lateral pale
stripe; proboscis of male with ventral cluster of setae at
false joint . . . . . . . . . . . . . . . . . . . . . . . . . . antennatus (p. 48)
Abdominal terga with basal pale bands; proboscisof male
without ventral clusterof setae at false joint . . . . . . . . . . . . . . . . . . 13
13(12).
Two to 4 lower mesepimeral setae present (note presence of
alveoli if setae are missing). . . . . . . . . . . . . . . . . . laticinctus (p. 67)
Only one lower mesepimeralseta present . . . . . . . . . . . . . . . . . .14
14(13).
Scales of forecoxa all pale; wing with short line of pale scales
at base of costa; proboscisall dark or faintly pale beneath. . . . . . . . . . 15
Forecoxa with some dark scales; wing entirely dark-scaled;
proboscisusually distinctlypale beneath in middle . . . . . . . . . . . . . 16
15(14).
Thoracic integument pale; scutal scales light yellowish brown;
scutum with short diagonal line of whitish scales along
prescutal suture; dorsal dark stripe of hindfemur beginning
beyond base and expanded over distal 0.2-0.3 of anterior
surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . laticinctus(p. 67)
Thoracic integument and scutal scales darker; scutum with
curved line of pale scales along prescutal suture and
posterior dorsocentral setae; dorsal dark stripe of
hindfemur beginning at base and expanded over
distal 0.3-0.4 of anterior surface . . . . . . . . . . . . . . mattinglyi (p. 72)
16(14).
Scutal scales more or less buff-colored; basal bands of
abdominal terga nearly white, usually slightly paler than
sternal scaling; females: tergal bands slightly if at all
darker than basolateral white spots, subcosta normally
intersects costa before furcation of R2+3; males: dorsal
arms of phallosome nearly parallel in dorsal aspect
(Fig. 5), DV/D usually more than 0.6*. . . . . . qulnquefasciatus
(p. 31)
*These characters are unreliable for separating pipiens and quinquefasciatus
in central and
northeastern areas of the Arabian Peninsula where introgressed populations (hybrids) occur.
16
Contrib.Amer. Ent. Inst.,vol. 24, no. 1, 1988
Scutal scales golden brown with reddish tint; basal bands of
abdominal terga yellowish, usually same color as sternal
scaling; females: tergal bands distinctly darker than basolateral white spots; subcosta normally intersects costa
at or beyond furcation of R2+3; males: dorsal arms of
phallosome divergent in dorsal aspect (Figs. 3, 9);
DV/D usuallyless than zero* . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
17( 16).
Females: cell R2 more than 4.0 length of vein R2+3; integument
and scales between supraala and posterior dorsocentral
setae usually noticeably darker than surrounding integument
and scales, evident as pair of ovoid spots; males: cell R2
2.4-3.6 length of vein R2+3; dorsal arm of phallosome
truncate in dorsal aspect, lateral plate without apicolateral
spiculate lobe (Fig. 3) . . . . . . . . . . . . . . . . . . . . . . . pipiens (p. 23)
Females: cell R2 less than 4.0 length of vein R2+3; integument
and scales between supraalar and posterior dorsocentral
setae not appreciably darker than surrounding integument
and scales; males: cell R2 2.0-2.5 length of vein R2+3;
dorsal arm of phallosome pointed in dorsal view, lateral
plate with apicolateral spiculate lobe (Fig. 9) . . . . . . . torrentium (p. 40)
sitiens group
18(l).
Wing with pattern of pale spots (Fig. 2E); midtibia with anterior
pale stripe(Fig. 2H) . . . . . . . . . . . . . . . . . . . . . . mimeticus (p. 97)
Wing either speckled, entirely dark-scaled or with variable
amount of pale scaling along anterior veins; no anterior
pale stripeon midtibia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .19
19(18)
Anterior surfaces of femora and tibiae with rows of small pale
spots (Fig. 2J); lower proepisternal scales present . . . poicilipes (p. 92)
Femora and tibiae without rows of pale spots; lower
proepistemalscales absent . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
20(19)
Abdominal terga with apical pale bands; wing with numerous
scattered pale scales (Fig. 2F); probosciswith pair of
dorsolateral pale spots before labella. . . . . bitaeniorhynchus (p. 101)
Withoutthese characters. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .21
21(20)
Femora with numerous scattered pale scales on anterior surfaces;
scutum with indefinite mottled pattern; cell MI long, furcation
of vein Ml+2 proximal to furcation of vein R2+3 . . . . . . . . sitiens (p. 87)
Withoutthese characters . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22
These
‘
characters are unreliable for separating pipiens and quinquefasciatus
in central and
northeastern areas of the Arabian Peninsula where introgressed populations (hybrids) occur.
Harbach: Subgenus Culex in SouthwesternAsia
22(21).
17
Females: pale ring of proboscis extended proximally on ventral
surface*; forked scales of vertex dirty yellow to brown in
middle; cell R2 more than 3.0 length of vein R2+3; males:
probosciswith ventral cluster of 10 or more setae at false
joint; forked scales as in female. . . . . . . . . tritaeniorhynchus (p. 106)
Females: pale ring of proboscis without proximal extension on
ventral surface; forked scales of vertex white to yellow in middle;
cell R2 less than 3.0 length of vein R2+3; males: proboscis
without ventral cluster of setae (1 or 2 setae sometimes
present); forked scales as in female . . . . . . . . pseudovishnui (p. 111)
PUPAE
1.
Median area of scutum, metanotum and terga I and II with conspicuous vesicles; seta 5-VI as long as terga VII and VIII
combined, usually single (Fig. 29). . . . . . . . . . . . . . . . duttoni (p. 82)
Median area of scutum, metanotum and terga I and II without
vesicles; seta 5-W distinctly shorter than combined length
of terga VII and VIII, normallydouble or triple . . . . . . . . . . . . . . . . . . 2
2(l)-
Seta 8-CT usually with 3 or more branches, seldom double; seta
6-III-VI also usually with 3 or more branches, sum of their branches on one sideof abdomen equals gormore, usually more . . . . . . .3
Set 8-CT usually double, rarely single or triple; seta 6-III-VI normally single or double, sum of their branches on one side
of abdomen not exceeding 8 . . . . . . . . . . . . . . . . . . . . . . . . . . .14
pipiens group
3(2)*
Seta 6-I,11short, not much longer than 7-l,ll; trumpet strongly
flared, index normally less than 4.0; tracheoid area short,
0.3orlessoftrumpetlength . . . . . . . . . . . . . . . . . . . . . . . . . . . .4
Seta 6-I,11long, at least twice as long as 7-l,ll; trumpet not so
strongly flared, index normally more than 4.0; tracheoid
area long, usually more than 0.3 of trumpet length . . . . . . . . . . . . . . 5
4(3).
Seta 6-l single or double; seta 5-11with 4-7 branches (usually 5);
seta 4-VIII double or triple (Fig. 21) . . . . . . . . . . . . . laticinctus (p. 67)
Seta 6-l with at least 3 branches; seta 5-11double; seta 4-VIII
single (Fig. 23) (based on a single specimen) . . . . . . mattinglyi (p. 72)
5(3).
Trumpet sharply bent at distal end of tracheoid area; seta 5-IV
at least 1.5 length of tergum V (Fig. 7) . . . . . . . . . . . . . vagans (p. 36)
Trumpet gently curved; seta 5-IV less than 1.5 length of
tergumV........................................6
*In many specimens, the ventral extension is weak or absent in the middle and the proboscis bears
an isolated spot proximal to the ring.
18
Contrib.Amer. Ent. Inst.,vol. 24, no. 1, 1988
6(5).
Setae 6-111
and 3-VII normally double or triple, sum of their branches
on both sides of abdomen rarely exceeding 12 . . . . . . . . . . . . . . . .7
Setae 6-111
and 3-VII normallywith 3-5 branches, sum of their
branches on both sides of abdomen rarely 12 or less . . . . . . . . . . . . 10
7(6).
Seta l-11with at least 25 branches, usually more; trumpet long and
slender, index usually more than 6.0 (Fig. 11); occurring in
southern part of Arabian Peninsula . . . . . . . . . . . . . . decens (p. 44)
Seta 1-II with fewer than 25 branches, usually fewer than 20;
trumpet moderate, index usually less than 6.0 . . . . . . . . . , . . . . . . .8
8(7).
Seta 5-VI at least 1.5 length of tergum VII; tracheoid area usually
more than 0.36 of trumpet length; pinna short, usually less
than 0.25 of trumpet length (Fig. 9); occurring in Turkey,
(?) Iran and (?) Iraq . . . . . . . . . . . . . . . . . . . . . . torrentium (p. 40)
Seta 5-VI less than 1.5 length of tergum VII; tracheoid area
usually less than 0.36 of trumpet length; pinna longer,
normally more than 0.25 of trumpet length; more
widespreadspecies . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
g(8).
Trumpet distinctly flared, index less than 5.0; pinna long, about
0.4 or more of trumpet length; seta 8-CT usually triple*
(Fig. 5). . . . . . . . . . . . . . . . . . . . . . . . . quinquefasciatus (p. 31)
Trumpet widened very little distally, index more than 5.0;
pinna shorter, usually less than 0.35 of trumpet length;
seta 8-CT with 4-7 branches* (Fig. 3) . . . . . . . . . . . . . pipiens (p. 23)
1O(6).
Caudolateral angle of tergum VIII bluntly rounded; trumpet
lightly to moderately tanned, length usually more than
0.58 mm; seta 4-CT usuallywith 3 or 4 branches . . . . . . . . . . . . . . . 11
Caudolateral angle of tergum VIII produced and acute, if not
produced, then trumpet heavily tanned and short, usually
less than 0.58 mm, and seta 4-CT normallydouble . . . . . . . . . . . . . 16
ll(l0)
Seta 1-II dendritic, usually with fewer than 12 branches;
trumpet flared, length more than 0.7 mm (Fig. 19) . . . . . . theileri (p. 61)
Seta 1-II fanlike, with 12 or more branches (usually more);
trumpet almost cylindrical or very little widened distally,
lengthusuallyless than 0.7 mm . . . . . . . . . . . . . . . . . . . . . . . . . 12
12(11)
Seta 5-IV shorter than l-IV; seta 5-V,VI about as long as
following tergum; paddle ovoid, inner part not noticeably
narrowed toward base (Figs. 15,17) . . . . . . . . . . . univittatus (p. 52)
perexiguus (p. 57)
*These characters are unreliable in central and northeastern areas of the Arabian Peninsula where
introgressed populations (hybrids) occur.
Harbach:SubgenusCulex in SouthwesternAsia
19
Seta 5-W at least as long as 1-IV, usually longer; seta 5V,VI about
1.5 or more length of following tergum; paddle more or less
pear-shaped, inner part distinctlynarrowed toward base . . . . . . . . . . 13
13(12).
Abdominal length usually more than 2.6 mm; paddle length
usually more than 0.8 mm; pinna of trumpet usually longer
than 0.16 mm (Fig. 13). . . . . . . . . . . . . . . . . . . . antennatus (p. 48)
Smaller species, these measurements usually less than
indicated (Figs. 25,27) . . . . . . . . simpsoni (p. 75); sinaiticus (p. 78)
sitiens group
14(2).
Trumpet very strongly flared, funnel-shaped; pinna long, more
than 0.36 mm; setael 1-CT, 3-I-111and 1-VII normally single
(Fig. 37) . . . . . . . . . . . . . . . . . . . . . . . bitaeniorhynchus (p. 101)
Trumpet not so strongly flared; pinna shorter, less than
0.36 mm; setae 1l-CT, 3-I-111
and l-VII usuallydouble . . . . . . . . . . . . 15
15(14).
Seta 1-II with l-4 branches; setae 6-III-VI usually single;
seta 9-VIII with 4-9 branches (Fig. 35). . . . . . . . . . . . mimeticus (p. 97)
Seta l-11with 4-10 branches; setae 6-III-VI usually double;
seta 9-VIII with 9-14 branches (Fig. 31) . . . . . . . . . . . . . sitiens (p. 87)
16(10).
Paddle with conspicuous dark spot on inner part; trumpet
length more than 0.75 mm; paddle length more than
0.87 mm; seta l-VI with 3 or 4 branches (Fig. 33). . . . . polcilipes (p. 92)
Paddle without dark spot on inner part; trumpet length less
than 0.75 mm; paddle length less than 0.87 mm;
seta l-VI with5-9 branches. . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
17(16).
Seta l-11with more than 18 branches; seta 8-CT with 6 or 7
branches; caudolateral angle of tergum VII bluntly
rounded (Fig. 39) . . . . . . . . . . . . . . . . . tritaeniorhynchus (p. 106)
Seta 1-II with fewer than 18 branches; seta 8-CT with
3-5 branches; caudolateral angle of tergum VIII
acute. . . . . . . . . . . . . . . . . . . . . . . . . . . pseudovishnui (p. 111)
LARVAE
1.
Siphon strongly swollen, sides markedly convex, with subapical
dark band and short spine anteriorly near apex (Fig. 30) . . duttoni (p. 82)
Siphonotherwise . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2(l).
All scales of comb evenly fringed at sides and apex . . . . . . . . . . . . . . . .3
Some or all scales of comb spinelike, with pointed apex
and fringe at sides proximally. . . . . . . . . . . . . . . . . . . . . . . . . . . 13
3(2)*
Distal pecten spines with 2-5 ventral denticles of different sizes
arising proximally; seta 1-C thin, scarcely if at all thicker than
20
Contrib.Amer. Ent. Inst.,vol. 24, no. 1, 1988
branchesof setae 5,6-C . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
Distal pecten spines with 7 or more ventral denticles of similar
size arising along entire length; seta 1-C stouter, distinctly
thickerthan branchesof setae 5,6-C . . . . . . . . . . . . . . . . . . . . . . 16
pipiens group
4(3).
Siphon very long and slender, index more than 8.5; seta 1-IV,V
long and single, much longer than length of 2 abdominal
segments (Fig. 12) . . . . . . . . . . . . . . . . . . . . . . . . decens (p. 44)
Siphon not as long and slender, index less than 8.5; seta 1-IV,V
shorter, less than length of 2 abdominal segments, with
one or more branches,usually more. . . . . . . . . . . . . . . . . . . . . . . 5
5(4).
Siphon with 6-8 pairs of seta 1-S, with one pair arising laterally
and 5-7 irregular pairs arising relatively close to posterior
midline;with 2 pairs arisingwithin pecten (Figs. 22, 24). . . . . . . . . . . . 6
Siphon with 3-6 pairs of seta 1-S, with l-3 pairs arising laterally
and 2-4 pairs arising posterolaterally; sometimes with one
pair arisingwithin pecten. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .7
6(5).
Seta 2-X with 4 or 5 branches; seta 6-VI single; ventral brush
with 13 or 14 setae, usually14 (Fig. 22). . . . . . . . . . laticinctus (p. 67)
Seta 2-X only double or triple; seta 6-VI double; ventral brush
with 1l-13 setae, usually 12 (Fig. 24). . . . . . . . . . . . mattinglyi (p. 72)
7(5).
Seta 1-S no longer than diameter of siphon at point of attachment,
usually in 5 pairs; seta 6-VI normally single; seta 5-C usually
double or triple (occasionallywith 4 branches) . . . , . . . . . . . . . . . . . 8
Seta 1-S longer than diameter of siphon at point of attachment,
usually in 4 pairs; seta 6-VI normally double; seta 5-C with
4-8 branches. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
8(7).
Seta 1-C long and slender, length about 0.1 mm; siphon with 3
lateral pairs of seta 1-S; seta 1-M usually double or triple,
sometimes single (Fig. 14) . . . . . . . . . . . . . . . . . antennatus (p. 48)
Seta 1-C not as long, slightly stouter; siphon with 2 lateral
pairs of seta 1-S; seta 1-M usually single, sometimes
double..........................................g
9(8).
Seta 1-S distinctly shorter than diameter of siphon at point of
attachment (Fig. 16); occurring in southern part of Arabian
Peninsula . . . . . . . . . . . . . . . . . . . . . . . . . . . . univittatus (p. 52)
Seta 1-S about as long as diameter of siphon at point of
attachment (Fig. 18); occurring throughout southwestern
Asia and Egypt . . . . . . . . . . . . . . . . . . . . . . . . perexiguus (p. 57)
1O(7)
Seta 1-Ill-V with 3-6 branches (usually 4 or 5), sum of their
branches on one side of abdomen 10 or more (usually
Harbach: Subgenus Culex in SouthwesternAsia
21
more); seta 1-M normally double or triple; seta 1-X usually
double (Fig. 10) . . . . . . . . . . . . . . . . . . . . . . . . torrentium (p. 40)
Seta l-Ill-V with l-4 branches (usually 1 or 2), sum of their
branches on one side of abdomen not exceeding 10
(usually 6 or less); seta 1-M normally single; seta 1-X
usuallysingle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
ll(10)
Seta 1-C pigmented, usually spiculate in middle; integument of
thorax and abdomen with rows of minute vesicles; seta 13-T
as long as 12-T (Fig. 8); occurring in Pakistan, and perhaps
Afghanistanand Iran . . . . . . . . . . . . . . . . . . . . . . . vagans (p. 36)
Seta 1-C not pigmented, sides smooth; integument of thorax
and abdomen without vesicles; seta 13-T distinctly
shorterthan 12-T; more widespread species . . . . . . . . . . . . . . . . . 12
12(11)
Siphonal saddle index less than 3.45; seta 1-III,IV usually
single* (Fig. 6). . . . . . . . . . . . . . . . . . . . quinquefasciatus (p. 31)
Siphon al saddle index greater than 3.45; seta 1-III,IV
usually double* (Fig. 4) . . . . . . . . . . . . . . . . . . . . . . pipiens (p. 23)
13(2).
Seta 7-l distinctly shorter than 6-1, usually double; seta 14-C
with 2 or more branches, rarely single; seta 1-C slender,
usually not thickerthan branches of setae 5,6-C . . . . . . . . . . . . . . . 14
Seta 7-l about as long as 6-1, usually single; setal4-C single;
seta 1-C stout, usually much thicker than branches of
setae5,6-C . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .17
14(13).
Seta 5-C with 3 or 4 branches; siphon with most elements of
seta 1-S arising relatively close to posterior midline,
these with 4-11 branches; seta 14-P single (Fig. 20) . . . . theileri (p. 61)
Seta 5-C with 1 or 2 branches (usually single);
siphon with all elements of seta 1-S arising laterally
and posterolaterally, these with 1-4 branches; seta
14-P double on at least one side . . . . . . . . . . . . . . . . . . . . . . . . 15
15(14).
Pecten borne on proximal third or less of siphon; all scales of
comb spinelike (Fig. 26); siphon index usually less than
6.0; occurring in southern part of Arabian Peninsula . . simpsoni (p. 75)
Pecten longer, extending about 0.4 of siphon length;
some anterior scales of comb evenly fringed at sides
and apex (Fig. 28); siphon index usually greater
than 6.0; more widespread species . . . . . . . . . . . . sinaiticus (p. 78)
*These characters are unreliable in central and northeastern areas of the Arabian Peninsula where
introgressed populations (hybrids) occur.
22
Contrib.Amer. Ent. Inst.,vol 24, no. 1,1988
sitiens group
16(3).
Seta 5-C with 3 or 4 branches; seta 1-S only slightly longer
than diameter of siphon at point of attachment; seta 1-X
with 2-4 branches (Fig. 40); siphon index greater
than 5.5 . . . . . . . . . . . . . . . . . . . . . . tritaeniorhynchus (p. 106)
Seta 5-C with 6-8 branches; seta 1-S nearly twice as
long as diameter of siphon at point of attachment;
seta 1-X single (Fig. 32); siphon index less
than5.5 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .sitiens(p.87)
17(13)
Comb with more than 25 scales; seta 2-S long and curved;
most elements of ventral brush with 4 branches
(Fig. 36). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . mimeticus (p. 97)
Comb with no more than 10 scales; seta 2-S long or short,
always straight; most elements of ventral brush with
morethan 4 branches. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
18(17)
Median labral plate indistinguishablyfused with dorsal
apotome; dorsomentum a straight-sided triangle with
minutely serrated margins; pecten very short, less
than 0.1 length of siphon (Fig. 38) . . . . . . bitaeniorhynchus (p. 101)
Median labral plate distinct, separated from dorsal
apotome by well-developed suture; dorsomentum
an imperfect triangle with large teeth; pecten
much longer, about 0.3 length of siphon . . . . . . . . . . . . . . . . . . . .19
19(18)
Anterior margin of siphon distinctly curved in lateral view;
length of seta 1-S 2.5-3.0 diameter of siphon; pecten
spines with 4-9 basal denticles; setae 2,3-A inserted
at apex of antenna (Fig. 34). . . . . . . . . . . . . . . . . . poicilipes (p. 92)
Anterior margin of siphon more or less straight; length of
seta 1-S not more than 1.5 diameter of siphon; pecten
spines with complete ventral row of denticles; setae
2,3-A inserted subapically (Fig. 42). . . . . . . . pseudovishnui (p. 111)
Harbach: SubgenusCulex in SouthwesternAsia
THE
Culex
SPECIES
(Culex)
23
TREATMENTS
pipiens
Linnaeus
pipiens Linnaeus, 1758: 602. +Neotype male: Silvakra farm, Veberod,
Scania, Sweden; designated by Harbach et al., 1985: 9 (NMNH).
molestus Forskal, 1775: 85. +Neotype male: Rosetta, Buhayrah Gov.,
Eygpt; designated by Harbach et al. 1984: 523 (NMNH). Synonymy
with pipiens by Harbach et al., 1984: 540.
bicolor Meigen, 1818: 7. +Type female: ? Africa (MNHP). Synonymy
with pipiens by Edwards, 1921: 345.
pallipes Macquart, 1838a: 37. Type specimen(s) (non-extant): Egypt.
Synonymy with pipiens by Edwards, 1932a: 210.
melanorhinus Giles, 1900: 342. Nomen novum for pallipes Macquart,
1838b, non Waltl, 1835: 110.
longefurcatus
Becker, 1903: 68. Type female: Cairo, Egypt (ZM).
Synonymy with pipiens by Edwards, 1932a: 209.
For complete synonymy, see Appendix B.
Culex fatigans Wiedemann of Theobald, 1904: 76 (in part; Egypt); Theobald, 1905b: 92
(Egypt); Gough, 1914: 135 (Egypt).
Culex molestus Forskal of Marshall and Staley, 1937: 17-26 (Egypt, M, F, L, beh.,
physiol.); Buttiker,l981: 473 (in part; Saudi Arabia, bionomics).
Culex (&flex) fatigans Wiedemann of Abdel-Malek, 1960: 113-l 15, 118 (Syria, L
bionomics).
Culex (Culex) laticinctus Edwards of S&guy, 1924: 37 (in part; L* in Figs. 20-23, PI. II,
only).
Culex (Culex) molestus Forskal of Parr, 1943: 246-251 (Syria, Lebanon, A, P, L keys,
bionomics); Knight, 1951: 354 (Mediterranean Subregion, M, F, L, E, beh.,
physiol, distr.); Senevet and Andarelli, 1954: 49 (North Africa, M, F, A key); Senevet and Andarelli, 1959: 185 (North Africa, A, M gen., P, L, keys, biol.); AbdelMalek, 1960: 113-121 (Syria, bionomics, distr.); Lotfi, 1970: 401 [author listed as
“(Wiedemann)“] (Iran); Lotfi, 1973: 206 (Iran); Harbach et al., 1984: 523 (Egypt,
M*,F”, P*, L*, biol., beh./physiol.).
Culexpipiens Linnaeus of Theobald, 1901 b: 132 (World, M*, F, syn., bionomics, distr.);
Theobald, 1904: 76 (Egypt); Patton, 1905: 635 (P.D.R. Yemen, L habitat);
Theobald, 1905a: 28 (Egypt, syn.); Theobald, 1905b: 93 (Egypt, Israel, Jordan);
Edwards, 1913a: 49 (Israel, A); Edwards, 1913b: 53 (Africa, Syn., A, M gen.‘);
Gough, 1914: 135 (Egypt); Storey, 1918(1919): 87-99 (Egypt, A, L keys, A,
bionomics); Barraud, 1920: 324 (Iraq); Barraud, 1921: 393 (Egypt, Israel, Jordan, A, L, distr.); Edwards, 1922: 324 (Oriental Region, A key); Buxton, 1923:
317 (Israel, Jordan, bionomics); Theodor, 1923: 341 (Israel, Jordan, P’, key);
Kirkpatrick, 1924 (1925): 368, 371 (Egypt, A, L keys); Salem, 1938: 27 (Egypt);
Lewis, 1945: 17 (Sudan, M, F, L, distr.); Knight and Abdel Malek, 1951: 178
(Egypt, M, F, E, L, beh., physiol., bionomics); Dow, 1953: 688 (Iran, L assoc.);
Gad, 1956: 135 (Egypt, bionomic note); Gad and Darwish, 1957: 538 (Sinai);
Senevet et al., 1957a: 86 (N. Africa, L*); Hussain, 1963: 64 (Iraq); Gad and Salit,
1972: 581 (Egypt, bionomics); Margalit et al., 1973: 32 (Israel); Doyle et al., 1980:
24
Contrib.Amer. Ent. Inst.,vol. 24, no. 1, 1988
676 (Egypt, crossings); El-Said and Kenawy, 1983a (Egypt); Zimmerman et al.,
1985: 84 (Egypt, bionomics).
Culex pipiens molestus Forskal of Lewis, 1945: 17 (Sudan, M, F, L, bionomics); Theodor, 1952: 113 (Middle East, zoogeogr.); Margalit and Tahori, 1970a: 142 (Israel,
bionomics); Margalit et al., 1971: 323 (Israel, bionomics); Margalit and Tahori,
1973: 90 (Sinai, L assoc.); Zaini et al., 1983: 117 (Iraq).
Culex pipiens pipiens Linnaeus of Zaini et al., 1983: 117 (Iraq).
Culex (Cdex) pipiens Linnaeus of Edwards, 1921: 345 (Palaearctic Region, A, L keys,
tax., syn., distr.); Seguy, 1924: 35 (Palaearctic Region, A, L keys, M’, L’, tax.,
distr., med. imp., syn., distr.); Kirkpatrick, 1925: 133 (Egypt, A, P, L keys, syn.,
M*, F, P*, L*, bionomics); Edwards, 1926: 142 (Palaearctic Region, A, L keys,
syn., A, L, distr.); Stackelberg, 1927: 166 (Palaearctic Region, M, F keys, A, M
gen.*, distr.); Martini, 1931: 374 (Palaearctic Region, A, L keys, syn., M*, F, L*,
distr.); Edwards, 1941: 314, 420, 482 (Afrotropical Region, A key, M’, F’, P,
distr.); Senevet, 1947a: 119 (N. Africa, L*, distr., L assoc., L key); Knight, 1951:
359 (Mediterranean Subregion, M, F, L, E, beh., physiol., distr.); Knight, 1953a:
230 (Yemen Arab Republic, M, L, toll. rec., bionomics); Senevet and Andarelli,
1954: 48 (N. Africa, M, F, A key); Khattat, 1955: 166, 183 (Iraq, L*, distr., syn.,
bionomics); Lewis, 1956: 710 (Egypt, Sudan, Eritrea, L, toll. rec., bionomics);
Abdel-Malek, 1956: 102 (Sinai, bionomics, L key); Mattingly and Knight, 1956:
102 (Saudi Arabia, P.D.R. Yemen, United Arab Emirates, Yemen Arab Republic,
A, L’ keys, distr., bionomics); Senevet et al., 1957b: 92 (N. Africa, L); Parrish,
1959: 266 (Turkey); Senevet and Andarelli, 1959: 167 (North Africa, A, L, P keys,
M*, F, P’, L*); Abdel-Malek, 1960: 121 (Syria, bionomics, distr.); Khalaf, 1962: 46
(Iraq, A, M*, P*, L, bionomics); Derwesh, 1965: 44 (Iraq); DuBose and Curtin,
1965: 352 (Mediterranean area, A, L keys); Abul-hab, 1968: 246 (Iraq, L key);
Lotfi, 1970: 401 (Iran); Lotfi, 1973: 206 (Iran); Margalit and Tahori, 1974: 87
(Israel); El-Said and Kenawy, 1983b (Egypt, distr.); Danilov, 1985a: 71 (Afghanistan, A key); Danilov, 1985b: 55 (Afghanistan, L key); Harbach et al., 1985: 9
(Sweden, M’, F*, P*, L*, tax.); Harbach, 1985a: 86, 95, 105 (SW Asia [excluding
Pakistan], Egypt, distr., A, L keys).
Culex (Culex) pipiens var. mofestus Forskal of Mattingly and Knight, 1956: 103 (Saudi
Arabia, A, L keys, zoogeogr.).
Culex (Culex) pipiens molestus Forskal of Monchadskii, 1951: 274 (Middle East, L key,
L’, biol.); Hopkins, 1952: 302 (Afrotropical Region, L key, L’, bionomics); Khalaf,
1962: 49 (Iraq, M, F, L, biol.); Margalit and Tahori, 1970b: 152 (Israel); Lotfi, 1976:
84 (Iran, L key, L’); lbrahim et al., 1983: 92 (Iraq, L* key).
Culex (Culex) pipiens pipiens Linnaeus of Monchadskii, 1951: 272 (Middle East, L’,
key, biol.); Hopkins, 1952: 300 (Afrotropical Region, L key, L’, bionomics);
Senevet and Andarelli, 1959: 184 (North Africa, A, M gen.‘, P, L, biol.); Khalaf,
1962: 48 (Iraq, M, F, L, biol.); Lotfi, 1976: 83 (Iran, L key, L’); lbrahim et al., 1983:
92 (Iraq, L* key).
Culex quinquefasciatus Say of Dow, 1953: 688 (in part; northern localities in Iran, L
bionomics).
Culex (Culex) quinquefasciatus Say of Parrish, 1959: 266 (Turkey).
Adult.
A medium-sized mosquito without striking features and special
ornamentation. This species closely resembles quinquefasciatus with which it has
introgressed in certain areas of the Arabian Peninsula. It can be distinguished from
quinquefasciatus by the following combination of characters: (1) scutal scales golden
Harbach: Subgenus Culex in SouthwesternAsia
25
brown with reddish tint, (2) integument and scales between supraalar and posterior
dorsocentral setae noticeably darker, forming a pair of more or less conspicuous spots,
(3) cell R2 at least 4.0 length of vein R2+3, subcosta intersecting costa just beyond
furcation of R2+3; (4) basal bands of abdominal terga yellowish, usually same color as
sterna, contrasted with white basolateral spots in females.
FEMALE. Dark scaling with metallic blue reflections, reflections lost in aged
specimens. Head: Length of antenna1 flagellum about 2.0 mm, entirely dark,
flagellomere 1 yellowish to brownish on proximal 0.5, with some tiny, inconspicuous pale
scales mesally; pedicel yellowish to brownish, darker mesal surface with or without some
tiny pale scales. Proboscis length 1.9-2.3 mm, mean 2.1 mm; dorsal surface dark, darker
distally; ventral surface with cream-colored scales on proximal 0.7, distal 0.3 dark; labella
yellowish. Maxillary palpus entirely dark-scaled; length 0.4 mm, about 0.2 proboscis
length. Forked scales of vertex rather short, dark, some pale medially; falcate scales
narrow, pale yellow, paler laterally; lateral spatulate scales creamy white. Ocular setae
dark, reddish brown to black, antrorsely curved. Interocular space narrow, with pale
falcate scales continuous with those of vertex, and 2 large yellowish-brown setae
projecting ventrally over clypeus. Thorax: Pleural integument yellowish brown to
reddish, faded posteriorly; scutal integument dark brown. Scutal scales fine, golden
brown with reddish tint; finer on fossae and supraalar areas; integument and scales
between supraalar and posterior dorsocentral setae usually noticeably darker, evident as
a pair of ovoid spots, particularly when viewed in dorsal aspect, acrostichal and fossal
scales often as dark as ovoid spots; whitish to pale yellow scales on anterior promotory
and outer margins of supraalar and prescutellar areas. Scutal setae dark reddish
brown to nearly black. Scutellum with whitish to pale yellow falcate scales; 3-6 large
setae on each lateral lobe, 7 or 8 on median lobe. Antepronotum with upper and lower
patches of whitish to pale yellow falcate scales, scales of lower patch coarser and paler;
setae mainly dark, pale ventrally. Postpronotum with golden-brown falcate scales,
usually paler and slightly coarser posteriorly; usually with 6 setae (5-9) on posterodorsal
margin, longer posteriorly. Pleural setae golden to golden brown: 7-15 upper
proepisternal in more or less single row, 5-l 2 prealar, 3-7 upper mesokatepisternal, 5-13
lower mesokatepisternal, 4-12 upper mesepimeral, and 1 (rarely 2) lower mesepimeral.
Pleural spatulate scales nearly white: few below upper proepisternal setae, patches on
upper corner and lower posterior border of mesokatepisternum, anterior patch on
mesepimeron at level of upper mesokatepisternal patch, and small patch before upper
mesepimeral setae. Wing: Length 3.6-4.5 mm, mean 4.0 mm; length of cell R2/length
of vein R2+3 4.6-6.0, mean 5.3; subcosta intersects costa at or just beyond furcation of
vein R2+3; length of cell Ml/length of cell R2 0.7-0.8; scales entirely dark. Dorsal scaling:
broad squame scales on costa, subcosta, R, Rl, and CuA; relatively narrow squame
scales on R4+5, Ml, M2, M3+4, mcu, and proximally on 1A; linear plume scales on Rs,
R2+3, R2, R3, M, Ml+2, and distally on 1A; remigium with 2 distinct rows of scales and
usually 2, sometimes 3, setae distally. Ventral scaling: squame scales on costa,
subcosta, base of Rl, Rs, R2+3, bases of R2 and R3, Ml+& and bases of Ml and M2;
plume scales on other veins and parts of veins except CuA before mcu and proximal 0.5
of 1A which are without scales. Halter: Yellowish, capitellum usually darker with pale
scales. Legs: Anterior surface of forecoxa mainly black-scaled, with small basal patch of
yellowish scales, anterior surface also with many long golden-brown to nearly black
ventrally curved setae, apex with 3 or 4 shorter setae on posterior margin, most proximal
seta projecting nearly perpendicular to surface, others project ventrally; midcoxa with
midlateral longitudinal row of 4 or 5 long golden to golden-brown setae, setae margined
anteriorly by longitudinal patch of white spatulate scales, anterior surface with small patch
26
Contrib.Amer. Ent. Inst.,vol. 24, no. 1, 1988
of black scales and several ventrally projecting setae at apex; posterolateral surface of
hindcoxa with longitudinal row of 5-l 1 golden to golden-brown setae becoming
gradually shorter from base to apex, anterolateral surface with narrow longitudinal row of
whitish spatulate scales and about 4 short ventrally projecting setae at apex, mesal
surface with 2 dark setae at apex. Ventral surfaces of trochanters with white spatulate
scales; anteroventral surface of fore- and midtrochanters usually with some black
spatulate scales. Apices of all femora with narrow dorsal knee spot of yellowish scales;
anterior surface of forefemur black-scaled, posterior surface white-scaled; midfemur like
forefemur but black scales extend over dorsal surface toward apex; hindfemur mainly
white-scaled, with complete anterodorsal stripe of black scales gradually widening to
extend over entire anterior and posterior surfaces just before apex. Foretibia mainly
black-scaled, posteroventral surface with whitish scales; midtibia mainly black-scaled,
posteroventral surface with whitish scales, anterior and dorsal surfaces with whitish spot
at apex. Tarsi black-scaled, slightly paler ventrally, particularly tarsomere 1. Pulvilli pale.
Ungues small, simple, black. Abdomen: Terga mainly black-scaled; tergum I golden
setose, with posteromesal patch of black scales; posterior margins of terga II-VII with row
of golden setae, median setae about length of basal band of next tergum, lateral setae
longer, about 0.75 of tergum length; tergum II with basomedian spot of yellowish scales
(often nearly or entirely lost) and lateral patches of white scales; terga III-VII with basal
bands of yellowish scales and basolateral spots of white scales, spots becoming
gradually larger on succeeding posterior terga and actually cover entire lateral surfaces
of tergum VII, bands 0.15-0.35 tergum length, bands usually convex on terga Ill-V and
separated from or just touching basolateral spots, bands normally straight or slightly
concave on terga VI and VII and contiguous with basolateral spots, bands of terga III and
VII sometimes nearly or entirely lost; tergum VIII golden setose, usually with whitish
scales, sometimes with yellowish scales, scales normally paler than those of basal bands
of preceeding terga, sometimes with some dark scales posteriorly in middle. Sterna II-VII
mainly or entirely yellowish-scaled (scales same color as basal bands of terga), often with
some median black scales, particularly in less arid areas; sternum VIII with whitish scales
on lateral margins, broad median area without scales; all sterna golden setose.
MALE. Like female except as follows. Metallic reflections of dark scaling
subdued or scales without reflections, sometimes more opalescent than metallic. Head:
Length of antenna1 flagellum 1.4-l .6 mm; flagellomeres l-l 2 pale between whorls;
flagellomeres 13 and 14 0.6-0.7 mm, approximately 0.4 flagellum length. Proboscis
without ventral cluster of setae near false joint; false joint about 0.6 from base; ventral
pale scales at level of false joint, usually extending 0.5-0.7 from base, sometimes
covering most of ventral surface. Length of maxillary palpus 2.2-3.0 mm, mean 2.6 mm,
approximately 1.3 proboscis length, extending beyond tip of proboscis by about length
of palpomere 5; palpus mainly dark-scaled, integument between palpomeres 2 and 3
pale; lateral surface of palpomere 3 with stripe of cream-colored to white scales in middle,
stripe bordered ventrally by row of long black setae on distal 0.5 of palpomere, ventral
surface devoid of scales and pale, with ventromesal row of small antrorsely curved pale
setae; palpomere 4 with nearly complete ventral stripe of white scales; base of
palpomere 5 with small ventral patch of white scales; lateral surfaces of palpomeres 4 and
5 with long black setae that are longest at base of 4 and become gradually shorter toward
apex of 5. Thorax: Ovoid spots of scutum usually distinct. Proepisternum with 9-25
setae in more or less double row. Wing: Length 2.8-3.7 mm, mean 3.3 mm; length of
cell R2Ilength of vein R2+3 2.4-3.6, mean 2.8; subcosta intersects costa at or before
furcation of vein R2+3; length of cell Ml/length of cell R2 0.70-0.85. Legs: Ungues
black; anterior foreunguis larger than posterior foreunguis, both stout, anterior foreun-
Harbach: SubgenusCulex in SouthwesternAsia
27
guis with small ventral tooth near midlength, posteriorforeunguis with small tooth nearer
base; anterior midunguis like that of foreleg, posterior midunguis shorter and more
slender than posterior unguis of foreleg but with tiny ventral tooth near midlength;
hindungues very small, simple. Abdomen: Terga without basolateral spots; basal bands
0.25-0.40 tergum length, usually straight, or nearly so, and produced posteriorly along
lateral scale-free areas, particularly on terga V-VII, giving bands a concave appearance;
lateral scale-free areas with long laterally-projecting golden setae. Sterna more
frequently with median dark scales; sternum VIII (dorsal in position) entirely pale or with
dusky scales posteriorly in middle. Genitalia (Fig. 3): Ninth tergal lobes small, each with
7-l 4 unevenly-spaced setae.
Gonocoxite normal, ventrolateral setae strongly
developed, these longer and stouter than lateral setae, mesal surface with 5 rows of
small setae extending from base to level of subapical lobe; subapical lobe undivided,
setae a-f in more or less straight row with gap between c and d, seta g immediately
lateral to d-f, seta h lateral to g; a-c slightlycurved, a stout and rodlike with blunt apex, b
and c each with stout base, tapering distally and with apex hooked and pointed; d-f
shorter than a-c, hooked apically, d and e slender, f laterally flattened and appearing
broad in lateral view; gfoliform, longer than broad, slightly asymmetrical, apex not sharply
produced; h slender, bent distally. Gonostylus stout, curved, dorsal surface concave
with 2 small slender setae on distal 0.3; gonostylar claw short, broadest apically,
troughlike. Phallosome longer than broad with lateral plates and aedeagus of nearly
equal length; lateral plate with definite dorsal, lateral and ventral arms, dorsal arm broad,
apex nearly truncate, diverging laterally from its mate of the opposite side, appearing
slightly sinuous in lateral view; lateral arm broad in lateral view, its posterior margin more
or less trilobed, the ventral lobe more prominent than the others and bent
ventrolaterally, base of lateral arm with thumblike basal articulatory process (articulates
with basal piece), base of this process continuous mesally with dorsal aedeagal bridge;
ventral arm narrow and sharply curved with apex directed laterally; DV/D minus0.14 to
zero, mean minus 0.09. Aedeagus subcylindrical, narrowed distally; ventral aedeagal
bridge relatively wide, joining aedeagal sclerites just beyond midlength. Proctiger
without distinctivefeatures; paraproctwith small, conical basal lateral arm, crown dark with
numerous short, spinelike spicules. Cereal sclerite elongate, irregularly shaped; 3-6
cereal setae. Tergum X rectangular, adjoining paraproct below and behind basal lateral
arm.
Pupa (Fig. 3). Placement and attributes of setae as figured, range and modal
number of branches in Table 1; diagnostic characters as in the key. Cephalothorax:
Lightly tanned, legs, scutum, metanotum and metathoracic wings darker. Setae 1,2-CT
with 3-5 branches, most often with 4; 3,4-CT usually triple (2-5); 5-CT usually with 5
branches (3-7); 6-CT usually triple (l-5); 7,9,1 l-CT double; 8-CT most often with 4
branches (4-7); lo-CT frequently with 8 branches (5-13); 12-CT usually triple (2-5).
Trumpet: Moderately tanned, subcylindrical, gradually widened distally; index 4.9-6.8,
mean 5.4; tracheoid area darker, extending about 0.33 from base; pinna oblique,
about length of tracheoid area. Abdomen: Lightly tanned, terga I-VI darker in middle;
length 2.3-3.7 mm, mean 3.1 mm. Seta 6-I,11normally single; 7-I,11usually double; l-11
multiple (10-24); l-Ill-V frequently with at least 6 branches, 1-III,IV most often with 7
branches (5-10) 1-V most often with 6 branches (4-8); 2-ll,Vll lateral to seta 1, l-Ill-VI
mesal to 1; 5-IV often with 4 branches, rarely double (2-6); 5-V-VII almost always double;
6-lll,lV most often triple, 6-V,VI most often with 4 branches. Genital lobe: Lightly tanned;
length about 0.20 mm in female, 0.34-0.39 mm in male. Paddle: Lightly tanned, midrib
and buttress darker; outer margin without distinct spicules; midrib distinct except at
apex; length 0.80-l .30 mm, mean 0.93 mm, width O-62-0.78 mm, mean 0.69 mm, index
28
Contrib.Amer. Ent. Inst.,vol. 24, no. 1, 1988
1.2-l .6, mean 1.4.
Larva (Fig. 4). Character and positions of setae as figured, range and modal
number of branches in Table 21; diagnostic characters as in the key. Head: Wider than
long; length 0.70-l .04 mm, mean 0.87 mm; width 1.00-l .39 mm, mean 1.25 mm; mainly
lightly tanned, labiogula and posterior portions of lateralia and dorsal apotome
moderately tanned, dorsal apotome often with variably developed, moderately tanned
spots as follows: crescentic spot just anterior to each seta 8-C, small median spot
immediately posterior to bases of seta 5-C median transverse oval spot midway
between bases of seta 5-C and posterior margin of head, latter with small spot on either
side laterally. Median labral plate narrow but distinct, anterior margin slightly emarginate
between insertions of seta 1-C. Labiogula narrower anteriorly than posteriorly, length
about same as posterior width; hypostomal suture complete, extended posterolaterally
from posterior tentorial pit to near collar. Collar moderately developed along lateralia,
heavily tanned. Dorsomentum most often with 10 teeth (8-14) on either side of median
tooth. Seta 1-C slender, tapered distally, slightly bent mesad, not tanned; 2-C normally
absent; 3-C distinct; 4-C single, rather long; 5-C frequently with 5 branches (4-7); 6-C
most often with 4 branches (3-6); 7-C resembles 5,6-C, most often with 10 branches
(8-13); 8-C usually double (2-4); 9-C most often with 5 branches (3-7); 10-C usually
double (2,3); 11,12,13-C double or triple, 13-C infrequently with 4 branches; 14-C
normally single, rarely double. Antenna: Length 0.38-0.55 mm, mean 0.45 mm, about
0.5 length of head; moderately tanned, mesal surface with dark spot at base; seta 1-A
0.7 from base; part proximal to seta 1-A with strongly developed aciculae mainly on
dorsal and lateral surfaces, distal part slender and smooth except for few short aciculae
laterally near seta 1-A. Seta 1-A with about 22 branches (1 l-27). Thorax: Integument
hyaline, tubercles of all large setae moderately tanned; setae l-3-P and 9-l 2-P,M,T on
common tubercles. Setae l-3-P single, nearly of equal length; 4,7,8-P normally double,
occasionally triple; 11-P most often with 5 branches (3-8). Seta 1-M single, about 0.5
length of 3-M; 3-M single; 4-M usually double, occasionally single. Seta 1-T short, 0.5 or
less length of 2-T, frequently triple (2-5); 2-T usually single or double, infrequently triple.
Abdomen: Integument hyaline, tubercles of setae 7-1, 6-I-VI and 1,3-VIII moderately
tanned, tubercle of setae 2,3-X heavily tanned. Seta 3-I,VII usually single, occasionally
double; seta 6-I-VI long, 6-I,11most often triple, sometimes with 4 branches, 6-11rarely
double, 6-III-VI double, 6-111rarely triple; 7-l resembles 6-1, almost always double (l-3);
l-Ill-VI usually double, one branch longer than the other. Segment VIII: Comb with
33-58 scales, mean 45; scales short, evenly fringed on sides and apex, arranged in 4
irregular rows. Seta l-VIII most often with 5 branches (4-9); 3-VIII frequently with 8
branches (6-9); 5-VIII commonly with 4 branches (3-5). Siphon: Index 3.00-5.80, mean
4.58; subcylindrical, broadest at base, slightly sigmoid in lateral view; moderately tanned,
darker at base; acus attached, longer on posterior side of attachment. Pecten on basal
0.3 of siphon, with lo-18 spines, mode 14; spines increasing in size from base of
siphon, larger spines with 3 or 4 long basal denticles. Seta 1-S usually in 4 pairs (a fifth
seta sometimes occurs on one or both sides of siphon),la-S very near most distal
pecten spine, lc-S distinctly out of line with the others. Segment X: Saddle complete;
moderately tanned, darker dorsally; posterodorsal area with minute spicules; length
0.31-0.45 mm, mean 0.36 mm, siphon/saddle index 3.48-4.63, mean 4.08. Seta 1-X
usually single (l-3); 2-X usually double, infrequently triple; 4-X almost always in 6 pairs, all
setae borne on grid, setae increasing in length posteriad, most posterior seta about 7.5
of most anterior. Anal papillae elongate, subacutely tapered, length variable.
Systematics.
Culex pipiens is a highly plastic species which is widely distributed in the Holarctic Region and cooler parts of Africa. It occurs throughout Egypt and
Harbach: Subgenus Culex in SouthwesternAsia
29
generally north of a line transecting Saudi Arabia between Jeddah in the west and Al
Hasa Oasis in the east. It occurs north of about 30” N. in countries east of the Persian
Gulf. Some isolated populations occur in the southwestern mountains of the Arabian
Peninsula.
This species exhibits considerable variation in all stages and it cannot always be
distinguished from quinquefasciatus and torrentium with certainty. In the adult stage,
pipiens is reliably differentiated from these species by the character of the male genitalia.
The immatures are usually separable by the characters given in the keys. Additional
characters for separating the larvae of pipiens and torrentium are discussed by Harbach
et al. (1985). The separation of all stages of pipiens and quinquesfasciatus is
complicated by the fact that these species introgress in certain areas of Saudi Arabia and
Iraq (see discussion under quinquefasciatus ).
A relatively small, dark variant of pipiens occurs in southwestern Asia and Egypt
which might be confused with antennatus. This variant consists chiefly of females in
which the basal pale bands of the abdominal terga are reduced or absent. The
occurrence of similar specimens were reported previously by Edwards (1921) and
Harbach et al. (1985). Unbanded specimens also occur in Africa (Edwards, 1941). A
sporadic larval variant in which the head and siphon are darker than usual is sometimes
found in the region. Some dark larvae develop into dark adults (Harbach et al., 1984). A
single female from northern Sinai deserves special mention. This specimen (collected
by E.E. Austen at El Arish and deposited in the BM) is unusually pale, the wing has a well
developed patch of pale scales at the base of the costa, the pale bands of the abdominal
terga are white and convex as in quinquefasciatus, and the scutal scales are somewhat
coarser and paler than usual.
There is tremendous variation in the length of the siphon in the larval stage.
Specimens from polluted water tend to have a shorter siphon than specimens from
cleaner water, but there is a considerable range in siphon length in almost every
collection.
Despite the extensive amount of morphological variation which occurs in all life
stages of this species, there is no indication of subspecific or racial differences in the
various geographically separated populations examined. The only exception to this
might be those populations which occur at higher elevations in the southwestern
mountains of the Arabian Peninsula and the mountainous countries of eastern Africa.
Insufficient material was available for a critical examination of these populations, but they
do not appear to introgress with populations of quinquefasciatus, indicating that they
may not be as closely related to this species as populations occurring at more northern
latitudes. lntrogression occurs where populations of pipiens and quinquefasciatus
overlap in the lower Tigris-Euphrates Valley, the coastal plains east of the Persian Gulf,
and a narrow zone extending across the Arabian Peninsula from the Gulf of Bahrain to
the Red Sea coast in the vicinity of Jeddah. The parental epiphenotypes are distinct
outside the zone of introgression, indicating that species-specific traits are being
selected outside the zone. This is evidence for independent species cohesion. It
supports the taxonomic conclusion that pipiens and quinquefasciatus are distinct
species. The genetic data provided by Urbanelli and Bullini (1985) accords this
conclusion. These authors found that allopatric populations exhibit an average Nei’s
genetic distance of 0.17. Values of similar magnitude have been found among some
members of the Aedes (Ochlerotatus) caspius (Pallas, 1771) Anopheles (Celia)
gambiae Giles, 1902, and Anopheles (Anopheles) maculipennis Meigen, 1818 species
complexes (Bullini and Coluzzi, 1982).
Bionomics. Culex pipiens is the most common species of the subgenus Culex
30
Contrib.Amer. Ent. Inst.,vol. 24, no. 1, 1988
in southwestern Asia. It utilizes numerous and variable breeding places ranging from
highly polluted cesspits to clear water pools and containers. The species usually breeds
in stagnant water in either shaded or unshaded situations. The water may be fresh or
slightly brackish. Populations reach their highest densities in areas occupied by man. It
is not uncommon to find larvae in underground habitats such as drains, wells, and septic
tanks.
Culex pipiens feeds on a variety of warm-blooded vertebrates from birds to man.
Specific host feeding is influenced by host density and availability. In southwestern
Asia, the species feeds predominantly on mammals. Females feed at night and readily
attack man both indoors and outdoors.
Culex pipiens is an important vector of human pathogens nearly everywhere it
occurs. It has been found naturally infected with Sindbis and West Nile viruses in Israel
(Samina et al., 1986) and West Nile and Rift Valley Fever viruses in Egypt (Taylor et al.,
1953; Taylor et al., 1955; Tahori et al., 1955; Meegan et al., 1980; Meegan, 1981; Turell
et al., 1982). The importance of pipiens as a primary vector of periodic Bancroftian
filariasis in Egypt is well known (see Southgate, 1979 for a thorough review). Culex
pipiens is a very serious pest in southwestern Asia because of its close association with
man.
Distribution.
Culex pipiens inhabits the temperate regions of Europe, Asia,
Africa, Australia, and North and South America.
Material examined.
7,994 specimens. A total of 6,384 specimens (1,386
females, 1,059 males, 205 male genitalia, 1,889 pupal exuviae, 1 ,129 larval exuviae,
and 716 fourth-instar larvae) were examined from southwestern Asia - AFGANISTAN:
(Bolla Quichi, Garwargin, He’ral, Kataghan); EGYPT (including the neotype series of
molestus ): Aswan (Abu Simbil, Aswan, Dar el Salam, El Aqaba el Saqhira, Ezbet el
Silsila, Khour Abu Subeira, Kom Ombo, Nag el Ashbab, Nag el ldwa el Bahari, Nag el
Ritag, Nag el Shalabab, Nag Barn? el Gezira, Nag’ Tingar, Sahara City), Buhayrah
(Alexandria, El Tarh, Kafr el Dauwar, Rosetta), Eastern Deserts (Farouk Field), N Fayyum
(Abhit el Haggar, El Nazla, Tubhar), El Giza (Abu Rauwash, Barnasht, Dokki, El
Badrshein, El Rahawi, Ezbet Bosna Sharawy, Imbaba, Kafr Tohormos, Mena Road,
Moneeb, Nazlet el Ashtar, Nazlet el Saman, Saqqara, Talbia, Tanash, Tirsa), El Isma’diya
(Abu Khalifa, Isam’ iliya, Nifisha), El &hi/a (Abbassia, Cairo, Ezbet Said), El Qalyubiya
(Bahtim, El Ga’afara, El Qash-Shish, El Salmaniya, Hashim, Kafr Abo-Seer, Kaha,
Khanka, Qalyubiya, Sindiwa), El Suweis (Kantara, Suez), Luxor (Luxor), Marsa Matruh
(Siwa Oasis - Abul el Leef, Aghurmi, Ain Bundi, Masouse), Port Said (Bur Fu’ad, El
Raswa, Port Said), Red Sea (Bernece, Ghubbetel Bos, Marsa Alaam, Quseir, Wadi
Qiseib, St. Anonious Monastery), Sinai Peninsula (Dahab, El Arish, Wadi Feiran),
unknown localities; IRAN: (Afzulabad, Alchangi, Amirabad, Ardebil, Behbehan,
Bisotun, Bushehr, Cham Asbi, Enzeli, Ghasem Gheshlari, Kamroud, Kermanshah,
Khorramshuhr, Khorroniabad, Kivi, Natchi Bahran, Rasht, Sari, Shemiran, Teheran,
Zarjub); IRAQ: (Al Basrah); ISRAEL: (Acre, Beer Sheva, Beit Gubrin, Ben Sheman,
Coastal plains, Deir el Belah, Dhahiriya, Ein Gedi, Ein Hajla, Ein Meshoshim, El Athroun,
En Aqev, Gennesaret, Golan Nafech, Hadera, Haifa, Jaffa, Jericho, Jerisheh, Jerusalem,
Kafr Shammai, Kalia, Kindana, Kishon Marsh, Lake Huleh, Lake Tiberias, Ludd, Nahr
Namin, Nazareth, Nulata, Ramat Razi’el, Revivim, Sarona, Shafat, Tell el Nagilla, Tul
Keram, Wadi Ghuzze, Wadi Latron, Wadi Shikma, Wassat, Yazur, unknown localities);
JORDAN: (Amman, AZ Zarqa); LEBANON: (Amioum, unknown localities); P.D.R.
YEMEN: (Seiyum); SAUDI ARABIA: (Al Hasa Oasis, Al Hufuf, Al Kharj, Al Qatif, Jeddah);
TURKEY: (Istanbul, Izmir, Konya, Sinop, Tatvan); YEMEN ARAB REPUBLIC: (Hada,
Hamman A
‘ li, San’a, Ta’izz). An additional 1,610 specimens (599 females, 354 males, 45
Harbach: Subgenus Culex in SouthwesternAsia
31
male genitalia, 344 pupal exuviae, 237 larval exuviae, and 31 fourth-instar larvae) were
examined from ALGERIA, ETHIOPIA, KENYA, LIBYA, MAURITANIA, MOROCCO,
SOUTH AFRICA, SUDAN, SWEDEN (including the neotype series), TANZANIA,
TUNISIA, and ZIMBABWE.
Culex
(Culex)
quinquefasciatus
Sa y
quinquefasciatus Say, 1823: 10. +Neotype male: New Orleans, Louisiana, USA; designated by Sirivanakarn and White, 1978: 362
(NMNH).
fatigans Wiedemann, 1828: 10. Lectotype female:
East Indies
[Indonesia]; designated by Belkin, 1968a: 68 (NMW). Synonymy
with quinquefasciatus by Stone, 1956( 1957): 342.
pallidocephala
Theobald, 1904: 73. Holotype female: Sennar, Blue
Nile, Sudan (BM). Synonymy with fatigans by Edwards, 1941: 316.
cartroni Ventrillon, 1905b: 429. +Lectotype male, hereby designated,
bearing following data: “6 // MUSEUM PARIS / MADAGASCAR /
VENTRILLON 1904 // Culex / cartroni ventr. / type”; genitalia on
acetate strip (MNHP). Synonymy with fafigans by Edwards, 1932a:
208.
pygmaeus Neveu-Lemaire, 1906: 256. Syntypes (non-extant): Imi,
Ehtiopia. Synonymy with fatigans by Edwards, 1932a: 208.
zelt neri Neveu-Lemaire, 1906: 251. Syntypes (non-extant) : Harar,
Ethiopia. Synonymy with quinquefasciatus by White,1 975: 322.
For complete synonymy, see Appendix C.
Culex fatigans Wiedemann of Theobald, 1901 b: 151 (World, A key, syn., M*,F*, distr.);
Theobald, 1904: 76 (in part; Sudan); Patton, 1905: 635 (P.D.R. Yemen, L
habitat); Giles, 1906: 131 (Bahrain); Edwards, 1913b: 55 (Africa, syn., M’, F,
distr.); Barraud, 1920: 324 (Iraq); Edwards, 1922: 279 (Oriental Region, A key);
Barraud, 1924b: 1264 (India, syn., M*, F, distr.); Barraud, 1924c: 430 (India, L*);
Lewis, 1945: 19 (Sudan, M, F, L, distr.); Leeson and Theodor, 1948: 228
(Socotra, L habitat; A, L keys); Hussain, 1963: 64 (Iraq).
Culex (Culex) fatigans Wiedemann of Edwards, 1921: 345 (Iran, Iraq, A, L keys, tax.,
distr.); Seguy, 1924: 35 (Iran, Iraq, A, L keys, A, M gen.‘, tax., distr., med. imp.,
syn.); Edwards, 1926: 141 (Iran, Iraq, A, L keys, syn., A, distr.); Stackelberg,
1927: 165 (Iran, Iraq, M, F keys, A, M gen.‘, distr.); Martini, 1931: 364 (Palaearctic
Region, A, L keys, M*, F, L*, distr.); Barraud, 1934: 420 (India, A, L keys, M*, F,
L*, syn., L habitat, distr.); Edwards, 1941: 316, 421, 482 (Afrotropical Region, A
key, M*, F’, P, syn., distr., bionomics); Lewis, 1943a: 282 (Eritrea, L);
Monchadskii, 1951: 261 (Middle East, L*, biol., distr.); Hopkins, 1952: 304
(Afrotropical Region, L*, bionomics); Senevet and Andarelli, 1959: 187 (North
Africa, A, P, L keys, A, M gen.‘, P’, L’, biol.); Derwesh, 1965: 44 (Iraq); Abul-hab,
1968: 249 (Iraq, L key); Lotfi, 1970: 401 (Iran); Lotfi, 1973: 206 (Iran).
Culex molestus Forskal of Buttiker, 1981: 473 (in part; Saudi Arabia, bionomics).
Culex pipiens fatigans Wiedemann of Zaini et al., 1983: 117 (Iraq, bionomics).
Culex (Culex) pipiens fatigans Wiedemann of Lewis, 1956: 715 (Sudan, Eritrea (?), toll.
rec.); Mattingly and Knight, 1956: 103 (Arabian Peninsula, A, L keys, distr.,
zoogeogr., bionomics); Khalaf, 1962: 50 (Iraq, M*, F, L, A biol., L habitat): Ward,
32
Contrib.Amer. Ent. Inst.,vol. 24, no. 1, 1988
1972: 96 (Afghanistan); Lotfi, 1976: 84 (Iran, L key, L*); lbrahim et al., 1983: 92
(Iraq, C key).
Culex quinquefasciatus Say of Dow, 1953: 688 (in part; southern localities in Iran, L
bionomics).
Culex (Culex) quinquefasciafus Say of Knight, 1953a: 230 (Yemen Arab Republic, M
gen., L, toll. rec., L habitat); Sirivanakarn, 1976: 30 (Oriental Region, A, P, L
keys, M*, F*, P*, L’, tax., distr., bionomics, med. imp.); Danilov, 1985a: 71
(Afghanistan, A key); Danilov, 1985b: 55 (Afghanistan, L key); Harbach, 1985a:
86, 95, 105 (SW Asia, distr., A, L keys).
Adult. A medium-sized species closely resembling Culex pipiens with which it
has introgressed in northeastern and central areas of the Arabian Peninsula. This
species is distinguishable from pipiens by the following combination of characters: (1)
scutal scales more or less buff-colored, relatively long and sparse, giving scutum a rather
shaggy appearance, (2) integument and scales between supraalar and posterior
dorsocentral setae usually not markedly shorter or noticeably darker than surrounding
scales, (3) length of cell R2 less than 4.0 length of vein R2+3, subcosta intersects costa
before furcation of vein R2+3; (4) basal bands of abdominal terga whitish to
cream-colored, slightly if at all darker than basolateral white spots, usually slightly paler
than sterna.
FEMALE. Dark scaling with metallic blue reflections, reflections lost in aged
museum specimens. Head: Antenna1 length 1.6-2.0 mm, mean 1.8 mm; flagellum dark,
base of flagellomere 1 and most of pedicel whitish to yellowish orange; mesal surface of
pedicel dark with some inconspicuous scales and/or setae. Proboscis length 1.7-2.1
mm, mean 1.9 mm; mainly black-scaled, ventral surface with cream-colored scales on
proximal 0.7, pale scaling most pronounced broadly in middle. Maxillary palpus short,
less than 0.2 proboscis length, entirely black-scaled. Falcate scales of vertex narrow,
whitish to pale yellow, slightly paler along margin of eye; forked scales pale medially, dark
laterally; lateral spatulate scales narrow, white; ocular setae golden brown. Interocular
space narrow, with some ventrally projectingfalcate scales coarser and slightly paler than
those of vertex, with 2 long golden to golden-brown setae projecting downward over
clypeus. Thorax: Integument yellowish brown, scutum slightly darker than pleura.
Scutal scales slightly coarser and less dense than in pipiens, more or less buff-colored
(at low magnification), slightly paler on margins and prescutellar area; integument and
scales between supraalar and posterior dorsocentral setae usually not distinctly shorter
or darker than surrounding integument and scales. Each lobe of scutellum with falcate
scales concolorous with prescutellar scales; lateral lobes each with 5 or 6 large setae and
several smaller setae, large setae in 2 rows with 2 setae in dorsal row and 3 or 4 in ventral
row; middle lobe with 6 or 7 large setae in single row and several scattered small setae.
Antepronotum with whitish to pale yellow falcate scales confined mainly to dorsal and
ventral patches, dorsal patch coarser; setae concolorous with scutal setae, paler
ventrally. Postpronotum with pale yellow to golden-yellow falcate scales, scales paler
and coarser posteriorly;with curved posterodorsalrow of 7 or 8 setae, setae more or less
same color as scutal setae, posterior 2 or 3 setae paler. Pleural setae pale yellow to
golden yellow, numbers follow: usually 6 upper proepisternal, 8-10 prealar, 5 or 6 upper
mesokatepisternal, 7-9 lower mesokatepisternal, usually about 7 upper mesepimeral
and 1 lower mesepimeral. Proepisternum with small number of elongate white spatulate
scales below upper proepisternal setae, usually few dark scales mesal to setae.
Mesopleuron with the usual patches of white spatulate scales in upper and lower
positions on katepisternum and anterior and upper positions on anepimeron. Wing:
Harbach: Subgenus Culex in SouthwesternAsia
33
Length 3.1-3.8 mm, mean 3.4 mm; length of cell R2 2.8-3.3 length of vein R2+3;
subcosta usually intersects costa before furcation of vein R2+3; length of cell Ml 0.8
length of cell R2; entirely dark-scaled. Dorsal scaling: broad squame scales on costa and
subcosta; narrow spatulate to near fusiform (decumbent) scales on R, Rl, R3+4, distally
on Ml, M2, mcu, M4+5, CuA, and all but distally on 1A; linear plume scales on Rs, R2+3,
R2, R3, M, Ml+2, proximally on Ml, and distally on 1A; remigiumwith 2 rows of scales and
2 or 3 setae distally. Ventral scaling: broad squame scales on costa, Rs, R2+3, Ml,
Ml+2, and proximally on Ml ; linear plume scales on R, Rl, R2, R3, R4+5, distally on Ml,
M2, M2+3, mcu, CuA distad of mcu, and about distal 0.3 of 1A; CuA proximal to mcu and
about proximal 0.7 of 1A without scales. Halter: Pedicel and scabellum pale; capitellum
dark with pale scales. Legs: Forecoxa with patch of whitish scales at base continuous
with predominantly black scales of anterior surface, variable number of whitish scales
usually near middle of anterior surface, ventrally curved anterior setae golden brown,
usually with 2 more or less prominent laterally projecting and several less conspicuous
ventrally projecting setae apically on posterolateral margin; midcoxa with narrow
longitudinal patch of white spatulate scales on anterior side of longitudinal row of usually
4 large golden-brown and several smaller paler setae on lateral midline, pale scales
continuous with small apical patch of black scales, number of small ventrally projecting
pale setae anterolaterally at apex; hindcoxa with somewhat inconspicuous narrow
longitudinal patch of whitish scales on anterolateral margin, with 6-8 golden
posterolateral setae and about 4 short golden setae at apical end of scale patch.
Trochanters with white scales on ventral surface extended onto anterior and posterior
surfaces; fore- and midtrochanters often with some black scales anteromesally. Femora
with inconspicuous narrow yellowish knee spots; fore- and midfemora black-scaled
anteriorly, white-scaled posteriorly, black scaling of midfemur gradually expanded over
dorsal and onto posterior surface toward apex; hindfemur white-scaled with anterodorsal
stripe of black scales gradually expanded distally over anterior and posterior surfaces,
sometimes nearly forming complete subapical ring. Tibiae mainly black-scaled, foretibia
with whitish scales ventrally; midtibia with whitish scales posteroventrally; hindtibia like
foretibia with somewhat indistinct apical pale spot mainly on dorsal and anterior surfaces.
Tarsi dark, slightly paler ventrally; tarsomeres l-4 with hint of pale scaling dorsally at apex.
Pulvilli pale. Ungues small, simple. Abdomen: Tergum I golden setose, with median
posterior patch of black, white or mixture of black and white scales; terga II-VII with long
golden setae on lateral and posterior margins, setae short medially on posterior margin;
tergum II with median basal triangular patch of whitish or very pale yellow scales and
lateral patches of white scales; terga Ill-VII with nearly white convex basal bands and
white basolateral spots, bands about 0.4 tergum length, spots becoming larger on
succeeding posterior terga, bands not touching or narrowly continuous with spots on
terga Ill-V, bands continuous with spots on terga VI and VII, tergum VII with some pale
scales on posterior border; tergum VIII entirely pale-scaled. Sterna II-VII nearly white to
yellowish, usually more yellow (slightly darker) than basal bands of terga, often with some
median black scales which sometimes form rather large patch not reaching posterior
border, occasionally with few black scales on posterolateral corners.
MALE. Like female except as follows. Head: Proboscis without ventral cluster of
setae near false joint; false joint about 0.6 from base; pale scaling mainly in middle
ventrally, forming indistinct more or less complete narrow ring at false joint. Length of
maxillary palpus about 2.0-2.6 mm, mean 2.4 mm, extending beyond tip of proboscis by
length of palpomere 5 (palpus of some males from Djibouti only slightly longer than
proboscis or exceed its length by no more than 0.5 length of palpomere 5); integument
pale across joint between palpomeres 2 and 3; middle of palpomere 3 with elongate
34
Contrib.Amer. Ent. Inst.,vol. 24, no. 1, 1988
patch of cream-colored scales on dorsal and lateral surfaces, palpomere with about 14
long ventrolateral setae on distal 0.33 and longitudinal row of short antrorsely curved
setae on ventromesal margin; palpomere 4 with complete ventral line of white scales,
distal scales usually darker with bluish hue; palpomere 5 with small ventral patch of white
scales at base. Thorax: Scutal scales slightly longer and narrower, of more uniform size
than in female. Propleuron with 2 irregular ventromesally oriented rows of upper
proepisternal setae. Wing: Length 2.6-3.2 mm, mean 3.0 mm; length of cell R2 1.8-2.2
length of vein R2+3; length of cell Ml 0.75-0.85 length of cell R2. Legs: Anterior unguis
longer than posterior unguis on fore- and midlegs, each with ventral tooth near middle.
Abdomen: Terga without basolateral spots, basal bands produced posteriorly on lateral
margins, particularly on terga VI and VII; tergum VIII (ventral in position) largely or entirely
pale-scaled. Sterna more frequently with median black scales than in female; sternum
VIII (dorsal in position) variable, often like tergum VII, sometimes largely pale-scaled.
Genitalia (Fig. 5): Form as figured; essentially as in pipiens, differing chiefly in details of
the phallosome. Ninth tergal lobe with 4-14 setae (mode 8). Seta f of subapical lobe of
gonocoxite sometimes duplicated, distinctly hooked. Lateral plate of phallosome
conspicuously broader at base; dorsal arm not as broad, apex pointed, nearly parallel
with its mate of the opposite side; dorsal and ventral lobes of lateral arm more prominent;
ventral arm broad and long; DV/D 0.56-1.89, mean 1.03. Paraproct with short, slender
basal lateral arm.
Pupa (Fig. 5). Character and placement of setae as figured, range and modal
number of branches in Table 2; very similar to pipiens, but usually separable by the
character of the trumpet and seta 8-CT. Cephalothorax: Seta 5-CT usually with 4
branches (2-4); 6-CT often double (2-4); 8-CT usually triple (2,3). Trumpet: Relatively
short and flared apically; index 2.9-4.8, mean 4.2; pinna oblique, long, length 0.4 or
more of trumpet length. Abdomen: Length 3.2-4.0 mm, mean 3.6 mm; seta l-Ill-V
frequently with 4 or 5 branches, l-111most often with 5 branches (4-7), l-IV,V most often
with 4 branches (3-6); 5-IV usually double or triple, sometimes with 4 branches; 6-lll,lV
double or triple, more often double. Paddle: Length 0.98-l .16 mm, mean 1.04 mm;
width 0.74-O-87 mm, mean 0.82 mm; index 1.1-l .4, mean 1.3.
Larva (Fig. 6). Character and positions of setae as figured, range and modal
number of branches in Table 22; closely resembling pipiens from which it cannot always
be separated, the most reliable diagnostic features from pipiens include the
siphon/saddle index and character of seta 1-III,IV. Head: Length 0.74-0.88 mm, mean
0.83 mm; width 1.13-l .30 mm, mean 1.21 mm. Dorsomentum most often with 12 teeth
(1O-13) on either side of median tooth. Seta 2-C normally present, very small; 6-C most
often with 5 branches (3-5); 7-C most often with 8 branches (7-10); 8-C often triple (2-4);
10-C most often triple (l-3). Antenna: Length 0.43-0.50 mm, mean 0.45 mm. Thorax:
Seta 1-T frequently double (2-5). Abdomen: Seta 1-III,IV normally single, seldom
double (more seldom double on both segments). Siphon: Generally shorter and
stouter than in pipiens, index 2.88-4.64, mean 3.70. Pecten with 5-17 spines, mode
10. Seta 1 -S usually in 4 pairs, occasionally in 3 pairs, seldom with a fifth seta on one
side; number of branches variable, tending to have a greater number than in pipiens,
particularly la- and 1b-S (cf. in Tables 21 and 22). Segment X: Saddle length 0.35-0.48
mm, mean 0.41 mm; siphon/saddle index 2.77-3.41, mean 3.11.
Systematics. Culex quinquefasciatus is a variable but fairly well differentiated
species which has introgressed with pipiens in areas where the two species come
together. During this study, 180 mongrels (57 females, 67 males, and 56 male genitalia)
were examined from the lower Tigris-Euphrates Valley in Iraq, the eastern coastal region
of Saudi Arabia along the Persian Gulf, Al Kharj Oasis near Riyadh, and Jeddah on the
Harbach: Subgenus Culex in SouthwesternAsia
35
Red Sea coast. Hybrids are morphologically more variable than the parental species.
Some hybrids are recognizable in having the scutal scaling of one species and the
abdominal banding of the other species, but in general the only reliable means of
distinguishing hybrids from the recurrent parental species is by the male genitalia. The
form of the dorsal and ventral arms of the phallosome is intermediate between that found
in pipiens and quinquefasciatus. Hybrid males examined from southwestern Asia have
DV/V ratios ranging from 0.08 to 0.43 with a mean of 0.18. Mongrels of partial hybrid
ancestry may be indistinguishable from the parental species. For this reason, some
hybrid specimens collected in and around the zone of introgression were probably
misidentified as parental epiphenotypes during the course of this study. There is no
doubt, however, that the parental epiphenotypes show independent species cohesion
outside the zone. The two epiphenotypes are closely related, distinct species. The
capacity for pipiens and quinquefasciatus to interbreed should not be considered as
prima facie evidence for incomplete speciation.
Culex quinquefasciatus is widely distributed in the tropical and subtropical zones
of the world where it has no doubt been spread by commerce. However, the taxon has
not been sufficiently studied thoughout its range and it is possible that it consists of two
or more biological species. There is currently no evidence to suggest that more than
one species occurs in southwestern Asia.
Blonomlcs. Culex quinquefasciatus is a common domestic and peridomestic
species which has received considerable attention in the literature. It breeds mainly in
polluted water but is also found in clean fresh water and brackish water habitats. The
immatures occur in virtually all types of man-made and natural breeding places. Females
are undoubtedly anthropophilic but readily feed on a variety of other mammals and birds
depending on the availability of hosts. Biting occurs both indoors and outdoors at night,
and engorged females can be found resting either inside or outside dwellings and
shelters.
Although it is well known that quinquefasciatusis an important vector of filarial and
viral agents of human disease in many parts of the world, there are no published reports
of natural infection in southwestern Asia. Culex quinquefasciatus is the predominant
vector of urban filariasis in areas of Africa and the Orient. It has been found infected with
Japanese encephalitis virus in Vietnam (Nguyen et al., 1974) and Chikungunya virus in
Tanzania (White, 1971). It is capable of transmitting West Nile virus in India (Rao, 1975)
and Murray Valley encephalitis virus in Australia (Karabatsos, 1985).
Distribution.
Culex quinquefasciatus is widely distributed in the tropical and
subtropical regions of the world.
Material
examined.
1 ,113 specimens. A total of 907 specimens (306
females, 308 males, 256 male genitalia, 10 pupal exuviae, 5 larval exuviae, and 22
fourth-instar larvae) were examined from southwestern Asia - BAHRAIN: (specific
localities unknown); IRAN: (Alchangi, Bushehr, Enzeli, Halileh, Hizan, Kish Island,
Manyuhi, Nahr Mohsen, Shush); IRAQ: (Amara, Baghdad, Basrah); OMAN: (Muscat,
Salalah); PAKISTAN (including localities east of the lndus River): (Jhang, Jhelum,
Lahore, Peshawar, Rawalpindi); P.D.R. YEMEN: (Anag, Lahej, Makhzan, N’Air, Wadi
Duan); SAUDI ARABIA: (Al Hasa Oasis, Al Khobar, Al Qatif, Al Ta’if, Dhahran, Jeddah,
Mecca, Saihat); YEMEN ARAB REPUBLIC: (Murawah). An additional 206 specimens
(102 females, 46 males, 31 male genitalia, 12 pupal exuviae, 12 larval exuviae, and 3
fourth-instar larvae) were examined from DJIBOUTI, ETHIOPIA, MAURITANIA,
MOZAMBIQUE, NEPAL, SENEGAL, SOCOTRA, SOUTH AFRICA, SUDAN, UGANDA,
and the UNITED STATES (the neotype series).
36
Contrib.Amer. Ent. Inst.,vol. 24, no. 1, 1988
Culex (Culex)
wagans W i e d e m a n n
vagans Wiedemann, 1828: 545. Syntypes (non-extant); Foochow
(Barraud, 1934: 416) China.
tipuliformis Theobald, 1901 b: 325. +Holotype female: Bakloh, N. W.
Provinces (? Pakistan) (BM). Synonymy with vegans by Edwards,
1926: 136.
virgatipes Edwards, 1914b: 126. +Holotype male: Hong Kong (BM).
Synonymy with vegans by Edwards, 1926: 136.
exilis Dyar, 1924: 127.
+Lectotype male: Vladivostok, [Maritime
Territory], USSR; designated by Stone and Knight, 1957: 49
(NMNH). Synonymy with vagans by Edwards, 1926: 136.
Culex tipuliformis Theobald of Barraud, 1924b: 1269 (India, A key, syn., M*, F, distr.);
Barraud, 1924c: 430 (India, L’).
Culex vagans Wiedemann of Theobald, 1901 a: 411 (Oriental Region, A key, F*).
Culex (Culex) vagans Wiedemann of Edwards, 1926: 136 (Palaearctic Region, A, L
keys, syn., A, distr.); Martini, 1931: 391 (Palaearctic Region, A, L keys, M*, F,
syn.); Barraud, 1934: 416 (India, A, L keys, syn., A, M gen.*, L*, L habitat, distr.);
Monchadskii, 1951: 283 (Middle East, L key, L*, bionomics, distr.); (?) Parrish,
1959: 266 (Turkey); DuBose and Cur-tin, 1965: 352,354 (Mediterranean area, A,
L keys); (?) Lotfi, 1970: 400, 402 (Iran); Aslamkhan, 1971: 155 (Pakistan); (?)
Lotfi, 1976: 73, 76 (Iran, L key, L*); Sirivanakarn, 1976: 37 (Oriental Region, A, P,
L keys, M*, F, P*, L’, tax., bionomics, distr.); Harbach, 1985a: 86, 94, 105
(Afghanistan, Iran [not Egypt], Pakistan, A, L keys).
Adult. This medium-sized species resembles theileri in having anterior pale
stripes on the fore- and midfemora and all tibiae, but is easily distinguished by the
absence of postspiracular and prealar scales. The species otherwise resembles pipiens,
which of course lacks the striped legs. The male genitalia are strikingly similar to those of
torfentium.
FEMALE. Head: Length of antenna about 2.2 mm; pedicel usually mainly yellow,
mesal surface black, both pedicel and flagellomere 1 with some small white scales on
mesal surface; flagellum normal, whorls normally with 6 setae. Proboscis length 2.0-2.4
mm, mean 2.2 mm; sides and venter with cream-colored scales extending from 0.15 to
0.70 from base, proximal and distal portions distinctly black-scaled. Maxillary palpus
dark-scaled with white scales mainly confined to dorsal stripe on palpomeres 3 and 4;
length about 0.4 mm, about 0.2 proboscis length. Vertex with whitish to pale yellow
falcate scales, usually paler medially and along ocular line; forked scales mainly brown,
some pale ones usually in middle; lateral spatulate scales white or nearly so. Interocular
space with few pale scales, concolorouswith ocular scales. Ocular setae dark brown with
golden or reddish sheen; 2 yellowish interocular setae project ventrally over clypeus.
Thorax: Pleural integument yellowish brown to brown; scutal integument brown. Color
of scutal scales ranging from brownish yellow to golden brown, often mainly yellowish
brown, surface sometimes appearing somewhat mottled with fossal, sublateral supraalar
and occasionally posterior acrostichal scales darker; scales pale yellow, whitish or even
white on anterior and lateral margins and on prescutellar area. Scutal setae dark brown
with golden or reddish sheen. Scales of scutellum same color as prescutellar scales;
lateral lobes each with 4 or 5 large setae, median lobe with 6-8. Antepronotum mainly
Harbach: Subgenus Culex in SouthwesternAsia
37
with dark brown setae, some yellowish ones ventrally; with rather coarse falcate scales in
upper and lower patches, upper patch brownish yellow, lower whitish. Postpronotum
with brownish-yellow falcate scales, some paler (frequently whitish) posteriorly; with
usually 6 or 7 dark setae on posterodorsal margin. Pleural setae golden: 8-12 upper
proepisternal, 1O-l 2 prealar, 4-6 upper mesokatepisternal, 9-13 lower mesokatepisternal, lo-14 upper mesepimeral, and 1 lower mesepimeral. Pleura with white,
elongate spatulate scales in patches as follows: small patch below upper proepisternal
setae, patches on upper corner and lower posterior border of mesokatepisternum,
anterior patch on mesepimeron at level of upper mesokatepisternal patch, and patch
before and among upper mesepimeral setae; lower mesokatepisternal and upper
mesepimeral scales semierect, others decumbent. Wing: Length 3.5-5.4 mm, mean
4.5 mm; length cell R2 2.9-3.8 length of vein R2+3, mean 3.3; length cell Ml 0.8 length
of cell R2; scales predominantly dark, pale scales on subcosta and posterior side of costa
on dorsal and ventral surfaces, dorsal surface also with pale scales on anterior and
posterior sides of Rl distal to furcation of vein R2+3. Halter: Integument usually
yellowish, dorsal areas of pedicel and capitellum often dark; scales entirely pale. Legs
(midleg, Fig. 2G): Anterior surface of forecoxa dark-scaled in middle, with patches of
white scales at base and apex; midcoxa with longitudinal patch of white scales on
anterior side of midlateral row of strong setae, anterior surface with spot of scales at
apex; hindcoxa with longitudinal patch of white scales on anterior side of lateral midline,
posterolateral margin with irregular row of strong setae. Lateral surfaces of trochanters
without scales, other surfaces white-scaled, anterior surface of foretrochanter often with
some dark scales. Femora mainly white-scaled, apices with pale knee spots, spots often
indistinct on fore- and midfemora; anterior and dorsal surfaces of fore- and midfemora
dark-scaled, anterior surface with narrow longitudinal white stripe, forefemur sometimes
with ventral 0.5 of anterior surface completely white-scaled, or nearly so, because
anteroventral dark scaling partially or completely absent; hindfemur with complete
anterodorsal stripe of dark scales, stripe gradually widening distally, expanded over
approximately distal 0.5 of dorsal surface but may or may not be expanded over whole of
anterior surface very near apex. Anterior and posterior surfaces with dark scales; dorsal
surface with spot of white scales at apex; ventral dark scales of foretibia sometimes partly
or entirely replaced by white scales. Tarsi dark-scaled with faint anterior and posterior
pale stripes on tarsomere 1. Abdomen: Tergum I with median posterior patch of
yellowish scales; terga II-VII mainly clothed in dark brown to blackish scales; terga II-VII
with rather large basolateral patches of white scales, sometimes covering most or all of
lateral surface, particularly on terga II, VI and VII; tergum II with basomedian patch of
yellowish scales, patch sometimes lengthened posteriorly; terga Ill-VII with basal bands
of yellowish scales, bands 0.3-0.5 of tergum length, bands of terga Ill-V (and often VI
and sometimes VII) convex, those of VI and VII often straight or nearly so, bands of terga
III and IV (and frequently V and VI) not reaching basolateral white patches; tergum VIII
usually entirely pale-scaled, sometimes with some dark scales posteriorly. Sterna II-VII
mainly clothed in yellowish scales, with a median area of dark scales; sternum VIII with
lateral patches of pale scales, median area without scales.
MALE. Like female except for the following principal differences. Head:
Proboscis with indefinite pale band (narrowly incomplete dorsally) about 0.5-0.7 from
base, pale scaling of ventral surface extending from band to about 0.2 from base; false
joint 0.6 from base; without ventral cluster of setae at false joint. Maxillary palpus mainly
dark-scaled above; lateral surface of palpomere 3 with rather broad stripe of white scales
0.2-0.8 from base, mesal surface with complete or nearly complete narrow line of white
scales which is broader and more noticeable near apex, ventrolateral margin with row of
38
Contrib.Amer. Ent. Inst.,vol. 24, no. 1, 1988
about 28 long dark setae on distal 0.5, ventromesal margin with complete row of short,
pale, antrorsely curved setae; palpomere 4 with complete ventral stripe of white scales
and dorsal spot of rather subtle pale scales at base; dorsal surface of palpomere 5 like
that of 4, ventral surface with distinct basal spot of white scales; lateral and mesal
surfaces of palpomeres 4 and 5 densely setose; palpus exceeds length of proboscis by
length of palpomere 5. Thorax: Scutal scales finer. Upper proepisternal setae more
numerous. Wing: Length of cell R2 1.5-2.0 length of vein R2+3; subcosta intersects
costa before furcation of vein R2+3; length of cell Ml 0.8-0.9 length of cell R2.
Abdomen: Terga II-VII without lateral white patches; basal bands of terga Ill-VII either
convex or nearly straight, bands of terga V-VII produced posteriorly along lateral
scale-free areas but only slightly so on V; tergum VIII (ventral in position) with pale scales
before median posterior setose area. Sterna II-VII usually with median dark scales
forming distinct stripe; sternum VIII (dorsal in position) entirely pale-scaled. Genitalia (Fig.
7): Form as figured; in general as in pipiens and related species. Ninth tergal lobe short
and broad, with 6-14 unevenly spaced setae. Gonocoxopodite essentially as in pipiens;
subapical lobe indistinctlydivided, seta g on small prominence with h at its base laterally;
usually 2 setae in group d-e, sometimes only 1, sometimes 3; seta g not as broad,
pointed; h flattened and shorter. Phallosome longer than broad with lateral plates
slightly longer than aedeagal sclerites; dorsal arm slightly twisted and pointed apically,
diverging laterally from its mate of the opposite side but not appearing sinuous in lateral
view; lateral arm longer than broad, with prominent triangular laterally directed apical lobe
and a shot-l dorsal process, apical lobe minutely spiculate on posterior surface, dorsal
process relatively short and borne on dorsal margin between apical lobe and rather
weakly developed basal articulatory process; ventral arm short, bent as in pipiens.
Aedeagus short, somewhat conical; aedeagal sclerites about as long as broad; ventral
aedeagal bridge rather narrow, joining aedeagal sclerites near midlength. Proctiger
normal; paraproct with long, rather stout, ventrally curved basal lateral arm, crown with
numerous spinelike spicules and some short lateral blades. Cereal sclerite elongate,
with 2-5 cereal setae. Tergum X developed as usual.
Pupa (Fig. 7). Placement and attributes of setae as figured, range and modal
number of branches in Table 3; differing little from pipiens and allied species, but usually
recognized without much difficulty by the bent trumpet and relatively long seta 5-IV.
Cephalothorax: Seta 1,2,4,6-CT usually with 3 or 4 branches; 3-CT usually triple (2,3);
7-CT double; 8-CT tending to have more branches than usual (3-9); 9-CT with 2-4
branches; 11-CT usually double, sometimes single. Trumpet: Distinctly bent at distal
end of tracheoid area; index 5.0-6.9, mean 5.6; pinna short, 0.15-0.20 length of
trumpet. Abdomen: Length 2.1-2.6 mm, mean 2.4 mm. Seta l-Ill-VII tends to have
more branches than in pipiens and related species, l-111frequently with 9 or 10 branches
(6-l l), 1-IV often with 8 branches (5-10); 5-IV as strong as 5-V,VI, normally double or
triple (2-4). Paddle: Length 0.73-0.81 mm, mean 0.78 mm; width 0.51-0.60 mm, mean
0.56 mm; index 1.2-l .6, mean 1.4.
Larva (Fig. 8). Placement and character of setae as figured, range and modal
number of branches in Table 23; very similar to pipiens, but usually easily recognized by
the character of seta 1-C and the presence of rows of minute vesicles on the thoracic
and abdominal integument. Head: Length 0.76-0.92 mm, mean 0.82 mm; width
1.17-l .30 mm, mean 1.24 mm; lightly to moderately tanned, darker behind eyes, gula
also usually darker; collar and dorsomentum heavily tanned. Dorsomentum usually with
9 teeth (8-l 1) on either side of median tooth. Seta 1-C long, tapered, tanned, and
usually spiculate in middle; seta 2-C absent. Antenna: Generally slightly longer than in
p@iens, length 0.52-0.58 mm, mean 0.57 mm. Seta 1-A with about 25 branches
Harbach: Subgenus Culex in SouthwesternAsia
39
(20-33). Thorax: Integument with rows of minute vesicles. Seta 1-M single, normally
slightly longer than 0.5 length of 3-M. Seta 1-T about 0.5 length of 2-T; 2-T frequently
single, sometimes double; 13-T rather long, about same length as 12-T. Abdomen:
Integument with rows of tiny vesicles, changing to rows of tiny spicules on segment VIII.
Seta 6-l usually triple (2-4), 6-11usually with 3 or 4 branches (3-5), 6-III-VI almost always
double, occasionally triple. Segment VIII: Comb with 36-47 scales, mean 41. Siphon:
Generally longer than in pipiens, index 3.92-6.72, mean 5.39; lightly to moderately
tanned. Pecten with 1O-l 5 spines, mode 13; larger spines with 2-5 denticles, usually 3
or 4. Seta 1-S usually in 4 pairs, occasionally with a fifth seta on one side; 1b-S often
inserted laterally in line with lc-S. Segment X: Saddle length 0.38-0.47 mm, mean 0.41
mm; siphon/saddle index 3.79-4.59, mean 4.19. Seta 1-X single or double, more often
single. Anal papillae equal in length, slightly longer than saddle.
Systematlcs.
Culex vagans and the next species (torrentium ) bear a
pronounced resemblance to members of the pipiens subgroup, but based on the male
genitalia they unmistakingly belong to the trifilatus subgroup of Mattingly and Rageau
(1958). Culex vagans is the most distinctly marked member of the subgroup, the adults
being readily distinguished from all other species of the group, as well as from pipiens
and quinquefasciatus, by the striped femora and tibiae. This species bears a superficial
resemblance to theileriwith which it has been confused in the past.
This species occurs in a relatively broad belt which stretches from the Middle East
to Hong Kong, Japan, and Korea. Its exact distribution in southwestern Asia is
unknown. I have seen specimens only from Afghanistan and Pakistan, but the species
has been reported to occur in Iran (Lotfi, 1970; 1976) and Turkey (Parrish, 1959). I am
somewhat skeptical about the records from Iran because they are based on limited
collections of larvae. Characters which have been used to separate the larva of vagans
from the larvae of pipiens, quinquefasciatus, and torrentium are certainly not as reliable
as they appear to be in previously published keys (see the list of earlier works following
the synonymy). In fact, the characters used in my key are probably not completely
reliable because of the very limited number of specimens examined. Concerning the
record from Turkey, it is not clear whether Parrish (1959) based his identificationon larval
or adult specimens. Parrish apparently collected much of his material in light traps. It is
possible that his record of vagans from Turkey is based on damaged and/or partially or
completely denuded females of theileri and/or males of torrentium collected in light
traps. I would have the same concern about the identification of larvae of vagans from
Turkey that I have for the identification of larvae from Iran. Mattingly (1955a: 31)
identified three male genitalia and a larva of torrentium from Turkey which were labelled
previously as vagans by Martini (1931).
The adult and immature specimens examined during this study seem to conform
quite well with the recent descriptions of this species based on material from eastern
Asia (Sirivanakarn, 1976; Tanaka et al., 1979). The only obvious difference is the
development of seta 1-C in the larva, and this may be within the normal range of
geographic variation. At present there seems to be no reason to suspect that more than
one species exists under the name of vagans, but this will not be known for certain until
considerably more material from southwestern and central Asia is studied in detail.
Blonomics. Little specific information is available on the bionomics of vagans.
In eastern Asia, the immature stages are found in permanent or semipermanent bodies
of fresh ground water, including streams, stream pools, ponds, lakes, and marshes.
Larvae have been collected in Iran in ponds with or without vegetation (Lotfi, 1976). In
Japan and Korea, females are known to bite man at night (Hsiao and Bohart, 1946;
Barnett and Toshioka, 1951). Culex vagans has been reported to be naturally and
40
Contrib.Amer. Ent. Inst.,vol. 24, no. 1, 1988
experimentally infected with larvae of Wuchereria bancrofti (Yamada, 1927, as
tipuliformis Theobald; Manson-Bahr, 1959; Hsiao, 1948; Hsiao and Bohart, 1946) and
should be regarded as a potential vector of this pathogen in southwestern Asia.
Distribution. This species is widespread in eastern and southcentral areas of
the Palaearctic Region. It is known to occur in eastern Siberia, Korea, Japan, Hong
Kong, China, Mongolia, Bangladesh, Nepal, northern India, Pakistan, (?) Iran,
Afghanistan, and (?) Turkey.
Material examined. 83 specimens. A total of 68 specimens (11 females, 10
males, 12 male genitalia, 8 pupal exuviae, 19 larval exuviae, and 8 fourth-instar larvae)
were examined from southwestern Asia - AFGHANISTAN: Bamiyan (Sayedabat);
PAKISTAN (including areas in eastern Pakistan): (Gharian, Gujranwalia, Jhelum,
Khanoharni, Lahore, Sattoki). An additional 15 specimens (3 females, 6 males, and 6
male genitalia) were examined from HONG KONG (the holotype male of virgatipes ),
INDIA (including the holotype female of tipuliformis), and the USSR (the lectotype male
of exilis).
Culex (Culex) torrentium Mart i n i
torrentium Martini, 1925: 336. +Hololectotype male: Schwartza River,
[Thueringia], Germany; designated by Mattingly, 1955a: 31 (BM).
pavlovsky Shingarev, 1928: 52. Type specimen(s) (male): Naryn
District, Romsk, Siberia, USSR (TMS). Synonymy with torrentium by
Edwards, 1932a: 110.
Culex (Culex) exilis Dyar of Martini, 1931: 386 (Palaearctic Region, A, L keys, syn., M’, F,
L); Monchadskii, 1951: 260 (Middle East, L, key, syn., distr.).
Culex torrentiumMartini of Zaini et al., 1983: 117 (Iraq).
Culex (Cdex) torrentium Martini of Edwards, 1926: 136 (Palaearctic Region, A key, A,
distr.); Senevet et al., 1957b: 92 (E. Mediterranean (?), L); Parrish, 1959: 266
(Turkey, distr.); Senevet and Andarelli, 1959: 203 (Mediterranean Basin, A, L, P
keys, syn., M*, F, P*, L*, distr., biol., L assoc.); DuBose and Cur-tin, 1965: 352,
354 (Mediterranean area, A, L keys); Abul-hab, 1969: 249 (Iraq, L key); Lotfi,
1970: 402 (Iran); Lotfi, 1976: 73, 77, 83 (Iran, L key, ecol. note, L*); lbrahim et al.,
1983: 91 (Iraq, L key); Harbach, 1985a: 86, 94, 105 (Iran, Iraq, Turkey, distr., A, L
keys).
This description is based almost entirely on specimens from Sweden.
Adult. A medium-sized species closely resembling pipiens, but slightly larger,
with prealar scales and different male genitalia.
FEMALE. Description based on specimens from Sweden. Head: Antenna1
length about 2.2 mm; pedicel yellow or orange, mesal surface black with some tiny setae
and whitish scales; flagellum normal, whorls with 6 setae, proximal part of flagellomere 1
with some tiny whitish scales on mesal surface. Proboscis mainly black-scaled, ventral
surface with pale scales extending from 0.1 to 0.7 from base. Maxillary palpus usually
entirely black-scaled, palpomere 4 sometimes with some indistinct pale scales on
dorsomesal surface; length 0.4 mm, about 0.2 proboscis length. Falcate scales of
vertex mainly pale yellow, narrower and darker laterally, whitish along margin of
compound eye and on interocular space; forked scales yellow medially, black postero-
Harbach: Subgenus Culex in SouthwesternAsia
41
laterally; lateral spatulate scales yellowish white. Ocular setae dark reddish brown;
interocular setae golden brown. Thorax (Fig. 2C): Pleural integument yellowish brown,
faded posteriorly; scutal integument dark brown. Scutal scales uniformly golden brown
with slight reddish tint, paler on anterior promontory, among supraalar setae, and on
margins of prescutellar area; scales finer on fossa and supraalar area between supraalar
and posterior dorsocentral setae, but not appreciably darker. Scutal setae dark reddish
brown. Scutellum with fine pale yellow falcate scales; 4-7 large setae on each lateral
lobe, 7 or 8 on median lobe. Antepronotum with yellowish falcate scales in upper and
lower patches, upper patch often darker, sometimes golden brown; setae dark reddish
brown dorsally, becoming golden brown ventrally.
Postpronotum with fine
golden-brown falcate scales on dorsal 0.5, paler and slightly coarser posteriorly; with
5-10 dark setae on posterodorsal margin, longer posteriorly. Pleural setae golden
brown: 7-9 upper proepisternal, 8-l 1 prealar, 5-7 upper mesokatepisternal, 7-11 lower
mesokatepisternal, 9-12 upper mesepimeral, and 1 lower mesepimeral. Pleura with
white or nearly white elongate spatulate scales in patches as follows: small patch below
upper proepisternal setae, few to small patch below prealar setae, patches on upper
corner and lower posterior border of mesokatepisternum, anterior patch on
mesepimeron about same level and size as upper mesokatepisternal patch, and patch
mainly before upper mesepimeral setae; rarely with some postspiracular scales. Wing:
Length about 4.7 mm; length of cell R2 3.0-3.8 length of vein R2+3; subcosta intersects
costa at or slightly beyond furcation of vein R2+3; length of cell Ml 0.7-0.8 length of cell
R2; scales entirely dark. Dorsal scaling: broad decumbent scales on costa, R and RI;
narrow decumbent scales on R4+5, M3+4, mcu, CuA and proximal 0.5 of 1A;
subcosta with broad semidecumbent scales; linear plume scales on Rs, R2+3, R2,
R3, M, and Ml+2; near-linear semiplume scales on Ml, M2 and distal 0.5 of 1A; remigium
with 2 distinct rows of broad decumbent scales and 2 or 3 setae distally. Ventral scaling:
narrow decumbent scales on costa, subcosta, Rs, R2+3, M, Ml+2, bases of Ml and M2,
and distal 0.5 of 1A; linear plume scales on Rl, R2, R3, R4+5, beyond bases of Ml and
M2, mcu, M3+4, and CuA distal to mcu; R, CuA proximal to mcu, and proximal 0.5 of 1A
without scales. Halter: Entirely pale, capitellum slightly darker dorsally. Legs: Anterior
surface of forecoxa mainly black-scaled, with small patch of cream-colored scales at base
and usually some scattered pale scales distally, also with many long, dark, ventrally
curved setae, posterior margin with 3 or 4 shorter setae at apex, most proximal of these
more or less perpendicular to surface, others project ventrally; midcoxa with midlateral
longitudinal row of 4 or 5 long golden-brown setae and several shorter ones margined
anteriorly by longitudinal patch of whitish spatulate scales, anterior surface with small
patch of black scales and several short ventrally projecting setae at apex; posterolateral
surface of hindcoxa with longitudinal row of 6-8 golden-brown setae of different lengths,
anterolateral surface with narrow longitudinal row of white spatulate scales and several
ventrally projecting setae at apex. Lateral surfaces of trochanters without scales,
foretrochanter with black scales anteriorly and white scales posteriorly, anterior, mesal
and posterior surfaces of mid- and hindtrochanters with white scales. Apices of femora
with knee spots of yellowish scales; forefemur black-scaled anteriorly, white-scaled
posteriorly; midfemur like forefemur but distal portion with black scales spread over
dorsal surface; hindfemur with complete anterodorsal stripe of black scales, stripe
gradually widening distally to cover about distal 0.5 of dorsal surface but not covering
whole of anterior surface except near apex. Foretibia mainly black-scaled, with
posteroventral stripe of white scales, apex with inconspicuous dorsal spot of whitish
scales; anterior surface of midfemur black-scaled with some faint pale scales at apex,
posterior surface white-scaled; anterior and dorsal surfaces of hindtibia black-scaled with
42
Contrib.Amer. Ent. Inst.,vol. 24, SKI.1, 1988
conspicuous spot of whitish scales at apex, posterior and ventral surfaces with whitish
scales. Tarsi black-scaled, with some indistinct pale scaling posteriorly, particularly on
tarsomere 1 of fore- and midlegs. Pulvilli pale. Ungues black, small, simple. Abdomen:
Terga mainly black-scaled; tergum I golden-brown setose, with posteromesal patch of
black scales; posterior and lateral margins of terga II-VIII with golden to golden-brown
setae; tergum II with small basomedian yellowish spot and lateral patches of white scales;
terga Ill-VII with yellowish basal bands and white basolateral patches, spots slightly
increasing in size on succeeding posterior terga, bands of terga Ill-V convex and usually
not reaching spots, bands of terga VI and VII reaching spots, band of VI usually straight,
that of VII slightly concave; lateral scales of tergum VIII white, dorsal scales entirely
yellowish or black posteriorly. Sterna II-VII mainly yellowish-scaled, with blackish scales
forming a median spot or streak; sternum VIII with lateral patches of whitish scales,
devoid of scales medially; all sterna golden to golden-brown setose.
MALE. Differs from female as follows. Head: Proboscis length about 2.4 mm;
false joint 0.6 from base; ventral pale scales usually extending from 0.4 to 0.7 from base;
without ventral cluster of setae at false joint. Maxillary palpus length about 3.4 mm,
usually extending beyond tip of proboscis by more than length of palpomere 5; mainly
dark-scaled; ventral surface of palpomeres 2 and 3 without scales, integument pale;
palpomere 3 with or without some indistinct pale scaling on lateral surface, with
ventrolateral row of 15-25 long setae on distal 0.33 and a complete row of short
antrorsely curved setae on ventromesal margin; palpomere 4 with complete or nearly
complete ventral line of white scales, distal scales usually losing their color; palpomere 5
with some white scales at base ventrally; lateral and mesal surfaces of palpomeres 4 and
5 densely setose. Falcate scales of vertex of uniform size and color; forked scales less
numerous and shorter, particularly posterolaterally. Thorax: Scutal scales slightly
coarser, scales and integument between supraalar and posterior dorsocentral setae
noticeably darker in dorsal view. Propleuron with 11-l 6 upper proepisternal setae more
or less in 2 rows. Wing: Length about 3.9 mm; length of cell R2 2.0-2.5 length of vein
R2+3; length of cell Ml 0.75-O-85 length of cell R2; subcosta intersects costa before
furcation of vein R2+3. Legs: Forecoxa without scattered pale scales distally. Anterior
unguis longer than posterior unguis on fore- and midlegs, each with midventral tooth;
posterior unguis of midleg smaller than posterior unguis of foreleg, both with small
ventral tooth before middle; hindungues small, simple. Abdomen: Terga without
basolateral spots; basal bands usually 0.4 tergum length (0.25-0.40); bands of terga Ill-V
usually straight, those of terga VI and VII produced posteriorly along lateral scale-free
margins, particularly on tergum VII; tergum VIII (ventral in position) pale-scaled before
posteromesal setose area. Sterna II-VII frequently with dark scales confined to median
area and posterolateral corners; sternum VIII (dorsal in position) pale-scaled, often with
some subtle dark scales which tend to form a pair of submedial spots. Genitalia (Fig. 9):
Form as figured; very much as in vagans, differing in minor details as follows. Ninth tergal
lobe with 5-l 1 setae. Subapical lobe of gonocoxite always with 2 setae in group d-e;
seta g slightly longer, rounded apically. Dorsal arm of lateral plate of phallosome broader,
0.25 distinctly twisted and curved mesad; lateral arm also broader, apical lobe with more
numerous and larger spicules on posterior surface, dorsal process more strongly
developed.
Pupa (Fig. 9). Character and arrangement of setae as figured; range and modal
number of branches in Table 4; closely resembling pipiens from which it cannot be
distinguished with certainty, seta 5-VI apparently stronger than in pipiens, trumpet
exhibiting slight differences in length of tracheoid area and pinna. Cephalothorax:
Lightly tanned, scutum, legs, and metathorax darker. Setae 1,4,6,8,12-CT usually triple,
Harbach:SubgenusCulex in SouthwesternAsia
43
l-CT with 2-4 branches, 4,6-CT with 3-5 branches, 8-CT with l-4 branches and 12-CT
with 2-4 branches; 2,5-CT usually with 4 branches, 2-CT with 2-5 branches and 5-CT
with 3-6 branches; 3,7,1 l-CT usually double, sometimes triple; 9-CT double. Trumpet:
Moderately tanned, subcylindrical, gradually widened distally; index 5.2-6.1, mean 5.5;
tracheoid area darker, extending about 0.4 from base; pinna oblique, shorter than in
pipiens, 0.5-0.7 length of tracheoid area. Abdomen: Lightly tanned, terga I-IV dark in
middle, particularly on anterior margin; length 2.6-3.3 mm, mean 2.9 mm. Seta l-11 most
often with 15 branches (12-16); 1-Ill-V frequently with at least 5 branches (5-9 on IV; 4-9
on V); 5-IV usually triple, sometimes with 4 branches; 5-V-VII normally double, 5-VII rarely
single; 6-III-VI most often triple, 6-lll,lV double or triple, 6-V with 3 or 4 branches, 6-VI
usually with 3 or 4 branches, infrequently with 2 or 5 and generally longer than 6-111-V.
Genital lobe: Tanning and length same as in pipiens. Paddle: Essentially as in pipiens;
length 0.80-l .05 mm, mean 0.91 mm; width 0.53-0.74 mm, mean 0.63 mm; index
1.3-1.7, mean 1.5.
Larva (Fig. 10). Placement and character of setae as figured, range and modal
number of branches in Table 24; extremely similar to pipiens and vagans, differing
chiefly in having setae l-Ill-V and 1-M more branched, also usually distinguishable from
pipiens by the character of setae l-T, 3-I,Vll, and 1-X. Head: Length 0.76-0.86 mm,
mean 0.83 mm; width 1.06-l .23 mm, mean 1.14 mm; moderately tanned, lateralia
slightly darker posterior to eyes; black spot area, dorsomentum and collar heavily
tanned. Dorsomentum usually with 8 teeth (8-10) on either side of median tooth. Seta
7-C most often with 8 or 9 branches (8-10); 9-C usually triple (2-5). Antenna: Generally
longer than in pipiens, length 0.50-0.64 mm, mean 0.57 mm; lightly tanned before seta
l-A, darker at level and distal to l-A, base with dark spot on mesal surface. Seta 1-A with
about 26 branches (21-29). Thorax: Seta 1-M usually double or triple, infrequently with
4 branches, length normally about 0.6 length of 3-M. Seta 1-T normally slightly longer
than 0.5 length of 2-T, often with 4 branches (3-7); 2-T double. Abdomen: Seta 3-I,Vll
usually double, sometimes single on one side, seldom single on both sides; 6-l usually
triple (3,4), 6-11 most often with 3 or 4 branches, sometimes with 5 branches; l-Ill-V
usually with 4 branches, l-VI normally triple, sum of branches for l-Ill-VI always greater
than for pipiens and vagans. Segment VIII: Comb with 31-46 scales, mean 39. Seta
l-VIII frequently with 6 branches (5-9); 3-VIII most often with 7 branches (6-9). Siphon:
Index 4.40-5.67, mean 5.23; shape and tanning as in pipiens. Pecten with 1O-16
spines, mode 13. Seta 1-S usually in 4 pairs; siphon often with a fifth seta 1-S on one or
both sides, rarely with 3 on one side. Segment X: Saddle length 0.32-0.38 mm, mean
0.37 mm; siphon/saddle index 3.92-4.65, mean 4.17. Seta 1-X usually double,
sometimes single on one or both sides, rarely triple; 2-X usually double, occasionally
triple; 4-X in 6 pairs; anal papillae subequal, longer than saddle.
Systematics. The occurrence of torrentium in southwestern Asia is based on
the examination of a few very poor specimens from Iran and reports from the literature.
The description here is based almost entirely on material from Sweden. Although the
occurrence of torrentium in Turkey is probably correct (see Mattingly, 1955a: 31) some
doubt remains about the occurrence of this species in Iran and Iraq. Previous records
from these countries are based on the identification of larvae (Lotfi, 1970, 1976; lbrahim
et al., 1983). Since it has been almost impossible to differentiate the larvae of torrentium
and pipiens using previously published keys, it is not unlikely that these records actually
refer to pipiens. I have seen only one denuded male with associated genitalia, an
unassociated male genitalia, and a single fourth-instar larva from Iran, and I am not
absolutely certain that these are specimens of torrentium. The genitalia may be those of
vagans and the larva that of pipiens. These specimens are only provisionally assigned to
Contrib.Amer. Ent. Inst.,vol. 24, no. 1, 1988
44
torfentiun7.
The female and larva of torrentium have never been easy to distinguish from
pipiens. These species, however, are not as closely related as the outward similarity
would suggest. The two species are well separated in the structure of the male
phallosome. If the morphological variation observed in the different life stages of
torrentium during this study is representative of the holomorphology of the species,
then the characters mentioned in the keys should reliably distinguish most specimens of
torrentium from pipiens in southwestern Asia, assuming of course that torrentium
indeed occurs there.
Bionomics. Very little is known about the biology of torrentium. The immatures
frequently occur with pipiens in Europe. Larvae are commonly found in ground pools,
but are also known to occur in artificial containers such as tin cans, metal tanks, rain
barrels, and concrete ornaments (Service, 1968). Harbach et al. (1985) reported their
occurrence with pipiens in an old boat. This species apparently tolerates some organic
matter and salt in its breeding sites. Collections have been made in rock pools in which
the water was dark brown with decaying leaves (Ribeiro et al., 1977) and in ground pools
with salinities of 0.3-0.4 that of sea water (Lever, 1954). Females are not attracted to
man and apparently feed on birds (Callot, 1957; Service, 1968).
Distribution.
This species is widely distributed in Europe and western Asia,
includingTurkey, (?) Iran, and (?) Iraq.
Material examined.
449 specimens. Only 4 specimens (1 male, 2 male
genitalia, and 1 fourth-instar larva) were examined from southwestern Asia - IRAN:
(Kivi, Sarab-bahran Kazeroon). A total of 445 specimens (73 females, 71 males, 36
male genitalia, 140 pupal exuviae, 102 larval exuviae, and 23 fourth-instar larvae) were
examined from EAST GERMANY (the hololectotype male, allolectotype female, and 2
paralectotypes), PORTUGAL, SWEDEN, and WEST GERMANY.
Culex
(Culex)
decens
Theobald
decens Theobald, 1901c: vii. +Lectotype male, hereby designated,
bearing following data: “Bonny [Nigera] / 29 V 00 // Type // Culex /
decens / (Type). Theobald.“, genitalia on acetate strip (BM).
masculus Theobald, 1901 b: 125. +Lectotype male, hereby designated,
bearing following data: “Wilberforce, / Free Town. / 3.9. 99. / Fr. larva
in / wood-side puddle. // Type // Free Town. / Sierra Leone. / IX. 99. /
E.E. Austen. 99. 267. / Culex / masculus / (Type) Theobald”;
genitalia on acetate strip (BM). Synonymy with decens by Edwards,
1911: 263.
minutus Theobald, 190%: 30. +Lectotype male, hereby designated,
bearing following data: “Type // Transvaal / Dr. Simpson // Culex /
minutus. / (Type). F.V.T. // Recd. from / F.V. Theobald / 1907-29.“;
genitalia on acetate strip (BM). Synonymy with decens by Edwards,
1911: 263.
nigrocostalis
Theobald, 1909: 16. +Lectotype
male, hereby
designated, bearing following data: “Accra / 7-6-08 / c in latrine /
WMG // Culex / nigrocostalis/ Type. [male symbol]. F.V.T / 30. 3. 09.
// Accra, / Gold Coast, / W. Africa. / 7-VI. 1908. / Dr. W.M. Graham. /
1909-271.“; genitalia on acetate strip (BM). Synonymy with decens
by Edwards, 1911: 263.
Harbach: Subgenus Culex in SouthwesternAsia
45
lividocostalis
Graham, 1910: 269.
+Lectotype male, hereby
designated, bearing following data: “Yaba / 3-7-09 / fr. larvae / No 16
I WMG /I Type /I Yaba, I Lagos, 1 W. Africa. I 3. VII. 1909. I D.W.M.
Graham. / 1910-80.“; genitalia on acetate strip (BM). Synonymy with
decens by Edwards, 1911: 263.
Culex decens Theobald of Edwards, 1912c: 381 (Africa, L key); Edwards, 1913b: 58
(Africa, M gen.).
Culex (Culex) decens Theobald of Edwards, 1941: 336, 425, 484 (Afrotropical Region,
A key, M*; F’; P*, syn., distr.); Lewis, 1943a: 282 (Eritrea, L bionomic note);
Lewis, 1943b: 72 (Sudan, A beh.); Lewis, 1945: 21 (Sudan, distr.); Abbott,
1948: 45 (Sudan, L habitat); Lewis, 1948: 144 (Sudan, L habitat); Hopkins, 1952:
320 (Afrotropical Region, L*, bionomics); Knight, 1953a: 229 (Yemen Arab
Republic, A, L, distr.); Lewis, 1956: 715 (Sudan, distr.); Mattingly and Knight,
1956: 110, 113, 120, 127-128, 132 (P.D.R. Yemen, Yemen Arab Republic, A, L
keys, distr., zoogeogr., bion’omics); Mekuria, 1968: 78 (Ethiopia); Harbach,
1985a: 86, 94, 103 (P.D.R. Yemen, Yemen Arab Republic, distr., A, L key).
Adult. Culex decens is distinguished from all other species of the subgenus
Culex occurring in southwestern Asia by the following combination of characters: (1)
proboscis entirely dark-scaled, (2) scutal scales uniform golden brown with reddish tint,
(3) abdominal sterna with broad apical black bands, and (4) maxillary palpus of male
completely dark-scaled. Specimens examined from the Sudan and Uganda differ from
Arabian material in having finer scutal scales which are more evenly dispersed and darker
in color.
FEMALE. Head: Antenna1 length 1.6-2.0 mm, mean 1.8 mm; entirely dark.
Proboscis length 1.8-2.0 mm, mean 1.9 mm; entirely clothed in brownish-black scales,
ventral surface slightly paler; labella yellowish brown. Maxillary palpus entirely
dark-scaled; length about 0.4 mm, approximately 0.2 of proboscis length. Forked scales
of vertex dark; falcate scales pale yellow; lateral spatulate scales off-white; with 2 very
long interocular setae projecting downward over clypeus. Thorax: Integument brown,
pleura paler. Scutum mainly with golden-brown scales with reddish tint, with pale yellow
or whitish scales on anterior and lateral margins and prescutellar area (scutum of
Sudanese and Ugandan specimens with finer, more evenly dispersed, uniform
reddish-brown scales; without distinctly paler scales at margins); setae dark reddish
brown, prominent. Scutellum with pale yellow falcate scales on all 3 lobes; 3 or 4 large
dark setae on each lateral lobe, 5-8 on median lobe. Antepronotum with rather broad
pale yellow falcate scales and dark setae, setae longer anteriorly; postpronotum with
pale yellow to golden-brown falcate scales somewhat intermediate in form between
scutal and antepronotal scales, usually with 4 (sometimes more) well-developed setae in
row along posterior margin at level of mesothoracic spiracle. Pleura with creamy-white
spatulate scales as follows: patch below and mesal to upper proepisternal setae,
patches on upper corner and lower posterior border of mesokatepisternum, and anterior
and upper patches on mesepimeron, upper mesepimeral patch larger than in most
species, these scales being among and around the upper mesepimeral setae; numbers
of pleural setae: 7-l 1 proepisternal, 7-12 prealar, 5-9 upper mesokatepisternal, lo-14
lower mesokatepisternal, 7-13 upper mesepimeral, and 1 (rarely 2) lower mesepimeral.
Wing: Length 3.3-3.6 mm, mean 3.5 mm; length ofcell R2/length of vein R2+3 2.4-3.0,
mean 2.6; length of cell Ml 0.8-l .O length of cell R2; entirely dark-scaled. Halter: Pale
brown. Legs: anterior surface of forecoxa dark-scaled, dark scales continuous with
46
Contrib.Amer. Ent. Inst.,vol. 24, no. 1, 1988
patch of creamy-white spatulate scales at base, coxa with few pale yellow spatulate
scales posterolaterally at apex; mid- and hindcoxae with longitudinal patch of
creamy-white spatulate scales on anterior side of lateral midline. Posterior surface of
trochanters with small, pale yellow spatulate scales. Femora with narrow pale knee
spots; forefemur dark-scaled anteriorly, pale-scaled posteriorly; midfemur dark-scaled
anteriorly and dorsally, pale-scaled posteroventrally; hindfemur largely pale-scaled, with
more or less complete dorsal stripe of dark scales that widens distally onto anterior and
posterior surfaces. Foretibia mainly dark-scaled, paler ventrally, with pale spot
anterodorsally at apex; midtibia also mainly dark, paler posteriorly,with spot of pale scales
dorsally at apex; hindtibia like foretibia but with prominent spot of pale scales at apex.
Tarsi entirely dark-scaled. Abdomen: Tergum I with posteromedial patch of brownish
scales; terga II-VII brown with narrow basal pale bands of off-white scales connected to
large lateral spots of same color, bands 0.01-0.25 tergum length; tergum VIII with narrow
basal pale band. Sterna II-VII with broad apical black bands; sternum VII with pale scales
laterally, without scales medially.
MALE. Differing from female as follows. Head: Maxillary palpus entirely dark;
palpomere 3 with 7-10 ventral setae at apex; palpomeres 4 and 5 densely setose. Falcate scales of vertex less numerous and paler in color. Thorax: Scutal scales paler and
longer. Wing: Length 3.0-3.2 mm, mean 3.1 mm; length of cell R21.8-1.9 length of vein
R2+3; length of cell Ml 0.7-0.9 length of cell R2; scales usually paler, particularly along
anterior veins. Abdomen: Terga essentially as in female, terga without basolateral pale
spots, basal bands produced posteriorly along lateral scale-free margins; tergum VIII
(ventral in position) mainly dark-scaled, with narrow basal pale band, posterior margin
slightly emarginate, densely setose, posterolateral setae very long. Apical dark bands
of sterna II-VII broader, more or less convex, proximal sterna with only basolateral corners
pale-scaled; sternum VIII (dorsal in position) usually nearly completely pale-scaled,
caudal margin narrowly dark-scaled, particularly in middle. Genitalia (Fig. 11): Ninth tergal
lobe small, with 9-l 7 setae (mode 10) in 1 or 2 irregular rows. Gonocoxite normal,
ventrolateral setae strongly developed, longer and stouter than lateral setae, mesal
surface with 4 irregular rows of small setae extending from base to level of subapical
lobe; subapical lobe indistinctly divided, setae d-f borne on slight prominence on mesal
side of main lobe; setae a-c rodlike, a shorter than b and c and more or less straight, b
and c each with stout base, tapered and slightly curved, b slightly stouter than the
others; 3 setae in group d-e, 2 simple, 1 flattened and resembling r; f longer than d-e,
flattened, slightly broader distally; g foliform, broad and strongly asymmetrical; h long,
slender, bent in middle. Gonostylus slender, curved, slightly widened before tip on
lateral side, tip rugose dorsally. Phallosome not much longer than broad, lateral plates
slightly longer than aedeagus; dorsal arm long, tip pointed and bent mesad, diverging
from its mate of the opposite side; lateral arm with 3-5 denticles (mode 4) in more or less
straight line on mesal side of caudal margin, also with a prominent flat lateral lobe, a short
conical dorsal process, and a weak basal articulatory process; ventral arm a sinuous
mesal process with tip pointed and bent caudad. Aedeagus semispherical; ventral
aedeagal bridge narrow. Proctiger normal; paraproct with long curved basal lateral arm,
crown dark with numerous short spinelike spicules and some shorter simple blades.
Cereal sclerite elongate, irregular in shape; 3-5 cereal setae, usually 4. Tergum X
straplike,joining base of paraproct below basal lateral arm.
Pupa (Fig. 11). Character and positions of setae as figured, range and modal
number of branches in Table 5; similar to pipiens and closely related species, the most
obvious distinctions being the form of the trumpet, which is long and slender, and the
character of seta l-11, which has at least 25 branches. Cephalothorax: Setae 3,4CT
Harbach: Subgenus Culex in SouthwesternAsia
47
usually triple, sometimes with 4 branches; 6-CT with 3 or 4 branches, more often with 4;
12-CT usually with 3 or 4 branches (3-5). Trumpet: Long and slender, index 5.5-9.1,
mean 7.1; tracheoid area about 0.4 trumpet length; pinna 0.15-0.25 trumpet length.
Abdomen: Length 2.6-2.9 mm, mean 2.8 mm. Seta l-11 with 25 or more branches,
usually more (25-32); seta 5-IV of same order of thickness and length as 5-V,VI, with 3 or
4 branches, more often with 3; 6-111double or triple, more often double, 6-IV-VI usually
triple, sometimes with 4 branches, rarely double. Genital lobe: Rather short in male,
length about 0.33 mm. Paddle: Length 0.74-0.92 mm, mean 0.82 mm; width 0.48-0.60
mm, mean 0.57 mm; index 1.3-1.5, mean 1.4.
Larva (Fig. 12). Placement and character of setae as figured, range and modal
number of branches in Table 25; immediately distinguished from all other Culex in
southwestern Asia by the long, slender siphon and very long, single seta l-IV,V. Head:
Length 0.70-0.81 mm, mean 0.76 mm; width 1.00-l .21 mm, mean 1.13 mm; lightly
tanned, darker behind eyes, dorsal apotome sometimes with small dark median spot
posterior to bases of seta 5-C, less frequently with a pair of dark sublateral spots before
median spot. Dorsomentum moderately tanned; usually with 8 teeth (6-8) on either side
of median tooth, lateral teeth longer. Seta O-C well developed, distinct; 1-C lightly to
moderately tanned, tapered to sharp point, length less than 0.1 mm; 2-C absent; 5,6-C
double, 5-C seldom triple; 7-C usually with 6 or 7 branches (5-7); 14-C normally double
with strongly divergent branches (l-3). Antenna: Length 0.52-0.61 mm, mean 0.55
mm; lightly tanned, distal part darker, base with dark spot on mesal surface. Seta 1-A
with about 27 branches (21-32). Thorax: Seta 7-P triple. Seta 1-M usually single,
seldom double, short, about 1.5 length of 2-M; 9-M with 3 or 4 branches, more often
with 4. Seta 1-T usually single, sometimes double; 2-T normally double (1,2).
Abdomen: Seta 3-I,VII double, 3-VII rarely triple; 6-I,11triple, 6-lll,V,VI single; 6-IV normally
double (l-3); 1-III,VI developed as usual, l-111either single, double or triple, l-VI usually
triple, sometimes double; 1-IV,V strongly developed, single and very long, much longer
than length of 2 abdominal segments. Segment V//I: Comb with 43-65 scales, mean 52.
Siphon: Very long and slender, index 8.75-l 1.27, mean 9.96; lightly to moderately
tanned. Pecten on basal 0.2, with 11-16 spines, mode 14; larger spines with 4 or 5
denticles. Seta 1-S usually in 4 pairs (a fifth seta occasionally occurs on one or both
sides of siphon), usually double or triple (l-3), length about same as diameter of siphon
at point of attachment. Segment X: Saddle lightly to moderately tanned; length
0.36-0.48 mm, mean 0.39 mm; siphon/saddle index 5.43-7.20, mean 6.36. Seta 1-X
usually double, occasionally single. Anal papillae subequal, slightly shorter than saddle.
I am recognizing decens as the nominotypical member of a
Systematics.
subgroup which includes invidiosus Theobald, 1901 c, antennatus, ornatothoracis
Theobald, 1909, trifoliatus Edwards, 1914a, perfuscus Edwards, 1914a, perfidiosus
Edwards, 1914a, telesilla De Meillon and Lavoipierre, 1945 (in De Meillon et al., 1945)
and litwakae Harbach, 1985b. Except for antennatus, the members of this group are
restricted to the Afrotropical Region. The group as a whole is poorly known and it may
be necessary to divide it into two or more subgroups when the species are studied in
detail. The subgroup is recognized primarily on the basis of overt similarities in adult
ornamentation and the structure of the male genitalia. There appear to be some striking
differences in the larvae.
The occurrence of decens in southwestern Asia (the Yemen Arab Republic) was
first recorded by Knight (1953a) who noted that Yemenite material resembles the African
material examined by Edwards (1941) except that the ventral surface of the proboscis
and the scutal scaling are lighter in some specimens. I have noticed that the scutal
scales of some specimens are slightly coarser and without the distinct reddish tint found
48
Contrib.Amer. Ent. Inst.,vol. 24, no. 1, 1988
in specimens from Sudan and Uganda.
The larva of decens is quite different from other Culex occurring in southwestern
Asia. It is easily recognized by the long, slender siphon and the long, single seta 1-IV,V.
Hopkins (1952) did not mention seta 1-IV,V in his description of decens so it is possible
that this character may serve to distinguish decens from invidiosus, a species which is
supposedly indistinguishable from decens in Africa. Seta 1-IV,V is similarly developed in
larvae of decens examined from several African countries. No specimens identified as
invidiosuswere examined during this study.
Bionomics. Larvae of decens have been taken from ground waters, such as
pools, swamps, rock holes, ditches, and streams, and a variety of artificial containers,
including cement basins, tubs, drains, wells, troughs, and household pots. The early
reports of decens breeding in tree holes, cut bamboo, and crabholes probably refer to
another species. This species has been collected in association with laticinctus and
Culex (Lutzia) tigripes De Grandpre and De Charmoy in the Yemen Arab Republic
(Knight, 1953a). Collection data seen during this study indicate that resting adults have
been collected in a house and an unused well. Lewis (1943b) observed that decens is
common in the Anglo-Egyptian Sudan, but is never seen biting man. Lumsden and van
Someren (1953) reported that a single female was collected during landing catches
made 11 m above the ground in riverain forest in Uganda. This species probably is not
involved in disease transmission.
Distribution. Culex decens is very widely distributed in the Afrotropical Region.
Material examined.
129 specimens. A total of 50 specimens (8 females, 9
males, 11 male genitalia, 9 pupal exuviae, 10 larval exuviae, and 3 fourth-instar larvae)
were examined from southwestern Asia - P.D.R. YEMEN: (Mukalla); YEMEN ARAB
REPUBLIC: (Ta’izz). An additional 79 specimens (17 females, 18 males, 15 male
genitalia, and 29 fourth-instar larvae) were examined from GHANA (type specimens of
nigrocostalis ), IVORY COAST, KENYA, LIBERIA, MAURITANIA, NIGERIA (type
specimens of decens and lividocostalis ), SENEGAL, SIERRA LEONE (including the
type series of masculus ), SOUTH AFRICA (the type male of minutus ), SUDAN,
UGANDA, and ZAIRE.
Culex
(Culex)
antennatus
(Becker)
antennatus Becker, 1903: 68 (Anopheles ). Type specimen(s) (male):
Cairo, Egypt (ZM).
laurenti Newstead, 1907 (in Newstead et al., 1907: 24). +Lectotype
male, hereby designated, bearing following data: “SYN- / TYPE //
COTYPE / [male symbol] // SCHOOL OF TROP. MED., /
LIVERPOOL UNIVERSITY / Culex laurenti. / Leopoldville, / Belgian
Congo, / December 1903. / Drs. Dutton & / Todd // SYNTYPE of /
Culex laurenti / Newstead / det. J. Chainey 1975”; genitalia on acetate strip (BM). Synonymy with antennatus by Hopkins, 1936: 224.
Culex antennatus (Becker) of Lewis, 1945: 20 (Sudan, distr.); Theodor, 1952: 113
(Middle East, zoogeogr.); El-Said and Kenawy, 1983a (Egypt, distr.); Zimmerman
et al., 1985: 84 (Egypt, bionomics); Kitron and Pener, 1986 (Israel, L bionomics).
Culex (Culex) antennatus (Becker) of Edwards, 1941: 333, 425, 484 (Afrotropical
Region, A key, M’; F*; P, distr.); Lewis, 1943b: 72 (Sudan, A beh.); Lewis, 1948:
145-148 (Sudan, A beh., L habitat); Hopkins, 1952: 320 (Afrotropical Region, L
Harbach: Subgenus Culex in SouthwesternAsia
49
key, L, bionomics); Lewis, 1956: 715 (Sudan, distr., zoogeogr.); Ovazza et al.,
1956: 176 (Ethiopia); Senevet et al., 1957b: 92 (North Africa, L); Senevet and
Andarelli, 1959: 215 (North Africa, A, P, L keys, syn., M*, F, P*, L*, distr., biol., L
assoc.); DuBose and Cur-tin, 1965: 352, 354 (Mediterranean area, A, L keys);
Mekuria, 1968: 77 (Ethiopia, distr.); Lotfi, 1970: 401 (Iran); Margalit and Tahori,
1970b: 151 (Israel, toll. sites); Margalit and Tahori, 1974: 56 (Israel, toll. rec.);
Lotfi, 1976: 73, 76, 82 (Iran, L key, ecol. note, L*); El-Said and Kenawy, 1983b
(Egypt, distr.); Harbach, 1985a: 86, 94, 104 (Egypt, Iran, Israel, Egypt, distr., A, L
keys).
Culex (?) invidiosusTheobald of Gough, 1914: 135 (Egypt, bionomic note, toll. sites).
Culex invidiosusTheobald of Storey, 1918( 1919): 100 (Egypt, A, L keys, A, bionomics).
Culex laurenti Newstead of Buxton, 1923: 317 (Israel, Jordan, toll. rec.); Kirkpatrick,
1924(1925): 369, 372 (Egypt, A, L keys); Salem, 1938: 27 (Sinai); Abdel-Malek,
1956: 105 (Sinai, L key).
Culex (Culex) laurenti Newstead of Edwards, 1921: 344 (Egypt, Israel, Jordan, A key,
tax., distr.); Seguy, 1924: 39, 190 (Egypt, Israel, Jordan, A key, M*, F, tax., distr.,
syn.); Kirkpatrick, 1925: 129 (Egypt, A, P, L keys, M’, F, P*, L’, bionomics);
Edwards, 1926: 141 (Egypt, Israel, Jordan, A, L keys, A, distr.); Stackelberg,
1927: 164 (Middle East, M, F keys, A, M gen.*, distr.); Martini, 1931: 370 (Egypt,
Jordan, A, L keys, M’, F, L*, distr.); Monchadskii, 1951: 265 (Middle East, L key,
L*, bionomics, distr., syn.).
Adult. A small pale brown species easily recognized by the prominent lateral
pale stripes on abdominal terga VI and VII and lateral pale spots on the other terga. The
proboscis of the male bears a ventral cluster of setae near the middle. This species
superficially resembles pusillus Macquart of the subgenus Barraudius, but the
abdominal terga of the latter are darker, black or nearly so, and the proboscis of the male
lacks a midventral cluster of setae.
FEMALE. Head: Antenna blackish; pedicel with some tiny scales and/or setae
on mesal side. Proboscis dark-scaled dorsally, with paler scaling ventrally, particularly on
distal 0.5; length 1.6-l .8 mm, mean 1.7 mm. Maxillary palpus dark-scaled, darker at tip;
length approximately 0.2 proboscis length. Forked scales of vertex numerous, usually
dark; falcate scales pale yellowish white; lateral spatulate scales narrow, white. Thorax:
Scutum brown; uniformly clothed in fine, golden-brown falcate scales, with some pale
yellow or whitish scales on anterior promontory and prescutellar area. Scutellum with
very fine pale yellow falcate scales on lateral and median lobes. Pronotum brown;
antepronotum with fine pale yellow falcate scales; similar scales on postpronotum but
slightly darker anteriorly. Pleura brownish yellow; numbers of pleural setae: 6-l 2 upper
proepisternal, 6-9 prealar, 4 or 5 upper mesokatepisternal, 5-9 lower mesokatepisternal,
5-8 upper mesepimeral, and 1 lower mesepimeral; patches of narrow cream-colored
spatulate scales present as follows: small patch below upper proepisternal setae, upper
and lower patches on mesokatepisternum, and anterior and upper patches on
mesepimeron. Wing: Length 2.7-3.0 mm, mean 2.9 mm; infrequently with some pale
scales on posterior side of costa before humeral crossvein, otherwise entirely
dark-scaled; length of cell R2 approximately 4.5 length of vein R2+3; length of cell Ml
about 0.8 length of cell R2. Halter: Pale, more or less same color as pleura. Legs:
Anterior surface of forecoxa with dark scales in middle, off-white scales at base and apex;
mid- and hindcoxa each with longitudinal patch of small cream-colored scales on anterior
side of lateral midline. Forefemur dark brown-scaled anteriorly, pale-scaled posteriorly,
with inconspicuous knee spot of yellow scales; midfemur same but dark brown scaling of
50
Contrib.Amer. Ent. Inst.,vol. 24, no. 1, 1988
anterior surface gradually expanded over dorsal surface on distal 0.5; hindfemur mainly
pale-scaled, with dorsal line of dark scales (often faint proximally; widened distally,
particularly on distal 0.5) and narrow yellow knee spot (usually more distinct than on foreand midfemora). Foretibia dark brown to blackish-scaled dorsally, with off-white scales
ventrally; anterior surface of midtibia dark brown-scaled, posterior surface with yellowish
white scales; hindtibia same as foretibia but sometimes with faint apical spot of pale
scales. All tarsi dark-scaled, somewhat paler scaling ventrally. Abdomen: Tergum I with
inconspicuous median posterior patch of cream-colored scales; terga II-VII almost
entirely brown-scaled; terga II-V with small basolateral patches of cream-colored scales
which are usually not visible in dorsal aspect, those of tergum V sometimes partly visible;
lateral borders of terga VI and VII with stripes of cream-colored scales; tergum VIII largely
pale-scaled. Sterna with cream-colored scales; sternum VIII with lateral patches only,
median area without scales.
MALE. Like female except as follows. Head: Ventral surface of proboscis usually
entirely pale-scaled, conspicuously so on distal 0.5; with ventral cluster of setae on
proximal side of false joint; length 1.7-l .9 mm, mean 1.8 mm. Maxillary palpus dark,
palpomeres 4 and 5 darker; length about 1.3 proboscis length, extending beyond tip of
proboscis by length of palpomere 5; palpomere 3 with lateral patch of pale scales before
apex; ventral surface of palpomere 4 with spot of white scales at base and short line of
similar scales before apex, these sometimes joined; palpomere 5 noticeably longer than
4 (about 1.4 length of 4), with spot of white scales ventrally at base. Lateral spatulate
scales of head slightly smaller and more numerous. Thorax: Scutal scales slightly finer.
Wing: Length 2.5-3.0 mm, mean 2.8 mm; length of cell R2 about 1.6 length of vein
R2+3. Abdomen: Terga II and III normally entirely dark-scaled; tergum VIII (ventral in
position) and sternum VIII (dorsal in position) largely or completely with cream-colored
scales. Genitalia (Fig. 13): Form as figured; basic structure as in decens, differing chiefly
as follows. Ninth tergal lobe with 6-14 setae (mode 8). Subapical lobe of gonocoxite
with 3 setae in group d-e, 2 longer setae with apical series of 2 or 3 (usually 3) recurved
projections, shorter seta simple and slightly flattened; seta g much as in decens,
acuminate. Gonostylus broader distally. Phallosome longer than broad with lateral
plates slightly longer than aedeagus; dorsal arm slightly sinuous in lateral view; caudal
margin of lateral arm with tight row of 3-5 denticles (usually 4) projecting over prominent
lateral ridge, end of ridge differentiated as a conspicuous dorsal process with narrowed
and slightly bent apex, ventral arm without prominent flat lateral lobe, basal articutatory
process more strongly developed; ventral arm a prominent rounded ventrocaudal lobe
with a short dorsally directed hornlike projection, ventrocaudal surface with many minute
spinules, dorsal projection slightly bent caudad and resembling denticles of lateral arm in
dorsal view. Aedeagus conical; ventral aedeagal bridge slightly broader. Basal lateral
arm of paraproct shorter, flattened, and broad. Cereal sclerite usually with 2 setae (l-3)
on posterolateral corner.
Pupa (Fig. 13). Character and positions of setae as figured, range and modal
number of branches in Table 6; resembling pipiens in most respects, but generally
smaller and setae 3-VII and 6-111
with more branches; extremely similar to members of the
univittatus and simpsoni subgroups, from which it cannot be distinguished with
confidence. Cephalothorax: Seta 5-CT frequently with 5, 6, or 7 branches (4-8); 8-CT
frequently with 6 or 7 branches (4-9); 9-CT often with 3 branches (2-4); 12-CT often with
5 branches (2-5). Trumpet: Much as in pipiens, index 4.3-6.5, mean 5.2; tracheoid area
about 0.4 trumpet length; pinna oblique, 0.2-0.3 trumpet length. Abdomen: Length
2.6-3.3 mm, mean 2.9 mm. Seta l-11 with considerable variation in number of branches
(7-41); 5-IV not developed quite as strongly as 5-V,VI, often with 5 branches (4-6); 6-111,
Harbach: Subgenus Culex in SouthwesternAsia
51
IV most often with 3 or 4 branches (3-6), 6-V,VI most often with 5 or 6 branches (3-6).
Paddle: Length 0.80-l .Ol mm, mean 0.90 mm; width 0.56-0.70 mm, mean 0.63 mm;
index 1.3-l .5, mean 1.4.
Larva (Fig. 14). Placement and form of setae as figured, range and modal
number of branches in Table 26; differing from pipiens and allied species and
resembling univitfatus and perexiguus in having seta 5-C usually double or triple, seta
6-VI normally single, and seta 1-S no longer than diameter of siphon. Head: Length
0.64-0.75 mm, mean 0.67 mm; width 1.02-l .16 mm, mean 1.lO mm; tanning, including
spots on dorsal apotome, almost exactly as in pipiens. Dorsomentum usually with 8
teeth (7-9) on either side of median tooth. Seta 1-C long, slender, moderately tanned,
length about 0.1 mm; 2-C absent or represented by a small bump; 5,6,1 I-C double or
triple; 7-C with 6, 7, or 8 branches occurring in nearly equal frequencies; 10-C usually
triple, sometimes with 4 branches; 14-C single or double, branches strongly divergent
when double. Antenna: Length 0.44-0.58 mm, mean 0.50 mm; moderately tanned at
base and on distal part from just before seta 1-A; spicules rather broad, not true aciculae;
scape developed; puncture not evident. Seta 1-A with about 20 branches (17-28).
Thorax: Seta 7-P triple. Seta 1-M most often double, frequently triple, sometimes
single, very short, about same length as 2-M. Seta 1-T short, about same length as l-M,
usually double or triple (l-3); 2-T double or triple. Abdomen: Segments III and V with
black pigment granules giving abdomen a 2-banded appearance to the unaided eye.
Seta 3-l usually triple (2-5), 3-VII usually with 4 or 5 branches (3-5); 6-I,11normally triple, 6-11
occasionally with 4 branches, 6-111usually double (l-3), 6-IV triple, 6-V double or triple,
more often double, 6-VI single; 1-III with 3 or 4 branches, l-IV-VI usually triple, 1-IV,V
seldom with 4 branches, l-V,VI sometimes double. Segment VIII: Comb with 32-46
scales, mean 38. Siphon: Long, slender, slightly tapered; index 5.47-8.14, mean 6.78.
Pecten on approximately basal 0.25, with lo-14 spines, mode 13; larger spines with 2 or
3 basal denticles. Seta 1-S normally in 5 or 6 pairs, 2 or 3 posterolateral and 3 lateral,
length about 0.5 diameter of siphon at point of attachment. Segment X: Saddle length
0.36-0.40 mm, mean 0.38 mm; siphon/saddle index 3.77-4.99, mean 4.35. Seta 1-X
with 3 or 4 branches, more often with 4; seta 2-X double or triple, more often double.
Anal papillae subacutely tapered, about length of saddle.
Systematics. Culex antennatus is an interesting species which shares some
features with the adults of both the pipiens and the decens subgroups. It is included as
a member of the decens subgroup because the male genitalia suggest a closer affinity
with decens than pipiens. The larva of this species bears a close resemblance to those
of univiftatus and perexiguus which suggests a relationship with these species. It is
possible, therefore, that antennatus has retained some generalized features of an
ancestral stock which may have given rise to the pipiens, decens and univittatus
subgroups. There is some suggestion of a distant relationship with members of the
sitiens group in the structure of the male phallosome. Once the Culex fauna of Africa
has been studied thoroughly and species affinities are more clearly understood, it may
be necessary to include antennatus and the closely related litwakae Harbach, 1985b, in
a separate subgroup.
The adults of antennatus are superficially very similar to those of Culex
(Barraudius ) pusillus Macquart, 1850, but are easily distinguished by the smaller
forefemur/proboscis ratio and the much longer hindtarsomere 1. These species are
sympatric thoughout the range of antenna&s in southwestern Asia and Egypt. Females
of antennatus are almost indistinguishable from unbanded females of pipiens, but the
males, larvae, and pupae of these species are easily differentiated. The larva of
antennatus is extremely similar to that of perexiguus which usually, but not always, dif-
52
Contrib.Amer. Ent. inst.,vol. 24, no. 1, 1988
fers in the character of setae 1-C, l-M, and 1-S. Live larvae of antennatus are readily
distinguished from those of perexiguus by the two-banded appearance of the
abdomen.
Bionomics. Culex antennatus mainly breeds in stagnant bodies of fresh water,
but is sometimes found in brackish water pools. it rarely occurs in polluted water.
Common larval habitats include rice fields, stream pools, springs, ponds, swamps,
ditches, seepages, and animal footprints. The breeding places usually contain
emergent vegetation. Larvae are often found in association with theileri, poicilipes,
perexiguus, and pipiens. Females feed on mammals. They appear to feed primarily on
bovids, but also attack sheep, goats, horses, donkeys, and man (Zimmerman et al.,
1985). The species bites both indoors and outdoors in the evening, and seeks resting
sites away from buildings (Lewis, 1948; Zimmerman et al., 1985). Culex antenna&s has
been found naturally infected with West Nile virus in Egypt (Taylor et al., 1953; Taylor et
al., 1956) and Rift Valley Fever virus in Nigeria (Lee, 1979). Successful experimental
transmission has been demonstrated for both of these viruses in the laboratory (Taylor
et al., 1953; Gad et al., 1987, respectively). Arumowat virus was isolated from a pool of
unfed females in the Sudan (Karabatsos, 1985), and there is serological evidence for
human infection of this virus in Egypt, Sudan, and Somalia (Tesh et al., 1976). immature
filariai infections have been found in this species in Tanzania (Smith, 1955) and Egypt
(A. Gad, personal communication).
Distribution.
This species occurs in the Middle East and the Afrotropical
Region south to Botswana.
Material examined. 885 specimens. A total of 849 specimens (211 females,
175 males, 13 male genitalia, 310 pupal exuviae, 112 larval exuviae, and 28 fourth-instar
larvae) were examined from southwestern Asia - EGYPT: Aswan (Aswan, El Aqaba ei
Saghira, Ezbet ei Silsiia, Khour Abu Subeira, Nag ei Kagug, Nag ei Ritag el Qibii), Asyut
(Beni Shiqeir, El Qusiya), El Fayyum (Abhit ei Haggar, Birket Qarum, Biyahmu, El Nazia,
Sinero ei Bahria, Tubhar, unknown localities), El Foadia (Baitim, Nimra ei Basal), El
Gharbiya (Saft Turab), El Giza (Ezbet Bosna Sharawy, Harania, Koniessa, Naziet ei
Ashtar, Talbia), El Isma’iliya (Abu Khaiifa, Nifisha), N Qahira (Cairo), El Qalyubka (Bahtim,
El Salmaniya, Ga’afara, Kaha, Kafr Hamza, Sindiwa), El Sharqiya (inshas et Rami,
Khirbetrama), Kafr el Sheikh (Kafr ei Sheikh, Makallet ei Qasab); IRAN: (Bisotun,
Kamroud, Kernianshah); IRAQ: (unknown localities); ISRAEL: (Lake Tiberias, Ludd,
Rosh Hal Ayn, many unknown localities); JORDAN: (AZ Zarqa, unknown localities);
TURKEY: (unknown localities). An additional 36 specimens (18 females, 7 males, 9
male genitalia,1 pupal exuviae, and 1 larval exuviae) were examined from ALGERIA,
MAURITANIA, SENEGAL, SUDAN, TUNISIA, ZAIRE (type specimens of /awent/), and an
unknown country.
Culex (Culex) univittatus
T he o b a Id
univittatus Theobald, 1901 b: 29. +Lectotype female: Saiisbury, Zimbabwe; designated by White, 1975: 321 (BM).
simplex Theobald, 1903a: 337 (Heptaphlebomyia ). +Hoiotype female:
Salisbury, Rhodesia (BM). Synonymy with univittatus by Edwards,
1911: 262.
montforti Ventriiion, 1905a: 448 (Heptaphlebomyia ). +Lectotype male:
Andajobe, Madagascar; designated by White, 1975: 321 (BM).
Synonymy with univittatus by Edwards, 1911: 262.
Harbach: Subgenus Culex in SouthwesternAsia
53
ataeniatus Theobald, 1911: 261. +Holotype female: Onderstepoort,
Transvaal (BM). Synonymy with univittatus by White, 1980: 137.
Culex univittatus Theobald of Edwards, 1912b: 28, 32 (in part; Afrotropical Region, A
key, distr.); Edwards, 1912c: 381 (in part; Africa, L key); Edwards, 1913b: 58
(inpart; Africa, A, M gen.); Edwards, 1914a: 67 (Africa, M gen?); Mattingly, 1954:
56 (typical form, in part; Africa, zoogeogr., M gen.*).
Culex (Culex) univittatus Theobald of Seguy, 1924: 39, 190 (in part; Africa, tax., M
gen.*, distr.); Edwards, 1941: 306, 419, 482 (typical form, in part; A key, M*, F*,
P, syn., distr.); Lewis, 1943a: 282 (in part; Eritrea, toll. rec.); Hopkins, 1952: 291
(? in part; Afrotropical Region, L); Mattingly and Knight, 1956: 104, 122 (in part;
Yemen Arab Republic, A, L keys); Jupp, 1970: 9-l 6 (in part, Highveld form;
South Africa, M’, F, distr., crossmatings, beh.); Jupp, 1971: 339-356 (South
Africa, M gen.*, F*, distr., crossmatings); Jupp, 1972 (excluding short-spined
form; Africa, M gen.*, F); White, 1975: 320-322 (Afrotropical Region, syn., distr.,
A key); Harbach, 1985a: 86, 93, 104 (P.D.R. Yemen, Yemen Arab Republic,
distr., A, L keys).
Culex (Culex) univittafus var. neavei Theobald of Knight, 1953a: 232 (Yemen Arab
Republic, toll. rec.).
The description of this species is based largely on the examination of topotypic
material and the type series. The occurrence of univittatus in southwestern Asia is
based on the examination of a single male from the P.D. R. Yemen and a single female
from the Yemen Arab Republic.
Adult. This species might be confused with sinaiticus or simpsoni, but differs in
the character of the scutal scaling and by the presence of an anterior pale stripe on the
hindtibia. It is closely allied to perexiguus from which it is usually distinguished by the
presence of a complete anterior pale stripe on the midfemur, and a line of scales on vein
2A in the female, and slightly different genitalia in the male.
FEMALE. Head: Length of antenna 1.5-l .9 mm, mean 1.7 mm; pedicel pale
laterally, dark mesally with patch of white scales; flagellum dark, flagellomere 1 with some
white scales mesally just above base, most flagellar whorls with 4 long dark setae.
Proboscis black-scaled, with whitish scales ventrally in middle; length 1.8-2.3 mm, mean
2.0 mm. Maxillary palpus black-scaled, with some white scales mesally at apex,
palpomeres 3 and 4 often with some white scales laterally; length about 0.2 proboscis
length. Falcate scales of vertex pale golden brown to white, usually paler medially;
interocular scales concolorous with white or whitish ocular scales; forked scales
cream-colored medially, golden brown to brown posterolaterally; lateral spatulate scales
white. Thorax: Integument brown. Scutal setae golden brown to brown; scutal scales
narrow, somewhat dense, golden brown; some pale yellow to white scales on margins
and prescutellar area, pale scales often forming pair of faint to distinct submedian spots
near middle of scutum. Scutellar scales same color as prescutellar scales. Falcate scales
of antepronotum golden brown medially, paler and slightly coarser dorsally and
especially ventrally, color of setae same as scales; postpronotum with narrow,
golden-brown scales becoming paler posteriorly, with 4 or 5 long dark setae along
posterodorsal margin. Pleural setae golden: 6 or 7 upper proepisternal, 10 or 11
prealar, 4-6 upper mesokatepisternal, 7-10 lower mesokatepisternal, 7 or 8 upper
mesepimeral, and 1 lower mesepimeral. Pleura with broad white spatulate scales: small
patch below upper proepisternal setae extending well mesad of setae on anterior
surface, patch of narrower scales on upper portion of postspiracular area sometimes se-
54
Contrib.Amer. Ent. Inst.,vol. 24, no. 1, 1988
parated into anterior and posterior clusters, small patch below prealar setae continuous
with upper mesokatepisternal scales, large patches on upper corner and lower posterior
border of mesokatepisternum, large anterior patch on mesepimeron at level of upper
mesokatepisternal scales and patch before and among upper mesepimeral setae. Wing
(Fig. 2D): Length 3.6-4.1 mm, mean 3.8 mm; length of cell R2/length of vein R2+3
2.7-3.8, mean 3.3; length of cell Ml/length of cell R2 0.75-0.79, mean 0.78; scales
entirely dark except for short line of white scales on posterior margin of costa at level of
humeral crossvein; vein 2A usually with scales on dorsal surface, scales usually
extending entire length of vein. HaIter: Pedicel and scabellum whitish; capitellum
blackish. Legs (mid- and hindlegs, Fig. 21,K): Mainly black-scaled. Anterior surface of
forecoxa with white scales at base and apex and black scales in middle, with many long
golden-brown to brown setae among median black and distal white scales, with 3 or 4
shorter setae and few pale scales posteriorly at apex; midcoxa with midlateral row of 5 or
6 well developed golden-brown to brown setae margined anteriorly by longitudinal patch
of white spatulate scales, with small patch of black scales and several short ventrally
projecting setae anteriorly at apex; hindcoxa with posterolateral row of normally 8 long
golden to golden-brown setae, anterolateral surface with longitudinal patch of white
spatulate scales and several short ventrally projecting setae at apex. Trochanters with
white scales on posterior and dorsal surfaces. Each femur with narrow pale knee spot:
anterior surface of forefemur black-scaled, usually with complete or incomplete narrow
ill-defined or broken anteroventral white stripe, posterior surface entirely white-scaled;
midfemur like forefemur but with complete distinct or ill-defined anteroventral white
stripe and black scales extending over dorsal surface toward apex; hindfemur largely
white-scaled, with anterodorsal stripe of black scales beginning beyond base and
widening distally, stripe abruptly expanded over anterior and posterior surfaces on distal
0.2 to form subapical band. Anterior and dorsal surfaces of foretibia black-scaled,
posterior and ventral surfaces whitish-scaled; frequently with small dorsal whitish spot at
apex; midtibia mainly black-scaled, whitish-scaled posteriorly, usually with narrow
anterodorsal stripe of whitish scales that usually reaches small dorsal whitish spot at
apex; hindtibia black-scaled with narrow anterior and posterior whitish stripes on proximal
0.8, with distinct white spot at apex. Tarsi black-scaled, tarsomeres 1 and 2 usually
noticeably paler ventrally. Abdomen: Terga mainly black-scaled; tergum I with median
posterior patch of black scales usually bordered anteriorly by some whitish scales; terga
II-VII with convex basal bands and basolateral spots of white scales, bands normally
0.25-0.35 tergum length; bands usually not reaching spots on tergum II, barely touching
spots on terga II-IV and joining them on terga V-VII, spots progressively larger from
anterior to posteror terga; tergum VIII frequently with posterior band of black scales, but
often entirely whitish-scaled. Sterna II-VII mainly with cream-colored to white scales,
usually with black scales on midline and posterolateral corners; sternum VIII with dark
integument and lateral patches of white scales.
MALE. Differing from the female as follows. Head: Proboscis usually entirely
black-scaled, occasionally with some whitish scales at false joint, without ventral cluster
of setae at false joint. Maxillary palpus mainly dark; midregion of palpomere 3 with narrow
ventrolateral line of dingy-white scales, with few whitish scales ventromesally at apex;
palpomere 4 with ventral white stripe narrowing to apex, also with indistinct or faint
dorsolateral line of whitish scales; palpomere 5 with ventral spot of white scales at base.
Falcate scales of vertex coarser, forked scales shorter. Thorax: Submedial spots of
scutum extremely faint or absent. Wing: Length 3.1-3.6 mm, mean 3.3 mm; length of
cell R2/length of vein R2+3 1.6-2.8, mean 2.1; basal line of white scales on costa ex-
Harbach: Subgenus Culexin SouthwesternAsia
55
tremely reduced or absent; vein 2A usually without scales. Legs: Anterior white stripe
of midfemur sometimes less distinct. Abdomen: Basal white bands of terga generally
broader, less convex; basolateral spots absent; terga V-VII with posterior extension of
bands along lateral scale-free margins. Black scaling of sterna more extensive, pale
scaling often reduced to large basolateral spots; sternum VIII (dorsal in position) with
basal pale band or entirely pale-scaled. Genitalia (Fig. 15): Form as figured; lateral plate
of phallosome generally constructed as in vagans and torrentium. Ninth tergal lobe
small, with 6-16 (mode 11) setae. Gonocoxite normal; ventrolateral and mesal surfaces
with usual complement of large and small setae respectively, lateral surface with 1 or 2
rows of moderately long slender setae and several rows of small setae between these
and the large ventrolateral setae; subapical lobe slightly divided, proximal part
prominent, bearing setae a-c of usual form, b slightly stouter than a and c; setae dand e
fine, hairlike and inconspicuous, shorter than f, fa blunt rod with slightly enlarged tip; g
foliform as usual, asymmetrical, longer than broad, apex not sharply produced; h
slender, bent in middle. Gonostylus slightly expanded laterally near tip. Phallosome
longer than broad, aedeagal sclerite shorter than lateral plate; dorsal arm of lateral plate
rather long, slender, and slightly sinuous in dorsal view; lateral arm strongly developed,
with conspicuous angled scooplike caudal extremity and prominent dorsal process;
ventral arm spinelike, curved dorsad, and projecting slightly beyond caudal margin of
lateral arm. Proctiger normal; paraproct with long, curved, distally flattened basal lateral
arm, crown mostly of small spinelike spicules but with some inconspicuous lateral blades.
Cereal sclerite and tergum X unmodified; l-3 cereal setae, usually 2.
Pupa (Fig. 15). Not associated with adults from southwestern Asia, specimens
examined primarily from South Africa; character and placement of setae as figured, range
and modal number of branches in Table 7; not immediately distinguishable from pipiens
and related species, differing chiefly in the number of branches of setae 6-111and 3-VII,
resembling theileri, perexiguus, antennatus, simpsoni, and sinaiticus in this respect.
Cephalothorax: Seta 5-CT rather short, about same length and thickness as 4-CT; 6-CT
with 3-5 branches occurring in nearly equal frequencies; 9-CT double or triple; 1 l-CT
single or double; 12-CT triple. Trumpet: Generally longer than in pipiens, index 5.6-7.3,
mean 6.4; pinna shorter than tracheoid area, usually about 0.25-0.30 trumpet length;
tracheoid area normally about 0.35-0.40 trumpet length. Abdomen: Length 2.7-3.2
mm, mean 3.0 mm; seta 6-111normally with 3 or 4 branches, rarely double, 6-IV-VI most
often with 4 branches (3-5); 5-IV very nearly same length and thickness as 5-V,VI, with 5
or 6 branches, 5-V with 3 or 4 branches, 5-W double or triple; 3-VII most often with 4
branches (3-5). Paddle: Length 0.82-0.94 mm, mean 0.88 mm; width 0.56-0.71 mm,
mean 0.63 mm; index 1.3-l .5, mean 1.4; outer margin with tiny serrations on proximal
0.5.
Larva (Fig. 16). Not associated with adults from southwestern Asia, specimens
examined include topotypic material; form and positions of setae as figured, see Table
27 for range and modal number of branches; very closely resembling antennatus, but
usually distinguished by the combination of shorter seta l-C, siphon with 2 lateral pairs
of seta l-S, seta 1-M usually single, and abdomen without obvious 2-banded
appearance in life. Head: Length 0.69-0.84 mm, mean 0.76 mm; width 1.02-1.24 mm,
mean 1.15 mm; lightly tanned, darker behind eyes. Dorsomentum usually with 7 teeth
(6-8) on either side of median tooth. Seta 1-C normally shorter than in antennatus,
tapered, not heavily tanned; 2-C occasionally present; 5,6,11-C as in antennatus but
5-C occasionally with 4 branches. Antenna: Length 0.50-0.64 mm, mean 0.57 mm;
lightly tanned, distal part slightly darker, dark on mesal surface at base. Seta 1-A with
about 27 branches (17-30). Thorax: Integument hyaline; with rows of vesicles on ventral
56
Contrib.Amer. Ent. Inst.,vol. 24, no. 1, 1988
surface. Seta 7-P almost always triple, seldom double; 8-P normally double, rarely triple.
Seta 1-M single or double, more often single, not much longer than 2-M. Seta 2-T most
often triple (2-4). Abdomen: Ventral surface with rows of vesicles; segments III and V
without conspicuous pigmentation. Chaetotaxy resembles that of antennatus in nearly
all respects; seta 6-VI usually single, occasionally double. Segment VIII: Comb with
32-55 scales, mean 43. Siphon: More lightly tanned and generally slightly shorter than
in antennatus; index 4.93-7.15, mean 5.72. Pecten with 8-15 spines, mode 11; larger
spines with 3 or 4 basal denticles. Seta 1-S in 5 or 6 pairs, la-S (when present) usually
borne within pecten, 2 pairs (Id, le) borne laterally; length distinctly less than diameter of
siphon, usually 0.5 or less diameter at point of attachment; branching variable, distal
elements usually with more branches than proximal elements (see table). Segment X:
Saddle length 0.37-0.42 mm, mean 0.40 mm; siphon/saddle index 3.56-4.30, mean
3.82. Seta 1-X most often with 3 or 4 branches, occasionally double; 2-X usually double,
sometimes triple.
Systematics.
Culex univittatus is a member of a very interesting species
aggregate. A thorough comparative study of this group has not been made yet but it
appears from a superficial examination of available material from Africa that three nominal
taxa and a possible fourth species are involved. The group includes univittatus, which
appears to be restricted to the temperate highlands of southern and eastern Africa and
the southwestern corner of the Arabian Peninsula; neavei Theobald, 1906, which
seems to occur throughout the subtropical and tropical lowlands south of the Sahara;
perexiguus, which inhabits the arid areas of northern and eastern Africa and
southwestern Asia; and another species which occurs in the evergreen rain forest of the
Congo Basin. Jupp (1972) identified specimens of univittatus from Benin, Niger, and
Upper Volta, but I do not believe that this species occurs in western Africa. These
records probably refer to neaveior an undescribed species.
The members of the univittatusaggregate are remarkably similar in all life stages.
The three nominal forms are characterized by slight and variable differences in adult
ornamentation and features of the male genitalia. These characters are contrasted for
univittatus, perexiguus, and neavei in Table 41. Some of these were noted previously
(Jupp, 1970, 1971, 1972) while others are noted here for the first time. No definite
differences have been found in the immature% but the larva of univittatus, based on the
examination of topotypic material, appears to be separable from perexiguus and neavei
by the development of seta 1-S. It is hoped that this information will provide a starting
point for a critical objective study of the group throughout its range. Few specimens are
currently available from most areas, and there are very few individual rearings from
anywhere other than Egypt. A comprehensive study of the group must include
extensive sampling throughout Africa and southwestern Asia.
The occurrence of univittatus in southwestern Asia is based on the examination
of a male and a female from the Yemen Arab Republic and the P.D.R. Yemen,
respectively. These are undoubtedly the specimens which Mattingly and Knight (1956)
recorded from the Yemen Arab Republic, the male erroneously. These specimens
agree well with the concept of univittatus presented here.
A very strikingfeature of univitiatusis the presence of a line of scales marking the
position of vein 2A. To my knowledge, this character is unique among mosquitoes and
appears to be a primitive feature in the family Culicidae. Culex univittatus appears to be
somewhat of a relict species.
Bionomics. Very little is known about the biology of univittatus, partly because
it has been confused with neaveiin the past. The species seems to be fairly abundant in
more temperate areas of the Afrotropical Region. Larvae are found in fresh water
Harbach: Subgenus Culex in SouthwesternAsia
57
ground pools, marshes, ditches, swamp margins, stream margins, and cattle hoofprints
under some degree of shade. Females feed primarily on birds (Anderson, 1967), but
will enter houses and bite humans (Jupp and McIntosh, 1967). Sindbis, West Nile, and
Wesselsbron viruses have been isolated from wild-caught specimens in South Africa
(Weinbren et al., 1956; McIntosh et al., 1967; Worth and De Meillon, 1960;
respectively). McIntosh et al. (1967) regard univittatus as the primary feral vector of both
Sindbis and West Nile infections in that country.
Distribution.
Culex univittatus is generally distributed in the temperate
highlands of the Afrotropical Region.
Material examined. 306 specimens. Only 3 specimens (1 F, 1 M, and 1 M
genitalia) were examined from southwestern Asia - P.D.R. YEMEN: (Mukaira); YEMEN
ARAB REPUBLIC: (Usaifira). A total of 303 specimens (110 females, 76 males, 41 male
genitalia, 6 pupal exuviae, 3 larval exuviae, and 67 fourth-instar larvae) were examined
from ANGOLA, ETHIOPIA, KENYA, MADAGASCAR (the type series of montforti ),
SOUTH AFRICA (the holotype female of ataeniatus ), TANZANIA, ZAIRE, and
ZIMBABWE (including the type series of univittatusand the holotype female of simplex).
A total of 253 specimens (111 females, 52 males, 44 male genitalia, 14 pupal
exuviae, 18 larval exuviae, and 14 fourth-instar larvae) of Culex neavei from the
following countries were examined and compared with univiftatus and perexiguus
BENIN, ETHIOPIA, GAMBIA, GHANA, KENYA, MALAWI,
during this study:
MOZAMBIQUE, NIGER, NIGERIA, SOUTH AFRICA, SUDAN (including the type series of
neavei), TANZANIA, UGANDA, UPPER VOLTA, ZAIRE, and uncertain locations.
Culex
(&flex)
perexiguus
Theobald
perexiguus Theobald, 1903a: 199. +Lectotype male:
[Lebanon]; designated by White, 1975: 318 (BM).
Sidon, Palestine
Culex decens Theobald of Storey, 1918(1919): 100 (Egypt, A, L keys, A); ? Abul-hab,
1968: 249 (Iraq, L key).
Culex (Culex) decens Theobald of ? Abdel-Malek, 1960: 113-l 18 (Syria, L bionomics,
distr.); lbrahim et al., 1983: 91 (Iraq, L key, L*).
Culex pallidocephalus Theobald of Gough, 1914: 135 (Egypt, toll. sites).
Culex perexiguus Theobald of Theobald, 1905a: 28 (Israel, Jordan); Edwards, 1922:
278 (Oriental Region, A key); Buxton, 1923: 316 (Israel, Jordan, toll. rec., A, L
bionomics); Theodor, 1923: 344 (Israel, Jordan, P’, key); Barraud, 1924b: 1263
(India, A key, M*, F, distr.); Barraud, 1924c: 431 (India, L*); Kirkpatrick,
1924(1925): 368, 372 (Egypt, A, L keys).
Culex (Culex) perexiguus Theobald of Edwards, 1921: 342 (Egypt, Israel, Jordan,
Oman, A, L keys, L’, tax., distr.); Seguy, 1924: 38, 190 (Lebanon, Egypt, A, L
keys, L*, tax., distr., syn.); Kirkpatrick, 1925: 125, 191 (Egypt, A, P, L keys, M’, F,
P’, L*, bionomics, distr.); Stackelberg, 1927: 163 (Middle East, A key, A, M gen.*,
distr.); Senevet and Andarelli, 1954: 63 (Egypt, A key, M, F); White, 1975: 318,
322 (Africa, SW Asia, A key); Harbach, 1985a: 86, 93, 104 (SW Asia, Egypt, distr.,
A, L keys).
Culex univittatus Theobald of Barraud, 1921: 394 (Israel, Jordan, Egypt, L habitat,
distr.); Parr, 1943: 247-250 (Syria, Lebanon, A, P, L keys, bionomics); Theodor,
1952: 113 (Middle East, zoogeogr.); Gad, 1956: 135 (Egypt, distr., bionomic
note); Senevet et al., 1957a: 86 (North Africa, L*); Nielsen and Nielsen, 1958:
Contrib.Amer. Ent. Inst.,vol. 24, no. 1, 1988
58
(Iraq, swarming beh.); Margalit and Tahori, 1970a: 142 (Israel, bionomics);
Margalit et al., 1971: 323 (Israel, bionomics); Margalit et al., 1973: 32 (Israel, toll.
rec.); El-Said and Kenawy, 1983a (Egypt, distr.); Kitron and Pener, 1986 (Israel, L
bionomics).
Culex univittatus form perexiguus Theobald of Mattingly, 1954: 56 (eastern
Mediterranean form, zoogeogr.).
Culex (Culex) univitfatusTheobald of Edwards, 1926: 138 (Egypt, Israel, Jordan, Oman,
A, L keys, syn., A, L*, distr.); Martini, 1931: 389 (Turkey, Oman, A, L keys, M*, F,
L*, distr.); Barraud, 1934: 418 (India, A, L keys, M*, F, L*, L habitat, distr.); Lewis,
1943b: 72 (in part; bionomic note); Senevet, 1947a: 126 (Turkey, Egypt, L’,
distr., L habitat, bionomics, L key); Abbott, 1948: 45 (Sudan, L); Senevet, 1949:
64 (North Africa, M gen.*, key); Monchadskii, 1951: 279 (Middle East, L’, key,
bionomics, distr., syn.); Hopkins, 1952: 291 (in part; Afrotropical Region, L*, key,
bionomics); Abdel-Malek, 1956: 101 (Sinai, L bionomics, L key); Lewis, 1956,
708 (at least in part: Sudan, toll. sites); Mattingly and Knight, 1956: 104, 122 (in
part; Oman, A, L keys); Senevet et al., 1957b: 92 (North Africa, L); Parrish, 1959:
266 (Turkey, distr.); Abdel-Malek, 1960: 113-125 (Syria, toll. sites, L bionomics);
DuBose and Cur-tin, 1965: 252, 254 (Mediterranean area, A, L keys); Abul-hab,
1968: 246 (Iraq, distr., L key); Lotfi, 1970: 402 (Iran, toll. rec.); Margalit and
Tahori, 1970b: 153 (Israel); Aslamkhan, 1971: 155 (Pakistan); Jupp, 1972:
105-l 13 (short-spined form, M gen.‘, F, distr.); Ward, 1972: 96 (Afghanistan);
Lotfi, 1973: 207 (Iran, toll. sites); Gutsevich et al., 1974: 385 (Middle East, A, L
keys, A, M gen.*, L*, L bionomics, med. imp.); Margalit and Tahori, 1974: 88
(Israel, toll. rec.); Lotfi, 1976: 73, 76, 82 (L key, ecol. note, L*); Sirivanakarn,
1976: 48 (Oriental Region, A, P, L keys, M*, F*, P, L*, tax., distr., bionomics);
El-Said and Kenawy, 1983b (Egypt, toll. rec.); Danilov, 1985a: 71 (Afghanistan, F
key); Danilov, 1985b: 55 (Afghanistan, L key).
Culex (Culex) univittatusform perexiguus Theobald of Lewis, 1956: 708 (Sudan, distr.);
Senevet and Andarelli, 1959: 212 (North Africa, Mediterranean basin, A, L keys,
syn., A distr., bionomics).
Culex (Cdex) univittatusvar. perexiguus Theobald of Mattingly and Knight, 1956: 104,
135 (Oman, distr., zoogeogr.); Ovazza et al., 1956: 171 (Ethiopia, M gen.*, toll.
sites); Mekuria et al., 1978: 80 (Ethiopia, distr.).
311
Adult. This common and widespread species closely resembles univittatus in
almost all respects, but the midfemur lacks a complete anterior pale stripe, vein 2A is
normally without scales, and the ventral arm of the male phallosome is short.
FEMALE. Head: Antenna dark, length 1.4-l .6 mm; pedicel with patch of pale
scales on dorsomesal surface; flagellomere 1 with few pale scales on mesal surface near
base. Proboscis dark-scaled, pale-scaled ventrally except at base, weakly pale on distal
0.25; length 1.7-1.8 mm. Maxillary palpus dark-scaled; palpomere 4 with pale scales
dorsomesally at apex, palpomeres 3 and 4 often with some lateral pale scales; length
0.3-0.4 mm, about 0.2 proboscis length. Falcate scales of vertex narrow, pale ochreous
to whitish, those of ocular line off-white and concolorous with interocular scales; forked
scales pale yellow medially, becoming brownish laterally; lateral spatulate scales white.
Thorax: Integument brown. Scutal scales narrow, somewhat dense, golden brown;
scales very pale yellow or whitish along extreme margins and around margin of
prescutellar area; pair of submedial, very faint to distinct pale spots on middle of scutum;
occasionally with trace of median anterior pale stripe. Falcate scales of antepronotum
pale yellow, paler ventrally; postpronotum with golden-brown falcate scales. Pleural
Harbach: Subgenus Culex in SouthwesternAsia
59
setae golden: 6-8 proepisternal, 12-l 6 prealar, 4-6 upper mesokatepisternal, 7-l 1
lower mesokatepisternal, 9 or 10 upper mesepimeral, and 1 (rarely 2) lower mesepimeral.
Pleura with white spatulate scales as follows: small patch below upper proepisternal
setae extending mesad onto anterior surface above level of base of coxa; postspiracular
area with patch on dorsal 0.5; posterior anepisternum with small patch below prealar
knob, usually more or less contiguous with upper katepisternal scales; katepisternum
with patches on upper corner and posterior border above coxa, extensive but well
separated; anterior and upper patches on anepimeron. Wing: Length 3.0-3.5 mm,
mean 3.2 mm; length of cell R2 2.2-2.8 length of vein R2+3, mean 2.4; length of cell
Ml 0.7-0.8 length of cell R2; dark-scaled with short line of dorsal pale scales on
posterior margin of costa at humeral crossvein; rarely with few pale scales at extreme
base of R; vein 2A occasionally with few scales distally, rarely with scales on entire
length. Halter: Pedicel and scabellum pale yellow; capitellum brown. Legs (hindleg,
Fig. 2K): Anterior surface of forecoxa with white spatulate scales at base and mixed pale
and dark scales distally, dark scales mostly in middle; mid- and hindcoxae with vertical
patch of white spatulate scales on anterior side of lateral midline. Each femur with narrow
pale knee spot; forefemur dark-scaled anteriorly, pale-scaled posteriorly, sometimes with
trace of anteroventral pale stripe; midfemur dark-scaled anteriorly, dark scales
progressively expanding over dorsal surface toward apex, with or without narrow
incomplete anteroventral longitudinal pale stripe, stripe faint, broken or distinct when
present; hindfemur pale-scaled with dorsal dark stripe usually beginning about 0.3 from
base and broadening apically. Anterior and dorsal surfaces of foretibia dark-scaled,
posterior and ventral surfaces pale-scaled; midtibia dark-scaled with anterodorsal and
posteroventral pale stripes; hindtibia dark-scaled dorsally, with broad anterior and ventral
pale stripes on proximal 0.8, stripes joined ventrally for at least part of their length distally,
separated on proximal 0.5 or less by narrow, often faint, distally diminished, dark stripe,
with distinct apical pale spot. Tarsi dark-scaled dorsally, pale-scaled ventrally; with very
faint indication of paler scales at apices of tarsomeres l-4. Abdomen: Tergum I with
median posterior scale patch usually comprised of pale and dark scales; terga II-VIII with
slightly convex basal white bands, bands 0.20-0.35 length of terga and continuous with
elongate basolateral patches of same color. Sterna usually completely pale-scaled,
occasionally with dark scales on posterolateral corners, less frequently also with median
patch of dark scales.
MALE. Differing from female as follows. Head: Proboscis essentially lacking
midventral pale scales; generally slightly longer, length 1.8-2.0 mm, mean 1.9 mm.
Maxillary palpus dark brown-scaled, length 2.4-2.7 mm, mean 2.6 mm; palpomere 2 with
some lateral pale scales; lateral surface of palpomere 3 with pale scales beginning above
base and few pale scales ventromesally at apex; palpomere 4 with narrow line of pale
scales tapering distally on ventral surface and dorsolateral line of pale scales ending just
short of apex; palpomere 5 with spot of white scales ventrally at base and somewhat faint
dorsolateral line of pale scales tapering distally to near apex. Falcate scales of vertex
very nearly white; forked scales paler than in female but still brownish laterally; lateral
spatulate scales broader dorsally than ventrally and posteriorly. Thorax: Pair of pale
spots extremely faint or absent on middle of scutum. Upper proepisternal scales more
extensive, usually well extended mesally on anterior surface. Wing: Length of cell R2
shorter, length of cell R2/length of vein R2+3 variable, 1.5-2.1, mean 1.8. Legs:
Midfemur usually without anterior pale stripe. Abdomen: Terga as in female except VI
and VII only with middle area dark-scaled, borders with pale scales. Sterna usually with
median steak and posterolateral spots of dark scales. Genitalia (Fig. 17): Almost exactly
as in univittatus, the only obvious distinctions being the following: seta g of subapical
60
Contrib.Amer. Ent. Inst.,vol. 24, 110.1, 1988
lobe of gonocoxite broader; ventral arm of lateral plate of phallosome shorter, not
projecting beyond caudal margin of lateral arm.
Pupa (Fig. 17). Character and placement of setae as figured, range and modal
number of branches in Table 8; no specific differences from univitiatus are apparent,
setae 6,12CT and 6-III-VI tend to have fewer branches. Cephalothorax: Seta 6-CT
usually triple (2-4); 12-CT most often double (2-4). Trumpet: As in univittatus, index
5.4-7.5, mean 6.5. Abdomen: Length 2.7-3.0 mm, mean 2.8 mm; setae l-Ill-VI and
5IV-VI tend to be rather shorter than in univitiatus; 6-111usually triple, seldom double or
with 4 branches, 6-IV-VI most often triple (3-5). Paddle: Length 0.84-0.94 mm, mean
0.89 mm; generally not as broad as in univittatus, width 0.53-0.61 mm, mean 0.57 mm;
index 1.5-l .7, mean 1.6.
Larva (Fig. 18). Form and positions of setae as figured, range and modal number
of branches in Table 28; characters almost the same as in univittatus,the only apparent
distinction being the development of seta 1-S; the siphon/saddle index is generally
larger. Head: Length 0.71-0.86 mm, mean 0.78 mm; width 1.12-l .38 mm, mean 1.28
mm. Dorsomentum most often with 7 teeth (6-9) on either side of median tooth.
Antenna: Length 0.54-0.64 mm, mean 0.60 mm. Seta 1-A with about 22 branches
(19-27). Thorax: Seta 7-P normally triple, seldom with 4 branches. Seta 2-T usually
double, sometimes triple. Abdomen: Setae 3-I-VII and 7-VI tend to have fewer
branches than in univiftatus. Segment VIII: Comb with 37-62 scales, mean 51. Siphon:
Generally slightly longer than in univittatus (see); index 5.45-7.87, mean 6.77. Pecten
with 9-15 spines, mode 13. Seta 1-S normally in 5 pairs, sometimes only 4.5 pairs
present, all normally borne beyond pecten; length about as long as diameter of siphon
at point of attachment; always with fewer branches than in univittatus, each element
usually double or triple (cf. tables). Segment X: Saddle length 0.35-0.46 mm, mean
0.41 mm; siphon/saddle index 4.20-4.87, mean 4.44. Seta 1-X double or triple in nearly
equal frequencies.
Systematics.
The recognition of perexiguus as a species distinct from
univittatus is somewhat provisional. The taxonomic status of these forms cannot be
settled unequivocally without a comprehensive study of individually reared material from
sympatric populations. An important analogy exists between these forms and pipiens.
The distribution of these forms coincides exactly with the distribution of pipiens in
southwestern Asia and Africa, yet pipiens is considered to be a single plastic species
while univittatus and perexiguus are regarded as separate species. The reason for this
is that univittatus and perexiguus exhibit slight anatomical differences (see Table 41)
whereas none are apparent in pipiens. For the present I am treating perexiguus as a
distinct species primarily because these differences, although variable, were observed
in a limited number of specimens from Ethiopia where the forms appear to be sympatric.
However, these forms seem to occur at different altitudes, and the observed differences
may be due in large part to environmental influences.
Bionomics.
Culex perexiguus is a common mosquito which is particularly
abundant during the summer and autumn months. The immature stages are found in a
variety of standing water habitats including swamps, springs, stream pools, ponds, and
wells. Breeding occurs in clean to moderately polluted water which usually contains
quantities of emergent vegetation. This species tolerates some degree of salinity and
occasionally utilizes artificial containers. It generally breeds away from dwellings, but has
the ability to adapt to breeding places modified by man. It is frequently found in
association with pipiens, antenna&s, and Aedes (Ochlerotafus) caspius (Pallas).
Although perexiguus probably feeds mainly on birds, it apparently occasionally
enters houses and bites man (Buxton, 1923; Kirkpatrick,1925). West Nile virus has
Harbach: Subgenus Culex in SouthwesternAsia
61
been isolated from this species in Israel (Nir et al., 1968; Samina et al., 1986) and Egypt
(Taylor et al., 1953; Taylor et al., 1956). Isolations of Sindbis virus have been made in
Israel (Samina et al., 1986) Egypt (Taylor et al., 1955), and Saudi Arabia (Wills et al.,
1985). Experimental transmission of Sindbis virus has been demonstrated for Egyptian
populations (Taylor et al., 1955).
Distribution.
Culex perexiguus is widely distributed in northern Africa and
southwestern Asia, and extends eastward into India.
Material examined.
2,696 specimens. A total of 2,546 specimens (689
females, 445 males, 65 male genitalia, 889 pupal exuviae, 429 larval exuviae, and 29
fourth-instar larvae) were examined from southwestern Asia - AFGHANISTAN:
(Garwargin); EGYPT: Aswan (Aswan, El A
‘ qaba el Saghira, Ezbet el Silsila, Khour Abu
Subeira, Kom Ombo, Nag el Higab, Nag el ldwa el Bahari, Nag el Kagug, Nag el Ritag el
Qibli, Nag el Shalabab, Nag’ Tingar, Sahara City), Asyut (Beni Shiqeir), Beni Suef(Biba),
Eastern Deserts (Bir el Maskhara, Wadi Digla, Farouk Field), El Buhayra (Wadi Natrun), N
Fay~um (Abhit el Hagar, Birket Qarum, Sanhur Bahariya, Sinero el Bahria), El Girgha (El
Birba), N Giza (Abu Ragwan el Bahari, Abu Rauwash, El Badrshein, Ezbet Bosna
Sharawy, Harania, Kafr el Ghataty, Koneissa, Nazlet el Ashtar, Saqqara, Talbia), N lsma’
i/iya (Abu Khalifa, Isma’iliya), N Qahira (Abbassia, Kirdasa, Ezbet Said), N Qalyubiya
(Ga’afara, Kaha, Khanka), ElSharqiya (Inshas el Raml), Port Said(Porl Said), Qena (Nag’
(Andimeshk, Behbehan, Bisotun, Ghosse, Kermanshah,
Hammadi); IRAN:
Khorramabad, Natchi Bahran, Shirin, Teheran); IRAQ: (Daurah); ISRAEL: (Akka,
Baniass, Beer Sheva, Beisan, Deir el Belah, Ein el Howl, Eliot, Galilee, Gaza, Genin,
Hadera, Inlerl, Jerisheh, Khirbet Hadrah, Kishon River, Lake Huleh, Latron, Lhallal,
Ludd, Nahala, Nahr es Zerga, Nahr Rubin, Shivta, Sidrat el Mallaha, Tel Aviv, Wassat,
Zichron); JORDAN: (Amman, Tafile, Taflah); LEBANON:
(the lectotype male and
paralectotype female); OMAN: (Itelba, Se’ Harr); PAKISTAN: (Peshawar); SAUDI
ARABIA: (Al Hasa Oasis, Al Khobar, Al Qatif, Riyadh). An additional 150 specimens (46
females, 29 males, 27 male genitalia, 6 pupal exuviae, 17 larval exuviae, and 25
fourth-instar larvae) were examined from ALGERIA, CHAD, DJIBOUTI, ETHIOPIA,
MOROCCO, SENEGAL, SUDAN, TUNISIA, UPPER VOLTA, and unknown localities.
Culex (Cu/ex)
theileri
Theobald
t heileri Theobald, 1903a: 187. +Lectotype male: Pretoria, Transvaal,
[South Africa] (BM), designated by Sirivanakarn, 1976: 45.
creticus Theobald, 1903a: 189. +Holotype female: Crete, Greece (BM).
Synonymy with theileri by Edwards, 1932a: 210.
pettigrewii Theobald, 191 Oa: 15. +Holotype female: Ukhrul, Manipur,
6400 ft., [Assam], India (BM). Synonymy with theilefi by Edwards,
1911: 262.
+Holotype female:
onderstepoortensis
Theobald, 1911: 265.
Onderstepoott, Transvaal, [South Africa] (BM). Synonymy with
theileri by Edwards, 1932a: 210.
annulata Theobald, 1913: 321 (theileri var.). Type specimens (female):
Onderstepoort, Transvaal, [South Africa] (? BM, see Harbach, 1983:
103, 104). Synonymy with theileri by Edwards, 1932a: 210.
alpha Skguy, 1924: 41. Lectotype larva, hereby designated, larva
illustrated by Seguy (1924: PI. VII, Fig. 34), specimen non-extant (?)
(see Harbach, 1983: 99). NEW SYNONYMY.
62
Contrib.Amer. Ent. Inst.,vol. 24, no. 1, 1988
Culex thei/eriTheobald of Gough, 1914: 135 (Egypt, toll. site); Storey, 1918(1919): 98
(Egypt, A, L keys, bionomics); Barraud, 1924b: 1266 (India, A key, syn., M*, F,
distr.); Barraud, 1924c: 429 (India, L*); Scott, 1927: 88 (Ethiopia, toll. rec.);
Salem, 1938: 27 (Sinai); Salem, 1940: 15 (Egypt, A, M gen.*); Parr, 1943:
246-251 (? Syria, Lebanon, A, P, L keys, bionomics); Senevet, 1947: 212 (North
Africa, L key); Theodor, 1952: 113 (Middle East, zoogeogr.); Dow, 1953: 688
(Iran, toll. rec., L bionomics); Mattingly, 1954: 58 (Africa, Asia, zoogeogr.); Gad,
1956: 135 (Egypt, distr., bionomic note); Senevet et al., 1957a: 85-86 (North
Africa, L*); Nielsen and Nielsen, 1958: 312 (Iraq, swarming beh.); Derwesh, 1965:
44 (Iraq); Margalit and Tahori, 1970a: 142 (Israel, bionomic note); Margalit et al.,
1971: 323 (Israel, bionomics); Gad and Salit, 1972: 581 (Egypt, toll. site); Margalit
and Tahori, 1973: 92, 93 (Sinai, distr., L assoc.); Margalit et al., 1973: 32 (Israel,
toll. rec., L assoc.); Buttiker, 1981: 476 (Saudi Arabia, toll. site); El-Said and
Kenawy, 1983a (Egypt, distr.); Zaini et al., 1983: 117 (Iraq, bionomic note); Kitron
and Pener, 1986 (Israel, L bionomics).
Culex (Culex) theileri Theobald of Edwards, 1926: 133 (Mediterranean Region, A, L
keys, A, L*, syn., distr.); Martini, 1931: 384 (Egypt, Israel, Jordan, Syria, Turkey,
A, L keys, M*, F, L*, distr.); Barraud, 1934: 414 (India, A, L keys, M’, F, L*, syn., L
habitat, distr.); Edwards, 1941: 305,419, 482 (Afrotropical Region, A key, M*, F’,
P, syn., distr.); Lewis,l943a: 281 (Eritrea, toll. rec., L bionomic note); Lewis,
1945: 15 (Sudan, A, distr.); Senevet, 1947a: 124 (Egypt, L*, distr., L bionomics,
L key); Senevet, 1949: 57, 64 (North Africa, M gen.*, key); Monchadskii, 1951:
275 (Middle East, L’, key, L assoc., distr.); Hopkins, 1952: 289 (Afrotropical
Region, L*, key, bionomics); Knight, 1953a: 232 (Yemen Arab Republic, toll.
rec.); Senevet and Andarelli, 1954: 60 (North Africa, A key, M, F); Khattat, 1955:
165 (Iraq, syn., L bionomics, distr., L’); Abdel-Malek, 1956: 101 (Sinai, L
bionomics, distr.); Lewis, 1956: 708-718 (Sudan, Egypt, Eritrea, L, toll. sites,
distr.); Mattingly and Knight, 1956: 106-129 (P.D.R. Yemen, Yemen Arab
Republic, L*, A, L keys, distr., zoogeogr.); Ovazza et al., 1956: 171 (Ethiopia, L*,
toll. sites); Parrish, 1959: 266 (Turkey, distr.); Senevet and Andarelli, 1959: 195
(North Africa, Mediterranean basin, A, P, L keys, syn., M*, F*, P*, L*, distr.,
bionomics); Abdel-Malek, 1960: 113-123 (Syria, toll. sites, L bionomics, distr.);
Khalaf, 1962: 54 (Iraq, A, L, L habitat); Derwesh, 1965: 44 (Iraq); DuBose and
Cur-tin, 1965: 352, 354 (Mediterranean area, A, L keys); Abul-hab, 1966: 281
(Iraq, distr., L bionomics); Abul-hab, 1968: 245 (Iraq, L habitat, distr., L key); Lotfi,
1970: 402 (Iran, toll. rec.); Margalit and Tahori, 1970b: 153 (Israel, toll. sites);
Aslamkhan, 1971: 155 (Pakistan); Ward, 1972: 96 (Afghanistan); Lotfi, 1973: 206
(Iran, toll. sites); Gutsevich et al., 1974: 388 (Asia, A, L keys, M*, F, L’, distr.,
bionomics); Margalit and Tahori, 1974: 87 (Israel, toll. rec.); Lotfi, 1976: 72, 74,
80 (Iran, L key, ecol. note, L’); Sirivanakarn, 1976: 43 (Oriental Region, A, P, L
keys, M’, F*, P, L*, tax., distr., bionomics); El-Said and Kenawy, 1983b (Egypt,
toll. rec.); lbrahim et al., 1983: 91 (Iraq, L*, key); Danilov, 1985a: 71 (Afghanistan,
F key); Danilov, 1985b: 55 (Afghanistan, L key); Harbach, 1985a: 86, 92, 106
(SW Asia, Egypt, dish., A, L keys).
Culex tipuliformis Theobald of Edwards, 1912b: 31 (Africa, A key); Barraud, 1920: 324
(Iraq, bionomic note); Barraud, 1921: 394 (Israel, Jordan, Egypt, distr., L assoc.);
Edwards, 1922: 278 (Oriental Region, A key); Buxton, 1923: 317 (Israel, Jordan,
COIL rec., bionomic note);Theodor, 1923: 344 (Israel, Jordan, P*); Kirkpatrick,
1924(1925): 367, 371 (Egypt, A, L keys).
Culex (Cutex) tipuliformisTheobald of Edwards, 1921: 339 (Mediterranean Region, A, L
Harbach: Subgenus Culex in SouthwesternAsia
63
keys, L*, tax., distr.); Seguy, 1924: 40, 192 (Egypt, Iran, Turkey, A, L keys, L*,
tax., distr., syn.); Kirkpatrick, 1925: 115, 188 (Egypt, A, P, L keys, syn., M’, F, P*,
L*, distr., bionomics); Stackelberg, 1927: 159 (Middle East, M, F keys, A, M
gen.*, distr.).
Adult. A distinct species with postspiracular and prealar scales and anterior white
stripes on all tibiae and the fore- and midfemora. Easily distinguished from all other
Culex species of southwestern Asia and Egypt, except perhaps univittatus, perexiguus,
and vagans. Culex univittatus and perexiguus are normally without an anterior white
stripe on the foretibia and vagans is without postspiracularand prealar scales.
FEMALE. Scaling of proboscis, maxillary palpus, legs, and abdominal terga
predominantly brownish black, legs blacker. Head: Antenna with some nearly white
scales on mesal surface of pedicel and flagellomere 1; length 1.6-2.0 mm. Proboscis
length 2.2-2.7 mm, mean 2.5 mm; dorsal surface largely black-scaled but often with
scattered pale scales, particularly in middle; ventral surface entirely with cream-colored
scales except at base. Maxillary palpus about 0.2 of proboscis length, with variable
amount of pale scaling; palpomere 4, and usually 3, with distinct dorsomesal patch of
white scales at apex, sometimes entire dorsomesal surface of both white-scaled;
scattered pale scales laterally, sometimes lateral surface entirely white-scaled. Falcate
scales of vertex whitish to light brown, usually largely pale yellow posteriorly, narrower
and darker anteriorly and laterally, narrowly white along margin of eye; forked scales pale
yellow medially, brown to blackish laterally; lateral spatulate scales white, narrow. Thorax
(Fig. 2B): Scutal integument dark brown; scales dense, rather fine to moderately coarse,
yellowish brown to golden brown, whitish or pale yellow on extreme anterior and lateral
margins and prescutellar area. Lobes of scutellum with scales same as prescutellar
scales. Falcate scales of antepronotum yellow, becoming pale yellow to white
posteriorly; falcate scales on anterior 0.67 of postpronotum fine, golden yellow, larger
and pale yellow to white on posterior 0.33. Pleural integument light to moderate brown,
faded posteriorly, frosted with dense covering of aculeae, with patches of white narrow
spatulate scales on upper area of proepisternum, postspiracular area near mesothoracic
spiracle, lower part of prealar area, upper and lower areas of mesokatepisternum, and
anterior and upper areas of mesepimeron, prealar patch continuous with upper
mesokatepisternal patch, latter usually continuous with lower mesokatepisternal patch,
anterior and upper mesepimeral patches often continuous; pleural setae golden: 1O-16
upper proepisternal, 7-14 prealar, 4-7 upper mesokatepisternal, 7-13 lower
mesokatepisternal, 5-16 upper mesepimeral, and 1 lower mesepimeral. Wing: Length
3.9-4.8 mm, mean 4.4 mm; length of cell R2 2.8-3.5 length of vein R2+3, mean 3.3;
length of cell M2 about 0.8 length of cell R2; scales almost entirely black or nearly black,
pale scales on posterior side of costa, distal 0.67 of subcosta, anterior side of distal 0.5
of R, and anterior side and approximately distal 0.5 of posterior side of Rl. Halter: Same
color as thoracic integument, with pale yellow to white scales on capitellum. Legs
(midleg, Fig. 2G): Anterior surface of forecoxa largely white-scaled, with black scales in
middle; midcoxa with small patch of white scales at base, these more or less distinctly
separated anteriorly from longitudinal patch of concolorous scales on anterior side of
lateral midline; hindcoxa with patch of white spatulate scales on anterior side of lateral
midline. All femora with narrow knee spots of white scales; posterior surface of fore- and
midfemora white-scaled, anterior surface dark-scaled with narrow longitudinal stripe of
white scales, white stripe of forefemur placed anteroventrally and sometimes more or
less broken up into spots, white stripe of midfemur borne medially with dark scales not
reaching base and anterodorsal line of dark scales progressively expanding onto dorsal
64
Contrib.Amer. Ent. Inst.,vol. 24, no. 1, 1988
surface toward apex; hindfemur white-scaled with dorsal and anteroventral stripes of
dark scales, dorsal stripe beginning beyond base and gradually widening distally,
anteroventral stripe on approximately distal 0.5. Anterodorsal surface of foretibia
dark-scaled with complete longitudinal stripe of white scales, posteroventral surface
white-scaled; anterior and posterior surfaces of mid- and hindfemora with complete
stripe of white scales, narrowly dark-scaled on dorsal and ventral surfaces. Tarsomere 1
of all tarsi more or less distinctly pale-scaled with anterodorsal longitudinal stripe of dark
scales; other tarsomeres more or less completely dark-scaled, scales appearing opalescent under light of certain angles. Abdomen: Tergum I with median posterior patch of
pale scales; tergum II with median longitudinal patch of creamy-white scales that is
broadest basally and may extend entire length of tergum; terga Ill-VII with more or less
triangular basal patches of creamy-white scales, lateral patches of white scales and
usually with narrow band of pale scales on posterior border, triangular patches broader,
produced less in middle and often continuous with lateral patches on posterior terga,
sometimes posterior angle of triangular patches extends to posterior margin or this area
with pair of submedial spots of pale scales; tergum VIII largely or entirely pale-scaled.
Sterna usually with scattered dark scales but sometimes with all whitish or yellowish
scales or a mixture of both; dark scales sometimes fairly numerous and arranged in more
or less distinct submedial longitudinal stripes and posterolateral spots that are often
joined posteriorly.
MALE. Differing from female as follows. Head: Maxillary palpus approximately
1.25 length of proboscis; integument dark brown; palpomeres 1 and 2 with scattered
pale scales ventrally; palpomere 3 with extensive patch of white scales on dorsolateral
margin of distal 0.67 and small mesal patch near apex, setose ventrally; dorsal and
ventral surfaces of palpomere 4 each with line of white scales; palpomere 5 white-scaled
dorsally, with small patches of white scales ventrally at base and apex; palpomeres 4 and
5 densely setose, each with lateral rows of dark setae. Dorsal falcate scales of vertex
more numerous; color more uniformly pale yellow. Thorax: Scutal scales finer; with
22-27 upper proepisternal setae. Wing: Length of cell R2 1.9-2.3 length of vein R2+3,
mean 2.1. Abdomen: Terga Ill-VII without lateral patches; basal triangular patches
generally broader; posterior terga usually with pale scales along lateral scale-free areas
and posterior border, these sometimes rather broad and more or less continuous with
submedial pale spots, thus occasionally significantly reducing dark areas. Genitalia (Fig.
19): Form as figured; structure of phallosome somewhat as in decens. Ninth tergal
lobes developed as usual, each with 7-14 (mode 9) irregularly spaced setae.
Gonocoxite normal, ventrolateral surface with usual complement of strong setae, mesal
setae longer and denser at base than usual, mesal surface also with 4 or 5 setae at apex,
lateral surface with cluster of moderately long setae at and below level of subapical lobe
(similar to laticinctus and matting/y/); subapical lobe slightly divided, setae a-c long and
tapered, a shorter and blunt at tip, b and c hooked apically; seta d absent, e simple, f
flattened and bladelike with one edge thickened; seta g foliform, rather short and
narrow, rounded apically; h simple, about same length as g, slightly curved distally.
Gonostylus developed as usual. Phallosome longer than broad with lateral plates and
aedeagal sclerites of nearly equal length; dorsal arm straight or bent laterad, tapered
distally, apex pointed; lateral arm with 2-5 (mode 3) laterally-directed denticles at tip, a
stubby dorsal process, and a rather weakly developed basal articulatory process; ventral
arm poorly developed, slender, sinuous, only slightly longer than one of the larger
denticles of lateral arm. Aedeagus rather conical; crest of aedeagal sclerite rather long,
about same length as proximal part. Proctiger not unusual; paraproct with moderately
long curved basal lateral arm and prominent ventral acetabulum, crown dark with numer-
Harbach: Subgenus Culex in SouthwesternAsia
65
ous spinelike spicules mesally and short blades laterally. Cereal sclerite elongate,
irregular in outline; cereal setae 2-6 (mode 3). Tergum X straplike, articulating with
paraproct below basal lateral arm.
Pupa (Fig. 19). Form and positions of setae as figured, range and modal number
of branches in Table 9; resembling univittatus, pefexiguus, antennatus, simpsoni, and
sinaiticus in the character of setae 6-111and 3-VII, but usually distinguishable in having
seta 1-II dentritic and the trumpet longer and flared. Cephalothorax: Lightly tanned; scuturn with pair of irregular submedial moderately tanned spots, these sometimes
subdivided; metanotum moderately tanned. Setae 1,2-CT usually with 4 branches (3-5);
6-CT with 3 or 4 branches; 8-CT usually with 5 branches (45); 9-CT double or triple;
lo-CT generally with fewer branches than other species of the pipiens group (2-4);
1 l-CT double; 12-CT usually with 4 or 5 branches (3-5). Trumpet: Moderately tanned,
pinna darker than usual; rather long and flared distally, index 4.1-6.2, mean 4.4; pinna
oblique, rather long, 0.2-0.3 total length of trumpet. Abdomen: Lightly tanned, often
with dark V-shaped stripe beginning at base of tergum I and gradually tapering to
posterior margin of tergum IV, width of stripe on tergum I equals width of metanotum;
length 3.0-3.7 mm, mean 3.5 mm. Seta 1-II dendritic, usually with fewer than 12
branches (7-16); l-Ill-VII multiple, number of branches progressively fewer on each
succeeding posterior segment, 1-III often with 10 or 11 branches (6-13), l-VII frequently
with 4 branches (3-6); 5-IV usually with 4 branches (2-4), 5-V,VI double, both distinctly
longer than 5-111,about 1.5 length of following tergum; 6-III-VI variable, often with 3-5
branches (2-6). Paddle: Developed and tanned as usual; length 0.79-l .08 mm, mean
0.94 mm; width 0.72-0.82 mm, mean 0.74 mm; index 1.2-l .4, mean 1.3.
Larva (Fig. 20). Character and placement of setae as figured, range and modal
number of branches in Table 29; resembles simpsoni and sinaiticus but differs from
other members of the pipiens group in having comb scales with a distinct sharp apical
spine, differs conspicuously from simpsoni and sinaiticus in the character of setae 5-C
and 1-S. Head: Length 0.78-l .05 mm, mean 0.88 mm; width 1.28-l .54 mm, mean 1.37
mm; mainly lightly tanned, lateralia with rather distinctly delineated dark area behind each
eye, dorsal apotome with O-9 dark spots of variable size and distinctness behind seta
5-C (in 3 rows of 3 when all are present, largest at bases of seta 5-C and medially
between bases of seta 5-C and posterior margin of head, 3-5 spots usually present).
Dorsomentum most often with 6 teeth (5-8) on either side of median tooth. Seta 1-C
stout, moderately tanned, sometimes barbed; 2-C absent; 4-C often double or triple,
sometimes single; 5-C usually triple, occasionally with 4 branches; 6-C almost always
double, seldom triple; 7-C often with 7 branches (6-10); 10-C with 3 or 4 branches
occurring in nearly equal frequencies; 11-C normally double, seldom single; 12-C most
often with 4 branches (3-6); 14-C usually double, infrequently single. Antenna: Length
0.60-0.80 mm, mean 0.70 mm; proximal part lightly tanned, distal part moderately
tanned, with dark spot mesally at base; proximal part aciculate, aciculae coarser just
before seta 1-A. Seta 1-A with about 22 branches (18-32). Thorax: Integument hyaline,
surface smooth, tubercles of pleural setal groups moderately tanned; with distinctive
somewhat hourglass-shaped internal body of black pigment extending from anterior
margin to middle of metathorax, easily seen in larva resting at water’s surface. Seta 7-P
usually triple, infrequently with 4 branches; 8-P most often double, frequently triple.
Seta 12-T usually double, sometimes single. Abdomen: Integument hyaline, lightly
tanned, sometimes slightly darker dorsally; tubercles of larger setae lightly to moderately
tanned. Seta 3-l usually double or triple (2-5), 3-VII most often with 5 branches (3-8);
6-I-VI almost always triple, 6-I-111distinctly longer than 6-IV-VI; 1-III,IV usually with 4
branches, l-111occasionally with 5 branches, l-IV infrequently with 3 or 5 branches,
66
Contrib.Amer. Ent. Inst.,vol. 24, no. 1, 1988
l-V,VI usually triple, 1-V often with 4 branches, l-VI sometimes double and infrequently
with 4 branches. Segment VIII: Comb with 28-45 fringed scales (mean 35) with sharp
apical spine. Seta l-VIII most often with 7 branches (5-9); 3-VIII usually with 8 branches
(7-10). Siphon: Index 4.15-5.36, mean 4.79; straight or very slightly sigmoid in lateral
view; moderately tanned, sometimes slightly darker in middle, proximal 0.5 often very
dark. Pecten of 8-13 long slender spines (mode 12) on basal 0.3 of siphon; spines with
3-5 short basal denticles. Seta 1-S usually comprised of 10 setae (7-l 3), two forming lateral pair, others imperfectly paired close to posterior midline. Segment X: Saddle
complete; moderately tanned, sometimes dark along posterior margin; length 0.44-0.56
mm, mean 0.48 mm, siphon/saddle index 3.15-3.46, mean 3.33. Seta 1-X usually triple,
occasionally double, sometimes with 4 branches; 2-X most often with 4 branches, often
triple, sometimes with 5 branches; 4-X usually in 7 pairs, sometimes less 1 or 2 setae.
Systematics. Sirivanakarn (1976) placed theileri in a subgroup of its own and
suggested that Afrotropical species such as andersoni Edwards, 1914a, and simpsoni
might also be members of the group. It is still not known whether andersoni belongs to
this group, but simpsoni and its close allies are sufficiently distinct from theileri to be
placed in a separate subgroup. There are a number of intermediate Afrotropical forms
which are not definitely assignable to either subgroup, so these subgroups may have to
be redefined once the species are better known. The affinities of laticinctus and
mattinglyi are not clear, but they resemble theileri in a number of characters and I am
placing them provisionally in the theileri subgroup.
Culex theileri is a clearly marked species which is widely distributed in the
Afrotropical, southern Palaearctic, and western Oriental regions. The larvae and adults,
excluding the male genitalia, are extremely variable and exhibit the same degree of
variation within local populations that is observed throughout the entire range.
Therefore, it appears that only a single widespread and variable species is involved, yet it
is possible that the taxon actually consists of an aggregate of biologically distinct,
isomorphic species. This form needs to be thoroughly studied throughout its range.
I am certain that Culex alpha Seguy, 1924 is conspecific with theileri. There are
two possible type larval specimens labelled as alpha in the Museum National d’Histoire
Naturelle, Paris (Harbach, 1983) which are easily identified as theileri. Since there is no
clear evidence to indicate that these larvae are type specimens, I have designated the
larva illustrated by Seguy as the lectotype of alpha. This figure agrees in all essential
details with the current concept of theileri.
Bionomics. Culex theileri is an adaptable species which utilizes a wide range of
breeding sites, but it is also a focal species, reaching high densities in some areas while
appearing to be rare or absent in others. The immature stages can breed in permanent
or temporary bodies of fresh, brackish, or foul water. They are commonly found in
stagnant or slow streams and irrigation ditches, swamps, marshy pools, ground pools,
ponds, springs, disused wells, open cisterns, and seepage water. The breeding sites
are usually sunlit and often contain little or no floating or emergent vegetation. Larvae
are often found alone or in association with pipiens, perexiguus, or antennatus.
In southwestern Asia, this species reaches peak densities in late spring and early
summer. Females feed mainly on mammals and are known to enter houses and bite man
(Kirkpatrick, 1925; Gad, 1956; Anderson, 1967). This species does not seem to be
involved in the transmission of any human diseases in southwestern Asia, but its
importance as a vector has not been thoroughly studied. McIntosh et al. (1967) found it
naturally infected with West Nile and Sindbis viruses in South Africa, but easy
experimental transmission has been demonstrated only for Sindbis virus (Jupp et al.,
1966).
Harbach: SubgenusC&x in SouthwesternAsia
67
Distribution.
Culex theileri seems to have a discontinuous distribution in
southern and eastern Africa, lands around the Mediterranean, southwestern Asia,
southern USSR, southern China, India, and Burma.
Material examined.
1,452 specimens. A total of 1,337 specimens (311
females, 221 males, 62 male genitalia, 282 pupal exuviae, 161 larval exuviae, and 300
fourth-instar larvae) were examined from southwestern Asia - AFGHANISTAN: (Angur
Abajhr, Bolla Quchi, Garwargin, Karukh, Sayedabat); EGYPT: Aswan (Aswan, El A
‘ qaba
el Saghira, Nag’ Tingar, Sahara City), N Burhayra (Wadi Natrun - Bir Hooker, Ezbet Kafr
Dawud), N I%yyum (Abhit el Haggar, Birket Qarun), El Giza (Saqqara), El lsma’ iliya
(Isma’iliya), Mafsa Matfuh (Siwa Oasis - Abul el Leef, Aghurmi, Ain Bundi, Masouse,
Mshendit), Sinai Peninsula (unknown localities); IRAN: (Bisotun, Cham Asbi, Dorud,
Ghasem Gheshlari, Kivi, Maragheh, Mazandaran, Mehrdasht, Mish Kazeroon, Natchi
Bahran, Semnan, Sharaf Khaneh); IRAQ: (Abu Ghraib, Amara, Baghdad, Barari Bela
Musal, Basrah, Daurah, Erbil Town, Hadji Geul, Rawanduz Nahia, Shubaichas, unknown
localities); ISRAEL: (Acre, Deir el Belah, Dhahiriya, Ein Ghridyan, Khedeira, Kishon
River, Latron, Lake Huleh, Ludd, Marakeb, Ramat Razi’el, Shivta, Wadi Sharia);
JORDAN: (AZ Zarqa); PAKISTAN (localities east of the lndus River): (Gujranwalia,
Jhelum, Lahore); P.D.R. YEMEN: (Jebel Jihaf); SAUDI ARABIA: (Gatib Oasis, Sabo);
SYRIA: (Damascus, Deir el Aachair, Doumar, Kissoue); TURKEY: (Adana, Istanbul,
Konya, Koyna-Eregli, Sinop-Boyobat, Tatvan); YEMEN ARAB REPUBLIC: (El Amra,
Hada, San’s, Ta’izz).
An additional 115 specimens (61 females, 38 males, 5 male
genitalia, 1 pupal exuviae, 6 larval exuviae, and 4 fourth-instar larvae) were examined
from ALGERIA, GREECE (CRETE) (including the holotype female of creticus ), KENYA,
INDIA (the type series of pettigrewii), LIBYA, NEPAL, SOUTH AFRICA (including the
types of theileri and onderstepoortensis), TUNESIA, ZIMBABWE, and unknown
localities.
Culex (Culex)
laticlnctus
Edwards
laticinctus Edwards, 1913a: 49. +Holotype male: Tiberias, Palestine,
[Israel] (BM).
Culex sp. no. 2518 of Storey, 1918(1919): 101 (Egypt, A, L keys, A, M gen.‘, L*).
Culex laticincfus Edwards of Barraud, 1921: 394 (Israel, Jordan, Syria, Egypt, L habitat,
distr.); Buxton, 1923: 317 (Israel, Jordan, toll. rec., L bionomics); Theodor, 1923:
341 (Israel, Jordan, P’, key); Kirkpatrick, 1924(1925): 368, 371 (Egypt, A, L
keys); Salem, 1938: 27 (Sinai); Parr, 1943: 246-250 (Lebanon, toll. sites,
bionomics, A, P, L keys); Lewis, 1945: 17 (Sudan, distr.); Senevet, 1946:
317-321 (? North Africa, L’); Leeson and Theodor, 1948: 228 (Socotra, L habitat,
toll. sites, A, L keys); Theodor, 1952: 113 (Middle East, zoogeogr.); Senevet et
al., 1957a: 86 (North Africa, L’); Abul-hab, 1968: 249 (Iraq, L key); Margalit and
Tahori, 1973: 90-93 (Sinai, distr., toll. sites, L bionomics); Margalit et al., 1973: 32
(Israel, COILrec., L assoc.); Dimentmand and Margalit, 1981: 124-l 27 (Israel, L
bionomics); Kitron and Pener, 1986 (Israel, L bionomics).
Culex (Culex) laticinctus Edwards of Edwards, 1921: 342 (Egypt, Israel, Jordan, Oman,
A, L keys, tax., distr.); Seguy, 1924: 37, 190 (Egypt, Israel, Jordan, Oman, A, L
keys, M gen.‘, tax., distr., syn., not L in Figs. 20-23, PI. II); Kirkpatrick, 1925: 119,
190 (Egypt, A, P, L keys, M*, F, P*, L’, bionomics, distr.); Edwards, 1926: 137
(Egypt, Israel, Jordan, Oman, A, L keys, A, L, distr.); Stackelberg, 1927: 160
68
Contrib.Amer. Ent. Inst.,vol. 24, no. 1, 1988
(Middle East, M, F keys, A, M gen.*, distr.); Martini, 1931: 368 (Arabia ?, Egypt,
Israel, Jordan, Syria, A, L keys, M*, F, L*, distr.); Edwards, 1941: 313, 420, 482
(Afrotropical Region, excluding records from Dhahran and San’a, A key, M’, F*, P,
distr.); Lewis, 1943a: 282 (Eritrea, toll. rec., bionomic note); Senevet, 1947a:
123 (Egypt, Saudi Arabia, Syria, Turkey, L*, distr.); Abbott, 1948: 45 (Sudan, L, L
habitat, distr.); Senevet, 1949: 56 (North Africa, M gen.*, key); Monchadskii,
1951: 263 (Middle East, L*, key, L assoc., distr.); Hopkins, 1952: 298 (Afrotropical Region, L*, key, bionomics); Knight, 1953a: 229 (Yemen Arab Republic, toll.
rec.); Senevet and Andarelli, 1954: 58 (Israel, Jordan, A key, M, F); Abdel-Malek,
1956: 100 (Sinai, L bionomics, L key); Lewis, 1956: 710 (Sudan, Egypt, Eritrea,
toll. sites, L habitat, distr.); Mattingly and Knight, 1956: 102-132 (Oman, P.D.R.
Yemen, Socotra, Yemen Arab Republic, L’, A, L keys, distr., bionomics); Senevet
et al., 1957b: 92 (North Africa, L); Parrish,1959: 266 (Turkey, distr.); Senevet and
Andarelli, 1959: 158 (North Africa, Mediterranean basin, A, P, L keys, M*, F*, P*,
L’, distr., bionomics); Abdel-Malek, 1960: 113-l 20 (Syria, toll. sites, L bionomics,
distr.); DuBose and Cut-tin, 1965, 352, 354 (Mediterranean area, A, L keys);
Mekuria, 1968: 78 (Ethiopia, distr.); Lotfi, 1970: 401 (Iran, toll. rec.); Margalit and
Tahori, 1970b: 152 (Israel, toll. sites); Lotfi, 1973: 206 (Iran, toll. site); Gutsevich
et al., 1974: 345 (Middle East, L key); Lotfi, 1976: 73, 76, 83 (Iran, L key, ecol.
note, L*); Margalit and Tahori, 1974: 87 (Israel, toll. rec., L assoc.); Harbach,
1985a: 86, 95, 103 (SW Asia, Egypt, distr., A, L keys).
Culex (Culex) mattinglyi Knight of ? Abdel-Malek, 1960: 120 (Syria, toll. sites, L
bionomics, distr.); ? Abul-hab, 1968: 248 (Syria [based on report by Abdel-Malek,
1960: 1201, L key).
Adult. A medium-sized mosquito easily recognized by the light-colored thorax,
dark proboscis, absence of postspiracular and prealar scales, and the relatively broad
basal pale bands of the abdominal terga. Individuals usually possess 2 or 3 lower
mesepimeral setae.
FEMALE. Head: Length of antenna about 2.0 mm; flagellum normal; pedicel and
flagellomere 1 with some small pale scales on mesal surface. Proboscis blackish, slightly
paler on proximal 0.6 of ventral surface; length 2.2-3.1 mm, mean 2.7 mm. Maxillary
palpus usually entirely black-scaled, palpomere 3 sometimes with few apical pale scales;
length 0.4-0.6 mm, about 0.2 proboscis length. Forked scales of vertex long and
slender, light yellowish brown; falcate scales white to pale yellowish, distinctly white
along margin of eye; lateral spatulate scales white. Thorax: Pleural integument yellowish
with tinge of brown, scutum not much darker. Scutal scales moderately coarse, light
yellowish brown, white scales on anterior promontory, lateral margins, along prescutal
suture and on prescutellar area; scutal setae prominent, brown, well contrasted with
light-colored scales. Scutellum with narrow whitish falcate scales on all 3 lobes. Anteand postpronota with long whitish to yellowish falcate scales; with 4-8 setae on
posterodorsal margin. Pleura with yellowish to golden-brown setae and patches of
narrow white spatulate scales: upper part of proepisternum with small patch of scales
below 5-l 0 setae, postspiracular and prealar areas without scales, prealar area with 7-14
setae, upper part of mesokatepisternum with scale-patch and 5-7 setae, lower posterior
border of mesokatepisternum with patch of semierect scales and row of 5-13 setae,
mesepimeron with anterior and upper scale-patches and 5-14 setae on upper area;
number of lower mesepimeral setae variable, commonly 2 or 3, often 1 (at least on one
side) and occasionally 4. Wing: Dark-scaled with short line of whitish scales on posterior
margin of costa near humeral crossvein; length 3.7-5.1 mm, mean 4.4 mm; length of cell
Harbach: SubgenusC&x in SouthwesternAsia
69
R2 2.6-4.3 length of vein R2+3, subcosta intersects costa beyond furcation of vein R2+3;
length of cell Ml 0.65-0.78 length of cell R2. Halter:
Pedicel and scabellum pale;
capitellum dark. Legs: Anterior surface of forecoxa with whitish scales, mid- and
hindcoxae with longitudinal patch of narrow whitish scales on anterior side of lateral
midline. All femora with inconspicuous whitish knee spots; anterior surface of fore- and
midfemora with blackish scales, posterior surface white-scaled; hindfemur largely whitescaled, with dorsal stripe of black scales beginning 0.1-0.3 from base and gradually
widening to encircle approximately distal 0.2. All tibiae black-scaled anteriorly,
whitish-scaled posteriorly, apices of fore- and midtibiae inconspicuously pale-scaled,
apex of hindtibia with conspicuous pale spot. Tarsi black-scaled with hint of paler scales
on posterior surface of tarsomere 1. Abdomen: Tergum I with median posterior patch of
white to light brown scales; terga II-VII black-scaled with basal 0.4-0.6 white-scaled;
tergum VIII with lateral patches of cream-colored scales, broad median area without
scales. Scaling of sterna I-VII entirely cream-colored; sternum VIII usually entirely whitescaled.
MALE. Differing from female as follows. Head: Antenna1 length 1.5-2.1 mm,
mean 1.8 mm. Length of proboscis 2.3-2.9 mm, mean 2.6 mm; without ventral cluster of
setae. Maxillary palpus barely if at all longer than proboscis, length 2.5-2.9 mm, mean
2.7 mm; scaling mainly dark, palpomere 3 with subtle subapical stripe of pale scales on
ventrolateral margin, palpomere 4 with ventral stripe of white scales, and palpomere 5
with ventral spot of white scales at base. Wing: Length 3.3-4.2 mm, mean 3.7 mm;
length of cell R2 1.3-2.3 length of vein R2+3, subcosta intersects costa before furcation
of vein R2+3; length of cell Ml/length of cell R2 0.6-0.8. Abdomen: Basal bands
0.45-0.75 tergum length; posterior edge of bands irregular, usually with submedial
indentations of posterior dark scaling; bands produced posteriorly along lateral setae,
particularly on terga VI and VII. Genitalia (Fig. 21): Form as figured; phallosome much as
in theileri. Ninth tergal lobe small, with 4-9 (mode 5) small setae. Gonocoxite stout,
ventral and mesal surfaces with usual complement of large and small setae respectively,
mesal surface also with conspicuous clump of moderately long setae near base of
gonostylus, lateral surface with rather dense covering of moderately long setae and
some long slender setae at base; subapical lobe distinctly divided but not prominent,
setae a-c stout, slightly flattened and bent distally, a about half as long as c, b
intermediate; seta d absent, e and f resemble a-c but are more slender and distinctly
flatter and broader distally, e about as long as a, f about as long as c; seta g a
crescent-shaped blade with pointed tip; h slender, curved, and inconspicuous.
Gonostylus relatively short, rather more sharply bent in middle than usual, with crest of
small sharp ridges before tip. Phallosome constructed as in theileri but lateral plate
longer than aedeagal sclerite, crest of aedeagal sclerite shorter, lateral arm of lateral plate
with 3-7 (mode 4) apical denticles, and ventral arm more strongly developed, much
longer than denticles of lateral arm. Proctiger almost as in theileri; ventral acetabulum of
paraproct inconspicuous; cereal setae more numerous, 5-8 (mode 7).
Pupa (Fig. 21). Form and placement of setae as figured, range and modal
number of branches in Table 10; distinguished from all the other species except
mattinglyi by the strongly flared trumpet with its short tracheoid area and unusually short
seta 6-l,ll. Cephalothorax: Lightly to moderately tanned, scutum, legs, and metathorax
darker. Setae 1,3,7,9-CT double or triple, more often double; 8-CT usually with 4 or 5
branches, sometimes with 6; lo- and 11-CT close set, 1O-CT with variable number of
branches (4-l 0), 11-CT usually double, occasionally single; 12-CT with 3 or 4 branches.
Trumpet: Moderately tanned, tracheoid area darker; rather short and distinctly flared,
index 2.8-3.7, mean 3.2; pinna 0.3-0.4 trumpet length; tracheoid area less than 0.3
70
Contrib.Amer. Ent. Inst.,vol. 24, no. 1, 1988
trumpet length. Abdomen: Lightly tanned, anterior terga and sterna darker in middle,
anterior and posterior margins of these darker still; length 3.5-4.0 mm, mean 3.8 mm.
Seta 6-I,11 short, only slightly longer than 7-l,ll, single or double; 7-l usually triple,
sometimes double; 7-11usually double, sometimes single; l-11 with short stem and only
7-14 branches; 1O-II occasionally present, often only on one side, usually single, sometimes double or triple; alveolus of 1 l-11 present; 6-III-VI usually triple (2-4); 5-IV shorter
than 5-V,VI, about length of following tergum, most often with 4 branches (2-5); 5-V,VI
normally double, distinctly longer than following tergum. Genital lobe: Moderately
tanned; larger than usual, length about 0.25 mm in female, about 0.50 mm in male;
female with short ventromesal pointed process projecting cephalad from base; male with
pair of bumps at base ventrally. Paddle: Very lightly tanned, buttress and midrib darker;
inner part widest distally; length 1.00-l .I5 mm, mean 1.09 mm, width 0.83-l .OO mm,
mean 0.91 mm, index 1.1-l .3, mean 1.2.
Larva (Fig. 22). Character and positions of setae as figured, range and modal
number of branches in Table 30; easily distinguished from all other Culex in
southwestern Asia by the combination of seta 6-III-VI normally single and 2-X with 4 or 5
branches, resembling decens and duftoni in the character of seta 6-III-VI, and many
specimens of theileri, Miens, poicilipes, and bitaeniorhynchus in the branching of seta
2-X. Head: Length 0.77-l .04 mm, mean 0.91 mm; width 1.10-l .54 mm, mean 1.33 mm;
mainly lightly tanned, darker ventrally (mainly gula) and posteriorly (particularly lateralia);
ventral ecdysial line distinct from level of seta 15-C to posterior margin of gula.
Dorsomentum usually with 7 or 8 teeth (7-9) on either side of noticeably larger median
tooth. Seta 1-C long and rather stout; 2-C present or absent; 5,6-C shorter than usual,
reaching only slightly beyond anterior margin of median labral plate; 5-C usually with 4 or
5 branches, seldom with 6; 6-C usually with 4 branches, sometimes triple; 8-C frequently
with 4 or 5 branches (3-6); 10-C usually triple (2-5); 14-C double or triple, more often
triple. Antenna: Length 0.53-0.68 mm, mean 0.62 mm; lightly tanned before seta l-A,
distal part darker, with usual dark mesal spot at base; aciculate as usual, aciculae slightly
shorter and broader near seta 1-A. Seta 1-A with about 23 branches (19-26); 2,3-A
distinctly subapical. Thorax: Integument hyaline; surface smooth. Seta 7-P with 3 or 4
branches, more often with 3; 8-P double. Setae 1,2-M about same length, both
frequently triple (l-4 and l-5 respectively). Seta 12-T double or triple, more often
double; 13-T smaller than usual, with 7-l 1 branches. Abdomen: Integument hyaline;
surface smooth. Seta 3-l usually with 3 or 4 branches (2-7) 3-VII frequently double or
triple, sometimes with 4 branches; 6-I,11 usually triple, infrequently with 4 branches,
6-lll,VI normally single, 6-lll,lV occasionally double; l-111most often single, sometimes
double, 1-IV,V single, long, about 1.5 length of segment, 1-VI almost always single and
long, seldom double, l-VII with fewer branches than usual, double or triple, more often
double. Segment VIII: Comb with 24-38 (mean 32) evenly fringed scales, fringe longer
at apex. Siphon: Index 3.07-4.18, mean 3.48; subcylindrical, broadest at base and
tapering distally; moderately tanned, slightly darker at base and apex. Pecten on
approximately basal 0.4, composed of 1O-l 6 (mode 12) long curved spines with 3 or 4
basal denticles. Seta 1-S usually represented by 14 setae but 12-15 may be present, 2
paired laterally near tip, others more or less paired along posterior midline, 2 proximal
pairs within pecten, lateral and most distal pair smaller and with fewer branches than the
others. Segment X: Saddle complete; moderately tanned, darker on anterior and
posterior margins; length 0.36-0.48 mm, mean 0.42 mm, siphon/saddle index
2.78-3.24, mean 3.02. Seta 1-X single or double, more often double; 2-X with 4 or 5
branches (Kirkpatrick, 1925 reported 5 or 6); 4-X almost always in 7 pairs, infrequently
with 6 paired and 1 unpaired seta.
Harbach: SubgenusCulex in SouthwesternAsia
71
Systematics. The adults of laticinctus bear little resemblance to those of theileri
except for a remarkable similarity in the structure of the male phallosome. The larva of
this species is generally similar to that of theileri, but the comb scales and setae 1,6-III-VI
are very different. In spite of these differences I am including both laticinctus and
maffinglyj in the theileri subgroup because maRing/yj, which seems to be very closely
related to laticinctus, exhibits some characters in the adult and larval stages which are
intermediate in development between those in laticinctus and theileri.
Culex laticinctus and theileri are sympatric throughout the arid lands of northern
Africa and southwestern Asia where they were once equally common species. Today,
laticinctusseems to be less common than theileri. The reason for this is unknown, but it
may be due to human influence. In 1983 I had the opportunity to collect mosquitoes all
over the Nile Delta area of Egypt with Bruce A. Harrison and we never found laticinctus ,
not even in the vicinity of Alexandria where it was once quite common (Kirkpatrick,
1925). We examined a number of specimens in the Dokki Institute and the Entomological Society of Egypt in Cairo which came from near Alexandria during Kirkpatrick’s
time, but no other specimens of laticinctuswere present in either repository.
The presence of two or three lower mesepimeral setae has been used
extensively to distinguish this species from other species of Culex known to occur in the
Afrotropical and Mediterranean regions. This character, however, is not as reliable as it
was previously thought to be. Many specimens examined during this study, especially
from Israel, have only one lower mesepimeral seta on both sides of the thorax. This
condition is reflected in the key presented above.
Bionomics. Culex laticinctus seems to have been more common in the past
than it is today. It was once collected frequently in artificial containers such as cisterns,
tanks, barrels, wells, concrete basins, and similar structures. Now it seems to be found
more often in stream pools, rock pools, swamps, springs, irrigation ditches, and
temporary ground water. It also uses seepages (Kitron and Pener, 1986) and has been
found in animal hoofprints supplied with water by a spring (Abdel-Malek, 1960). This
species usually occurs in fresh water, but is found occasionally in slightly brackish water.
It is frequently collected in association with Anopheles (Anopheles) claviger (Meigen)
and Culiseta (Allotheobaldia) longiareolata (Macquart). It has also been found in
association with Anopheles (Cellia) hispaniola (Theobald), ? An. (Gel.) multicolor
Cambouliu, An. (Gel.) sergentii (Theobald), Aedes (Ochlerotatus) caspius (Pallas),
Culiseta (Culiseta) annulata (Schrank), Uranotaenia (Pseudoficalbia) unguiculata
Edwards, Culex (Barraudius) pusillus Macquart, Cx. (Maillotia) hortensis Ficalbi, and
several members of the subgenus Culex, including mimeticus, perexiguus, pipiens,
sinaiticus, theileri, and tritaeniorhynchus. Culex laticinctus is mainly a summer species
but specimens can be found during most of the year. Resting adults have been
captured in tents, a tub, and an ice factory, but it is not known whether females enter
houses or bite man.
Distribution.
This species is apparently widespread from the Canary Islands
eastward through lands around the Mediterranean, Equatorial Africa, Sudan, Ethiopia,
Somalia, the Arabian Peninsula, and the Middle East.
Material examined. 481 specimens. A total of 400 specimens (103 females,
109 males, 21 male genitalia, 67 pupal exuviae, 53 larval exuviae, and 47 fourth-instar
larvae) were examined from southwestern Asia - EGYPT: Sinai Peninsula (Mt. Abas),
unknown (Ibrahimien, Moharem Bey); IRAN: (Nanrizac Borazjun); ISRAEL (including the
holotype male): (Ein Turieba, Haifa, Jericho, Jerusalem, Mont Fort, Mt. Garnish, Rahm
Alla, Lake Tiberias, Wadi Faria, Yeroham); JORDAN: (Aqaba, Wadi Ram); LEBANON: (N.
Damdur); OMAN: (Muscat, unknown localities); P.D.R. YEMEN: (Amd Town, Anag,
72
Contrib.Amer. Ent. Inst.,vol. 24, no. 1, 1988
Awabil, Geidun, Khirbat Bakarman, Khalla, Mikhuras, Mukalla, Nafhun, N’Air, Saha,
Shu’abat Amudi, Wadi Amd, Wadi Duan); SAUDI ARABIA: (Hima, Jeddah, Qunfida);
SYRIA: (Baalbelk); YEMEN ARAB REPUBLIC: (Ta’izz). An additional 81 specimens (35
females, 24 males, 4 male genitalia, 1 pupal exuviae, 6 larval exuviae, and 11
fourth-instar larvae) were examined from ALGERIA, GREECE (CRETE), ETHIOPIA,
LIBYA, MOROCCO, SOCOTRA, SUDAN, TUNISIA, and an unknown locality.
Culex (Culex) maftinglyl
K n i g ht
mattinglyi Knight, 1953b: 320. +Holotype male:
San’a, Yemen Arab Republic (NMNH).
Birket Shiekh Kunnaf,
Culex (Culex) laticinctus of Edwards, 1941: 313 (records from Zahran, Saudi Arabia, and
San’a, Yemen Arab Republic).
Culex (Culex) matting/Vi Knight of Knight, 1953b: 214-219 (Yemen Arab Republic, toll.
rec.); Mattingly and Knight, 1956: 94-134 (Saudi Arabia, Yemen Arab Republic,
A, L keys, distr., bionomics); Harbach, 1985a: 86, 95, 103 (Saudi Arabia, Yemen
Arab Republic, distr., A, L keys).
Described chiefly from the type series.
Adult. A medium-sized yellowish-brown species markedly similar to laticinctus,
differing in having normally only 1 lower mesepimeral seta and a line of pale posterior
scutal fossal scales that is continous with a posterior line of pale outer dorsocentral
scales. The thoracic integument is generally darker and the male genitalia are distinctive.
FEMALE. Head: Antenna1 length 2.0-2.5 mm, mean 2.3 mm; pedicel and first
flagellomere largely yellow or yellowish brown, rest of flagellum dark; mesal surface of
pedicel dark with patch of tiny setae and pale scales; ventromesal surface of first
flagellomere with patch of whitish scales. Proboscis dark-scaled, labella lighter; with 3-5
large basal labial setae and number of smaller decumbent setae just anterior to them;
long and slender, length 2.9-3.2 mm, mean 3.1 mm. Maxillary palpus dark-scaled; length
about 0.6 mm, approximately 0.2 proboscis length; palpomere 2 usually with 2 or 3
strong lateral setae. Vertex with narrow pale yellow falcate scales, whitish on ocular line
and interocular space; forked scales brownish, darker posteriorly; lateral spatulate scales
narrow, yellowish white. Thorax: Integument yellowish brown, darker dorsally. Scutum
with narrow golden-brown falcate scales; scales whitish to pale yellow on margins and
prescutellar area; pale line of posterior scutal fossal scales continuous with concolorous
posterior line of outer dorsocentral scales; broad strip of bare integument between
distinct lines of acrostichal and dorsocentral scales; approximately 13 well developed
lateral prescutellar setae on each side. Scutellum with narrow pale yellow falcate scales;
lateral lobes each with 4-10 and median lobe with 7-10 large setae. Ante- and
postpronota with mainly broad yellowish-white spatulate scales, some narrow ones
anteriorly; postpronotum with 5-8 setae on posterodorsal margin. Pleura with rather
small patches of yellowish-white spatulate scales below upper proepisternal setae, on
upper corner and lower posterior border of mesokatepisternum, at middle of anterior
area of mesanepimeron and among upper mesepimeral setae; pleural setae golden
brown: 5-8 upper proepisternal, 8-13 prealar, 5-7 upper mesokatepisternal, 5-10 lower
mesokatepisternal, lo-16 upper mesepimeral, and normally 1 lower mesepimeral (2 on
one side of thorax in 1 female and 1 male examined). Wing: Length 5.4-5.7 mm, mean
5.5 mm; length of cell R2 3.3-4.4 length of vein R2+3, mean 3.6; length of cell Ml
Harbach: Subgenus Culex in SouthwesternAsia
73
0.8-0.9 length of cell R2; dark-scaled with small patch of pale scales on posterior margin
at base of costa, all scales narrow, those on veins Rs, R2+3, Rl and R2 noticeably longer.
Halter: Integument dark with whitish scales covering scabellum and capitellum. Legs:
Anterior surface of forecoxa pale-scaled, with about 6 small setae and several
indistinct pale scales in short row posteriorly at apex; midcoxa with anterior patch of pale
scales; hindcoxa with narrow longitudinal stripe of pale scales on anterior side of lateral
midline. Ventral and posterior surfaces of trochanters covered with small white spatulate
scales. Fore- and midfemora dark-scaled anteriorly, pale-scaled posteriorly, with narrow
knee spots of pale scales; hindfemur mainly pale-scaled with dorsal stripe of dark scales
beginning at base and expanding over anterior and posterior surfaces on distal 0.3-0.4,
dark scaling sometimes forming complete ring distally, apex with knee spot of yellowish
scales. Tibiae dark-scaled, slightly paler posteriorly, apex of each with ring of yellowish
scales. Tarsi entirely dark-scaled. Abdomen: Tergum I with median posterior patch of
pale scales; tergum II with more or less convex basal pale band not reaching lateral
margins (one female in BM with median posterior extension of pale band); terga Ill-VII
with broad basal white bands, bands 0.3-0.5 tergum length, straight on terga Ill-V,
slightly concave on terga VI and VII; tergum VIII with concave basal pale band, band
sometimes obsolete in middle. Sterna entirely whitish-scaled; sternum VIII with lateral
scale-patches only, broad median area void of scales.
MALE. Differing from female as follows. Head: Length of antenna about 2.0-2.5
mm; pedicel and flagellomere 1 without scales. Maxillary palpus dark-scaled; with some
indistinct pale scales in dorsomesal line on distal 0.33 of palpomere 3, on ventral surface
of palpomere 4, and ventrally at base of palpomere 5; palpomeres 4 and 5 sparsely
setose, long setae mainly on palpomere 4; length about same as proboscis. Wing: Pale
scales absent at base of costa (few present at humeral crossvein in one specimen
examined). Abdomen: Terga and sterna same as in female except those of segment VIII
entirely pale-scaled. Genitalia (Fig. 23): Form as figured; phallosome resembling that of
theileriand laticinctus. Ninth tergal lobes more widely separated than usual, each with
3-8 unevenly spaced setae. Gonocoxite enlarged, ventral and mesal surfaces with
setae developed as usual, dorsolateral surface with very dense vestiture of moderately
long hairlike setae; subapical lobe prominent, very near tip of gonocoxite, undivided;
seta a well removed proximally from b and c, rather slender and curved, b long, sinuous
and stout at base, c shorter, more slender and slightly hooked at tip; d-f represented by
3 fine simple setae on mesal side of lobe; g narrow and pointed; h rather short, slender
and nearly straight. Gonostylus notably expanded, with a conspicuous recurved
thornlike process on mesal side at tip. Lateral plate of phallosome differing from that of
laticinctusin having a smaller dorsal arm, stouter ventral arm, and lateral arm with variable
number of tiny elevations in addition to 2-6 (mode 5) distinct small denticles. Proctiger
not much different than in laticinctus; basal lateral arm of paraproct not as large; cereal
setae generally fewer in number (2-6, mode 4).
Pupa (Fig. 23). Known from a single specimen in the type series; form and
placement of setae as figured, number of branches in Table 11; apparently similar to
laticinctus, seemingly differing in most of the following. Cephalothorax: Setae 8,12-CT
double; 11-CT with 4 branches; 13-CT present, small. Trumpet: Index about 4.0; pinna
slightly longer. Abdomen: Length 3.8 mm. Seta 6-I,11multiple; lo-11 with distinct stem
and many branches; alveolus of 11-II absent; 5-V,VI single, 5-VII much shorter than 4-VII.
Paddle: Length about 1.20 mm; width about 0.95 mm; index about 1.3.
Larva (Fig. 24). Character and positions of setae as figured, range and modal
number of branches in Table 31; resembles laticinctus but differs constantly in having
the thorax covered with minute spicules, seta 6-VI double, and seta 2-X double or triple.
74
Contrib.Amer. Ent. Inst.,vol. 24, no. 1, 1988
Head: Length 0.94-l .05 mm, mean 0.99 mm; width 1.26-l .48 mm, mean 1.37 mm;
moderately tanned, slightly to distinctly darker behind eyes. Dorsomentum with 9-12
(mode 10) teeth on either side of median tooth. Seta 1-C long and slender; 2-C
present; 5,6-C long, 5-C usually with 4 branches (3-5), 6-C normally triple, infrequently
with 4 branches; 8-C most often triple (2-4); 10-C forked, often triple (2-5); 14-C single.
Antenna: Long and slender, length 0.70-0.78 mm, mean 0.73 mm; lightly to moderately
tanned, distal part darker, basal rim and mesal spot at base more heavily tanned. Seta
1-A with about 21 branches (18-26); 2,3-A subapical as in laticinctus. Thorax:
Integument hyaline; covered with tiny spicules. Seta 7-P appreciably longer than usual,
usually double (2-4). Seta 1-M longer than 2-M, single; 2-M most often with 4 branches
(2-4). Seta 12-T single; 13-T short as in laticinctus but with fewer branches (4-7).
Abdomen: Integument hyaline, smooth. Seta 3-l usually triple, seldom double, 3-VII
usually double, sometimes single; 6-I-IV with 2-5 branches, 6-l,lV usually double, 6-11,111
usually triple; 6-V,VI usually double, sometimes triple; 1-IV-VI single or double, 1-VI more
often single. Segment VIII: Integument with minute spicules, particularly around comb.
Comb with 29-48 (mean 38) scales. Siphon: Index 3.40-4.55, mean 4.03; more or less
distinctly swollen before middle, apex narrowed. Pecten with 13-22 (mode 16) spines;
most spines with 2 basal denticles (l-3). Seta 1-S usually composed of 14 setae (7
pairs) but 11-l 6 may be present, paired and developed as in laticinctus. Segment X:
Saddle complete, broader than long; moderately tanned, darker dorsally; posterodorsal
margin conspicuously spiculate; length 0.37-0.42 mm, mean 0.39 mm, siphon/saddle
index 3.90-4.64, mean 4.19. Seta 1-X double or triple, more often triple; 2-X double or
triple; 4-X usually comprised of 12 setae (6 pairs), sometimes 11 or 13 are present.
Systematlcs.
Culex mattinglyi is clearly indigenous to the southwestern
mountains of the Arabian Peninsula where it is restricted to higher terrain. This species
is undoubtedly derived from the same stock as laticinctus. The two species are so similar
that Edwards (1941) failed to recognize specimens of mattinglyifrom San’a, Yemen Arab
Republic, and Zahran, southwestern Saudi Arabia, as a species different from laticinctus.
In fact, the author of mattinglyi did not realize that the type series was distinct from
laticinctus until he removed and examined the male genitalia (Kenneth L. Knight,
personnal communication). The adults of mattinglyi differ in normally having only one
lower mesepimeral seta and a diffuse line of pale scales on the scutum. The males differ
markedly in the development of the gonocoxite and gonostylus. Conspicuous
differences in the larval stage include the minutely spiculate thorax and the differently
developed setae 6-W and 2-X. The pupa of mattinglyi is known from only one specimen
in the type series so it is probably much more difficult to distinguish from that of
laticinctusthan it appears to be in the key.
Culex mattinglyi and laticinctus differ conspicuously from theileri in the
ornamentation of the adult, the structure of the comb scales of the larva, and the
development of setae 1,6-II of the pupa. Culex mattinglyi seems to be intermediate
between theileriand laticinctuswith respect to the general pigmentation of the adult, the
structure of the gonocoxopodite of the male, and the development of setae 1,6-III-VI of
the larva. I consider mattinglyi and laticinctus to be members of the theileri subgroup
because of the intermediate nature of mattinglyi. The male phallosome of these species
is undoubtedly of the type found in theileri, and it is difficult to believe that this striking
similarity is due to convergence.
Bionomics.
Knight (1953b) found the immature stages of this species in a
cement watering trough, a broad open well, a large cement basin, and a large cement
tank. Adults misidentified as laticinctus by Edwards (1941) were collected in a house.
Other than this nothing is known about the bionomics or disease relations of this
Harbach: SubgenusCulex in SouthwesternAsia
75
species.
Distribution. Culex mattinglyi is known only from the highlands of the Yemen
Arab Republic and a single locality near its northern border in Saudi Arabia.
Material
examined.
42 specimens (11 females, 5 males, 5 male
genitalia, 1 pupal exuviae, 11 larval exuviae, and 9 fourth-instar larvae) were examined
from southwestern Asia - SAUDI ARABIA: (Zahran); YEMEN ARAB REPUBLIC
(mostly type specimens): (Al-Asr, Rauda, San’a, Wadi Dhahr).
Culex (Culex)
simpsoni
Theobald
simpsoni Theobald, 1905c: 28. +Lectotype male, hereby designated,
bearing following data: “46 // Type // Transvaal / Simpson // Culex /
simpsoni / (Type) F.V.T. N Recd. from / F.V. Theobald, / 1907-29.“;
genitalia on acetate strip (BM).
richteri Ingram and De Meillon, 1927: 72. +Lectotype male, hereby
designated, bearing following data: Museum type label with “Culex”
written above and “richteri / [illegible mark]” written below the word
“Type” N “[Begamuzi River], Zululand. / 1927. / Dr. A. Ingram. / B.M.
1928-27.” (BM). Synonymy with simpsoni by Edwards, 1932a: 210.
mauritanicus Callot, 1940: 362. +Cotypes (larvae): Taghjicht, Morocco
(IPS; slide with 3 larvae retained in NMNH). NEW SYNONYMY.
Culex simpsoni Theobald of Edwards, 1913b: 58 (Africa, A, M gen.); Lewis, 1945: 16
(Sudan, distr.).
Culex (Culex) simpsoni Theobald of Edwards, 1941: 309, 420, 482 (Afrotropical
Region, A key, M*, F, P, syn., distr.); Lewis, 1943a: 282 (Eritrea, toll. rec., L
bionomic note); Hopkins, 1952: 293 (Afrotropical Region, L key, L*, bionomics);
Knight, 1953a: 215, 218, 230 (Yemen Arab Republic, toll. rec., M, distr.); Lewis,
1956: 709 (Sudan, toll. sites); Mattingly and Knight, 1956: 94-132 (Yemen
Arab Republic, A, L keys, distr., bionomics); Ovazza et al., 1956: 172 (Ethiopia,
toll. sites, L assoc.); Harbach, 1985a: 86, 93, 106 (Yemen Arab Republic only,
distr., A, L keys).
Described chiefly from specimens collected in eastern Africa.
Adult. This and the following species resemble univittatus and perexiguus in
most respects, but are easily distinguished by the absence of an anterior pale stripe on
the hindtibia. They also lack a spot of pale scales at the base of the costa.
FEMALE. Proboscis, maxillary palpus, legs and abdominal terga largely clothed in
dark brown scales. Head: Antenna dark, pedicel with pale scales on dorsomesal surface;
length 1.6-1.9 mm, mean 1.7 mm. Proboscis with pale scaling in middle of ventral
surface, sometimes forming more or less complete ring; length 1.6-2.0 mm, mean 1.8
mm. Maxillary palpus with some pale scales dorsomesally at apex; length about 0.4 mm,
approximately 0.2 proboscis length. Falcate scales of vertex coarse, yellowish white;
forked scales long, yellowish medially, dark laterally; ocular scales elongate, white,
continuous with elongate white spatulate scales of postgenal area. Thorax: Integument
brown. Scutum with yellowish and golden-brown falcate scales, often forming pattern
consisting of median stripe on anterior 0.5, patches on fossae and pair of diffuse
patches behind middle; with paler scaling on anterior and lateral margins, posterior
dorsocentral area and lateral prescutellar area. Scutellum with white to very pale yellow
76
Contrib.Amer. Ent. Inst.,vol. 24, no. 1, 1988
falcate scales mainly on median lobe, scales larger on median lobe and projecting laterad
on either side of midline. Ante- and postpronota with some golden-yellow falcate scales,
longer on antepronotum. Pleura with patches of broad white spatulate scales as follows:
patch ventral and mesal to upper proepisternal setae, patch on postspiracular area near
spiracle, small patch below prealar knob, larger patches on upper corner and lower
posterior border of mesokatepisternum, anterior patch on mesepimeron about same
level and size as upper mesokatepisternal patch, and small patch before upper
Pleural setae golden brown, numbers follow: 4-6 upper
mesepimeral setae.
proespisternal, 6-l 0 prealar, 4-6 upper mesokatepisternal, 4-7 lower mesokatepisternal,
3-5 upper mesepimeral, and 1 lower mesepimeral. Wing: Length 3.0-3.5 mm, mean 3.2
mm; length of cell R2 2.0-2.5 length of vein R2+3; length of cell Ml about 0.8 length of
cell R2; entirely dark-scaled. Halter: Pedicel and scabellum pale; capitellum dark brown.
Legs (hindleg, Fig. 2L): Forecoxa pale-scaled with dark scales in middle; mid- and
hindcoxae with longitudinal line of pale scales on anterior side of lateral midline, scales
slightly more numerous on hindcoxa. Femora with narrow knee spots of white scales;
anterodorsal surface of forefemur dark-scaled, posteroventral surface white-scaled;
midfemur dark anteriorly, dark scaling expanded over dorsal surface toward apex,
posterior surface white-scaled; hindfemur largely white-scaled, with narrow dorsal stripe
of dark scales beginning near base and abruptly widening to encircle femur on distal 0.2.
Tibiae with spots of white scales dorsally at apex, large and conspicuous on hindtibia;
fore- and midtibiae dark-scaled anteriorly, pale-scaled posteriorly; hindtibia very dark,
nearly black on dorsal surface. Tarsi entirely dark-scaled. Abdomen: Tergum I with large
median patch of brown scales; terga II-VII with distinct, slightly convex, white basal bands
that are narrowly joined to basolateral spots of same color, bands 0.20-0.35 tergum
length; tergum VIII normally entirely white-scaled. Sterna with broad dark apical bands
(0.45-0.50 sternum length), usually produced anteriorly in middle; sternum VIII with
lateral patches of pale scales only.
MALE. Like female except as follows. Head: Proboscis sometimes with few pale
scales ventrally at base; length shorter, about 1.4 or 1.5 mm. Maxillary palpus dark
brown; lateral surface of palpomere 3 with median patch of white scales; palpomere 4
with ventral line of white scales on proximal 0.5; palpomere 5 with ventral spot of white
scales at base. Scales of vertex generally paler. Wing: Slightly shorter, 2.6-3.1 mm,
mean 2.9 mm; length of cell R2 1.6-2.1 length of vein R2+3. Legs: Midfemur sometimes
with very narrow line of pale scales on anterior surface. Abdomen: Terga without
basolateral pale spots. Sterna sometimes almost entirely dark-scaled, with basolateral
pale spots; tergum VIII (ventral in position) entirely dark-scaled or with narrow posterior
band of pale scales. Genitalia (Fig. 25): Form as figured; general structure of
phallosome as in decens. Ninth tergal lobe not produced, with 5-10 (mode 7) short
setae. Gonocoxite normal, ventral and mesal setae developed as usual, lateral setae in 2
rows of 4 or 5 setae each; subapical lobe undivided, setae a-cof the usual form, group
d-e absent, fstout (on order of magnitude of a-c), tapered and flattened apically, g
asymmetrical (triangular) and rounded apically, h gently curved. Gonostylus narrow,
tapered; gonostylar claw rather broad, flattened and curled. Phallosome longer than
broad, lateral plate larger than aedeagal sclerite; dorsal arm more or less straight,
diverging slightly from its mate of the opposite side; lateral arm with graded series of
laterally directed denticles on caudal margin, 2-5 (mean 4) larger denticles project over
13-23 (mean 18) smaller denticles which grade into small bumps laterally, small denticles
extend onto prominent dorsal process; ventral arm much as in decens. Proctiger
unmodified; paraproct with long, curved, rodlike basal lateral arm and distinct ventral
acetabulum. Cereal setae most often 3 (2-4).
Harbach:SubgenusCulexin SouthwesternAsia
77
Pupa (Fig. 25). Not associated with adults from southwestern Asia; form and
placement of setae as figured, range and modal number of branches in Table 12; much
the same as antennatus, but generally smaller, as reflected in the length of the abdomen
and paddle, and pinna of trumpet usually shorter. Cephalothorax: Not distinguishable
from antennatus. Trumpet: Index 4.6-6.3, mean 5.2; pinna short, usually less than 0.2
trumpet length. Abdomen: Length 2.2-2.6 mm, mean 2.4 mm. Seta l-11 not so variable,
generally with fewer branches (12-19); 6-lll,lV like 6-V,VI, often with 5 branches (4-6).
Paddle: Length 0.63-0.77 mm, mean 0.69 mm; width 0.48-0.59 mm, mean 0.50 mm;
index 1.3-1.6, mean 1.4.
Larva (Fig. 26). Not associated with adults from southwestern Asia; character
and positions of setae as figured, range and modal number of branches in Table 32; this
and the next species are readily separable from all other species of Culex in
southwestern Asia by the combined character of seta 5-C seta 14-P, and the comb
scales. Head: Length 0.58-0.72 mm, mean 0.64 mm; width 1.01-l .14 mm, mean 1.06
mm; lightly tanned, darker behind eyes. Dorsomentum with 4-6 teeth on either side of
median tooth, third from center distinctly larger. Seta 1-C long, slender, pale; 2-C
absent; 4-C rather smaller than usual; 5-C single or double, most often single; 6-C
usually double, occasionally single; 1O-C usually triple (2,3); 11 -C double; 13-C most
often triple, sometimes with 4 branches; 14-C normally double with markedly divergent
branches (2-4). Antenna: Length 0.50-0.66 mm, mean 0.60 mm; proximal part lightly
tanned with dark basal ring contiguous with dark basomesal spot, distal part moderately
tanned. Seta 1-A with about 27 branches (17-33). Thorax: Integument hyaline,
smooth. Setae 4,7-P with 3 or 4 branches, 4-P more often with 4, 7-P most often triple;
14-P normally double, sometimes single on one side. Seta 1-M usually single, seldom
double, longer than 2-M; 5-M shorter than usual. Seta 1-T normally single, seldom
double, 0.5 or less length of 2-T; 2-T usually double, rarely single, occasionally triple;
13-T usually with 5 or 6 branches (5-7). Abdomen: Integument hyaline, smooth. Seta
3-I-IV double (3-l occasionally triple), 3-V,VI single, 3-VII variable (2-5); 6-I-VI normally
triple (see table); 7-l usually double, sometimes single; l-II-V with 4 or 5 branches, 6-VI
with 3 or 4 branches, 6-VII often with 5 or 6 branches (4-7). Segment VIII: Comb with
17-36 scales, mean 28; most scales with sharp apical spine, some anterior scales fringed
entirely, apical spine usually larger on posterior scales. Siphon: Index 5.14-5.67, mean
5.40; slender, tapered, broadest at base; lightly tanned, basal rim and acus moderately
tanned. Pecten on proximal 0.33 or less, composed of 9-14 (mode 12) rather long
slender spines with tiny basal denticles. Seta 1-S in 4 pairs, la-S occasionally within
pecten, lb- and lc-S out of line with the others; usually double or triple (see table).
Segment X: Saddle lightly tanned, with very inconspicuous posterodorsal spicules;
length 0.35-0.39 mm, mean 0.37 mm, siphon/saddle index 2.98-3.97, mean 3.85. Seta
1-X usually triple, sometimes double; 2-X double, triple, or with 4 branches occurring in
nearly equal frequencies; 4-X almost always in 6 pairs.
Systematics. The simpsoni subgroup is erected for simpsoni and sinaiticus,
and also undoubtedly includes striatipes Edwards, 1941, terzii Edwards, 1941, and
seldeslachtsi Wolfs, 1947 from the Afrotropical Region. The group is homogeneous in
all essential adult and larval features. The adults resemble univittatus in ornamentation
and coloration, but differ markedly in the male genitalia. The male genitalia resemble the
type found in the decens subgroup, but the species are otherwise distinct in all life
stages. The larvae share some characteristics with theileri, particularly the form of the
comb scales, but differ conspicuously in the position and development of seta 1-S.
Culex simpsoni shows relatively little variation in adult characters, particularlythe
male genitalia. The larva is somewhat variable in details of the comb and siphon, but
78
Contrib.Amer. Ent. Inst.,vol. 24, no. 1, 1988
there is no indication that more than one species is involved in the various separated
populations examined. During this study, several individually reared specimens from
Morocco previously identified as Culex mauritanicus Callot, 1940, and three cotype
larvae of this nominal species, were examined and found to be remarkably similar to the
type specimens of simpsoni and richteri in all important features. The similarities are so
evident in all stages that for the present I cannot consider mauritanicus as a distinct
species.
The occurrence of simpsoni in southwestern Asia is based upon two males
collected in the Yemen Arab Republic (Knight, 1953a). The association of the immature
stages with the adults is based on a small number of individual rearings from Tanzania
and Morocco, and the type series of richteri. The unknown stages of simpsoni in
southwestern Asia should agree with the descriptions above based on the overall
conformity of the males with African specimens.
Bionomics. Culex simpsoni is found in a wide variety of habitats, but its original
breeding places were probably sunlit pools of slow or stagnant water along streams or
streambeds. Besides these habitats, simpsoni is commonly found in lakes, ponds,
marshes, puddles, furrows, seepage pools, and artificial pools such as tubs, wells,
troughs, and tanks. Ribeiro and Mexia (1966) collected larvae in relatively saline
marshes and ground pools with somewhat turbid water and muddy bottom in Angola.
Larvae have been found in association with Anopheles (Celia) cinereus Theobald, An.
(Gel.) pharoensis Theobald, An. (Gel.) sergentii (Theobald), Culex fheileri, and Cx.
(Maillotia) arbieeni Salem (as jenkinsi Knight) in southwestern Asia (Knight, 1953a), with
An. (Gel.) arabiensis Patton (as gambiae Giles) in Uganda (Harris, 1942) and with Culex
antenna&s, Cx. species near infula Theobald (as bitaeniorhynchus Giles), and Cx.
(Culex) thalassius Theobald in Angola (Ribeiro and Mexia, 1966). Nothing is known
about the vector potential of this species, but it is not likely to be of importance in
disease transmission because it only rarely bites man (Mattingly and Brown, 1955).
Distribution. This species is widely distributed in the Afrotropical Region and is
known to occur in the northwest African countries of Mauritania and Morocco.
Material examined. 229 specimens. Only 4 specimens (1 female, 2 males,
and 1 male genitalia) were examined from southwestern Asia - YEMEN ARAB
REPUBLIC: (El-Hauban , Wadi el-Malak near Ta’izz). A total of 225 specimens (49
females, 52 males, 24 male genitalia, 15 pupal exuviae, 13 larval exuviae, and 72
fourth-instar larvae) were examined from African countries which include ETHIOPIA,
KENYA, MALAGASY REPUBLIC, MAURITANIA, MOROCCO (including 3 cotype larvae
of mauritanicus ), SOUTH AFRICA (the type specimens of simpsoni and richten],
TANZANIA, ZAIRE, ZIMBABWE, and unknown countries.
Culex (Culex)
sinaiticos
Kirkpatrick
sinaiticus Kirkpatrick, 1924(1925): 383. +Syntypes (male; female):
Kossaima, El Moweilleh, Ain Kadeis, Ain Gedeirat, and Ain Musa,
Sinai , Egypt (DI; ESE).
Culex sinaiticus Kirkpatrick of Salem, 1938: 27 (Sinai, toll.); Lewis, 1945: 16 (Sudan, L
distr.); Leeson and Theodor, 1948: 228 (Socotra, A, L keys, toll. sites, L habitat);
Senevet et al.,l949: 46, 47 (Egypt [probably Sinai], A, M gen., L); Theodor,
1952: 113 (Middle East, zoogeogr.); Senevet et al., 1957a: 88 (North Africa, L);
Margalit and Tahori, 1973: 90-93 (Sinai, distr., L bionomics); Margalit et al., 1973:
Harbach: Subgenus Culex in SouthwesternAsia
79
32 (Israel, toll. rec., L assoc.).
Culex (Culex) sinaiticus Kirkpatrickof Kirkpatrick,1925: 122, 191 (Sinai, A, P, L keys, M’,
F, P*, L’, bionomics, distr.); Edwards, 1926: 138 (Sinai, A, L keys, A, L, distr.);
Stackelberg, 1927: 162 (Sinai, M, F keys, A, M gen.*, distr.); Martini, 1931: 381
(Sinai, A, L keys, M*, F, L’); Edwards, 1941: 311,420, 482 (P.D.R. Yemen, Sinai,
Somalia, Sudan, A key, M’, F, P, distr.); Lewis, 1943a: 282 (Eritrea, toll. rec., L
morphol. note, L bionomic note); Lewis, 1943b: 69 (Sudan); Abbott, 1948: 45
(Sudan, L, L habitat, distr.); Hopkins, 1952: 295 (Afrotropical Region, L key, L,
bionomics); Knight, 1953a: 216, 230 (Yemen Arab Republic, coll. rec., L, L
habitat, distr.); Lewis, 1956: 709 (Sudan, Eritrea, A, toll. sites, distr.);
Abdel-Malek, 1956: 100 (Sinai, L bionomics, distr., L key); Mattingly and Knight,
1956: 94-132 (Oman, P.D.R. Yemen, Saudi Arabia, Socotra, Yemen Arab
Republic, A, L keys, distr., bionomics); Senevet et al., 1957b: 92 (North Africa, L);
Senevet and Andarelli, 1959: 192 (SW Asia, A, P, L keys, M*, F, P, L, distr.,
bionomics); DuBose and Curtin, 1965: 352, 354 (Mediterranean area east of
Suez Canal, A, L keys); Lotfi, 1970: 401 (Iran, toll. rec.); Lotfi, 1973: 206 (Iran,
coll. sites); Margalit and Tahori, 1974: 87 (Israel, toll. rec.); Lotfi, 1976: 72, 74, 79
(Iran, L key, ecol. note, L*); Harbach, 1985a: 86, 93, 106 (SW Asia, Egypt, A, L
keys).
Adult.
A rather small species closely resembling simpsoni, but with an
incomplete dark dorsal stripe on the distal half or less of the hindfemur, the abdominal
sterna largely pale-scaled, and different maxillary palpi and genitalia in the male.
FEMALE. Head: Antenna dark, pedicel paler; pedicel and flagellomere 1 with
some pale scales on mesal side; length about 1.5 or 1.6 mm. Proboscis dark with
indefinite broad area of pale scaling in midregion of ventral surface, sometimes forming
more or less complete ring; length about 1.6 or 1.7 mm. Maxillary palpus about 0.2
proboscis length; dark-scaled with pale scales dorsally on palpomere 3 and base and
apex of palpomere 4. Falcate scales of vertex coarser than in simpsoni, yellowish white;
ocular and interocular scales white; forked scales mainly yellowish, paler anteriorly, dark
posterolaterally; lateral spatulate scales white. Thorax (Fig. 2A): Integument brown.
Scutum with pale golden-brown falcate scales, whitish on margins; scutal setae dark
brown with reddish and golden tint. Scutellar scales whitish, few on lateral lobes,
confined mainly to median lobe. Antepronotum with large, almost white falcate scales
ventrally; postpronotum with similar but slightly yellowish scales, with 4 golden setae on
posterodorsal border. Pleural setae golden to golden brown, numbers follow: 4 or 5
upper proepisternal, 6-9 prealar, 3-5 upper mesokatepisternal, 2-6 lower mesokatepisternal, 3-5 upper mesepimeral, and 1 lower mesepimeral. Pleura with broad white
spatulate scales as follows: patch ventral and mesal to upper proepisternal setae, patch
on anterodorsal border of postspiracular area just behind spiracle, small patch below
prealar knob, larger patches at upper corner and on lower posterior border of
mesokatepisternum, mesepimeron with anterior patch about same level and size as
upper mesokatepisternal patch, and small upper patch before upper mesepimeral setae.
Wing: Length about 3.0 mm; length of cell R2 2.0-2.3 length of vein R2+3, mean 2.1;
length of cell Ml/length of cell R2 0.7-0.8; entirely dark-scaled; dorsal scaling: elongate
spatulate scales on costa, subcosta, R4+5, mcu, M3+4, CuA and proximal 0.3 of IA,
linear or near linear scales on other veins and distal part of 1A; ventral scaling: elongate
spatulate scales on costa, subcosta, Rl, R2+3, bases of R2 and Ra, M, Ml+& and
proximally on Ml and M2, scales absent from R, CuA before mcu and proximal 0.5 of 1 A,
other veins and distal parts of CuA and 1A with linear or near linear scales. Halter: Pedi-
80
Contrib.Amer. Ent. Inst.,vol. 24, no. 1, 1988
ccl and scabellum pale; capitellum dark. Legs (hindleg, Fig. 2M): Anterior surface of
forecoxa completely white-scaled or with dark scales in middle; mid- and hindcoxae with
line of white or slightly yellow scales on anterior side of lateral midline. Femora with
distinct white knee spots; fore- and midfemora with anterior dark scaling expanded
distally over dorsal surface; forefemur often with narrow anteroventral line of white scales
on proximal 0.5; midfemur with nearly complete white anteroventral stripe that terminates
before apex, stripe often reduced to narrow line on distal 0.5; hindfemur largely white,
with dorsal dark stripe on distal 0.5 or less (usually about 0.4) that abruptly expands over
anterior and posterior surfaces on distal 0.1-0.2 (normally about 0.2). Tibiae with
conspicuous white anterodorsal spots at apex; anterior and dorsal surfaces of foretibia
dark-scaled, posterior and ventral surfaces pale-scaled; midtibia pale-scaled dorsally,
dark-scaled ventrally; hindtibia mainly dark-scaled, posteriorly with narrow longitudinal
stripe of pale scales not reaching base or apex (about 0.2 from both). Tarsi mainly
dark-scaled; posteroventral surface of tarsomere 1, and often 2, with pale scaling.
Abdomen: Tergum I with median posterior patch of black, black and white or white
scales; terga II-VII with white basal bands 0.3-0.4 tergum length, slightly produced
laterally, especially on terga VI and VII; tergum VIII almost entirely pale-scaled. Sterna
pale-scaled, infrequently with some subtle dark scales on posterolateral corners of
sterna VI and VII; sternum VIII with scales on lateral borders only.
MALE. Like female except as follows. Head: Antenna shorter, length 1.2-l .5
mm, mean 1.4 mm. Length of maxillary palpus 1.7-l .9 mm, mean 1.8 mm, extending
beyond tip of proboscis by length of palpomere 5; palpomere 3 with broad line of white
scales on dorsolateral margin not reaching base or apex, also with 3 or 4 long setae and
several shorter ones distally beyond pale scaling on ventrolateral margin; palpomere 4
with ventral line of white scales not always reaching apex; palpomere 5 with ventral spot
of white scales at base. Legs: Anteroventral pale stripe of midfemur poorly developed
or absent. Abdomen: Sterna usually with some dark scaling on posterior margin or
posterolateral corners. Genitalia (Fig. 27): Form as figured; phallosome developed as in
simpsoni. Ninth tergal lobe with 4-7 (mode 5) setae. Gonocoxite unmodified, lateral
setae more numerous than in simpsoni (about 21 in 4 rows, increasing in length toward
base of gonocoxite), apex with conspicuous cluster of about 9 moderately long setae on
mesal side of gonostylus; subapical lobe undivided, setae &similarly developed, rather
short, all about same length (fthe shortest), all flattened and hooked at tip; g a rather
long, crescentic blade; h long and sharply bent distally. Gonosylus rather short and
narrow, distinctly flattened distally, rugose dorsally before tip. Phallosome very much as
in simpsoni, lateral plate with fewer denticles, 3-5 (mode 4) larger denticles project over
7-16 (mode 11) smaller denticles, smaller denticles not as well developed, often
reduced to mere bumps. Proctiger and cereal setae as in simpsoni.
Pupa (Fig. 27). Form and positions of setae as figured, range and modal number
of branches in Table 13; not separable from simpsoniunless by the possible distinctions
noted below. Cephalothorax: As in antennatus and simpsoni. Trumpet: Index 4.0-7.3,
mean 5.6; pinna usually longer, 0.2-0.3 length of trumpet. Abdomen: Length 2.1-2.6
mm, mean 2.4 mm. Setae l-Ill-VI, 5-111-V,and 6-III-VI generally shorter. Paddle: Length
0.72-0.82 mm, mean 0.77 mm; width 0.49-0.58 mm, mean 0.54 mm; index 1.3-l .6,
mean 1.4.
Larva (Fig. 28). Character and placement of setae as figured, range and modal
number of branches in Table 33; resembles simpsoni in most respects, differing mainly
in characters of the siphon. Head: Length 0.61-0.75 mm, mean 0.69 mm; width
1.01-l .18 mm, mean 1.08 mm; lightly tanned, not noticeably darker behind eyes.
Dorsomentum usually with 5 teeth (4,5) on either side of median tooth, shape as in
Harbach: Subgenus Culex in SouthwesternAsia
81
simpsoni. Seta 1-C long, very slender, pale; 5-C single, smaller than in simpsoni; 10-C
more often double (2-4); 11-C usually double, seldom single; 13-C with 3 or 4 branches,
more often with 4. Antenna: Length 0.60-0.69 mm, mean 0.63 mm. Seta 1-A with
about 25 branches (21-28). Thorax: Seta 4-P double, triple or with 4 branches, seldom
single. Abdomen: Segments III and V with pigmentation giving abdomen a 2-banded
appearance similar to antennatus. Seta 3-l double or triple, 3-II-IV double (3-11,111
seldom
single), 3-V,VI single (3-V seldom double), 3-VII with 3 or 4 branches; 1-III usually with 3
or 4 branches (3-5), 1-IV,V with 4 or 5 branches, l-VI often with 4 branches (2-5).
Segment VIII: Comb with 32-44 scales, mean 37; anterior scales small and evenly
fringed, posterior scales larger with long apical spine. Siphon: Index 5.97-8.07, mean
6.97; long, slender, very slightly expanded apically; lightly tanned, slightly darker
distally, basal rim dark. Pecten on proximal 0.4, composed of 11-19 (mode 17) spines
similar to those in simpsoni. Seta 1-S in 4 pairs, first pair occasionally within pecten,
individual elements in straight row on posterolateral margin; normally double or triple
(see table). Segment X: Saddle longer than broad, length 0.33-0.40 mm, mean 0.38
mm, siphon/saddle index 3.35-3.80, mean 3.60. Seta 1,2-X double or triple; 4-X usually
in 6 pairs, occasionally in 7, sometimes with 1 seta missingon 1 side.
Systematics.
Culex sinaiticus is very similar in all stages to simpsoni, but is
normally distinguishable in the adult and larval stages by the characters given in the
keys. The larva also apparently differs consistently in the positions of seta 1-S. The
pupae cannot be differentiated at present. The males are easily recognized by the
uniquely developed setae of the subapical lobe of the gonocoxite. The latter is
probably the most derived feature of the species.
Three larvae from southeastern Iran were available during this study which differ in
vague details from typical sinaiticus. The comb scales are larger than usual and more
closely resemble those of simpsoni. These specimens were not examined critically, but I
suspect that another species may be involved. The single male from Muscat, Oman,
noted by Mattingly and Knight (1956) seems to differ little from specimens collected
elsewhere in Arabia.
Culex beta Seguy, 1924 could be a synonym of sinaiticus. This nominal form was
described from an undisclosed number of larvae collected in Algiers. The partial
illustration of the larva published with the original description bears a definite
resemblance to sinaiticus and I would regard beta as a synonym of this species except I
have never seen a specimen identified as beta. There are no labelled specimens in the
alleged type depository (Harbach, 1983) and sinaiticus has never been recorded from
northern Africa. A thorough survey of the type locality is necessary before it can be
decided whether beta is a valid species or a synonym of sinaiticus.
Blonomlcs.
The immature stages of sinaiticus are commonly found in quiet
permanent or semipermanent bodies of clean water such as streams, stream pools,
springs, wells, and seepages. The breeding sites often contain green algae or higher
plants. The species tolerates water with relatively high salt content, but occurs most
frequently in fresh water. Associated species of the subgenus Culex in southwestern
Asia include bitaeniorhynchus, laticinctus, mimeticus, perexiguus, pipiens, theileri, and
tritaeniorhynchus. The species has also been collected in association with Anopheles
(Celia) arabiensis Patton, An. (Gel.) dfhali Patton, An. (Gel.) hispaniola (Theobald), An.
(Gel.) multicolor Cambouliu, An. (CeL) pharoensis Theobald, An. (Gel.) pretoriensis
(Theobald), An. (Gel.) rhodesiensis Theobald, An. (Gel.) sefgentii (Theobald), An. (Gel.)
superpictusGrassi, An. (Gel.) tutkhudi Liston, Culex (Maillotia) arbieeni Salem, Cx. (Ma/.)
deserticola Kirkpatrick, Culiseta (Allotheobaldia) longiareolata (Macquart), and
Uranotaenia (Pseudoficalbia) unguiculata Edwards. Kirkpatrick(1925) found adults of
82
Contrib.Amer. Ent. Inst., vol. 24, no. 1,1988
sinaiticus in a rest-house and reported that females bite at night. Nothing else is known
about the bionomics of the adults, but the species does not appear to be involved in
disease transmission.
Distribution. Culex sinaiticus is known from the Arabian Peninsula, the Sinai
Peninsula of Egypt, Israel, Jordan, Iran, Sudan, Ethiopia, Somalia, and Socotra.
Material examined.
196 specimens. A total of 171 specimens (38 females,
40 males, 19 male genitalia,1 2 pupal exuviae, 10 larval exuviae, and 52 fourth-instar
larvae) were examined from southwestern Asia - EGYPT (including type specimens) :
Sinai Peninsula (Ain Gedeirat, Ain Musa, Moweilleh, Wadi Feiran, unknown localities);
IRAN: (Hajiabad, Sarbaz Zahedan, Shah-Kahoor Zahedan); ISRAEL: (En Avedat,
Kadesh Baruea, Neot Hakikar, St. Caltrie, Tel Nagila, unknown localities); JORDAN:
(Amman, Aqaba, Wadi Hasa); OMAN: (Muscat); P.D.R. YEMEN: (Harshiyat, Hami, Wadi
Ma’adin); SAUDI ARABIA: (Al Hasa); YEMEN ARAB REPUBLIC: (unknown locality). An
additional 25 specimens (5 females, 9 males, 7 male genitalia, and 4 fourth-instar larvae)
were examined from ETHIOPIA, SOCOTRA, and SUDAN.
Culex (Culex)
duttoni
T h e o b a Id
duttoni Theobald, 1901c: v. +Lectotype female, hereby designated,
bearing following data: “Type // Culex / Duttoni [sic] / (Type)
Theobald N C. No II / fr. larvae / 15.4.00 / Canoes Creek / Duke Town
[Nigeria]” (BM).
dissimilis Theobald, 1901 a: 376. +Lectotype male, hereby designated,
bearing following data: “Mt. Aureol, / Free Town. / 9. 9. 99. / Bred fr.
water / in drain. // Type // Free Town. / Sierra Leone. / VIII. 99. / E.E.
Austen. / 99. 267. // Culex / dissimilis / (Type). Theobald” (BM).
Synonymy with dutfuni by Edwards, 1911: 259.
hirsutipalpis
Theobald, 1901a: 378. +Lectotype male, hereby
designated, bearing following data:
“Type / SALISBURY /
Mashonaland [Rhodesia] / Feb. 1900 / G.A.K. Marshall / 79 / Culex /
hirsutipalpis / (Type) Theobald.” (BM). Synonymy with duttoni by
Edwards, 1911: 259.
anarmostus Theobald, 1903a: 170.
+Holotype female: Freetown,
Sierra Leone (BM). Synonymy with dutfoni by Edwards, 1911: 259.
bifoliata Theobald, 1905c: 31. +Lectotype male, hereby designated,
bearing following data: “Type // Culex / bifoliata. / Type. F.V.T. //
Transvaal / Dr. Simpson // 54- // Recd. from / F.V. Theobald, /
1907-29.” (BM). Synonymy with duttoni by Edwards, 1941: 300.
condylodesmus
Grunberg, 1905: 385. Syntypes (male; female):
Victoria, Kamerum, [Nigeria] (ZM). Synonymy with duttoni by
Edwards, 1911: 259.
albovirgatus Graham, 1910: 264. +Lectotype male, hereby designated,
bearing following data: “Yaba / 26-5-09 / from larva in / a fetish pot /
WMG N Type N Yaba, / Lagos, / W. Africa. [Nigeria] / 26. V. 1909. / Dr.
WM. Graham. / 1910-80.” (BM). Synonymy with duttoniby Edwards,
1911: 259.
Culex dissimilisTheobald of Wesche, 1910: 40 (W. Africa, P, L keys, P*, L*).
Culex dufloniTheobald of Wesche, 1910: 34 (W. Africa, P, L keys, P*, L*); Lewis, 1945:
Harbach: Subgenus Culex in SouthwesternAsia
83
15 (Sudan, distr.).
Culex (Culex) duftoniTheobald of Edwards, 1941: 300, 418, 482 (Afrotropical Region,
A key, M*, F’, P*, syn., distr.); Lewis, 1943a: 281 (Eritrea, coll. rec., L bionomic
note); Lewis, 1943b: 69, 72 (Sudan, L habitat, bionomic note); Abbott, 1948: 45
(Sudan, distr.); Hopkins, 1952: 287 (Afrotropical Region, L key, L*, bionomics);
Lewis, 1956: 707 (Sudan, Eritrea, toll. sites); Mattingly and Knight, 1956: 94-135
(P.D.R. Yemen, A, L keys, distr. bionomics); Ovazza et al., 1956: 168 (Ethiopia,
colt sites); Mekuria, 1968: 78 (Ethiopia, distr., rec.); Harbach, 1985a: 86, 92, 102
(SW Asia, distr., A, L keys).
Adult. A rather large species easily distinguished from all other species of Culex
known to occur in southwestern Asia by the presence of an indefinitely ringed
proboscis, ringed tarsi, postspiracular scales, an anterior white stripe on the midtibia, and
usually 2 to 4 lower mesepimeral setae.
FEMALE. Head: Antenna dark, length about 2.4 mm; pedicel pale laterally,
mesal surface usually with some pale scales; flagellum normal, most whorls with 5 or 6
setae. Proboscis black-scaled at base and apex, middle with ring of whitish scales
beginning 0.2-0.4 from base and extending to point about 0.7 from base, proximal
boundary of ring ill-defined, distal boundary sharply defined, ring sometimes incomplete
or nearly so on dorsal surface; length about 2.2 mm. Maxillary palpus mainly
black-scaled, palpomere 4 with dorsomesal stripe of white scales; length about 0.5 mm,
about 0.2 length of proboscis. Forked scales of vertex mainly pale, darker laterally;
falcate scales mainly pale, usually yellowish white in middle, somewhat brownish laterally
and white along margins of compound eyes; lateral spatulate scales white, often dark
posteriorly and in middle of lateral surface. Thorax: Pleural integument brown, paler on
middle of mesokatepisternum and posterior area of mesepisternum, pale areas and
scale-patches separate dark areas to give pleura a spotted appearance; scutal
integument dark brown. Scutum with fine, rather densely matted falcate scales, scales
mainly golden brown to brown but with an indefinite and irregular mottling of pale yellow
to golden scales, usually with whitish scales on anterior and lateral margins, prescutellar
area and forming indefinite middorsal pair of submedial spots; scutal setae prominent
only on supraalar area. Scutellar scales same as prescutellar scales; lateral lobes each
usually with 4 or 5 large setae, median lobe normally with 8 large setae. Antepronotum
with upper and lower patches of falcate scales, upper patch golden to golden brown,
lower patch paler, usually whitish. Postpronotum mainly with golden-brown scales,
scales usually whitish posteriorly; with 6-9 dark setae in posterodorsal row, often paler
posteriorly. Pleural setae golden to golden brown: 5-9 upper proepisternal, 12-14
prealar, 5 upper mesokatepisternal, 5-7 lower mesokatepisternal, 6-12 upper mesepimeral and l-4 (usually 2; often 3) lower mesepimeral. Pleural scales white, anterior
part of postspiraculararea with falcate scales, other areas with narrow spatulate scales as
follows: few below upper proepisternal scales, upper and lower patches on mesokatepisternum and anterior and upper patches on mesepimeron. Wing: Length 4.2-4.5
mm, mean 4.4 mm; length of cell R2/length of vein R2+3 3.0-3.9, mean 3.4; subcosta
intersects costa just before furcation of R2+3; length of cell Ml/length of cell R2 slightly
more than 0.8; dark-scaled with line of pale scales on posterior margin of costa at base.
Dorsal scaling: narrow spatulate scales on costa, subcosta, R and Rl; very narrow
spatulate scales on R4+5, Ml, M2, mcu, M3+4, CuA and approximately proximal 0.5 of
1A; linear scales on Rs, R2+3, R2, R3, M, Ml+2 and distally on 1A. Ventral scaling:
narrow spatulate scales on costa, subcosta, Rs, R1+2, M, M1+2, CuA proximal of mcu and
proximal 0.5 of IA; linear scales on R, Rl, R2, R3, R4+5, mar, M3+4 and CuA
a4
Contrib.Amer. Ent. Inst.,vol. 24, no. 1, 1988
beyond mcu. Halter: Entirely pale. Legs (midleg, Fig. 2H): Mainly with dark brown to
black scales. Anterior surface of forecoxa mainly dark-scaled, with basal and distal
patches of white scales; anterolateral margin of midcoxa with longitudinal stripe of white
scales margined anteriorly by longitudinal stripe of dark scales, white stripe broadest at
base, dark stripe broadest at apex; integument of hindcoxa nearly colorless, lateral
surface with longitudinal stripe of white scales and small spot of dark scales at apex.
Trochanters mainly with white scales, anteroventral surface of fore- and midtrochanters
usually with some dark scales. Femora each narrowly pale at base and with indistinct
narrow pale knee spot; forefemur with narrow anteroventral line of pale scales on
proximal 0.5, posterior surface with posterodorsal pale stripe and more or less dintinct
narrow median line of pale scales; posterior surface of mid- and hindfemora pale-scaled
except on distal 0.3 or less. Tibiae each with narrow white ring at base; posterior surface
of foretibia with complete white stripe; midtibia with narrow white anterodorsal and
broader posterior white stripes; hindtibia with incomplete posterior pale stripe. Tarsi with
narrow white rings across joints, rings may be indistinct or absent between tarsomeres 4
and 5; tarsomere 1 of each leg with narrow basal white ring, tarsomere 1 of fore- and
midlegs with pale scales on posterior surface, hindtarsomere 1 with pale scales on
ventral surface. Pulvilli distinct, white. Ungues dark, small, simple. Abdomen: Tergum I
with median posterior patch of black scales; terga II-VIII mainly dark brown- to
black-scaled, tergum II with small basomedian triangular patch of cream-colored scales,
terga Ill-VIII with narrow, convex, cream-colored bands 0.20-0.35 tergum length and
elongate basolateral white patches which become longer on more posterior terga,
bands and spots not connected on anterior terga, narrowly if at all connected on terga
V-VIII, band usually present and continuous with spots on tergum VIII. Sterna II-VII mainly
with cream-colored scales, with black scales on posteriorcorners; sternum VIII with lateral
patches of cream-colored scales, broad median area without scales.
MALE. Like female except as follows. /-lead: Antenna strongly plumose, pale
between flagellar whorls; length about 1.8 mm. Proboscis with narrow pale ring
0.55-0.70 from base; false joint about 0.65 from base; length 2.3-2.9 mm, mean 2.5 mm.
Length of maxillary palpus 3.1-3.4 mm, mean 3.3 mm, extending beyond tip of
proboscis by more than length of palpomere 5; palpus mainly black-scaled, integument
between palpomeres 2 and 3 pale; palpomere 3 white-scaled in middle, ventromesal
margin at apex with few white scales, ventral surface without scales, with ventrolateral
row of about 20 long black setae on distal 0.5 and complete or nearly complete
ventromesal row of short antrorsely curved setae; palpomeres 4 and 5 with small dorsal
and ventral patches of white scales at base, ventral surface of palpomere 4 also with
some white scales just beyond middle, palpomere 5 also with dorsomesal line of white
scales on distal 0.4-0.5; lateral and mesal surfaces of palpomeres 4 and 5 densely
setose. Thorax: Upper proepisternal setae more numerous, 16-26. Wing: Length
3.8-4.0 mm, mean 3.9 mm; length of cell R2/length of vein R2+3 1B-2.4, mean 2.1;
subcosta intersects costa well before furcation of R2+3. Abdomen: Terga without
basolateral patches of white scales, basal pale bands normally 0.35-0.45 tergum length;
tergum II with basomedian triangular pale patch or narrow basal pale band produced
posteriorly in middle; basal bands of terga Ill-VII usually straight, sometimes slightly
produced posteriorly in middle on terga V-VII, bands of terga V-VII and occasionally IV
produced posteriorly along lateral scale-free areas, particularly on terga VI and VII;
tergum VIII (ventral in position ) with broad concave basal whitish band. Sternum VIII
(dorsal in position) variable, similar to tergum VII but basal band sometimes obsolescent
in middle or produced medially to posteriorborder. Genitalia (Fig. 29): Ninth tergal lobe
poorly developed, with 3-8 (mode 6) rather widely spaced setae. Gonocoxite rather long
Harbach: Subgenus Culex in SouthwesternAsia
85
and slender, mesal and ventral setae developed as usual, lateral setae in 3 or 4
longitudinal rows, these shorter in each row toward large ventral setae; subapical lobe
indistinctly divided, setae a-c equally developed, long and tapered with hooked tips; d-e
slender with hooked tips, fflattened with hooked tip; g more or less symmetrical, rather
narrow, rounded apically, borne on distinct prominence; h straight or bent apically,
tapered with narrow pointed distal part. Gonostylus normal; gonostylar claw flat, slightly
broader distally. Phallosome as broad as long with lateral plate and aedeagal sclerite
nearly completely fused into a single simple sclerite, crest of aedeagal sclerite not
distinguished, dorsal arm of lateral plate absent, ventral arm a long slender dorsally
directed and laterally bent sickle-shaped process. Proctiger slightly elongated;
paraproct long and slender, without basal lateral arm, crown of spinelike spicules grading
into short blades and series of bumps laterally. Cereal sclerite with ribbonlike
caudolateral extension joining paraproct near base of crown; 3-6 (mode 4) cereal setae.
Tergum X developed as usual.
Pupa (Fig. 29). Not definitely associated with adults; form and placement of
setae as figured, range and modal number of branches in Table 14; immediately
distinguished from all other species of Culex in the region by the vesiculation of the
scutum, metanotum, and first two abdominal terga. Cephalothorax: Lightly tanned,
scutum and metathorax moderately tanned; middle of scutum behind trumpets and
metanotum covered with distinct vesicles. Setae 1,2,8,9CT frequently double,
sometimes triple; 3,4,6,11 ,12-CT often triple (2-5); 7-CT double. Trumpet: Lightly to
moderately tanned, tracheoid area darker; relatively short and flared, index 3.3-4.5,
mean 3.8; pinna with distinct shallow rounded emargination at tip. Abdomen: Lightly
tanned, terga I-IV darker medially; middle of terga I and II and intersegmental membrane
between terga II-III and III-IV with conspicuous vesicles. Seta 6-I,11short as in laticinctus
and mattinglyi; 7-I,11 usually with 3 or 4 branches, 7-l occasionally double or with 5
branches; 9-11frequently anterior to 7-11;l-III,IV most often with 4 or 5 branches (see
table), l-V-VII variable (see table) but usually double or triple, more often double; 5-IV
usually multiple (l-4) and about length of following tergum, very different in appearance
from 5-V,VI which are usually single and much longer than following tergum; 6-111-V
normally single, seldom double (6-V triple on 1 side in 1 specimen examined), 6-VI most
often triple (l-3). Genital lobe: Lightly tanned; length 0.2-0.3 mm in female, about 0.4
mm in male. Paddle: Very lightly tanned, buttress and midrib darker; midrib with obvious
curvature, inner part not as broad as outer part; length 0.98-l .16 mm, mean 1.07 mm,
width 0.72-l .02 mm, mean 0.88 mm, index 1.1-l .4, mean 1.2. Seta 1-P usually double,
sometimes single; 2-P almost always absent (double on 1 paddle in 1 specimen
examined).
Larva (Fig. 30). Not definitely associated with adults; positions and attributes of
setae as figured, range and modal number of branches in Table 34; easily recognized by
the unusual siphon. Head: Length 0.87-l .07 mm, mean 0.95 mm; width 1.14-l .54 mm,
mean 1.35 mm; moderately tanned, darker posteriorly. Labiogula very nearly
parallel-sided; hypostomal suture rather poorly indicated but complete, extended to
collar from posterior tentorial pit. Dorsomentum more strongly produced anteriorly in
middle than usual; with 12-15 teeth on each side of median tooth, teeth progressively
larger toward sides. Seta 1-C long, slender, tapered, and tanned; 2-C absent; 5,6-C
usually with 7 or 8 branches (see table); 7-C usually with 8 or 9 branches (7-l 1); 8-C
normally double, seldom single; 9-C variable (l-6); 10-C forked, usually bifurcate,
infrequently trifurcate. Antenna: Length 0.53-0.57 mm, mean 0.55 mm; lightly to
moderately tanned, distal part and basal rim darker. Seta 1-A with about 23 aciculate
branches (18-26); 2,3,4-A of equal length, distinctly aciculate. Thorax: Integument
86
Contrib.Amer. Ent. inst.,vol. 24, no. 1, 1988
hyaline; rather sparsely covered with fine spicules. Seta 4-P double or triple; 7-P
double; 8-P most often with 4 branches (3-6). Seta 1-M single; 2-M variable (l-4); 6,7-M
on common tubercle. Seta 1-T very small, usually double (l-3); 2-T long, single.
Abdomen: Integument hyaline; smooth. Seta 3-I-VII single, 3-VII sometimes double; 6-l
usually double, infrequently triple, 6-11not as strong as 6-1, with 2-4 branches occurring in
nearly equal frequencies; 7-l most often triple (2-5), nearly as long as 6-1, 7-11more
strongly developed than in other species, resembling 7-l but shorter, with 3 or 4
branches; 5III-VII and 6-III-VI long and single, 5III-VI distinctly longer than 2 abdominal
segments; 13-111-Vnormally double (see table). Segment VIII: Comb with 25-44 (mode
42) rather short fringed scales. Setae 1,5-VIII distinctly shorter than 2,4-VIII, most often
with 5 or 6 branches. Siphon: Index 3.37-4.24, mean 3.70;strongly inflated, narrowed
distally, with short spine anteriorly near apex; moderately tanned, with dark subapical
band, basal rim heavily tanned. Pecten reduced, beginning well above base of siphon,
composed of 3-5 (Hopkins, 1952 reported 3-6) slender spines with ventral row of short
denticles. Seta 1-S in 4 or 5 pairs, usually 4, lc-S often absent; la,b,c,d-S long and
single (Id-S double on 1 side in 1 specimen examined), 1e-S usually triple (l-4); 1a-S
within pecten, Id-S either within or below dark band, le-S within band. Segment X:
Relatively small; saddle complete, length 0.44-0.51 mm, mean 0.48 mm, siphon/saddle
index 3.76-4.63, mean 4.09. Seta 1-X long and single; 2-X single; 4-X in 6 pairs. Anal
papillae stout, about twice length of saddle, ventral pair shorter than dorsal pair.
Systematics. Culex duttoni is a bizarre species which combines features of the
pipiens and sitiens groups and is recognized here as the name-bearing member of a
separate group. The group also includes the closely related Culex wafti Edwards, 1920.
These are rather large species which are characterized by the indefinitely ringed
proboscis, ringed tarsi, and the usual presence of two to four lower mesepimeral setae in
the adult. The male genitalia are distinctive: the lateral plate is a simple sclerite without a
dorsal arm, the ventral arm is enormously developed, yet reminiscent of the type found
in the pipiens and trifilatussubgroups; and the paraproct has no trace of a basal lateral
arm. The larva resembles members of the pipiens group in most respects, but setae 1-C
and 3-P and the pecten spines approach the condition in the sitiens group and setae
8-P, 7-11, and 1-S are uniquely developed. The distinct form of the larval siphon is
suggestive of that in Culex hutchinsoni Barraud, 1924b of the Oriental Region. The
larva of watti has never been described, but is said to be indistinguishable from duttoni
(Hopkins, 1952). The pupal stage seems to differ from most members of the pipiens
and sitiens groups in having seta 7-I,11usually with 3 or 4 branches, setae 6-111-Vand
5-V,VI usually single, 1-P usually double, and 2-P normally absent. The pupa of waftiis
unknown.
The association of the immature stages with the adults of this species is
presumptive only. No individual rearings are available, but the association seems
correct. It is based on the sympatric occurrence of larvae and adults in the P.D.R. Yemen
and associated larval and pupal exuviae from Ghana. There is a possibility,however, that
the unknown larva and pupa of wattiwere mixed in the material examined.
Bionomics. Culex duttoni is a semidomestic species which uses a wide variety
of breeding places. It has been found in pools in streams, ground pools (including
puddles), and a variety of artificial containers, including drums, pots, sunken barrels,
boats, and wells. It is possible that the dispersal of this species to the Arabian Peninsula
was accomplished by man. The occurrence of this species in southwestern Asia is
based on two larvae and two adults from the P.D.R. Yemen. The adults were captured in
a cave. According to Lewis (1943b), females are never seen biting man. Culex duttoni
is probably not a medically important species.
Harbach: Subgenus Culexin SouthwesternAsia
87
Distribution.
Culex duttoni is widely distributed in the Afrotropical Region. It
has been recorded and described from a single larva collected in the south of Morocco
(Bailly-Choumara, 1968).
Material examined. 85 specimens. Only 5 specimens (1 female, 1 male, 1
male genitalia, and 2 fourth-instar larvae) were examined from southwestern Asia P.D.R. YEMEN: (Mlkhuras, Wadi Gheil el Khumeira). A total of 80 specimens (36
females, 22 males, 5 male genitalia, 9 pupal exuviae, 4 larval exuviae, and 4 fourth-instar
larvae) were examined from ANGOLA, BENIN, CAMEROON, ETHIOPIA, GHANA,
KENYA, MALAWI, NIGERIA (including the type specimens of duttoniand albovirgafus),
SENEGAL, SIERRA LEONE (the type specimens of dissimilisand anarmustus ), SOUTH
AFRICA (the lectotype male of bifoliata ), SUDAN, TANZANIA, UGANDA, ZIMBABWE
(the type specimens of hirsutipalpis), and unknown localities.
Culex (Culex) Miens
W i e d em a n n
sitiens Wiedemann, 1828: 542. Holotype female: Sumatra, Indonesia
(ZMC).
microannulatus
Theobald, 1901 a: 353. +Lectotype male: Quilon,
Travancore, [Madras State], India (BM), designated by Bram, 1967a:
243. Synonymy with sitiens by Edwards, 1913c: 232.
somaliensis
Neveu-Lemaire, 1906: 254.
Syntypes (male; female):
Djibouti, Djibouti (non-extant?). Synonymy with sifiens by Edwards,
1913c: 232.
+Lectotype male, hereby designated,
salus Theobald, 1908b: 256.
mounted on two slides (see Harbach, 1983: 104) bearing following
data: Slide 1 (with maxillary palpi, proboscis, and part of head capsule) - “The / Salt Water. / Mosquito. / TYPE // Culex / salus. /
Theobald / Port Sudan // Recd. from F.V. Theobald. / 1910-396”;
Slide 2 (with genitalia) - “Salt water / mosquito. / TYPE // Culex /
salus / Theobald / Port Sudan. // Recd. fr[om] / F.V. Theo[bald] /
1910-395.” (BM). Synonymy with somaliensis by Edwards, 1911:
261.
For complete synonymy, see Appendix D.
Culex annulus Theobald of Becker, 1907( 1910): 140 (Socotra, A).
Culex impel/ens Walker of Giles, 1901 a: 606 (India, A’, L’, distr.); Theobald, 1901 a: 362
(India, F*); Christophers, 1906: 8 (? India, L*).
Culex sitiens Wiedemann of Edwards, 1913c: 232 (Oriental Region, syn., tax., A);
Edwards, 1922: 277, PI. VI (Oriental Region, A key, M gen.‘); Barraud, 1924a:
993 (India, A key, syn., M’, F, distr.); Barraud, 1924c: 427 (India, L*); Lewis, 1945:
15 (Sudan, L, distr.); Leeson and Theodor, 1948: 227 (Socotra, A, L keys, L, L
bionomics, toll. sites).
Culex (Culex) sitiens Wiedemann of Barraud, 1934: 398 (India, A, L keys, M*, F, L’, L
habitat, distr., syn.); Edwards, 1941: 296, 417, 482 (Afrotropical Region, A key,
M*, F, P, syn., distr.); Hopkins, 1952: 284 (Afrotropical Region, L key, L,
bionomics); Knight, 1953a: 231 (Yemen Arab Republic, A, L, distr., L habitat);
Lewis, 1956: 707 (Sudan, L, toll. rec., distr.); Mattingly and Knight, 1956:
103-137 (Arabian Peninsula, L*, A, L keys, distr., bionomics); Ovazza et al., 1956:
167 (Ethiopia, M gen.*); Lotfi, 1970: 401 (Iran, toll. rec.); Aslamkhan, 1971: 155
88
Contrib.Amer. Ent. Inst.,vol. 24, no. 1, 1988
(Pakistan); Lotfi, 1973: 206 (Iran, toll. rec.); Lotfi, 1976: 72, 75, 81 (Iran, L key,
ecol. note, L*); Sirivanakarn, 1976: 95 (Oriental Region, A, P, L keys, M*, F*, P*,
L*, tax., distr., bionomics); Harbach, 1985a: 86, 97, 102 (SW Asia, distr., A, L
keys).
Culex somaliensis Neveu-Lemaire of Edwards, 1911: 261 (Africa, A, key); Edwards,
1912~: 28 (Africa, A key).
Culex thalassius Theobald of Leeson and Theodor, 1948: 228 (in part; Socotra, L key).
Adult. A medium-sized species distinguished from all other Culex occurring in
southwestern Asia by having speckled femora and base of cell R2 distal to base of cell
Ml.
FEMALE. Head: Antenna blackish; pedicel yellow, mesal surface dark brown to
black with tiny setae and scales; length 1.7-2.2 mm, mean 1.9 mm. Proboscis dark, with
distinct median whitish ring about 0.2 proboscis length, extending from 0.4 to 0.6 from
base; frequently with scattered pale scales proximal to ring, particularly ventrally;
sometimes also with scattered pale scales faintly evident distal to ring; with indefinite
narrow pale fringe of scales at apex of prementum; length 1.8-2.1 mm, mean 1.9 mm.
Maxillary palpus dark with pale scales laterally at apex, sometimes also with pale scales
laterally at base or on entire lateral surface; palpomeres 2 and 3 with 3-5 long lateral
setae; length about 0.6 mm, approximately 0.3 proboscis length. Forked scales of
vertex pale yellow to yellowish brown, darker posterolaterally; falcate scales pale yellow
to brownish, darker laterally; interocular scales yellowish white and projecting ventrally
between eyes; lateral spatulate scales yellowish white. Thorax: Integument dark brown
to black. Scutum mottled with variable spots and blotches of coarse dark and pale
scales; dark brown to golden-brown scales usually predominant with pale yellow to
golden scales on anterior and lateral margins, across middle and on prescutellar area;
acrostichal and dorsocentral setae short, slender and indistinct; supraalar setae robust
and dense. Scutellum with rather numerous pale yellow falcate scales; lateral lobes each
with 4 or 5 strongly developed setae, 6-9 weaker ones; median lobe with 5-8 large
setae, 9-l 2 smaller ones. Antepronotum with coarse yellowish falcate scales, often with
some brownish ones in middle; postpronotum with dark brown or golden-brown scales
in middle, pale yellow scales anteriorly and posteriorly, posterior scales coarser.
Proepisternum with elongate subfusiform scales laterally below upper proepisternal
setae; shorter apically rounded pale yellow spatulate scales among and extending mesal
to setae. Mesopleuron with white spatulate scales: distinct upper and lower patches on
katepisternum, distinct anterior patch on anepimeron, and few scales usually present
among upper mesepimeral setae. Numbers of pleural setae: 6-8 upper proepisternal,
9-17 prealar, 4-8 upper mesokatepisternal, 5-l 0 lower mesokatepisternal, 7-l 2 upper
mesepimeral, no lower mesepimeral. Wing: Length 3.2-4.0 mm, mean 3.7 mm; length of
cell R2 2.1-2.5 length of vein R2+3, mean 2.3; length of cell Ml/length of cell R2 about
0.9; furcation of vein R2+3 normally distal to furcation of vein Ml+2; mainly dark-scaled,
with pale spatulate scales along posterior margin of costa (except at apex), posterior
margin of approximately distal 0.5 of subcosta, and anterior margin of Rl; usually with
small spot of pale scales at base of remigium. HaIter: Pedicel and scabellum pale;
capitellum dark, with white scales. Legs: Anterior surface of forecoxa white-scaled, with
few pale scales posterolaterally at apex; midcoxa with broad longitudinal patch of white
scales on anterior side of lateral midline, extending onto anterior surface distally;
hindcoxa with slightly curved lateral longitudinal line of pale scales. Trochanters with
white spatulate scales on ventral and posterior surfaces; mid- and hindtrochanters also
with white scales on dorsal surface. Femora with whitish knee spots; anterior surface of
Harbach: SubgenusCulex in SouthwesternAsia
89
forefemur predominantly dark-scaled with more or less definite whitish stripe on proximal
0.5, distal 0.5 speckled with pale scales, posterior surface whitish-scaled; anterior
surface of midfemur dark-scaled exept at base, speckled with pale scales, posterior
surface whitish-scaled; hindfemur primarily whitish-scaled, with narrow band of dark
scales at apex, sometimes with indistinct dorsal dark stripe extending proximad from
band, frequently with scattered dark scales on anterior surface. Dorsal surface of
foretibia dark-scaled with indistinct spot of pale scales at apex, pale-scaled ventrally;
anteroventral surface of midtibia with dark scales confluent with patch of dark scales on
0.25 to 0.50 of dorsal surface except at extreme base, with spot of pale scales dorsally at
apex; hindtibia mainly dark-scaled dorsally, with dark scales dissipated in middle and
extended over sides at ends,with anterodorsal spot of pale scales at apex. Tarsi mainly
dark-scaled; tarsomere 1 pale-scaled ventrally; joints with narrow whitish rings mainly on
bases of tarsomeres, rings distinct between tarsomeres l-2 and 2-3, often indistinct
between 3-4 and 4-5, tarsomere 1 with narrow pale ring at base; hindtarsomere 2 faintly
pale beneath. Abdomen: Tergum I with median posterior patch of scales, scales usually
dark anteriorly and pale posteriorly; terga II-VIII with basal white bands continuous with
elongate basolateral spots of same color, band of tergum II produced in middle, others of
even width (0.2-0.4 length of tergum); terga II-VIII also with very narrow white bands on
posterior margin, that of tergum VII often strongly produced in middle and occasionally
united with basal band. Sternum I bare; sternum II usually somewhat sparsely clothed in
white scales, particularly on median and lateral areas; sterna Ill-VII white-scaled with
posterolateral dark spots, median area of posterior margin with yellowish scales; sternum
VIII with lateral patches of white scales, median area void of scales.
MALE. Like female except as follows. Head: Proboscis with ventral cluster of
setae at base of pale ring; false joint about 0.55 from base of proboscis. Length of
maxillary palpus about 2.1-2.8 mm, mean 2.6 mm, extending beyond tip of proboscis by
length of palpomere 5; integument of palpomere 2 pale apically, with pale scales on
dorsal and lateral surfaces; palpomere 3 with broad median pale ring comprised of
whitish scales on dorsal and lateral surfaces and pale bare integument on ventral and
mesal surfaces, ventromesal margin with row of about 25 short apically curved and
flattened setae on distal 0.8, ventrolateral margin beyond median pale ring with about 15
long setae; palpomeres 4 and 5 with basal pale rings, densely setose; palpomere 5 with
apical 0.5 or less (usually 0.33) pale-scaled. Thorax: Upper proepisternal setae more
numerous, 9-12. Wing: Pale scales usually absent at base of remigium. Abdomen:
Terga without basolateral pale spots; basal bands of terga V-VII produced posteriorly
along lateral scale-free areas; tergum VII (and occasionally VI) with large triangular
posteromedian pale spot, spot often connected medially to basal band; tergum VIII
(ventral in position) with broad basal pale band. Sterna I and II same as in female; sterna
Ill-VII with narrow posterior band of white scales, rather broad subapical dark band and
broad basal pale band, dark band of more posterior sterna often divided in middle by
narrow longitudinal line of pale scales connecting basal and apical bands; sternum VIII
(dorsal in position) like tergum VII. Genitalia (Fig. 31). Ninth tergal lobe poorly
developed, narrowly connected with its mate of the opposite side, with line of 2-l 0
(mode 6) short setae. Gonocoxite normal, lateral area with 3 or 4 rows of moderately
developed setae extending from above base to point just above subapical lobe;
subapical lobe undivided, setae a-c rodlike as usual, a shorter than b and c and blunt at
tip, b and c hooked at tip; setae d-e with small barblike denticle before recurved tip, f
hooked at tip; seta g a simple, symmetrical, apically rounded leaf; h about length of g,
flattened, bent distally. Gonostylus rather stout basally, distinctly tapered apically,
slightly rugose dorsally before tip; gonostylar claw short, broadened apically, troughlike.
90
Contrib.Amer. Ent. inst.,vol. 24, no. 1, 1988
Phallosome not much longer than broad, aedeagal sclerite shorter than lateral plate;
lateral plate with definite inner and outer divisions, inner division (= ventral arm) a broad
minutely spiculate lobe borne on ventromesal margin of outer division (= lateral arm),
ventrocaudal angle rounded, dorsocaudal angle rather pointed, dorsal margin with 3
prominent caudally directed teeth before apex; outer division rather simple,
posteromesal margin with a conspicuous laterally curved tooth which appears broad in
lateral view, ventrocaudal angle giving rise to a heavy flattened laterally directed and
dorsally curved pointed process, dorsal process a prominent dorsolaterally projecting
earlike lobe; dorsal arm absent (unless represented by the posteromesal tooth of the
outer division). Proctiger unmodified; paraproct with moderately long, gently curved
basal lateral arm and distinct ventral acetabulum. Cereal sclerite more or less rectangular;
2-5 (mode 3) cereal setae present. Tergum X developed as usual.
Pupa (Fig. 31). Character and positions of setae as figured, range and modal
number of branches in Table 15; without obvious distinctive features, most closely
resembling mimeticus from which it can usually be separated by the larger number of
branches of setae l-II, 6-III-VI, and 9-VIII. Cephalothorax: Lightly tanned, legs and
metathorax darker. Setae l-9,1 1,12-CT usually double or triple (see table); 1-CT longer
and slightly stronger than 2,3-CT; lo-CT frequently with 7 or 8 branches (59), distinctly
metathorax darker. Setae l-9,1 1,12-CT usually double or triple (see table); l-CT longer
and slightly stronger than 2,3-CT; lo-CT frequently with 7 or 8 branches (5-g), distinctly
longer than 11 ,12-CT. Trumpet: Moderately tanned, tracheoid area only slightly darker;
index 4.1-5.7, mean 4.7; pinna long, 0.3-0.4 trumpet length. Abdomen: Lightly
tanned, anterior terga darker; length 3.1-3.7 mm, mean 3.5 mm. Seta 7-l triple, 7-11
double or triple, more often double; l-11 with 4-10 aciculate branches; l-Ill-VI moderately
developed, with fewer branches on each successive posterior tergum, l-111usually with 5
or 6 branches (5-8), l-IV normally with 4 or 5 branches (3-5), 1-V with 3 or 4 branches,
l-VI double or triple; 6-III-VI single or double, more often double, 6-V,VI larger than
6-lll,lV; 5-IV-VI equally developed, usually double, longer than following tergum; 9-VIII
commonly with 1O-l 2 branches (9-l 4). Genital lobe: Lightly tanned; length about 0.20
mm in female, about 0.35 mm in male. Paddle: Lightly tanned, buttress and midrib
darker; length 0.90-1.00 mm, mean 0.94 mm, width 0.68-0.80 mm, mean 0.74 mm,
index about 1.3.
Larva (Fig. 32). Form and placement of setae as figured, range and modal
number of branches in Table 35; bearing a strong resemblance to theileri, laticinctus,
and mattinglyi but immediately distinguished by the form of seta 1-C and the pecten
spines. Head: Length 0.72-0.88 mm, mean 0.80 mm; width 1.13-1.35 mm, mean 1.24
mm; lightly tanned, labiogula and posterior areas of lateralia and dorsal apotome darker.
Hypostomal suture strongly indicated, diverging posteriorly from its mate of the opposite
side, ending about halfway between posterior tentorial pit and collar. Dorsomentum with
7 or 8 teeth on either side of center. Seta 1-C short, thick, usually spiculate at sides,
dark; 2-C present or absent; 5-C usually with 6 or 7 branches (6-8); 6-C often with 4,5 or
6 branches (4-7); 7-C frequently with 9 or 10 branches (7-l 1); lo,1 1 ,12,15-C double or
triple, lo,1 1-C occasionally with 4 branches; 13-C normally double (1-3). Antenna:
Length 0.41-0.56 mm, mean 0.49 mm; lightly tanned, distal part darker, with dark ring at
base; proximal part with aciculae on dorsal and lateral surfaces, distal part with few
spicules laterally. Seta 1-A with about 27 branches (22-32); 2,3-A subapical. Thorax:
Integument hyaline, smooth. Seta 4-P double; 7-P double or triple; 8-P single or
double, distinctly shorter than 4,7-P. Seta 1-M single, about twice length of 2-M; 4-M
single or double. Seta 1-T single, nearly as long as 2-T which is usually single,
sometimes double; 13-T distinctly longer than 12-T, with 2-5 branches. Abdomen:
Harbach: Subgenus Culex in SouthwesternAsia
91
Integument like that of thorax. Seta 3-l usually single, seldom double, 3-II-VI single, 3-VII
usually double (l-4); 6-I,11usually triple (2-4), 6-III-VI normally double; 7-l usually long and
single, about as long as 6-1, seldom double; l-Ill-VII rather short, about 0.5 length of
segment, double or triple, more often double; 13-Ill-V,Vll developed like l-III-VII, usually
double (see table). Segment VIII: Comb with 30-54 (mean 42) rather short, fringed
scales. Siphon: Index 3.61-4.55, mean 3.80; lightly tanned, basal ring and acus darker.
Pecten composed of 9-17 (mode 13) spines with complete ventral row of denticles,
larger spines with 7-13 (mode 10) denticles, tendency for most basal spines to fuse.
Seta 1-S in 6, 6.5 or 7 pairs, 1 distal pair (1 e-S) borne laterally, others imperfectly paired
near posterior midline, proximal 3 or 4 pairs often with 6 or 7 branches and about twice as
long as diameter of siphon at point of attachment. Segment X: Saddle complete; lightly
tanned; posterodorsal margin with distinct short spicules; length 0.29-0.35 mm, mean
0.33 mm, siphon/saddle index 3.03-4.29, mean 3.71. Seta 1-X single; 2-X usually with 4
or 5 branches, occasionally triple; 4-X usually in 6 pairs. Anal papillae short, globose.
Systematics. Culex sitiens is the nominotypical member of a large and variable
group of species which probably should be divided into several different groups. The
group is widespread in the warmer parts of the Old World with species occurring in the
Afrotropical, Oriental, and Australian regions, and the Mediterranean and Manchurian
subregions of the Palaearctic. The group is currently subdivided into six subgroups
(Sirivanakarn,
1976).
Six members belonging to the sitiens, mimeticus,
bitaeniorhynchus, and vishnui subgroups occur in southwestern Asia. The sitiens
subgroup is represented by sitiens and poicilipes.
Culex sitiens is a highly variable species which breeds in brackish coastal waters
from East Africa to the South Pacific. It is very likely that this nominal form is actually
composed of several distinct species. There is considerable variation in all stages and
there seems to be many isolated populations. Adults from southwestern Asia have
much lighter scutal scales and more extensive pale scaling on the wings than specimens
from the Oriental Region. There is an extreme range of variation in larval characters,
especially in the size and shape of seta 1-C and the siphon, but there are no apparent
differences in the form of the male genitalia.
There should be no difficulty in identifying specimens of sitiens in southwestern
Asia unless the very similar Culex thalassius Theobald, 1903a also occurs in the
Afrotropical part of the Arabian Peninsula. Adults of this species differ in having darker
scutal scales and no speckling of pale scales on the femora. The male genitalia differ
slightly in details of the phallosome. The larva of thalassius at present cannot be
distinguished with certainty from that of sitiens. The record of thalassius from Syria
(Abdel-Malek, 1960) cannot be correct, and probably is based on misidentified larvae of
tritaeniorhynchus.
Bionomics. Culex sitiens breeds primarily in coastal brackish ground pools,
mangrove swamps, and salt marshes, but is also found in artificial containers and
collections of fresh water not far from the seacoast. Specimens from the Yemen Arab
Republic have been collected in open cement basins some distance inland and in
brackish water seepage pools near the seashore (Knight, 1953a). Females feed mainly
on birds and pigs (Colless, 1959), but are also known to bite cattle, dogs, and man.
Culex sitiens is a potential vector of human pathogens. Experimental transmission of
Japanese encephalitis virus has been demonstrated in the laboratory (Hodes, 1946, as
jepsoniTheobald) and it has been found naturally infected with larvae of Brugia malayi in
Thailand (lyengar, 1953).
Distribution.
This species occurs in the coastal lands of the Oriental Region,
southwestern Asia, eastern Africa, and Madagascar. Populations are also known from
92
Contrib.Amer. Ent. Inst.,vol. 24, no. 1, 1988
the Ryukyu Archipelago, Korea, northern Australia, and many islands in the South
Pacific.
Material examined. 544 specimens. A total of 455 specimens (90 females,
85 males, 31 male genitalia, 105 pupal exuviae, 105 larval exuviae, and 39 fourth-instar
larvae) were examined from coastal areas of southwestern Asia - OMAN: (Dhofar,
Itelba, Salalah, Trucia); P.D.R. YEMEN: (Al Ais, Buaish, Dis Town, Geregeri, Gheil
Bawazir, Hami, Hamial Sharq, Harshiyab, Khon, Makhzan, Meifa, Mukalla, Qaiti, Shihr,
Sukhal al Dis, Thrilla, Urfat Subai, Wadi Maseila, Wasira, unknown localities); SAUDI
ARABIA: (Jeddah); UNITED ARAB EMIRATES: (Sharja); YEMEN ARAB REPUBLIC:
(Hodeidah). An additional 89 specimens (40 females, 15 males, 4 male genitalia, 10
pupal exuviae, 4 larval exuviae, and 16 fourth-instar larvae) were examined from
DJIBOUTI, INDIA (the type specimens of microannulatus ), SOCOTRA, SUDAN
(including the type specimens of salus ), and unknown localities. An unknown number
of specimens from Southeast Asia and the type specimens of impel/ens, gnophodes,
and jepsoni were also examined.
Colex (Culex)
polcilipes
(Theobald)
poicilipes Theobald, 1903b: ix (Lasioconops ). +Holotype female:
Bonny, [Eastern Provinces], Nigeria (BM).
quasigelidus
Theobald, 1903a: 181. +Lectotype female, hereby
designated, bearing following data: “Type // Entebbe / Dr. Row //
Culex / quasigelidus / (Type) Theobald // Entebbe. / bred from /
larvae. / Sep. 20. 02”; left wing mounted on slide (BM). Synonymy
with poicilipes by Edwards, 1932a: 203; incorrectly considered as
senior synonym of poicilipes by Edwards, 1911:258.
Giles, 1904: 369. +Holotype female: Benguela,
taeniorhynchoides
Angola (BM). Synonymy with quasigelidus by Edwards, 1911: 258.
maculipes Theobald, 1904: 79 ( Taeniorhynchus tenax var.). +Holotype
female: Kenissa, White Nile and Middle Sobat, Sudan (BM).
Synonymy with quasigelidus by Edwards, 1911: 258.
madagascariensis
Ventrillon, 1905b: 427 (Pseudoheptaphlebomyia).
+Lectotype female, hereby designated, bearing following data: “9 //
MUSEUM PARIS / [Tananarive,] MADAGASCAR / VENTRILLON
1905 N Pseudohepta- / phlegomyia [sic] / madagasca- / riensis Ventr.
/ type.” (MNHP). Synonymy with quasigelidus by Edwards, 1911:
258.
par Newstead, 1907 (in Newstead et al., 1907: 25). +Holotype female:
Tshumbiri, [Leopoldville], Belgian Congo (BM). Synonymy with
quasigelidus by Edwards, 1912b: 29.
punctipes
Theobald, 1907: 316 (Aporoculex ). +Holotype female:
Chinde, [Zambezia, Mozambique] (BM). Synonymy with quasigelidus by Edwards, 1911: 258.
auritaenia Enderlein, 1920: 49. +Lectotype female, hereby designated,
bearing following data: “Madagaskar / Tanararivo [sic] / Friederichs
S.V. // Type // Culex / auritaenia / Type Enderl. [female symbol] / Dr.
Enderlein det. 1920 // 1920. 375.” (BM). Synonymy with poicilipes
by Edwards, 1932a: 203.
Harbach: Subgenus Culex in SouthwesternAsia
93
Culex poicilipes (Theobald) of Lewis, 1945: 14 (Sudan, dish.); Theodor, 1952: 113
(Egypt, zoogeogr.); Senevet et al., 1957a: 86 (North Africa, L*); El-Said and
Kenawy, 1983a (Egypt, dish-.).
Culex (Culex) poicilipes (Theobald) of Edwards, 1941: 289, 415, 482 (Afrotropical
Region, A key, M*, F*, P*, syn., distr.); Lewis, 1943a: 281 (Eritrea, toll. rec.);
Lewis, 1943b: 68 (Sudan); Abbott, 1948: 44 (Sudan, distr.); Lewis, 1948:
144-148 (Sudan, A, L bionomics); Hopkins, 1952: 280 (Afrotropical Region, L
key, L*, bionomics); Mattingly, 1954: 58 (Africa, zoogeogr.); Lewis, 1956: 705
(Sudan, distr., bionomics); Ovazza et al., 1956: 167 (Ethiopia, toll. sites);
Senevet et al., 1957b: 92 (North Africa, L); Senevet and Andarelli, 1959: 135
(Egypt, A, L, P keys, syn., M’, F*, P’, L*, distr., biol., L assoc.); DuBose and
Curtin, 1965: 351-354 (Mediterranean area, A, L keys); Mekuria, 1968: 78
(Ethiopia, distr.); El-Said and Kenawy, 1983b (Egypt, toll. rec., distr.); Harbach,
1985a: 86, 96, 107 (Egypt, Israel, A, L keys.).
Culex quasigelidus Theobald of Wesche, 1910: 38 (West Africa, P*, L’, keys); Edwards,
1912c: 381 (Africa, L, key); Gough, 1914: 135 (Egypt, toll. rec.); Storey,
1918(1919): 86-98 (Egypt, A, L keys, A); Ingram and Macfie, 1919: 66 (W. Africa,
P*, L); Kirkpatrick, 1924( 1925): 367, 371 (Egypt, A, L keys); Scott, 1927: 88
(Ethiopia, toll. rec.); Abdel-Malek, 1956: 105 (Sinai, L key).
Culex (Culex) quasigelidus Theobald of Edwards, 1921: 337 (Egypt, A, L keys, tax.,
bionomic note, distr.); Seguy, 1924: 29, 185 (Egypt, A key, A, L, tax., distr.,
syn.); Kirkpatrick, 1925: 107, 186 (Egypt, syn., M*, F, P’, L*, distr., bionomics);
Edwards, 1926: 129 (Egypt, A, L keys, A, L, distr.); Stackelberg, 1927: 152
(Egypt, M, F keys, A, M gen.*, distr.); Martini, 1931: 380 (Egypt, A, L keys, M*, F,
L, distr.).
Adult.
A rather large species easily distinguished from other species of the
subgenus Culex by the presence of rows of pale spots on the femora and tibiae. It bears
a superficial resemblance to tigripes De Grandpre and De Charmoy of the subgenus
Lufzia which also has spotted legs, but these species do not occur together in
southwestern Asia and are distinguished without difficulty by the characters given in the
key to subgenera.
FEMALE. Head: Antenna dark; pedicel with small patch of pale scales on mesal
surface; length 1.5-l .9 mm, mean 1.7 mm. Proboscis dark-scaled with sharply defined
median pale ring extending from 0.4 to 0.6 from base; length 1.8-2.1 mm, mean 1.9 mm.
Maxillary palpus dark-scaled, with pale scales at apex and also usually on dorsal surface
of palpomere 2; length 0.4-0.5 mm, about 0.25 proboscis length. Forked scales of
vertex dark; narrow falcate scales yellowish and yellowish brown; postocular and
postgenal areas with white spatulate scales, these often divided into dorsal and ventral
patches by anterior extension of posterior black scales, white patches usually connected along margin of eye. Thorax: Integument usually black, sometimes dark brown.
Scutum mainly with dark brown scales, with variable amount of whitish or yellowish to
yellowish-brown scaling, principal areas of pale scaling include median area of anterior
0.33, paired sublateral areas at middle, and transverse band behind middle; small but
conspicuous patch of long white near-linear scales on antealar area; some whitish or pale
yellow scales around bare part of prescutellar area. Scutellum largely covered with
yellowish-brown falcate scales. Ante- and postpronotawith golden-brown falcate scales,
antepronotum with paler scales dorsally and/or ventrally, postpronotum with few slightly
lighter scales at extreme ventral angle. Proepisternum with lower patch of pale scales in
addition to usual upper patch, upper patch rather large, extending ventromesally on
94
Contrib.Amer. Ent. Inst.,vol. 24, no. 1, 1988
anterior surface; with 9-13 setae in upper area. Mesopleuron with usual patches of white
spatulate scales on katepisternum and epimeron; with numbers of setae as follows:
8-14 prealar, 3-8 upper mesokatepisternal, 6-l 2 lower mesokatepisternal, 6-l 0 upper
mesepimeral, and no lower mesepimeral. Wing: Length 3.4-4.6 mm, mean 4.0 mm;
length of cell R2 3.6-4.7 length of vein R2+3, mean 4.0; length of cell Ml/length of cell
R2 0.75-0.85, mean 0.81; mainly brown-scaled, with variable pale scaling; pale scaling
sometimes absent or indistinct, sometimes limited to patches at base and apex of costa,
but usually more extensive (particularly in dark individuals) as follows: patch at base of
costa before humeral crossvein; scattered along posterior margins of costa and
subcosta; line on posterior margin of costa distal to intersection of subcosta; scattered
along anterior margin of Rl before level of furcation of R2+3; lines on anterior and
posterior margins of Rl beyond furcation of R2+3. Halter: Pedicel and scabellum yellow;
capitellum dark. Legs (midleg, Fig. 2J): Anterior surface of forecoxa with mixture of dark
and pale scales, sometimes almost completely pale-scaled, often nearly all dark-scaled;
anterolateral surface of midcoxa with mixture of dark and pale scales; hindcoxa with
lateral line of pale scales. Femora with pale scales encircling base and covering posterior
surface; anterior surfaces dark brown-scaled with row of 7-9 yellowish-white spots, spots
fewer and larger on hindfemur; with narrow pale knee spots. Tibiae like femora; pale
spots more distinct on fore- and midtibiae than on hindtibia; spots slightly dorsal on
foretibia and in more or less double row; with pale scales at apex. Tarsi with pale rings on
base and apex of tarsomeres l-3 and base of tarsomere 4, basal rings broader.
Abdomen: Terga II-VII mainly dark brown-scaledwith variable basal pale bands and lateral
pale spots; basal bands of terga II and III produced medially; most of middorsal surface
pale on terga VI and VII; basal bands continuous with lateral spots except perhaps on
more proximal terga; lateral spots progressively larger from anterior to posterior terga.
Sterna largely pale-scaled; often with indistinct V-shaped patch of dark scales, base of V
at middle of anterior margin.
MALE. Same as female except as follows. Head: Proboscispaler; with many pale
scales mixed with dark ones; mostly pale beneath; often pale at sides before labella.
Maxillary palpus with 6 pale areas as follows: (1) lateral pale area at base of palpus without
scales, (2) narrow pale ring across junction of palpomeres 2 and 3 without scales, (3)
broad dorsolateral pale area at middle of palpomere 3 with pale scales, (4) pale area at
apex of palpomere 3 and base of palpomere 4, latter with pale scales dorsally and
ventrally, (5) pale area at apex of palpomere 4 and base of palpomere 5, latter with small
patch of pale scales dorsally and ventrally; (6) pale area on approximately distal 0.5 of
palpomere 5 with line of pale scales on dorsal and ventral surfaces; palpomeres 4 and 5
often with scattered pale scales on ventral surface; length 2.6-3.3 mm, mean 3.0 mm.
Forked scales of vertex pale medially, becoming dark laterally and posteriorly; falcate
scales paler, whitish to yellow; spatulate scales becoming yellowish ventrally. Thorax:
Upper proepisternal scales numerous, about 35. Scutal scales smaller, more numerous
and paler, those of fossae largely golden yellow; band behind middle pale yellow.
Scutellar scales narrower and paler. Pleural scale-patches generally paler. Wing:
Length 3.1-4.2 mm, mean 3.7 mm; length of cell R2/length of vein R2+3
variable, 2.1-4.1, mean 2.7; pale scaling less distinct, wing paler overall; pale scaling
reduced along posterior margin of Rl. Abdomen: Terga II and Ill with medial
extension of basal band often reaching posterior margin; terga VI and VII sometimes
entirely pale-scaled. Genitalia (Fig. 33): Form as figured; lateral plate of phallosome
somewhat as in sitiens. Ninth tergal lobe scarcely produced, with row of 5-9 (mode 8)
irregularly spaced setae. Gonocoxite rather elongate, lateral setae extending from near
base to apex, in 2 or 3 rows increasing to about 5 rows at level of subapical lobe; subapi-
Harbach: Subgenus Culex in SouthwesternAsia
95
cal lobe undivided, not very prominent; setae a-c slender, about same length, a blunt
at tip, b and c hooked at tip; only 2 simple setae in group d-f, one acute, the other (f?)
slightly flattened and hooked at tip; g and h essentially as in sitiens; a noteworthy
auxiliary seta (aux) present basally on proximal side of lobe. Gonostylus not unusual.
Phallosome longer than broad; aedeagal sclerite slightly longer and narrower than lateral
plate, ventral margin deeply excavated in middle; inner division of lateral plate closely
fused to ventromesal margin of outer division, shape much as in sitiens but with 3-5
more massive teeth arising between dorsocaudal angle and mesal surface of outer
division, teeth resemble a hand with stubby fingers directed dorsolaterally, dorsomesal
margin with tiny bumps and spicules; outer division with large concave-convex flap
arising from dorsocaudal margin, dorsal process and basal articulatory process poorly
developed. Proctiger large; paraproct long and slender, without basal lateral arm, crown
a large mass of short spinelike spicules. Cereal sclerite elongate, somewhat triangular;
2-6 (mode 4) cereal setae. Tergum X small.
Pupa (Fig. 33). Form and positions of setae as figured, range and modal number
of branches in Table 36; usually easily recognized by the presence of a darkened area
on the paddle (not illustrated). Cephalothorax: Setae 1,3,8-CT usually with 3 or 4
branches (see table); 2-CT usually with 4 or 5 branches (3-5); 4,6-CT usually triple (2-4);
5-CT usually with 4 or 5 branches (4-6), not much longer than 4-CT; 7,ll -CT double or
triple, more often double; 1O-CT multiple (7-16); 12-CT with 3-6 branches occurring in
nearly equal frequencies, longer than lo,1 l-CT.
Trumpet: Lightly to moderately
tanned, tracheoid area and pinna darker; index 5.7-7.5, mean 6.8; pinna short, 0.1-0.2
length of trumpet. Abdomen: Length 3.1-3.6 mm, mean 3.4 mm; caudolateral angle of
terga VII and VIII acute. Seta 7-l double or triple, 7-11usually double, sometimes single or
triple; l-Ill-V strongly developed, multiple, about length of following tergum; 6-III-VI
moderately developed, usually with 4 or 5 branches (3-6), 6-VI distinctly longer than the
others; 5-IV usually triple (3-5), about length of l-IV, shorter than 5-V,VI which are
double and considerably longer than following terga. Paddle: Lightly tanned, with
darkened area on distal 0.5 or less and mainly on inner part; length 0.91-l .16 mm, mean
1.00 mm, width 0.68-0.82 mm, mean 0.74 mm, index 1.2-1.5, mean 1.4.
Larva (Fig. 34). Character and placement of setae as figured, range and modal
number of branches in Table 36; recognized at once by the shape of the siphon, and
black pigmentation of posterior part of thorax and abdominal segments III and V which
give the larva a 3-banded appearance in life. Head: Length 0.82-0.95 mm, mean 0.89
mm; mainly lightly tanned, darker posteriorly, ventral part of lateralia adjacent to
hypostomal suture sometimes with dark triangular spot. Median labral plate noticeably
produced forward at point of insertion of seta 1-C. Dorsomentum usually with 6 teeth
(5-7) on either side of middle tooth. Seta 1-C short, stout, pointed, dark; 2-C
occasionally present; 4,5,6-C inserted more posterior than usual, 5,6-C only extending
to near base of 1-C; 4-C short, usually double (l-3); 5-C most often with 4 branches
(3-5); 6-C distinctly shorter than 5-C, often with 4 branches (4-6); 7-C strongly
developed, usually with 10, 11 or 12 branches (10-15); 8,9,1 O-C generally with more
branches than usual, 1O-C usually with 4 or 5 branches (3-5); 11-C double, seldom triple;
14-C rather spiniform, normally single (1,2). Antenna: Length 0.76-0.86 mm, mean
0.80 mm; tanning and spiculation as usual; scape developed. Seta 1-A large, with
about 34 branches (29-37); 2,3-A inserted near apex. Thorax: Integument hyaline,
smooth; with concentration of black pigment granules under dorsal surface of meso- and
metathorax. Setae 4,8-P double, about same length; 7-P with 3 or 4 branches, perhaps
slightly longer than 4,8-P. Setae 1,4-M similar, both with more branches than usual, 1-M
often with 5 branches (5-8), 4-M usually with 4 or 5 branches (4-6). Seta 1-T multiple
96
Contrib.Amer. Ent. Inst.,vol. 24, 170.1, 1988
about 0.5 length of 1-T; 2-T also multiple, most often with 4 branches (3-5); 12-T double
or triple; 13-T with more branches than usual (12-23) not much longer than 12-T.
Abdomen: Integument hyaline, smooth; dark pigment granules present, particularly
noticeable in segments III and V (as in antenna&s and Sinai&s). Seta 3-l multiple (2-7)
infrequently double or triple, 3-II-IV usually with 3 or 4 branches (see table), 3-V,VI
double or triple, 3-VII highly branched (7-10); 6-I-VI normally triple, some occasionally
with 4 branches, 6-V usually distinctly shorter than the others; 7-l usually double (1,2)
but as long as 6-1, 7-II-VII generally with more branches than usual (see table); l-Ill-VII
usually with 7 or 8 branches (see table). Segmenf VI//: Comb consisting of 6-l 0 (mode
7) large spinelike scales, fringed on either side at base. Seta 2-VIII double or triple; 5VIII
generally with more branches than usual (5-7). Siphon: Index 3.59-4.66, mean 4.22;
moderately tanned; slightly but distinctly bent anteriorly, posterior side longer than
anterior side; spiracular lobes particularly large. Pecten composed of 8-14 (mode 11)
short spines with 4-9 (mode 5) basal denticles, most basal denticles tend to be fused.
Seta 1-S in 5 pairs, middle pair borne laterally, others on posterior midline, length of latter
more than twice diameter of siphon at point of attachment. Segment X: Saddle
complete, rather long, length 0.43-0.52 mm, mean 0.48 mm, siphon/saddle index
2.54-3.07, mean 2.78. Seta 1-X usually with 4 branches (2-5); 2-X often with 5 branches
(3-7); 4-X in 6 or 7 pairs, branches numerous. Length of anal papillae variable, ventral
pair usually, but not always, longer than dorsal pair.
Systematics.
Culex poicilipes is widely distributed south of the Sahara and
along the Nile River into Egypt. The species occurs in the Sinai Peninsula and the
Negev of Israel where it probably spread from the Nile Delta in relatively recent times.
The species is strikingly distinct in all stages and cannot be confused with any other
species of Culexoccurring in Egypt or Israel.
This is one of the most variable species of Culex in Africa. Remarkable variation
occurs in the ornamentation of the adults and the chaetotaxy of the larva. Much of this
variation is undoubtedly individual, but there are indications that some populations differ
constantly from others. A noticeably darker adult form with less extensive pale markings
is common in some areas. Whether this is due to genetic of environmental factors
cannot be determined without sufficient individually reared material collected
throughout the recorded range of the taxon. The adult and immature stages of
poicilipes in Egypt and Israel are not so variable and it is unlikely that more than one
species is involved in these countries.
The placement of poicilipes in the sitiens subgroup is provisional. It is included
chiefly because of overt similarites in the structure of the male phallosome. There are
important differences in all life stages. The adults differ in having proepisternal scales
and ornate markings. The larval and pupal stages differ in the development of many
setae. Despite the differences in chaetotaxy, the larval head capsule, comb scales, and
siphon are constructed almost like those in the aberrant Culex whitmorei (Giles, 1904)
which is currently recognized as a member of this subgroup (Sirivanakarn, 1976). It is
very likely that both poicilipes and whitmorei will have to be placed in separate
subgroups after members of the sitiens group are studied more thoroughly.
Bionomics.
The immatures of this species generally inhabit clean bodies of
standing water with thick vegetation. The breeding places are normally exposed to full
sunlight. Rice fields, shallow pools, and ditches are common breeding sites in Egypt.
This mosquito is often associated with antenna&s, perexiguus, and Anopheles (Celia)
pharoensis Theobald, but rarely with other species. Adults enter houses in Egypt
(Kirkpatrick, 1925) and readily attack man near swamps in the Sudan (Lewis, 1956)) but
are not known to vector any pathogens of human disease.
Harbach: Subgenus Culex in SouthwesternAsia
97
Distribution.
Culex poicilipes is widely distributed south of the Sahara and
enters Egypt along the Nile, Populations occur in the Sinai Peninsula and the Negev
Desert of southern Israel.
Material examined. 294 specimens. A total of 122 specimens (74 females,
14 males, 5 male genitalia, 10 pupal exuviae, 9 larval exuviae, and 10 fourth-instar
larvae) were examined from southwestern Asia - EGYPT: Beni Suef (Biba), El
Bahayrah (Alexandria), El Buhayra (Mansura, Wadi Natrun), El Isma’iliya (Abu Khalifa,
Isma’iliya), El Fayyum (Birket Qarum), N Giza (Harania, Kafr el Ghataty, Kerdasa), El
Qulyubiya (Ezbet Ayoub), El Sharqiya (Inshas el Raml); ISRAEL: (Ha’arana, Iddan, Neot
Hakikar, Yeroham, Zikim). An additional 172 specimens (95 females, 39 males, 7 male
genitalia, 3 pupal exuviae, 5 larval exuviae, and 23 fourth-instar larvae) were examined
from ANGOLA (the holotype female of taeniorhynchoides ), BENIN, ETHIOPIA,
GAMBIA, GHANA, KENYA, MADAGASCAR
(including the types series of
madagascariensis and auritaenia), MALAWI, MALI, MAURITANIA, MOZAMBIQUE (the
holotype female of punctipes), NIGERIA (including the holotype female), SENEGAL,
SOUTH AFRICA, SUDAN (including the holotype female of maculipes), TANZANIA,
UGANDA (including the holotype female of par), ZIMBABWE, and unknown localities.
Culex (Culex) mimeticus
No&
mimeticus No&, 1899: 240. Type(s) (female): Grassano in Basilicata,
[Italy] (non-extant).
pseudomimeticus
Sergent, 1909: 445. Type(s) (adult): Oued Chiffa,
Algeria (non-extant). Synonymy with mimeticus by Edwards, 1932a:
205.
Culex mimeticus No& of Giles, 1901 a: 605 (India, A, distr.); Theobald, 1901 a: 329 (India,
A key, F’); Christophers, 1906: 9 (? India, L); Joyeux, 1918: 533 (Macedonia,
~011.);Waterston, 1918: 9 (Macedonia, A, L habitat); Barraud, 1921: 395 (Israel,
Jordan, Syria); Edwards, 1922: 284 (Oriental Region, A, key); Barraud, 1923: 941
(India, L’); Buxton, 1923: 316 (Israel, Jordan, toll. rec., biol. note); Theodor,
1923: 341 (Jordan, Israel, P’, key); Barraud, 1924a: 991 (India, A key, M*, F);
Parr, 1943: 246-250 (Syria, Lebanon, A, P, L keys, bionomics); Senevet, 1947b:
212 (North Africa, L key); Theodor, 1952: 113 (Middle East, zoogeogr.); Dow,
1953: 688 (Iran, toll. rec.); Senevet et al., 1957a: 86 (North Africa, L’); Margalit
and Tahori, 1973: 90-93 (Sinai, distr., L assoc.); Margalit et al., 1973: 32 (Israel,
toll. rec., L assoc.); Kitron and Pener, 1986 (Israel, L bionomics).
Culex (Culex) mimeticus No6 of Edwards, 1921: 337 (E. Mediterranean Region, Israel,
Jordan, A, L keys, tax., distr.); S&guy, 1924: 34, 186 (E. Mediterranean Region,
Israel, Jordan, A, L keys, L’, tax., distr., syn.); Edwards, 1926: 130 (E. Mediterranean Region, Israel, Jordan, A, L keys, A, L, L assoc., distr.); Stackelberg,
1927: 155 (Middle East, M, F keys, A, M gen., distr.); Martini, 1931: 371 (Israel,
Jordan, Turkey, A, L keys, M*, F, L’, distr.); Barraud, 1934: 409 (India, A, L keys,
M*, F, L*, distr.); Edwards, 1934: 451 (India, F, distr.); Senevet, 1947a: 121
(Egypt, L’, distr., L assoc.); Senevet, 1949: 56 (North Africa, M gen.*, key);
Monchadskii, 1951: 266 (Middle East, L’, L assoc., distr.); Senevet and Andarelli,
1954: 57 (North Africa, M, F, A key); Senevet et al., 1957b: 92 (North Africa, L);
Parrish, 1959: 266 (Turkey, distr.); Senevet and Andarelli, 1959: 151 (North
Africa, A, P, L keys, syn., M*, F’, P*, L’, distr., biol., L assoc.); Abdel-Malek, 1960:
98
Contrib.Amer. Ent. Inst.,vol. 24, no. 1, 1988
112-121 (Syria, COILsites, L bionomics); Derwesh, 1965: 44 (Iraq); DuBose and
Curtin, 1965: 351, 354 (Mediterranean area, A, L keys); Abul-hab, 1966: 283
(Iraq, distr., L bionomics); Abul-hab, 1968: 246 (Iraq, L key, distr.); Lotfi, 1970:
401 (Iran, toll. rec.); Margalit and Tahori, 1970b: 152 (Israel, COILsites, distr.);
Aslamkhan, 1971: 154 (Pakistan); Lotfi, 1973: 206 (Iran, toll. sites); Gutsevich et
al., 1974: 375 (Middle East, F, M gen., L keys, M*, F*, L*, distr., bionomics);
Margalit and Tahori, 1974: 87 (Israel, toll. rec.); Lotfi, 1976 (Iran, L key, L’, ecol.
note); Sirivanakarn, 1976: 145 (Oriental Region, A, P, L keys, M’, F’, P*, L*, syn.,
tax., distr., bionomics); lbrahim et al., 1983: 91 (Iraq, L*, key); Harbach, 1985a: 86,
96, 107 (SW Asia [excluding records for Afghanistan and Saudi Arabia], Egypt,
distr., A, L, key).
Adult. The presence of pale spots on the wings of this medium-sized species
readily distinguishes it from all other species of Culex known to occur in southwestern
Asia and Egypt. This character evokes a superficial resemblance to Anopheles species
of the subgenus Cellia.
FEMALE. Head: Antenna dark; length 2.0-2.3 mm, mean 2.1 mm; pedicei with
small patch of tiny pale scales on dorsal surface. Proboscis black-scaled with well
defined median whitish ring extending from 0.4 to 0.6 from base; length 1.9-2.1 mm,
mean 2.0 mm. Maxillary palpus dark with pale scales at apex. Forked scales of vertex
pale yellow medially, dark brown to black laterally; falcate scales white, yellowish
dorsolaterally; lateral spatulate scales white. Thorax: Integument brown, pleura with
darker spots on proepimeron, postspiracular area, prealar area, anterior and posterior
margins of mesokatepisternum, and anterior and upper areas of mesepimeron; meron
also dark. Scutal scales coarse, predominantly pale yellow to golden yellow, with white
to pale yellow scales on anterior promontory,lateral margins and prescutellar area, scales
on fossa and middorsal area sometimes golden brown to brown. Scutellum with white
falcate scales, large patch on median lobe, few on lateral lobes. Ante- and postpronota
with golden-yellow falcate scales; postpronotum with about 6 setae on posterior border.
Pleura with patches of white spatulate scales as follows: proepisternum with patch
immediately above forecoxa, mesopleuron with upper and lower patches on
katepisternum and anterior and upper patches on anepimeron, scales on katepisternum
more elongate; numbers of pleural setae: 4-8 upper proepisternal, 1O-l 2 prealar, 5-7
upper mesokatepisternal, 9-l 3 upper mesepimeral; without lower mesepimeral seta.
Wing (Fig. 2E): Length 3.8-4.0 mm, mean 3.9 mm; black-scaled with white or yellowishwhite spots, pattern follows: costa dark with sector, subcostal and preapical pale spots;
Rl dark with subcostal and preapical pale spots; pale scales at apex of R2+3 continuous
with pale scales at base of R2 and R3; R2 with preapical pale spot; R4+5 with dark base
followed by large median pale spot and small apical dark spot; pale spot at furcation of
Ml+2; M3+4 with large median pale spot; CuA with very small pale spot at apex
confluent with pale fringe spot; 1A with large pale spot near middle. Halter: Pedicel and
scabellum yellow; capitellum brown with pale scales. Legs (Fig. 2H): Forecoxa
black-scaled with patch of white scales at base; mid- and hindcoxae with longitudinal
stripe of white scales on anterior side of lateral midline. Anterior surface of forefemur
entirely black-scaled with ill-defined narrow line of white scales on proximal 0.5, with
distinct posterodorsal stripe of white scales not reaching apex; anterior surface of
midfemur black-scaled with white scales at base, posterior surface white-scaled with
small subapical area of black scales; hindfemur white-scaled except for dorsal stripe of
black scales beginning near base and broadening distally to cover approximately distal
0.25 of anterior and posterior surfaces. Tibiae and tarsi mainly black-scaled; all tibiae with
Harbach: Subgenus Culex in SouthwesternAsia
99
posterior white stripe, mid- and hindtibiae with narrow anterior white stripe, anterior stripe
of midtibia incomplete distally, anterior stripe of hindtibia incomplete at ends.
Tarsomeres 1-4 of all legs with basal and apical white rings, basal rings broader.
Abdomen: Terga mainly black-scaled; tergum I with submedian patches of black scales
on posterior border that may be separated by a few median pale scales; terga II-VII with
basal bands of yellowish-white scales and large basolateral spots of white scales, spots
may cover entire lateral surface, bands straight or slightly convex, 0.15-0.35 tergum
length and barely if at all reaching spots. Sternum I with some dark scales near
posterolateral corners; sterna II-VII with posterolateral dark spots that may join to form
broad apical band.
MALE. Resembling female, differences follow. Head: Maxillary palpus with broad
subapical ring of pale scales on palpomere 3, narrow ring of pale scales at bases of
palpomeres 4 and 5, and pale scales on dorsal surface of palpomere 5. Vertex with
fewer forked scales; falcate scales fewer, longer and all yellowish white; larger patch of
pale lateral spatulate scales. Thorax: Scutal scales longer, predominantly whitish to
golden yellow with spots of golden-brown scales on fossae and middorsal area, lightest
scales on lateral margins and posterior 0.33 of scutum. Scutellar scales fewer and
longer. Upper proepisternal setae more numerous, usually 8-l 0. Wing: Length 2.9-3.3
mm, mean 3.1 mm; pattern of spots differs as follows: sector pale spot on anterior
margin of subcosta; pale spots at apex of costa, RI and R2; pale spots absent from
furcation of R1+2, furcation of Ml+2 and apex of CuA; M3+4 with small basal dark spot
followed by small pale spot and large dark spot; 1A pale spot extending further proximad
than in female. Abdomen: Without basolateral spots; basal pale bands concave,
0.35-0.45 tergum length on dorsal midline. Genitalia (Fig. 35): Form as figured;
structure much as in sitiens, differing chiefly as follows. Ninth tergal lobe poorly
differentiated, with 4-7 short setae. Subapical lobe of gonocoxite with 3 or 4 setae in
group d-e, 3 usually present, these flattened with recurved apex and 1 or 2 recurved
denticles before tip; seta f flattened and bladelike, rounded apically; h longer. Inner
division of phallosome more closely fused to outer division, not as broad, dorsocaudal
angle elongate and slender, with 1-3 teeth (usually 2) arising very near tooth on
posteromesal margin of outer division; outer division with much smaller process
projecting laterally from ventrocaudal margin, dorsal process not as prominent. Proctiger
essentially as in Miens; basal lateral arm of paraproct generally shorter and more slender.
Cereal sclerite with 2 or 3 setae on posterolateralcorner. Tergum X small.
Pupa (Fig. 35). Placement and form of setae as figured, range and modal
number of branches in Table 17; without striking distinctive features, closely resembles
sitiens but setae l-11, 6-III-VI, and 9-VIII usually with fewer branches. Cephalothorax:
Lightly tanned. All setae except lo-CT usually double or triple, more often double;
lo-CT usually with 4 or 5 branches (4-6). Trumpet: Moderately tanned; slender,
pinna slightly widened; index 4.7-7.0, mean 5.9; pinna 0.25-0.40 trumpet length.
Abdomen: Lightly tanned; length 2.6-3.9 mm, mean 3.2 mm. Seta 6-I,11short, not more
than twice length of 7-l,ll; 7-I,11usually double (l-3); l-11 weak, with only l-4 branches,
l-Ill-V generally with progressively fewer branches on each successive posterior tergum,
l-111frequently with 4 or 5 branches (3-6) l-IV most often triple (2-5), 1-V usually double
(l-3), 1-VI commonly single (1,2), l-VII like 1-V; 6-III-VI usually single (see table), 6-lll,lV
distinctly shorter than 6-V,VI; 5IV-VI strongly developed, about 1.5 length of tergum
following, 5-IV most often double (2-4) 5-V,VI double; 2-VII usually mesal to l-VII; 9-VIII
usually with 6 or 7 branches (4-9). Paddle: Lightly tanned; length 0.76-l .03 mm, mean
0.94 mm; width 0.59-0.78 mm, mean 0.67 mm; index 1.3-l .6, mean 1.5.
Larva (Fig. 36). Character and positionsof setae as figured, range and modal
100
Contrib.Amer. Ent. Inst.,vol. 24, no. 1, 1988
number of branches in Table 37; resembling theileri, simpsoni, and sinaiticus in
the form of the comb scales but differing conspicuously from these species in the
placement of seta 1-S and in having setae 2,3-A inserted approximately midway
between seta 1-A and apex of antenna, easily distinguished from other members
of the sitiens group by the character of the comb scales and the reduced
branching of seta 4-X. Head: Length 0.60-0.72 mm, mean 0.66 mm; width
0.95-1.10 mm, mean 1.05 mm; lightly tanned, darker posteriorly. Hypostomal
suture ending before collar; collar well developed, heavily tanned. Dorsomentum
almost shape of an equilateral triangle; usually with 7 or 8 teeth (6-8) on each side
of middle tooth. Seta 1-C moderately long, stout, pointed, tanned; 2-C absent;
4-C double, triple, or with 4 branches occurring in nearly equal frequencies; 5-C
with 3 or 4 branches, more often with 4; 6-C double or triple, more often double;
14-C rather large and spinelike. Antenna: Length 0.46-0.62 mm, mean 0.55 mm;
lightly tanned, distal part dark, with dark ring at base. Seta 1-A with about 25
branches (21-27); 2,3-A inserted almost halfway between 1-A and apex. Thorax:
Integument hyaline, smooth. Setae l-6-P equally long and single; 7-P usually
triple, seldom double; 8-P double or triple in nearly equal frequencies, distinctly
shorter than 7-P. Seta 1,2-T usually single, occasionally double, 2-T 2 or 3 times
as long as 1-T; 13-T weak, shorter than 12-T, usually double, occasionally single
or triple. Abdomen: Integument hyaline, smooth. Seta 3-l,lV single or double,
3-ll,lll,V,VI almost always single, 3-VII often double but frequently triple or with 4
branches; 6-l-V usually triple, 6-I,11often with 4 branches, 6-VI usually double,
occasionally triple; 7-l as in sitiens; 1-III short and single, 1-IV-VI usually double,
6-V,VI stronger than 6-IV, l-VII triple or with 4 branches, more often with 4;
13-Ill-V,Vll weakly developed, much shorter than seta 1 of same segment, almost
always single or double. Segment VIII: Comb with 27-50 (mean 33) sharply
pointed scales with short lateral fringes on proximal part. Siphon: Index
4.55-7.41, mean 6.07; lightly tanned, slightly darker at apex, basal rim moderately
to heavily tanned. Pecten composed of 13-20 (mode 15) long slender spines
with rather short line of fused denticles at base. Seta 1-S in 5-7 pairs (lo-14
setae), 1 subapical pair borne laterally, others along posterior midline, proximal 4
or 5 pairs stronger, most often with 4 or 5 branches (see table), lateral pair and
most distal pair shorter and usually double or triple; 2-S rather long and curved.
Segment X: Saddle complete; lightly tanned, slightly darker posteriorly; with rows
of minute spicules; length 0.37-0.42 mm, mean 0.41 mm, siphon/saddle index
3.90-4.65, mean 4.22. Seta 1-X normally double (l-3); 2-X normally double,
occasionally triple; 4-X usually in 6 pairs, 1 seta sometimes unpaired, most setae
commonly with 4 branches (3-6). Anal papillae long and slender, about twice
length of saddle.
Systematics.
This species is the nominate member of an interesting
subgroup which is largely confined to the Oriental Region. Culex mimeticus is the
only species of the group which is known to occur in the Mediterranean
Subregion of the Palaearctic. It is apparently mainly a chaparral and temperate
grassland species which is excluded from the Arabian Peninsula and the Sahara.
The adults of mimeticus are unique among the Culexof southwestern Asia
in having spotted wings much as in species of the subgenus Celia of Anopheles.
The larva is variable but distinctfrom all other species in the region. It bears a very
superficial resemblance to the larva of Culex deserticola Kirkpatrick, 1924( 1925),
but is distinguished at once from this species by the placement of setae 2,3-A
and the character of the comb scales.
Harbach: Subgenus Culex in SouthwesternAsia
101
It is possible that two or more species are currently confused under the concept
of mimeticus in the Mediterranean Subregion. Topotypic material from Italy, for example,
differs noticeably in the size and shape of the pupal trumpet. There also seems to be a
number of geographically isolated populations in the region. Individual rearings are
lacking from almost all localities and the available material is much too scanty to permit a
detailed comparative analysis. The above diagnosis is based mainly on reared material
from the Negev of Israel.
Blonomics. The natural breeding sites of mimeticus appear to be springs and
residual pools in drying streams. The water in these habitats is fresh and clear and
usually contains growths of Spirogyra. Other collections have been made at the margins
of streams, pools in swampy ground, irrigationditches, and animal hoofprints. Larvae are
frequently found with Anopheles (Cellia) hispaniola (Theobald), An. (Gel.) sergentii
(Theobald), and An. (Gel.) superpictus Grassi. Adults apparently occasionally enter
human habitations (Hsiao and Bohart, 1946; Ribeiro et al., 1977), but females have
never been seen landing on or biting man or animals. Nothing is known about the
biology of the adults.
Distribution.
This species occurs in the southern Palaearctic and Oriental
regions. Its range extends from lands around the Mediterranean to eastern and
southeastern Asia. Culex mimeticus seems to be less common than it once was,
probably as a result of pesticide usage, organic pollution, and the destruction of
preferred breeding places. The species has a very spotted distribution.
Material examined. 215 specimens. A total of 128 specimens (26 females,
27 males, 24 male genitalia, 15 pupal exuviae, 17 larval exuviae, and 19 fourth-instar
larvae) were examined from southwestern Asia -- IRAN: (Amirabad, Chalus,
Izeh-Ahwaz, Kanroud, Kazeroon, Khorramabad, Natchi Bahran, Tehran, Zanjan); IRAQ:
(Mirgasur); ISRAEL: (Dalia, Ein Amud, En Avedat, Jericho, Jirou, Latron, N. Bet
Shearim, N. Moran, Wadi Kabalah, Wadi Kurn); JORDAN: (Jerash, Tafile, Tafilan, Wadi
Hasa, Wadi Musa, unknown localities); TURKEY: (Yenice). An additional 87 specimens
(24 females, 28 males, 12 male genitalia, 10 pupal exuviae, 11 larval exuviae, and 2
fourth-instar larvae) were examined from ALGERIA, GREECE (CRETE), ITALY (topotypic
material), MOROCCO, and TUNISIA.
Culex (Culex)
bitaeniorhynchus
G i Ie s
bitaeniorhynchus
Giles, 1901 a: 607 . Type(s) (adult): Travancore,
India (non-extant).
ager Giles, 1901 b: 196 (Taeniorhynchus ). Type(s) (female): Madras,
Presidency, India (non-extant, lectotype designation of Bram,l967a:
263 invalidated by Harbach, 1983: 105). Synonymy with bitaeniorhynchus by Edwards, 1913c: 231.
ethiopicus Edwards, 1912b: 30 (Taeniorhynchus ager var.). +Holotype
female: Bole, Gold Coast, West Africa (BM). NEW SYNONYMY.
abdominalis Taylor, 1913: 53 (Culicelsa ). Type(s) (female): Ayr and
Townsville, Queensland, Australia (non-extant). Synonymy with
bitaeniorhynchus by Edwards, 1913~: 231.
maculipesarabiensls
Patton, 1905: 635 (Taeniorhynchus tenax var.).
Type(s) (adult; larva): D’thala, Hardeba and Nobat, P.D.R. Yemen
(non-extant). Synonymy with ethiopicus by Edwards, 1941: 291.
karatsuensls Mochizuki, 1913: 28. Type(s) (male; female; egg): Karat-
102
Contrib.Amer. Ent. Inst.,vol. 24, no. 1, 1988
su, Kyushu, Japan (location unknown).
rhynchus by Edwards, 1932a: 202.
Synonymy with bitaenio-
Culex ager var. ethiopicus Edwards of Edwards, 1912~: 381 (Africa, L’, key); Ingram and
Macfie, 1919: 65 (W. Africa, P*, L*, L habitat).
Culex bitaeniorhynchus Giles of Barraud, 1923: 936 (India, L*); Barraud, 1924a: 984 (in
part; India, A key, syn., M*, F, distr.).
Culex bitaeniorhynchusvar. ethiopicus Edwards of Edwards, 1920: 135 (Africa, M gen.).
Culex (Cu/ex) bitaeniorhynchus Giles of Edwards, 1921: 337 (in part; Palaearctic
Region, A, L keys, tax., bionomics, distr.); Edwards, 1926: 129 (in part;
Palaearctic Region, A, L keys, syn., A, distr.); Stackelberg, 1927: 153 (in part; S.
Asia, M, F keys, A, M gen.*, distr.); Martini, 1931: 362 (in part; S. Asia, A, L keys,
M*, F, L*, distr.); Barraud, 1934: 391 (in part; India, A, L keys, M*, F, L’, syn., L
habitat, distr.); Monchadskii, 1951: 258 (in part; S. Asia, L”, L assoc., distr.); Lotfi,
1970: 401 (Iran, toll. rec.); Aslamkhan, 1971: 154 (in part; Pakistan); Lotfi, 1973:
206 (Iran, toll. rec.); Gutsevich et al., 1974: 367 (in part; S. Asia, M*, F, L’, distr.,
bionomics, med. imp.); Lotfi, 1976: 72, 74, 79 (Iran, L key, bionomic note, L’);
Sirivanakarn, 1976: 65 (Oriental Region, A, P, L keys, M*, F*, P*, L*, tax., distr.,
bionomics, med. imp.); Harbach, 1985a: 86, 98, 107 (Iran, Pakistan, P.D.R.
Yemen, Yemen Arab Republic, A, L keys).
Culex ethiopicus Edwards of Lewis, 1945: 14 (Sudan, L, distr.).
Culex (Culex) ethiopicus Edwards of Edwards, 1941: 291, 415, 482 (Afrotropical
Region, A key, M*, F, P*, syn., distr.); Lewis, 1943a: 281 (Eritrea, toll. rec., L
bionomic note); Lewis, 1943b: 69 (Sudan, bionomic notes); Abbott, 1948: 44
(Sudan, distr.); Lewis, 1948: 144 (Sudan, L habitat); Hopkins, 1952: 282
(Afrotropical Region, L key, L*, bionomics); Knight, 1953a: 229 (Yemen Arab
Republic, toll. rec., L habitat); Lewis, 1956: 706 (Sudan, toll. rec.); Mattingly and
Knight,l956: 106-132 (P.D.R. Yemen, Yemen Arab Republic, L*, A, L keys,
distr., bionomics); Mekuria, 1968: 78 (Ethiopia, distr.).
Adult.
A rather large species easily recognized by the speckled wings and
rather broad apical yellow bands of the abdominal terga.
FEMALE. Head: Antenna dark, pedicel and flagellomere 1 paler; pedicel
sometimes with some pale scales on mesal surface; length 1.8-2.4 mm, mean 2.0 mm.
Proboscis with distinct median pale ring about 0.25 proboscis length, pair of dorsolateral
pale spots at apex before labella and scattered pale scales proximal to ring; length
1.6-2.1 mm, mean 1.9 mm. Maxillary palpus dark-scaled; palpomere 4 with pale scales
apically, sometimes with scattered pale scales proximally; length 0.4-0.5 mm, slightly
more than 0.2 length of proboscis. Dorsal falcate scales of vertex coarse, very pale
yellow; forked scales largely pale yellow to yellowish brown, some darker
posterolaterally; lateral and ventral spatulate scales yellowish white. Thorax: Integument
brownish yellow, scarcely darker dorsally. Scales on anterior 0.7 of scutum mainly pale
yellow to golden yellow, somewhat mottled by presence of some golden-brown to
brownish scales, particularly on anteromedian area and sublaterally at middle; posterior
0.3 clothed in golden-brown and dark brown scales with some coarser pale scales on
antealar area and posteriorly on prescutellar area; scutal setae golden brown except for
distinctly paler (yellowish) antealar setae, supraalar setae dark, dense (numerous) and
long. Scutellum with pale scales on lateral lobes, median lobe mainly pale-scaled with
some brownish-yellow to dark brown scales anteriorly; 5 or 6 long setae on lateral lobes,
7-l 1 on median lobe. Ante- and postpronota with coarse yellowish to golden falcate
Hatbach: Subgenus Culex in SouthwesternAsia
103
scales, scales golden brown to brown centrally. Pleural setae mainly pale, usually
entirely pale: 8-l 1 upper proespisternal, 6-l 3 prealar, 5-7 upper mesokatepisternal,
6-10 lower mesokatepisternal, 4-14 upper mesepimeral, and no lower mesepimeral.
Pleural scales yellowish white: proepisternum with upper patch comprised of falcate
scales laterally and spatulate scales anteriorly, mesokatepisternumwith upper and lower
patches of elongate spatulate scales, mesepimeron with anterior patch of elongate
spatulate scales at level of upper katepisternal patch and patch of similar scales before
upper mesepimeral setae. Wing (Fig. 2F): Length 3.6-4.6 mm, mean 4.1 mm; length of
cell R2 2.4-3.5 length of vein R2+3, mean 3.0; length of cell Ml about 0.9 length of cell
R2; dorsal surface with broad apically rounded yellowish-white and brown spatulate
scales evenly intermixed on all veins; ventral surface with mixture of pale and dark
spatulate scales on costa, subcosta, Rs and M, other veins with pallid brownish linear to
very narrow spatulate scales, scales on Rl, R4+5, M2 and M3+4 linear and near linear,
scales on other veins near linear and very narrow spatulate. Halter: Integument
yellowish to brownish yellow; capitellum pale-scaled, distal part of scabellum and base of
capitellum usually with some dark scales. Legs: Forecoxa with dark and pale spatulate
scales anteriorly, latter dominant at base and apex; mid- and hindcoxae with anterior
patches of pale scales, some dark scales distally. Anterior, posterior, and ventral
surfaces of trochanters with mixture of pale and dark scales. Femora with more or less
even mixture of dark and pale scales on anterior surface, posterior surface
predominantly pale with interspersed dark scales mainly on distal 0.5 or less. Dorsal
surface of foretibia and anterior surface of mid- and hintibiae with interspersed dark and
pale scales, ventral surface of foretibia and posterior surface of mid- and hindtibiae
mainly pale-scaled with intermingled pallid brown scales, especially proximally. Tarsi
dark-scaled; tarsomeres l-4 with narrow apical and broader basal rings of pale scales;
tarsomere 1 of all legs and tarsomere 2 of hindleg with some scattered pale scales.
Tergum I with rather large median posterior patch of yellow scales,
Abdomen:
specimens from Asita, Wallo Province, Ethiopia with some rather diffuse dark and pale
scales on lateral borders; terga II-VIII with broad apical yellow bands (about 0.33 tergum
length) not reaching lateral margins, scattered yellowish scales among proximal dark
scales, and large basolateral white spotsdiagonally opposed to apical bands, tergum VIII
also with basal pale band contiguous with basolateral spots that may join apical band
medially. Sterna II-VII with narrow yellow apical bands joined medially to broad whitish
basal bands, apicolateral corners with dark scales, sometimes with some scattered pale
scales, particularly on proximal sterna where dark patches are larger and less definite;
sternum VIII with lateral pale patches, middle devoid of scales.
MALE. Like female, differing mainly in sexual characters, major differences follow.
Head: Median pale ring of proboscis narrow, 0.1-0.15 proboscis length; with ventral
setal cluster at base of ring before false joint; false joint about 0.55 from base. Maxillary
palpus mainly dark-scaled; with narrow ring of pale integument across junction of
palpomeres 2 and 3; palpomere 3 with rather long median pale patch with dispersed dark
scales on dorsal and lateral surfaces, sometimes with scattered pale scales distal to pale
patch, dense patch of long setae on ventrolateral surface immediately distal to pale
patch, and ventromesal surface with line of short antrorsely curved setae along entire
length; palpomeres 4 and 5 densely setose with basal pale rings, palpomere 4 with
scattered pale scales distal to ring, distal 0.5 of palpomere 5 with pale setae and scales;
length 2.6-2.9 mm, mean 2.8 mm, extending beyond tip of proboscis by length of
palpomere 5 and part of 4. Wing: Length of cell R2 1.6-l .8 length of vein R2+3; length
of cell Ml 1.0-l .l length of cell R2; scales fewer and smaller, particularly posteriorly.
Abdomen: Terga same as in female but basolateral white spots smaller and sometimes
104
Contrib.Amer. Ent. inst.,vol. 24, no. 1, 1988
less distinct (comprised of fewer scales in loose cluster) on anterior terga (II-IV or V).
Sterna as in female, tergum VIII (ventral in position) like sterna. Genitalia (Fig. 37): Form
as figured; closely resembling those of infula Theobald of the Oriental Region. Ninth
tergal lobe not produced, indicated by line of 3-7 (mode 6) setae. Gonocoxite rather
short, lateral setae as in poicilipes; subapical lobe moderately prominent, part bearing
setae a and b more strongly produced; setae a and b rather short and stout, c slender
and removed from a and b, all usually hooked at tip; 4 or 5 simple setae in group d-f, 1 (I
?) longer and heavier (flattened) than the others; seta g short and lanceolate (sometimes
duplicated in specimens from the Oriental Region); h not distinctly differentiated, short
and acuminate. Gonostylus rather short, simple, tapered distally; gonostylar claw of the
usual form. Phallosome much longer than broad; aedeagal sclerite larger than lateral
plate, angled ventrad, crest about as large as proximal part, proximal part with patch of
minute bumps along ventral margin of mesal surface; lateral plate with strongly
differentiated inner and outer divisions, inner division (= ventral arm) a laterally
compressed lobe with a fuzzy covering of minute spicules, dorsocaudal angle rounded,
ventrocaudal angel produced ventrad and pointed, dorsal margin with a flat laterally
directed single or bilobed process (= dorsal arm ?); outer division (= lateral arm)
developed as a large ventrolateral winglike flap, dorsal process and dorsal articulatory
process indicated by a thickened ridge. Proctiger rather long; paraproct long and
narrow, basal lateral arm short and slender, crown a relatively small cluster of rather short
spinelike spicules and a few small lateral blades. Cereal sclerite with 2 or 3 setae on
posterolateral margin. Tergum X small.
Pupa (Fig. 37). Character and positions of setae as figured, range and modal
number of branches in Table 18; well distinguished from other species of the subgenus
in the region by the darkened cuticle around the bases of most setae, differs from other
species of the sitiens group in having setae 1 l-CT, 3-I-111,and l-VII normally all single.
Cephalothorax: Lightly to moderately tanned a bright yellowish color. Setae short, all
except 5,10,11 -CT usually double (see table); 5CT double or triple, more often triple;
lo-CT frequently with 7 branches (510); 1 l-CT usually single, sometimes double.
Trumpet: Moderately to heavily tanned, brown, tracheoid area darker; strongly flared,
funnel-shaped, slightly bent at tracheoid area; index 4.0-5.7, mean 4.6; pinna large,
about 0.4 trumpet length; tracheoid area about 0.3 trumpet length. Abdomen: Lightly
to moderately tanned a bright yellowish color, cuticle around bases of almost all setae
strongly tanned a dark brown (not shown in figure); length 3.3-3.9 mm, mean 3.6 mm;
caudolateral corners of terga VII and VIII sharply produced as in poicilipes. Seta 7-l single
to triple, 7-11 single or double; l-11 single or double, aciculate; l-Ill-VI moderately
developed, 1-III often with 6 or 7 thin branches (6-l 0), branches of 1-IV-VI thicker, 1-IV
very often with 4 branches (4-6) 1-V double or triple, l-VI single or double; l-VII single,
thinner than 1-IV-VI; 5-IV-VI resemble l-IV-VI respectively, slightly longer, about length
of following tergum; 6-III-VI single, 6-lll,lV relatively short and thin, 6-V,VI longer and
thickened, markedly different from 6-lll,lV.
Paddle: Inner part darkened; length
0.98-1.05 mm, mean 1.02 mm, width 0.76-0.81 mm, mean 0.79 mm, index 1.3-1.4.
Larva (Fig. 38). Form and placement of setae as figured, range and modal
number of branches in Table 38; bearing little resemblance to other members of the
subgenus in southwestern Asia, easily recognized by the triangular dorsomentum with
its minutely serrated edges and the absence of a discrete median labral plate. Head:
Length 0.61-0.64 mm, mean 0.63 mm; width 0.90-1.02 mm, mean 0.97 mm; lightly
tanned, dorsomentum and collar moderately to heavily tanned. Median labral plate
indistinguishably fused to cranium. Hypostomal suture complete, extended posteriorly
from posterior tentorial pit but ending well before collar. Collar large and well developed,
Harbach: Subgenus Culex in SouthwesternAsia
105
heavily tanned only on posterior margin. Dorsomentum an equilateral or isosceles
triangle with numerous minute teeth on lateral sides. Seta I-C long, stout, slightly
swollen just beyond midlength, sharply pointed; 2-C absent; 4-C usually double (l-4);
5-C double or triple; 6-C double; 7-C distinctly shorter than 5,6-C, branches thinner
(4-7); 10,15-C double or triple; 11,13-C both usually single (see table). Antenna:
Length 0.36-0.48 mm, mean 0.41 mm; lightly tanned, distal part darker near tip; proximal
part with few spicules on dorsal and ventral surfaces; seta I-A inserted near middle, with
about 19 branches (16-27); 2,3-A subapical. Thorax: Integument hyaline, smooth.
Seta 4-P short, about half length of I-P, usually double, sometimes single; 7-P triple;
8-P usually double (2,3), only slightly larger than 4-P. Tubercles of meso- and
metathoracic pleural setal groups large and heavily tanned; seta 13-T small, with 3-6
branches. Abdomen: Integument like that of thorax. Dorsal and ventral setae all shorter
than usual, especially 1 and 13; seta 3-l usually double (l-3), 3-II-VII usually double or
triple (see table); 6-l-V normally triple (see table), 6-VI normally double; I-III-VII normally
double or triple; 13-Ill-V,Vll usually double, sometimes triple, 13-VII seldom with 4
branches. Segment VIII: Comb with 5-7 (mode 6) large spinelike scales with tiny
spicules on sides of expanded base. Setae 2,4-VIII normally double, 2-VIII seldom triple.
Siphon: Index 5.01-8.35, mean 6.83; long and slender, broadest at base, sigmoid in
lateral view; lightly tanned, basal rim darkened before acus. Pecten very short, confined
to basal 0.1; comprised of 5-8 (mode 7) inconspicuous spines with O-3 tiny spicules on
ventral side at base. Seta I-S in 4 pairs, widely spaced, in line, usually double or triple
(see table), no longer than diameter of siphon at point of attachment; 2-S rather longer
than usual, straight, stiff, darkly tanned. Segment X: Saddle complete; lightly tanned;
smooth; length 0.38-0.45 mm, mean 0.43 mm, siphon/saddle index 3.89-5.69, mean
4.84. Seta I-X often triple (2-4); 2-X often with 3 or 4 branches (2-5); 4-X usually in 6
pairs. Anal papillae subequal, longer than saddle.
Systematics.
The concept of bitaeniorhynchus presented here is essentially
that of Sirivanakarn (1976). This concept includes populations in southwestern Asia and
Africa which Edwards (1941) and all later workers recognized under the name of Culex
ethiopicus Edwards, 1912b. I could find no distinct differences in any stage to separate
these populations from typical bitaeniorhynchus in the Oriental Region. The form which
Edwards (1941) and later authors confused for bifaeniorhynchus in Africa appears to be
an unnamed species related to Culex infula Theobald, 190la which occurs in countries
east of Pakistan. This species remains unnamed because none of the currently
available material is individually reared and is inadequate for description.
Culex bitaeniorhynchus and Taeniorhynchus ager Giles, 1901 b are objective
synonyms. They are based on the same name-bearing type female which is apparently
lost (Harbach, 1983). Nevertheless, there are two males in the British Museum which
were labelled as types of ager by Theobald (1901 b). These are the specimens which
Edwards (1922) Bram (1967b), and Sirivanakarn (1973; 1976) used to establish the
concept of “typical” bitaeniorhynchus in the Oriental Region. Bram (1967b) designated
one of these specimens as the lectotype of ager. Unfortunately, these specimens have
absolutely no taxonomic standing and the lectotype designation is invalid (see Harbach,
1983). But since these specimens already serve as a standard for the current
application of the name of this species, the specimen bearing the invalid lectotype label
of ager is hereby designated as the neotype of Culex bitaeniorhynchus Giles. This
action ensures taxonomic stability and restricts the type locality to Madras, India. The
neotype is identified by the following labels: “Paddy field / [two illegible marks] /
24.12.99 // Taeniorhynchus / ager / (Type). Theobald // Capt. Cornwall. / Madras. // Type
N LECTOTYPE / Taeniorhynchus / ager Giles / By R. A. Bram 6‘ 6”; with genitalia on an
106
Contrib.Amer. Ent. Inst.,vol. 24, no. 1, 1988
acetate strip attached to the pin that supports the specimen.
Culex bitaeniorhynchus is widely distributed in the Afrotropical and Oriental
regions. Its range also includes part of the Australian Region and the eastern part of the
Mediterranean Subregion of the Palaeactic. It is possible that bitaeniorhynchusconsists
of more than one biological species, but there is no indication of geographical
differentiation. The same degree of variation occurs within individual populations that
occurs throughout the known range of the taxon.
Bionomics. The immature stages of bitaeniorhynchus occur in permanent or
semipermanent bodies of standing water containing quantities of filamentous green
algae (Spirogyra). This species apparently bites man in the Oriental Region where it is
suspected of being involved in disease transmission. It has been found naturally
infected with larvae of Wuchereria bancrufti in India (lyengar, 1938) and Brugia malayi in
Sri Lanka (Carter, 1948). It also is associated with Murray Valley encephalitis in Australia
and with Batai virus in India (Karabatsos, 1985). Laboratory studies have shown that
Japanese encephalitis virus can be transmitted transovarially (Soman and Mourya, 1985)
and Sindbis virus has been isolated from populations in the Philippines and Australia
(Rudnick et al., 1962; Doherty et al., 1963). Females have not been reported to bite
man in southwestern Asia or Africa (as ethiopicusEdwards).
Distribution. Culex bitaeniurhynchus has a very wide distribution in the tropical
and subtropical areas of Palaeogaea. Populations occur in the Afrotropical Region,
eastern and southern areas of the Palaearctic Region, and the Oriental and Australian
regions. They occur in the southern coastal lands of southwestern Asia.
Material examined. 276 specimens. A total of 76 specimens (20 females, 18
males, 8 male genitalia, 9 pupal exuviae, 6 larval exuviae, and 15 fourth-instar larvae)
were examined from southwestern Asia - IRAN: (Hajiabad, Shah-Kahoor); PAKISTAN
(locations east of the lndus River): (Kahnakacha, Kasur, Lahore, Sargodna, Sattoki);
P.D.R. YEMEN: (Wadi Ma’adin); YEMEN ARAB REPUBLIC: (Wadi Siham near O
‘ bal).
Another 200 specimens (81 females, 47 males, 27 male genitalia, 3 pupal exuviae, 8
larval exuviae, and 34 fourth-instar larvae) were examined from BENIN, ETHIOPIA,
GAMBIA, GHANA (including the type series of ethiopicus ), INDIA (including the
neotype), MAURITANIA, NEPAL, NIGERIA, SUDAN, TANZANIA, UGANDA, ZAIRE,
ZIMBABWE, and unknown localities in Africa.
During this study 56 specimens (18 females, 13 males, 12 male genitalia, 1 pupal
exuviae, 6 larval exuviae, and 6 fourth-instar larvae) representing a species previously
regarded as bitaeniorhynchus in Africa were examined from BENIN, ETHIOPIA,
GAMBIA, GHANA, SENEGAL, SUDAN, TANZANIA, UGANDA, UPPER VOLTA, ZAIRE,
and ZIMBABWE.
Culex (Culex)
tritaeniorbynchus
Giles
tritaeniorhynchus
Giles, 1901 a: 606. +Holotype female: Travancore,
[Madras State], India (BM).
biroi Theobald, 1905b: 82. Syntypes (male; female): Bombay, India
(HNM). Synonymy with tritaeniorhynchusby Edwards, 1913~: 233.
summorosus Dyar, 1920: 180. Holotype male: Los Banos, [Laguna,
Luzon], Philippines (NMNH). Synonymy with tritaeniorhynchus by
Bram, 1967a: 225.
siamensis Barraud and Christophers, 1931: 283.
Lectotype male:
Chiang Mai, Thailand; designated by Mattingly, 1956: 37 (BM). Syn-
Harbach: Subgenus Culex in SouthwesternAsia
107
onymy with summorosus by Colless, 1957: 98.
Culex tritaeniorhynchus Giles of Theobald, 1901 a: 364 (India, A key, M*, F*); Edwards,
1913c: 233 (Oriental Region, syn., A, tax.); Barraud, 1920: 324 (Iraq, toll. rec.);
Austen, 1921: 116 (Israel, Jordan, toll. rec.); Barraud, 1923: 940 (India, L’);
Buxton, 1923: 316 (Israel, Jordan, toll. rec., A, L bionomics); Barraud, 1924a:
995 (India, syn., M*, F, distr.); Kirkpatrick, 1924(1925): 367, 372 (Egypt, A, L
keys); Parr, 1943: 247-251 (Syria ?, Lebanon, L bionomics, A, P, L keys); Lewis,
1945: 23 (Sudan, L key); Theodor, 1952: 113 (Middle East, zoogeogr.); Dow,
1953: 689 (Iran, toll. rec., L bionomics); Gad, 1956: 136 (Egypt, distr., bionomic
note); Reuben, 1969: 650 (India, A key, M*, F’, L’); Margalit et al., 1971: 323
(Israel, bionomics); El-Said and Kenawy, 1983a (Egypt); Zaini et al., 1983: 117
(Iraq, bionomic note); Kitron and Pener, 1986 (Israel, L bionomics).
Culex (Culex) thalassius Theobald of ? Abdel-Malek, 1960: 113-l 23 (Syria, L toll. sites,
bionomics).
Culex (Culex) tritaeniorhynchus Giles of Edwards, 1921: 339 (Iraq, Israel, Jordan, A, L
keys, tax., distr.); Edwards, 1926: 132 (Iraq, Israel, Jordan, A, L keys, A, L, distr.);
Barraud, 1934: 404 (India, A, L keys, M*, F, L’, syn., L habitat, distr.); Seguy,
1924: 32, 186 (Iraq, Israel, Jordan, A key, M, F, L, tax., distr., syn.); Kirkpatrick,
1925: 111 (Egypt, A, P, L keys, syn., M*, F, P*, L’, distr., bionomics);
Stackelberg, 1927: 157 (Middle East, M, F keys, A, M gen.*, distr.); Martini, 1931:
387 (Egypt, Iraq, Israel, Jordan, Turkey, A, L keys, M*, F, L’, distr.); Edwards,
1941: 299, 282 (Afrotropical Region, A key, M, F*, P, distr.); Monchadskii, 1951:
277 (Middle East, L key, L*, L assoc., distr.); Hopkins, 1952: 286 (L key, L,
bionomics); Knight, 1953a: 232 (Yemen Arab Republic, toll. rec., L habitat,
distr.); Khattat, 1955: 165 (Iraq, L*, L habitat, distr.); Mattingly and Knight, 1956:
104-137 (Oman, P.D.R. Yemen, Saudi Arabia, Socotra, Yemen Arab Republic, A,
L keys, L*, distr., bionomics); Parrish, 1959: 266 (Turkey); Senevet and Andarelli,
1959: 142 (North Africa, A, P, L keys, syn., M’, F, P*, L*, distr., bionomics, med.
imp.); Abdel-Malek, 1960: 113-124 (Syria, L, bionomics); Khalaf, 1962: 52 (Iraq,
M, F, L, bionomics); Derwesh, 1965: 44 (Iraq); DuBose and Curtin, 1965: 352,
354 (Mediterranean area, A, L keys); Abul-hab, 1966: 281 (Iraq, distr., L
bionomics); Abul-hab, 1968: 245 (Iraq, L key, L habitat, distr.); Lotfi, 1970: 402
(Iran, toll. rec.); Margalit and Tahori, 1970b: 153 (Israel, toll. sites); Aslamkhan,
1971: 155 (Pakistan); Lotfi, 1973: 206 (Iran, toll. sites); Gutsevich et al., 1974:
382 (Middle East, M, F, L keys, M”, F’, L*, distr., bionomics, med. imp.); Margalit
and Tahori, 1974: 88 (Israel, toll. rec.); Lotfi, 1976: 73, 76, 82 (Iran, L key, ecol.
note, L’); Sirivanakarn, 1976: 129 (Oriental Region, A, P, L keys, syn., M’, F*, P*,
L*, tax., distr., bionomics, med. imp.); lbrahim et al, 1983: 91 (Iraq, L*, key);
Harbach, 1985a: 86, 97, 102 (SW Asia, Egypt, distr., A, L keys).
Adult. A rather drab species resembling sitiens in most respects, but readily
distinguished by its smaller size, shorter cell Rl of the wing, lack of speckling on the foreand midfemora, and the presence of ventral pale scaling proximal to the median pale ring
of the proboscis.
FEMALE. Head: Antenna dark, pedicel yellowish to light brown; length 1.2-l .9
mm, mean 1.5 mm. Proboscis dark-scaled with narrow median pale ring, ring 0.15-0.20
proboscis length, proximal margin of ring about 0.45 from base; proximal part of
proboscis usually with scattered pale scales above and proximal extension of median
pale ring below (extension of ring often weak or absent in middle leaving an isolated spot
108
Contrib.Amer. Ent. Inst.,vol. 24, no. 1, 1988
proximal to ring), sometimes giving appearance of a second pale ring; labella yellowish;
length 1.4-l .7 mm, mean 1.6 mm. Maxillary palpus dark-scaled, pale scales at apex of
palpomere 4; length about 0.3 mm, about 0.2 length of proboscis. Forked scales of
vertex dirty yellow to brown, darker posteriorly and laterally, nearly linear anteriorly;
falcate scales pale yellow, long and relatively narrow; lateral spatulate scales yellowish
white, rather broad above, narrower below. Thorax: Integument reddish brown to
brown. Scutum with shiny golden-brown falcate scales, with paler scales on anterior
promontory, prescutellar area and often on lateral margin of fossa and supraalar area.
Scales of scutellum same as prescutellar scales. Antepronotum with some pale to dirty
yellow falcate scales confined largely to anterior area, with 2 more or less distinct rows of
setae; postpronotum with fine falcate scales more or less same color as scutal scales,
with 3-5 setae in vertical row on posterior margin. Pleura with pale scales as follows:
proepisternum with small patch of narrow spatulate scales below and mesal to upper
proepisternal setae; mesokatepisternum with rather small patches of spatulate scales on
upper corner and lower posterior border at level of katepimeron; mesepimeron with small
anterior patch of spatulate scales adjacent to upper katepisternal scales, and sometimes
with few near linear or fusiform scales among upper mesepimeral setae. Pleural setae:
5-7 upper proepisternal, 4-9 prealar, 3 or 4 upper mesokatepisternal, 7-10 lower
mesokatepisternal, 5-9 upper mesepimeral, and no lower mesepimeral. Wing: Length
2.6-3.4 mm, mean 2.9 mm; length of cell R2 2.6-3.4 length of vein R2+3, mean 2.9;
length of cell Ml 0.8-0.9 length of cell R2; dark-scaled with short line of pale scales at
base of costa, pale line often weakly developed or absent; dorsal surface with spatulate
scales on costa, subcosta, R and Rl, R3, MS+4 and CuA, dorsal surface of other veins
with long linear scales; ventral surface with linear scales on all veins except costa and
subcosta. Halter: Pale with distal part of scabellum and base of capitellum usually
dark-scaled, capitellum sometimes largely dark-scaled. Legs: Forecoxa always with pale
scales at base, rest of scales either entirely dark, dark proximally and pale distally, or with
dark and pale scales mixed, frequently with few narrow pale scales posterolaterally at
apex; midcoxa with vertical patch of pale scales on anterior side of lateral midline and
small patch of dark scales distally on anterior surface; hindcoxa with vertical line of
indistinct pale scales on anterior side of lateral midline. Trochanters each with small
patch of dark scales on posteroventral surface. Forefemur largely dark-scaled, poorly
contrasted but darker anteriorly than posteriorly, with narrow rather indistinct knee spot;
midfemur dark-scaled anteriorly, pale-scaled posteriorly and ventrally, dark and pale
areas better contrasted than on forefemur, dark area forming narrow band at apex, with
very indistinct knee spot; hindfemur pale-scaled with variable, often indefinite,
anterodorsal dark stripe that abruptly widens distally to form more or less distinct narrow
apical band, with very narrow knee spot. Foretibia dark-scaled dorsally (darker than
femur), pale-scaled ventrally, with narrow dorsal pale spot at apex; midtibia dark-scaled
anteriorly, pale-scaled posteriorly, with faint indication of pale scaling at apex, particularly
on anterior surface; hindtibia dark-scaled dorsally, pale-scaled ventrally, with indistinct
dorsal pale spot at apex and indication of dorsal pale spot at base. Tarsi with narrow,
indistinct basal and apical pale rings on tarsomeres 1-4; if pale ring present on apex of
midtarsomere 4, then pale ring usually also evident at base of midtarsomere 5; rings
between midtarsomeres 3-4 and 4-5 often absent. Abdomen: Tergum I with median
posterior patch of dark scales; terga II-VII with narrow, slightly convex basal pale bands
(0.15-0.35 tergum length) and basolateral pale spots that become larger in progression
from anterior to posterior terga, tergum VII often with median posterior patch of pale
scales; tergum VIII largely pale-scaled with narrow apical dark band. Sterna pale-scaled,
frequently with dark scales on posterolateral corners; sternum VIII with lateral patches of
Harbach: Subgenus Culex in SouthwesternAsia
109
pale scales, without scales medially.
MALE. Like female except as follows. Head: Proboscis with ventral cluster of
setae on proximal side of false joint. Maxillary palpus extending beyond tip of proboscis
by at least length of palpomere 5, length 1.8-2.8 mm, mean 2.4 mm; palpomere 3 with
subapical stripe of pale scales on dorsal and lateral surfaces, often only subapical line on
lateral surface, ventromesal margin with short setae in row extending length of pale
scaling, ventral surface with long setae distal to pale scaling; palpomeres 4 and 5 each
with narrow basal ring of pale scales and numerous long ventrolateral setae, basal rings
actually very narrowly broken on lateral and mesal margins by row of setae, palpomere 5
also usually with some pale scales dorsally at apex. Forked scales of vertex broader and
somewhat shorter. Wing: Length 2.2-3.2 mm, mean 2.8 mm; length of cell R2 1.6-2.7
length of vein R2+3, mean 2.2; scaling same as in female but scales smaller and less
dense, spatulate scales not as short and broad. Abdomen: Basal pale bands of terga
generally slightly broader, 0.22-0.40 tergum length. Genitalia (Fig. 39): Form as figured;
very much as in sitiens and mimeticus. Ninth tergal lobe with row of 3-10 (mode 6) setae.
Gonocoxite with 3 or 4 rows of lateral setae extending from just above subapical lobe to
point about midway between subapical lobe and base of gonocoxite; subapical lobe with
3 setae in group d-e, all flattened and hooked, 2 with subapical denticles, f flattened and
broad, broadest distally. Inner division of phallosome with 3-5 (mode 4) caudally curved
teeth. Proctiger not unusual; paraproct with slightly larger basal lateral arm, smaller
crown, and well developed ventral acetabulum. Cereal sclerite usually with 2 setae (l-4)
on posterolateral corner.
Pupa (Fig. 39). Form and placement of setae as figured, range and modal
number of branches in Table 19; different from other members of the sitiens group in
southwestern Asia in having setae 8-CT and l-11 more branched. Cephabthorax: Lightly
tanned. Setae 1-3-CT usually triple (see table); 4,7,11 -CT double, 4-CT rarely single;
8-CT shorter and more branched than usual, same length as g-CT, with 6 or 7 branches;
g-CT usually double (2,3); 12-CT longer than 10,l l-CT, usually with 4 or 5 branches
(3-5). Trumpet: Dark and rather short; index 5.88-6.25, mean 6.04; pinna 0.2-0.3
trumpet length. Abdomen: Lightly tanned; length 2.19-2.42 mm, mean 2.32 mm. Seta
7-l usually with 3 or 4 branches (2-5), 7-11usually double or triple (2-4); l-II-VII and 6-III-VI
tend to be more branched than usual, especially l-11 (see table); l-III-VII about length of
tergum following; 5-IV multiple, different in appearance from 5-V,VI which are double
with heavier and longer branches; 6-V,VI developed like 6-lll,lV but distinctly longer;
4-VIII with 3 or 4 branches. Paddle: Pale; length 0.66-0.77 mm, mean 0.70 mm, width
0.46-0.54 mm, mean 0.51 mm, index 1.3-1.5, mean 1.4.
Larva (Fig. 40). Character and positions of setae as figured, range and modal
number of branches in Table 39; resembling members of the pipiens group, but
differing conspicuously in the character of seta 1-C and the pecten spines; more closely
resembling Miens than other members of the sitiens group occurring in southwestern
Asia, but easily distinguished by the characters given in the key. Head: Length
0.68-0.78 mm, mean 0.72 mm; width 1.03-1.11 mm, mean 1.07 mm; lightly to
moderately tanned, darker behind eyes, collar moderately to heavily tanned, dorsal
apotome with variable number of moderately tanned spots distributed somewhat as in
pipiens. Hypostomal suture complete, extended to collar from posterior tentorial pit.
Dorsomentum with 6-9 teeth (mode 7) on either side of center. Seta 1-C tapered,
sharply pointed, dark; 2-C absent; 5-C normally triple (3,4); 6-C normally double (2,3);
7-C with 5-10 branches, about as long as 5,6-C; 10,12,13-C usually triple (2-4); 11-C
double. Antenna: Length 0.52-0.58 mm, mean 0.55 mm; tanning and spiculation as in
sitiens. Seta 1-A with about 24 branches (21-35); 2,3-A subapical. Thorax: Integument
110
Contrib.Amer. Ent. Inst.,vol. 24, no. 1, 1988
normal. Seta 4-P double; 7-P normally triple (2,3), slightly longer than 4-P; 8-P double,
weaker than 4-P. Seta 1-M single, longer than 2-M; 2-M multiple (4-8); 4-M usually triple
(2,3); 5-M shorter than usual. Seta 1-T single, not more than 0.5 length of 2-T; 2-T
double or triple; 13-T with 7-l 1 branches, not much longer than 12-T. Abdomen:
Integument normal. Seta 3-l,ll,lV usually triple (2,3), 3-I,11 rarely with 4 branches; 3-111
commonly double (2,3), 3-V single or double, 3-VII with 4-6 branches, often with 5; 6-I,11
normally triple (2,3), 6-11shorter than 6-l; 6-III-VI usually double (2,3), 6-lll,VI same length
as 6-11,6-IV,V shorter; 7-l most often double (l-3); l-III-VII about 0.5 length of segment,
l-111most often triple (2-4), 1-IV,V with 4 branches, l-VI most often with 4 branches (3-6),
l-VII usually with more than 4 branches (4-7); 13-111-Vresemble l-Ill-V but usually with
more branches (see table). Segment VIII: Comb with 26-60 (mode 38) evenly fringed
scales. Siphon: Index 5.77-7.00, mean 6.30; moderately tanned, slightly darker at
apex, basal ring and acus darker. Pecten of 12-18 (mode 14) spines with ventral row of
denticles, larger spines with 7-12 denticles. Seta 1-S normally in 6 pairs, occasionally in
5 or 5.5 pairs, 1 subapical pair (le-S) borne laterally, others borne posterolaterally, 3
proximal pairs (1 a,b,c-S) commonly with 4 branches (2-6), 3 distal pairs (ld,e,f-S) most
often triple (2-4), only slightly shorter than proximal pairs, all a little longer than diameter
of siphon at point of attachment. Segment X: Saddle complete; lightly to moderately
tanned, darker posterodorsally; with conspicuous spicules posteriorly; length 0.29-0.36
mm, mean 0.33 mm, siphon/saddle index 4.23-6.14, mean 4.73. Seta 1-X often triple
(2-4); 2-X usually with 3 or 4 branches, seldom double; 4-X usually in 6 pairs, an
additional unpaired seta occasionally present. Anal papillae elongate, subacutely
tapered, about as long as saddle.
Systematics.
This and the next species are members of the complicated
vishnui subgroup of Bram (1967b). The group is almost wholly Oriental, but some
members also occur in adjacent areas. Culex tritaeniorhynchusand pseudovishnui are
the only members of the group which occur in southwestern Asia.
Culex
tritaeniorhynchusis common throughout the area and many coastal countries of Africa.
Culex tritaeniorhynchus should not be confused with any other species in
southwestern Asia except pseudovishnui. It is readily differentiated from this species by
the development of the comb scales and pecten spines of the larva. The pupa differs in
the character of setae 8-CT and l-11 and the posterolateral corners of tergum VIII. There
is considerable overlap in the adults, but specimens of tritaeniorhynchus usually can be
recognized relatively easily by the more extensive ventral pale scaling of the proboscis.
The male genitalia are very nearly congruent, differing essentially only in the size of the
teeth of the inner division of the phallosome.
Culex tritaeniorhynshus is a peculiar species which, as pointed out by
Sirivanakarn (1976), is similar to members of the barraudi subgroup in the larval stage.
This situation parallels that of poicilipes and whitmoreiwhich are included as members of
the sitiens subgroup in spite of conspicuous larval disparity. It is apparent that the
present system of grouping species of the Miens group does not accurately reflect
species affinities based on all life stages. This group more than any other in the
subgenus Culex brings to mind Belkin’s (1962) contention that hybridization seems to
have been an important mechanism of speciation in mosquitoes.
Bionomics.
Culex tritaeniorhynchus is a common species in southwestern
Asia. The immature stages are typically found in rice fields, flood waters, and marshy
areas with floating or emergent vegetation. They also are found in ponds, swamps,
streams, springs, irrigation ditches, grassy pools, seepages, and animal hoofprints. The
water may be fresh or slightlybrackish but is always clear and clean.
Females of tritaeniorhynchus are known to enter dwellings and bite man during
Harbach: Subgenus Culex in SouthwesternAsia
111
any time at night (Kirkpatrick, 1925; Khalaf, 1962). In Iran and Iraq, tritaeniorhynchus is
considered to be a vector of Japanese encephalitis virus (Travis and Labadan, 1967).
The susceptibility and transmission efficiency for West Nile virus in the laboratory
indicate that tritaeniorhynchus is an excellent potential vector of this pathogen in
Pakistan (Hayes et al., 1980). A single isolation of Sindbis virus was made from a pool of
specimens collected at Al Khobar, Saudi Arabia (Wills et al., 1985). This species has
been associated with various viruses and filarial pathogens in many areas of eastern and
southeastern Asia.
Distribution.
Culex tritaeniorhynchus occurs in eastern Asia, the Oriental
Region, southwestern Asia, and Africa.
Material examined. 570 specimens. A total of 536 specimens (221 females,
137 males, 69 male genitalia, 23 pupal exuviae, 40 larval exuviae, and 46 fourth-instar
larvae) were examined from southwestern Asia - IRAN: (Aliabad, Basra, Cham Asbi,
Chomaisaray, Hassan Kiadeh, Kanroud, Karken River, Sabzab, Shour River, Zahedan,
Zarjub); IRAQ: Basrah, Galala, Magil, Rawasnduz, Tal A’far, Thallal); ISRAEL: (Beisan,
Galilee, Khirbet Hadrah, Neot Hokikar);JORDAN: (Yabes); OMAN: (Salalah); PAKISTAN
(including localities east of the lndus River): (Balloki, Kasur, Khulna, Lahore, Peshawar);
P.D.R. YEMEN: (Bateis, Dis Town, Hami, Mukalla, Urfat Subai); SAUDI ARABIA: (Alajam,
Al Khobar, Al Qatif, Dhahran, Saihat); TURKEY: (Adana, Silifke). An additional 34
specimens (19 females, 6 males, 5 male genitalia, and 4 fourth-instar larvae) were
examined from BANGLADESH, ETHIOPIA, INDIA (the holotype female), KENYA,
SOCOTRA, and unknown localities.
Colex
(&flex)
pseudovishnui
C 0 I Ie s s
pseudovishnui Colless, 1957: 88. Holotype female: Singapore (BM).
neovishnui
Lien, 1968: 230.
Holotype male: Peiyuan, Tungho,
Taitung, Taiwan (TMRI).
Synonymy with pseudovishnui by
Sirivanakarn, 1976: 116.
Culex pseudovishnuiColless of Reuben, 1969: 648 (India, M’, F’, L’, A key); Zaim and
Cranston, 1984: 179 (Iran, L, toll. sites).
Culex (Culex) pseudovishnui Colless of Aslamkhan, 1971: 155 (Pakistan); Sirivanakarn,
1976: 116 (Oriental Region, M*, F, P*, L’, syn., distr., bionomics, med. imp.);
Harbach, 1985a: 86, 97, 107 (Iran and Pakistan only, distr., A, L keys).
Culex VishnuiTheobald of Barraud, 1923: 938 (India, L*); Barraud, 1924a: 996 (in part;
India, A key, syn., M*, F, distr.).
Culex (Culex) VishnuiTheobald of Edwards, 1921: 339 (Iraq, A key, bionomics, distr.);
Edwards, 1926: 132 (Iraq, A, L, keys, A, distr.); Seguy, 1924: 30, 186 (Iraq, A
key, M, F, L, tax., distr., syn.); Stackelberg, 1927: 158 (in part; S. Asia, M, F keys,
A, M gen.*, distr.); Martini, 1931: 392 (in part; S. Asia, A, L keys, M*, F, L*);
Barraud, 1934: 400 (in part; India, A, L keys, M*, F, L*, syn., L habitat, distr.); ?
Monchadskii, 1951: 284 (in part; S. Asia, L key, L*, bionomic note, distr.); Khalaf,
1962: 55 (Iraq, A, M gen., L); Abul-hab, 1968: 246 (Iraq); Lotfi, 1973: 206 (Iran,
bionomics, distr., toll. sites); Gutsevich et al., 1974: 371 (in part; S. Asia, L key);
Lotfi, 1976: 72, 78, 79 (Iran, L key, ecol. note, L’).
Adult. A small brown species closely resembling tritaeniorhynchus, but the
median pale ring of the proboscis is not extended proximally on the ventral surface and
112
Contrib.Amer. Ent. Inst., vol. 24, no. 1, 1988
there is often an indefinite anterior pale stripe on the hindtibia. The proboscis of the
male may have one or two short setae at the base of the median pale ring but is without a
distinct ventral cluster of setae.
FEMALE. Head: Pedicel of antenna yellowish laterally, mesal surface dark brown
with some tiny inconspicuous scales; flagellum normal, dark, length about 1.8 mm.
Proboscis length 1.5-l .8 mm, mean 1.7 mm; dark brown to blackish-scaled with median
pale ring of cream-colored scales 0.40-0.65 from base. Maxillary palpus dark-scaled with
few pale scales at apex; length about 0.3 mm, approximately 0.2 proboscis length.
Vertex with whitish to pale golden falcate scales, paler along margin of eye, sometimes
darker laterally; forked scales white to yellow in middle, dark brown to black
posterolaterally; lateral spatulate scales white, or nearly so. Interocular space with few
scales, same color as ocular scales. Ocular setae golden brown to brown, often paler
near middorsal line of cranium; 2 golden interocular setae project ventrally over clypeus.
Thorax: Scutal integument brown; pleural integument yellowish brown or paler, with
dark brown areas on proepimeron, anterior margin of mesokatepisternum, postspiracular
area, prealar area, between upper and lower mesokatepisternal scale-patches, below
anterior mesepimeral scales and on upper portion of mesepimeron. Scutal scales rather
long and narrow, usually predominantly pale, color ranging from silvery white to golden,
frequently pale golden with whitish scales along margins and on prescutellar area; darker
scales, when present, either indistinct or distinct, occurring on acrostichal, median scutal
fossal and/or posterior dorsocentral areas, usually golden brown. Scutal setae dark
brown with reddish and/or golden sheen. Lobes of scutellum with scales resembling
prescutellar scales; each lateral lobe with 3-5 large setae, median lobe with 5-7.
Antepronotum with brown setae, some paler, often yellowish, ventrally; with whitish to
pale golden falcate scales, coarser and paler ventrally. Postpronotum with whitish to
pale golden falcate scales on about dorsal 0.5, slightly coarser and paler posteriorly;with
5 or 6 dark setae on posterodorsal margin. Pleural setae yellow to brownish yellow: 4-6
proepisternal, 8 or 9 prealar, 4 or 5 upper mesokatepisternal, 7 or 8 lower mesokatepisternal, 4 or 5 upper mesepimeral; lower mesepimeral seta absent. Pleura with the
usual patches of elongate white spatulate scales: small patch below proepisternal setae,
patches on upper corner and lower posterior margin of mesokatepisternum, anterior
patch on mesepimeron at level of upper patch on mesokatepisternum, and small patch
before upper mesepimeral setae; upper proepisternal and upper mesepimeral patches
usually with only 2 or 3 scales, or sometimes absent. Wing: Length 2.8-3.2 mm, mean
3.0 mm; length of cell R2 1.5-2.8 length of vein R2+3, mean 2.3; length of cell Ml
0.8-0.9 length of cell R2; scales predominantly dark, with variable amount of pale scaling
on costa, subcosta, R and RI, pale scales always present on posterior side of costa near
humeral crossvein. Halter: Integument and scales entirely pale. Legs: Anterior surface
of forecoxa entirely white-scaled or with variable amount of black scaling in middle, most
often with black scales; midcoxa with longitudinal patch of white scales on anterior side
of midlateral row of strong setae; hindcoxa with longitudinal patch of white scales on
anterolateral surface, posterolateral surface with irregular row of strong setae. Lateral
surfaces of trochanters without scales, other surfaces white-scaled, anterior surface of
foretrochanter sometimes with dark scales. Apices of all femora with narrow white knee
spots; anterior surface of forefemur brown-scaled, posterior surface white-scaled;
midfemur like forefemur but brown scaling gradually expanded over dorsal surface
toward apex and white scaling sometimes expanded over ventral surface onto lower part
of anterior surface; hindfemur mainly white-scaled, with anterodorsal stripe of brown
scales, stripe gradually spreading over dorsal surface distally and then abruptly
expanded onto anterior and posterior surfaces near apex. Foretibia whitish-scaled with
Harbach: Subgenus Culex in SouthwesternAsia
113
complete dorsal stripe of brown scales; midtibia mainly whitish-scaled, ventral surface
with complete stripe of brown scales, dorsal surface with narrow, usually obscure brown
stripe that is largely obsolescent in middle; hindtibia mainly brown-scaled, narrowly pale
at base and apex, often with ill-defined anterior and posterior stripes of whitish scales
usually not reaching ends of tibia and normally joined ventrally for most or part of their
length. Tarsi with narrow pale basal bands on tarsomeres l-3, frequently on 4, and
occasionally on 5; tarsomeres 1 and 2 also frequently with very narrow pale apical bands;
posterior surface of tarsomere 1 usually pale-scaled. Abdomen: Tergum I with
posteromesal patch of blackish scales; terga II-VII mainly clothed in dark brown to
blackish scales, with basolateral patches of white scales, tergum II with basomedian spot
of pale scales; terga Ill-VII with straight or convex basal bands of white or nearly white
scales, bands usually about 0.25 tergum length but often narrower and sometimes not
reaching basolateral pale patches, particularly on more anterior terga; tergum VIII usually
with broad basal pale band and narrow apical dark band. Sterna II-VII mainly pale-scaled
with large apicolateral dark patches, dark patches sometimes more or less joined mesally,
particularly on more posterior sterna; sternum VIII with lateral pale patches, broad median
area devoid of scales.
MALE. Resembling female except for following sexual differences. Head:
Proboscis with narrow pale band about 0.50 to 0.65 or 0.70 from base, sometimes with 1
or 2 long setae ventrally at base of band; false joint inconspicuous. Maxillary palpus with
bands of pale scales and/or pale integument on about distal 0.5 of palpomere 2, middle
of palpomere 3, bases of palpomeres 4 and 5 and apex of palpomere 5; ventrolateral
margin of palpomere 3 with lo-20 long dark setae at apex; ventromesal margin with
complete row of short, translucent, antrorsely curved, scalelike setae that increase
progressivly in length from base to apex of palpomere; lateral and mesal surfaces of
palpomeres 4 and 5 densely setose; longer than proboscis by length of palpomere 5
and part of 4. Thorax: Scutal scales finer, more uniformly colored. Wing: Length of cell
R2 1.5-2.1 length of vein R2+3, mean 1.8; subcosta intersects costa before furcation of
R2+3. Abdomen: Terga II-VII without basolateral pale patches; basal bands of terga Ill-VII
broader, usually 0.30-0.45 tergum length, bands usually straight or nearly so, those of
terga VI and VII produced posteriorlyalong lateral scale-free areas; tergum VII sometimes
with pale scales apically in middle; tergum VIII (ventral in position) with approximately
basal 0.5 pale-scaled. Sterna II-VII with dark apical bands that are broadest in middle,
sometimes with few pale scales centrally on posterior border; sternum VIII (dorsal in
position) like tergum VIII of female but sometimes with pale scales along posterior
margin. Genitalia (Fig. 41): Form as figured; constructed as in sitiens, mimeticus, and
tritaeniorhynchus. Ninth tergal lobe with row of 3-7 (mode 4) short setae. Gonocoxite
and phallosome closely resembling those of tritaeniorhynchus, inner division of
phallosome differs in having distinctly larger teeth (4 or 5, mode 5) and the dorsocaudal
angle usually notched. Proctiger as in tritaeniorhynchus; cereal sclerite with 2 or 3,
usually 2, setae on posterolateral corner.
Pupa (Fig. 41). Character and positions of setae as figured, range and modal
number of branches in Table 20; similar to tritaeniorhynchus, distinguished by setae
8-CT and l-11 with fewer branches and posterolateral angle of tergum VIII acute.
Cephalothorax: Lightly tanned. Seta 2-CT usually with 3 or 4 branches, more often with
4 (3-5); 8-CT with 3-5 branches, distinctly longer than g-CT; 12-CT about as long as
lo,1 1-CT, usually triple (2-5). Trumpet: Moderately tanned, tracheoid area darker; index
5.45-8.10, mean 6.61; pinna 0.15-0.35 trumpet length. Abdomen: Lightly tanned;
length 2.34-2.96 mm, mean 2.69 mm. Seta 7-l usually double (2-4); I-II-VII distinctly
shorter than tergum following; 6-III-VI commonly with 4 or 5 branches (see table), all
114
Contrib.Amer. Ent. Inst.,vol. 24, no. 1, 1988
about same length; 5IV shorter and W,VI about the same as in tritaeniorhynchus;4-VIII
usually double or triple (2-4). Paddle: Very lightly tanned; length 0.72-0.83 mm, mean
0.75 mm, width 0.47-0.56 mm, mean 0.53 mm, index 1.4-1.6, mean 1.4.
Larva (Fig. 42). Form and placement of setae as figured, range and modal
number of branches in Table 40; distinct from sitiens, tritaeniorhynchus, and members
of the pipiens group in southwestern Asia in the character of the comb scales, easily
distinguished from other members of the sitiens group in the region by the characters
given in the key. Head: Length 0.71-0.88 mm, mean 0.76 mm; width 1.08-l .22 mm,
mean 1.16 mm; lightly tanned, darker posteriorly. Dorsomentum heavily tanned; teeth
large and distinct, 5-7 (mode 6) on either side of center. Seta 1-C long, relatively
slender, tapered, dark; 2-C absent; 5-C usually triple (2-4); 6-C usually double (l-3); 7-C
often with 8 or 9 branches (8-l 2), about as long as 5,6-C; 1O-C with 3 or 4 branches on
short stem; 11,13-C usually double (l-3); 12-C frequently double but often more
branched (2-5). Antenna:
Length 0.60-0.71 mm, mean 0.66 mm; tanning and
spiculation as usual. Seta 1-A with about 30 branches (27-30); 2,3-A distinctly
subapical. Thorax: Integument hyaline, with rather distinct thumbprint pattern, lateral
surfaces and dorsal surface anteriorly with minute spicules. Seta 4-P normally double
(l-3) nearly as long as 7-P; 7-P triple; 8-P normally double (1,2). Seta 1-M single or
double in nearly equal frequencies; 4-M usually triple (2,3). Seta 1-T single or double,
much shorter than 2-T; 2-T double or triple; 13-T often with 9 or 10 branches (7-12), not
much longer than 12-T. Abdomen: Hyaline, thumbprint pattern evident. Seta 3-l
variable, with 3-7 branches, most often with 3; 3-II-IV usually double or triple, 3-11,111
occasionally with 4 branches; 3-V usually single (1,2); 3-VII with 3-7 branches; 6-I,11triple,
6-III-VI usually triple (see table), 6-l distinctly longer than the others, 6-lll,lV not much
shorter than 6-ll,V,VI; 7-l single; l-Ill-VII slightly longer than 0.5 length of tergum, 1-III,IV
frequently with 5 or 6 branches, 1-V,VI often with 6 or 7 and l-VII often with 7 or 8 (see
table). Segment VIII: Comb with 5-10 (mode 6) scales developed as in poicilipes and
bitaeniorhynchus. Siphon: Index 4.41-6.87, mean 5.40; lightly tanned, basal rim dark;
surface with rows of minute vesicles; very slightly bent forward before apex. Pecten
composed of 8-12 (mode 11) large distally curved spines with complete ventral row of
denticles. Seta 1-S in 6.5, 7 or 7.5 pairs, middle pair (1 d-S) borne laterally, others borne
posterolaterally; most proximal pair (la-S) 1.5-2.0 times as long as diameter of siphon at
point of attachment, others gradually shorter toward tip of siphon, most distal pair (lg-S)
no longer than diameter of siphon at point of attachment. Segment X: Saddle lightly
tanned, with prominent spicules on posterodorsal margin; length 0.36-0.42 mm, mean
0.38 mm, siphon/saddle index 3.13-4.29, mean 3.61. Seta 1-X with 3 or 4 branches;
2-X usually triple (2,3); 4-X in 6 pairs, often with 7-9 branches. Anal papillae rather
slender, longer than saddle.
Systematics.
This species has been confused in the past with Culex vishnui
Theobald, 1901 a, which is mainly restricted to the Oriental Region and occurs no farther
west than eastern India. Although pseudovishnui closely resembles vishnui, it is
definitely a distinct species which is well differentiated in the larval stage. Zaim and
Cranston (1984) corrected the previous record of vishnui in Iran on the basis of larval
characters. All life stages from populations in southwestern Asia seem to agree in all
essential respects with pseudovishnui in the Oriental Region, but too few specimens
are currently available for a detailed comparative study.
To my knowledge, pseudovishnui has never been recorded from Afghanistan.
My previous report (Harbach, 1985a) was in error, but the species probably occurs there
because it has been collected very near its borders in both Iran and Pakistan.
Culex pseudovishnui is very similar to tritaeniorhynchus in the adult stage, and
Harbach: Subgenus Culex in SouthwesternAsia
115
the two forms are often difficult to differentiate. The most reliable characters that will
separate the females of these forms are the less extensive pale scaling of the proboscis
and the shorter vein R2+3 in pseudovishnui. The male genitalia cannot be differentiated
with absolute certainty even though the teeth of the inner division of the phallosome are
definitely larger in this species. The larvae are strikingly different in the form of the comb
scales and pecten spines, and the pupae differ in the development of setae 8-CT and
I-II and in having more acute posterolateralcorners on tergum VIII.
Bionomics.
This species breeds in permanent standing water and transient
ground pools. Larvae are commonly found in rice fields, ditches, stream margins,
marshes, and ponds in association with tritaeniorhynchus and bitaeniorhynchus.
Females are attracted to birds, pigs, and other domestic animals and occasionally bite
man (Colless, 1959; Reuben 1971a,b). Culex pseudovishnui is a potential vector of
Japanese encephalitis virus and has been associated with Kunjin and Tembusu viruses
in Southeast Asia (Karabatsos, 1985).
Distribution. Culex pseudovishnui is widely distributed in the Oriental Region
and occurs in Japan, Korea, China, Pakistan, and Iran.
Material examined. 73 specimens. Only 38 specimens (6 females, 7 males, 7
male genitalia, 2 pupal exuviae, 7 larval exuviae, and 9 fourth-instar larvae) were
examined from southwestern Asia - IRAN: (Shemian, Zahedan); PAKISTAN (including
localities east of the lndus River): (Kohat, Lahore, Peshawar, Sargodha, Wahga).
Another 35 specimens (17 females, 10 males, 5 male genitalia, and 3 pupal exuviae)
were examined from INDIA.
Contrib.Amer. Ent. Inst.,vol. 24, no. 1, 1988
116
REFERENCES
CITED
(in the text and appendices)
Abbott,
P.H.
1948. The Culicidae (Diptera) of Darfur Province, Angloegyptian Sudan, with
observations on the geography and zoogeographical relations of the region.
Proc. R. Entomol. Sot. Lond. Ser. B Tax. 17: 37-48.
Abdel-Malek,
A.
1956. Mosquitoes of north-eastern Sinai. Bull. Sot. Entomol. Egypte 50: 97-107.
1960. The culicine mosquitoes of the northern region of the United Arab Republic.
Bull. Sot. Entomol. Egypte 44: 11 l-l 28.
Abul-hab,
J.
1966. Larvae of culicine mosquitos in north Iraq (Diptera, Culicidae). Bull. Entomol.
Res. 57: 279-284.
1968. Larvae of culicine mosquitoes of Iraq with a key for their identification. Bull.
Endem. Dis. 10: 243-256.
Aiken, J. and E.D. Rowland
1906. Preliminary notes on the mosquitoes of British Guiana. Br. Guiana
Ann. 1905: 13-38.
Med.
Anderson,
D.
1967. Ecological studies on Sindbis and West Nile viruses in South Africa. III. Host
preferences of mosquitoes as determined by the precipitin test. S. Afr. J. Med.
Sci. 32: 34-39.
Aslamkhan,
M.
1971. The mosquitoes of Pakistan I. A checklist. Mosq. Syst. News Lett. 3:
147-l 59.
Austen,
E.E.
A contribution to knowledge of the blood-sucking Diptera of Palestine, other
than Tabanidae. Bull. Entomol. Res. 12: 107-l 24, 1pl.
1921.
Bailly-Choumara,
H.
1960. Une espece nouvelle d’Anophele du golfe d’Aden, Anopheles (Myzomyia)
azaniae (Diptera, Culicidae); notes morphologiques, systematiques et ecologiques. Bull. Sot. Pathol. Exot. 53: 531-542.
1968. Contribution a I’etude des moustiques du Maroc (Diptera, Culicidae) six
especes nouvelles pour le pays. Cah. ORSTOM Ser. Entomol. Med.
6:139-144.
Harbach: Subgenus Culex in SouthwesternAsia
117
Baisas, F.E.
1938. Notes on Philippine mosquitoes, VII. A - Culex (Culex) with banded
proboscis and tarsi. B - Anopheles: The pupae of three rare species; the
leucosphyrus subgroup. Mon. Bull. Bur. Health Philipp. Manila 18: 175-232,
18 pl.
Barnett, H.C. and S. Toshioka
1951. The bloodsucking insects, mites and ticks of Korea and their relation to
disease transmission. 406th Med. Lab. 25 pp.
Barraud, P.J.
1920. Notes on some Culicidae collected in lower Mesopotamia. Bull. Entomol.
Res. 10: 323-325.
1921. Mosquitos collected in Palestine and adjacent territories. Bull. Entomol.
Res. 11: 387-395.
1923. A revision of the culicine mosquitoes of India. Part II. The larvae of some
Indian species of Culex. Indian J. Med. Res. 10: 934-942, 9 pl.
1924a. A revision of the culicine mosquitoes of India. Part Xl. Some Indian species
of Culex L. Indian J. Med. Res. 11: 979-998, 4 pl.
1924b. A revision of the culicine mosquitoes of India. Part XII. Further descriptions
of Indian species of Culex L. including two new species. Indian J. Med. Res.
11: 1259-1274, 3 pl.
1924c. A revision of the culicine mosquitoes of India. Part XVII. Further descriptions of the larvae of Indian species of Culex. Indian J. Med. Res. 12:
427-434, 5 pl.
1934. The fauna of British India, including Ceylon and Burma. Diptera. Vol. V.
Family Culicidae. Tribes Megarhinini and Culicini. Taylor and Francis, London.
xxviii + 463 pp., 8 pl., 1 map.
Barraud, P.J. and S.R. Christophers
1931. On a collection of anopheline and culicine mosquitoes from Siam.
Malaria Surv. India 1: 269-285.
Rec.
Becker, T.
1903. Agyptische Dipteren. Mitt. Zool. Mus. Berl. 3: 67-195, 4 pl.
1907(1910). Dipteren aus Siidarabien und von der lnsel Sokotra. Denkschr. Acad.
Wiss. Wien 71: 131-160.
Belkin, J.N.
1962. The mosquitoes of the South Pacific (Diptera, Culicidiae). 2 vols. University
of California Press, Berkeley and Los Angeles. xii + 608 pp. and 412 pp.
1968a. Mosquito studies (Diptera, Culicidae) VII. The Culicidae of New Zealand.
118
Contrib.Amer. Ent. Inst.,vol. 24, no. 1, 1988
Contrib. Am. Entomol. Inst. (Ann Arbor) 3(l): 1-182.
1968b. Mosquito studies (Diptera, Culicidae). IX. The type specimens of New
World mosquitoes in European museums. Contrib. Am. Entomol. Inst. (Ann
Arbor) 3(4): l-69.
Bigot, J.M.F.
1857. Dipteros, pp. 328-349. In: R. de la Sagra, Historia fisica, politica y natural de
la lsla de Cuba. Vol. 7. Paris.
1859. Dipteres de Madagascar.
115-135,
2 pi.
Premiere pat-tie. Ann. Sot. Entomol. Fr. 7:
1861. Trois dipteres nouvelleaux de la Corse.
227-229.
Ann. Sot. Entomol. Fr. (4) 1:
1888. Description des especes nouvelles. Expl. Scient. Tunisie Dipt. 1888: 7-l 1.
1889. Communications. -M. J.M.F. Bigot envoie de Quincy, une note sur une
nouvelle espece du genere Culex. Ann. Sot. Entomol. Fr. 9: cxxii.
Bram, R.A.
1967a. Contributions to the mosquito fauna of Southeast Asia. - II The genus
Culex in Thailand (Diptera: Culicidae). Contrib. Am. Entomol. Inst. (Ann Arbor)
2( 1): l-296.
1967b. Lectotype assignments for several species of the genus Culex in
Southeast Asia. Proc. Entomol. Sot. Wash. 69: 327-328.
Brunetti,
E.
1912. Annotated catalogue of Oriental Culicidae - Supplement. Rec. Indian Mus.
4: 403-517.
Bullini, L. and M. Coluzzi
1982.
Evolutionary and taxonomic inferences of electrophoretic studies in
mosquitoes, pp. 465-482. In: W.W.M. Steiner, W.J. Tabachnick, KS. Rai, and
S. Narang (eds.), Recent developments in the genetics of insect disease
vectors. Stipes Publishing, Co., Champaign, Illinois.
Bijttiker, W.
1981. Observations on urban mosquitoes in Saudi Arabia, pp. 472-479. In: W.
Wittmer and W. Buttiker (eds.), Fauna of Saudi Arabia. Vol. 3. Ciba-Geigy Ltd.,
Basle.
Buxton,
P.A.
1923. Applied entomology of Palestine, being a report to the Palestine government. Bull. Entomol. Res. 14: 289-340.
119
Harbach: Subgenus Culex in SouthwesternAsia
Callot, J.
1940. Sur quelques moustiques du Maroc. Arch. inst. Pasteur Maroc 2: 361-365.
1957. Sur Culex torrentium Martini. Ann. Parasitol. Hum. Comp. 32: 438-442.
Cambouliu,
M.
1902. Contribution a I’etude des Anopheles de I’isthme de Suez. C.R. Acad. Sci.
Paris 135: 704-706.
Carter, H.F.
1948. Records of filaria infections in mosquitoes in Ceylon.
Parasitol. 42: 312-321.
Ann. Trop. Med.
Christophers,
S.R.
1906. On the importance of larval characters in the classification of mosquitoes.
Sci. Mem. Med. Sanit. Dep. Gov. India No. 25, 18 pp., 3 pi.
1924. Some further varieties of Indian species of Anopheles with notes on the
species A. pallidus Theobald and A. philippinensis Ludlow. Indian J. Med. Res.
12: 295301.
Colless,
D.H.
1957. Notes on the culicine mosquitoes of Singapore II. - The Culex vishnuigroup
(Diptera, Culicidae), with descriptions of two new species. Ann. Trop. Med.
Parasitol. 51: 87-l 01.
1959. Notes on the culicine mosquitoes of Singapore. VII. - Host preferences in
relation to transmission of disease. Ann. Trop. Med. Parasitol. 53: 259-267.
Coluzzi, M. and A. Sabatini
1968. Divergenze morfologiche e barriere de sterilita’ nel complesso Aedes mariae
(Diptera, Culicidae). Riv. Parassit. 29: 49-70.
Curtis, J.
1837. A guide to an arrangement of British insects. Edition 2. London. 294 pp.
Danilov,
V.N.
1985a. Komaty (Diptera, Culicidae) Afganistana. Soobshchenie I. Opredelitel’naja
tablitsa somak. Medits. Parasitol. Parazitar. Bolezn. 2: 67-72.
1985b. Komary (Diptera, Culicidae) Afganistana. Soobshchenie 2. Opredelitel’naja tablitsa lichinok IV stadii. Medits. Parasitol. Parazitar. Bolezn. 4:
51-55.
De Grandpre, A.D. and D. D’E. De Charmoy
1900(1901). Les moustiques: anatomie et biologie. Contribution a I’etude des
Culicides et principalement des genres Culex et Anopheles, de leur role darts
la propagation de la malaria et de la filariose et des moyens de s’en preserver.
Port-Louis. 69 pp., 5 pl.
120
Contrib.Amer. Ent. Inst.,vol. 24, no. 1, 1988
Delfinado,
M.D.
1966. The culicine mosquitoes of the Philippines, tribe Culicini (Diptera, Culicidae).
Mem. Am. Entomol. Inst. 7: l-252.
De Meillon, B., M. Parent and O’C. Black
1945. Descriptions of new larvae and pupae of Ethiopian Culicini. Bull. Entomol.
Res. 36: 85-l 01.
Derwesh,
A.I.
1965. A preliminary list of identified insects and some arachnids of Iraq. Minist. Agr.
Baghdad, Bull. No. 112. 123 pp.
Dimentmand, Ch. and J. Margaiit
1981. Rainpools as breeding and dispersal sites of mosquitoes and other aquatic
insects in the Central Negev Desert. J. Arid Environ. 4: 123-129.
Doherty, R.L., J.G. Carley, M.J. Mackerras and E.N. Marks
1963. Studies of arthropod-borne virus infections in Queensland III. Isolation and
characterization of virus strains from wild-caught mosquitoes in North
Queensland. Aust. J. Exp. Biol. Med. Sci. 41: 17-40.
Doleschall,
C.L.
1856. Eerste bijdrage tot de Kennis der dipterologische Fauna. Natuurk. Tijdschr.
Ned.-lnd. 10: 403-414.
Ddnitz, W.
1902. Beitrage zur Kenntniss der Anopheles. Z. Hyg. Infektionskr. 41: 15-88, 2 pl.
Dow, R.P.
1953. Notes on Iranian mosquitoes. Am. J. Trop. Med. Hyg. 2: 683-695.
Doyle, P.E., S. Sadek Bekheit and D.S. Ellis
1980. Further studies of crossing type in Egyptian members of the Culex pipiens
complex. Trans. R. Sot. Trop. Med. Hyg. 74: 676.
DuBose, W.P. and T.J. Curtin
1965. Identification keys to the adult and larval mosquitoes of the Mediterranean
area. J. Med. Entomol. 1: 349-355.
Dyar, H.G.
1907. Report on the mosquitoes of the coast region of California, with descriptions
of new species. Proc. U.S. Nat. Mus. 32(1516): 121-129.
191711918). A second note on the species of Culex of the Bahamas (Diptera,
Culicidae). Inset. Inscit. Menst. 5: 183-187.
1918. A revision of the American species of Culex on the male genitalia (Diptera,
Culicidae). Inset. Inscit. Menst. 6: 86-l 11, 2 pl.
1920. A collection of mosquitoes from the Philippine Islands (Diptera, Culicidae).
Harbach: Subgenus Culex in SouthwesternAsia
121
Inset. Inscit. Menst. 8: 175-186.
1924. A new mosquito from Siberia (Diptera, Culicidae). Inset. Inscit. Menst. 12:
127-131.
Dyar, H.G. and F. Knab
1906. The larvae of Culicidae classified as independent organisms.
Entomol. Sot. 14: 169-230, 12 pl.
J. N.Y.
1908. Descriptions of some new mosquitoes from tropical America. Proc. U.S. Nat.
Mus. 35( 1632): 53-70.
1909a. On the identity of Culex pipiens Linnaeus. Proc. Entomol. Sot. Wash. 11:
30-39.
1909b.
Descriptions of some new species and a new genus of American
mosquitoes. Smithson. Misc. Collect. 52(1822): 253-266.
1914. New mosquitoes from Peru (Diptera, Culicidae).
58-62.
Inset. Inscit. Menst. 2:
1915. Notes on the species of Culex of the Bahamas.
112-115.
Inset. Inscit. Menst. 3:
Edwards,
F.W.
The African species of Culex and allied genera.
241-268.
1911.
1912a. Notes on the British mosquitos (Culicinae).
260-264.
Bull. Entomol. Res. 2:
Entomologist (Lond.) 45:
1912b. A synopsis of the species of African Culicidae, other than Anopheles. Bull.
Entomol. Res. 3: l-53.
1912c. Revised key to the known larvae of African Culicinae. Bull. Entomol. Res.
3: 373-385.
1913a. Tipulidae and Culicidae from the Lake of Tiberias and Damascus. J. Proc.
Asiatic Sot. Bengal 9: 47-51.
1913b. Further notes on African Culicidae. Bull. Entomol. Res. 4: 47-59.
1913~. New synonymy in Oriental Culicidae. Bull. Entomol. Res. 4: 221-242.
1914a. New species of Culicidae in the British Museum, with notes on the genitalia
of some African Culex. Bull. Entomol. Res. 5: 63-81.
1914b. New Culicidae from Borneo and Hong Kong.
125-l 28.
Bull. Entomol. Res. 5:
122
Contrib.Amer. Ent. Inst.,vol. 24, no. 1, 1988
1915. New and little-known East African Culicidae.
273-281,
Bull. Entomol. Res.
5:
1920. Mosquito notes. Bull. Entomol. Res. 10: 129-137.
1921. A revision of the mosquitoes of the Palaearctic Region. Bull. Entomol. Res.
12: 263-351.
1922. A synopsis of adult Oriental culicine (including megarhinine and sabethine)
mosquitoes. Part I. Indian J. Med. Res. 10: 249-293, 3 pl.
1923. Mosquito notes. -
IV. Bull. Entomol. Res. 14: l-9.
1924. A synopsis of the adult mosquitos of the Australasian Region. Bull. Entomol.
Res. 14: 351-401.
1926. Una revisione delle zanzare delle regioni paleartiche. Riv. Malariol. 5: l-l 52.
1932a. Genera Insectorum. Diptera. Fam. Culicidae.
Verteneul, Bruxelles. 258 pp., 5 pl.
1932b. Mosquito notes. -
Fascicle 194.
Desmet-
Xl. Bull. Entomol. Res. 23: 559-562.
1934. Appendix, pp. 425-453.
In: P.J. Barraud, The fauna of British India,
including Ceylon and Burma. Diptera. Vol. V. Family Culicidae. Tribes
Megarhinini and Culicini. Taylor and Francis, London.
1941. Mosquitoes of the Ethiopian Region III. - Culicine adults and pupae. British
Museum (Natural History), London. viii + 499 pp., 4 pl.
El-Said, S. and M. Kenawy
1983a. Geographical distribution of mosquitoes in Egypt. J. Egypt. Public Health
Assoc. 58: 46-76.
1983b. Anopheline and culicine mosquito species and their abundance in Egypt.
J. Egypt. Public Health Assoc. 58: 108-l 42.
Enderlein,
G.
1920. Die Culiciden-Fauna Madagascars. Wien. Entomol. Z. 38: 47-52.
Evans, A.M.
1933. Anopheles demeilloni sp. n., a new name for Anopheles transvaalensis of
authors. Ann. Trop. Med. Parasitol. 27: 265-269.
Fabricius,
J.C.
1794. Entomologia systematica emendata et aucta, secundum classes, ordines,
genera, species adjectis synonymis, locis, observationibus, descriptionibus.
Vol. 4. Hafniae. vi + 472 pp.
Harbach: Subgenus Culex in SouthwesternAsia
123
Favre, V.V.
1903. An attempt in the study of malaria in Russia. Kharkov, Russia. 344 pp.
Ficalbi, E.
1889a. Alcune generalita. - Descrizione di una specie nouva (Culex hortensis ).
Notizie preventive sulla zanzare italiane. - la. (Nota preventiva). Boll. Sot.
Entomol. Ital. 21: 20-30.
1889b. Descrizione di una specie nouva. (Notizie preventive sulle zanzare italiane.
Ila. Nota preventiva). Boll. Sot. Entomol. Ital. 21: 50-53.
1889( 1890). Notizie preventive sulle zanzare italiane. Via. Nota preventiva.
Quistioni zoologiche intorno al Culex pipiens e descrizione di una specie
nouva (Culex phyfophagus). Boll. Sot. Entomol. Ital. 21: 124-131.
1890. Descrizione di una specie nouva (Notizie preventive sulle zanzare italiane.
Vlla. Nota preventive). Boll. Sot. Entomol. Ital. 22: 81-84.
1893. Revisione della especie Europee della famiglia della zanzare.
Entomol. Ital. 25: 48-61, 136-144.
Boll. Sot.
ForskBI, P.
1775. Descriptiones animalium avium, amphibiorum, piscium, insectorum, vermium,
quae in itinere orientali observavit. Mdlleri, Hauniae. 164 pp.
Gad, A.M.
1956. Mosquitoes of the oases of the Libyan Desert of Egypt. Bull. Sot. Entomol.
Egypte 40: 131-136.
Gad, A.M. and R.O. Darwish
1957. Mosquitoes collected in southern Sinai. Bull. Sot. Entomol. Egypte 41:
535-538.
Gad, A.M. and A. Salit
1972. The mosquitoes of the Red Sea area, Egypt. J. Med. Entomol. 9: 581-582.
Gad, A.M., M.M. Hassan, S. El Said, M.I. Moussa and O.L. Wood
1987. Rift Valley fever virus transmission by different Egyptian mosquito species.
Trans. R. Sot. Trop. Med. Hyg. 81: 694-698.
Germar, E.F.
1817. Reise nach Dalmatien und in das Gebiet von Ragusa. Leipzig, Brockhaus.
323 pp.
Giles, G.M.
1900. A handbook of the gnats or mosquitoes giving the anatomy and life history of
the Culicidae. John Bale, Sons & Danielsson, Ltd., London. xi + 374 pp.
1901 a. A plea for the collective investigation of Indian Culicidae [sic], with suggestions as to moot points for enquiry, and a prodromusof species known to the
124
Contrib.Amer. Ent. Inst.,vol. 24, no. 1, 1988
author. J. Bombay Nat. Hist. Sot. 13: 592-610, 2 pl.
1901 b.
Six new species of Culicidae from India.
192-l 97.
Entomologist (Lond.)
34:
1901c. Description of four new species of Anopheles from India. Entomol. Mon.
Mag. 37: 196-198.
1902. A handbook of the gnats or mosquitoes, giving the anatomy and life history
of the Culicidae, together with descriptions of all species notices up to the
present date. 2nd edition. John Bale, Sons & Danielsson, Ltd., London. xii +
530 pp. + 17 pl.
1904. Notes on some collections of mosquitoes, &c., received from the Philippine
Islands and Angola; with some incidental remarks upon classification. J. Trop.
Med. 7: 365-369.
1906. Mosquito notes. I. - Note on a small collection of mosquitoes from Barhain
[sic] in northern Arabia (shores of Persian Gulf). J. Trop. Med. 9: 130-132.
Goug h, L.H.
1914. Preliminary notes on Egyptian mosquitoes. Bull. Entomol. Res. 5: 133-l 35.
Graham, W.M.
1910. On new species of West-African Culicidae. Ann. Mag. Nat. Hist. 5: 264-273.
Grassi, G.B.
1899. Ancora sulla malaria. R.C. Accad. Lincei 8: 559-561.
Griinberg,
K.
1905. Zur Kenntnis der Culicidenfauna von Kamerun und Togo. Zool. Anz. 29:
377-390.
Gutsevich,
A.V., A.S. Monchadskii
and A.A. Shtakel’berg
1974. Fauna of the U.S.S.R. Diptera. Volume 3, No. 4. Mosquitoes Family Cullcidae [sic]. Keter Publishing House Jerusalem Ltd., Jerusalem. iii + 409 pp.
Hackett, L.W.
1934. The present status of our knowledge of the subspecies of Anopheles
maculipennis. Trans. R. Sot. Trop. Med. Hyg. 28: 109-128.
Haliday, A.H.
1833. Catalogue of Diptera occurring about Holywood in Downshire.
Mag. 1: 147-180.
Entomol.
Harbach,
R.E.
1983. Notes on some types of Culex (Culex) (Diptera, Culicidae) deposited in
England and France. Mosq. Syst. 15: 98-l 10.
1985a. Pictorial keys to the genera of mosquitoes, subgenera of Culex and the
Harbach: Subgenus Culex in SouthwesternAsia
125
species of Culex (Culex) occurring in southwestern Asia and Egypt, with a note
on the subgeneric placement of Culex deserticola (Diptera: Culicidae). Mosq.
Syst. 17: 83-l 07.
1985b. A new species, Culex Culex) litwakae (Diptera: Culicidae), from the coastal
region of Kenya. Mosq. Syst. 17: 254-265.
1986( 1987). Culex verutus, a new species of the subgenus Culex (Diptera:
Culicidae) from Sierra Leone. Mosq. Syst. 18: 216-229.
Harbach, R.E. and K.L. Knight
1980. Taxonomists’ glossary of mosquito anatomy.
Martton, New Jersey. xi + 415 pp.
Plexus Publishing, Inc.,
Harbach, R.E., E.L. Peyton and W.L. Jakob
1983( 1984). Synonymy of Culex (Culex) oswaldoi with Culex (Culex) maxi (Diptera,
Culicidae). Mosq. Syst. 15: 31 O-317.
Harbach, R.E., B.A. Harrison and A.M. Gad
1984. Culex (Culex) molestus (Forskal) (Diptera: Culicidae): neotype designation,
description, variation and taxonomic status. Proc. Entomol. Sot. Wash. 86:
521-542.
Harbach, R.E., C. Dahl and G.B. White
1985. Culex (Culex) pipiens Linnaeus (Diptera: Culicidae): concepts, type
designations and description. Proc. Entomol. Sot. Wash. 87: l-24.
Harris, W.V.
1942. Notes on culicine mosquitos in Tanganyika Territory. Bull. Entomol. Res.
33: 181-193.
Hayes, C.G., A. Basit, S. Bagar and R. Akhter
1980. Vector competence of Culex tritaeniorhynchus (Diptera: Culicidae) for West
Nile virus. J. Med. Entomol. 17: 172-l 77.
Hodes, H.L.
1946. Experimental transmission of Japanese B encephalitis by mosquitoes and
mosquito larvae. Bull. Johns Hopkins Hosp. 79: 358-360.
Hopkins,
G.H.E.
1936. Mosquitoes of the Ethiopian Region I. - Larval bionomics of mosquitoes and
taxonomy of culicine larvae. British Museum (Natural History), London. 250 pp.
1952. Mosquitoes of the Ethiopian Region I. - Larval bionomics of mosquitoes and
taxonomy of culicine larvae. 2nd edition. British Museum (Natural History),
London. viii + 355 pp.
Hsiao, T.Y.
1948. Epidemiology of the diseases of naval importance in Manchuria. U. S. Nav.
Med. P-1289. 89 pp.
126
Contrib.Amer. Ent. Inst.,vol. 24, no. 1, 1988
Hsiao, T.Y. and R.M. Bohart
1946. The mosquitoes of Japan and their medical importance. Navmed. (Dep. of
the Navy, Washington, D.C.) 1095: l-44.
Hussain,
A.A.
1963. Provisional list of insect pests and bibliography of insect fauna of Iraq. Bull.
Coil. Sci. Jami Baghdad Kulligat 7: 43-83.
Ibrahim, I.K., T.Y.M. Al-Samarae, M.A. Zaini and S. Kassal
1983. Identification key for Iraqi culicine mosquitoe [sic] larvae (Culicinae-Diptera).
Bull. Endem. Dis. 12-13: 89-l 13.
Iglisch, I.
1977. “Hausmucken” (Culex-pipiens - Gruppe) als mdgliche ubertrager des
Serumhepatitis-Virus. Untersuchungen zur Genitalmorphologie. Z. Ang. Zool.
64: 257-290.
Ingram, A. and B. De Meillon
1927. A mosquito survey of certain parts of South Africa, with special reference to
the carriers of malaria and their control. (Part I). Publ. S. Afr. Inst. Med. Res. 4:
I-81.
Ingram, A. and J.W.S. Macfie
1919. The early stages of West African mosquitos - IV. Bull. Entomol. Res. 10:
59-60.
lyengar,
M.O.T.
1938. Studies on the epidemiology of filariasis in Travancore.
Mem. 30: l-179 pp.
Indian Med. Res.
1953. Filariasis in Thailand. Bull. W. H. 0. 9: 731-766.
Izquierdo,
J.J.
1916. lnvestigaciones sobre paludismo en Puebla. Tesis Col. Est. Puebla. 100
PP.
James, S.P.
1902. Malaria in India. Sci. Mem. Med. Sanit. Dep. India 2: l-106.
Joyeux, C.
1918. Note sur les Culicides de Macedoine. Bull. Sot. Pathol. Exot. 2: 530-547.
Jupp, P.G.
1970. The taxonomic status of Culex (Culex) univittatus Theobald
Culicidae) in South Africa. Mosq. Syst. News Lett. 2: 9-16.
(Diptera:
1971. The taxonomic status of Culex (Culex) univittatus Theobald (Diptera:
Culicidae) in South Africa. J. Entomol. Sot. South. Afr. 34: 339-357.
1972. A morphological study of Culex (Culex) univittatus Theobald and Culex
(Culex) neavei Theobald from various African countries. Mosq. Syst. 4: 103-
Harbach: Subgenus Culex in SouthwesternAsia
127
113.
Jupp, P.G. and B.M. McIntosh
1967. Ecological studies on Sindbis and West Nile viruses in South Africa. II. Mosquito bionomics. S. Afr. J. Med. Sci. 32: 15-33.
Jupp, P.G., B.M. McIntosh and R.G. Brown
1966. Laboratory transmission experiments with West Nile and Sindbis viruses and
Culex (Culex) thei/eriTheobald. S. Afr. J. Med. Sci. 31: 95-97.
Karabatsos, N. (ed .)
1985. International catalogue of arboviruses including certain other viruses of
vertebrates. 3rd edition. Am Sot. Trop. Med. Hyg., San Antonio. 1137 pp.
Khalaf, K.T.
1962. Handbook of the mosquitoes recorded from Iraq. Shafik Press, Baghdad. v
+ 62 pp.
Khattat,
F.H.
1955. An account of the taxonomy and biology of the larvae of culicine mosquitoes
in Iraq I. Central Iraq. Bull. Endem. Dis. 1: 154-183.
Kirkpatrick,
T.W.
1924( 1925). Preliminary notes on the determination of Egyptian mosquitos, with
descriptions of three species new to science. Bull. Sot. Entomol. Egypte 8:
362-391.
1925. The mosquitoes of Egypt. Egyptian Government Anti-Malaria Commission.
Government Press, Cairo. xi + 224 pp., 33 pl., 2 maps.
1926. Culex (Lasiosiphon) adairi, nom. nov. Bull. Sot. Entomol. Egypte 10: 112.
Kitron, U. and H. Pener
1986. Distribution of mosquitoes (Diptera: Culicidae) in northern Israel: a historical
perspective II. Culicine mosquitoes. J. Med. Entomol. 23: 182-187.
Knight,
K.L.
1951. A review of the Culex pipiens complex in the Mediterranean subregion
(Diptera, Culicidae). Trans. R. Entomol. Sot. Lond. 102: 354-364.
1953a. The mosquitoes of the Yemen (Diptera, Culicidae).
Wash. 55: 212-234.
Proc. Entomol. Sot.
1953b. Two new species of mosquitoes from the Yemen (Diptera: Culicidae). J.
Wash. Acad. Sci. 43: 320-325.
Knight, K.L. and A.A. Abdel Malek
1951. A morphological and biological study of Culex pipiens in the Cairo area of
Egypt. Bull. Sot. Fouad ler Entomol. 35: 175-185.
128
Contrib.Amer. Ent. Inst.,vol. 24, no. 1, 1988
Koidzumi,
M.
1920. Preventive medical study of mosquitoes in Formosa.
Hokoku (Rep. Formosa Gov.) 8: 1-158.
Daiwan Kenkyujo
Latreille,
P.A.
1810. Considerations generales sur I’ordre nature1 des animaux composant les
classes des Crustaces, des Arachnides, et des Insectes; avex un tableau
methodique de leur genres, disposes en familles. Schoell, Paris. 444 pp.
Laveran, A.
1900. Sur un Anopheles provenant de Madagascar.
109-l 10.
C. R. Sot. Biol. Paris 52:
Leach, W.E.
1825. Description of thirteen species of Formica and three species of Culex, found
in the environs of Nice. Zool. J. Lond. 2: 289-293.
Lee,
V.H.
1979. Isolation of viruses from field populations of Culicoides (Diptera: Ceratopogonidae) in Nigeria. 3. Med. Entomol. 16: 76-79.
Leeson, H.S. and 0. Theodor
1948. Mosquitoes of Socotra. Bull. Entomol. Res. 39: 221-229.
Leicester,
G.F.
1908. The Culicidae of Malaya. Stud. Inst. Med. Res. F.M.S. 3(3): 18-261.
Lever,
R.J.A.W.
1954. The mosquito Culex torrentium Mar. (Dipt., Culicidae) new to Dorset.
Entomol. Mon. Mag. 90: 65.
Lewis, D. J.
1943a. The culicine mosquitos of Eritrea. Bull. Entomol. Res. 34: 279-285.
1943b. Mosquitoes in relation to yellow fever in the Nuba Mountains, Angloegyptian Sudan. Ann. Trop. Med. Parasitol. 37: 65-76, 3 pl.
1945. Observations on the distribution and taxonomy of Culicidae (Diptera) in the
Sudan. Trans. R. Entomol. Sot. Lond. 95: l-24, 2 maps.
1948. The mosquitos of the Jebel Auliya Reservoir of the White Nile.
Entomol. Res. 39: 133-157, 1 pl.
Bull.
1956. The Culex mosquitos of the Sudan. Bull. Entomol. Res. 47: 703-721.
Lien, J.C.
1968. New species of mosquitoes from Taiwan (Diptera: Culicidae). Part V. Three
new subspecies of Aedes and seven new species of Culex. Trop. Med. 10:
217-262.
Harbach: Subgenus Culex in SouthwesternAsia
129
Linnaeus,
C.
1758. Systema naturae per regna tria naturae, secundum classes, ordines, genera,
species, cum characteribus, differentiis, synonymis, locis. Ed. 10, Vol. 1.
Holmiae. 824 pp.
1762. Zweyter Theil, enthalt Beschreibungen verschiedener wichtiger Naturalien,
pp. 267-606. In: F. Hasselquist, Reise nach Palastina inden Jahren von 1749
bis 1752. Rok, Germany.
Liston, W.G.
1901. A year’s experience of the habits of Anopheles in Ellichpur. The description
of the species of Anopheles found in Ellichpur during the year. Indian Med.
Gaz. 36: 441-443,l pl.
Lotfi, M.D.
1970. Iranian species of genus Culex (Culinae [sic]: Diptera). Bull. Sot. Pathol.
Exot. 63: 399-403.
1973. Iranian species of genus Culex (Culicinae: Diptera) II. - Report of four
species of larvae (including three new records) and 14 adult species. Bull. Sot.
Pathol. Exot. 66: 204-207.
1976. Key to Culicinae larva [sic] of Iran genus Culex and their biology (Culicidae:
Diptera). Iran. J. Publ. Health 5: 71-84.
Lumsden, W.H.R. and E.C.C. van Someren
1953. Records of Culex species (Diptera: Culicidae) from West Nile District,
Uganda, with notes on their behavior. Proc. R. Entomol. Sot. Lond. B Taxon.
22: 19-22.
Lynch Arribalzaga,
F.
1891 a. Dipterologia argentina. Revta Mus. La Plata 1: 345-377.
1891 b. Dipterologia argentina. Revta Mus. La Plata 2: 131-174, 5 pl.
Macquart, J.
1838a. Dipteres exotiques nouveaux ou peu connus.
9-225, 25 pl.
Mem. Sot. Sci. Lille 2:
1838b. Dipteres. Vol. I: 33. In: P. Barker-Webb and S. Berthelot 1835-1844,
Histoir naturelle des iles Canaries. 3 vols. Paris.
1850. Dipteres exotiques nouveaux ou
Sci. Lille 1850: 309-479.
peuconnus, 4e Supplement. Mem.Soc.
Manson-Bahr,
P.
1959. The story of Filaria bancrofti. Part IV.
bancrofti. J. Trop. Med. Hyg. 62: 138-145.
Mosquito intermediaries of W.
130
Contrib.Amer. Ent. Inst., vol. 24, no. 1, 1988
Margalit, J. and A.S. Tahori
1970a. Population dynamics of Culex pipeins molestus Forskal and of Culex
univittatusTheobald in Israel. Isr. J. Entomol. 5: 141-150.
1970b. Species of mosquitoes found in Israel during a survey (1955-58).
Entomol. 5: 151-l 59.
Isr. J.
1973. The mosquito fauna of Sinai. J. Med. Entomol. 10: 89-96.
1974. An annotated list of mosquitoes in Israel. Isr. J. Entomol. 9: 77-91.
Margalit, J., A.S. Tahori and Y. Nir
1971. Seasonal abundance of some mosquito species in Israel during a survey
1965-8. Isr. J. Entomol. 6: 323-325.
Margalit, J., M. Avrahami and A.S. Tahori
1973. Mosquitoes (Diptera: Culicidae) breeding in the Dead Sea area. Isr. J. Zool.
22: 27-37.
Marsh, F.
1933. A new species of Anopheles (Myzomyia group) from Southwest Persia.
Stylops 2: 193-197, 1 pl.
Marshall, J.F. and J. Staley
1937. Some notes regarding the morphological and biological differentiation of
Culex pipiens Linnaeus and Culex molestus Forskal (Diptera, Culicidae). Proc.
R. Entomol. Sot. Lond. A 12: 17-26.
Martini, E.
1922. Zur Nomenclatur der fur Mitteleuropa Wichtigsten Stechmiicken. Entomol.
Mitt. 11: 106-126.
1925. Zwei bemerkenswerte Culiciden von einem eigenartigen Biotop. Int. Rev.
Hydrobiol. Hydrogr. 12: 333-337.
1926. Uber die Stechmucken der Umgebung von Saratow. Arb. Biol. Wolga Sta.
8: 189-227.
1927.
Uber zwei neue Stechmuckenarten
Tropenhyg. 31: 386-390.
aus Anatolien.
1931. 11. u. 12. Culicidae. E. Schwizerbart’sche
Nagele) G.M.B.H. 398 pp., 1 pl.
Arch. Schiffs.
Verlagsbuchhandlung (Erwin
Mattingly,
P.F.
1954. The distribution of some African mosquitoes. Proc. Linn. Sot. Lond. 165:
49-61.
1955a. Mosquitoes (Diptera: Culicidae) from the Tropical Institute at Hamburg.
Proc. R. Entomol. Sot. Lond. Ser. B Taxon. 24: 27-33.
Harbach: Subgenus Culex in SouthwesternAsia
131
1955b. Le sous-genre Neoculex (Diptera, Culicidae) dans la sous-region
Mediterraneenne. I. - Espece, sous-espece et synonymie nouvelles. Ann.
Parasitol. Hum. Comp. 30: 374-388.
1956. Lectotypes of mosquitoes (Diptera: Culicidae) in the British Museum. Proc.
R. Entomol. Sot. Lond. A 31: 37-44.
Mattingly, P.F. and E.S. Brown
1955. The mosquitos (Diptera: Culicidae) of the Seychelles. Bull. Entomol. Res.
46: 69-l 10, 3 pl.
Mattingly, P.F. and K.L. Knight
1956. The mosquitoes of Arabia. I. Bull. Br. Mus (Nat. Hist.) Entomol. 4: 91-141.
Mattingly, P.F. and J. Rageau
1958. Culex (Culex) iyengari n. sp., a new species of mosquito (Diptera, Culicidae)
from the South Pacific. Pac. Sci. 12: 241-250.
McIntosh,
B.M., P.G. Jupp, D.B. Dickinson, G.M. McGillivray and J.
Sweetnam
1967. Ecological studies on Sindbis and West Nile viruses in South Africa. I. Viral
activity as revealed by infection of mosquitoes and sentinel fowls. S. Afr. J.
Med. Sci. 32: l-l 4.
Medschld, [no initials]
1928. ijber Aedes lepidonotus Edw. und Aedes refiki n. sp. Arch. Schiffs.
Tropenhyg. 32: 306-315.
1930. Culex martiniin. sp. Arch. Schiffs. Tropenhyg. 34: 364-369.
Meegan,
J.M.
1981. Rift Valley Fever in Egypt: an overview of the epizootics in 1977 and 1978.
Contrib. Epidem. Biostatist. 3: 100-l 13.
Meegan, J.M., G.M. Khalil, H. Hoogstraal and F.K. Adham
1980. Experimental transmission and field isolation studies implicating Culex
pipiens as a vector of Rift Valley fever virus in Egypt. Am. J. Trop. Med. Hyg.
29: 1405-1410.
Meigen,
J.W.
1804. Klassification und Beschreibung der europaischen zweiflijglichen lnsekten
(Diptera Linn.). Vol. 1, Pt. 1. Braunschweig. xxviii + 152 pp., 8 pl.
1818. Systematische Beschreibung der bekannten europaischen zweiflugligen
Insekten. Vol. 1. Aachen. xxxvi + 333 pp., 11 pl.
1830. Abbildung aller bis jetzt bekannten europaischen zweifltigligen Insekten.
Vol 6. Hamm. iv + 401 pp., 12 pl.
Contrib.Amer. Ent. inst., vol. 24, no. 1, 1988
132
Mekuria, Y.
1968. A catalogue of the culicine mosquito fauna of Ethiopia and their distribution
in the country. Ethiop. Med. J. 6: 73-80.
Mekuria, Y., S. Asseta and Y. Mebrahtu
1978. Culicine mosquitoes (Diptera: Culicidae) of Ethiopia:
distribution. Ethiop. Med. J. 16: 71-93.
Checklist and
Mochizuki,
D.
1913. Culicidae of the Fukuoka district. Fukuoka lkadaigaku Zasshi 7: l-65.
Monchadskii,
A.S.
1936. Les larves des moustiques (Fam. Culicidae) de I’URSS
limitrophes. Tabl. Anal. Faune U.R.S.S. 24: l-383.
et des pays
1951. Lichinki krovososuchehlkh komarov SSSR i sopredel’ni’kh stran (podsem.
Culicinae). Akad. Nauk SSSR Opred. Faun. SSSR 37: l-288 + 2 unnumbered
PP.
Mulligan, H.W. and I.M. Puri
1936. Description of Anopheles (Anopheles) habibi n. sp. from Quetta, Baluchistan. Rec. Malariol. Surv. India 6: 67-71, 1 pl.
Miiller,
O.F.
1764. Fauna insectorum Fridrichsdalina, sive methodica descriptio insectorum agri
Fridrichsdalinensis, cum characteribus genericis et specificis, nominibus
trivialibus, locis natalibus, iconibus allegatis, novisque pluribus speciebus
additis. Hafniae et Lipsiae. xxiv + 96 pp.
Neveu-Lemaire,
M.
1906. Etude des culicides africains. Arch. Parasitol. 10: 238-288.
Newstead, R., J.E. Dutton and J.L. Todd
1907. Insects and other Arthropoda collected in the Congo Free State. Ann. Trop.
Med. Parasitol. 1: l-l 12, 6 pl.
Nguyen Thi Kim Thoa, Ngo Thi Vien, Tran Tuyet Mai and Nguyen Thi
Ngoc Xuan
1974. Japanese encephalitis vectors: isolation of virus from culicine mosquitoes
in the Saigon area. Southeast Asian J. Trop. Med. Public Health 5: 408-412.
Nielsen, E.T. and H.T. Nielsen
1958. Observations on mosquitoes in Iraq. Entomol. Medd. 28: 282-321, 4 pl.
Nir, Y., R. Goldwasser, Y. Lasowski and J. Margalit
1968. Isolation of West Nile virus strains from mosquitoes in Israel.
Epidemiol. 87: 496-501.
No&,
Am. J.
G.
1899. Contribuzione allo studio dei Culicidi. Boll. Sot. Entomol. Ital. 31: 235-262.
Harbach: Subgenus Culex in SouthwesternAsia
133
Olivier, G.M.
1791. Encyclopedic methodique. Histoire naturelle. Insects. Vol. 6. Paris. 704
PP.
Ovazza, M., J. Hamon and P. Neri
1956. Contribution a I’etude des Dipteres vulnerants de I’empire d’Ethiopie.
Culicidae. Bull. Sot. Pathol. Exot. 49: 151-l 82.
Pallas, P.S.
1771. Reise durch verschiedene Provinzen des Russischen Reichs.
(1967 reprint, Akademische Druck-u. Verlagsanstalt, Graz, Austria).
I.
Vol. 1.
Parr, H.C.M.
1943. The culicine mosquitos of Syria and the Lebanon. Bull. Entomol. Res. 34:
245-251.
Parrish, D.W.
1959. The mosquitoes of Turkey. Mosq. News 19: 264-266.
Patton, W.S.
1905. The culicid fauna of the Aden Hinterland, their haunts and habits. J. Bombay
Nat. Hist. Sot. 16: 623-637.
Philippi,
R.A.
1865. Aufzahlung der chilenischen Dipteren. Verh. Zool. Bot. Ges. Wien
595-782, 7 pl.
15:
Rao, T.R.
1975. Arboviruses and their vectors in India. Indian J. Med. Res. 63: 1219-1233.
Reaumur, R.A.F. de
1738. Memoires pour servir a I’histoire des insectes. Vol. 4. Paris. xxxvi + 636
pp.944 pl.
Reuben,
R.
1969. A redescription of Culex vishnui Theo., with notes on C. pseudovishnui
Colless and C. trifaeniorhynchus Giles, from southern India. Bull. Entomol.
Res. 58: 643-652, 3 pl.
197la. Studies on the mosquitoes of North Arcot District, Madras State, India. Part
3. Host preferences for pigs, birds and some small mammals. J. Med. Entomol.
8: 258-262.
1971 b. Studies on the mosquitoes of North Arcot District, Madras State, India. Part
4. Breeding places of the Culex vishnuigroup of species. J. Med. Entomol. 8:
363-366.
Ribeiro, H. and J.T. Mexia
1966. Research on the mosquitoes of Angola (Diptera: Culicidae) III - A culicine
survey in the Lobito-Catumbela region. An. Inst. Hig. Med. Trop. 23: 167-l 82.
134
Contrib.Amer. Ent. Inst.,vol. 24, no. 1, 1988
Ribeiro, H., H.C. Ramos, C.A. Pires and R.A. Capela
1977. Research on the mosquitoes of Portugal (Diptera, Culicidae) I - Four new
culicine records. An. Inst. Hig. Med. Trop. 5: 203-214.
Rioux, J.-A. and J.-M. Pech
1959. Le biotype autogene de Culex pipiens L. ne doit pas &re nomme Culex
molestus Forskal. Cah. Nat. 15: 115-l 17.
Robineau-Desvoidy,
J.B.
1827. Essai sur la tribu des culicides. Mem. Sot. Hist. Nat. Paris 3: 390-413, 1 pl.
Rondani, C.
1872. Sulle specie ltaliane del genere Culex Lin. Boll. Sot. Entomol. Ital. 4: 29-31.
Rossi, P.
1790. Fauna Etrusca sistens lnsecta quae in provinciis Florentina et Pisana
praesertim collegit. Pt. 1. Liburni. xxii + 348 pp., 6 unnumbered pp., 10 pl.
Roubaud,
E.
1935. La microstructure du flotteur de I’oeuf dans les races biologiques de Culex
pipiens L. Bull. Sot. Pathol. Exot. 28: 443-445.
1945. Le probleme de I’espGce chez le moustique commun Culex pipiens L. Bull.
Sot. Pathol. Exot. 38: 47-60.
1957. Biologie des moustiques - Sur I’existence de deux entities biologiques
distinctes dans la definition du moustique commun rural, Culex pipiens pipiens.
C.R. Acad. Sci. Paris 244: 3115-3116.
Rudnick, A., W. McD. Hammon and G.E. Sather
1962. A strain of Sindbis virus isolated from Culex bitaeniorhynchus mosquitoes in
the Philippines. Am. J. Trop. Med. Hyg. 11: 546-549.
Salem, H.H.
1938. The mosquito fauna of Sinai Peninsula (Egypt) with a description of two new
species. The Egyptian University, the Faculty of Medicine, Publ. No. 16. 31
pp. +19 pl.
1940. Further observations on Anopheles rupicolus Lewis, Culex arbieeni Salem,
and Culex theileri Theobald. Bull. Sot. Fouad ler Entomol. 24: 1l-l 6.
Samina, I., J. Margalit and J. Peleg
1986. Isolation of viruses from mosquitoes of the Negev, Israel. Trans. R. Sot.
Trop. Med. Hyg. 80: 471-472.
Sanchez,
J.
1885. Nota 10s moscos zancudos que han invadido la capital en el presente ano.
La Naturaleza Mexico 7: 203-213.
Harbach: Subgenus Culex in SouthwesternAsia
Say,
135
T.
1823. Descriptions of dipterous insects of the United States. J. Acad. Nat. Sci.
Philadelphia 3: 9-54.
Schrank, F. von P.
1776. Beytrage zur Naturgeschichte. Fritsch, Leipzig. 137 pp.
Scott, H.
1927. Notes on the distribution and habits of Culicidae in central Abyssinia. Bull.
Entomol. Res. 18: 83-89, 1 pl.
Seguy,
E.
1921. Description d’un nouveau moustique Francais du groupe de Aedes mariae
et synopsis des especes de ce groupe. Bull. Sot. Entomol. Fr. 1921:
192-195.
1924. Les moustiques de I’Afrique mineure de I’Egypte et de la Syrie etude
comparative des moustiques des regions Mediterraneennes, de I’Europe
centrale et septentrionale leurs parasites suivi de catalogue des Culicides
Nearctiques et Palearctiques. Encycl. Entomol. 1: l-257.
Senevet,
G.
1946. Contribution a I’etude des larves de Culicides la structure du prementum.
Arch. Inst. Pasteur Alger. 24: 315-329.
1947a. Le genre Culex en Afrique du Nord 1. - Les larves. Arch. Inst. Pasteur
Alger. 25: 107-136.
1947b. Le genre Culex en Afrique du Nord Ibis. - Clef de determination des
larves. Arch. Inst. Pasteur Alger. 25: 212-213.
1949. Le genre Culex en Afrique du Nord 2. - Les armures genitales males. Arch.
Inst. Pasteur Alger. 27: 48-65.
Senevet, G. and L. Andarelli
1954. Le genre Culex en Afrique du Nord. III. Les adultes. Arch. Inst. Pasteur
Alger. 32: 36-79.
1959. Les moustiques de I’Afrique du Nord et du bassin Mediterraneen. Les
genres Culex, Uranotaenia, Theobaldia, Otthopodomyia et Mansonia. Encycl.
Entomol. 37: l-383.
Senevet, G. and M. Prunnelle
1927. Une nouvelle espece d’anophele en Algerie, Anopheles marteri n. sp. Arch.
Inst. Pasteur Alger. 5: 529-533.
Senevet, G., L. Andarelli and R. GraUls
1957a. Les spines preclypeales des larves de moustiques leur utilisation dans la
diagnose. Arch. Inst. Pasteur Alger. 35: 83-89.
136
Contrib.Amer. Ent. Inst.,vol. 24, no. 1, 1988
1957b. Les soies dorsales prothoraciques chez les larves de Culex. Arch. Inst.
Pasteur Alger. 35: 90-98.
Senevet, G., J. Gaud and A. Millet
1949. Validite de I’espece Culex mauritanicus Callot, 1940. Arch. Inst. Pasteur
Alger. 27: 42-47.
Sergent,
E.
1909. Liste des mostiques de I’Afrique du Nord. Ann. Sot. Entomol. Fr. 78:
440-448.
Service,
M.W.
1968. The taxonomy and biology of two sympatric sibling species of Culex, C.
pipiens and C. torrentium (Diptera, Culicidae). J. Zool. (Lond.) 156: 313-323.
Shlngarev,
NJ.
1928. Notes on Culicidae III. Trop. Med. Vet. Moscow 6: 47-53.
Slrivanakarn,
S.
1973. The forms of Culex (Culex) bifaeniorhynchusGiles in Southeast Asia. Mosq.
Syst. 5: 235251.
1976. Medical entomology studies - III. A revision of the subgenus Culex in the
Oriental Region (Diptera: Culicidae). Contrib. Am. Entomol. Inst. (Ann Arbor)
12(2): 1-272.
Sirivanakarn,
S. and G.B. White
1978. Neotype designation of Culex quinquefasciatus Say (Diptera: Culicidae).
Proc. Entomol. Sot. Wash. 80: 360-372.
Skuse,
F.A.A.
1889. Diptera of Australia. Part V. - The Culicidae. Proc. Linn. Sot. N.S.W. 13:
1717-1764, 1 pl.
Smith, A.
1955. The transmission of bancroftial filariasis on Ukara Island, Tanganyika. III. Biting - incidences on man and filarial infections in wild-caught mosquitos. Bull.
Entomol. Res. 46: 495-504.
Soman, R.S. and D.T. Mourya
1985. Transovarial transmission of Japanese encephalitis virus in Culex bitaeniorhynchus mosquitoes. Indian J. Med. Res. 81: 257-259.
Southgate,
B.A.
1979. Bancroftian filariasis in Egypt. Trop. Dis. Bull. 76: 1045-l 068.
Stackelberg,
A.
1927. Les Culicides de I’URSS et des pays limitrophes. Tabl. Anal. Faune URSS
1: 1-169.
Harbach: Subgenus Culex in SouthwesternAsia
137
Stephens,
J.F.
1825. Some observations on the British Tipulidae, together with descriptions of
the species of Culex and Anopheles found in Britain. Zool. J. Lond. 1:
448-457.
1828. Note on a memoir of Mr. Haliday, on insects taken in north of Ireland, with the
description of a new species of Anopheles. Zool. J. Lond. 3: 502-504.
Stone, A.
1956(1957).
Corrections in the taxonomy and nomenclature of mosquitoes
(Diptera, Culicidae). Proc. Entomol. Sot. Wash. 58: 333-344.
1957. Notes on types of mosquitoes in the Hungarian National Museum (Diptera,
Culicidae). Ann. Entomol. Sot. Am. 50: 171-174.
1958. The identity of Cuiex aestuans Wiedemann (Diptera, Culicidae).
Entomol. Sot. Wash. 60: 186.
Proc.
Stone, A. and K.L. Knight
1957. Type specimens of mosquitoes in the United States National Museum: IV,
The genus Culex (Diptera, Culicidae). J. Wash. Acad. Sci. 47: 42-59.
Stone, A., K.L. Knight and H. Starcke
1959. A synoptic catalog of the mosquitoes of the world (Diptera, Culicidae).
Thomas Say Found. 6: l-358.
Storey, G.
1918( 1919). Keys for the determination of Egyptian mosquitoes and their larvae.
Bull. Sot. Entomol. Egypte 5: 84-l 06, 2 pl.
Tahori, A.S., V.V. Sterk and N. Goldblum
1955. Studies on the dynamics of experimental transmission of West Nile virus by
Culex molestus. Am. J. Trop. Med. Hyg. 4: 1015-1027.
Tamayo, M.O. and C.A. Garcia
1907. Las aquas de Huacachina informe presentado a la Sociedad Geografica de
Lima. Mem. Municip. Lima 1906: l-63.
Tanaka, K., K. Mizusawa and E.S. Saugstad
1979. A revision of the adult and larval mosquitoes of Japan (including the Ryukyu
Archipelago and the Ogasawara Islands) and Korea (Diptera: Culicidae).
Contrib. Am. Entomol. Inst. (Ann Arbor) 16: l-987.
Taylor, F.H.
1912. Contribution to a knowledge of the Culicidae and lxodidea of North
Queensland and the Torres Straits. Rep. Comm. Public Health Qd. pp. 27-29.
1913. Report of the entomologist. Rep. Aust. Inst. Trop. Med. 1911: 49-74, 3 pl.
1914a. The Culicidae of Australia. I. Trans. R. Entomol. Sot. Lond. 1914:
Contrib.Amer. Ent. Inst.,vol. 24, no. 1, 1988
138
683-708, 4 pl.
1914b. Culicidae from Papua. Trans. R. Entomol. Sot. Lond. 1914: 185205, 2 pl.
1919. Contributions to a knowledge of Australian Culicidae. No. IV. Proc. Linn.
Sot. N.S.W. 43: 826-843,4 pl.
Taylor,
R.M., H.S. Hurlbut, H.R. Dressler, E.W. Spangler and D.
Thrasher
1953. Isolation of West Nile virus from Culex mosquitoes. J. Egypt. Med. Assoc.
36: 199-208.
Taylor,
R.M., H.S. Hurlbut, T.H. Work, J.R. Kingston and T.E.
Frot hlngham
1955. Sindbis virus: a newly recognized arthropod-transmitted virus. Am. J. Trop.
Med. Hyg. 4: 844-862.
Taylor, R.M., T.H. Work, H.S. Hurlbut and F. Rlzk
1956. A study of the ecology of West Nile virus in Egypt. Am. J. Trop. Med. Hyg. 5:
579-620.
Tesh, R.B., S. Saldl, S.Y. Gaidamovich,
R.Roldhain and JH. Vesenjak
1976. Serological studies on the epidemiology of sandfly fever in the Old World.
Bull. W.H.O. 54: 663-674.
Theobald,
F.V.
1901 a. A monograph of the Culicidae or mosquitoes. Vol. 1. British Museum
(Natural History), London. xviii + 424 pp.
1901 b. A monograph of the Culicidae or mosquitoes. Vol. 2. British Museum
(Natural History), London. viii + 391 pp.
1901c. Notes on a collection of mosquitoes from West Africa, and descriptions of
new species. Mem. Liverp. Sch. Trop. Med. 4: i-xiv, 3 pl.
1902. A short description of the Culicidae of India, with descriptions of new species
of Anopheles. Proc. R. Sot. Lond. Ser. B 69: 367-394, 1 pl.
1903a. A monograph of the Culicidae or mosquitoes. Vol. 3. British Museum
(Natural History), London. xviii + 359 pp., 17 pl.
1903b. Report on a collection of mosquitoes or Culicidae, etc., from Gambia, and
descriptions of new species. Mem. Liverp. Sch. Trop. Med. lO(App.): i-xi, 2 pl.
1904. The mosquitoes of Egypt, the Sudan and Abyssinia. Rep. Wellcome Res.
Lab. Gordon Mem. Coil. Khartoum 1: 62-83, 2 pl.
1905a. Genera Insectorum. Diptera, Fam. Culicidae. Fascicle 26. M.P. Wytsman,
Bruxelles. 50 pp., 2 pl.
139
Harbach: Subgenus Culex in SouthwesternAsia
1905b. A catalogue of the Culicidae in the Hungarian National Museum with descriptions of new genera and species. Ann. Mus. Nat. Hung. 3: 61-l 20,4 pl.
1905~. New Culicidae from India, Africa, British Guiana, and Australia. J. Econ. Biol.
1:17-36, 2 pl.
1906. Report on economic entomology. Part I. Second report on the mosquitoes
or Culicidae of the Sudan. Rep. Wellcome Lab. Gordon Coil. Khartoum 2:
67-83.
1907. A monograph of the Culicidae or mosquitoes. Vol. 4.
(Natural History), London. xix + 639 pp., 16 pl.
British Museum
1908a. First report on the collection of Culicidae and Corethridae in the Indian
Museum, Calcutta, with descriptions of new genera and species. Rec. Indian
Mus. 2: 287-302.
1908b. New mosquitoes from the Sudan and list and synoptic table of all the known
Sudanese species. Rep. Wellcome Res. Lab. Gordon Mem. COIL Khartoum 3:
249-267, 2 pl.
1909. Descriptions of the new mosquitoes collected by Dr. Graham in Ashanti.
Colonial Office, Gold Coast, Miscellaneous No. 237. 31 pp.
1910a. I. Second report on the collection of Culicidae in the Indian Museum,
Calcutta, with descriptions of new genera and species. Rec. Indian Mus. 4:
l-33, 3 pl.
1910b. The Culicidae of Fiji, including two new species. Entomologist (Lond.) 43:
155-159.
191Oc. A monograph of the Culicidae or mosquitoes. Vol. 5.
(Natural History), London. xiv + 646 pp., 6 pl.
1911. The Culicidae or mosquitoes of the Transvaal.
232-272, 11 pl.
British Museum
S. Afr. Vet. Div. Rep. 1:
1912. Three new Culicidae from the Transvaal. Entomologist (Lond.) 45: 92-93.
1913. Second report on the mosquitoes of the Transvaal. S. Afr. Vet. Div. Rep. 2:
315-342, 2 pl.
Theodor,
0.
1923. Pupae of some Palestinian culicines. Bull. Entomol. Res. 14: 341-345.
1952. Diptera in the Middle East.
107-l 19.
Istanbul Univ. Fen Fak. Mecm. Seri B 17:
Travis, B.V. and R.M. Labadan
1967. Arthropods of medical importance in Asia and the European USSR. Part I.
140
Contrib.Amer. Ent. inst.,vol. 24, no. 1, 1988
U.S. Army Natick Lab. Tech. Rep. 67-65-ES. 322 pp.
Tureil, M., T. Gargan and C. Bailey
1982. Culexpipiens as a vector of Rift Valley fever. [Abstracts of papers presented
at the Fifteenth Annual Meeting of the Society for Epidemiologic Research.]
Am. J. Epidemioi. 116: 556.
Urbaneill, S. and L. Builini
1985. Eiectrophoretic studies on Culex quinguefasciatus Say from Africa: genetic
variability and divergence from Culex pipiens L. (Diptera: Cuiicidae). Bull.
Entomoi. Res. 75: 291-304.
Ventrillon,
E.
1905a. Cuiicides nouveaux de Madagascar. Arch. Parasitoi. (Paris) 9: 441-450.
1905b. Cuiicides nouveaux de Madagascar.
427-431.
Bull. Mus. Hist. Nat. Paris
11:
Walker, F.
1848. List of the specimens of dipterous insects in the collection of the British
Museum. Part I. British Museum, London. 484 pp.
1856. lnsecta Saundersiana: or characters of undescribed insects in the collection
of William Wilson Saunders. Vol. I., Diptera. London. 474 pp.
1859. Cataiogue of the dipterous insects collected at Makessar in Celebes, by Mr.
A.R. Wallace, with descriptions of new species. J. Proc. Linn. Sot. Lond. Zool.
4: 90-172.
Waiti, J.
1835. Reise durch Tyroi, Oberitaiien und Piemont nach dem sudlichen Spainen. . .
nebst einem Anhang zoologischen inhaits. Vol. 2. Passau, Pustet. 120 pp.
Ward, R.A.
1972. Mosquitoes of Afghanistan - an annotated checklist. Mosq. Syst. 4: 93-97.
Waterston,
J.
1918. On the mosquitos of Macedonia. Bull. Entomol. Res. 9: l-l 2.
Weinbren, M.P., R.H. Kokernot and K.C. Smithburn
1956. Strains of Sindbis-like virus isolated from culicine mosquitoes in the Union of
South Africa. S. Afr. Med. J. 30: 631-535.
Wesche, W.
1910. On the larval and pupal stages of West African Culicidae. Bull. Entomoi. Res.
1: 7-50, 7 pi.
Wesenburg-Lund,
C.
1921. Contributions to the biology of the Danish Culicidae.
Seisk. Skr. Nat. Math. Afd. (8) 7: l-210, 21 pi.
D.K. Dan. Vidensk.
Harbach: Subgenus Culex in SouthwesternAsia
Weyenbergh,
S.H.
1882.
Los habitantes del Rio Primero, por el doctor H. Weyenbergh.
Montevideo, lmprenta y encuadernacion de Rius y Becchi. 27 pp.
141
. .
White, G.B.
1971. The present importance of domestic mosquitoes (Culex pipiens fatigans
Wiedemann) in East Africa and recent steps towards their control. E. Afr. Med.
J. 48: 266-274.
1975. Notes on a catalogue of Culicidae of the Ethiopian Region. Mosq. Syst. 7:
303-344.
1980. Family Culicidae, pp. 114-148. In: R.W. Crosskey (ed.), Catalogue of the
Diptera of the Afrotropical Region. British Museum (Natural History), London.
Wiedemann,
C.R.G.
1828. Aussereuropaische zweiflugelige Insekten. Vol. 1. Hamm. xxxii + 608 pp.
Wills,
W.M., W.L. Jakob, D.B. Francy,
R.E. Oertley,
E. Anani,
C.H.
Calisher and T.P. Monath
1985. Sindbis virus isolations from Saudi Arabian mosquitoes. Trans. R. Sot. Trop.
Med. Hyg. 79: 63-66.
Wolfs, J.
1947. Culex (Culex) seldeslachtsi, sp. nov. Rev. Zool. Bot. Afr. 40: 224-228.
Worth, C.B. and B. De Meillon
1960. Culicine mosquitoes (Diptera: Culicidae) recorded from the province of
Mocambique (Portuguese East Africa) and their relationship to arthropodborne viruses. An. Inst. Med. Trop. (Lisb.) 17: 231-256.
Yamada, S.
1927. An experimental study on twenty-four species of Japanese mosquitoes
regarding their suitability as intermediate hosts for Filaria bancrofti Cobbold.
Sci. Rep. Gov. Inst. Infect. Dis. Tokyo 6: 559-622, 3 pl.
Zaim, M. and P.S. Cranston
1984. The occurrence of Culex pseudovishnui in Iran. Ann. Trop. Med. Parasitol.
78: 179-l 80.
Zaini,
M., I. lbrahim and T. Al-Samarrae
1983. The culicine mosquitoes of the city of Baghdad (Culicinae - Diptera). Bull.
Endem. Dis. 12 & 13: 115-l 24.
Zimmerman, J.H., H.A. Hanafi and M.M. Abbassy
1985. Host-feeding patterns of Culex mosquitoes (Diptera: Culicidae) on farms in
Gharbiya Governorate, Egypt. J. Med. Entomol. 22: 82-87.
Cot-Mb.Amer. Ent. inst.,vol. 24, no. 1, 1988
FIGURE
1
Mapof southwestern Asia and Egypt
20
30
40
50
60
I
I
I
I
BLACK
SEA
U.
S.
S.
40
30
BA
20
\r
I
ARABIAN
SUDAN
SOUTHWESTER
AND
10
E
144
Contrib.Amer. Ent. Inst.,vol. 24, no. 1, 1988
FIGURE
2
Some Adult Vestiture
A-C.
Thoraces, lateral (left) aspects. MeSL = lower mesanepimeral seta; MScL = lower
mesokatepisternal scales; MScU = upper mesokatepisternal scales; PoSc = postspiracular scales; PrSc = prealar scales.
D-F.
Wings, dorsal aspects of right wings. 2A = vein 2A.
G-J.
Midlegs, anterior aspects.
K-M.
Hindlegs, anterior aspects.
Fig.2
POSC
I----
20+
2A
unlvittatus
sinaiticus
PrSc
mlmetlcus
bitaenlarhynchus
Hindlegs
Midlegs
~-20------l
j---20-+
theilerl
vagans
mimeticus
thelleri
duttoni
perexlguus
Culex
(Cr flex)
Israel
I
02
/
I
1
p&ens
Fig.3
Culex (Culex) pipiens
Israel
Culex
(Culex I quhquefasciafus
Fig.5
Culex
(Culex
)
quinquefasciafus
Pakistan
Fig.6
Ii
1
Culex
(Culex) vagans
Pakistan
/
I
’
I
,
Fig. 8
Fig. 9
N\-\ Culex
(Culexl forrenfium
Fig.10
1
Sweden
/
e-55
\
Culex fCu/ex) decens
Yemen
Arab
Republic
\\I,
Fig.11
02
r-
I Culex
I
(Culex)
Egypt
antennatus
Fig.14
M
I
10
Culex (Culex ) perexiguus
Egypt
dorsal
dorsal
ii
Culex (Culexl pefexiguus /
Fig.18
mesal
dorsal
i
Fig. 22
f,
.
Culex (Culex) ma f tinglyi
Fig. 23
\
k
Id
Fig. 28
Culex
(Culex 1 sinaiticus
Israel
n
1/
Culex
CT
genitalia
fCulex
- P.D.R.
)
dut
toni
Yemen and Ethiopia
pupa - Uganda
c___)
._
B
.‘
*--
mesal
Culex (Culex) dut toni
P D R
Yemen
and
Ethlopla
Fig. 30
Culex
pupa
genitalia
dorsal
(Culex)
- Yemen
P.D.R
Arab
Yemen
sitiens
Republic
and
Saud1
Arabia
Culex
(Culex
I poicil@e
Fig. 33
s
I
’I
VIII
\ I
Fig. 34
Culex (Culex) poich@es
Egypt
/
12
T
1
10
I
/w
mesal
Q
dorsal
Culex (Culex ) bifaeniorbynchus
\
Pakistan
Fig. 38
Culex kulex I frifaeniorbynchus
Pakistan
Culex (Culex ) pseudovishnui
Pakistan
Fig.42
-
Culex (Culex) pseudovishnui
Pakistan
I
\
Contrib.Amer. Ent. Inst.,vol.24, no. 1, 1988
186
Table
Seta
1. Number of branches for pupalsetae of Culexpipiens Linnaeus.a
Abdominal Segments
Cephalothorax
No.
0
1
2
3
4
5
6
7
8
9
10
11
12
13
14
I
Ill
IV
1
lo-24(18)
1
1,2(l)
5-lO(7)
1
I
CT
1,2(2)
5-13(8)
W1)
1
2,3(2)
3-8(4)
4-lO(6)
2-4(3)
4-8(5)
3-5(3)
1
2,3(2)
2-5(3)
l-3(2)
2,3(2)d
3-5d
2,3(2)
1
1
5-9(7)
1
3-8(5)
2-5(2)
2-6(4)
2-5(3)
3-5(3)
l-4(3)
1
2
1
1
1
36-75
l-4(1)
3-5(4)b
3-5(4)
2-5(3)
2-5(3)
3-7(5)
l-5(3)
2,3(2)
2-7(4)
4-9(5)
1
2,3(2)
4-8(5)
l-8(5)
1,2(l)
2-4(2)
2,3(2)
4-7(4)
2,3(2)
_C
Seta
AbdominalSegments
No.
V
VI
0
1
2
3
4
5
6
7
8
9
10
11
12
13
14
1
4-8(6)
1
3-7(5)
1
2,3(2)
3-6(4)
Paddle
VII
VIII
-
3-:(5)
1,2(l)
l-3(2)
-
1IV)
l-3(2)
l-3(2)
3-lO(5)
-
1,W)
W(1)
2-4(3)
1
1
2-5(3)
1,2(l)
l-3(2)
2-5(4)
4-6(5)
1
l-4(2)
1
1,W)
P
1
1,2(l)
2-4(2)
2-6(4)
l-3(2)
3-7(4)
2,3(2)
3-6(4)
3-6(4)
XJ
1
-
_
5-ll(7)
1
a Based primarily on specimens from Egypt and Israel.
bRange(mode).
C Alveolus only.
d Infrequently present.
-
-
1,2(l)
1
187
Harbach: Subgenus Culex in SouthwesternAsia
Table
Seta
2.
Number of branches for pupal setae of Culex quinquefasciafus Say.a
Abdominal Segments
Cephatothorax
I
CT
No.
0
1
2
3
4
5
6
7
8
9
10
11
12
13
14
_C
2,3(2)b
3,4(3)
2,3(3)
2-5(3)
2-4(4)
2-4(2)
0
1
2
3
4
5
6
7
8
9
10
11
12
13
14
1
1
4-7( 5)
1
1,2(2)
3-7(5)
3-5( 5)
2,3(2)
3-7(5)
2-7( 3)
1
l-3(2)
1
1SW
3,4(3)
3,4(4)
1
1,2(l)
2,3(2)
1,2(2)
2,3(3)
1
1y
1,W)
4-l l(7)
l-3(1)
1,2(2)
2-4(3)
IV
1
4-6(4)
1
3-5(4)
l-4(2)
2-4(2)
2,3(3)
2-4(3)
2-4(3)
1
1,W)
1
1
1
AbdominalSegments
Seta
No.
Ill
12-22(16)
1
2,3(2)
4-7(6)
3-9(5)
l-3(2)
tt
V
VI
1
3-5( 4)
1
1,W)
3-5(3)
1,2(2)
2-4( 3)
3-5(4)
2-5(3)
1
1
1,2(l)
1
3-5(4)
1
2,3(2)
2-5(3)
1,W)
3-5(3)
1
2-4(3)
1
1
l-3(2)
1
1
Paddte
VII
VIII
Xl
3-i(5)
1
1,w
2,3(2)
1
-
1
2
2
3-8(4)
1
2-4(3)
3-5(4)
6-8(7)
1
2-4(3)
-
1
1
P
1
1
-
a Based on specimens from southwestern India.
b Range (mode).
c With numerous secondary branches arising from 4-9(6) primary stems.
d Alveolus only.
Contrib.Amer. Ent. Inst.,vol. 24, no. 1, 1988
188
Table
Seta
3.
Number of branches for pupal setae of Culex vagans Wiedemann.a
Cephalothorax
No.
Abdominal Segments
I
I
CT
1
9-l 6( 10)
1
2,3(2)
2-5(4)
4-6(5)
1
1,2(2)
0
1
2
3
4
5
6
7
8
9
10
11
12
13
14
_C
3,4(4)b
3,4(4)
2,3(3)
3,4(3)
4-7(4)
2-4(3)
2
3-9(7)
2-4( 2)
5-l 4(7)
1,2(2)
4-6(4)
1,2(l)
2-4( 2)
4-7( 6)
2-6( 3)
1
2,3(2)
0
1
2
3
4
5
6
7
8
9
10
11
12
13
14
IV
1
6-ll(l0)
1
1,2(2)
4-7( 5)
5-8( 6)
2-5(3)
4-9(6)
3-5(3)
1
1,w
1
1
1”;‘ ’
1,2(l)
5-l&8)
1
4-7( 7)
2-4(3)
2-4( 3)
2-5(4)
3-5(4)
2-4( 2)
1
1,2(2)
1
1
1
AbdominalSegments
Seta
No.
Ill
Paddle
VI
VII
VIII
xl
P
1
4-7(5)
1
-
1
1,2(l)
1
4-6( 4)
l-3(2)
1
1
1
5:(6)
1
2
3,4(4)
2
4,5(4)
1av
W(2)
1
1
1,2(l)
1
1
V
1
5-9( 7)
1
1,2(2)
3-6( 4)
1 Z(2)
3-6( 3)
1
1,W)
1,w
VW
-
-
1
2,3(3)
3-6(4)
1
2,3(2)
7-l l(8)
-
1
1
2-5(2)
4-7(5)
a Based on specimens from Pakistan.
b Range (mode).
CWith numerous secondary branches arising from 6-l O(7) primary stems.
d Alveolus only.
Harbach: Subgenus Culex in SouthwesternAisa
Table
Seta
4. Number of branches for pupal setae of Culex torrentiumMartini.a
Abdominal Segments
Cephalothorax
No.
I
CT
0
1
2
3
4
5
6
7
8
9
10
11
12
13
14
189
_C
2-4(3)b
2-5(4)
2,3(2)
3-5(3)
3-6(4)
3-5(3)
2,3(2)
I
III
IV
1
1
6-l O(7)
1
5-9( 5)
1
2
2-6(4)
l-3(1)
12-16(15)
1
2,3(2)
3-5(3)
3,4(3)
12(l)
2
1
2,3(2)
3-5( 3)
3-5( 4)
1
2,3(2)
4,5(4)
2-6(5)
3,4(3)
2,3(3)
2-5(3)
2-4(2)
1
2,3(2)
5,6(5)
2,3(3)
1
2
1
l-4(3)
5-1?2(5)
2,3(2)
2-4(3)
1,2(2)
1
1
1
AbdominalSegments
Seta
No.
V
0
1
2
3
4
5
6
7
8
9
10
11
12
13
14
1
4-9( 5)
1
1 ,w
2-6(4)
2
3,4(3)
3-7(5)
2-5(3)
1
1
1
1
Paddle
VI
VII
VIII
1
1
2-5(4)
1
2-6(4)
1,2(l)
1
1,2(2)
1,2(2)
1,2(2)
3-6(6)
1
2-4(3)
3-5(4)
1
l-3(2)
1
1
1,2(l)
1,2(2)
2,3(2)
2
2-5(3)
1
2-4(3)
1
1
Xl
1
P
1
1,2(l)
1,2(2)
-
5-lO(7)
-
1
a Based on specimens from Sweden.
b Range (mode).
c With numerous secondary branches arising from 5,6(5) primary stems.
d Alveolus only.
190
Table
Seta
Contrib.Amer. Ent. Inst.,vol. 24, no. 1, 1988
5. Number of branches for pupal setae of Culex decens Theobald.a
Cephalothorax
No.
0
1
2
3
4
5
6
7
8
9
10
11
12
13
14
Abdominal Segments
CT
I
l-3(1)
2
4-7( 5)
4-6( 5)
1,2(l)
2,3(2)
l-3$)
l-3(1)
III
IV
6-i(8)
1
2
3-6(4)
4-6( 5)
2,3(2)
3-6( 6)
3-6(4)
1
1,2(2)
1,2(l)
1
4-7( 6)
1
4-6( 5)
2,3(2)
3,4(3)
2-4( 3)
2-5( 3)
3,4(3)
1
2
1,2(l)
1
AbdominalSegments
V
Paddle
VI
VII
VIII
xl
P
1
3,4(4)
1
2,3(2)
2
2
4-9(6)
1
3-5(4)
2-5(4)
1
l-4(2)
1
-
1
1,2(l)
1
1,2(2)
1
4-6(4)
1
2
24(4)
2
3,4(3)
1
3,4(4)
1
1
l-3(2)
1
1
1
0
1
2
3
4
5
6
7
8
9
10
11
12
13
14
1
25-32(30)
1
2
3-6(4)
3-6(5)
l-3(1)
2
_C
3-5(3)b
3-5(4)
3,4(3)
3,4(3)
3-6(4)
3,4(4)
2
3-6(4)
2,3(2)
5-l O(7)
2,3(2)
3-5(4)
Seta
No.
I
3-i(5)
1
2
3-5( 4)
2
3,4(3)
3,4(4)
2,3(3)
1
2
-
5-8(8)
-
1
a Based on specimens from the Yemen Arab Republic.
b Range (mode).
c With numerous secondary branches arising from 6-9(7) primary stems.
d Alveolus only.
Harbach: Subgenus Culex in Southwestem Asia
Table
Seta
6.
Number of branches for pupal setae of Culex antennatus (Becker).a
Abdominal Segments
Cephalothorax
No.
CT
I
3-5(3)b
_C
0
1
2
3
4
5
6
7
8
9
10
11
12
13
14
1
2-5( 2)
5-9( 7)
4-l O(8)
1
2,3(2)
3-5(4)
2-5(3)
l-4(3)
4-8(6)
2-5(4)
2,3(2)
4-9(7)
2-4( 3)
3-l O(9)
1,2(2)
2-5(5)
2
3
4
5
6
7
8
9
10
11
12
13
14
Ill
IV
1
7-42(31)
1
2,3(2)
3-6(4)
5-7(6)
1
12(2)
1
6-l 3(7)
1
2-4(2)
2-8(4)
6,7(7)
3-6(4)
2-8( 6)
2-5(4)
5-h(8)
2
1
5-i(6)
2-6(4)
4-6(5)
3-6(3)
2-6(3)
2-4(4)
1
2
1
1
1
AbdominalSegments
Paddle
V
VI
VII
VIII
Xl
P
1
5-i(6)
1
2-4( 3)
48(7)
1
2,3(3)
3-6(4)
2
3-6(5)
1
2-5(4)
1
1
l-3(2)
1
4-7(5)
1
3-5(5)
2
Gw
4-12(10)
1
-
12V)
1 a1 1
1,2(l)
0
1
I
_
Seta
No.
191
4-8(4)
2,3(2)
3-6(6)
4-8(6)
2-5(4)
1
1
1,m
1
1
2,3(2)
-
-
1
2-4(3)
2-4(3)
6-8(8)
1
l-5(2)
-
1
1
-
a Based on specimens from Egypt.
b Range (mode).
c With numerous secondary branches arising from 7-1 O(8) primary stems.
d Alveolus only.
-
Contrib.Amer. Ent. Inst.,vol. 24, no. 1, 1988
192
Table
Seta
7.
Number of branches for pupal setae of Culex univitiatus Theobald.a
Abdominal Segments
Cephalothorax
CT
I
I
1
1
1
2-3(3)b
3-6(4)
3,4(3)
3-5(3)
3-6(4)
3-5(5)
2
3-5(5)
2,3(2)
4-l O(4)
1,2(2)
3
_C
14-20(17)
1
7-l l(9)
1
2
4-7(5)
5-9( 6)
2-4(4)
5-9( 6)
3-5(3)
1
2
1
5-8( 6)
5-L(6)
l-3( 2)
5,6(6)
3,4(4)
2,3(3)
2-4( 3)
1
1,2(2)
1
1
1
No.
0
1
2
3
4
5
6
7
8
9
10
11
12
13
14
1
1,2(2)
5-8(5)
6-9(7)
1
2,3(2)
0
1
2
3
4
5
6
7
8
9
10
11
12
13
14
1,2(2)
3-5(4)
5-7(5)
1
2,3(2)
1
IV
Abdominalsegments
Seta
No.
Ill
V
VI
5-i(7)
1
2,3(2)
4-6( 5)
3,4(3)
4,5(4)
4-7( 6)
l-4( 3)
1
1
1
5-:(5)
1
Paddle
VII
VIII
x1
P
1
-
1
2,3(3)
1
3-5(4)
1
3-5(4)
1
1
-
35(4)
2,3(3)
3-5(4)
1
24(3)
1
1
1
2
2
5-7(6)
1
2-4(3)
3,4(4)
1
1
2,3(2)
1
1
1
-
:
7,8(8)
-
1
a Based on specimens from South Africa.
b Range (mode).
c With numerous secondary branches arising from 8-l 2(9) primary stems.
d Alveolus only.
Harbach: Subgenus Culex in SouthwesternAsia
Table
Seta
8.
Number of branches for pupal setae of Culex perexiguus Theobald.a
Cephalothorax
Abdominal Segments
CT
I
3,4(3)b
3-5(3)
2-4(3)
_C
No.
0
1
2
3
4
5
6
7
8
9
10
11
12
13
14
1
2
5-7(5)
4-7( 6)
1
l-3(2)
2,3(3)
3-5(4)
2-4(3)
2
4-6(4)
2,3(2)
4-9(6)
112.‘;’
1 ,w
1
0
1
2
3
4
5
6
7
8
9
10
11
12
13
14
I
Ill
1
13-22( 16)
1
2
3-6(5)
5-7(6)
1
l-3(1)
1
7-l 2(B)
1
IV
3-f(5)
6-8(6)
2-4(3)
4-7( 6)
3-6(3)
1
2-4(2)
1
5-:(5)
1
4-7( 5)
2,3(2)
4-6( 5)
3-5( 3)
2-5(3)
2-4(3)
1
2
1
1
1
2-4(2)
Seta
No.
193
Abdominalsegments
Paddle
V
VI
VII
VIII
1
4-B(5)
1
2
3-5(4)
2-4(3)
3,4(3)
2-5( 4)
2,3(3)
1
1,2(l)
1
1
4-6(5)
1
2-4(3)
1
1
1,2(l)
1
4,5(4)
1,2(l)
3,4(4)
1,2(2)
2
4-6(5)
1
2-4(3)
3-6(3)
1
l-3( 2)
1
1
1
2
24(3)
2,3(2)
3-5(3)
:
2,3(2)
xl
P
1
1
1
-
-
5-B(7)
-
1,2(l)
-
a Based on specimens from Egypt.
b Range (mode).
c With numerous secondary branches arising from 6-1 O(8) primary stems.
d Alveolus only.
Contrib.Amer. Ent. Inst.,vol. 24, no. 1, 1988
194
Table
Seta
9. Number of branches for pupal setae of Culex thei/eriTheobald.a
CT
I
II
Ill
3,4(4)b
3-5(4)
3,4(3)
2-5(4)
2-6(4)
3,4(3)
1,2(2)
4,5(5)
2,3(2)
2-4( 2)
2
3-5(5)
_C
1
7-16( 10)
1
6-l 3( 10)
1
2,3(2)
5-8(7)
7-l O(9)
2-5(3)
5-7( 7)
3-7(4)
1
1,w
1
No.
0
1
2
3
4
5
6
7
8
9
10
11
12
13
14
Abdominal Segments
Cephalothorax
1,2(l)
2
5-8(6)
3-9(4)
2
48(4)
5-l O(6)
1
2
2,3(2)
1
_
IV
1
Seta
No.
0
1
2
3
4
5
6
7
8
9
10
11
12
13
14
5:(6)
2,3(2)
2-4(4)
2-6( 3)
2-5( 3)
3-5(4)
1
1 ,W)
1
1
AbdominalSegments
V
5-L(7)
Paddle
VI
VII
VIII
xl
1
4-7( 6)
1
2,3(2)
5,6(6)
2
2-6(3)
4-7( 6)
2-6( 5)
1
1
1,2(2)
1
3-6(5)
1
1,2(2)
4,5(4)
2
3-6(3)
1
2-6(5)
1
1,2(l)
l-3(2)
1
3-6(4)
1
2-5(4)
2
2
6-9(8)
1
3-5(4)
4-8(4)
1
2,3(2)
1,2(l)
-
1
2,3(2)
-
1
1
1
-
5-lO(6)
-
-
1
a Based on specimens from Egypt and the Yemen Arab Republic.
b Range (mode).
c With numerous secondary branches arising from 7-l O(7) primary stems.
d Alveolus only.
P
1,2(l)
1,2(l)
Harbach: Subgenus Culex in SouthwesternAsia
Table
Seta
10. Number of branches for pupal setae of Culex laticinctusEdwards.a
Abdominal Segments
Cephalothorax
CT
I
I
Ill
IV
_C
1
7-l 4(9)
1
1
213(2)b
2-4(3)
4-7(5)
1
4
1
4-7(7)
l-3(2)
2-5(4)
No.
0
1
2
3
4
5
6
7
8
9
10
11
12
13
14
l-3(1)
2
6-9(6)
58(5)
1,2(l)
2,3(3)
2,3(2)
4-6(4)
3-8(4)
4-6(4)
2,3(2)
4-6(4)
2,3(2)
4-lO(7)
2
_d
1,2(2)
3,4(3)
1,2(l)
0
1
2
3
4
5
6
7
8
9
10
11
12
13
14
1,2(l)
2,3(2)
4,5(4)
4-8(5)
3,4(3)
4-9(8)
2,3(2)
3-6(4)
4-7(5)
1,2(T)
1,2(2)
2,3(3)
3-5(4)
2-4(3)
1
4,5(4)
1
l-3(2)
1
2,3(2)
1
1
1
Abdominalsegments
Seta
No.
195
V
VI
Paddle
VII
VIII
xl
P
1
1
1
1
1
1
3-5(4)
1
3,4(4)
1,2(l)
1,2(2)
3,4(3)
1,2(l)
2-4(3)
-
1
_
2-4(3)
2,3(2)
2-4(3)
1
2,3(2)
-
3,4(3)
1
1
1
1,2(2)
1,2(2)
4-8(6)
1
3-5(4)
3-5(4)
1
l-3(1)
1
1
1,2(2)
3-5(4)
2
2-4(3)
4-6(4)
l-3(2)
1
1,2(l)
1
1
-
6-13(6)
-
1
a Based on specimens from the Yemen Arab Republic.
b Range (mode).
c With numerous secondary branches arising from 7-9(7) primary stems.
d Alveolus only.
* Frequently absent, often present on one side only.
196
Table
Seta
Contrib. Amer. Ent. Inst.,vol. 24, no. I, 1988
II.
Number of branches for pupalsetae of Cu/exmatting/yiKnight.a
Cephalothorax
No.
Abdominal Segments
CT
I
0
I
2
3
4
5
6
7
8
9
IO
II
I2
I3
I4
9/6 c
I/I
2f2
5l4
6'6
713
3l4
3/- b
3l3
3J3/7l7
W2
2/2/2
2/2
8fIO
4/4
2l2
3l2
Ill
IV
l/I
17114
I/I
2l2
W2
2f2
5l5
I/I
I/I
14/10
I/I
2l2
2l3
3f2
5/5
5L3
I/I
III
2l2
III
III
7/7
I/I
412
I/I
2J2
5f3
I/I
2/I
I/I
2l2
I/I
I/I
I/I
I/I
IO/9
3f2
-dk
Il-
Seta
I
AbdominalSegments
No.
V
VI
VII
0
I
2
3
4
5
6
7
8
9
IO
II
I2
I3
I4
I/I
414
III
2/I
2f3
I/I
314
412
312
I/I
I/2
I/I
I/I
3l4
I/I
2/2
2/2
I/I
4/3
-/I
2/3
I/I
-II
I/I
I/I
3l2
III
I/2
I/I
2l4
-t9
-II
-I3
-I2
III
I/I
I/I
I/I
Paddle
VIII
xl
P
I/I
I/I
I/I
III
I/I
-/I2
a Based on a single specimen inthetype series from the Yemen Arab Republic.
b Branches of setae on right/left sides.
c Numberofprimary stems,with numerous secondary branches.
d Alveolus only.
Harbach: Subgenus Culex in SouthwesternAsia
Table
Seta
12. Number of branches for pupal setae of Culex simpsoniTheobald.a
Abdominal Segments
Cephalothorax
No.
I
I
CT
_C
415(4)b
4-6(5)
3
3-5(4)
4-6(5)
2-6(4)
2,3(2)
4-6(5)
3,4(3)
3-8(8)
2
3-5(4)
1
2
4-6( 6)
2-6( 5)
1
1,2(2)
0
1
2
3
4
5
6
7
8
9
10
11
12
13
14
IV
1
12-15(12)
1
2
47(5)
7-9(7)
4-6(5)
5-8(7)
l-3(2)
?
?
3-6(4)
1
2
1
1
8-12(12)
1
4-7(5)
24(2)
46(5)
46(5)
35(3)
3,4(3)
1
2
1
1
1
AbdominalSegments
Seta
No.
Ill
1
12-l 9( 16)
1
2
3-7(6)
5-7(6)
1
2
0
1
2
3
4
5
6
7
8
9
10
11
12
13
14
197
Paddle
V
VI
VII
VIII
xl
P
1
7-9( 8)
1
l-3(2)
5,6(6)
2,3(2)
4-6( 5)
4-7( 6)
3,4(4)
1
l-3(1)
1
1
3-7(7)
1
2,3(2)
3-5(4)
2
1
-
1
1
1
46(5)
1
3-5(4)
1
1
1,2(l)
1
3,4(4)
1
2-4(4)
2,3(2)
2,3(2)
6-8(7)
1
3-5(4)
4-6(4)
1
7-9(8)
-
2,3(2)
-
1
1
1
1
2,3(3)
-
-
-
a Based on specimens from Ethiopia and Tanzania.
b Range (mode).
c With numerous secondary branches arising from 6-9(8) primary stems.
Contrib.Amer. Ent. Inst., vol. 24, no. 1, 1988
198
Table
Seta
13. Number of branches for pupal setae of Culex sinaiticusKirkpatrick.a
Abdominal Segments
Cephalothorax
No.
I
CT
0
1
2
3
4
5
6
7
8
9
10
11
12
13
14
_C
3,4(4)b
4,5(5)
2,4(2)
2-4(4)
3-6(5)
3-5(4)
2
4-6( 5)
2,3(3)
4-7(5)
l,W
1,W)
4-6( 6)
2-5(3)
1
2
1
15-22( 18)
1,2(l)
1,2(2)
4-7(6)
5,6(5)
1
142)
1
8-16(11)
1
2
4-6(5)
5-8(6)
3-5(4)
4-7(6)
2-6(4)
1
2
1,2(l)
1
7-l 2(7)
1
1
1
_
1
IV
5&5)
l-3(2)
5,6(6)
3-6(5)
2-5(3)
2-5(3)
1
1,2(2)
1,2(l)
3,i(4)
AbdominalSegments
Paddle
VII
VIII
XI
P
1
-
1
1
1
1
2,3(2)
-
1
l-3(1)
1
3-5(4)
1,2(l)
3,4(4)
2,3(2)
2
5-10(6)
1
3-5(4)
3-5(4)
1
l-3(2)
1
1
V
VI
5-l&7)
1
l-3(2)
4-6( 5)
2,3(2)
3-5(5)
4-6( 6)
2-4(4)
1
1
1,2(l)
5-:(5)
1
2
3,4(3)
2
4-6(5)
1
3-5(3)
0
1
2
3
4
5
6
7
8
9
10
11
12
13
14
Ill
1-3_(i)
Seta
No.
I
6-10(7)
-
-
_
1
1
a Based on specimens from Egypt (Sinai) and Israel.
b Range (mode).
c With numerous secondary branches arising from 5-8(8) primary stems.
d Alveolus only.
Harbach: Subgenus Culex in SouthwesternAsia
Table
Seta
14. Number of branches for pupal setae of Culex duttoniTheobald.a
Cephalothorax
No.
0
1
2
3
4
5
6
7
8
9
10
11
12
13
14
Abdominal Segments
I
CT
_C
2,3(2)b
2,3(2)
2-4(3)
2-5(3)
2-6(4)
2-4(3)
2
2,3(2)
2,3(2)
5-l 3(8)
2-5(3)
2-5(3)
V(1)
2
6-l l(9)
3-6(4)
1,2(l)
2-5(3)
0
1
2
3
4
5
6
7
a
9
10
11
12
13
14
I
Ill
IV
1
17-26( 18)
1,2(l)
1aq
4-7(6)
3-5(3)
1,2(l)
3,4(4)
1
4-6(4)
1
2
3-6(4)
4,5(4)
1,w
3-8(4)
2-5(4)
1
2,3(2)
l-3(1)
1
2-6(4)
1
4-8(6)
l-3(2)
1-w)
1,w 1
3-5(4)
1-w)
1
2
1,2(l)
1
1
1t”d2’
2-5(2)
Seta
No.
199
AbdominalSegments
Paddle
V
VI
VII
VIII
1
l-4(2)
1
1,W)
4-6( 5)
l-3(1)
l-3(1)
4-9( 6)
2-4(4)
1
1
l-3(1)
1
2,3(2)
1
l-3(2)
3-5(4)
1
2-4( 2)
1
2
2
l-3(2)
6-16(11)
1
4-9(4)
3-6(3)
V(1)
V(2)
1
1
1
l-3(3)
1
2-5(4)
1
l-3(1)
2-4(2)
Xl
P
1
1a1 >
02(O)
-
-
7-ll(10)
-
-
a Based on specimens from Uganda and Cameroon.
b Range (mode).
c With numerous secondary branches arising from 7-l 2(7) primary stems.
d Alveolus only.
Contrib.Amer. Ent. Inst.,vol. 24, no. 1, 1988
200
Table
Seta
15. Number of branches for pupal setae of Culex Miens Wiedemann.a
Abdominal Segments
Cephalothorax
No.
CT
I
II
Ill
1,2(l)
1a1 1
_C
l-3-$)
1
5-8(6)
1
2
4-7(5)
4-6(5)
1,2(l)
5-9(7)
3-5(3)
1
1,2(2)
1
3-5(5)
1
2,3(2)
58(7)
4-7(6)
1
3
4-:0(6)
1
1,2(2)
4-6(4)
4,5(4)
1
2,3(2)
1
1
0
1
2
3
4
5
6
7
8
9
10
11
12
13
14
213(2)b
3,4(4)
l-3(2)
2-4(3)
3,4(3)
2,3(2)
2
2
2,3(2)
59(7)
2-4(3)
2-5(3)
1,2(l)
5:(6)
2,3(3)
2,3(2)
1,2(2)
3-5(5)
3,4(3)
1
l-3(2)
1
AbdominalSegments
Seta
No.
IV
V
VII
VIII
1
1
1,2(l)
141)
1
2
1
2-4(2)
1,2(l)
1,2(2)
6-ll(8)
1
2-4(4)
3-5(4)
1
2,3(2)
1
1
0
1,2(l)
1,2(l)
1
2
3
4
5
6
7
8
9
10
11
12
13
14
3,4(4)
1
2,3(2)
1
1,2(2)
24(3)
2
1,2(2)
5,62(5)
1,2(2)
1,2(2)
5-7( 6)
2-4(3)
1
1,2(l)
1,2(l)
1
Paddle
VI
2-w
2,3(2)
xl
P
1
1
1
-
-
9-14(12)
-
1
a Based on specimens from the Yemen Arab Republic.
b Range (mode).
c With numerous secondary branches arising from 8-12(9) primary stems.
d Alveolus only.
Harbach: Subgenus Culex in SouthwesternAsia
Table
Seta
16. Number of branches for pupal setae of Culex poicilipes (Theobald).a
2
3
4
5
6
7
8
9
10
11
12
13
14
Abdominal Segments
Cephalothorax
No.
0
1
I
II
Ill
IV
l-3(1)
2,3(2)
8-17(14)
4-13(4)
1
2,3(2)
1
8-15(11)
1,2(l)
2,3(2)
5-l O(7)
7-l 4(8)
1
l-3(2)
1’2;’)
1
1
7-l 4(9)
1
2
6,7(6)
8-20( 10)
3-5(3)
6-l O(6)
5-8(5)
1
2-4(3)
1,2(2)
_
5:(8)
1
5-l l(6)
3-6(4)
3-5(3)
3-6(4)
48(7)
4-7(5)
1
2,3(3)
1,2(l)
CT
_C
3-5(4)b
3-5(4)
2-5(3)
2-4(3)
4-6( 6)
2-4(3)
2,3(2)
2-4(4)
2-4(2)
7-l 6(9)
2,3(2)
3-6(4)
l,W)
1
Seta
AbdominalSegments
No.
V
VI
VII
0
1
1
3,4(3)
1
V(3)
1,2(l)
3-5(4)
2-4(3)
2-4(4)
S-17(8)
1
4-8( 7)
2-5(3)
1
2-4(3)
1
2
3
4
5
6
7
8
9
10
11
12
13
14
201
5,W)
1a1 )
2-5(3)
5-l O(6)
2
3-6(4)
6-13(7)
3-7(5)
1
1
1,w
1,2(l)
W(3)
3-7(5)
2
3-6(5)
1,2(l)
3-5(4)
1
1
l-3(3)
Paddle
VIII
l-me
2-4(3)
-
5-lO(8)
xl
P
1
_
1
1
-
-
-
1
a Based on specimens from Egypt.
b Range (mode).
c With numerous secondary branches arising from 8-l l(9) primary stems.
d Alveolus only.
e Many branches.
Contrib.Amer. Ent. Inst.,vol. 24, no. 1, 1988
202
Table
Seta
17. Number of branches for pupal setae of Culex mimeticus NO&~
Abdominal Segments
Cephalothorax
CT
No.
I
I
Ill
IV
1
_C
l-4(3)
1
3-6(4)
1
2
5-l O(6)
2-5(3)
1,2(l)
5-9(6)
2,3(3)
1
1,W)
1
1
25(3)
1
2-7(4)
1
1
0
1
2
3
4
5
6
7
8
9
10
11
12
13
14
213(2)b
2-4(3)
2-4( 2)
l-3(2)
2,3(2)
2-4( 2)
2
2,3(2)
2-4( 2)
4-6(5)
2,3(2)
2-4( 2)
1
1,2(2)
l-3(2)
1
0
1
2
3
4
5
6
7
8
9
10
11
12
13
14
1
2/1e
_d
1
VII
V
VI
1
l-3(2)
1
1,2(l)
2-5(4)
2
l-3(1)
2-8(3)
l-3(2)
1
1
1
1
1,2(l)
1
1m
2-v)
2
1,2(l)
1
2,3(2)
1
1
1
1
l-3(2)
1
l-3(2)
l-3(1)
1
3-7(3)
1
2-4( 2)
2-4( 2)
1
1,W)
1
1
1
24(2)
2-4(2)
1
2-6(3)
l-4(3)
1
1 ,w
1
Paddle
AbdominalSegments
Seta
No.
3-6(4)
3-5(3)
1
l-3(2)
5-i(5)
2-7(3)
VIII
xl
P
1
-
1
1
1
1,2(l)
-
-
4-9(6)
-
1
a Based on specimens from Israel.
b Range (mode).
c With numerous secondary branches arising from 5-8(6) primary stems
d Alveolus only.
e Branches of setae on right/left sides of one specimen.
Harbach: Subgenus Culex in SouthwesternAsia
Table
Seta
18. Number of branches for pupal setae of Culex bitaeniorhynchusGiles.a
Cephalothorax
No.
0
1
2
3
4
5
6
7
8
9
10
11
12
13
14
Abdominal Segments
CT
I
0
1
2
3
4
5
6
7
8
9
10
11
12
13
14
I
1
1-w
1
1
_C
1-3(2)b
2-4( 2)
l-4( 2)
l-3(2)
2,3(2)
l-4(2)
1,2(2)
1,2(2)
1,2( 2)
5-l O(7)
1,2(l)
l-3(2)
1
1
4-8(6)
3-9(3)
1,2(l)
l-3(2)
Ill
IV
1
1
2-6(4)
6-1 O(7)
1
1
1
3-7(4)
2,3(3)
2,3(3)
1
l-5(3)
24(3)
1,2(l)
2
1,w
1
1
2-6(4)
2-5(4)
1,2(l)
4-6(6)
3-6(5)
1,W)
3-6( 5)
2-5( 4)
1
1
1
1-y)
2,3(2)
1
Seta
No.
203
AbdominalSegments
Paddle
V
VI
VII
VIII
1
1
1,2(2)
1
1,2(l)
2,3(3)
2,3(2)
1
1,2(l)
2-4(3)
1
1
1,2(l)
1
1,W)
1
1,2(l)
l-3(2)
1,2(l)
3-8(5)
1
2-4(3)
3-5(3)
1,2(l)
l-3(2)
1
2,3(2)
1
l-3(1)
2-5(4)
2,3(2)
4-:(5)
l-3(2)
1
1
1
xl
P
1
1
1
2
_
-
6-lO(6)
-
-
1
a Based on specimens from Pakistan and the Yemen Arab Republic.
b Range (mode).
c With numerous secondary branches arising from 5-l O(7) primary stems.
d Alveolus only.
Contrib.Amer. Ent. Inst.,vol. 24, no. 1, 1988
204
Table
Seta
19. Number of branches for pupal setae of Culex tritaeniorhynchus Giles.a
Abdominal Segments
Cephalothorax
No.
0
1
2
3
4
5
6
7
8
9
10
11
12
13
14
_C
3,4(3)b
3-5(3)
2,3(3)
1,W)
4-6(5)
3-6(4)
1,w
WV)
2,3(2)
6-l 2(9)
2
3-5(4)
7-l:(9)
4-7(6)
1
2-5(4)
0
1
2
3
4
5
6
7
8
9
10
11
12
13
14
Ill
IV
1
20-36( 28)
1
2
4-7(5)
5-9(6)
1
2-4(2)
1
lo-15(11)
1
2
4-7(6)
68(8)
4-7(6)
4-8(5)
2-6(5)
1
1,2(2)
1,2(l)
1
1
7-12(10)
1
4-7(6)
l-4(3)
4-7(5)
3-6(5)
2-5(3)
3,4(3)
1
2
1
1
AbdominalSegments
Seta
No.
II
I
CT
Paddle
V
VI
VII
VIII
xl
P
1
7-1 O(8)
5,i(5)
1
4-6(5)
W
‘ (3)
1,2(2)
9-13(12)
1
4-6(4)
3-5(5)
1
2-5(2)
1-
1
1
1
3,4(3)
:
1,2(l)
1
7-9(7)
1
2,3(2)
4-6(5)
2
6-8(7)
1
2-5(3)
1
1
1,2(l)
1
1
2,3(2)
5-7( 6)
2
6,7(6)
5,6(6)
2-5(4)
1
-
4-8(5)
-
-
a Based on specimens from Pakistan.
b Range (mode).
c With numerous secondary branches arising from 6-9(7) primary stems.
d Alveolus only.
Harbach: Subgenus Culex in SouthwesternAsia
Table
Seta
20.
Number of branches for pupal setae of Culex pseudovishnuiColless.a
Abdominal Segments
Cephalothorax
No.
0
1
2
3
4
5
6
7
8
9
10
11
I
CT
2,3(2)
6-l 2(6)
3-7(4)
1
2-4(2)
2
2-5(3)
2
3
4
5
6
7
8
9
10
11
12
13
14
IV
1
lo-16(10)
1
3-28(S)
5-9(6)
3-5(4)
3-9(7)
3-6(4)
1
2
1
1
2
_d
2,3(2)
3-l O(9)
Ill
1
9-15(12)
1,2(l)
2,3(2)
48(7)
68(7)
1
2
1
1,2(l)
1
Paddle
V
VI
VII
VIII
1
1
58(6)
1
2,3(2)
4,5(4)
2,3(2)
5-7(5)
1,2(l)
4,5(4)
1
1
l-3(1)
1
3-5(4)
1
3-5(4)
2,3(2)
1,2(2)
5-11(B)
1
2-5(3)
3-6(4)
1
2-4(3)
1
1
1
4-B(7)
1
2,3(2)
5-7(5)
2-4(2)
5,6(5)
5-8( 7)
3-5(4)
1
1
1,2(l)
2-4(2)
Xl
P
1
1,2(l)
1
-
6-9(B)
-
-
_
1
7-l i(9)
1
2-9(4)
24(4)
5-7(6)
4-6(5)
3-6(5)
3-5(4)
1
2
1
1
AbdominalSegments
Seta
0
1
II
_C
2-4(3)b
3-5(4)
3
l-3(2)
3-5(4)
2-5(3)
2
3-5(5)
12
13
14
No.
205
1
a Based on specimens from Pakistan and India.
b Range (mode).
c With numerous secondary branches arising from 7-12(11) primary stems
d Alveolus only.
Contrib.Amer. Ent. Inst.,vol.24, no. 1, 1988
206
Table
21. Number of branches forfourth-instarlarvalsetae
of Culex pipiens
Linnaeusa
Seta
Abdominal Segments
Thorax
Head
No.
C
P
M
0
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
1
9-26(17)b
1
1
1
-
-
1
2-5(3)
1
2-5(3)
l-3(2)
2-5(4)
2-5(4)
O-:(o)
1
1
4-7(5)
3-6(4)
8-13(10)
2-4(2)
3-7(5)
2,3(2)
2,3(2)
2,3(2)
2-4(3)
1,2(l)
2,3(2)
1
1
2,3(2)
2,3(2)
1
1
3-8(5)
1
1A1 >
0
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
1,2(2)
1
1
1
5-9(5)
4-8(5)
1
l-4(2)
1
14-27(23)
15-26(20)
I
l-5( 5)
1
1,2(l)
7-12(8)
2-7(5)
1
6-lO(10)
8-18(14)
4-9(6)
1
l-3(2)
1
3-6(5)
3,4(3)
l-3(2)
l-5(2)
l-3(1)
3-lO(4)
l-3(2)
l-3(2)
II
II
1
l-4(2)
1
1
l-4(2)
1
1,2(l)
1,2(l)
l-4(2)
l-4(2)
3-7(4)
l-3(2)
2-4(3)
3-6(4)
1
1
1
2-5(3)
l-3(2)
7-23(15)
2,3(2)
4-9(6)
1
1
1
l-3(2)
l-3(1)
2-5(3)
1
3-5(3)
Abdominal Segments
Seta
No.
T
IV
1
2-4(2)
1
1a1 )
1,2(l)
l-3(2)
2
5-lO(7)
1
1
1
l-4(3)
1a1 1
2-4(3)
1
V
1
l-3(2)
1
1
2-5(3)
l-4(2)
2
4-8(6)
1,2(l)
1
1
l-4(3)
1
2-4(3)
1
VI
VII
VIII
1
1
2-5(3)
1
1
4-9(5)
1
6-9(8)
1
3-5(4)
2,3(2)
1,2(l)
1
l-3(2)
l-4(3)
2
1,2(l)
1
1-w)
1,2(l)
lo-20(10)
1
2-5(3)
1
1
2-6(4)
28(3)
1
2-4(3)
1
22-43(28)
1
24(2)
1
2-4(3)
1
a Based primarily on specimens from Egypt and Israel.
b Range (mode)obtained from counts made on at least40 setae.
c Siphon occasionally with a fifth seta 1-S on one or both sides.
la-SC
lb-S,
lc-s,
Id-S,
X
l-3(1)
2,3(2)
5-l&6)
Harbach: Subgenus Culex in SouthwesternAsia
Table
Seta
22.
Number of branches for fourth-instar larval setae of Culex quinquefasciatus
Say?
Head
No.
C
1
1
1
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
0
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
M
T
I
1,w
2-5(2)
2-6(2)
1
1
l-4(2)
1,2(l)
l-3(2)
2-4(3)
3-6(5)
1,2(l)
6-lO(8)
5-lO(6)
1
2-5(4)
l-3(2)
1
6-8(7)
g-16(11)
4-7(5)
1
l-3(3)
1
3-7(6)
3,4(3)
2
3,4(3)
2-5(4)
1
1
1
4-6(5)
4-6(5)
1
2-4( 2)
1
16-36( 22)
15-33( 19)
1
II
J,W)
1,w
4-:(5)
3-5(5)
7-lO(8)
2-4(3)
2-7(4)
l-3(3)
2-4(3)
l-4(2)
2-5(4)
1,2(l)
2-4(3)
Abdominal Segments
Thorax
P
Seta
No.
207
1,2(2)
1
1
3-7( 6)
l-3( 2)
2-4(3)
1,2(l)
1,2(2)
lo-25(17)
ltl
1
1
1
1
l-4(1)
1,2(2)
2
4-7(6)
1
1
1
l-3(2)
1
2-4(3)
1
Abdominal Segments
IV
V
VI
VII
VIII
1
1,2(2)
1
2
1
3-5(3)
1
4-8(6)
1
1
1
1
1 ,W)
2-4(3)
l-3(2)
2-6(3)
1
1,2(l)
1
2-5(4)
1
5-9(7)
1
3-6(5)
2
5-lO(6)
5-:(6)
1
1,2(l)
1
1m )
1,2(l)
1,2(l)
1
1
2,3(2)
1,2(l)
l-4(3)
1
1,2(l)
1
1
2-4(2)
1
2-4(3)
1
1,w
2
1
2-5(3)
1
1
2-4(2)
1
22-46(31)
1
X
1
2,3(2)
1
3-lO(6)
-
8-15(12)
1
2-5(4)
3-6(4)
1
2,3(2)
1
2-4(3)
1
aE3asedon specimensfromtheYemen Arab Republicandlndia.
b Range (mode)obtainedfrom
counts made on at least10 setae.
c Siphon infrequently with only 3 pairs of setae; rarely with an additional unpaired seta.
208
Table
Seta
No.
0
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
Contrib.Amer. Ent. Inst.,vol. 24, no. 1, 1988
23. Number of branches for fourth-instar larval setae of Culex vegans
Wiedemann.a
Thorax
Head
C
P
M
T
I
1
1
9-13(12)b
1
1
1
2
1
1
2-4(2)
1
l-3(2)
1
l-3(2)
1
1
1
4-8(6)
4-7(5)
1
2-5(3)
3-lO(6)
1
1,2(l)
5-8(6)
4-8(5)
7-l2(8)
2-4(3)
3-7(7)
2-4(2)
2-4(2)
2,3(3)
2,3(3)
1,2(l)
3-5(3)
2,3(2)
1
1
4-7(5)
1
1,w 1
0
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
1,2(l)
2-6(4)
3-6(4)
1
1
6-9(6)
ll-17(11)
4-8(6)
1
1,2(l)
1,2(l)
8-12(9)
4-7(6)
2-4(3)
2,3(2)
l-3(2)
1
4-9(6)
l-3(1)
l-3(2)
2,3(2)
1
3-7(4)
-
2,3(2)
1
lo-31(24)
15-30(24)
II
1
2-4(3)
1
1
3,4(4)
l-3(2)
3-5(4)
3-6(4)
1
1
1
2,3(2)
1,2(2)
lo-25(18)
Ill
1
l-3(2)
1
1
l-3(2)
l-3(2)
2,3(2)
4-8(5)
1
1
1
l-3(2)
1,2(l)
2-6(4)
1
Abdominal Segments
Seta
No.
Abdominal Segments
IV
1
2
1
1
1
l-4(3)
2
4-9(6)
1
1
1
l-3(2)
1,2(l)
3-5(3)
V
1
2
1
1
2-4(2)
2-5(3)
2
5-7(5)
1,2(l)
1
1
2,3(2)
1
3-5(3)
1
VI
VII
VIII
1
1
3-5(3)
1
1
1
5,6(5)
1
6-9(7)
1
4-i(5)
4,5(4)
-
1,2(l)
1
1
1,2(2)
24(3)
2,3(2)
1,2(l)
l-4(3)
1,2(l)
2-4(2)
1
27-36(29)
1
2-4(3)
lo-16(10)
1
12(l)
3-6(6)
3-6(5)
1
1,3(2)
1
3-5(4)
1
a Based on specimens from Pakistan.
b Range (mode) obtained from counts made on at least IO setae.
c Siphon occasionally with an additional unpaired seta.
X
1
1,2(l)
l-3(2)
Harbach: Subgenus CulexinSouthwestem Asia
Table
Seta
No.
0
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
24. Number of branches for fourth-instar larvalsetaeof Culex torrentium
Martini.a
0
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
Abdominal Segments
Thorax
Head
C
P
M
T
I
1
1
9-13(11)b
1
1
1
2
1
1
2-4(3)
l-4(2)
1
3-7(4)
2
2-6(5)
1
l-3(2)
7-12(9)
4-7(5)
1
1,2(l)
4-7(5)
3-5(4)
I3-lO(8)
2-4(2)
2-5(3)
2,3(2)
2-4(3)
2,3(3)
2,3(3)
l-3(1)
2,3(2)
2
1
1
4-8(6)
1
1
3,4(4)
2-4(3)
1
1
2,3(2)
1
1,2(l)
1
3-7(5)
4-6(6)
1
l-3(2)
1
13-20(18)
12-25(19)
4-8(6)
8-15(12)
5-7(6)
1
1,2(2)
1
Ill
1
l-3(2)
1
4,5(4)
1
1
3,4(3)
2,3(2)
1,2(l)
3-5(4)
l-4(1)
3-5(4)
3-5(4)
1,2(l)
1
l-3(1)
2
3-7(4)
1
l-3(2)
1
4-6(4)
1
1
2-4(3)
1,2(l)
1
1,2(2)
1,2(l)
38(5)
II
2,3(2)
12-19(17)
2,3(2)
1,2(l)
4-6(4)
1
3,4(4)
Abdominal Segments
Seta
No.
209
IV
1
3-6(4)
1
1,2(l)
1
l-4(2)
2
4-7(6)
1
1,2(l)
1
1,2(2)
1
4-6(4)
1
V
VI
VII
VIII
1
3-5(4)
1
1
2-5(3)
1
4-6(4)
5-i(6)
1
1
1,2(2)
2-5(3)
2
1
1
2
1,2(l)
l-4(3)
8-13(10)
1
3-i(3)
l-4(2)
2
4-7(5)
1,2(l)
1,2(l)
1
2
1
3-5(4)
1
24(Z)
1,2(l)
1
l-3(2)
1
17-34(28)
1
1
6-9(7)
1
3-5(4)
38(6)
2-6(3)
1
la-S,c
lb-S,
lc-s,
Id-S,
X
1,2(2)
2,3(2)
1
3-8(6)
2,3(3)
l-3(3)
2
l-3(2)
1,2(l)
1
3,4(3)
1
aBasedonspecimensfromSweden.
b Range (mode)obtained from counts made on at least10 setae.
CSiphon often with a fifth seta 1-S on one or both sides;seldomwith3
on one side.
210
Table
Seta
Contrib.Amer. Ent. Inst.,vol. 24, no. 1, 1988
25.
Number of branches for fourth-instar larval setae of Culex decens
Theobald.a
Thorax
Head
No.
C
P
0
1
13-19(15)b
1
1
1
2,3(2)
1
1
3
2
1
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
1
2,3(2)
5-;6)
4-6(5)
4-8( 7)
l-3(2)
1,2(2)
2-4(3)
3-5(4)
l-3(2)
3-6(4)
Abdominal Segments
M
5-:(6)
1
1
T
I
-
1,2(l)
2-4(3)
1
1,2(2)
1
1
1
4,5(4)
3,4(4)
1
2-4(3)
1
18-28(24)
16-24(21)
_
1,2(l)
1,2(2)
l-3(1)
3-5(3)
4-7(6)
1,2(l)
2
6-l l(9)
7-9( 8)
3
2
5-:(6)
IO-18(11)
5,6(6)
1
24(3)
1,2(l)
5-7(6)
12(l)
3,4(4)
1
4-8( 5)
1,2(2)
l-3(2)
II
1
1
1,2(l)
1,2(2)
4-6(5)
2,3(2)
3
4-7(5)
1,qq
1
1
2-5(3)
2-4(3)
14-23(17)
IU
1
l-3(2)
1
1,2(l)
1,2(l)
l-4(2)
1
4-7(5)
1,2(2)
1
1
2,3(2)
1,2(l)
2-5(4)
1
Abdominal Segments
Seta
No.
IV
V
0
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
1
1
1
2
1
2-4(2)
l-3(2)
5-7(6)
2
1
1
2,3(2)
1
3-6(4)
1
1
1
1
1
4-6(5)
2,3(2)
1
4-8(6)
2
1
1
2,3(2)
1
3-5(4)
1
VI
1
2,3(3)
1
1
2,3(3)
24(3)
1
1
24(4)
1
1
2,3(3)
1
32-56(42)
1
VII
VIII
1
2-5(4)
1
2,3(2)
1
2-4(2)
II-21(15)
1
24(2)
1,W)
1
1
5-7( 6)
1
5-7( 7)
1
3,4(3)
1a-S,C
1b-S,
lc-s,
1d-S,
24(3)
1
2-4(3)
1
a Based on specimens from the Yemen Arab Republic (primarily) and Senegal.
b Range (mode) obtained from counts made on at least 10 setae.
c Siphon occasionally with a fifth paired or unpaired seta.
X
1,2(2)
2
1
3-9(7)
l-3(2)
2,3(2)
2,3(2)
l-3(2)
Harbach: Subgenus Culex in SouthwesternAsia
Table
Seta
26
Number of branches for fourth-instar larval setae of Culex antenna&s
(Becker).a
Abdominal Segments
Thorax
Head
No.
C
P
M
0
1
2
3
4
5
6
7
a
9
10
11
12
13
14
15
1
1
17-29(23)b
1
1
1
2
1
1
3
2
1
1,2(l)
6-W)
1
l-3(2)
4-a(5)
1
l-3(2)
1
1
1
4,5(5)
4,5(4)
1
2-4(2)
1
26-43(36)
14-27(21)
1
1a1 1
2,3(2)
2,3(2)
6-a(8)
3-5(5)
4-7( 5)
3,4(3)
2,3(2)
3-5(4)
2-4(3)
1,w
5-a(7)
1
0
1
2
3
4
5
6
7
a
9
10
11
12
13
14
T
II
I11
1
l-3( 2)
1
2-4(3)
5-9(6)
2,3(3)
3,4(3)
5-9(6)
1,2(l)
1
1
1
3,4(4)
1
2,3(2)
2-4( 2)
2-5(3)
l-3( 2)
7-1 O(9)
12(l)
1
1
2,3(2)
1,W)
4-7( 5)
1
I
-
242)
2,3(2)
4-7(5)
3-8(4)
1,2(l)
1
5-8(7)
g-16(12)
46(5)
l-4(4)
1
2-5(3)
a-14(1 1)
4-9( 5)
3
2
3,4(3)
1
4-6( 5)
l-4( 2)
2,3(2)
2-L(3)
l-3(2)
7-14(a)
2-4(3)
2,3(2)
15-30(20)
Abdominal Segments
Seta
No.
211
IV
V
VI
VII
1
3,4(3)
1
2-4(3)
l-3(2)
2,3(2)
3
6-l 2(9)
1,w
1
1
2,3(2)
1
2-4(3)
1
1,2(l)
5-a(6)
l-3(2)
1
2,3(3)
1
1
3,4(3)
2,3(2)
1
2,3(3)
1
3-6(5)
3-E(5)
1
U(2)
a-i 3(9)
2
1
1
1,W)
1
4-6(5)
1
142)
1
1
2,3(3)
1
31-47(39)
1
3-i(4)
1
2,3(2)
12-20( 15)
1a1 1
4-7(5)
24(2)
1
24(3)
1
3-6(4)
1
VIII
6-&a)
1
7-9(7)
1
4,5(4)
15
a Based on specimens from Egypt.
b Range (mode) obtained from counts made on at least 10 setae.
c Seta 1-S often only in 5 pairs. Seta 1a-S often absent; sometimes single.
X
3,4(4)
2,3(2)
5-L(a)
-
Contrib.Amer. Ent. Inst.,vol.24, no. 1, 1988
212
Table
Seta
27
Number of branches for fourth-instarlarval
Theobald.a
C
P
0
1
8-26(19)b
1
1
1
2
1
1
O,:(o)
1
1
2-4(3)
2,3(3)
6-8(6)
2-6(5)
3-9(7)
2-4(3)
2,3(2)
2-5(4)
3-5(4)
1,2(2)
3-8(6)
0
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
T
I
1w 1
l-3(2)
2-5(3)
2-7(2)
1
l-4(3)
1
1
1
2-w
1,2(l)
2-6(5)
2-7(4)
l-3(1)
1
5-7(6)
7-16(10)
2-4(2)
5-15(12)
4-12(5)
5,6(5)
2-4(2)
1
2-7(4)
l-3(2)
l-3(2)
M
2,3(3)
2,3(2)
1
1,2(l)
6-10(7)
1
1
5,6(5)
3-6(4)
1
l-4(2)
1
20-41(29)
lo-28(21)
II
1
l-3(2)
1
l-3(2)
3-7(5)
l-3(3)
3
2-7(5)
3,4(3)
2,3(2)
2-:(3)
l-5(2)
5-lO(7)
1,2(l)
1,2(l)
1
3,4(3)
l-3(2)
9-22(20)
I11
1
2-4(3)
1
l-3(2)
l-4(3)
l-4(2)
2,3(2)
4-9(8)
1,2(2)
1
1,2(2)
l-3(2)
l-3(2)
3,4(4)
1
Abdominal Segments
Seta
No.
Abdominal Segments
Thorax
Head
No.
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
setae of Culexunivittatus
IV
V
1
2-4(3)
1
2-4(3)
1
2-l(3)
1W)
1av
3-9(6)
l-3(2)
l-3(2)
2,3(2)
3-11(8
l-3(2)
1
1SW)
l-3(2)
1,2(l)
3-5(4)
1
VI
VII
VIII
1
1
3-7(5)
1
4-9(6)
1
3-6(4)
1
l-5(4)
7-23(16)
1
3-7(3)
2-5(4)
1
2-4(3)
1
3-6(4)
1
1
4-9(7)
1
2-5(5)
2,3(2)
1
3-lO(7)
la-SC
lb-S,
lc-s,
Id-S,
le-S,
If-S,
2-6(6)
2-6(6)
l-6(4)
l-5(4)
l-4(3)
l-3(3)
2-4(3)
1
l-3(2)
l-5(4)
W3)
1,2(l)
5-l:(8)
l-3(3)
2,3(2)
1
1
l-3(2)
1
3-5(4)
1
2-4(4)
1
1
2,3(2)
1
16-42(30)
1
L
1
a Based on specimens from Zimbabwe (type locality) and South Africa.
b Range (mode) obtained from counts made on at least10 setae.
CSeta la-S usually borne belowlastpecten spinewhen present.
X
2-4(3)
-
Harbach: Subgenus Culex in SouthwesternAsia
Table
Seta
28.
Number of branches for fourth-instar larval setae of Culexperexiguus
Theobald.a
Head
Thorax
Abdominal Segments
No.
C
P
M
0
1
1
lo-23(19)b
1
1
1
2
1
1
3,4(3)
2
1
1
4-8(5)
1
1,2(l)
3-5(4)
1,2(l)
l-3(2)
1
1
1
3-6(6)
3-5(3)
1,a1 1
l-4(2)
1
21-42(29)
15-28( 21)
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
1
1
2,3(3)
2,3(2)
4-8( 6)
3-5(4)
59(5)
2-4(3)
l-3( 2)
2-5(3)
2-4(2)
2
3-7( 6)
1
IV
0
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
T
I
2,3(2)
2-4( 2)
W(2)
3-5(4)
4-6(4)
1
1
5-8(8)
6-l 8(9)
4-6(5)
1
2-4( 2)
l-3(1)
5-8(6)
2-l(2)
6-l 4(8)
4-9( 5)
2,3(3)
1,W)
2-4( 3)
1
3-6( 4)
2
2,3(2)
II
II
1
1,2(2)
1
1,2(l)
4-8(5)
3,4(3)
1
3,4(3)
1
1,2(2)
l-3(2)
2,3(3)
2
5-8( 6)
1,2(2)
1
1,2(2)
1,2(2)
l-3(2)
3-5(3)
1
3-l5)
1,2(l)
1
1
2,3(2)
2,3(3)
l l-26( 15)
Abdominal Segments
Seta
No.
213
2-i(3)
1
l-3(2)
1
2,3(2)
2,3(2)
4-8( 7)
l-3(2)
1
1,2(l)
1,2(l)
1,2(l)
3-5( 4)
1
V
VI
VII
1
2-4(3)
1
1,2(l)
4-6( 6)
2,3(2)
1
3,4(3)
1
1,2(2)
24(3)
2,3(2)
1
3-5(4)
1
24(3)
1
2-4(3)
g-18(11)
1
5-l&5)
2
1
1
2
1
3-5(4)
1
1,2(l)
1,2(2)
233)
1
l-3(2)
1
26-44( 29)
1
4-6(4)
l-3( 2)
1
2,3(2)
1
2-5(3)
1
a Based on specimens from Egypt.
b Range (mode) obtained from counts made on at least 10 setae.
c Siphon sometimes with only 4 setae on one side.
VIII
5-A(6)
5-l(8)
1
3-5(4)
1a-SC
1b-S,
1c-s,
1d-S,
1e-S,
1
X
2,3(2)
2
1
4-9(7)
3
2,3(3)
2,3(2)
2,3(2)
1,W)
Contrib.Amer. Ent. Inst.,vol.24, no. 1, 1988
214
Table
Seta
29. Number of branches for fourth-instar larval setae of Culex fheileri Theobalda
Thorax
Head
Abdominal Segments
M
T
I
No.
C
P
0
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
1
1
13-25(16)b
-
-
-
1
1
1
2
1
1
l-4(1)
3-9(3)
1
2-4(2)
l-3(2)
4-6(4)
3-8(5)
3-6(4)
1
5-:(6)
2-i(3)
lo-18(14)
3-1115)
3
2
1
l-3(2)
3,4(3)
2,3(2)
6-10(7)
3-7(5)
5-9(6)
3,4(3)
2,3(2)
1
3,4(3)
1,2(2)
3-6(4)
l-3(3)
1,2(l)
5-9(5)
1
1,2(2)
5-9(6)
1
0
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
4-6(5)
4-6(5)
1
3-5(3)
1
30-48(32)
21-39(34)
1
2-6(3)
1
l-3(2)
4-7(6)
3-8(4)
3,4(3)
4-9(7)
12-19(15)
4-6(5)
1
-
1 ,W)
l-5(3)
2-6(3)
1,2(2)
6-10(7)
3-l(5)
l-3(3)
2-5(2)
1
1
2-7(3)
2,3(2)
21-35(29)
-
II
1
4,5(4)
1
1,2(2)
l-4(3)
3-6(5)
9-230(9)
2,3(2)
1
1,2(l)
2,3(2)
l-3(2)
3-5(5)
1
Abdominal Segments
Seta
No.
2,3(2)
1
1
1
II
IV
V
VI
VII
VIII
1
3-5(4)
1
3-8(5)
1
4-6(4)
14-22(18)
1
5-9(5)
2-7(4)
1
1
5-9(7)
1
7-10(8)
1,2(l)
6-l-L(8)
4,5(4)
-
1
3-5(4)
1
2-4(2)
1,2(l)
2-5(3)
1
1
3,4(3)
1
13(l)
3-9(7)
l-5(3)
8-l:(8)
2,3(3)
7-14(9)
2,3(2)
1
2,3(2)
1
1,2(l)
2-4(2)
1,2(l)
2-4(3)
1
3-5(4)
1
2-4(3)
1
1
2-5(4)
3-5(4)
3
1
3-6(3)
1
1
1,2(l)
3-5(4)
1
3,4(3)
1
38-52(46)
1
24(3)
1
2-6(3)
1
X
2-4(3)
3-5(4)
-
1
a Based on specimens from Egypt, Israel, and the Yemen Arab Republic.
b Range (mode) obtained from counts made on at least 10 setae.
c Siphon usually with 10 seta 1-S, not distinctlypaired; sometimes with as few as 7 or as
many as 13.
Harbach: Subgenus CulexinSouthwestem Asia
Table
30
Number of branches for fourth-instarlarvalsetae
215
of Culexlaticinctus
Edwards.a
Seta
No.
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
C
M
P
1
4,5(4)
2-5(3)
1,W)
2-4(3)
2,3(2)
2,3(2)
3-5(4)
3,4(3)
2
1
1,2(l)
2-6(4)
1
_
1
I
T
l-3(2)
l-4(3)
l-5(3)
1
0,l
1
1
4-6(5)
3,4(4)
7-ll(8)
3-6(5)
AbdominalSegments
Thorax
Head
2,3(2)
3-5(4)
3-6(5)
1
1
1,2(2)
1,2(l)
1
1
4-7(5)
4-6(5)
1
3-5(3)
1
20-30(24)
16-21(20)
II
I11
l-3(2)
1
2-7(3)
12-15(12)
4-6(4)
1,2(l)
1,2(l)
1
l-3(2)
3-8(5)
3-8(4)
1
l-3(2)
1,2(l)
3-6(5)
3,4(3)
2
3,4(3)
2-5(4)
1,2(l)
5-lO(7)
1,2(l)
1,2(l)
1
1
l-3(2)
1
3-6(4)
1
68(6)
8-18(13)
48(5)
1
2-4(2)
24(4)
2,3(2)
7-ll(10)
5-:(5)
l-3(1)
1
1
1
2,3(2)
2,3(3)
13-29(22)
-
AbdominalSegments
Seta
No.
IV
V
VI
0
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
1
1
1
1
1
1
1
4-6(4)
1,2(2)
1
1,2(l)
1,2(l)
S-10(9)
1,2(l)
1
1
l-3(2)
1
3-6(4)
1
1,2(l)
1
58(7)
1,W)
1
1
l-3(2)
1
2-6(4)
1
VII
VIII
1
1
1,2(l)
1
1
2,3(2)
1
1
6-8(7)
1,2(2)
1,2(l)
1
1
l-3(2)
1
1
l-3(2)
1
25-39(31)
1
24(2)
1
1,2(2)
lo-15(10)
1
8-12l(lO)
1
4-6(6)
4-6(4)
l-4(2)
1
l-3(1)
1
la-S,c
lb-S,
lc-s,
Id-S,
le-S,
If-S,
3-6(4)
1
lg-S,
1
a Based on specimens from lsraelandthe Yemen Arab Republic.
b Range (mode)obtainedfrom counts made on at least10 setae.
CSeta 1-S usually with 14(12-15)imperfectly
paired setae.
X
1,2(2)
4,5(4)
1
6-13(10)
6-ll(10)
7-ll(10)
7-ll(10)
7-13(10)
5-12(7)
24(3)
l-4(3)
]
Contrib.Amer. Ent. Inst.,vol.24, no. 1, 1988
216
Table
Seta
No.
0
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
31.
Number of branches for fourth-instarlarvalsetae
Head
C
1
1
1
1
1
3-5(4)
3,4(3)
7-10(8)
2-4(3)
4-7(4)
2-5(3)
2-4(3)
2-4(3)
M
7-12(9)b
1
1
1,2(2)
2
1
1
2-4(2)
2-4(2)
1
1
2-4(3)
1
2,3(2)
1
2-5(4)
I
T
1,2(2)
1
1
1
6,7(7)
5-8(6)
1
2-4(3)
1
12-22(17)
16-27(21)
1
1,2(2)
2,3(3)
6-10(7)
3-5(3)
2-5(2)
2,3(2)
1,2(2)
3-6(5)
1,2(l)
1
1,2(1)
1,2(l)
1
2-7(3)
1,2(l)
1,2(l)
12(l)
1,2(l)
2-5(3)
5-9(7)
1
1
1
2-6( 5)
1
2-4(3)
3-6(4)
3-5(3)
1
1
5-9(8)
9-14(12)
6-10(8)
1
Ill
1
l-3(2)
1
1,2(l)
1
2-4(4)
1
II
2,3(3)
1
4-7(5)
2-4(3)
3-5(4)
1
1
l-4(2)
2,3(2)
9-14(13)
1
1,2(l)
1
2-6(4)
1
_
Abdominal Segments
Seta
No.
AbdominalSegments
Thorax
P
of Cu/exmatting/yiKnight.a
IV
V
VI
VII
VIII
X
0
1
1
1
1
1
1
2
3
4
1,2(l)
1
1,2(l)
ld(2)
1,2(l)
1
1
3-5(4)
1,2(l)
1
1
2,3(2)
2
1
1,2(2)
1
58(6)
1
12-14(12)
1
2,3(3)
2,3(2)
1
7-12(9)
l-4(2)
2,3(2)
68(7)
1
1
1
l-3(2)
1
3-5(4)
1
l-3(2)
2,3(2)
l-3(2)
6-9( 8)
3,4(4)
-
5
6
7
8
9
10
11
12
13
14
15
2-5(2)
5-8(7)
1
1
1
l-3(2)
1,2(l)
4,5(5)
1
12(l)
l-4(2)
1
1
l-4(2)
1
16-29(21)
1
1,2(l)
2-4(3)
l-4(3)
1
l-3(1)
1
2-5(2)
1
la-S,c
lb-S,
lc-s,
Id-S,
le-S,
If-S,
1g-s
aBasedonspecimensfromtheYemen
Arab Republic.
h Range (mode) obtained from counts made on at least10 setae.
CSiphon normally with 14 seta 1-S (7 pairs), but ll-16 may be present.
1
5-10(8)
7-13(9)
7-12(10)
7-ll(l0)
7-12(10)
2-5(3)
3-7(6)
Harbach: Subgenus CulexinSouthwestem Asia
Table
217
32. Number of branches for fourth-instar larvalsetae of Culex simpsoni
Theobald.a
Seta
No.
0
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
C
P
M
1
1
13-19(16)b
1
1,2(l)
1
1
1,2(l)
1,2(2)
7-lO(9)
3-5(4)
4-8(6)
2,3(3)
2
4-7(5)
3,4(3)
2-4(2)
4-6(5)
3,4(4)
1
1
3,4(3)
2,3(2)
1
1
3-5(4)
1
1,W)
0
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
T
I
1,2(l)
l-6(3)
l-3(2)
1,2(l)
2,3(2)
9-13(10)
2-4(3)
1
2
1
1
1
4-6(6)
3-5(5)
1
l-3(2)
1
23-38(30)
ll-24(20)
3,4(3)
2-5(3)
1
1
2-6(4)
3
68(6)
8-19(11)
1,2(2)
48(5)
1
l-3(2)
2,3(3)
1
3-6(4)
1,2(l)
5-7(5)
1,2(l)
2
II
1
1
1
2
3-8(4)
l-3(2)
3
4-6(5)
1,2(l)
1
1
2-4(3)
2
15-24(19)
Ill
1
4,5(4)
1
2
l-3(2)
l-4(1)
3,4(3)
8-13(8)
2,3(2)
1
1,2(l)
2
2
3-5(4)
1
Abdominal Segments
Seta
No.
Abdominal Segments
Thorax
Head
IV
V
VI
VII
VIII
1
4-7(5)
5-i(5)
1
25(3)
1
1
s-10(7)
1
l-3(2)
14-25(19)
1
3,4(4)
1
1
1
4,5(4)
1
2
1,2(l)
4,5(5)
1
3-:(5)
3,4(4)
1,2(l)
1
l-3(2)
1,2(l)
3-5(3)
6-lO(9)
2
1
l-3(1)
24(3)
7-lO(8)
2
1
1
l-3(2)
3,4(3)
1
3
1
1
1,2(l)
W(2)
12u 1
3-6(4)
1
2,3(2)
1
3-5(4)
1
2,3(2)
1
29-45(33)
1
38(5)
l-3(1)
1
l-3(2)
1
3-6(4)
1
a Based on specimens from Ethiopia, Kenya, and Tanzania.
b Range (mode)obtained from counts made on at least 10 setae.
la-S,
lbS,
lc-s,
Id-S,
1
X
2,3(3)
24(2)
1
4-ll(10)
-
2,3(3)
243
2,3(2)
12(2)
218
Table
Seta
Contrib.Amer. Ent. Inst.,vol. 24, no. I, 1988
33.
Number of branches for fourth-instar larval setae of Culex sinaiticus
Kirkpatrick.a
No.
C
0
1
1
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
Thorax
Head
1
1
1
P
13-22(18)b
1
1
1
1,W)
l-4(3)
1
1
6-8(7)
3-7(4)
3-6(3)
2-4(2)
1,2(2)
1
1
1,V)
3-6(4)
3,4(4)
2-4(2)
4-6(5)
3
2-6(4)
1
1,2(2)
IV
T
I
II
-
1
2-4(3)
1
1
1
1
3-5(3)
1
W(2)
1,2(2)
8-ll(l0)
3-6(5)
l-3(2)
3
4-6(4)
1
1
1
X2(2)
2,3(2)
l-3(2)
1,2(l)
2
3-5(3)
3-5(4)
2-5(3)
1
l-3(2)
1
1
1
4-6(5)
1
5-i(5)
7-12(8)
4-6( 5)
3,4(4)
1
l-3(3)
1
19-28(23)
IO-23(15)
1,2(l)
2-6(3)
2,3(3)
1,2(2)
2-4(2)
1
l-3(2)
2-6(3)
1,2(l)
5-8(5)
2,3(2)
1,W)
2,3(2)
2,3(2)
14-22(17)
ill
6-l:(9)
2
1
1,2(l)
2
2
3,4(3)
1
_
V
VI
VII
1
2-5(4)
1
4-7(5)
1
1,2(l)
3,4(3)
1
2,3(2)
12-20(17)
1
4-7(4)
0
1
1
1
4,5(4)
1
2
1,2(l)
1,2(2)
4,5(5)
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
M
Abdominal Segments
Seta
No.
Abdominal Segments
7-l:(9)
2,3(2)
1
1,2(l)
2,3(2)
1,2(2)
3,4(3)
1
1,2(l)
1
4-6(5)
l-3(2)
3
6-9(7)
2,3(2)
1
1
2
2,3(2)
l-4(2)
3
1,2(l)
3,4(4)
1
3-i(3)
1
1
2,3(2)
1
22-43(35)
1
VIII
2,3(3)
1
2,3(3)
7,8(8)
1
3,4(3)
1
4-9(6)
la-S,
1,2(l)
1
2,3(2)
1
4,5(4)
1
a Based on specimens from Israel, Egypt (Sinai), and Iran.
b Range (mode) obtained from counts made on at least 10 setae.
X
1
-
Harbach: Subgenus Culex in SouthwesternAsia
Table
Seta
34.
Number of branches for fourth-instar larval setae of Culex duttoni
Theobald.a
Thorax
Head
Abdominal Segments
No.
C
P
0
1
1
ia-25(20)b
-
-
1
1
1
l-4(3)
141)
1
l-3(2)
1
4-6(6)
2-7(4)
1
2
3
4
5
6
7
a
9
10
11
12
13
14
15
1
1
M
T
I
l-5( 2)
1
1
lo-13(11)
5-l 4(7)
2,3(2)
2
6-a(7)
7-9(a)
7-ll(9)
1,w 1
1
1
1
1
1,W)
l-6(4)
3-6(4)
1
1
2-7(6)
1
1
5-10(5)
4-7(5)
1
2-4(2)
1
la-37(2a)
21-37(27)
5-:1(a)
16-24(20)
5-9(6)
1
l-3(2)
1
3-6(5)
2,3(2)
2,3(2)
3-5(3)
2-5(3)
1,2(l)
2
1
2,3(2)
2-5(3)
l-5(3)
2-:5(g)
1,2(l)
l-4(2)
VI
VII
1
1
1
1
2,3(2)
2-7(3)
1
1
1
1
1
l-7(5)
26(2)
1
1
1
1
2-5(2)
3-9(5)
a
9
10
a-16(9)
1
1
1
11
12
13
14
15
2-4( 2)
1,2(l)
2
1
_
7-:2(g)
1,2(l)
1,2(l)
1
24(3)
1
1,2(2)
1
1
1
1
1,2(l)
1
2-6(4)
12-27(20)
1
7-15(10)
2-5(3)
1
l-4(3)
1
2-5(4)
1
2
3
4
5
6
7
1
1
1
1
1
2,3(2)
2-5(4)
1
6,7(6)
1
1
1
l-3(2)
1
2,3(2)
1
1m
1
1
5-l O(7)
l-4(4)
2-4(2)
3,4(3)
1
1
1
2,3(2)
1
15-34( 25)
Abdominal Segments
V
1
II
-
IV
0
II
3-5(3)
Seta
No.
219
1
3-7(4)
1
1
2-5(3)
1
21-52(37)
1
VIII
1
4-9(6)
1
7-l l(9)
1
3-6(5)
1a-SC
lb-S,
lc-s,
Id-S,
1e-S,
X
1
1
1
4-7(5)
1
1
1
1
1a1 >
14(3)
I
1
a Based on unassociated specimens from P.D.R. Yemen, Ethiopia, Cameroon, Ghana,
and Uganda.
b Range (mode) obtained from counts made on 12 setae (in most cases).
c Five pairs of seta 1-S were observed in specimens from P.D.R. Yemen. Only four pairs
were present in specimens from other areas.
Contrib.Amer. Ent. Inst., vol. 24, no. 1, 1988
220
Table
Seta
35.
Number of branches for fourth-instar larval setae of Culex sitiens
Wiedemann.a
Abdominal Segments
Thorax
Head
No.
C
P
M
T
I
0
1
1
1
9-16(12)b
1
1
1
2
1
1
2,3(3)
1,2(2)
1
1
4-8(6)
1
1
2-4(3)
1
1,2(2)
1,2(l)
1
1
4-6(6)
4-6(5)
1
2-4(3)
1
13-21(16)
13-25(18)
1
1,2(l)
24(3)
3-6(4)
1
1
6-9(7)
8-15(14)
4-7(5)
1
2-5(3)
1,2(l)
2-5(3)
l-3( 1)
1
1,2(l)
5-l 4(9)
2-7(4)
3,4(3)
1,2(l)
2
3
4
5
6
7
8
9
10
11
12
13
14
15
1
1
6-8( 7)
4-7(4)
7-l l(9)
3-5(4)
3,4(3)
2-4( 2)
2-4( 2)
2,3(3)
l-3(2)
1
2,3(3)
01
2
3
4
5
6
7
8
9
10
11
12
13
14
15
II
1
1
1
1
1
2,3(2)
1
1
1,2(2)
2-4(3)
2-6(4)
2-4(3)
2-4(3)
4-6(5)
1
1
1
1,2(2)
1,2(2)
lo-21(12)
5-i(6)
l-3(1)
1
1
2
1,2(2)
2
1
Abdominal Segments
Seta
No.
1,2(2)
1
l-3( 2)
1,2(l)
l-3( 2)
u
IV
V
VI
VII
VIII
X
2,3(2)
1
1
1
2,3(2)
1
1
1
2-6(4)
2,3(2)
2,3(2)
1
1
1
l-3( 2)
24(2)
1,2(2)
1,2(l)
2-w
1
1
2,3(2)
1
14-36( 28)
1
2,3(3)
1
1
l-4(2)
5:(6)
1
8-l l(9)
1
4
1
3-5(4)
1,2(l)
S-12(10)
Gw)
U(2)
5-i(6)
2
1
1
2,3(2)
l-3( 2)
l-3(2)
1
5:(6)
2
1
1
2
1
24(2)
1
1,2(l)
2-5(3)
6-13(11)
1
3-6(4)
2,3(2)
1
l-3( 2)
1
2,3(2)
1
1a-S,C
1b-S,
1c-s,
1d-S,
1e-S,
1f-S,
1g-s
1
a Based on specimens from the Yemen Arab Republic, United Arab Emirates, and
Oman.
b Range (mode) obtained from counts made on at least 10 setae.
c Seta la-S may be absent on one or both sides of the siphon.
Harbach: Subgenus Culex in SouthwesternAsia
Table
Seta
36. Number of branches for fourth-instar larval setae of Culex poicilipes
(Theobald).a
Thorax
Head
Abdominal Segments
No.
C
P
M
T
0
1
1
1
15-30(22)b
1
1
1
2
1
1
5-8(5)
4-lO(5)
3-6(5)
3-5(4)
6-11(g)
5-lO(6)
2-4(3)
1,2(2)
2
3
4
5
6
7
8
9
10
11
12
13
14
15
1
l-3(2)
35(4)
4-6(4)
lo-15(10)
3-7(5)
6-12(8)
3-5(4)
2,3(2)
3-7(5)
2
1,2(l)
5,6(5)
3,4(3)
2
l-3(2)
1,2(l)
4-8(8)
1
1
I
7-lO(9)
3-5(3)
2-7(5)
ll-19(15)
5-lO(7)
3
5-8(6)
8-17(12)
4-7(5)
1
2-4(2)
2,3(2)
12-23(21)
-
1,2(2)
3-6(5)
l-3(2)
S-10(7)
2-4(3)
2-5(4)
II
Ill
1
5-9(6)
2-4(3)
2-4(3)
7-lO(8)
5-16(12)
6-8(7)
l-3(2)
3-5(3)
3-5(4)
7-13(11)
1
3,4(3)
8-18(11)
3,4(3)
9-27(17)
1,2(l)
1,2(l)
2,3(2)
1
5-A(6)
2-4(4)
18-40(29)
2,3(2)
4-6(4)
2-5(3)
5-9(6)
1
Abdominal Segments
IV
0
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
1,2(2)
4-6(4)
1
1
1
4-8(6)
4-6(4)
1
2-4(4)
1
24-45(32)
31-43(37)
Seta
No.
221
6-:0(g)
l-3(2)
3,4(4)
2-5(4)
6-lO(8)
3,4(3)
lo-16(13 )
3,4(3)
1,2(l)
1,2(2)
3-6(5)
2-4(3)
5-7(6)
1
V
VI
1
7-9(8)
13(l)
1
2,3(2)
6-12(g)
4-8(5)
3
lo-16(10
3-5(3)
1
1,2(l)
3-7(4)
l-3(1)
5-7(6)
1
6-9(7)
l-3(2)
2,3(2)
2-6(4)
3-7(5)
3,4(3)
2-6(3)
3-7(4)
ZW)
3-w
1
43-57(50)
1
VII
5-iO(7)
1,2(l)
7-lO(8)
l-3(3)
2-8(5)
13-27(19)
3,4(3)
5-12(7)
2-5(3)
4-6(5)
2-6(4)
1
6-lO(6)
1
a Based on specimens from Egypt.
b Range (mode) obtained from counts made on at least 10 setae.
VIII
X
5&
2,3(2)
7-l l(8)
2-5(4)
3-7(5)
1
5-7(4)
1
7-14(11)
-
Contrib.Amer. Ent. Inst.,vol. 24, no. 1, 1988
222
Table
Seta
37. Number of branches for fourth-instar larval setae of Culex mimeticus Noka
Head
No.
C
0
1
2
3
4
5
6
7
8
9
1
1
10
11
12
13
14
15
1
2-4(3)
3,4(4)
2,3(2)
5-8(6)
2-4(3)
3-6(4)
2-4(3)
2-4(3)
2,3(3)
2,3(2)
1
Thorax
P
8-16(11)b
1
1
1
0
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
1
2-7(4)
1
1
1
1
2,3(3)
2,3(2)
1
1
1
2,3(2)
1
4-:0(5)
5,6(5)
3-5(5)
1
1
T
I
II
Ill
1,2(l)
1,2(l)
1
1
1
1
1
1
1
2-4(3)
l-3(1)
1,2(l)
1
3-5(3)
3-6(4)
13(l)
1,2(l)
6-12(11)
2-6(3)
3,4(3)
1,2(l)
5:(8)
g-16(11)
5,6(5)
1
3,4(3)
1
20-34(25)
15-28(19)
1
2,3(2)
1
2-4(2)
l-3(2)
2-4(2)
2-4(3)
1,2(l)
l-3(2)
1,2(l)
3-7(4)
l-4(2)
3,4(3)
2-5(4)
1
1,2(l)
1
2,3(2)
1,2(2)
13-23(15)
5,:(6)
1
1
1
2
1,2(2)
1,2(2)
1
2,3(3)
Abdominal Segments
Seta
No.
Abdominal Segments
M
IV
V
1
VI
VII
1
1
1
1,2(2)
1
1,2(l)
2,3(2)
2,3(2)
1
1
4-6(4)
2,3(2)
1
1,2(l)
2,3(2)
3,4(4)
1
2-4(2)
1
l-3(1)
3,4(3)
5-9(6)
13(l)
2,3(3)
1,2(l)
2,3(2)
1
2-5(2)
lo-17(14)
1
2-5(3)
1,2(l)
1
1
2,3(2)
1,2(2)
1,2(l)
1
5-7(6)
2
1
1
l-3(2)
1
1,2(l)
1
1,2(2)
1
1
2,3(2)
1
25-41(34)
1
243)
1
2,3(2)
1
l-3(1)
1
VIII
X
5:(6)
l-3(2)
1,2(l)
6-8(7)
1
2,3(2)
1
3-6(4)
3,4(4)
-
1a-S,C
Its,
lc-s,
Id-S,
b-s,
If-S,
1
a Based on specimens from Israel and Iran.
b Range (mode) obtained from counts made on at least 10 setae.
c Siphon most often with 12 seta 1-S (6 pairs), but lo-14 may be present.
3-6(4)
3-7(4)
3-7(4)
3-6(4)
2-4(2)
l-3(2)
Harbach: Subgenus Culex in SouthwesternAsia
Table
Seta
No.
0
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
38. Number of branches for fourth-instar larval setae of Culex bitaeniorhynchus Giles.a
Thorax
Head
C
1
1
1
l-4(2)
2,3(3)
4-;(6)
2-4( 2)
4,5(4)
2,3(2)
1,2(l)
2-5(3)
l-3(1)
1
2,3(3)
P
0
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
Abdominal Segments
M
10-20(14)b
1
1
1
1,2(2)
1
1
3
2,3(2)
l-3( 2)
l-3(1)
3-7(5)
1
Seta
No.
223
T
I
1
l-3(2)
3,4(3)
26(5)
1,2(l)
1
6-9(7)
8-23( 16)
5-7(5)
1
2-6(4)
1,2(l)
3-6(4)
1-W
-
1,2(l)
3-7(4)
1
2,3(2)
1
1
1
4-6(5)
5-8(6)
1
2-4(2)
1
23-45(35)
16-34( 20)
W(l)
l-3( 2)
8-l l(8)
4-8(8)
3,4(3)
1
2,3(3)
1,W)
2-4(3)
1,2(2)
2,3(2)
#
Ill
1
1,2(l)
1
2,3(3)
3-8(8)
2-4(3)
2,3(3)
3-7(5)
1
1
1,2(l)
2-4(2)
2
13-18(15)
1
2,3(2)
WV
2,3(2)
l-3( 2)
2-4(3)
3,4(3)
5-lO(6)
1,2(l)
1
2
2,3(2)
2
2,3(2)
1
Abdominal Segments
IV
V
VI
VII
1
2,3(2)
1
2-4(3)
1,2(2)
2,3(2)
2-5(3)
4-l O(6)
2
1
1,2(2)
1,2(2)
2
2,3(2)
1
1
2,3(3)
1
l-3(2)
4-7(6)
2,3(2)
2-5(3)
4-9(7)
2
1
1,2(2)
2
1
2,3(2)
1
1
l-3( 2)
13(l)
2-5(3)
2-5(3)
2,3(2)
2,3(2)
2,3(2)
l-3(3)
1
1,W)
2,3(2)
1,w
23-35(25)
1
1
2,3(2)
1
2-4(2)
2,3(2)
243)
lo-19(11)
l-3( 2)
2-5(5)
l-3(1)
2-6(3)
1,2(l)
W(2)
242)
1
VIII
5-i(5)
2,3(2)
5-7(6)
2
3-5(3)
1
a Based on specimens from Iran, Pakistan, and the Yemen Arab Republic.
b Range (mode) obtained from counts made on at least 10 setae.
X
24(3)
2-5(3)
1
4-9(6)
-
Contrib.Amer. Ent. Inst.,vol. 24, no. 1, 1988
224
Table
Seta
39. Number of branches for fourth-instar larval setae of Culex tritaeniorhynchus Giles.a
No.
C
0
1
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
Thorax
Head
1
1
3,4(3)
2,3(2)
5-l O(8)
3-8(6)
3-7(3)
2-4(3)
2
2-4(3)
2-4(3)
1
3-5(4)
P
17-23(19)h
1
1
1
1,2(2)
1
1
2,3(3)
2
1
5-i(6)
1
1,2(l)
0
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
I
T
M
-
1a1 1
1,2(l)
4-8(5)
2,3(2)
1
58(5)
3-6(4)
2,3(3)
1
1
1
1
5-:(6)
3-5(5)
g-18(11)
3-5(4)
4,5(5)
1
1
2-5(3)
143)
1
l-3(2)
20-35( 28) 7-l l(9)
21-30( 25)
2-4(3)
1,2(l)
2-4(3)
ll-16(13)
4-8(5)
2,3(3)
l-3(2)
2-4(2)
3,4(3)
l-3( 2)
2,3(2)
II
Ill
1
1,2(2)
1
2-4(3)
6-l O(8)
2-4(2)
2,3(3)
5-8(7)
1
1
1
2-4(3)
2,3(2)
l l-23( 17)
1
2-4(3)
1
2,3(2)
l-3( 2)
2,3(2)
2,3(2)
6-l 2(9)
1,2(l)
1
1
1,2(2)
2
2-5(5)
1
Abdominal Segments
Seta
No.
Abdominal Segments
IV
V
VI
VII
1
4
1
2,3(3)
l-3( 2)
2,3(2)
2,3(2)
6-l l(9)
2,3(2)
1
1
2,3(2)
1,2(2)
4-6( 4)
1
1
4
1
1,2(l)
5-9(6)
2,3(3)
2,3(2)
S-10(10)
2,3(2)
1
1
1
3-6(4)
1
1
2-4(3)
24(3)
2
1
4-7(5)
1
24(2)
1
2-5(5)
1
24(3)
3-5(4)
1
1
2,3(2)
1
21-48(38)
1
VIII
4-6(5)
1,2(2)
4-:(5)
1,2(l)
7,8(7)
1,2(l)
3-w
W(5)
1o-1 7( 17)
2,3(2)
4-l 3(7)
2-4(3)
1,2(2)
24(3)
1
2-6(5)
1
a Based on specimens from Iran, Pakistan, and Saudi Arabia.
b Range (mode) obtained from counts made on at least 10 setae.
c Siphon with 1O-13 seta 1-S; usually with 12 (6 pairs).
X
24(3)
2-4(3)
6-111(8)
-
Harbach: Subgenus Culexin SouthwesternAsia
Table
Seta
No.
0
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
40
Number of branches for fourth-instarlarvalsetae
Colless.a
Head
C
1
1
l-3(1)
2-4(3)
l-3(2)
8-12(9)
2-6(4)
5-lO(5)
3,4(3)
l-3(2)
2-5(2)
l-3(2)
1
2-5(3)
0
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
Abdominal Segments
M
11-24(15)b
1
1
1
l-3(2)
1
1
3
1,2(2)
1
1
4-6(4)
1
1
T
I
II
II
1
4-8(5)
1
2-4(2)
-
-
-
1
12(l)
12(l)
l-3(2)
3-5(4)
1
2,3(2)
4-7(5)
2-5(3)
1,2(l)
3-7(3)
8-14(11)
3-5(4)
3
1
1,2(2)
1
2-4(2)
4-lO(7)
2,3(3)
1
1
1
3-5(4)
3-5(4)
1
1,2(l)
1
4-6(5)
ll-20(14)
3-6(4)
1
2-W
1
3-5(4)
2,3(3)
l-3(2)
7-12(9)
3,4(4)
1
35-44(40)
14-32(15)
1m
2,3(2)
2,3(2)
3
4-9(6)
1
1
1
2,3(3)
l-3(2)
13-32(21)
2,3(2)
2,3(2)
3,4(3)
7-12(8)
1,2(l)
1
1
2,3(2)
2,3(2)
3-6(4)
1
Abdominal Segments
Seta
No.
of Culex pseudovishnui
Thorax
P
225
IV
1
4-7(6)
1
2,3(2)
l-3(2)
2-4(2)
2-4(3)
6-12(8)
2-4(2)
1
1
2,3(2)
2,3(2)
4,5(5)
1
V
VI
VII
1
1
4-9(7)
1
48(6)
1
1
1
6-9(8)
1
3-7(4)
1
2-5(3)
12-23(16)
l-3(2)
4-ll(5)
l-5(2)
1
1,2(l)
4-7(6)
2,3(2)
3,4(3)
7-13(8)
2,3(2)
1
1
2-4(2)
1
4-7(5)
1
24(3)
2-4(2)
2,3(3)
3,4(3)
3-5(3)
1
1
2-5(2)
1
29-55(42)
1
2-4(3)
1
5-7(5)
1
a Basedon specimens from Pakistan.
b Range (mode)obtainedfromcounts
made on at least 10 setae.
c Siphon with 13-15 seta 1-S;usuallywith 14(7 pairs).
VIII
X
1
4,5(5)
5-l(7)
5-:(5)
la-SC
lbS,
lc-s,
Id-S,
le-S,
If-S,
1
3,4(3)
2,3(3)
1
4-lO(8)
4,5(4)
3-5(4)
3-5(4)
24(3)
3-5(4)
3-5(3)
Contrib.Amer. Ent. Inst.,vol. 24, no. 1,1988
226
Table 41.
Comparison of potentially diagnostic characters observed in members of the Culex
univittatus species aggregate.
character
univiltatus
perex@uus
neavei
1) Ventral surface
of proboscis
pale in middle
pale except at base,
weakly pale on distal
0.25
inconspicuously
pale in middle
2) Postspiracular
area
tendency for scales
to cover less than
dorsal 0.5
tendency for scales
to cover more than
dorsal 0.5
tendency for scales
to occur in small patch
near spiracle
3) Forefemur
sometimes with
indistinctanterior
pale stripe
usually with indistinct
anterior pale stripe
no anterior pale stripe
4) Midfemur
with complete distinct
or indistinctanterior
pale stripe
with or without incomplete faint or distinct
anterior pale stripe
normally without
anterior pale stripe,
weakly indicated
when present
5) Hindtibia
with distinctanterior
and posterior pale
stripes on proximal 0.8,
separated ventrally by
complete dark stripe;
with distinctapical
pale spot
with distinctanterior
and posterior pale
stripes on proximal 0.8,
partly separated on
proximal 0.5 or less by
weak ventral dark
stripe; with distinct
apical pale spot
with rather indistinct
anterior and posterior
pale stripes ending
before base; with
rather indistinct
apical pale apot
6) Wing - Costa
with short line of pale
scales at base
with short line of pale
scales at base
without pale scales
at base
female usually with
line of scales
female occasionally
with few scales
female occasionally
with few scales
7) Abdomen Pale bands
ofterga
normal
normal
reduced or absent
8) Male genitalia
-Setag
nanOw
broad
broad
long
short
long
- Vein 2A
- Ventral
arm
Harbach: Subgenus Culex in SouthwesternAsia
APPENDIX
227
A
Species of Culicidae other than Culex (Culex) known to occur in
southwestern Asia and Egypt
(subspecies and varieties not recognized)
1.
2.
3.
4.
5.
6.
7.
8.
9.
10.
11.
12.
13.
14.
15.
16.
17.
18.
19.
20.
21.
22.
23.
24.
25.
26.
27.
28.
29.
30.
31.
32.
33.
34.
35.
36.
37.
38.
39.
40.
41.
42.
Anopheles (Anopheles) algeriensis Theobald, 1903a
Anopheles (Anopheles) claviger (Meigen, 1804)
Anopheles (Anopheles) coustani Laveran, 1900
Anopheles (Anopheles) habibi Mulligan and Puri, 1936
Anopheles (Anopheles) hyrcanus (Pallas, 1771)
Anopheles (Anopheles) maculipennis Meigen, 18 18
Anopheles (Anopheles) marten! Senevet and Prunnelle, 1927
Anopheles (Anopheles) melanoon Hackett, 1934
Anopheles (Anopheles) plumbeus Stephens, 1828
Anopheles (Anopheles) sacharovi Favre, 1903
Anopheles (Anopheles) tenebrosus Donitz, 1902
Anopheles (Cell/a) apoci Marsh, 1933
Anopheles (Ce//ia) arabiensis Patton, 1905
Anopheles (Celia) azaniae Bailly-Choumara, 1960
Anopheles (Cell/a) cinereus Theobald, 1901 a
Anopheles (Cellia) culicifaciesGiles, 1901 c
Anopheles (Cellia) demeilloni Evans, 1933
Anopheles (Ce//ia) dthali Patton, 1905
Anopheles (Cellia) fluviatilisJames, 1902
Anopheles (Celia) hispaniola (Theobald, 1903a)
Anopheles (Ce//ia) moghulensis Christophers, 1924
Anopheles (Ce//ia) multicolorCambouliu, 1902
Anopheles (Ce//ia) pharoensis Theobald, 1901 a
Anopheles (Cell/a) pretoriensis (Theobald, 1903a)
Anopheles (Cell/a) pulcherrimus Theobald, 1902
Anopheles (Ce//ia) rhodesiensis Theobald, 190 1a
Anopheles (Ce//ia) sergentii (Theobald, 1907)
Anopheles (Ce//ia) splendidus Koidzumi, 1920
Anopheles (Ce//ia) squamosus Theobald, 1901 a
Anopheles (Celia) stephensi Liston, 1901
Anopheles (Cell/a) subpictus Grassi, 1899
Anopheles (Cell/a) superpictus Grassi, 1899
Anopheles (Ce//ia) turkhudi Liston, 1901
Aedes (Aedimorphus) hirsutus (Theobald, 190la)
Aedes (Aedimorphus) natronius Edwards, 1932b
Aedes (Aedimorphus) vexans (Meigen, 1830)
Aedes (Aedimorphus) vittatus (Bigot, 1861)
Aedes (Finlaya) echinus (Edwards, 1920)
Aedes (Finlaya) geniculatus (Olivier, 1791)
Aedes (Ochlerotatus) bertandi Seguy, 1921
Aedes (Ochlerotatus) cabal/us (Theobald, 1912)
Aedes (Ochlerotatus) caspius (Pallas, 1771)
Contrib.Amer. Ent. Inst.,vol. 24, no. 1, 1988
228
43.
44.
45.
46.
47.
48.
49.
50.
51.
52.
53.
54.
55.
56.
57.
58.
59.
60.
61.
62.
63.
64.
65.
66.
67.
68.
69.
70.
71.
72.
73.
74.
75.
76.
77.
78.
Aedes
Aedes
Aedes
Aedes
Aedes
(Ochlerotatus) chelli (Edwards, 19 15)
(Ochlerotatus) detritus (Haliday, 1833)
(Ochlerotatus) duplex Martini, 1926
(Ochlerotatus) flawescens (Miiller, 1764)
(Ochlerotatus) lepidonotus Edwards, 1920
Aedes(Ochlerotatus) leucomelas (Meigen, 1804)
Aedes (Ochlerotatus) nigrocanus Martini, 1927
Aedes (Ochlerotatus) phoeniciae Coluzzi and Sabatini, 1968
Aedes (Ochlerotatus) pulchritarsis (Rondani, 1872)
Aedes (Ochlerotatus) rusticus (Rossi, 1790)
Aedes (Rusticoidus) refiki Medschid, 1928
Aedes (Stegomyia) aegypti (Linnaeus, 1762)
Aedes (Stegomyia) cretinus Edwards, 1921
Culex (Barraudius) modestus Ficalbi, 1889( 1890)
Culex(Barraudius) pusillus Macquart, 1850
Culex (Culiciomyia) nebulosus Theobald, 1901 c
Culex (Lasiosiphon) adairi Kirkpatrick, 1926
Culex (Lutzia) tigripes De Grandpre and De Charmoy, 1900( 1901)
Culex (Maillotia) arbieeni Salem, 1938
Culex (Maillotia) deserticola Kirkpatrick,1924( 1925)
Culex (Mailotia) hortensis Ficalbi, 1889a
Culex (Maillotia) quettensis Mattingly, 1955b
Culex (Mail0 tia) salisburiensisTheo bald, 190 1a
Culex (Neoculex) impudicus Ficalbi, 1890
Culex (Neoculex) judaicus Edwards, 1926
Culex (Neoculex) martinii Medschid, 1930
Culex (Neoculex) teffitans Walker, 1856
Culiseta (Allotheobaldia) longiareolata (Macquart, 1838a)
Culiseta (Culicella) morsitans (Theobald, 1901 a)
Culiseta (Culicella) setivalva (Monchadskii, 1936)
Culiseta (Culiseta) annulata (Schrank, 1776)
Culiseta (Culiseta) subochrea (Edwards, 1921 [in Wesenberg-Lund,
19211)
Coquillettidia (Coquillettidia) buxtoni (Edwards, 1923)
Coquillettidia(Coquillettidia)richiardii(Ficalbi, 1889b)
Orthopodomyia pulcripalpis (Rondani, 1872)
Uranotaenia (Pseudoficalbia) unguiculata Edwards, 1913a
Harbach: Subgenus Culex in SouthwesternAsia
APPENDIX
229
B
Complete synonymy of Culex (Culex) pipiens Linnaeus
pipiens Linnaeus, 1758: 602. Type locality: Sweden, Scania, Veberod, Silvakra farm.
+Neotype male (NMNH), designated by Harbach et al., 1985: 9.
bifurcatus Linnaeus, 1758: 603. Type locality: France. Lectotype male (non-extant,
illustrated by Reaumur, 1738: PI. 40, Fig. 2), designated by Harbach et al., 1985:
7. Synonymy with pipiens by Martini, 1922: 107.
fasciatus Muller, 1764: 87. Type locality: none designated. Location of type(s) (adult)
unknown. Synonymy with pipiens by Edwards, 1921: 345.
molestus Forskal, 1775: 85. Type locality: Egypt, Buhayrah Gov., Rosetta. +Neotype
male (NMNH), designated by Harbach et al., 1984: 523. Synonymy with pipiens
restored by Harbach et al., 1984: 540; prior synonymy with pipiens by Edwards,
1921: 345.
trifurcatus Fabricius, 1794: 401. Type locality: “Europae paludosis.” Type(s) (adult)
non-extant. Synonymy with pipiens by Edwards, 1932a: 210.
luteus
Meigen, 1804: 6. Type locality: Europe. Type(s) (female) non-extant.
Synonymy with pipiens by Edwards, 1932a: 310.
domesticus Germar, 1817: 290. Type locality: Yugoslavia, Dalmatia. Location of
type(s) (adult) unknown. Synonymy with molestus by Marshall and Staley, 1937:
23; prior synonymy with pipiens by Edwards, 1921: 345.
rufus Meigen, 1818: 7. Type locality: none designated. +Syntypes (male; female)
(MNHP), see Harbach, 1983: 103. Synonymy with pipiens by Edwards, 1921:
345.
bicolor Meigen, 1818: 9. Type locality: ? Africa. +Type female (MNHP), see Harbach
1983: 99. Synonymy with pipiens by Edwards, 1921: 345.
marginalis Stephens, 1825: 455. Type locality: England, London. Type(s) (female)
(BM). Synonymy with pipiens by Edwards, 192 1: 345.
meridionalis
Leach, 1825: 292. Type locality: France, Nice. Location of type(s)
(adult) unknown. Synonymy with pipiens by Edwards, 1921: 345.
consobrinus Robineau-Desvoidy, 1827: 408. Type locality: U.S.A., Pennsylvania.
Type(s) (adult) non-extant. Synonymy with pipiens by Edwards, 1932a: 209.
calcitrans Robineau-Desvoidy, 1827: 409. Type locality: France, Paris; Germany.
Type(s) (female) non-extant. Synonymy with pipiens by Edwards, 1921: 345.
230
Contrib.Amer. Ent. Inst.,vol. 24, no. 1,1988
thoracicus Robineau-Desvoidy, 1827: 409. Type locality: France, Paris. Type(s)
(female) non-extant. Synonymy with pipiens by Edwards, 1921: 345.
pallipes Waltl, 1835: 110. Type locality: Spain. Type(s) (female) (MNHP). Synonymy
with pipiens by Edwards, 1921: 345.
unistriatus
Curtis, 1837: 231. Type locality: England.
Synonymy with pipiens by Stone et al., 1959: 254.
pallipes Macquart, 1838a: 37. Type locality: Egypt.
Synonymy with pipiens by Edwards, 1921: 345.
Type(s) (adult) (NMM).
Type(s) (adult) non-extant.
rufinus Bigot, 1888: 7. Type locality: Tunisia, El-Djerid. Type(s) (female) (non-extant).
Synonymy with pipiens by Edwards, 1921: 345.
agilis Bigot 1889: 122. Type locality: Algeria. Type(s) (female) (BC). Synonymy with
pipiens by Edwards, 1921: 345.
phytophagus
Ficalbi, 1889 (1890): 126. Type locality: Italy, Sardinia, Romagna,
Toscana. Location of types (male; female) unknown. Synonymy with pipiens by
Edwards, 1921: 345.
haematophagus Ficalbi, 1893: 143. Type locality: Europa (e d’ltalia). Location of
type(s) (adult) unknown. Synonymy with molestus by Marshall and Staley, 1937:
23; prior synonymy with pipiens by ? Edwards, 1921: 345.
melanorhinus Giles, 1900: 342. Nomen novum for pallipes Macquart, 1838b: 37, non
Waltl, 1835: 110.
varioannulatus Theobald, 1903a: 198. Type locality: Azores, St. Michaels. Type(s)
(female) (BM). Synonymy with pipiens by Edwards, 1913b: 53.
azoriensis Theobald, 1903a: 210. Type locality: Azores, St. Michaels. Types (male;
female) (BM). Synonymy with pipiens by Edwards, 1913b: 53.
longefurcatus Becker, 1903: 68. Type locality: Egypt, Cairo. Type(s) (female) (ZM).
Synonymy with pipiens by Edwards, 1932a: 209.
quasimodestus
Theobald, 1905b: 88 (bicolor var. ?). Type locality: Tunisia, Sfax.
Type(s) (female) (HNM). Synonymy with molestus by Stone,1957: 172; prior
synonymy with pipiens by Edwards, 1921: 246.
doliorum Edwards, 1912a: 263 (pipiens var.).
Kingswear. Type(s) (male) (BM).
Type locality:
England, Devon,
autogenicus Roubaud, 1935: 444 (pipiens ssp.). Type locality: France. Location of
type(s) (egg) unknown (probably lost), see Harbach,l983: 106. Synonymy with
molestus by Marshall and Staley, 1937: 18.
berbericus Roubaud, 1935: 445 (pipiens ssp.). Type locality: North Africa [Algeria].
Hatbach: Subgenus Culex in SouthwesternAsia
231
Location of type(s) (egg) unknown (probably lost), see Harbach 1983: 106.
Synonymy with molestus by Knight and Abdel-Malek, 1951: 184.
sternopallidus
Roubaud, 1945: 54 (autogenicus ssp.). Type locality: France,
Aigues-Mortes and Bages (Pyrenees-Orientales).
Location of type(s) (adult)
unknown (probably lost), see Harbach, 1983: 106.
sternopunctatus
Roubaud, 1945: 54 (autogenicus ssp.). Type locality: France,
Arles and Paris. Location of type(s) (adult) unknown (probably lost), see Harbach,
1983: 106.
disjunctus Roubaud, 1957: 3116 (pipiens ssp.). Type locality: France, Normandy, Ifs.
Type(s) non-extant. Synonymy with pipiens by Stone et al., 1959: 254.
calloti Rioux and Pech, 1959: 117 (pipiens ssp.). NEW SYNONYMY.
France and Tunisia. Location of type(s) (adult) unknown.
Type locality:
erectus Iglisch, 1977: 269 (pipiens ssp.). NEW SYNONYMY. Type locality: West
Germany, Berlin-Zehlendorf, Krummer Fenn. No type(s) designated.
torrldus Iglisch, 1977: 269 (pipiens ssp.). NEW SYNONYMY. Type locality: West
Germany, Berlin-Zehlendorf, Krummer Fenn. No type(s) designated.
232
Contrib.Amer. Ent. Inst.,vol. 24, no. 1, 1988
APPENDIX
C
Complete synonymy of Culex (Culex) quinquefasciatus Say
quinquefasciatus
Say, 1823: 10. Type locality: New Orleans, Louisiana, USA.
+Neotype male (NMNH), designated by Sirivanakarn and White, 1978: 362.
pungens
Wiedemann, 1828: 9. Type locality: New Orleans, Louisiana, USA.
Lectotype female (NMW), designated by Belkin, 1968b: 19. Synonymy with
quinquefasciatus by Edwards, 1932a: 208.
fatigans Wiedemann, 1828: 10. Type locality: East Indies [Indonesia]. Lectotype
female (NMW), designated by Belkin, 1968a: 68. Synonymy with quinquefasciatus by Dyar and Knab, 1909a: 34.
aestuans Wiedemann, 1828: 11. Type locality: Salvador, Bahia, Brazil. Lectotype
male (NMW), designated by Stone, 1958: 186. Synonymy with cpinquefasciatus
by Stone, 1958: 186.
acer Walker, 1848: 8. Type locality: New Zealand. Holotype female (BM). Synonymy
with fatigans by Edwards, 1924: 395.
cingulatus Doleschall, 1856: 405. Type locality: Ambarawa, Java. Location of type(s)
(adult) unknown. Synonymy with fatigans by Edwards, 1932a: 208.
cubensis
Bigot, 1857: 329. Type locality: Cuba. Lectotype female (MNHP),
designated by Belkin, 1968b: 15. Synonymy with fatigans by Edwards, 1932a:
208.
anxifer Bigot, 1859: 117. Type locality: Madagascar. Location of types (male; female)
unknown. Synonymy with fatigans by Edwards, 1932a: 208.
serotinus Philippi, 1865: 595. Type locality: Santiago and Valdivia, Chile. Types
(male; female) non-extant. Synonymy with fatigans by Edwards, 1932a: 209.
autumnalis Weyenbergh, 1882: 23. Type locality: Primer0 River, Cordoba, Argentina.
Location of type(s) (male; female; pupa; larva) uncertain (? Universidad de
Cordoba). Synonymy with fatigans by Edwards, 1932a: 208.
penafleli Sanchez, 1885: 213. Type locality: Mexico City, Mexico. Location of types
(male; female) unknown. Synonymy with fatigans by Edwards, 1932a: 208.
macleayi Skuse, 1889: 1746. King George’s Sound, Western Australia, Australia.
Type(s) (female) (MM). Synonymy with fatigans by Edwards, 1924: 395.
skusii Giles, 1900: 292. Type locality: Australia. Location of type(s) (male; female)
unknown. Synonymy with fatigans by Edwards, 1924: 395.
Harbach: Subgenus Culex in SouthwesternAsia
233
doleschallii Giles, 1900: 338. A/omen novum for cingulatus Doleschall, 1856: 405.
Synonymy with fatigans by Edwards, 1932a: 208.
albolineatus Giles, 1901 a: 609. Type locality: Shahjahanpur, India. Lectotype female
(BM), designated by Sirivanakarn, 1976: 33. Synonymy with fatigans by Barraud,
1924b: 1264.
quaslpipiens Theobald, 1901 b: 136. Type locality: Sambalpur, Central Provinces,
India. Lectotype female (BM), designated by Bram, 1967b: 327. Synonymy with
fatigans by Edwards, 1913b: 55.
fouchowensis
Theobald, 1901 b: 137. Type locality: Foochow, China. Lectotype
male (BM), designated by Bram, 1967a: 196. Synonymy with fatigans by
Edwards, 1913b: 55.
reesi Theobald, 1901 b: 145. Type locality: Hong Kong. Type(s) non-extant (Bram,
1967a: 196). Synoymy with fatigans by Edwards, 1913b: 55.
sericeus Theobald, 1901 b: 147. Type locality: Hong Kong. Location of type(s)
(female) unknown. Synonymy with fatigans by Edwards, 1913b: 55.
luteoannulatus
Theobald, 1901b: 159 (fatigans ssp.). Type locality: none
designated. Type(s) (adult) (BM). Synonymy with fatigans by Edwards, 1932a:
208.
trilineatus Theobald, 1901 b: 159 (fatigans ssp.). Type locality: none designated.
Location of type(s) (adult) unknown. Synonymy with fatigans by Edwards, 1932a:
208.
pallidocephala
Theobald, 1904: 73. Type locality: Sennar, Blue Nile, Sudan.
Holotype female (BM). ? Synonymy with fatigans by Edwards, 1941: 316.
cartroni Ventrillon, 1905b: 429. Type locality: Majunga, Moroudara, Madagascar.
+Lectotype male (MNHP), designated herein (see treatment of quinquefasciatus
Say). Synonymy with fatigans by Edwards, 1932a: 208.
barbarus Dyar and Knab, 1906: 210. Type locality: Trinidad.
(NMNH). Synonymy with fatigans by Edwards, 1932a: 208.
Holotype female
zeltneri Neveu-Lemaire, 1906, 251. Type locality: Harar and Comboltcha, Ethiopia.
Type(s) (male; female) non-extant. Synonymy with quinquefasciatus by White,
1975: 322.
didieri
Neveu-Lemaire, 1906: 257. Type locality: Leopoldville, Belgian Congo.
Type(s) (female) non-extant. ? Synonymy with fatigans by Edwards, 1932a: 208.
pygmaeus Neveu-Lemaire, 1906: 256. Type locality: Imi, Ethiopia. Type(s) (male;
female) non-extant, ? Synonymy with fatigans by Edwards, 1932a: 208.
234
Contrib.Amer. Ent. Inst., vol. 24, no.l,1988
quasiiineaiis Theobald, 1907: 415. Type locality: Adelaide, Australia. Type (female)
(BM). Synonymy with fatigans by Edwards, 1932a: 208.
stoehri Theobald, 1907: 419. Type locality: British Central Africa. Type (female) (BM).
Synonymy with fatigans by Edwards, 1932a: 209.
christophersii
Theobald, 1907: 453. Type locality: India. Lectotype male (BM),
designated by Bram, 1967b: 327. Synonymy with fatigans by Edwards, 1913b:
55.
raymondii Tamayo, 1907 [in Tamayo and Garcia, 1907: 361. Type locality: Huacachina,
Peru. Location of type(s) (male; female; pupa; larva) unknown. Synonymy with
fatigans by Edwards, 1932a: 209.
aikenii Dyar and Knab, 1908: 61. Type locality: New Amsterdam, Surinam. Lectotype
male (NMNH), designated by Stone and Knight, 1957: 42. Synonymy with
fatigans by Edwards, 1932a: 208.
minor Theobald, 1908a: 298. Type locality: Sylhet and Lungleh, Lushai Hills, Assam
and Calcutta, Bengal, India. Types (male; female) (BM). Synonymy with fatigans
by Barraud, 1924b: 1264.
revocator Dyar and Knab, 1909b: 256. Type locality: Hope Gardens, Newcastle,
Jamaica. Lectotype male (NMNH), designated by Stone and Knight, 1957: 55.
Synonymy with guinquefasicatus by Dyar, 1918: 99.
iachrimans Dyar and Knab, 1909b: 259. Nomen novum for aikenii Dyar and Knab,
1908: 61, non Aiken and Rowland, 1906: 34.
goughii Theobald, 1911: 268. Type locality: Onderstepoort, Transvaal. Lectotype
male (BM), designated by White, 1975: 321. Synonymy with fatigans by
Edwards, 1913b: 55.
fuscus
Taylor, 1914a: 699 (Culicelsa ). Type locality: Townsville, Queensland,
Australia. Type(s) ? non-extant. Synonymy with fatigans by Edwards, 1924: 395.
aseyehae Dyar and Knab, 1915: 112. Type locality: New Providence Island, Bahama
Islands. Lectotype male (NMNH), designated by Stone and Knight, 1957: 43.
Synonymy with fatigans by Edwards, 1932a: 208.
townsviiiensis Taylor, 1919: 836. Nomen novum for fuscus Taylor, 1914a: 699, non
Theobald, 191 Oc: 319.
hensemaeon
Dyar, 1920: 178. Type locality: Los Banos, Philippines.
female (NMNH). Synonymy with fatigans by Edwards, 1922: 278.
Holotype
nigrirostris Enderlein, 1920: 51 (fatigans var.). Type locality: Tananarive, Madagascar.
Type(s) (female) (ZM).
Harbach: Subgenus Culex in SouthwesternAsia
APPENDIX
235
D
Complete synonymy of Culex (Culex) sitiens Wiedemann
sitiens Wiedemann, 1828: 542. Type locality: Sumatra, Indonesia. Holotype female
(ZMC).
impellens Walker, 1859: 91. Type locality: Makessar, Celebes. +Holotype female
(BM). Synonymy with sitiens by Edwards, 1913c: 232.
microannulatus Theobald, 1901 a: 353. Type locality: Quilon, Travancore, [Madras
State], India. +Lectotype male (BM), designated by Bram, 1967a: 243. Synonymy with sitiens by Edwards, 1913~: 232.
gnophodes
Theobald, 1903a: 163. Type locality: Bruas, Dindings, [Malaysia].
+Holotype female (BM). Synonymy with sitiens by Edwards, 1913c: 232.
somaliensls Neveu-Lemaire, 1906: 254. Type locality: Djibouti, Djibouti. Syntypes
(male; female) non-extant (?). Synonymy with Miens by Edwards, 1913~: 232.
nlgricephala Leicester, 1908: 149. Type locality: Batu Gajah, [Perak], Malaysia.
Syntypes (male; female) non-extant. Synonymy with Miens by Edwards, 1932a:
204.
salus Theobald, 1908b: 256. Type locality: Port Sudan, [Kassala], Sudan. +Lectotype
male (BM), designated herein (see treatment of sitiens Wiedemann). Synonymy
with somaliensis by Edwards, 1911: 261.
jepsoni Theobald, 1910b: 158. Type locality: Suva, [Viti Levu], Fiji. +Holotype female
(BM). Synonymy with Miens by Edwards, 1924: 394.
saibaii Taylor, 1912: 28. Type locality: Saibai Island, Torres Strait. Type(s) (female)
(US). Synonymy with Miens by Edwards, 1924: 394.
paludis Taylor, 1913: 56 (Culicelsa ). Type locality: Townsville, Queensland, Australia.
Type(s) (female) (US). Synonymy with Miens by Edwards, 1924: 394.
annulata Taylor, 1914a: 689 (Culicada ). Type locality: Townsville, Queensland,
Australia. Syntypes (male; female) (US). Synonymy with sitiens by Edwards,
1924: 394.
milni Taylor, 1914b: 196 (Culicelsa annulirostrisvar.). Type locality: Milne Bay, Papua,
New Guinea. Syntypes (male; female) (US). Synonymy with sitiens Edwards,
1924: 394.
salinus Baisas, 1938: 204. Type locality: Caloocan, Rizal, Luzon, Philippines. Type(s)
(male, female, larva, pupa) non-extant. Synonymy with sitiens by Delfinado,
1966: 152.
236
Contrib.Amer. Ent. inst.,vol. 24, r~. 1, 1988
CONSPECTUS
OF TAXONOMIC
Lectotype
CHANGES
Designations
a/bovigatusGraham, 1910 ....................................
alpha%guy,1924 .........................................
autiaenia Enderlein, 1920 ....................................
bifoliataTheobald, 190% .....................................
cartroniVentrillon,1905b .....................................
decensTheobald,1901c .....................................
dissimilisTheobald, 1901a ....................................
duttoniTheobald,1901c .....................................
hiisti@a/pisTheobald,1901a ..................................
laurentiNewstead, 1907 .....................................
lAMocosta/is
Graham, 1910 ....................................
madagascariensis
Ventrillon,1905b ..............................
nzcu/usTheobald, 1901b ....................................
minUusTheobald, 1905c .....................................
n@focosta/isTheobald,1909 ..................................
quas@e/idus
Theobald, 1903a ..................................
richterilngramand De Meillon, 1927 ..............................
sabs Theobald, 1908b ......................................
s&psoniTheobald, 19059 ....................................
Neotype
.82
61
.92
82
31
44
.82
.82
.82
.48
45
.92
44
44
.44
92
.75
.87
75
Designation
bitaeniorhynchus
Giles, 1901a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 105
New
Synonymies
alpha Seguy, 1924, synonymywith thei/eriTheobald, 1903a ...............
calloti Rioux and Pech, 1959, synonymy with pipiens Linnaeus, 1758 .........
erectus Iglisch, 1977, synonymy with pipiens Linnaeus, 1758 .............
ethiopicus Edwards, 1912c, synonymy with biaeniorhynchusGiles, 1901 a ......
mauritanicusCallot, 1940, synonymy with SimpsoniTheobald, 1905c ..........
torridusIglisch, 1977, synonymy with pipiens Linnaeus, 1958 .............
61
231
-231
101
75
.231
Harbach: Subgenus Culex in SouthwesternAsia
237
SYSTEMATIC INDEX
Valid taxa are in roman type, valid taxa treated in detail are in bold roman type;
synonyms and invalid taxa are italicized. Boldface page numbers refer to species
treatments; figure numbers and chaetotaxy tables are italicized.
abdominalis (Culiselsa ) , 101
acef (Culex), 232
adairi (Culex), 228
Aedes, 5, 6
aegypti (Aedes), 228
aestuans (Culex),232
ager ( Taeniofhynchus ) , 101, 105
agilis (Culex),230
aikenii (Culex), 234
albolinea tus (Cu lex) , 233
albowifgatus (Culex), 82, 87, 236
algeriensis (Anopheles), 227
alpha (Culex), 61, 66, 236
anafmostus (Culex), 82,87
andersoni (Culex), 66
annulata (Culex), 61
annulata (Culicada ), 235
annulata (Culiseta), 71, 228
annulata ( Tfichopfonomyia), 10
annulus (Culex), 87
Anopheles, 5, 6, 98, 100
antennatus (Culex), 7, 11, 12, 15, 19, 20,
29, 47, 48-52, 55, 56, 60, 65, 66, 77,
78, 80, 81, 96; Figs. 73, 74; 797, 27 7
anxifef (Culex), 232
apoci (Anopheles), 227
Apofoculex, 10
arabiensis (Anopheles), 78, 81, 227
arbieeni (Culex), 78, 81, 228
aseyehae (Cu lex) ,234
ataeniatus (Culex), 53, 57
aufitaenia (Culex), 92, 97, 236
autogenicus (Culex), 230
autumnalis (Culex), 232
azaniae (Anopheles), 227
azoriensis (Culex) , 230
bafbafus (Culex), 233
barraudi subgroup, 110
Barraudius, 7, 8, 9, 49
befbeficus (Culex), 230
berlandi (Aedes), 227
beta (Culex), 81
bicolor (Culex), 23,229
bifoliata (Culex), 82, 87, 236
bifufcatus (Culex), 229
bifoi (Culex), 106
bitaenlorhynchus
(Culex) 11, 12,
13, 16, 19, 22, 70, 78, 81, 101-106,
114, 115; Figs. 2, 37, 38; 203,
223, 236
bitaeniorhynchus subgroup, 11, 91
buxtoni (Coquillettidia), 228
caballus (Aedes), 227
Cacoculex, 10
calcitfans (Culex), 229
calloti (Culex), 231, 236
caftfoni (Culex), 31, 233, 236
caspius (Aedes), 29, 60, 71, 227
Cellia, 98, 100
chelli (Aedes), 227
christophersii(Culex), 234
cinereus (Anopheles), 78, 227
cingulatus (Culex), 232, 233
claviger (Anopheles), 71, 227
condylodesmus (Culex) , 82
consobfinus (Culex), 229
Coquillettidia, 5, 6
coustani (Anopheles), 227
cretinus (Aedes), 228
cfeticus (Culex), 61, 67
cubensis (Culex), 232
Culex (Genus), 2, 5, 6, 7, 10
Culex (Subgenus), 2, 3, 7, 8, 9, 10, 11,
13,29,45,51,63,71,81,93,
110, 227
culicifacies (Anopheles), 227
Culicidae, 5, 227
Culiciomyia, 7, 8
Culiseta, 5, 6
debilis (Phalangomyia ) , 10
decens (Culex), 11, 12, 15, 18, 20,
44-48, 50, 57, 64, 70, 76; Figs
11,72; 790,270, 236
decens subgroup, 11, 47, 51, 77
238
Contrib.Amer. Ent. Inst., vol. 24, no. 1, 1988
demeilloni (Anopheles), 227
deserticola (Culex), 81, 100, 228
detritus (Aedes), 227
ciidieri(Culex), 233
disjunctus (Culex), 23 1
ciissimilis(Culex), 82, 87, 236
doleschallii (Culex), 233
doliorum (Culex) , 230
dolosa (Heteronycha ), 10
domesticus (Culex) , 229
dthali (Anopheles), 81, 227
duplex (Aedes), 228
duttoni (Culex), 8, 11, 12, 13, 17, 19, 70
82-87; Figs. 2, 29, 30; 199, 219,
236
duttoni group, 11, 13
echinus (Aedes), 227
eleuthera (Culex), 10
erectus (Culex), 231, 236
ethiopicus (Culex), 101, 102, 105, 106
236
exiles(Culex), 36, 40
fasciatus (Culex), 229
fatigans (Culex), 23, 31, 232, 233, 234
flavescens (Aedes), 228
fluviatilis (Anopheles), 227
fouchowensis (Culex), 233
fuscus ( Culicelsa ) , 234
gambiae (Anopheles), 29, 78
gelidus (Culex), 10
geniculatus (Aedes), 227
gnophodes (Culex), 92, 235
goughii (Cu lex) , 234
habibi (Anopheles), 227
habilitator (Culex), 10
haematophagus (Culex), 230
hensemaeon (Culex), 234
Heptaphlebomyia, 10, 52
Heteronycha, 10
hirsutipalpis (Culex), 82, 87, 236
hirsutus (Aedes), 227
hispaniola (Anopheles), 71, 81, 101, 227
hortensis (Culex), 71, 228
hutchinsoni (Culex), 86
hyrcanus (Anopheles), 227
impel/ens (Culex), 87, 92, 235
impudicus (Culex), 228
infula (Culex), 104, 105
invidiosus (Culex), 47, 48, 49
jenkinsi, 78
jepsoni (Culex), 92, 235
judaicus (Culex), 228
karatsuensis (Culex), 101
lachrimans (Culex), 234
Laiomyia, 10
Lasioconops, 10
Lasiosiphon, 7, 8, 9
laticinctus (Culex), 11, 12, 13, 14, 15,
17, 20, 23, 48, 64, 66, 67-72, 73,
74, 81, 85, 90; Figs. 27, 22; 195,
215
laurenti (Culex), 48, 49,52, 236
lepidonotus (Aedes), 228
leucomelas (Aedes), 228
Leucomyia, 10
litwakae (Culex), 47, 51
lividocostalis(Culex),45,48, 236
longefurcatus (Culex) , 23, 230
longiareolata (Culiseta), 71,81, 228
luteoannulatus (Cu lex) , 233
luteus (Culex), 229
Lutzia, 7, 8, 93
macleayi (Cutex), 232
maculipennis (Anopheles), 29, 227
maculipes ( Taeniorhynchus) , 92,
97
maculipesarabiensis (Pseudotaeniorhynchus), 101
madagascariensis (Pseudoheptaphlebomyia ), 10, 92, 97, 236
Maillotia, 7, 8, 9
marginalis (Culex), 229
marteri (Anopheles), 227
martinii (Culex), 7, 228
masculus (Culex), 44, 48, 236
mattinglyi (Culex), 11, 12, 15, 17, 20,
64, 66, 68, 69, 71, 72-75, 85, 90;
Figs. 23, 24; 796, 276
mauritanicus (Culex), 75, 78, 236
melanoon (Anopheles), 227
Harbach: Subgenus Culex in SouthwesternAsia
melanorhinus (Culex), 23, 230
meridional& (Culex), 229
microannulatus (Culex), 87, 92, 235
milni ( Culicelsa) , 235
mimeticus (Culex), 11, 12, 13, 16, 19,
22, 71, 81, 90, 97-101, 109, 113;
Figs. 2, 35, 36; 203, 222
mimeticus subgroup, 11, 71
minor (Culex) , 234
minutus (Culex), 44, 48, 236
modestus (Culex), 228
moghulensis (Anopheles), 227
molestus (Culex), 2,23,24,31,229,230,231
montfofli (Heptaphlebomyia ), 52,57
morsitans (Culiseta), 228
multicolor (Anopheles), 71, 81, 227
natronius (Aedes), 227
neavei (Culex), 53, 56, 57, 226
nebulosus (Culex),228
Neoculex, 7, 8, 9
neovishnui (Culex), 111
nigricephala (Cu lex) , 235
nigrirostris(Culex), 234
nigrocanus (Aedes), 228
nigrocostalis (Culex), 44, 48, 236
Oculeomyia, 10
onderstepoortensis (Cu lex), 61, 67
ornatothoracis (Culex), 47
Orthopodomyia, 5, 6
pallidocephala (Culex), 31,57, 233
palkpes (Culex), 23, 230
paludis (Culicelsa ), 235
par (Culex), 92, 97
pavlovsky (Culex), 40
penafieli (Cu lex) , 232
perexiguus (Culex), 11, 12, 13, 14, 18,
20,51,52,53,55,56,
57-61, 63, 65, 66,
71, 75, 81, 96; Figs. 77, 78; 793,
213,226
perfidiosus (Culex), 47
perfuscus (Culex), 47
pettigrewii (Culex), 61, 67
Phalangomyia, 10
pharoensis (Anopheles), 78, 81, 96, 227
phoeniciae (Aedes), 228
phytophagus (Culex), 230
239
pipiens(Culex),2,3,10,11,12,13,15,16
18,21, 23-31, 32, 34, 35, 36, 38, 39,
40,42,43,44,46, 50, 51, 52,55,60, 66
71,81 ;Figs3,4; 786,206,229,230,236
pipiens group,ll,l3,
14, 17, 20, 65, 86
109, 114
pipiens subgroup, 11, 39, 51, 86
plumbeus (Anopheles), 227
poicilipes (Culex), 10, 11, 12, 16, 19,
22, 52, 70, 91, 92-97, 104, 110,
114; Figs. 2, 33, 34; 207, 227
pretoriensis (Anopheles), 81, 227
Pseudoheptaphlebomyia, 10
pseudomime ticus (Cu lex) , 97
(Culex), 11, 12, 13,
pseudovishnui
17, 19, 22, 110, 111-115; Figs.
4 7, 42; 205, 225
pulcherrimus (Anopheles), 227
pulcripalpis (Orthopodomyia), 228
pulchritarsis (Aedes), 228
pungens (Culex), 232
punctipes (Aporoculex ), 10, 92, 97
pusillus (Culex), 49, 51, 71, 228
pygmaeus (Culex), 31, 233
quasigelidus (Culex), 92, 93, 236
quasilinealis (Culex) , 234
quasimodestus (Culex), 230
quasipipiens (Cu lex) , 233
quettensis (Culex), 228
quinquefasciatus
(Culex), 3, 11, 12,
13, 15, 16, 18, 21, 24, 29, 31-35,
39; Figs. 5,6; 187, 207, 232,233,234
raymondii (Culex), 234
reesi (Cu lex) , 233
refiki (Aedes), 228
revocator (Culex), 234
rhodesiensis (Anopheles), 81, 227
richiardii (Coquillettidia), 228
richteri (Culex), 75, 78, 236
rufinus (Cu lex) , 230
fufus (Culex), 229
rusticus (Aedes), 228
sacharovi (Anopheles), 227
saibaii (Culex), 235
salinus (Culex), 235
salisburiensis (Culex), 228
salus (Culex), 87, 92, 235, 236
sarawaki ( Oculeomyia ) , 10
240
Contrib.Amer. Ent. Inst.,vol. 24, no. 1, 1988
seldeslachtsi (Culex), 77
sergentii (Anopheles), 71,78,81, 101, 227
sericeus (Culex) , 233
serotinus (Culex), 232
setivalva (Culiseta) , 228
siamensis (Culex), 106
simplex (Heptaphtebomyia), 10, 52,
57
simpsoni (Culex), 11, 12, 14, 19, 21, 53,
55, 65, 66, 75-78, 79, 80, 81, 100;
Figs. 2, 25, 26; 197, 277, 236
simpsoni subgroup, 11, 50, 77
sinaiticus (Culex), 11, 12, 13, 14, 19, 21,
53, 55, 65, 71, 77, 78-82, 96, 100;
2, 27, 28; 198, 218
sitiens (Culex), 11, 12, 13, 16, 19, 22, 70,
87-92, 94, 95, 99, 100, 107, 109,113,
114; Figs. 31, 32; 200, 220, 235
sitiens group, 11, 13, 16, 19, 22, 51, 86,
96, 100, 104, 109, 110, 114
sitiens subgroup, 11, 91, 96, 110
skusii (Culex) , 232
somaliensis (Culex), 87, 88, 235
splendidus (Anopheles), 227
sp. near infula (Culex), 78
squamosus (Anopheles), 227
stephensi (Anopheles), 227
sternopallidus (Culex), 231
stemopunctatus (Culex), 231
stigmatosoma (Culex), 10
stoehri (Culex), 234
striatipes (Culex), 77
subochrea (Culiseta), 228
subpictus (Anopheles), 227
summorosus (Culex), 106, 107
superpictus (Anopheles), 81, 101, 227
taeniorhynchoides (Culex) , 92, 97
telesilla (Culex), 47
tenax ( Taeniorhynchus), 92
tenebrosus (Anopheles), 227
territans (Culex), 228
terzii (Culex), 77
thalassius (Culex), 78, 88, 91, 107
theileri(Culex),11,12,13,14,18,21,
39,52,55,61-67,69,70,71,73,74,77,
78, 81, 90, 100; Figs. 2, 79, 20; 194,
214, 236
theileri
subgroup, 11, 66, 71, 74
Theobaldiomyia, 10
thoracicus (Cu lex) , 230
tigripes (Culex), 48, 93, 228
tipuliformis(Culex), 36, 40, 62
torrentium (Culex), 11, 12, 13, 14, 16,
18, 21, 29, 36, 39, 40-44, 55; Figs.
2, 9, 10; 189, 209
torridus (Culex), 321, 236
townsvillensis (Cu lex) , 234
Transculicia,10
Trichopronomyia, 10
trifilatus subgroup, 10, 11, 39, 86
trifoliatus (Culex), 47
trifurcatus(Culex), 229
trilineatus (Culex), 233
tritaeniorhynchus
(Culex), 11, 12,
13, 17, 19, 22, 71, 81, 91, 10611 1, 113, 114, 115; Figs. 39, 40;
204, 224
turkhudi (Anopheles), 81, 227
unguiculata (Uranotaenia), 71, 81, 228
unistriatus (Culex), 230
univittatus aggregate, 56, 226
univittatus(Culex),11,12,13,14,18,20
51, 52-57, 58, 59, 60, 63, 65, 75,
77; Figs. 2, 75, 76; 792, 272, 226
univittatus subgroup, 11, 50, 51
Uranotaenia, 5, 6
vagans (Culex), 11, 12, 13, 15, 17, 21,
36-40, 42, 43, 55, 63; Figs. 2, 7, 8;
188, 208
varioannufatus(Culex), 230
vexans (Aedes), 227
virgatipes (Culex), 36, 40
vishnui (Culex), 111, 114
vishnui subgroup, 11, 91, 110
vittatus (Aedes), 227
watti (Culex), 86
whitmorei (Culex), 96, 110
36,
zeltneri (Culex), 31, 233