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Full text of "Mosquitoes; family Culicidae"

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ff ?* 5 0020 
Akademiya Nauk SSSR 



FAUNA OF THE U.S.S.R 

DIPTERA 

Volume III No. 4 



A.V. Gutsevich, A.S.Monchadskii, and 
A. A. Shtakel'berg 

Mosquitoes 
Family Culicidae 



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AKADEMIYA NAUK SSSR • ZOOLOGICHESKII INSTITUT 
Academy of Sciences of the USSR • Zoological Institute 

New Series No. 100 



FAUNA OF THE U.S.S.R 

(Fauna SSSR) 

DIPTERA 

(Nasekomye dvukrylye) 
Volume 3,No.4 

A. V. Gutsevich, A. S. Monchadskii, 
and A. A. Shtakel'berg 

MOSQUITOES 
Family Culicidae 



Chief Editor: B.E.Bykhovskii 
Editorial board: I.M.Gromov, A. S. Monchadskii, A. A. Shtakel'berg 
O.A.Skarlato, and A.A.Strelkov (Volume editor) 



Izdatel'stvo "Nauka" 

Leningradskoe Otdelenie 

Leningrad, 1971 



Translated from Russian 



Israel Program for Scientific Translations 
Jerusalem 1974 



The Publication of this Translation 

was Supported by 

THE UNITED STATES- ISRAEL 

BINATIONAL SCIENCE FOUNDATION 



Copyright © 1974 

Keter Publishing House Jerusalem Ltd. 

Cat. No.61014 4 

ISBN 7065 1475 



Translated by Rose Lavoott 
Edited by Prof. O. Theodor 



Printed in Jerusalem by Keter Press 
Binding: Wiener Bindery Ltd., Jerusalem 



Available from the 

U.S. DEPARTMENT OF COMMERCE 

National Technical Information Service 

Springfield, Va. 22151 



IV/7/3 



Table of Contents 

Preface 1 

Systematic index 2 

INTRODUCTION 5 

Morphological description 5 

A dult mosquitoes 5 

Eggs 16 

Larva 17 

Pupa 40 

Geographical distribution 41 

Importance of mosquitoes for man 45 

Mosquitoes as bloodsucking insects 45 

Mosquitoes as vectors of disease 46 

Methods of collection and of mounting and preserving collections 

of mosquitoes 54 

Adults 54 

Larvae 55 

Bibliography „ 57 

SYSTEMATIC PART 73 

Family Culicidae 73 

Key to genera 73 

I. Subfamily Anophelinae 77 

1 . Genus Anopheles 78 

II. Subfamily Toxorhynchitinae 119 

2. Genus Toxorhynchites 120 

III. Subfamily Culicinae 127 

3. Genus Uranotaenia 128 

4. Genus Orthopodomyia 132 

5. Genus Culiseta 136 

6 . Genus Mansonia 168 

7. Genus Aedes 175 

8. Genus Culex 339 

Index of scientific names 405 



in 



PREFACE 

Volumes on mosquitoes have been published in the series "Fauna of the 
U.S.S.R."byA.A. Shtakel'berg, 1937, Family Culicidae (Bloodsucking 
Mosquitoes of the Palaearctic Region) and in the series "Keys to the Fauna 
oftheU.S.S.R." by A. S. Monchadskii, 1936 and 1951, Larvae of Bloodsucking 
Mosquitoes of the U.S.S.R. and Adjacent Countries. Data from these 
volumes, and original research and material from the literature in the years 
after the publication of the above works, have been included in the present 
volume .. 

The large collections in the Zoological Institute of the Academy of 
Sciences of the USSR (ZIN) form the basis of our work. We are deeply 
indebted to all those who sent additional material to the Institute and who are 
too numerous to list. 

To avoid lengthiness and to facilitate the use of the keys, we have in- 
cluded 85 species which occur in the Soviet Union and also 9 Palaearctic 
species of Aedes which are vectors of diseases or species which probably 
occur in the USSR. 

The authors did not want to burden the book with specialized data or 
lengthy descriptions. The characters given are mainly of diagnostic 
importance. Only a few of the most common synonyms have been given. 
References have been cut to the minimum (only the more important and 
recent publications). The authors of the numerous works on which the 
geographical distribution of the species of mosquitoes is based could not 
be listed. 

Most of the illustrations are taken from the books mentioned above; 
some have been redrawn, and others borrowed from sources which have been 
acknowledged. 

We express our gratitude to E. A. Afanas 'eva, T. A. Lyakhtinen and 
K. G. Fedorova for their long years of work spent in making the preparations, 
and in arranging and preserving the collection of mosquitoes at the 
Zoological Institute. 

January 1968 



SYSTEMATIC INDEX" 

Fam. Culicidae —Bloodsucking mosquitoes 
I. Subfam. Anophelinae 
1 . GenusA nopheles Mg. - 

1. Sub gen. Anopheles Mg. 

1. An. (An.) algeriensis Theob 80 

2. An. (An.) plumbeus Steph °£ 

3. An. (An.) claviger Mg °° 

4. An. (An.) marteri Sen.-Prun «» 

A/z. m. marteri Sen.-Prun 90 

Arc. m. sogdianus Kesh 90 

5. Arc. (An.) maculipennis Mg 90 

Are. m. sacharovi Favre 99 

6. An. (A re J lindesayi Giles J 00 

An. Z. lindesayi Giles 103 

Aw. I. japonicus Yam 103 

7. An. fAreJ hyrcanus Pall 103 

2. Subgen. Myzomyia Blanch. 

8. An. (M.) pulcherrimus Theob 108 

9. A«. (M.) superpictus Grassi 113 

II. Subfam. Toxorhynchit inae 
2. GenusT oxorhynchites Theob. 

1. T. christophi Portsch H8 

III. Subfam. Culicinae 
3. Genus Uranotaenia Arrib. 

1. U. unguiculata Edw 126 

4. Genus Orthopodomyia Theob. 

1. O. pulchripalpis Rond < - 130 

5. Genus C u I i s e t a Felt 
!• Subgen. A llotheobaldia Brol. 

1. C. (All.) longiareolata Macq 138 

2. Subgen. Culiseta Felt 

2. C. (C.) glaphyroptera Schin 142 

3. C. (C.) alaskaensis Ludl 144 

C. a. alaskaensis Ludl 147 

C. a. indica Edw 147 

4. C. fCJ annulata Schr 147 

C. a. annulata Schr 150 

C a. subochrea Edw 150 

5. C. (C.) bergrothi Edw 151 



[ This index has been photographically reproduced from the Russian original. The pages indicated 
are Russian page numbers; these appear in the left-hand margin of the text. ] 



3.Subgen. Culicella Felt 

6. C. (C.J morsitans Theob 153 

7. C. (C.J ochroptera Peus 156 

8. C. (C.J fumipennis Steph 159 

9. C. (C.J setivalva Masl 160 

6. Genus M a n s o n i a Blanch. 

1. M. richiardii Fie • 164 

2. M. buxtoni Edw 167 

7. Genus A e d e s Mg. 
l.Subgen. Ochlerotatus Arrib. 

1. A. (O.J caspius Pall 191 

A. c. caspius Pall 192 

A. c. dorsalis Mg 195 

2. A. (O.J mariae Serg 196 

3. A. (O.J pulchritarsis Rond 199 

A. p. pulchritarsis Rond 200 

A. p. asiaticus Edw 201 

4. A. (O.J cantans Mg 202 

5. A. (O.J riparius D. K 205 

A. r. riparius D. K 208 

A. r. ater Guts 208 

6. A. (O.J behningi Mart 208 

7. A. (O.J excrucians Walk 211 

8. A. (O.J beklemishevi Den 214 

9. A. (O.J annulipes Mg 216 

10. A. (O.J flavescens Mull 218 

11. A. (O.J cyprius Ludl 221 

12. A. (O.J rusticus Rossi 223 

13. A. (O.J refiki Med 225 

14. A. (O.J lepidonotus Edw 227 

15. A. (O.J subdiversus Mart 230 

16. A. (O.J communis Deg 232 

17. A. (O.J pionips Dyar 235 

18. A. (O.J punctor Kirby 237 

19. A. (O.J hexodontus Dyar 240 

20. A. (O.J sticticus Mg 243 

21. A. (O.J nigrinus Eck 245 

22. A. (O.J hungaricus Mih 246 

23. A. (O.J diantaeus H. D. K 247 

24. A. (O.J intrudens Dyar 250 

25. A. (O.J pullatus Coq 252 

26. A. (O.J nigripes Zett 255 

27. A. (O.J impiger Walk 258 

28. A. (O.J cataphylla Dyar 261 

29. A . (O.J leucomelas Mg 263 

30. A. (O.J detritus Hal 264 

31. A. (O.J simanini Guts 267 

32. A . (O.J kasachstanicus Guts 269 

SUPPLEMENT (Aedes) 
2.Subgen.Aedimorphus Theob. 

33. A. (A.J vexans Mg 270 

A . v. vexans Mg 273 

A . v. nipponii Theob 273 

Appendix (Aedes) 

A. (O.J montchadskyi Dubitzky 270a 

A. (O.J stramineus Dubitzky 270b 

A. (O.J fitchii Felt et Young 270c 

A. (O.J rempeli Vock 270d 



3.Subgen. Finlaya Theob. 

34. A. (F.J geniculatus 01 274 

35. A. (F.) echinus Edw 27S 

36. A . (F.J nipponicus LaCasse et Yam 280 

37. A . (F.J seoulensis Yam 283 

38. A. (F.) alektorovi Stack 284 

39. A. (F.) togoi Theob 288 

40. A. (F.J japonicus Theob 290 

41. A. (F.) koreicus Edw 291 

4. Subgen. Stegomyia Theob. 

42. A. (S.J aegypti L 294 

43. A. (S.J cretinus Edw 298 

44. A. (S.J vittatus Big 299 

45. A. (S.J galloisi Yam 300 

46. A. (S.J albopictus Sk 303 

47. A. (S.J flavopictus Yam 305 

48. A. (S.J chemulpoensis Yam 306 

5. Subgen. Aedes Mg. 

49. A. (A.) cinereus Mg 308 

A. c. cinereus Mg 310 

A. c. rossicus D. G. M 311 

A . c. esoensis Yam 312 

50. A. (A.) nobukonis Yam 313 

51. A. (?) aureus Guts 313 

8. Genus C u I e x L. 
1. Subgen. Lutzia Theob. 

1. C. (L.J fuscanus Wied 322 

2. C. (L.J vorax Edw 324 

2. Subgen. Barraudius Edw. 

3. C. (B.J modestus Fie 325 

4. C. (B.) pusillus Macq 327 

3. Subgen. Neoculex Dyar 

5. C. (N.) territans Walk 331 

6. C. (N.) hortensis Fie 333 

7. C. (N.) martinii Med 335 

8. C. (N.J hayashii Yam 336 

4. Subgen. Culex L. 

9. C. (C.) bitaeniorhynchus Giles 341 

10. C. (C.J sinensis Theob 345 

11. C. (C.J whitmorei Giles 345 

12. C. (C.J mimeticus Noe 348 

13. C. (C.J jacksoni Edw 350 

14. C. (C.) orientalis Edw 352 

15. C. (C.J tritaeniorhynchus Giles 354 

16. C. (C.J univittatus Theob 358 

17. C. (C.J theileri Theob 360 

18. C. (C.J vagans Wied 362 

19. C. (C.J torrentium Mart 364 

20. C. (C.J pipiens L 365 



INTRODUCTION 



7 MORPHOLOGICAL DESCRIPTION 

ADULT MOSQUITOES 

The lateral surfaces of the usually spherical head are nearly completely 
occupied by the large, compound eyes which are situated close together or 
are contiguous in the upper and lower part of the head and are usually 
kidney- shaped. The number of ommatidia of one eye is 300—1,000. The 
size of each ommatidium is usually slightly greater in females than in 
males, and the number of ommatidia is therefore slightly larger in males. 

The clypeus is situated in the anterior part of the head before the base 
of the antennae, and the vertex above the antennae between the eyes. The 
posterior part of the head is named the occiput. 

The antennae are situated close together on the anterior part of the head 
(Figure 2). The number of antennal segments is 15 in both sexes. The 
first segment (scape) is ring-shaped or plate-shaped. The second segment 
(torus) is relatively large, especially in the male, and more or less spherical. 
It contains the organ of Johnston, which is the organ of hearing. Its structure 
9 varies in the different genera (Risler, 1955). The other 13 segments form the 
flagellum. These are generally of the same type, almost cylindrical and 
more or less of the same length, except the last segments in the male which 
are much longer. The hairs on the antennae form rosettes and are much 
denser and longer in the male (Figure 2, C). The hairs of some antennal 
segments form hornlike appendages in the males of some groups (subgenus 
Lophoceratomyia of the genus Culex). 

The antennal hairs, which are of various structure and function, are 
sensillae of different types. There are 5 types of hairs on the antennae of 
Aedes aegypti (Slifer and Sekhon, 1962). The thick-walled hairs are 
mechanoreceptors, the thin-walled hairs with numerous small openings are 
chemoreceptors. Openings in the walls of the hairs are visible under the 
electronic microscope. 

The buccal organs form a proboscis (Figure 2). The proboscis of the 
female consists of a labrum, a pair of mandibles, a pair of maxillae, a 
hypopharynx and a labium. The labrum forms a canal through which liquid 
food enters the intestine. The hypopharynx has a canal through which the 
saliva flows out during bloodsucking. In the resting state, all these parts 
are situated in the groove of the labium. 

Females of all Palaearctic mosquitoes, except Toxorhynchites, feed 
on plant juices and the blood of vertebrates. When sucking blood, the 
piercing parts penetrate the skin. The labium becomes folded in two and 
remains outside (Figure 2). The piercing parts move in different directions 



in the skin and penetrate a capillary, and then bloodsucking begins. Sucking 
of blood extravasated from the capillary is also possible. Bloodsucking 
lasts slightly longer in this case. 



(7) 



Proboscis 




Tarsus 



FIGURE 1. Structure of the mosquito: 

I - head and its appendages; II - thorax; III - abdomen; 
1—8 — segments of abdomen. 



(8) 





CH- 



<u P 
ca E 

3 3 



> D. 

ca >> 




M 


~H 


c 






Ifl 


o 

3 


I 


■R 


X 





b 



a e 




(U c 

■S - 



.- c 



X 5 



Male mosquitoes do not suck blood. Their proboscis differs from that 
of the female in the absence or reduction of the mandibles and maxillae. 
The degree of reduction of the mandibles and maxillae varies in males of 
different genera (Marshall, 1938). 

At the sides of the proboscis are the maxillary palps; these are seg- 
mented appendages which are long in both sexes of Anopheles and usually 
in the males of other genera, but are very short in the females of Culicinae, 
in the males of Uranotaenia and in the subgenus A e d e s (Palaearctic 
species). The palps consist of 5 segments but segment 5, sometimes also 
segment 4, may be absent or form a small rudiment. 



Thorax 

The thorax (Figure 3) is divided into 3 parts: prothorax, mesothorax 
and metathorax, each of these forming a ring which consists of several 
sclerites. The dorsal sclerites form the mesonotum, the lateral sclerites 
the pleurae of the thorax. The largest part is the mesothorax, in which the 
wings are situated. The dorsal surface of the thorax, the mesonotum, is 
10 uniformly convex. In the median line of the mesonotum extends a row of 
setae, the acrostichals, and lateral to them, in about the middle between the 
median line and the lateral margins, there is also a longitudinal row of 
setae, the dorsocentrals. Behind the mesonotum is the scutellum, which is 
3-lobed in the Culicinae, uniformly rounded in Anopheles. Below the 
scutellum and behind it is the postnotum. 




FIGURE 3. Thorax of mosquito, lateral: 



CS — cervical sclerite. Prothorax: p — pronotum; prepm — proepimeron; 
preps — proepisternum. Mesothorax: m — mesonotum; sc — scutellum; 
pn — postnotum; as — anterior spiracle; mesp — mesepisternum (dorsal part); 
stp - mesepisternum (ventral part — "sternopleuron"); w - attachment of 
wing; mepm — mesepimeron; mes — meron. Metathorax: ps — posterior 
spiracle; mteps — metepisternum; mtepm — metepimeron. Legs: Ic.IIc, 
IIIc - coxae. Abdomen: 1 1 - tergite 1; st 1 - sternite 1. 



The lateral parts of the thorax (pleurae) also consist of several sclerites 
separated by sutures. The lateral sclerites of each segment are divided 
into two parts: episternum (anterior part) and epimeron (posterior part). 
In lateral view, the thorax consists of the following parts (Figure 3): pro- 
notum and episterna and epimera of the mesothorax. The mesothorax of 
mosquitoes is divided into two parts: a small anterior dorsal and a large 
ventral part. The ventral part is wedged in the form of a triangle between 
the base of the fore and mid-legs and is called "sternopleuron." Further 
posteriorly are the epimera of the mesothorax (mesepimera) and the 
episternum and epimeron of the metathorax. The membranous area be- 
tween the base of the fore legs and the anterior ventral margin of the sterno- 
pleuron is called the postccxal area. The arrangement of the scales on the 
pleurae, particularly on the proepimeron, sternopleuron and mesepimeron, 
are important in the systematics of mosquitoes, especially in the genus 
A e d e s. 



(11) 




FIGURE 4. Arrangement of setae on the pleurae: 

A-Culiseta; B-Aedes; p - pronotal; preps - proepisternal; prepm - proepimeral; sp - spiracular; 
postsp-postspiracular; preal-prealar; stp - sternopleural ; dm - dorsal mesepimeral; vm -ventral mesepimeral. 



Between the epimeron of the prothorax, the episternum of the meso- 
thorax and lateral margin of the mesonotum is situated the anterior 
spiracle. At the border of the epimeron of the mesothorax and episternum 
of the metathorax is the posterior spiracle. The spiracles are important 
because not only air passes through them into the tracheae but water is 
11 evaporated. It has been shown experimentally in Anopheles maculi- 
pennis that evaporation takes place at the surface of the tracheae (Vino- 
gradskaya, 1953). The size of the spiracle depends on the intensity of 
evaporation. In mosquitoes living in arid zones, the spiracles are smaller 
than in mosquitoes in humid climates. An index of the size of the spiracle 
is the spiracle index, the ratio between the length of the anterior spiracle 
and the length of the thorax, expressed in percentages (Vinogradskaya, 1950). 

Near the spiracles and in other parts of the pleurae are groups of setae 
the arrangement of which is constant and distinct in many groups of the 
family. The arrangement of the setae is denoted as follows (Figure 4): 
pronotal (on the pronotum); proepisternal; proepimeral; spiracular (im- 
mediately before the anterior spiracle, between it and the margin of the 
proepimeron); postspiracular (behind the anterior spiracle in the dorsal 
part of the mesepisternum); sternopleural; dorsal and ventral mesepimeral. 



Legs 

There are 3 pairs of legs on the thorax — fore, mid- and hind legs, 
situated on the prothorax, mesothorax and metathorax. The legs consist of 
the following parts: a short coxa articulated with the thorax; the trochanter, 
which forms a small ring; the femur, which forms a long cylinder which is 
sometimes slightly thickened at the base; the tibia, also long, slender and 
cylindrical; the 5- segmented tarsus with paired claws at the end (Figure 5). 



12 



pulv. 



FIGURE 5. Terminal appendages of the tarsus (from E.N. Pavlovskii): 

A— praetarsus of Culex; B — claws of a female of Aedes; C — claws of afemale of Culex; 
pulv — pulvilli. 



On the ventral, concave side of the claws there may be 1 or 2 denticles. 
The arrangement of the claws is expressed by the formula 2.1; 2.1; 1.1, 
i. e.: one claw of the anterior tarsus has 2 denticles, the other one denticle; 
12 the claws of the mid-tarsus are the same; both claws of the hind tarsi have 
one denticle. The formula 0.0; 0.0; 0.0 indicates that all claws are simple. 



10 



The structure of the claws differs in different species (particularly in the 
genus Aedes), and this character is therefore important in the determina- 
tion of some mosquitoes (Vockeroth, 1950, Sazonova, 1958). The form of 
the claws is nearly the same on all legs but the size of the claws decreases 
from the fore to the hind legs, and the claws of the fore tarsus are there- 
fore mainly used. 

There are pulvilli, small, oval plates covered with hairs (Figure 5), under 
the claws in some genera. Between the claws is a single structure, the 
empodium, which often forms a feathered or branched seta. 

The following should be remembered about the topography of the legs. 
The articulation of the coxa, trochanter and femur is multiaxial so that it 
may occupy different positions with respect to the main axis of the body. 
Thus, for a convenient orientation, the knee or the monaxial articulation of 
femur and tibia should be used. If the knee is bent, the femur and tibia face 
each other with the ventral surface. If the leg is extended horizontally and 
laterally this surface is ventral and the opposite surface dorsal. The sur- 
faces perpendicular to these surfaces are the anterior and dorsal surfaces. 



Wings 

The wings are usually elongate -oval and supported by longitudinal veins 
and cross veins. The point of attachment of the wings to the thorax is their 
base. The distal end of the wings is the apex. When the wings are spread 
(i. e. when the long axis of the wing is perpendicular to the longitudinal axis 
of the body), the margin of the wing which is directed anteriorly forms the 
anterior margin and the opposite margin the posterior. 




«+5 



FIGURE 6. Wing of Culex: 

Veins: c - costa (marginal); sc - subcosta; r lt >r 2 ,r 3 r 4+5 - radius; m 1 ,m 2 , m 3 + 4 [or cu x ] - media; 

m x [or cu 2 ] — cubitus; an — anal, r-m — anterior cross-vein; m-cu — posterior cross- vein; h — humeral. 

The longitudinal veins are named as follows (Figure 6): costa, or 
marginal vein (c ) along the outer margin of the wing (it is present along 
the anterior and posterior margin in the mosquito); the next vein is the 
13 subcosta (sc), which begins at the base of the wing, extends nearly parallel 
to the anterior margin and ends in the costa in the apical third of the wing. 
Then follows the radius, which in the mosquito always has 4 branches (n, r2, 



11 



^3, r*4+5; ^4+5 indicates that this vein has been formed by the fusion of r$ 
and rs, which are present separately in some primitive forms related to 
mosquitoes). The radius begins in a common stem at the base of the wing, 
the radial veins then branch and the stem r2 + r3 + r4+5 branches from ri; 
from it branches r4+s; branches r2 and r3 form the anterior or radial fork 
of the wing; section r2+3 from the cross-vein to the base of the fork is 
called the stem of the radial fork. The next vein is the media (m); in the 
Culicidae it has 3 branches — mi, m2 and m3+4* the first two branches form 
the median fork, the last branch together with the cubitus (cu) forms the 
posterior fork in the apical part of the wing. The posterior longitudinal 
vein is called the anal vein (an). 

There are three cross-veins: the humeral (h), between costa and sub- 
costa near the base of the wing, the radio-medial (r— m) between the radius 
and media in the middle of the wing, and the medio-cubital or posterior 
cross-vein (m— cu) between media and cubitus. 

The longitudinal veins and the cross-veins form cells which are named 
after the veins anterior to the cell. The veins are indicated by lower-case 
letters, the cells by capitals, e. g. Ri, M, Cu, An. 

The veins are covered with scales which are more densely arranged in 
the female. The scales on the veins are usually narrow, linear or lanceolate, 
rarely wide, e. g. in the genus Mansonia. The wing membrane is 
covered with minute hairs — microtrichia — that are visible only under the 
microscope. 

The club-shaped appendages of the metathorax are characteristic for the 
Diptera; these are the halteres, which are the rudiments of the posterior 
14 pair of wings. The halteres function as a gyroscopic organ for the main- 
tenance of equilibrium. There are two groups of sensillae at the stem of 
the halteres. 



Abdomen 

The abdomen of mosquitoes is usually elongate-cylindrical and consists 
of 10 segments; the last two segments are markedly modified and included 
in the genitalia. The first eight segments consist of a dorsal sclerite, the 
tergite and a ventral sclerite, the sternite; they are connected by the 
pleural membrane on which the abdominal spiracles are situated on seg- 
ments 2—7. In the young female with undeveloped ovaries, the lateral 
margins of the tergites are curved ventrally and are not visible in dorsal 
view. The membrane connecting the tergites with the sternites forms 
folds. The abdomen of a dry mosquito has the same appearance. After 
feeding and the development of the ovaries, the tergites become straightened 
and the pleural membranes distended. 

Segments 9 and 10 form the genitalia. The male genitalia (Figures 7—9), 
the hypopygium, are of complicated structure, different in every species. 
The structure of the genitalia is therefore of great importance in the 
systematics of mosquitoes. The species of a mosquito can often be deter- 
mined with certainty only by the structure of the genitalia. There is no 
uniform nomenclature of the parts of the genitalia. 

* [In the usual interpretation the media is 2 -branched and the cubitus 2 -branched.] 



12 




FIGURE 7. Hypopygium of Aedes: 

gc — gonocoxite (coxite); gst — gonostyle (style); a.c. — 
apical claw; a.l. — apical lobe; b.l. — basal lobe; s.c. — 
stem of claspette; w.c. — wing of claspette; tg 9 — lobes of 
tergite 9. 

The hypopygium of males is rotated through 180° after hatching so that 
tergite 9 is then situated ventrally and sternite 9 dorsally. The terms 
"tergite" and "sternite" are kept for the morphologically dorsal and ventral 
sclerites of segment 9. 

Segment 9 consists of a dorsal and a ventral sclerite. The true tergite 9 
15 is a narrow sclerite at the sides of which, at the posterior margin, are 
processes named lobes of tergite 9. They usually bear setae or hairs at 
the apex. In some species of S t e go m y i a, the tergite has a median, 
conical or tongue- shaped lobe in addition to the lateral lobes. The true 
sternite 9 is usually a narrow strip which sometimes bears lateral 
processes. 

The largest part of the hypopygium is formed by the gonopods (valves), 
which are paired formations. Each gonopod consists of 2 segments, the 
gonocoxite and the gonostyle. These parts will be named simply coxite and 
style. The coxite is a large formation which usually has an apical and a 
basal lobe on the inner side (in most species of Aedes, Figure 7). There 
is sometimes a lobe in the middle of the coxite (Culex, Culiseta) or 
near the apex, as in the genus Culex (Figure 8). The inner side of the 
coxite usually bears spines or 1 — 3 large setae, the position, structure and 
number of which are characteristic for some species, particularly in the 
genus Anopheles (Figure 9). The spines, if present, are usually situated 
on the basal lobe in Aedes. The coxite is covered with hairs or scales 
on the outside, dorsally and ventrally. 

At the apex of the coxite or slightly subapically (subgenus Aedes and 
others) is situated the narrow, movable style, which is sometimes widened. 



13 



At the apex of the style is a finger-shaped or spine-shaped claw which is 
sometimes very short. 




FIGURE 8. Hypopygium of Culex: 

st — style; c — coxite; s.l. — subapical lobe; 
ph^ ph 2 — first and second part of phallosome; 
st 10 — sternite 10; b.p. — basal process of 
sternite 10. 



FIGURE 9. Hypopygium of A no pheles (after Coe, 
Freeman and Mattingly): 

st — style; c — coxite; cl — claspettes; ae — aedeagus. 



16 Inside of the coxites are the claspettes, which are processes of the basal 

part of the coxite. The claspettes are completely developed only in some 
groups; a special development is present in the subgenera Ochlerotatus 
and F inlay a of the genus Aedes in which the claspettes are 2 -segmented 
formations consisting of a basal part, the stem, and an apical part, the wings, 
the form of which varies in different species. There may be a transparent, 
platelike widening at the convex side of the wings. Claspettes are absent 
in Culex and other genera. 

Near the median line are the sclerites of the 10th (anal) segment which 
surround the anus. Tergite 10 is usually represented by 2 small sclerites 
which are connected with sternite 10 at the apex. Sternite 10 (paraproct) 
usually consists of two distinct, elongate sclerites which are sometimes 
covered with short hairs, especially in C ul e x, in which the spines or 
denticles at the apex of sternite 10 and the development of its basal process 



14 



are characteristic. The sclerotized apex of each sclerite of sternite 10 
forms a simple or denticulate claw. 



st 10 




st 9 




FIGURE 10. Inner sclerites of the hypopygium 
of A edes, lateral (after Matheson): 

t 9 - tergite 9; st 9 - sternite 9; t 10 - 
tergite 10; st 10 — sternite 10; ph — phallo- 
some; p - parameres; b.p. - basal plate. 



FIGURE 11. Abdomen of female (after Coe, 
Freeman and Mattingly): 

A - Anopheles; B-Culex; C-Aedes. 



The organ of copulation is the aedeagus (Figure 10), a complicated 
formation which consists of several sclerites: lateral sclerites, parameres 
and phallosome (mesosome). The phallosome of Culex is very compli- 
cated and usually consists of 2 parts which are divided into several denticles, 
The structure of the phallosome is of great systematic importance. 

The terminal segments of the abdomen of the female are relatively little 
17 modified. Sternite 8 is usually simple. In the subgenera Stegomyia 

and Finlaya of the genus A e d e s it attains a high degree of development. 
At the end of the abdomen are the paired cerci (Figure 11, C). They are 
sometimes distinctly developed as two small, elliptical sclerites, sometimes 
more or less covered. The development of sternite 8 and of the cerci in 
groups of the genus Aedes are characters connected by reverse 
correlation. 

In cleared preparations of the posterior end of the abdomen of the female, 
spermathecae in the form of sclerotized capsules are visible in segments 
7—8. Their number varies: there may be one (Anopheles, Urano- 
taenia and rarely in some species of Aedes), two (some species of 
Mansonia) or three (most Culicinae). 

The structure of the female genitalia is not very important in the 
systematics of mosquitoes, but is quite characteristic for some genera and 
subgenera (La Casse and Yamaguti, 1955). 

The structure of the cutaneous appendages is important in systematics. 
These appendages are scales (elongate, rarely broad, longitudinally striated), 



15 



spines (thick setae, for example at the basal lobe of the coxite, in definite 
numbers in each species), setae (thinner than spines), and hairs. 



EGGS 



Head 



The female of Anopheles deposits her eggs singly on the surface of 
the water, and they may form geometric patterns. The females of Culex, 

Mansonia and the subgenus Culi- 
seta deposit the eggs in compact rafts, 
the form of which is determined by the 
space between the crossed posterior 
legs of the mosquito. In A e d e s and in 
the subgenus Culicella of the genus 
Culiseta, the eggs are usually 
deposited on moist soil, in depressions 
where they may remain dry for a long 
time. The larvae hatch after the site 
of oviposition has become flooded (after 
rain or thawing of snow). The diapause 
in the egg is sometimes very long, 
especially in species of A e d e s . 

The eggshell consists of three layers. 
The inner layer is a thin, vitelline 
membrane which encloses the egg cell 
with yolk. The next layer is the chorion, 
which consists of two layers: a median 
layer, the endochorion, which is hard 
and opaque, and an outer layer, the exo- 
chorion, which is usually soft and trans- 
parent. On the exochorion are sculp- 
tural patterns which are characteristic 
for some genera and species. 

At the anterior end of the eggs 
(C u 1 e x, A e d e s ) or slightly subapically 
(Anopheles) is the micropyle, which 
permits the entry of the spermatozoa 
into the egg and fertilization. It con- 
sists of a rosettelike membrane sur- 
rounded by the exochorion which forms 
an annular ridge. In the center of the 
rosette is an opening with a canal leading 
inside; the canal is covered inside with 
a plug which opens only when the sperm 
enters. 

The eggs of Culicinae are oblong-oval 
or nearly cylindrical with a wider 
anterior end and a narrower, blunt 




FIGURE 12. Larva of A no phel es, dorsal. 
Roman figures — segments of abdomen. In 
parentheses — primitive segments. 



16 



posterior end. The surface often (A e d e s and others) has a reticulate, 
polygonal pattern. The reticulation is narrower in Culiseta and much 
wider in Mansonia. 

The eggs of Anophelinae are boat- shaped, with slightly concave upper 
and convex lower surface. They are usually surrounded by a narrow fringe 
which is interrupted in the middle by larger air floats with a varying 

number of cells in the different 
species. In Anopheles macu- 
lipennis sacharovi, the eggs 
of summer generations have only 
a fringe and small air floats in 
addition to the fringe in winter. 
The pattern of the exochorion of 
the upper surface of the eggs, 
especially in the subgenera of 
A. maculipennis, is important in 
systematics (see pages 97 and 98). 




19 



LARVA 

The body of the larva consists 
of three parts (Figures 12 and 13): 
the head, which is enclosed in a 
sclerotized capsule, the thorax, 
which consists of 3 fused segments, 
and the abdomen, with 9 segments. 
The last segment is curved ventral- 
ly and at its end is situated the 
anus, surrounded by 2 pairs of 
gills. On the ventral side of the 
last segment is the organ of move- 
ment, the fin or ventral brush. 
On the dorsal side of segment 8 of 
larvae of Anopheles is the only 
functional, posterior pair of 
spiracles surrounded by a valvular 
apparatus which forms a stigmatical plate; in the larvae of Culicinae, the 
posterior pair of spiracles with the stigmal plate is situated at the end 
of the siphon, which is situated on the dorsal side of segment 8. The thorax 
is the widest part. The abdominal segments are narrower. 

The 4th- stage larvae are 0.5- 0.7 cm long or small species (Urano- 
taenia, some species of C ul e x) and 1.5 cm long or longer in the large 
species (Culiseta, Aedes, T o x o r hyn c h it e s, and others). 

The color of the larvae varies. The cuticle, except the sclerotized 
parts (head, siphon, etc.), is transparent and colorless and the color of the 
body depends therefore on the color of the inner organs and tissue, mainly 
the fat body. The pigment is concentrated mainly on the dorsal side of the 



FIGURE 13. Larva of A edes cinereus. Lip. 



17 



fat body. The inner parts of the fat body are practically without pigment. 
The dorsal side is always darker than the ventral, which is important for 
species developing in the spring at low temperatures since the pigment 
absorbs heat rays and the development of the larvae takes place at more 
favorable temperatures. The larvae of the same species are therefore 
more strongly pigmented in the north than in the south. 

The larvae of Toxorhynchites and Orthopodomyia are wine-red 
The larvae of Ochlerotatus and Culiseta are light to dark brown or 
blackish; the larvae of Mansonia are yellowish; those of Culex light 
brown or grayish green. Many larvae which are transparent in the early 
stages become bluish green (C.territans)in the 4th stage. Greenish 
shades (yellowish green to brownish green) are characteristic for 4th-stage 
larvae of Anopheles; the earlier stages are nearly black. 



Hairs of the body 

The body of the larva is covered with numerous, different hairs. The 
longer hairs together form an extensive sensory zone around the body. They 
sense the slightest disturbance of the water caused by predators and induce 
the larva to change its position. Especially long hairs are present in 
larvae (Aedes alektorovi, Orthopodomyia p u 1 c h r i p a lp i s, and 
others) which live in tree holes, where they are so crowded and food is so 
scarce that cannibalism often occurs. In larvae of other species living in 
tree holes, the hairs are short and form spines (Toxorhynchites, 
Aedes geniculatus, A. echinus, A. g a 1 lo i s i, etc.). Larvae living 
on the surface of the water (A no ph e 1 e s) have short or specially modified 
20 hairs on the dorsal side and strongly developed lateral hairs, while larvae 
with a submerged mode of life have more or less uniformly developed hairs 
on all sides. 

Two types of cuticular formations are especially important. 

The first type are various sculptured appendages of the cuticle: small 
spines (chaetoids) which often cover the body densely (Aedes cyprius, 
Culex hortensis); short, pointed spines on the antennae or at the 
posterior margin of the saddle of the last abdominal segment whi'ch form 
combs; microscopic spines on the siphon, and others. This type of forma- 
tion includes the scales of the comb at the sides of segment 8 and the 
denticle of the pecten on the siphon since there are all transitions between 
them and the typical, small spines of the chetoid type. All these formations 
are very variable in number; the more specialized denticles of the pecten 
and the scales of the comb are less variable. 

The second type are the true hairs. Each hair is hollow with slightly 
widened base situated in a cup-shaped, articulated pit which forms a ring 
around the base. The cuticle surrounding the ring may be thickened in the 
form of a dark sclerite which sometimes encloses a group of hairs near it 
(Toxorhynchites, Anopheles, etc.) or it forms a tubercle with pro- 
cesses (lateral hairs on thorax and abdominal segments). These hairs are 
often of systematic importance. Their position and number are constant 
not only in larvae of the same species but often also in the genus or in the 



18 



whole family. The structure of the hairs and their branching are very 
different and often variable. Stronger branching of a hair in the same 
species is connected with the delicacy of its branches and vice versa. 

In the simple cases, the hairs form a long stalk which tapers apically; 
its surface may be smooth or bear secondary feathering; these secondary 
hairs may be short or long or situated on the same plane (Anopheles). 
The hairs may be divided at the end to a greater or lesser degree. The 
second group consists of branched hairs which may have a short stalk and 
then branch dichotomously, dendriform, irregularly (outer clypeal hairs of 
Anopheles) or form a fan on a short base. In some cases the branches 
are not situated in the same plane but in a circle, like the margin of a poly- 
hydral pyramid (A e d e s g e ni cu 1 at u s and others). The hairs may be 
compressed and each branch is oblong, leaf- shaped (palmate hairs of 
Anopheles, Figure 22). All hairs situated in one basal ring should be 
considered as separate. 

In addition to the hairs, the body of the larvae may bear large, dark 
sclerites of varying form. They are often restricted to areas where 
muscles are attached (dorsal abdominal plates of Anopheles) or may be 
important as supports (saddle of last abdominal segment, plates at sides of 
segment 8 in Uranotaenia and others). These plates together with the 
spines often protect the larvae from predators (Orthopodomyia, 
Toxorhynchites). 



21 Head 



The chitinized capsule of the head has a flattened ventral side and a 
convex dorsal side. Its form varies from oblong-ovoid (Anopheles, 
Figure 14) to broadly transverse-oval (Mansonia, C u 1 e x, etc., Figure 15). 
It is nearly rectangular in Toxorhynchites. These forms are con- 
nected with the nutrition of the larvae and the characteristics of their mouth 
parts. 

The dorsal side of the head is formed mainly by the large frontoclypeus, 
which is usually oblong and widest before the eyes. It is nearly square in 
larvae of Toxorhynchites (Figure 57). The frontoclypeus is laterally 
and posteriorly bordered by the frontal suture. Before it is situated the 
clypeus, which is separated from it by the epistomal suture. The form of 
the clypeus is connected with the structure and function of the labrum: it 
is narrower in the middle. The anterior part of the frontoclypeus is 
morphologically the clypeus because the muscles of the labrum originate 
there and the epipharyngeal musculature originates in its posterior part. 
The frontal sclerite is therefore really the frontoclypeus and the clypeus 
the preclypeus. 

Lateral to the frontoclypeus are the epicranial plates. They reach to 
near the median line posteriorly and form the short coronal suture. The 
epicranial plates extend on the ventral side to the base of the maxillae 
where the sutures begin which separate the genae from the ventral median 
sclerite, the gula. The anterior margin of the gula reaches to the base of 
the labium. 



19 




FIGURE 14. Head of larva of A no p he 1 es, dorsal (after Puii): 

A - 1st stage; B - 4th stage; fr - frontoclypeus; f rs . - frontal suture; o.sc. - occipital 
sclerite ("collar"); o.s. - occipital suture; c.s. - coronal suture; e.t. - egg tooth; 
2,3- inner and outer clypeal hairs; 4— postclypeal hairs; 5-7 - inner, median and 
outer frontal hairs; 8 - sutural hair; 9 - transsutural hair; 12 - hair at base of antenna; 
14 - supraorbital hair. 

In the middle of the gula of the 4th- stage larvae of Anopheles and 
22 others, especially posteriorly, is a longitudinal suture which does not extend 
to the base of the labium. During the molt, the head capsule becomes 
divided along the frontal suture into 3 parts: the frontoclypeus is raised 
and both epicranial plates are pushed aside (Figure 14). 

The posterior margin of the head forms the occipital foramen, which is 
wide in Anopheles and Toxorhynchites and narrow in other larvae . 
The foramen is surrounded by a dark, narrow, ring-shaped sclerite (collar). 
The occipital suture is interrupted dorsally near the coronal suture and 
ventrally near the median line. 

The occipital sclerite of Anopheles changes during development. In 
larvae hatching from the eggs it is very narrow but in lst-stage larvae it grows 
and forms 70% of the length of the head before the molt. This is repeated 
in the 2nd stage. During the 3rd stage, the length of the collar decreases 
to 30—32% of the length of the head and in the 4th stage to 11%. This is not 
observed in larvae of other genera. 

In lst-stage larvae there is an egg tooth in the middle of the posterior 
margin of the frontoclypeus (Figure 14, A) which facilitates exit of the 
larva from the egg. 

The color of the head capsule varies from pale and semitransparent 
(C u 1 e x) to yellowish brown, dark brown (many A e d e s) and black ( C u 1 i - 
seta longiareolata, Uranotaenia). If the head is dark, it is more 
or less of uniform color, if it is light, dark spots which form a character- 
istic pattern are present on the frontoclypeus and lateral plates, especially 
posteriorly. These are the points of attachment of the muscles which move 
the mouth parts and antennae and the dilator muscles of the pharynx 



20 



(Figure 14). Places with the most intense mechanical action are more 
strongly sclerotized. Further strengthening of the head capsule is also 
attained by the fusion of spots with the formation of dark, curved, transverse 
stripes in the form of two arcs which strengthen the posterior part of the 
head. As the pattern of the head becomes darker during development, it is 
of no importance as a systematic character. 



Hairs of the head 



23 




The head bears about 20 pairs of hairs the number, position and develop- 
ment of which are relatively constant and characteristic for most species. 

We describe only the main hairs 
which are important for the deter- 
mination of the species. 

At the anterior margin of the 
clypeus is a pair of preclypeal setae 
(Figure 15) which are strong, curved 
inward, pointed and of varying length 
and thickness. Behind them are 2 
pairs of clypeal hairs, an inner and 
an outer pair, at the anterior margin 
of the frontoclypeus. These are of 
great importance for the determina- 
tion of species of Anopheles 
larvae and are not present in other 
genera except in T o x o r hy n c h i t e s. 
Their arrangement, development and 
branching (Figure 14) are important 
in Anopheles larvae, 
fr - frontoclypeus; c.s. and f.s. - coronal and Some distance behind the clypeal 

frontal sutures; 1 - setae of clypeus; 2 - postclypeal hairs are the postclypeal hairs . They 
hairs; 3,4, 5 - inner, median and outer frontal hairs; are more weakly developed in Ano- 
6,7 - coronal and transsutural hairs. pheles larvae than the clypeal hairs, 

and in most other species are very 
short and situated more posteriorly, usually between the base of the strongly 
developed frontal hairs (frontal or postantennal). There are three pairs, 
an outer, a median and an inner pair, and they form a curved row (Ano- 
pheles, C u 1 i s e t a, Mansonia and some species of Aedes and Culex) 
or the median pair may be situated anterior to the posterior pair forming 
2 triangles with their apex laterally. The median hairs are usually more 
widely separated than the posterior pair (Ochlerotatus, some species 
of Culex, etc.). The frontal hairs are feathered in Anopheles larvae; 
they are very short and simple only in A.plumbeus. In the larvae of 
Culiseta, Mansonia, many Aedes and Culex they are strongly 
developed and branch from the base in a fan, often with secondary feathering. 
In other species, they are simple or weakly branched, the outer hairs usually 
branched more densely than the others. 



FIGURE 15. Head of a larva of A e d es, dorsal: 



21 



Behind the frontal hairs are 3 pairs of short hairs: the outer, supra- 
orbital, behind the simple eyes, the median, transsutural, between the inner 
margin of the compound eyes and the frontal suture, and the inner, sutural, 
at the sides of the posterior part of the frontoclypeus. The hairs on the 
ventral side are not of systematic importance. 



Eyes 

Larvae of the 4th stage have two pairs of eyes, simple (ocelli) and com- 
pound eyes. The ocelli are small, rounded pigment spots at the sides of the 
head where it is widest. These eyes are fully developed and functional only 
in newly hatched larvae of the 1st stage; they are present in adult mos- 
quitoes but are covered with scales. 

The compound eyes are situated at the sides of the head anterior to the 
ocelli. They are absent in lst-stage larvae and develop during meta- 
morphosis. In larvae living in weakly illuminated water (tree holes), 
development and pigmentation of the compound eyes begin later, often in the 
4th stage. The eyes function in the larvae because the part of the integument 
covering the eyes is always unpigmented. 



(24) 




FIGURE 16. Different types of antennae of larvae of Culicinae: 
1— Culiseta morsitans; 2— Aedes; 3 — Culex pipiens. 



The antennae (Figure 16) are one-segmented and are little mobile. Their 
length is very variable, from /j— j% of the length of the head (some species 
of Finlaya and Stegomyia) to 1.5—2 times as long (Ma n s o n i a, Fig- 
ure 94, Aedes d i a n t a e u s, Figure 155). The simplest, short antennae are 



22 



rod-shaped, slightly tapering toward the apex, and bear the sense organs: 
hairs and setae. The antenna is smooth, without small spines, and a hair is 
always present near its middle. The antennae do not extend anteriorly 
beyond the hairs of the lateral lobes of the labrum. 
24 The longer antennae are slightly curved and more distinctly divided into 
a basal part and a thinner apical part. At the border between them is a 
hair with more or less long, numerous branches which form a fan, often 
with secondary feathering. The antenna is covered with sparse or dense 
spines, especially in the basal part. The terminal hairs and setae form 
two groups: a terminal group at the apex of the antenna and a subapical 
group at a varying distance from it (Figure 16). Both groups may be long. 
This structure is characteristic for the larvae of Culex, Mansonia and 
others. The antennae are pigmented either uniformly or at the base or also 
at the apex (Culex and others). 

Toxorhynchites larvae have three hairs on the antenna; they bear 
rounded sense organs resembling the annular organs of some larvae of 
Tendipedidae. 



Mouth parts and feeding of larvae 

The structure of the mouth parts of larvae is connected with their method 
of feeding and their biology. The larvae of Culicidae may be divided into 
two groups according to this character: phytophagous (omnivorous) and 
predatory. Feeding on plants is primary. 

There are 3 different types of feeding in phytophagous larvae, l) Feeding 
on the surface of the water, which is characteristic for larvae of Ano- 
pheles and some species of Uranotaenia. 2) Feeding on the substrate 
by sucking or scraping on encrustations and overgrowths on objects in the 
water, periphyton, which consists of microorganisms of animal or plant 
origin. This type is common in species of A e d e s, C ul i s e t a and others. 
25 3) Filtration-planktonic feeding on suspended particles in the water (bacteria, 
algae, Infusorian, etc.). This type is present mainly in larvae of Culex, 
Man s o n i a, some species of A e d e s and Culiseta. Some species are 
adapted to feed on filamentous algae (Culex bitaeniorhynchus, 
C. sinensis). 

There are all possible transitions between the types of feeding, and larvae 
of a certain species may use one or another, but one type of feeding usually 
predominates. Thus, Anopheles larvae which feed on the surface may 
also feed on the substrate, and cannibalism of some larvae has been ob- 
served (A. plumbeus). Larvae with the filtration type of feeding may 
also feed on the substrate and on the surface. This is observed in typical 
scrapers. 

Predatory feeding and cannibalism of larvae of Culicidae are secondary, 
and develop much later in species adapted to feeding on plants. In species 
of different subfamilies and genera the transition of species or groups to 
predatory feeding has taken place independently and at different times. The 
larvae of T o xo r hyn c h i t e s and Lu t z i a in Russia are predatory. 



23 



The type of feeding is governed not only by the structure and function of 
the mouth parts but also by the structure of the antennae, the frontal hairs 
and other hairs on the head, and even by the structure of the siphon. All 
these structures form a single, trophic, functional complex of correlated 
characters which adapt the larvae to one or other mode of feeding. Without 
giving a detailed description of the mouth parts (see Monchadskii, 1936, 
1951), we describe only their main modifications and their connection with 
the structure of other elements of the trophic functional complex. 




FIGURE 17. Labrum (after Wesenberg-Lund): 

1 — Culiseta morsitans; 2— Culiseta annulata. 

The mouth parts of the larvae consist of the following. 1. The labrum 
(Figure 17) consists of a median lobe and two more strongly developed 
lateral lobes with the mouth brushes, or flabella, which consist of numerous 
flexible, long hairs. This is the main organ which draws the food into the 

26 mouth and is used to catch the prey in predaceous larvae. 2. The epipharynx 
borders the prebuccal cavity above and is a continuation of the inner surface 
of the labrum. It bears symmetrical groups of anteriorly directed hairs, 
setae and spines which scrape food particles from the lobe of the labrum 

and prevent their return. 3. The mandibles (Figure 18) are strongly 
sclerotized, with dark teeth at the distal end of the inner margin and with 

27 spines and hairs. During movement of the mandibles, their terminal teeth do 
not touch and they have no masticatory function. During accumulation of 

the food bolus in the preoral cavity before the hypopharynx, the terminal 
teeth break up the bolus into small particles. The hypopharynx serves as 
an anvil. The hairs and spines function as in the epipharynx. The bases 
of the mandibles border the prebuccal cavity on both sides. 4. The maxillae 
(Figure 19) are of a simpler structure; in form, development of palps and 
hairs and in their mobility, they vary markedly in the different species. 
They border the posterolateral parts of the prebuccal cavity. 5. The labium 
is situated between the bases of the maxillae and border the prebuccal 
cavity ventrally with the hypopharynx. It consists of 3 overlapping 



24 




FIGURE 18. Mandible: 

1— Anopheles (after Puri); 2 — Culiseta annulata; 3 — Culiseta morsitans (after 
Wesenberg- Lund). 



(26) 




FIGURE 19. Maxilla: 

1 — Anophel es (after Puri); 2 — Culiseta morsitans; 3 
(after Wesenberg- Lund). 



Culiseta annulata 



25 



triangular sclerites directed anteriorly, the two inner plates, the mentum 
and submentum, with denticulate margins and a larger apical tooth. 6. The 
hypopharynx is situated below the mouth and is separated from the base of 
the labium by the opening of the salivary ducts. This is a markedly sclero- 
tized sclerite of complicated structure which is covered with tubercles and 
processes. The accumulation of food is performed by the grinding teeth 
of the mandibles and then the food enters the pharynx through the mouth. 



Metamorphosis of the trophic functional complex 

In larvae feeding on the surface (A no p h e 1 e s), the head is oblong be- 
cause of the longitudinal arrangement of the muscles of the labrum, the 
lateral lobes of which move anteroposteriorly during feeding. 

During feeding, the head of the larvae is rotated through 180°, 
supported under the surface of the water by 2 pairs of special forma- 
tions which are absent in larvae which feed differently: l) two flat- 
tened, knife-shaped setae at the end of the antenna and 2) the large palps of 
the maxillae which bear a branched lateral hair to increase the area of 
support and leaf- shaped, flattened sense organs at the apex. The hairs of 
the lateral lobes of the labrum are not differentiated; they consist of hairs 
of the same type and length. The mandibles and maxillae receive the food 
particles which are transported by the labrum, remove them from the hairs 
and move them to the posterior part of the prebuccal cavity, but their move- 
ment is restricted. The row of branched frontal hairs, supplemented by the 
dendriform outer clypeal hairs (if present) is important in directing the 
passage of the food particles. The frontal and clypeal hairs of larvae of 
A. plumbeus, which feed on the substrate, are simple and short. The 
absence of a siphon ensures close contact of the larva with the surface. In 
larvae of Uranotaenia which live and feed below the surface, the siphon 
forms an obtuse angle with longitudinal axis of the body. 

During feeding on a substrate which is not flat, the movement of the lobes 
28 of the labrum becomes more complicated and the mandibles and maxillae 
take on a more active role in the procurement of food. The muscles of the 
labrum are therefore diagonal and the musculature of the jaws becomes 
stronger. 

The lateral parts of the epicranial plates develop more strongly, es- 
pecially in the posterior part so that the head becomes wider than long. A 
differentiation of the structure of the hairs of the lateral lobes of the labrum 
takes place. The apex of the inner hairs becomes more flattened and bears 
10—25 denticles which form a comb with which the larvae scrape off deposits 
from the substrate. If this type of feeding predominates, the hairs with 
denticulate ends become more numerous and their development stronger, 
in extreme cases forming a long, inner brush. The median lobe is narrower, 
with dense spines in the form of a brush. The mandibles and maxillae 
bear rows of short, thick setae which not only remove the food from the lobe 
of the labrum but also scrape the substrate. The teeth of the mandibles are 
strongly developed but the maxillary palp is very small. The antennae are 
always rodlike and short, never reaching beyond the hairs of the labrum. 



26 



Longer antennae would interfere with feeding. The median and inner 
frontal hairs are relatively short, simple or slightly branched. The siphon 
is usually short, usually 4—5 times longer than its width at the base. This 
type of feeding is present in species of the subgenus Ochlerotatus and 
of the genus Culiseta, but their adaptation to this type of feeding varies 
and does not exclude planktonic feeding. 

The filtration -planktonic type of feeding is connected with a stronger 
development of the head in width, so that it becomes transverse-oval. The 
complicated bending and rotatory movement of the labrum and maxillae 
during feeding inside the water and the collecting of planktonic food in the 
prebuccal cavity cause a diagonal direction of the musculature at a large 
angle with the longitudinal axis of the body. The lateral lobes of the labrum 
are widely separated and consist of two distinctly different bundles: an 
inner, broad and short bundle, directed anteriorly and inward, and an outer, 
longer bundle, directed obliquely, laterally and anteriorly. The hairs of the 
inner bundle do not have serrated ends. The median lobe is broad, half- 
round, not narrow as in larvae feeding on the substrate or on the surface. 
The teeth of the mandibles are thinner and the hairs denser and longer. 
The maxillae are markedly changed: they are conical, slightly tapering 
toward the base and the palps are strongly reduced. At the apex of the 
maxillae is a bundle of long, straight or slightly S- curved hairs, and their 
inner side is densely covered with short hairs. The maxillae are very 
mobile and take an active part together with the labrum in the movement of 
the food (terminal bundle) and its filtration (bundle of inner-lateral hairs). 
The antennae are usually long or very long and curved; the apical setae are 
often situated far from the subapical setae and all are long. The bundle 
forms a large fan with secondarily feathered branches. The frontal hairs 
are long and usually branched like a fan. Together with the bundle on the 
antennae they restrict the passage of food pushed by the lobes of the labrum. 
The siphon is usually thin and long, which increases the radius of the food- 
collecting area. 
29 As a secondary adaptation, the predatory type of feeding develops in 
larvae adapted to feed on the substrate or plankton. In most predatory 
larvae the organ for catching food has become the labrum, which drives the 
food to the mouth in the phytophagous forms. In the predatory larvae of 
Toxorhynchites and Lutzia which occur in the U SSR, the modified 
lateral lobes of the labrum catch the food. They are displaced markedly 
laterally, the median lobe is reduced, and the lateral parts of the clypeus are 
enlarged and project anteriorly. The hairs of the lobes are reduced in 
number (to 10 in Toxorhynchites and to 30— 40 in Lu t z i a) and are 
transformed into hook- shaped setae. In Toxorhynchites they have 
coarse denticles at the end and the combs of chitinized denticles are much 
stronger in Lutzia than in larvae which feed on the substrate. The 
mandibles grasp the food already caught by the setae of the labrum and pass 
it into the pharynx. The hairs on them are partly reduced and the teeth 
more strongly developed. The hairs on the maxillae are transformed into 
shorter setae and they become smaller and rectangular. The antennae are 
short and the head of Toxorhynchites is rectangular with a large 
occipital foramen. 



27 



Thorax 

The three segments of the thorax (prothorax, mesothorax and metathorax) 
are fused so completely that their borders can be determined only according 
to the arrangement of the hairs in 3 successive stripes. The thorax is 
dorsoventrally compressed, the ventral side more strongly than the dorsal. 
Its form varies from rounded-hexagonal to rounded-oval, according to the 
stage and age of the larvae. In the older stages the developing appendages 
of the thorax of the adult insect, the 5 pairs of imaginal discs, are visible 
through the transparent cuticle, especially on the ventral side. Three pairs 
(imaginal discs of the legs) are situated on the ventral side and extend to 
the median line toward the thoracic part of the ventral nerve cord. On each 
side near the dorsal side, are the imaginal discs of the wings and behind 
them those of the halteres. Dorsally, on both sides, at the anterior margin, 
are the imaginal discs of the respiratory horns of the pupa which begin to 
develop at the end of the 4th stage. Before the molt, they come into contact 
with the respiratory system of the larva, are filled with air, become dark 
and refract light. This is the first indication that the larva is ready for 
pupation. 




FIGURE 20. Thoracic hairs of larva of A nophele s (after Puri): 

A — anterior margin of prothorax with notched shoulder organ; 
2,3 — middle and outer middle hair; 4—6 — lateral shoulder 
hairs. B — base of lateral hairs of prothorax: 9, 10 — ventral and 
dorsal hair of anterior pair; 11, 12 — dorsal and ventral hair of 
posterior pair. 



30 All larvae of Anopheles have notched shoulder organs (Figure 20) 
which are situated at the sides of the anterior margin on the dorsal side 
of the thorax. These are hollow, transparent processes directed obliquely 
anteriorly and laterally and consist of 2 lobes on a broad base. From this 
point extends a retractor muscle attached to the ventral side. Extension 
of the organ takes place by pressure of the hemolymph. The shoulder 
organs, which are absent in all other larvae, are closely connected with the 
life of Anopheles larvae on the surface. They support the anterior end 



28 



of the larvae on the surface film and give the body stability if the head is 
rotated. These organs bear no relation to the developing prothoracic 
spiracles or to the respiratory system. 




FIGURE 21. Thoracic hairs of a larva of Anopheles (after Puri): 

A - ventral side; 8-12 - lateral hairs; 13, 14 - other hairs. B - dorsal side - prothorax: 
1-3 - middle hairs (inner, middle and outer); 4-7 - shoulder hairs; mesothorax: 1 - inner 
middle hairs; metathorax: 1 — middle hair, often palmate. 



Many thoracic hairs are of importance for the determination of Ano- 
pheles larvae (Figure 21): middle hairs (l, 2, 3) on the prothorax, lateral 
hairs of all three stripes (9—12) and hair 1 on the metathorax, which is 
often palmate. In the larvae of Culicinae, the group of middle hairs of the 
prothorax may be important for determination, and in Toxorhynchites 
the lateral hairs of all three stripes. However, these characters are not 
generally used since there are more convenient characters on the last 
segments of the abdomen. 



Abdomen 

The abdomen consists of 9 segments. The primitive number is 10, but 
the 8th and 9th segments became fused and form the complex 8th segment 
(Figure 12). The last, 10th, segment thus becomes the secondary 9th seg- 
ment. This fusion was caused by the formation of a stigmal plate as 



29 



the only functional organ which includes elements of the 8th segment 
(spiracles, anterior and lateral lobes or valves) and the 9th segment (poste- 
rior lobes or valves and the central plate with its processes). The posterior 
part of the 9th segment may be preserved in most species as an intermediate 
ring between the 8th and the last segments, and this rudiment disappears in 
some species of Uranotaenia (Figure 65) and others. 

As the structure and development of the hairs of the first 7 segments is 
relatively similar, but the 2 last segments are distinctly different, their 
description is given below separately. 




-' .....y 



FIGURE 22. Sixth abdominal segment of an Anopheles larva (after Puri): 

A — dorsal side. B — ventral side; p.t. and a.t. — posterior and anterior tergal plate; 1—13 — 
hairs (1 — palmate and 6 — lateral). C — palmate hair, high magnification. 



Each of the first 7 segments bears 13 pairs of hairs in an irregular, 
zigzag row (Figure 22). The lateral (pleural) hairs are most strongly 
developed. 

The palmate hairs on the dorsal side situated near the posterior margin 
of the Anopheles larvae are the most important for determination. 
They vary in form and development on the anterior segments. They are 
often rudimentary on the first segment - short, usually simple hairs. They 
are 3— 4- branched only in A. lindesayi. They are very distinct in most 
species of Myzomyia. From the 2nd segment onward, they are more or 
less developed, from the 3rd to the 7th segment they are fully developed 
except in A . t u r k h u d i, in which they are present only on the 4th and 5th 
segments. 

The palmate hairs (Figure 22) consist of a short stem from which extend 
leaf-shaped, flattened branches parallel to the surface of the body. The 
form of the branches, their pigmentation, denticulation of the margins, etc. 
vary in the different species. These hairs, together with the shoulder 
organs and the stigmal plate, form the support of the larva in the sus- 
pended state from the surface film of the water. If the hairs are open they 
lie on the surface. If the larva is submerged, the leaves become folded 
32 together and hold a bubble of air with the aid of which they break the surface 
film at the next rise to the surface. 



30 



The thorax with the shoulder organs and the stigmal plate are slightly 
raised above the surface of the 7th segment, and when the shoulder organs 
and stigmal plate touch the water surface, contact with the surface film of 
the palmate hairs near the thorax (first and second) and of the stigmal plate 
of the 7th and 8th segments is less likely than of the hairs on segments 
3—6. The fact that palmate hairs are developed in all Anopheles larvae 
on the 4th and 6th segments, exceptionally on the 1st segment, and are absent 
on the 7th segment is due to the great variation of their development on 
segments 1 — 3 of the abdomen. They are therefore only fully developed on 
segments 4—6. 

The lateral (pleural) hairs 6 (Figure 22) are also important for the 
determination of Anopheles larvae. Their important characters are 
their branching and relative development. 




FIGURE 23. Posterior end of larvae of Culicinae (A), lateral: 

VII, VIII - abdominal segments; 6-13 - hairs behind the comb (6,13 - extreme upper 
and lower, 9 - median, 7, 11 - intermediate); B - scale of comb; C - tooth of pecten; 
D - siphon of larva of C u 1 i s e t a , lateral. The arrows indicate the length of the 
siphon and its width at the base. 



The hairs on the first 7 segments of the abdomen of larvae of Culicinae 
are not used in systematics, but the hairs and other formations on the last 
2 segments are very important. The 8th segment of larvae of Culicini and 
Toxorhynchitini (Figure 23) bears a siphon in its posterior part. Before 
the siphon, near the posterior margin of the segment, is situated a comb 
which consists of a varying number of scales directed posteriorly. Their 
number varies from 5-7 to many tens. Each scale extends posteriorly and 



31 



often ends in a pointed spine. At its base are nearly always denticles on 

33 both sides which decrease in size toward the base. These lateral denticles 
are sometimes only a little smaller than the main spine. In many larvae 
(Culiseta, Culex, some species of Aedes), the main spine is absent 
and the scale or only its free end has a row of denticles. The structure, 
number and position of the scales vary in the different species, but they are 
relatively constant in the same stage and species. If there are few scales 
of the comb, they may form a straight, curved or zigzag row; if there are 
many, they form a group. The structure of the marginal scales (upper, 
lower or posterior) often differs from that of the scales in the middle. The 
scales of Uranotaenia are situated on the posterior margin of a sclerite . 
In Toxorhynchites, there are instead of scales 1 — 3 strong spines which 
extend from the sides of the sclerite. The elongate scales of Ortho- 
podomyia form 2 regular rows and are of complicated structure. 

The comb is surrounded by 5 hairs posteriorly and dorsally. The dorsal 
and ventral and especially the middle hairs are the largest and branched 
like a fan; the two intermediate hairs are usually simple and shorter. 

The siphon is of great importance in the systematics of the larvae. It 
is usually sclerotized to the base from the 3rd stage, and often darkly pig- 
mented. Only its distal end is sclerotized in the 1st stage. The scleroti- 
zation extends to near the base in the 2nd stage. The siphon is rarely 
cylindrical but usually tapers toward the apex; it is rarely slightly widened 
at the apex (Culex territans). It is widest in the middle in some 
species. In cross section the apex is round or longitudinally oval. The 
base is usually darkly pigmented and slightly thickened anteriorly and on 
both sides, and there is a more or less deep incision posteriorly. There 
are 2 small processes, auricles, near the posterior margin, which are 
points of attachment of a muscle extending at a right angle to the base, the 
contraction of which curves the siphon posteriorly. The incision at the base 
facilitates this movement which is important in detaching the larva from 
the surface of the water. Auricles are not present in larvae of Stegomyia 
and Toxorhynchites and others. 

One of the important characters for determination (Culex, some species 
of A e d e s ) is the siphonal index: the ratio between the length of the siphon 
and its width at the base. Measurement may give different values which 
are often not comparable in different species and with the data of various 
authors. The same method of measurement should therefore be used: 
1) measurement of the siphon along the side; 2) measurement of the length 
without the valves of the stigmal plate, from the base of the auricle of the 
basal ring to the posterior margin of the apex at the base of the posterior 
valves of the stigmal plate; 3) width of the base should be measured along 
the line extending from its anterior margin through the base of the auricles 
to the point where it crosses the posterior margin of the base or its con- 
tinuation if the incision of the posterior margin is distinct (Figure 23, D). 
In some cases (Culiseta), the ratio of the length of the siphon to the width 
of the apex is important. 

The structure of the pecten and the position, development and number of 
setae on the siphon are of systematic importance. 

34 Of the larvae of mosquitoes occurring in Russia, the pecten on the siphon 
is absent only in Orthopodomyia and Toxorhynchites; in all 



32 



others it is present as two symmetrical, longitudinal rows of denticles, 
spines or hairs at the sides of the posterior side of the siphon, beginning at 
the base. The denticles of the pecten are not of the same form; those near 
the base are small; farther away from the base the denticles become 
larger. Each denticle is like a pointed spine, usually with small, accessory 
denticles at the base; the distal denticles are the largest. The denticles 
are more or less regularly spaced or the distal denticles are more widely 
spaced. The pecten may extend beyond the middle of the siphon. Some- 
times (Culiseta 1 o n g i a r e o 1 at a) the pecten consists of a row of 
denticles which are widely separated along the entire siphon. In some 
larvae of the subgenus Cu 1 i c e 1 1 a, the short pecten continues in a row of 
large spines, and in the subgenus Culiseta there are denticles only near 
the base, which then become longer and form long, thin hairs. 




FIGURE 24. Posterior end of the abdomen of a larva of A nopheles, 
lateral (after Martini): 

0-13 - hairs. 



In most larvae, the siphon has only one pair of tufts of hairs in addition 
to a pair of short apical setae on the anterior side of the apex (A e d e s, 
Culiseta). The tufts are situated near the middle (A e d e s and others) 
or at the base of the siphon (Culiseta, Toxorhynchites). There are 
sometimes also 2—4 pairs of hairs on the anterior side of the siphon or at 
its apex (group rusticus of Aedes, A. cinereus). The siphon of 
larvae of Culex bears 4—8 or more pairs of tufts of hairs arranged sym- 
metrically or displaced to the posterior surface. Of systematic importance 
are the number, position and the branching of the tufts on the siphon and 
also their length, the width of the siphon at the position of the hairs (some 
species of Culex and Aedes). 

The hairs on the 8th segment of larvae of Anopheles resemble those 
of the Culicinae in number and position but are of no systematic importance. 
In lst-stage Anopheles larvae, as in the Culicinae, there is a tuft at the 
sides of the 8th segment which consists of 5—6 oblong scales. This is no 
longer present in the 2nd stage. 



33 



35 




FIGURE 25. Stigmal plate of an Anopheles 
larva (diagrammatic): 

p.l. — posterior lobe; 1.1. — lateral lobe; 
a.l.— anterior lobe; c.p.— central plate; 
p.p.— posterior part; m.p.— median part; 
a. p.— anterior part; sp — spiracles; p.s.— 
processes of spiracles; u.s.— upper surface 
of posterior lobe; l.s.— lower surface (base) 
of posterior lobe; a, b, c, d, f, 1—5 — hairs 
of stigmal plate. 



The absence of a siphon is character- 
istic for the larvae of Anopheles. 
The larvae have instead a special 
formation which supports the stigmal 
plate and is homologous to the siphon. 

This formation consists of a dark, 
chitinized arc which surrounds the 
base of the posterior lobes posteriorly 
and has 2 lateral processes (Figure 24). 
The ends of the arc pass on both sides 
into the triangular plate, the posterior 
margin of which bears a row of large 
spines. The dorsal spines of this row 
correspond to the pecten of the siphon 
of Culicinae and the ventral row to the 
comb at the sides of the 8th segment 
of the abdomen. The development, 
structure and number of spines in the 
comb vary in different species and also 
in the different stages. The lateral 
chitinized plates are not developed in 
lst-stage larvae. There is only a thin, 
chitinized arc and separate groups of 
scales corresponding to the pecten and 
comb. 

The structure of the stigmal plate 
is often of great systematic importance 
and gives distinct characters for the 
determination of Anopheles larvae. 

In the Anopheles larva (Fig- 
ure 25), the spiracles are surrounded 
by 5 lobes: a single anterior lobe and 



a pair of lateral and a pair of posterior 
lobes. In the middle of the base of the anterior lobe there is an oval, 
transparent formation. Lateral to its base are 3 pairs of short hairs. Its 
upper surface may be dark (larvae of the 2nd stage of A. claviger, 
A. plumbeus, A. superpictus, and the last 3 stages of A. pulcher- 
rimus) or the dark pigment is restricted -to the base and the anterior part 
is lighter (A. maculipennis, A. hyrcanus). 

On the dorsal surface of the lateral lobes is an elongate, dark, chitinized 
supporting plate with a sensory hair at the apex, its narrow base directed 
toward the spiracles. 

The apical surfaces of the posterior lobes are fused with the central 
plate. The latter consists of 3 parts: an anterior part, tapering into the 
anterior process between the spiracles, the base of which may continue 
posteriorly to the anterior or the median part; a median part, and a broader 
posterior part. The sculpture of the central plate is different in the lateral 
and median parts. The last is the point of attachment of the dorsoventral 
muscles, which causes its reticulate structure, the position of the cells and 
their pigmentation. From the anterior process extend anteriorly 



34 



36 longitudinal dorsal muscles of the 8th segment. This is the appearance of 
the open stigmal plate if the larvae are on the surface. 

After submersion, the anterior lobe turns posteriorly, covering the 
spiracles and the central plate to the median part. The spiracles are dis- 
placed anteriorly toward the base of the anterior lobe. The lateral lobes, 
the bases of which are connected with the spiracles but have no musculature, 
turn posteriorly and end at the ridge of the anterior margin. At the same 
time, the central plate, through the combined contraction of longitudinal 
muscles which extend from the anterior process and dorsoventral muscles 
extending from its median part, moves anteriorly and downward to deepen 
the cavity which opens dor sally in the posterior part. This cavity becomes 
filled with air during submersion. During the ascent this air pierces the 
surface film and the spiracles come into contact with the air. 




FIGURE 26. Structure of stigmal plate (diagrammatic): 

A - stigmal plate of Culiciae larva: p. v. - posterior valve; l.v. - lateral valve; a. v. - 
anterior valve; st - "stirrup"; a.ap. - anterior appendage of "stirrup"; p.ap. - posterior append- 
age of "stirrup"; p.p. - posterior process; sp - spiracle; s - shoulder; p.s. - process of spiracle; 
a.c.f, 1-5 - hairs. B - sagittal section through end of siphon of a Culiseta larva with open 
stigmal plate. 

In all lst-stage larvae of Anopheles (Figure 14), the structure of the 
stigmal plate is the same and strongly reduced. In 2nd stage larvae, all its 
elements are already present but the specific characters are still indistinct. 
Their development becomes distinct only in the 3rd stage and inlarvae of 
A. hyrcanus and A. maculipennis this takes place only in the 4th stage . 

The presence of a siphon in Culicine larvae restricts the movement of 
the muscles to a direction parallel to the longitudinal axis of the siphon so 
that the chitinized elements of the stigmal plate become more differentiated 
than in Anopheles. 



35 



On the stigmal plate of Culicinae (Figure 26) the spiracles are surrounded 
by 5 valves homologous to the lobes of the plate in Anopheles: a single 
anterior lobe, a pair of lateral and pair of posterior lobes. Between the 
37 posterior valves and the spiracles is a plate, the "stirrup" which is homolo- 
gous to the central plate in Anopheles; is connected with most of the 
elements of the plate by processes and coordinates their movement. 

The "stirrup" consists of a stem in the lumen of the siphon between the 
tracheal trunks. It is distinctly visible in lateral view. At the apex of the 
siphon, the stem widens and passes into the laterally compressed "stirrup" 
with a cavity which is open above. This cavity is formed by the develop- 
ment of the "stirrup" of the next larval stage which surrounds the old skin. 
The cavity of the "stirrup" is therefore absent in the lst-stage larvae. 
The "stirrup" ends in a longitudinal, saddle-shaped widening and its ends 
form the anterior and posterior appendage. There is often an anterior 
process on the anterior appendage which is connected with the anterior 
valve. The posterior appendage continues into the posterior process the 
form of which is characteristic in many genera and species. From the 
sides of the saddle the processes extend to the spiracles and often surround 
their posterolateral margins. The spiracles are oval, rarely rounded, and 
are situated at the sides of the "stirrup." Their outer margins are situated 
in the plane of the plate and the inner margins follow the curve of the saddle. 

During submersion, the "stirrup" is retracted ventrally because of the 
contraction of the retractor muscles which are attached to the stem. All 
parts of the stigmal plate are moved simultaneously and the spiracles face 
each other. All valves turn inward, come together above the spiracles and 
form a hollow, irregular pyramid. Air is kept in the cavity. The closing 
of the valves is usually not complete since the surface of the stigmal plate 
is not wettable, and water cannot enter the cavity to press the air out. 

During the ascent, when the upward-directed hairs of the posterior and 
lateral valves touch the surface film, the retractors relax and their 
antagonists, a pair of levators of the "stirrup" which extend from it to the 
upper margin of the siphon, are contracted. The plate opens and the air in 
the cavity enters above the half-open valves and pierces the surface film, 
and contact is then established between the tracheal and the atmospheric 
air and the valves are situated on the surface. 

Some characters of the tracheal system are of systematic importance. 
Two main tracheal trunks extend from the posterior spiracles to the pro- 
thorax. In most larvae, the main tracheal trunks, especially in the siphon 
and in the posterior segments, are oblong in cross section and appear 
ribbonlike in lateral view. In some larvae of Culex (especially in the 
subgenus Neoculex), Culicella and others they are thin and rounded 
in cross section. In larvae of Toxorhynchites and Orthopodomyia, 
the main tracheal trunks form a pair of widenings in the metathorax and 
partly in the first abdominal segment. In larvae of Mansonia, similar 
widenings are present on the branches of the main tracheal trunks. All 
these widenings have a hydrostatic function. 

The last abdominal segment is narrower than the others. It forms an 
angle to the ventral side and bears a chitinized plate, the saddle, dorsally 
and laterally. The saddle, especially in young larvae, covers only the dorsal 
part of the segment or, usually, extends to the sides. It sometimes extends 
to the ventral side, surrounding the segment like a ring with a posterior 



36 



38 incision around the base of the fin. If this incision is absent, part of the 
anterior fascicles of the fin seem to be situated inside the ring which, like 
the saddle, has a supporting function. The posterior margin of the saddle 
bears denticles and spines of varying form which are especially strongly 
developed in Toxorhynchites. 

The saddle always bears a moderately long or small lateral hair. On 
the dorsal side, in the middle of the posterior end of the last segment, behind 
the saddle, there are two pairs of hairs, the caudal hairs (Figure 23). The 
outer pair is situated on small sclerites and consists of longer and less 
branched hairs. The inner caudal hairs which are situated on the common 
sclerite are shorter, more strongly branched and often form asymmetrical 
fans. In Anopheles larvae (Figure 12), the hairs of the outer pair have 
curved, hooked ends which fix the larvae to objects floating on the water. 
Without them, the movements of the labrum would cause forward movement 
of the larvae. 

Two pairs of gills surround the anus. Their development, form and 
color are often used in determination. In spite of their name, the function 
of the gills in the gas exchange is not greater than that of any other part of 
the integument. Their main function is osmoregulation. The size of the 
gills varies in different species from a third of the length of the body 
(Aedes pulchritaris, A. ga 1 1 o i s i) to a small sphere. In some 
species the gills are longer when the concentration of salt in the water is 
low and they are smaller if it is higher. The gills may be sausage- shaped, 
with rounded or pointed ends, or lanceolate, leaf- shaped or spherical. 

The gills are thin- walled, saclike processes with a thin cuticle, which is 
slightly permeable to water and substances dissolved in it. Its hypodermal 
cells are often strongly vacuolized and may contain pigmented inclusions. 
The cavity of the gills communicates with the body cavity and is filled with 
hemolymph, the movement of which is directed by a membrane which divides 
the cavity longitudinally but does not reach its end. The trachea passes 
through the membrane with a few branches toward the hypodermal cells. 
The gills of the larvae of Culicidae are therefore blood gills and not tracheal 
gills. 

On the ventral side of the last segment is the fin or ventral brush, the 
main organ of movement which consists of a row of long, curved hairs 
situated in a zigzag line. They form a flat fan which is asymmetrically 
curved posteriorly. Each hair therefore usually branches asymmetrically 
posteriorly from the main stem. The fin is symmetrical only in larvae of 
Anopheles and Toxorhynchites and the hairs are two-branched. 

During movement, the main pressure on the fin is directed laterally and 
the base of each hair is therefore strenghened by transverse lateral pro- 
cesses. Because of the zigzag arrangement of the hairs, the lateral pro- 
cesses are asymmetrical: if the base of the hair is situated on the right of 
the median line, the left lateral process is longer and the right shorter, and 

39 vice versa. The ends of the processes are curved, fused, and form the 
common base of the fin. The hairs of the fin become shorter anteriorly 
and the lateral processes at the base weaker, but their ends are free. In 
some forms (many species of Aedes) there is a row of 2—10 much shorter, 
more weakly branched hairs, extending anteriorly from the base of the fin. 

The fin is absent in the 1st- stage larvae of most species. In many 
species there is a reduction of the number and branches of the hairs of the 



37 



fin and a reduction of the common base and the processes. This reduction 
of the fin is connected with the strong development of the gills (e. g. in 
Aedes pulchritarsis, A. galloisi, A. aegypti). The caudal hairs 
of the larvae curve down during movement into the space between the upper 
pair of gills and the hairs of the fin, which are curved dorsally, are situated 
between the lower pair of gills, and have a supporting function. The gills 
then become organs of movement. 



Growth of larvae, changes of characters during growth, 
determination of the stage and age in the stage 

The larvae molt 3 times, i.e. there are 4 stages. Their measurements 
increase by more than 8 times during development. All sclerotized forma- 
tions (head, siphon, saddle, etc.) attain their maximum size during the molt 
and their minimum growth during the stage, while the other part of the body 
with soft cuticle, grows mainly during the stage and minimally during the 
molt. 

The head, siphon and other hard parts are transparent and soft after the 
molt. They harden after a few hours and lose their transparency because 
of deposits of organic and inorganic substances. Their size increases 
slightly due to the stretching of the cuticle. Exceptions are the larvae of 
Anopheles (sometimes the siphon of Culicinae ), in which there is an 
increase of the size of the "collar" (p. 20 and Figure 27) during the stage. 
The other part of the body covered with soft cuticle increases 90% during 
the stage and only 10% during the molt. There is always a distinct increase 
in the size of the larvae in the first 6—16 hours after the molt because of 
the stretching of the cuticle; the increase then becomes smaller and growth 
proceeds uniformly to the next molt only from the beginning of the 2nd day. 

There are also morphological changes which are restricted to the molts, 
so that the larval stage or the age within the stage can be determined more 
or less accurately. 

The newly molted larvae, in every stage, differ from older larvae not 
only in the pale pigmentation of the head, siphon and other sclerotized forma- 
tions, but also in their relatively large head compared with the thorax. The 
hairs of the body, especially the lateral hairs, are not straight but have 
curved ends. Larvae approaching the molt or pupation are always dark, 
their head is always relatively small compared with the thorax, and the 
40 hairs, especially the lateral hairs, have straight ends. The hairs of the 
next stage, especially the long, lateral hairs of the segments, the hairs of 
the fin and caudal hairs which surround the segment under the old cuticle 
are visible through the cuticle. In larvae ready for pupation, the pupal 
respiratory horns, which are dark from the air filling them, are visible at 
the lateral angles of the prothorax. 

With each stage, the branching of the hairs increases. This also takes 
place with the number of scales of the comb and with the teeth of the pecten, 
in which the number of accessory denticles at the base increases. If the 
scales of the comb are few (Aedes riparius, A. pulchritarsis), 
their number does not increase with the 2nd stage. The species can be 



38 



determined at any stage by the quantitative changes of all characters, but 
these characters are unreliable and therefore of no use in determination. 




FIGURE 27. Outline of head and thorax of all stages of larvae of Anopheles 
maculipennis Mg. after the molt (1) and before the molt to the next stage (2): 

I, II, III and IV - larval stages (stages III and IV are shown 4 times smaller than 
stages I and II). 



41 There are a few constant characters in some larval stages. Character- 

istic for all larvae of the 1st stage is the egg tooth on the frontoclypeus 
(Figure 14) and the absence of a fin. All hairs of the body are simple except 
in a few cases. Larvae of the 4th stage show the developing pupal horns 
which are in contact with the tracheal system before pupation and filled 
with air. Characteristic for all larvae of the 2nd and 3rd stage of the sub- 
family Culicinae is the vesicular thickening of the posterior or postero- 
lateral margin of the spiracles which is distinct on the open plate and less 
so on the closed plate. 

The following method of determining the larval stage seems most con- 
venient. The larvae are placed in a white enamel bowl and around it four 



39 



small dishes. The largest larvae with the widest head are placed in one 
dish. The smallest larvae with the narrowest head are placed in another 
dish. The larvae with the widest and narrowest heads belong to the 4th and 
1st stage. Of the other larvae, smaller specimens of the 4th stage and 
larger specimens with the same head are collected; the same is done for 
the lst-stage larvae. The other larvae are divided into two groups, ir- 
respective of size. One group with a head wider than in the 1st stage and 
the other with a head narrower than in the 4th stage is placed in the middle 
dishes. Mistakes made will be recognized since larvae placed in the wrong 
dish will differ from the others in the width of the head. This method makes 
it possible to determine the larval stage with certainty. 



PUPA 

Larvae of the 4th stage pass into the pupal stage; processes of histolysis 
and histogenesis take place and the fully formed, adult mosquito develops in 
the pupa. The pupae do not feed and are connected with the outer environ- 
ment only by the gas exchange. They are usually suspended from the sur- 
face film and their horns are in contact with the atmospheric air. They 
are capable of rapid movement and descend to the bottom if disturbed. If 
they are not disturbed, they may remain on the surface for a long time. 
The duration of the pupal stage usually lasts only 2—2.5 days. The males 
hatch first. 
42 The pupae have the form of a comma in lateral view; they are irregularly 

oval seen from above. The body consists of two parts: a large cephalo- 
thorax and a narrower, dorsoventrally flattened abdomen. The dorsal and 
anterior parts of the cephalothorax are formed by the thorax, the ventral 
part by the head. The developing appendages of head and thorax are 
situated in the ventral and posterior parts of the cephalothorax. The pig- 
mented eyes are visible through the transparent cuticle. Above them, at the 
sides of the dorsum are the pupal horns which are wider apically, wider in 
the Anophelinae than in the Culicinae. If the pupae are at the surface, they 
lie in the plane of the surface. From the base of the horns extend the 
tracheae, which connect them with the prothoracic spiracles of the develop- 
ing insect. Direct contact of the pupal horns with the tracheae takes place 
in 4th- stage larvae a few hours before pupation and the horns become filled 
with air. In pupae of M a n s o n i a, in connection with their method of 
breathing, the horns are pointed and serrated at the end. 

The abdomen of the pupae consists of 9 segments, of which the last seg- 
ment, morphologically the 8th and 10th, consists of the fused 8th and 9th 
segments. On the posterior margin of the dorsal side of the 8th segment 
are two paddles, which are the organs of movement of the pupae. They 
are irregularly oval plates with a narrow base. The plates are attached by 
a longitudinal median ridge which does not reach the free margin which 
often bears a row of denticles or small spines. At the end of the ridge is a 
hair slightly inward from its base and a shorter hair. The paired primordia 
of the paddles of the pupa are situated in the larva in the posterior part of 
the siphon (Culicinae) or between the formations which support the stigmal 
plate which are homologous to the siphon in the Anophelinae. 



40 



The last segment is a narrow ring; from its ventral part extends a pair 
of terminal lobes between the bases of the paddles. These are the genital 
appendages, which are short in females and much longer in males. The 
sex of the pupa can be determined by their size. 

On the dorsal side of the first segment is a pair of palmate hairs re- 
sembling those on the dorsal side of the 1st and 7th segments of Ano- 
pheles larvae and which have the same function. They are absent in Ma n - 
sonia. The relative development and branching of the hairs of segments 
3—8 are important for the determination of pupae. Segments 3—7 have 
8 pairs of hairs on the dorsal side, segment 8,3 pairs of hairs. 

Since the determination of species is usually made with larvae and adult 
mosquitoes, we are not describing the generic and specific differences of 
the pupae. 



GEOGRAPHICAL DISTRIBUTION 

Paleontological data on the family Culicidae are so few and fragmentary 
that they do not help to determine their recent geographical distribution. 
43 Their geological age is not exactly known, but primitive characters of the 
Culicidae, both of larvae and adults, suggest a great antiquity. Blood- 
sucking mosquitoes appeared hardly later than the Jurassic (Edwards, 
1923). The essential absence of eurythermal vertebrates at that time did 
not exclude the possible evolution of the group. There are even now some 
species which prefer to feed on reptiles and amphibians, e. g. Culex 
territans. 

Reliable paleontological data on the Culicidae begin with the Eocene 
(particularly in North America), from which 2 species of the genus Culex 
have been described. Mosquitoes from the Oligocene are more numerous 
but bloodsucking forms have not been found in the Lower Oligocene amber 
in the Baltic area. The amber contains only species of related families 
which are not bloodsucking, mainly Dixidae and Chaoboridae. About ten 
species of mosquitoes of the recent genera Culex, Mansonia, Aedes 
and Anopheles are known from the Oligocene. 

The recent fauna of bloodsucking mosquitoes (Culicidae) is not very 
large. There are about 2,500 species of 30 genera (Stone, Knight and 
Starcke, 1959). This figure does not include all the forms, but in view of 
the intensive study throughout the world in recent years, it will probably 
not increase much. 

The mosquitoes are mainly a tropical group. Not only are there many 
more genera and species in the tropics but there is also a marked, ancient 
differentiation of the fauna which is reflected in the presence of endemic 
groups in each zoogeographical region in the tropics. 

The fauna of mosquitoes is especially well defined in the Neotropical 
region, which contains characteristic endemic genera and subgenera, some 
of them with numerous species. These are mainly genera of the tribe 
Sabethini, which is predominant in the Neotropical region: Trichopro- 
sopon (32 species), Phoniomyia (22 species), Sabethes (21 species), 
Limatus (8 species). The large genus Wyeomyia (about 90 species) 



41 



of this tribe is restricted mainly to this region. Very few species of this 
genus enter the southern parts of the Nearctic region. Endemic to the 
Neotropical region are also some groups of the subfamily Anophelinae, e.g. 
the genus Chagasia and the subgenera Stethomyia, Kerteszia, and 
Lophopodomyia of the genus Anopheles. Not strictly endemic but 
characteristic for the Neotropical region are the genera Psorophora 
and Deinocerites and the subgenera Melanoconion (Culex) and 
Ny s s o r h y n c hu s (Anopheles). 

The fauna of mosquitoes of the Ethiopian and Oriental regions is also 
very rich in species but their differentiation is not as marked as in the 
Neotropical region. There is only one endemic genus, Eretmapodites, 
with 23 species in the Ethiopian region and the subgenera Myzomyia 
(Anopheles), Stegomyia and Aedimorphus (Aedes) and the 
genera Ficalbia, Mansonia and Uranotaenia. 

Endemic in the Oriental region, in addition to some small groups, are the 
44 genera Topomyia (21 species) Heizmannia (18 species) and especially 
Armigeres (41 species). One species of the last two genera occurs in 
the Palaearctic region. The subgenus My z o m y i a, the genus Toxo- 
rhynchites and the subgenera Aedes, Finlaya and Stegomyia of 
the genus Aedes are well represented in the Oriental region. 

The Australian fauna of Culicidae is small and there are few endemic 
forms. The genus Opifex with one species and some of the small and 
little differentiated subgenera of the genus Aedes may be considered as 
endemic. 

In the Holarctic region (Nearctic and Palaearctic), the fauna of mosquitoes 
is also small compared with that in the tropics (in numbers of species). It 
is easier to characterize them by the absence of some groups than by their 
presence. Particularly characteristic for this region are the large number 
of species of the subgenus Ochlerotatus (genus Aedes) and the distinct 
role of the genera Culiseta and Anopheles. The most important 
negative characteristics are the small numbers of the subgenus Myzomyia 
(absence in the Nearctic ), the small role of the genera Toxorhynchites, 
Tripteroides, Uranotaenia and Armigeres and of the subgenera 
Aedes and Aedimorphus, the relatively few species of the genus 
Culex and the absence of many of the tropical groups of mosquitoes. 

Characteristic for the Holarctic is the wide distribution of many species, 
which often has a zonal character. 

The northern boreal region includes the Canadian region and the tundra 
and forest areas of the Palaearctic (except for the broadleaved forests of 
the Far East). The species of the subgenus Ochlerotatus (genus 
Aedes) predominate in number and "biomass" here. Also present are 
some species of Anopheles and Culiseta, several subgenera of Aedes 
and a few species of Culex, but the genera Toxorhynchites, Tripter- 
oides, Uranotaenia, Orthopodomyia and Armigeres and the 
subgenera Myzomyia and Stegomyia are absent and species of the 
genus Culex are few. 

The Arctic subregion is characterized by a few species of the genera 
Aedes (subgenus Ochlerotatus) and Culiseta and the absence of 
all other groups. 

The Mediterranean subregion, which includes the Mediterranean, Asia 
Minor and adjacent countries, the southern shore of the Crimea, 



42 



Transcaucasia and a large part of Middle Asia, is characterized by the 
presence of the group Myzomyia, which plays an important role in the 
southern parts, and the genera Uranotaenia, Orthopodomyia and 
C ul e x, the last of which in number of species surpasses Ochlerotatus, 
which is represented only by very few species. 

The Palaearchearctic subregion (Manchuria) includes the eastern and 
northeastern parts of China, Korea, Japan, the Maritime Territory and the 
southern Amur area. Characteristic for these areas are Toxorhyn- 
c h i t e s, numerous species of F inlay a and Stegomyia and Oriental 
species of Culex. 
45 The mosquitoes of Mongolia and the mountains and highland deserts of 

Central Asia are little known. The number of species of Culicidae is pos- 
sibly much greater than is known at present. 

The fauna of mosquitoes in the southern parts of the Nearctic (Sonorian 
region) is very characteristic and contains neotropical elements. 

The absence of endemic subgenera and genera in the Holarctic and part 
of the Palaearctic suggests that the fauna is very young. It is debatable 
whether under the same conditions the evolution of mosquitoes may be 
slightly retarded compared with organisms exposed to more varied environ- 
mental factors. 

The development of the recent fauna of Culicidae in the Palaearctic did 
not take place simultaneously in the different parts nor in one center. A 
study of the world fauna of Culicidae suggests the following scheme of its 
formation. Southern elements spread to the north in the preglacial period. 
Ethiopian elements spread into the western part of the Palaearctic and 
Oriental elements into the eastern part. 

The Mediterranean fauna of mosquitoes developed under the specific, 
and apparently continuing, influence of the Ethiopian fauna. This is also 
reflected in the specific composition of mosquitoes in the USSR, where 
numerous Mediterranean species (23) are represented. Some of them are 
also distributed in the Ethiopian region. Mediterranean species are well 
represented in the southern Ukraine, the Caucasus and in Middle Asia, some 
of them spreading far to the north. Anopheles algeriensis occurs 
in Estonia (Remm, 1957), and A e d e s geniculatus in Chuvash and Tatary 
("Volkova, 1956). The spread of some Ethiopian species far to the north and 
northeast, e.g. Culex theileri, to the Kharkov Region and Culiseta 
longiareolata to the southern regions of West Siberia, is remarkable. 

A similar, but less distinct phenomenon appears in the eastern Palae- 
arctic, in the Palaearchearctic subregion (Manchuria), where the fauna is 
mainly Oriental. In the extreme Southeast of the USSR, 18 species of 
mosquitoes have been found which are characteristic for the Manchurian 
subregion (including Oriental species). The distribution of this group is 
mainly restricted to the Maritime Territory and the southern Khabarovsk 
Territory, some of them only in the far south of the Maritime Territory. 
These include Toxorhynchites christophi, Aedes nipponicus, 
A. flavopictus, A. nobukonis, A. aureus and Culex hayashii. 
The occurrence of a large group of Oriental species of the genus Culex, 
including C. vorax, C. fuscanus, C. tritaeniorhynchus, 
C. bitaeniorhynchus, C. sinensis, C.whitmorei and C . jack- 
soni is noteworthy. 



43 



The fauna of the boreal and arctic zones, with its numerous species of 
the subgenus Ochlerotatus, is not connected with the tropical fauna of 
46 the eastern hemisphere. The boreal and arctic fauna of mosquitoes of the 
PalaearCtic is closely connected with the Nearctic, where groups character- 
istic for the boreal zone are numerous. For example, 23 species of 
Culicidae occur in both the Palaearctic and Nearctic regions. All of them 
occur in the USSR. The origin of the boreal and arctic fauna of mosquitoes 
of the USSR is connected with the Nearctic. 

It is not known how the boreal mosquitoes entered the Palaearctic from 
the Nearctic. The immigration apparently took place from the West and the 
East. Until the separation of America from Europe the western route was 
probably used by some forms of the more southern latitudes of the western 
Palaearctic. An example is the group of species of the genus Ortho- 
podomyia, which are related to O. pulchripalpis. This group is re- 
presented in the Palaearctic only by O. pulchripalpis, which is dis- 
tributed from the Mediterranean to southern England and Azerbaidzhan in 
the east. Two species related to this group are common in the eastern 
states of North America. The well defined group rusticus of the sub- 
genus Ochlerotatus, which is distributed over a large part of the Medi- 
terranean and the Volga area, West Siberia and the Baikal area, also has a 
representative in North America, A. trichurus Dyar . 

However, there must also have been an eastern route for the exchange of 
fauna between the Nearctic and Palaearctic. This is supported by the 
common distribution of some species in both regions in the western part of 
the Nearctic and the close resemblance of the specific composition of 
mosquitoes in Northeast Asia and Northwest America. Of the 27 species 
of mosquitoes found in Alaska (Gjullin, Sailer, Stone and Travis, 1961), 17 
also occur in Siberia. 

The geological period associated with the entry into the Palaearctic of 
some species is difficult to determine. It is only certain that the main 
contingent of mosquitoes entered the boreal zone of the Palaearctic in the 
Upper Tertiary, at any rate before the Pleistocene. This is proved by the 
presence in the mountains of the southern Palaearctic (Pyrenees, Caucasus, 
Tien Shan) of a number of Culicidae which belong to forest areas, e. g. 
Aedes pullatus and Culiseta alaskaensis. The spread of 
numerous southern elements, Ethiopian and Oriental, into the boreal fauna 
was probably more extensive than it is at present. The pressure of the 
southern elements in the USSR mentioned above proves this. 

When the geographical distribution of mosquitoes in the Palaearctic is 
studied, it should be remembered that most of the species of Culicidae are 
widely distributed and endemism in the various geographical regions is 
relatively weakly developed. The cosmopolitan distribution of Culex 
pipiens is often quoted as an example of a wide distribution. But this is 
a typical synanthropic species and its wide distribution is probably con- 
nected with man. A more convincing example would be the "feral" Aedes 
v e x a n s, which is distributed in Europe, Asia, Australia, Oceania, Africa 
and North America. 

We know little about the Palaearctic species of mosquitoes with a small 
distribution. Possible examples are Culiseta setivalva (Crimea, 
Western Transcaucasia, Bulgaria, Anatolia) and Aedes alektorovi 



44 



(Maritime Territory, southern Amur area), but their apparently narrow dis- 
47 tribution may be due to insufficient collections. 

It is characteristic that in a well studied area like the Caucasus, in which 
many endemic species of insects are found, there is not a single endemic 
species of mosquito. Endemism determined by mountains is usually not 
characteristic for mosquitoes. There is not a single species which occurs 
only in mountains in the USSR, but in other bloodsucking Diptera, Simuliidae, 
Ceratopogonidae and Tabanidae, such species are numerous. Aedes pul- 
latus, the only species which is usually considered as a mountain form, 
also occurs in the plains in the northern parts of its range. 

The fauna of mosquitoes of the USSR at present (1966) consists of 85 
species belonging to 8 genera, half of them species of Aedes. In the thirty 
years since the publication of the previous edition of this volume, the list of 
species has increased by 17 due to the description of 5 new species and the 
discovery of 17 species hitherto not known to occur in the USSR. On the 
other hand, 4 species have been omitted as their status is doubtful (Aedes 
duplex Mart., Cul i se t a silvestris Shing. ) or they have been con- 
sidered as subgenera (Aedes esoensis, A.rossicus). Aedes 
lepidonotus has been excluded from the list of mosquitoes of the USSR 
as its occurrence is doubtful. This list is not definitive. It will probably 
grow but not very much. 



IMPORTANCE OF MOSQUITOES FOR MAN 

MOSQUITOES AS BLOODSUCKING INSECTS* 

Mosquitoes are an important element of bloodsucking flies which are a 
scourge in the taiga, tundra and other landscapes. Their economic impor- 
tance is determined by the fact that they attack man and animals. The injury 
is done by the females, which feed on plant juices but also on the blood of 
man, mammals, birds and other animals. 

Mosquitoes are temporary parasites since their contact with the host is 
very brief, just long enough to suck blood. The bite is little noticed or not 
felt at all due to their thin mouth parts (the proboscis of Anopheles is 
only about 0.055 mm thick). At the beginning of bloodsucking, saliva is 
injected through the hypopharynx, the secretion of the salivary glands in 
48 the thorax. The amount of saliva is very small, 1 — 3/Ltg in Aedes 

a e gy Pt i, average 4.7^g (Devine, Venard and Myser, 1956), but the effect 
of the bite is often very marked. It depends on the species of mosquito, 
the sensitivity of the subject and the number of bites (single or multiple 
bites). 

People react differently to mosquito bites. The reaction varies from 
total insensitivity to high sensitivity, depending on the constitution of the 
subject and on its allergic state (anaphylaxis). In many people of low 
sensitivity the bites of Anopheles cause only slight reddening of the skin 
and a small papule at the site of the bite. After the bite of some species 
of A e d e s, a swelling appears in 10—20 minutes which may be 12 mm wide. 

* Written by E.N.Pavlovskii. 



45 



The subjective symptoms are itching and a burning sensation. Birds re- 
act severely to bites of Aedes but show little reaction to bites of Culex. 
The bite may cause local inflammation. This is usually secondary, because 
the mosquito had fed on infected material earlier and had then injected the 
infection into the host. Scratching may also result in secondary infections. 

In mass attacks, the general reaction of the skin and sensitivity increase 
correspondingly. The papules may fuse and the skin swells, for a day. If 
there is only a single bite, the pain is slight, but if there is a massive attack 
it may be intolerable. During the mass flight of mosquitoes (especially of 
Aedes), field work is impossible, at least during the hours the insects are 
active. This hinders work in the forest, building of dams, bridges, roads, etc. 

Domestic animals also suffer from mosquitoes. Cattle cannot work or 
pasture. Fattening and milk yield decrease. All this is due not to loss of 
blood but to the toxic effect of the saliva and to side effects of the bites. 
The toxic properties of the saliva are the primary cause of the skin reaction 
and general reactions (Pavlovskii, Shtein and Perfil'ev, 1928; Hecht, 1929). 



MOSQUITOES AS VECTORS OF DISEASE 

Mosquitoes are important vectors of human disease. Filariasis, yellow 
fever and dengue are common in the tropics but malaria, mosquito-borne 
encephalitis, etc. also occur in the temperate zone. Control of the vectors 
is important in prophylaxis, and its success depends upon an intimate 
knowledge of their specific composition and biology. 

There has been little study of the transmission of disease to domestic 
49 animals, and yet this is certainly important. Mosquitoes also transmit 
disease to wild animals, but this has been little studied. 

The causative agents which mosquitoes transmit are Plasmodia, filariae, 
bacteria and viruses. 



Mosquitoes and Plasmodia of malaria 

Species of Anopheles transmit all four species of plasmodia, which 
cause malaria in man: Plasmodium vivax, P. ovale, P.malariae 
and P. falciparum. People with gametocytes in the blood are the source 
of infection of mosquitoes. Fertilization of the plasmodia and part of their 
life cycle and reproduction, sporogony, take place in the mosquito and end 
in the appearance of sporozoites in the salivary glands. Infection takes 
place during the bite by the saliva which contains sporozoites. 

Most species of Anopheles are able under experimental conditions 
to transmit plasmodia from man to man to a greater or lesser extent, but 
the epidemiological importance of the different species is very irregular in 
nature, depending on the susceptibility to the infection by the various species 
of plasmodia, length of life of the mosquito, frequency of attack on man, 
duration of seasonal activity, etc. (Beklemishev, 1941). 



46 



Only a few species are effective (actual or potential) vectors of malaria: 



Region 



Species of Anopheles 



50 



Europe (except the Mediterranean), Siberia 
Southern Europe, Transcaucasia 

Middle Asia 



East Asia 
Southeast Asia 



Australia, New Guinea 
India 



Southwest Asia 



North Africa 



Africa south of the Sahara 



South America 



Central America 



North America 



A .maculipennis Mg . 

A.maculipennis Mg . 

A.superpictus Gr. 

A.maculipennis sacharovi Favre 

A.superpictus Gr. 

A . pulcherrimu s Theob. 

A.hyrcanus Pall, (s.l.) 

A . pattoni Christ. 

A. minimus Theob. 

A.maculatus Theob. 

A.sundaicus Rod. 

A .jeyporiensis James 

A.leucosphyrus Don. 

A . 1 etif er Sand. 

A.umbrosus Theob. 

A . barbirostris V.d.Wulp 

A.aconitus Don. 

A.hyrcanus Pall, (s.l.) 

A.punctulatus Don. 

A .farauti Lav. 

A. Stephen si List. 

A.culicifacies Giles 

A. minimus Theob. 

A .sundaicus Rod. 

A . f lu vi atilis James 

A .philippinensis Ludl. 

A.maculipennis Mg. (s.l.) 

A.superpictus Gr. 

A .stephensi List. 

A .maculipennis Mg. (s.l.) 

A.sergenti Theob. 

A. multicolor Camb. 

A.pharoensis Theob. 

A.gambiae Giles 

A .funestus Giles 

A.nili Theob. 

A.pharoensis Theob. 

A .rufipes Gough 

A .darlingi Root 

A.albimanus Wied. 

A .aquasalis Curry 

A.pseudopunctipennis Theob. 

A .a lb it arsis Arr. 

A.albimanus Wied. 

A.pseudopunctipennis Theob. 

A. aquasalis Curry 

A.quadrimaculatus Say 

A.freeborni Aitk. 



From different sources: Russell etal., 1946; Horsfall, 1955; Foote and Cook, 1959, and others. These data 
were obtained when malaria was widespread. Malaria has now been eradicated or become rare in many 



47 



In the past, when malaria was prevalent, 2 of the 9 species of Anopheles 
which occur in the USSR were considered as important vectors: A . ma cu- 
lipennis.in nearly the entire area in which malaria was present, and 
A. superpictus, in Middle Asia and Eastern Transcaucasia. Three or 
four species, A. pulcherrimus, A.hyrcanus, A. claviger and, 
possibly, A. plumbeus, were considered as vectors of local or minor 
importance. A. algeriensis, A. marteri and A. lindesayi do not 
transmit malaria. In India, only 7 of the 42 species of Anopheles which 
occur there are serious vectors of malaria (Roy, 1946). In Malaya, 6 of 
the 47 local species of Anopheles are considered as serious vectors 
(Sandosham, 1959). In Brazil, 4 of the 50 local species of Anopheles 
are the main vectors and 5 are minor vectors (Ferreira, 1964). 

The duration of the gonotrophic cycle of Anopheles is usually shorter 
than that of the sporogony of Plasmodium at the same temperature and 
sporozoites may appear only in females after 3—4 gonotrophic cycles 
(Beklemishev, 1944). The life of a mosquito is short and most die before the 
sporogony is completed (Detinova, 1962). 

In localities in which control includes residual insecticides, the population 
of mosquitoes becomes reduced and their life shortened. The females do 
not live to the infective stage, i. e. to the time when there are sporozoites in 
the salivary glands. This causes a reduced circulation of the plasmodia. 
The use of residual insecticides alone or combined with other methods has 
eradicated malaria in many parts of the world including the Soviet Union. 

Malaria has been eradicated in some countries but there are areas where, 
in spite of control, transmission of Plasmodia continues. These are 
designated problematic regions (Committee of Experts on Malaria, 1965). 
51 The difficulties may be connected with the "entomological factor:" resist- 
ance of mosquitoes to insecticides, changes in the behavior of the vectors, 
the appearance of new species or subspecies in a locality, the replacement 
of endophilic species of Anopheles by exophilic species, etc. These 
problems can be overcome only by a more detailed study of the specific 
ecology of the species of Anopheles. 

Control carried out for a long time has resulted not only in a lowering 
of their numbers but also in their eradication in many regions. An illustra- 
tion has been given by Kalmykov (1965) which shows the gradual reduction 
of the distribution ofA.maculipennis sacharovi and A. pulcher- 
rimus in Tadzhikistan. 

Mosquitoes also transmit plasmodia of animals. An exception is ap- 
parently Hepatocystis kochi, a parasite of African monkeys which is 
transmitted by Ceratopogonidae like probably also other species of this 
genus, but the genus Hepatocystis should be removed from the family 
of Plasmodiidae and transferred to the family Haemoproteidae. The 
vectors of the plasmodia of monkeys are species of Anopheles. Thus, 
Plasmodium cynomolgi bastianelli (which may also parasitize 
man) developed under experimental conditions to the stage of sporozoites 
in Anopheles hyrcanus, A. kochi Donitz, A. maculatus, 
A. philippinensis Ludlow, and A. sundaicus (Warren et al., 1963). 
The only carriers of plasmodia in Malaya are A. balabacensis Baisas, 
A. leucosphyrus and A. hackeri Edwards . P. brasilianum was 
successfully transmitted from Ateles geoffroyi to other monkeys of 



48 



the same species and also to human volunteers by A. freeborni 
(Contacos et al., 1963). 

Species of Anopheles are also the vectors of Plasmodia of rodents. 
Plasmodium berghei is the most thoroughly studied and most often 
used in experiments with parasites of malaria. This parasite was trans- 
mitted experimentally from one rodent to another by Anopheles 
(A. Stephens i, A . quadrimaculatus; see Yoeli, 1965) and also by 
Aedes aegypti (see Raffaele, 1965). Under natural conditions A.dur- 
eni Edwards (Vincke, 1964) transmitted P. berghei in the Congo 
(Kinshasa). 

Species of Culex are the vectors of plasmodia of birds; Plasmodium 
gallinaceum, P. lophurae, P. fallax and others are transmitted by 
species of Aedes. Some plasmodia of birds may possibly also develop in 
A nop he le s, for example P. gallinaceum in A. quadrimaculatus. 
Details have been given by Huff (1965). 

Plasmodia of animals are sometimes transmitted by mosquitoes which 
do not bite man, for example A. hacker i, a vector of some plasmodia of 
monkeys in Malaya (Wharton et al., 1964), but A.letifer is a vector of 
human plasmodia and of P. traguli, which parasitizes Tragulus 
javanicus (Wharton et al., 1963) in that area. Under these conditions, 
the evaluation of studies on the natural infection of mosquitoes with plas- 
modia is obviously difficult. 

There are also provisional data on the possible role of mosquitoes as 
vectors of plasmodia in reptiles. Oocysts were found in a few species 
52 (Culex t e r r i t a ns, etc.) which feed on lizards infected with 
P. floridense. 



Mosquitoes and filariae 

Mosquitoes are the intermediate hosts of some nematodes of the sub- 
order Filariata (referred to below as filariae). The mosquitoes ingest the 
microfilariae with the blood of the vertebrate. The microfilariae develop 
in the thoracic muscles, in the body cavity and fat body or in the Malpighian 
tubes. Infective larvae migrate to the proboscis of the mosquito and enter 
the skin during the bite. Filariae do not multiply in the vector. Only a few 
larvae ingested by the mosquito are infective. 

As the result of the long, related evolution of the parasite and its inter- 
mediate host, special adaptations developed, e. g. the periodicity of the 
microfilariae in the peripheral blood of the vertebrate which corresponds 
to the diurnal activity of the mosquito. Thus, the microfilariae of 
Wuchereria bancrofti appear in the peripheral blood of man at night, 
which corresponds to the nocturnal activity of most mosquitoes. An 
exception is the aperiodic variety of this parasite in Polynesia which appears 
in the blood during the day, and its intermediate host is Aedes poly- 
nesiensis Mag. (Mattingly, 1962). 

There are over 30 species of filariae, parasites of man and animals, 
transmitted by mosquitoes. Further details are given by Lavoipierre 
(1958), Hawking and Worms (1961) and Nelson (1964). However, the life 
cycle of Filiarata is known only in 10% of the species. 



49 



Of the filariae parasitizing man, Wuchereria bancrofti and 
Brugia malayi are transmitted by mosquitoes (for their vectors, see 
Infections Caused by Wuchereria and Brugia, 1963; Edeson and 
Wilson, 1964). The larvae of these species develop in the thoracic muscles 
of the mosquito. 

Wuchereria bancrofti infects millions of people in the tropics and 
subtropics in both hemispheres. The list of mosquitoes in which the larvae 
develop to the infective stage includes numerous species, mainly of Culex 
and Anopheles, but in many species of mosquitoes W. bancrofti does 
not develop or its development is retarded or ceases at some stage. Some 
species of different genera are "good" vectors while the microfilariae do 
not develop in related species of the same genera. There may also be 
distinct differences between the subspecies of the same species. Thus, 
Culex pipiens fatigans is a "classic" carrier of W. bancrofti 
although the larvae of this nematode do not develop in Culex pipiens 
pipiens. There may also be differences in susceptibility to infection 
53 between different populations of the same species of mosquitoes. Individual 
differences have also been observed in infection not only with filariae but 
also with other parasites, e. g. plasmodia, viruses. 

The role of mosquitoes as vectors of filariae depends on many factors. 
Under natural conditions, few species of mosquitoes transmit filariasis to 
man. Inmost countries in which W. bancrofti is endemic one of the 
main vectors is the circumtropical, synanthropic C. pipiens fatigans. 
A large role in the distribution of the filariae is apparently played by 
Anopheles gambiae (Africa ), A. hyrcanus and Aedes togoi 
(East Asia), A. minimus and A. nigerrimus Giles (Southeast Asia), 
A. farauti and A. bancrofti Giles (New Guinea and the Solomon 
Islands) and other species. W. bancrofti is known at present only as a 
specific parasite of man, and synanthropic species of mosquitoes or at any 
rate mosquitoes which feed mainly on man are therefore the main vectors. 

Brugia malayi is widely distributed in Southeast Asia, its main 
vectors are Aedes togoi, Anopheles hyrcanus, A. nigerrimus 
and species of the genus Mansonia. Man may be infected by mosquitoes 
ingesting microfilariae from animals since the filariae parasitize not only 
man but also domestic and wild animals (cats, monkeys, etc.). Species of 
Mansonia transmit the filariae among animals. 

Cases of wuchereriasis have been introduced in the USSR, but the local 
origin of cases cannot be excluded since species known as vectors of 
filariae occur in the USSR and the temperatures in Transcaucasia and 
Middle Asia permit the development of larvae of filariae in mosquitoes. 
The discovery of a focus of wuchereriasis in the mountains of East Anatolia 
(Yucel and Deschiens, I960) is of interest. 

Mosquitoes are also apparently the intermediate hosts of other species of 
Brugia which parasitize animals. A detailed study was made of the develop- 
ment of Brugia ceylonensis, a parasite of dogs, in Aedes aegypti, in 
which the larvae developed in the thoracic muscles (Jayewardene, 1963). 

The role of mosquitoes as intermediate hosts of filariae of other genera 
parasitic in animals has also been studied. The transmission of Diro- 
filaria of dogs (Dirofilaria immitis and D . r e p e n s ) has been 
investigated. The larvae develop in the Malpighian tubes of Anopheles 



50 



maculipennis, A. claviger, A. hyrcanus, Aedes aegypti, 
A. albopictus, A. vexans and others . Dirofilariasis of dogs is widely- 
distributed in the southern USSR (Ukraine, Caucasus, Middle Asia). Diro- 
filaria of monkeys is also transmitted by mosquitoes. The larvae of 
D.aethiops of monkeys do not develop in the Malpighian tubes but in the 
body cavity and fat body of the mosquito (Webber, 1955). 

Mosquitoes are the intermediate hosts of filariae which parasitize farm 
54 animals, particularly of species of the genus Setaria, S. equina, 

S. marshalli, S. digitata, S. labiatopapillosa. As to the last 
species, there are data that the larvae also develop in bloodsucking flies. 

Our knowledge of the vectors of filariae of wild animals is very in- 
complete. However, the role of mosquitoes of the genera Aedes, Armi- 
geres, Culex, Mansonia and Anopheles as intermediate hosts of 
filariae which parasitize wild mammals, birds, reptiles and amphibians has 
been established. Mosquitoes are the intermediate hosts of 5 species of 
the genus Foleyella which parasitize frogs. The development of the 
microfilariae in the body cavity and the fat body of Culex pipiens 
fatigans, C. p. molestus and Aedes aegypti has been established. 
The feeding of mosquitoes on Amphibia is thus not rare but common. 



Mosquitoes and arboviruses 

Among the arboviruses of man and animals are viruses that reproduce 
in bloodsucking arthropods and are transmitted to vertebrates by the 
saliva of insects or mites. There are many such viruses: there were 155 
in 1963, of which 117 were isolated from mosquitoes (Gordon Smith, 1964). 
This figure is increasing each year. Mosquitoes play a major role in the 
transmission of virus infections (see Mattingly, 1960; Chamberlain and 
Sudia, 1961; Reeves, 1962, 1 965; Sazonova, 1962; Gordon Smith, 1964; 
Gutsevich, 1964; Pratt, 1964; also Viruses Transmitted by Arthropods 
[Russian], 1962). 

Below is a list of the most important species of mosquitoes which trans- 
mit virus infections that have been well studied: 

Disease Main vectors 

Yellow fever In urban areas — A edes aegypti; in natural foci — 

A.africanus Theob., A . si m pso ni Theob. and 
other species of Stegomyia (Africa), Haemagogus 
sp.,Aedes (F inlay a) leucocelaenus D. and Sh., 
Sabethes chloropterus Humb. and other species 
(South and Central America) 

Dengue (dengue and Chikungunya viruses) Aedes aegypti, A. albopictus and other species 

of S t eg omy ia 

Japanese B encephalitis Culex t rit ae niorhy nchus, C .g elidus Theob., 

C .vishnui Theob., C .pipiens, Aedes togoi 
and others 

Murray Valley encephalitis Culex a nnul iros tri s Sk. and others 

West Nile encephalitis Culex pipiens molestus, C.univittatus, 

C.antennatus Beck., A edes aegypti and others 

St. Louis encephalitis Culex pipiens fa tiga ns, C . tarsalis Coq. 

Eastern equine encephalitis (America) Culiseta melanura Coq., Aedes sollicitans 

Walk., A. vexans, Culex restuans Theob. and 
others 

51 



Western equine encephalitis (America) 
Venezuelan equine encephalitis 

California encephalitis 
O'nyong-nyong fever (Africa) 



Culex tarsalis 

Mansonia perturbans Walk., M .t itillans 

Walk, and others 
Aedes melanimon D., Culex tarsalis (probably) 
Anopheles f unes tus, A .gambi ae 



55 The role of mosquitoes as vectors is determined by the intensive multi- 

plication of the virus in the insect and the accumulation of the virus in the 
salivary glands in an amount sufficient to infect the vertebrate. 

Characteristic for most arboviruses is a broad range of invertebrate 
hosts. Mosquitoes of different genera, rarely species of one genus or sub- 
genus, are potential carriers of the virus. The vectors of the virus of 
dengue are only a few species of Stegomyia. Under equal conditions the 
virus usually multiplies intensively in the mosquitoes of a certain species 
but less intensively or not at all in related species. Examples may be found 
in the publications cited. 

Nearly all virus infections transmitted by mosquitoes have natural foci. 
Species of mosquitoes that rarely if ever attack man may have a distinct 
role in circulating the virus in natural foci. An example is Culiseta 
melanura Coq., which is a vector of the virus of eastern equine encepha- 
litis among birds. Synanthropic species of mosquitoes spread the infection 
among people . This is the role of Aedes aegypti and A. albopictus 
as vectors of dengue. Species of mosquitoes with a broad range of hosts, 
feeding on man and on animals, may spread the virus among animals and 
also transmit it to man. Such species are Culex t a r s a 1 i s, the vector 
of western equine encephalitis in North America, and C . tritaenio- 
rhynchus and C. gelidus, vectors of Japanese B encephalitis in East 
and Southeast Asia. 

If the mosquito-vectors feed on wild animals and on man, the virus may 
spread from natural foci to populated zones. An example is the outbreak 
of yellow fever in Ethiopia in 1960- 1962, when more than 15,000 people were 
infected, with a mortality of about 85% (Serie et al., 1964). The connection 
between the infected animals (monkeys) and people was apparently Aedes 
simps oni, which lives in forests but also in habitations and areas near to 
habitations, and attacks both monkeys and man. 

The part played by mosquitoes in the spread of viruses among wild 
animals has been little studied. Since some viruses are prevalent mainly 
among birds and others among mammals (rodents), the possible role of 
mosquitoes as vectors is determined to a large extent by their food 
preference. 

Man is usually an accidental, facultative, link in the spread of viruses 
transmitted by mosquitoes, but this may take place on a wide scale in nature, 
as serological examinations of animals have shown. In determining the 
vectors of viruses, biological characters of mosquitoes, e.g. their mobility, 
frequency of bloodsucking and repetition of gonotrophic cycles, and possible 
mass development in a short time have to be considered. Some of these 



52 



characters may explain severe outbreaks of virus infections transmitted 
by mosquitoes. Such outbreaks were the epidemics of yellow fever in 
Europe and America and recently of O'nyong-nyong fever in East Africa 
56 in 1959—1960 (more than one million people infected), the epidemic of dengue 
and the form of hemorrhagic fever (also caused by the virus of dengue) in 
Southeast Asia. The vectors in these epidemics were species closely con- 
nected with man: Aedes aegypti (dengue), A n o p h e 1 e s funestus, 
and A.gambiae (O'nyong-nyong fever). 

Mosquitoes transmitting virus infections are most prevalent in the 
tropics but they also occur in temperate zones, e.g. some forms of en- 
cephalitis in North America and Japanese B encephalitis in East Asia. 

The transmission of West Nile encephalitis is of special interest. The 
virus is apparently transmitted by mosquitoes and probably also by mites 
and ticks, Argasidae and Ixodidae. Foci of West Nile fever are known in 
Africa, Southwest Asia, India and also in Southern Europe, particularly in 
the South of France (where the virus was isolated from Culex modestus) 
and in the Volga delta. 

In Czechoslovakia the role of mosquitoes in the transmission of tahynia 
virus (of the group of California encephalitis), isolated from Aedes 
caspius and A . ve x a n s, was studied (Bardos and Danielova, 1959). The 
transmis sion of the virus by A. vexans and A. sticticus was proved 
experimentally, but the results were negative in experiments with A . cin - 
ereus (Simkova et al., 1960; Danielova, 1962). A virus was isolated from 
Anopheles maculipennis which was named "Calovo" (Bardos and 
Danielova, 1962, Czechoslovakia) and belongs to the Bunyamvera group. 
Very similar if not identical viruses were isolated from Culex gelidus 
(batai virus ) in Malaya and from A. barbirostris, A. tesselatus 
Theob., A. subpictus and Culex bitaeniorhynchus in India 
(Chittoor virus; Singh and Pavri, 1966). Viruses of the group of California 
encephalitis and Bunyamvera virus transmitted by mosquitoes are apparently 
also widely distributed in the USSR. 

In the western Ukraine, some strains of lymphocytic choriomeningitis 
were isolated from Aedes communis, A. diantaeus, A. cantans 
and other species (Glushchenko et al., 1957; Vigovskii and Gutsevich, 1961). 
This virus may also belong to the group of arboviruses since its multipli- 
cation in arthropods had been proved (Rzhegachek, 1965). Many strains of 
viruses have been isolated from different species of mosquitoes in other 
parts of the USSR (Azerbaidzhan, Kazakhstan, etc.; Il'enko et al., 1962; 
Mirzoeva et al., 1964; Ananyan, 1965). 



Mosquitoes as mechanical vectors of disease 

In this form of transmission, the causative agent of the disease does not 
develop or multiply in the vector and it is transmitted without its 
participation. 

The best studied example is the transmission of the bacteria of tularemia 
by mosquitoes and other bloodsucking insects. The bacteria remain alive in 
mosquitoes feeding on infected rodents for a long time (to one month; 



53 



Olsuf'ev and Golov, 1938). The transmission takes place by mechanical 
inoculation, feeding of the infected mosquito. The longer the interval between 
feeding on the infected and the healthy animal (man), the greater the pos- 
sibility that the parasite will be transmitted. Transmission is also possible 
57 by crushing the infected mosquito on the scarified skin of the animal (man). 
Mosquitoes evidently have a prominent role in spreading the bacteria of 
tularemia in natural foci and also in the infection of people during outbreaks 
of the infection (Olsuf'ev and Rudnev, 1960). The vectors may be mosquitoes, 
particularly of the genus Aedes (A. caspius, A. vexans, A. cinereus, 
A. excrucians, A. flavescens and als o Mansonia richiardii 
and others). Infected mosquitoes have been repeatedly found in nature. 
Species of mosquitoes which feed preferably on rodents, which is especially 
characteristic for Aedes, are the main agents in the spread of tularemia. 
Mosquitoes are also mechanical vectors of viruses which cause disease 
in animals: fowl pox, infectious myxomatosis, and others. 



METHODS OF COLLECTION AND OF MOUNTING 
AND PRESERVING COLLECTIONS OF MOSQUITOES 

ADULTS 

Mosquitoes can be collected nearly everywhere and at any time of the 
year. In winter, the females of some species (especially of the genera 
Culex, Culiseta and Anopheles) hibernate in cellars, basements, 
caves and buildings containing animals. With the thawing of the snow, they 
leave their winter resting places and mass flights of mosquitoes which 
hibernated as eggs (Aedes) or as larvae (some Anopheles, Culiseta 
and others) take place at the end of spring. A new generation that hibernated 
as adults appears at the beginning of summer. 

Resting and biting females are collected with an aspirator, with test 
tubes, or with an insect net. The males are found on flowers, especially on 
white Umbelliferae, mountain ash, rhubarb, on leaves of shrubs and trees or 
during swarming. They decrease in numbers in summer but at this time new 
forms appear. Most species of Culex, Anopheles, Aedes and others 
are best collected at the end of summer. 

Methods ensuring complete, large and quantitative collections are widely 
used. The most exact results have been obtained by using a bell-trap net 
(Monchadskii and Radzivilovskaya, 1947), which is hung at a height of about 
2m. with an observer as bait under it. The net is lowered after five minutes 
and the insects in it collected. Less exact results are obtained by the use 
of an aspirator to catch the insects flying around the observer for a certain 
time. 

For quantitative calculation an insect net is used. The number of mos- 
quitoes caught with 100 swings is taken as a unit (Olsuf'ev, 1939). The 
58 mosquitoes should be removed from the net after every 10 swings to avoid 
damage. Less satisfactory results are achieved by swings through the 
vegetation as the insects are severely damaged. Other variations are huts, 
tents or canopies (Chinaev, 1959). People or animals are used as bait. 



54 



Light traps are used in quantitative catches. Electric lamps, ultraviolet 
or polarized light are the source of light (Breev, 1958, 1963; Zhogolev, 1959; 
Kovrov and Monchadskii, 1963; Zhogolev and Shcherbina, 1966). Light traps 
with suction devices (ventilators) are useful. Not all species of mosquitoes 
are attracted by light to the same degree, but this method gives valuable 
results since males are also collected. 

To obtain well preserved specimens, mosquitoes are bred from larvae 
and pupae. 

Potassium cyanide, chloroform and acetic and sulfuric ether are used to 
kill mosquitoes. The insects are placed in a jar with the above substances 
or placed in a test tube or aspirator together with absorbent cotton dipped 
in these liquids. Five to ten minutes (not longer) are sufficient to kill them. 
Mosquitoes hatching from pupae are killed a day after hatching when the 
integument has hardened. 

The mosquitoes are pinned on thin entomological pins (Nos. 0, 00) or 
short, thin pins (minuten pins). The insect is placed on a piece of white 
paper with its back turned upward. Holding it with a fine pair of tweezers, 
the pin is inserted in the middle of the thorax slightly lateral to the median 
line. The pin is pushed into a cork plate to about fa of its length. Mos- 
quitoes pinned on minuten pins are placed on a small rectangular piece of 
carton which is then fixed to an entomological pin (Nos. 1 — 2). Pinning 
mosquitoes from the side is also suitable. The label should give the 
locality, date, conditions of the locality and the name of the collector. The 
59 mosquitoes are examined with the binocular microscope at a magnification 
of X20-25. 

If it is not possible to pin the material, the mosquitoes are placed on 
cotton in a box and covered with paper with the data for the label. Naphtha- 
lene is placed in the box. Before dried mosquitoes are pinned, they are 
placed in a moist chamber for 3—6 hours. 

To determine the species of a mosquito it is often necessary to examine 
the genitalia of the male (hypopygium). The last segments of the abdomen 
are cut off and placed for a minute in 96° alcohol and then left for a few 
hours in 10% KOH. After they have been washed in water, they are passed 
through alcohols of increasing strength, dehydrated in 96° alcohol, cleared 
in oil of cloves or carbol xylene, and mounted in Canada balsam. The pre- 
paration is labeled with the same number or data as the pinned insect. 

Further details on methods of collection and preservation are given by 
E. N. Pavlovskii (1935, 1959), A. S. Monchadskii (1952), and P. A. Petrishcheva 
(1964). 



LARVAE 

The collection of larvae is necessary in any study of mosquitoes and 
gives the most complete data on their specific composition, times of develop- 
ment, and typical breeding places. All natural and artificial water bodies, 
permanent or temporary, should be examined. 

Inspections should begin in early spring, when the larvae of early spring 
species of A e d e s, which hatch from hibernating eggs, are found. 



55 



Hibernating larvae of Culicella, Culiseta, Anopheles claviger 
and Mansonia richiardii will be found only in the middle or at the 
end of spring, when the water is free of ice and its temperature is 5—12°. 
Until then, they lie immobile at the bottom. In the middle of spring, the 
larvae of Aedes appear. When the birdcherry begins to blossom, pupation 
of the early spring species takes place and the larvae of the late spring 
species appear. At the end of spring, when the lilac blossoms, larvae hatch 
from eggs deposited by hibernating females of Anopheles, Culiseta 
and C u 1 e x . 

Spring is the most important time for inspection, especially in the north 
and in the taiga, forest and forest-steppe of the USSR, because of the ap- 
pearance of larvae of Aedes. In southern parts of the USSR, most species 
of Culex and Anopheles are found in summer and autumn. The typical 

60 breeding places are inspected every 10—14 days, in the south (Caucasus and 
Middle Asia) every week. Other water bodies of the same type should be 
inspected every 3—4 weeks. Inspection for the larvae can be stopped from 
the middle of September, and in the south in early November. 

The breeding place is approached without noise and without shadowing 
it . The larvae of many species (Anopheles claviger, A. super- 
pictus, Aedes excrucians, Culiseta morsitans and others ) are 
very sensitive to disturbances and rapidly descend to the bottom where they 
may remain for 10—15 minutes. In small water bodies roiling may cause 
the larvae to rise to the surface. 

Nets are best for collecting larvae. The net should be 10— 15 cm wide and 
its wire frame 3— 5 mm thick and stiff. The rod should be 2 m long. The net is 
made of miller's gauze (Nos. 8—15) or fine-mesh, netting. Gauze is not 
suitable. The net should be rounded-conical. In conical nets the larvae 
accumulate at the bottom and are damaged. The net should not be more 
than 1.5 times longer than wide. 

The net is submerged two-thirds in the water at a right angle and drawn 
along the surface for 2— 3 m. It is then rapidly turned through 180°, at a 
depth of 10— 15 cm, against the current. 

To collect larvae in shallow or small water bodies, a wire hoop about 
10cm wide and covered with miller's gauze is effective. The larvae are 
washed in a dish with water. A dipper with a few small openings may 
also be used. 

Special pumps or small nets about 5 cm wide may be used for tree holes. 
Several pumpings are necessary and part of the water must be returned 
to the hole. 

The contents of the net are poured into a white enamel dish half filled 
with water from the breeding place. The larvae are placed in a jar and labeled. 

61 The jar should be protected against direct sunlight and onhot days covered with 

a wet cloth or put into a case with layers of gauze and with cotton between the jars . 

In the laboratory, the larvae are selected by species and stages. All 
pupae and some 4th-stage larvae are separated for the breeding of adult 
mosquitoes; the other larvae are preserved. 

For the development of mature mosquitoes the larvae and pupae are 
placed singly or in a group of several in a small cylinder or in an Erlen- 
meyer flask half filled with water, preferably water from the breeding place, 
with a fresh green leaf placed on it. The vessels are protected against 



56 



sunlight, covered with absorbent cotton and placed with their opening toward 
the source of light. The average length of the pupal stage is two days. The 
males pupate and hatch first. 

Larvae of Anopheles are examined from the dorsal side, where most 
characters are distinct. Few characters (comb of the 8th segment, fin, 
caudal hairs and gills) are visible in lateral view. 

Larvae of Culicinae are examined as follows: the anterior end from 
above, the posterior end laterally. The larvae are cut at abdominal segments 
4—5. Preparations of whole larvae are not suitable for determination. 

The larvae are preserved in 96° alcohol heated to 50—55°. This gives 
better preservation of the hairs and the stigmal plate. 

For permanent preservation and transport, the larvae are placed in a 
small tube with a label inside. The tube is filled with 96° alcohol and is 
then stopped with absorbent cotton to prevent air bubbles and damage by 
shaking. 

Gum solutions (Faure-Berlese) should not be used for permanent pre- 
62 parations. This method is suitable only for rapid determination in the field. 
The treatment of larvae with KOH is useless and causes deterioriation of 
the preparations. 

The larvae should be mounted in Canada balsam. They are pierced and 
transferred for 1 — 2 hours to absolute alcohol for complete dehydration. 
They are then placed in a clearing medium (oil of cloves, eugenol, terpineol) 
for 2—3 hours and mounted in Canada balsam. They are covered with a 
cover glass on pieces of wax about as high as the larvae are thick. 

It is often necessary to make preparations of the shed skin of larvae. 
The skin is placed for a few minutes in 96° alcohol and transferred to lactic 
acid, where it softens. It is then placed on a slide and straightened under 
the binocular; the lactic acid is then suctioned off and replaced with 96° 
alcohol. The skin is then fixed. Mounting is as in Canada balsam 
preparations. 

Larvae should be handled with care to avoid damage to the hairs, gills, 
etc. Live larvae are transferred with a pipette with a broad opening. 



Bibliography 

The literature on mosquitoes is very large. We have given only a short 
list. For details on the older literature on the USSR, the following are 
recommended: N. S. Shtakel'berg. Bibliografiya malyarii (epidemiologiya, 
perenoschiki, bor'ba s malyariei, organizatsiya protivomalyariinogo dela v 
SSSR) 1771—1935 (Bibliography of Mosquitoes (Epidemiology, Vectors, 
Control of Malaria, Organization of Anti-Malarial Measures in the USSR) 
1771-1935). -Moskva-Leningrad, Izd. AN SSSR. 1940. 963 pp. Auxiliary 
Indexes. 1941. 136 pp. The work contains a complete list of the Russian 
literature on mosquitoes, with notes. The journal of abstracts "Biologiya" 
should be consulted (issue "Zooparazitologiya"), and also the annual lists 
published in "Meditsinskaya Parazitologiya i Parazitarnye Bolezni" 
(N. N. Dukhanina). 



57 



Recent catalogues and books in non-Russian literature are: A.Stone, 
K. L. Knight and H. Starcke. A Synoptic Catalog of the Mosquitoes of the 
World (Diptera, Culicidae). 1959; 38-page bibliography, mainly on 
systematics and faunistics. W. R. Horsfall. Mosquitoes. Their Bionomics 
and Relation to Disease 1955, 79-page bibliography, mainly on the biology 
63 and medical importance of mosquitoes. M. Bates. The Natural History of 
Mosquitoes. 1949; 46-page bibliography, also mainly on the biology and 
medical importance of mosquitoes. A. N.Clements. The Physiology of 
Mosquitoes. 1963; 56-page bibliography, mainly on physiology and 
morphology. 

Extensive bibliographies are contained in the monographs listed below. 

For up-to-date non-Soviet references, see the following journals: Review 
of Applied Entomology, Ser. B (Medical and Veterinary), London; Tropical 
Diseases Bulletin, London. 



A. MONOGRAPHS AND REGIONAL REPORTS 
Publications in Russian 

Beklemishev, V.N. Ekologiya malyariinogo komara (Anopheles 

maculipennis Mgn. ) (Ecology of Anopheles maculipennis 
Mgn. ). -Moskva, Medgiz. 1944. 299 pp. 

Beklemishev, V. N. and N. K. S h ip i t s i n a (editors). Sezonnye yavleniya 
v zhizni malyariinykh komarov v Sovetskom Soyuze (Seasonal 
Phenomena in the Life of Anopheles in the Soviet Union). 
Moskva. 1957. 527 pp. 

Maslov, A. V. Komary roda Culiseta (Mosquitoes of the Genus 

Culiseta). - Moskva-Leningrad, Izd. "Nauka." 1967. 182 pp. 

Monchadskii,A.S. Lichinki komarov (sem. Culicidae) SSSR i sopredel' - 
nykh stran (Larvae of the Culicidae of the USSR and Adjacent 
Countries). - Opredeliteli po faune SSSR, 24: 1-379. Moskva- 
Leningrad, Izd. AN SSSR. 1936. 

Monchadskii, A. S. Lichinki krovososushchikh komarov SSSR i sopredel' - 
nykh stran (podsem. Culicinae) (Larvae of Mosquitoes of the USSR 
and Adjacent Countries (Subfamily Culicinae)). — Ibid., 37: 1—290. 
Moskva- Leningrad, Izd. AN SSSR. 1951. 

Monchadskii, A. S. and A. A. Shtakel'berg. Malyariinye komary 
Tadzhikistana i mery bor'by s nimi (Anophelines of Tadzhikistan 
and their Control). Stalinabad. 1943. 96 pp. 

Shtakel'berg, A. A. Krovososushchie komary (sem. Culicidae) Soyuza 
SSR i sopredel 'nykh stran (Culicidae of the Soviet Union and 
Adjacent Countries). — Opredeliteli po faune SSSR, 1: 1 — 170. 
Moskva-Leningrad, Izd. AN SSSR. 192 7. 

Shtakel'berg, A.A. Semeistvo Culicidae (The Family Culicidae). — 
Fauna SSSR, Nasekomye dvukrylye, 3(4): 1-257. Moskva- 
Leningrad, Izd. AN SSSR. 1937. 



58 



Publications in Other Languages 

Barraud.P. J, Family Culicidae, Tribes Megarhinini and Culicini. — 

The Fauna of British India Including Ceylon and Burma. Diptera, 

5: 1-463. London. 1934. 
Bates, M. The Natural History of Mosquitoes. New York. 1949. 379 pp. 
Carpenter, S. J. and W. J. La Casse. Mosquitoes of North America 

(North of Mexico). Berkeley and Los Angeles. 1955. 360 pp. 
Christophers, S. R. Family Culicidae. Tribe Anophelini. — The Fauna 

of British India. Diptera, 4:1-371. London. 1933. 
Christophers.S.R. Aedes aegypti(L.), the Yellow Fever 

Mosquito, its Life History, Bionomics and Structure. Cambridge. 

1960. 739 pp. 
Clements, A.N. The Physiology of Mosquitoes. International Series of 

Monographs on Pure and Applied Biology, Vol. 17: 1—393. Oxford. 

1963. 
Edwards, F.W. A Revision of the Mosquitoes of the Palaearctic Region. — 

Bull. Ent. Res., 12 (3): 263-351. 1921. 
Edwards, F. W. Una revisione delle zanzare delle regioni Palaeartiche. — 

Riv. Malar., 5(3/6): 1-152. 1926. 
E d w a r d s , F. W. Fam. Culicidae. Genera insectorum, diriges par 

Wytsman. Diptera, fasc. 194. Bruxelles. 1932. 258 pp. 
Edwards, F. W. Mosquitoes of the Ethiopian Region. III. Culicinae 

Adults and Pupae. London. 1941. 499 pp. 
Evans, A. M. Mosquitoes of the Ethiopian Region. II. Anophelini Adults 

and Early Stages. London. 1938. 404 pp. 
Feng,L. C. et al. Mosquitoes of China. Shanghai. 1958. 250 pp. (in 

Chinese.) 
Foote,R.H. and D. R. Cook. Mosquitoes of Medical Importance. — 

Agric.Handb. (U.S. Dep. Agric. ). 152. Washington. 1959. 158 pp. 
Gjullin, C. M., R. J. Sailer , A. Stone, and B. V. Travis. The 

Mosquitoes of Alaska. — Ibid., 182. Washington. 1961. 98 pp. 
Hopkins, G. H. Mosquitoes of the Ethiopian Region. I. Larval Bionomics 

of Mosquitoes and Taxonomy of Culicinae Larvae. 2nd edition. 

London. 1952. 355 pp. 
Ho r s f a 11 , W. R. Mosquitoes, their Bionomics and Relation to Disease, 

New York. 1955. 723 pp. 
Ki r kpat r ick , T. W. The Mosquitoes of Egypt. Cairo. 1925. 224 pp. 
Kramar, J. Komari bodavi — Culicinae. — Fauna C"SR, 13. Praha. 

1958. 286 pp. 
La Casse, W. J. and S. Yamaguti. Mosquito Fauna of Japan and 

Korea, pp. 1-268, 1-213. Kyoto. 1955. 
Lang, W. D. A Handbook of British Mosquitoes. London. 1920. 125 pp. 
Mar shall, J. F. The British Mosquitoes. London. 1920. 341pp. 
Martini, E. Uber Stechmiicken besonders deren europaische Arten und 

ihre Bekampfung. Leipzig. 1920. 267 pp. 
Martini, E. Culicidae. — In: E. Lindner, Die Fliegen der Palaearktischen 

Region, 11 and 12. Stuttgart. 1929-1931. 398 pp. 
Matheson, R. Handbook of the Mosquitoes of North America. 2nd edition. 

Ithaca. 1944. 314 pp. 
Mihalyi,F. and M. G ul y a s . Magyaroszag csipo szunyogjai. Budapest 

1963. 229 pp. (in Hungarian.) 



59 



Natvig, L. R. Contributions to the Knowledge of the Danish and Fenno- 

scandian Mosquitoes — Culicini. Oslo. 1948. 567 pp. 
Peus,F. Die Fiebermucken des Mittelmeergebiets. Leipzig. 1942. 150pp. 
Puri, J. M. Larvae of Anopheline Mosquitoes, with Full Description of 
those of the Indian Species. — Indian Med. Res. Mem., Vol.21: 
1-225. 1931. 
Rioux, J.A. Les Culicides du "Midi" Mediterranean. — Encycl. ent. (A), 

Vol. 35: 1-303. 1958. 
Seguy, E. Les moustiques de France. — Encycl. prat. Nat., Vol.19: 

1-225. 1923. 
Senevet,G. Les Anopheles du Globe. -Ibid., Vol.36: 1-215. 1958. 
Senevet, G. and L. Andarelli. Les Anopheles de 1' Afrique du Nord et 

du Bassin Mediterranean. -Ibid., Vol.33: 1-280. 1955. 
Senevet, G. and L. Andarelli. Les moustiques de 1' Afrique du Nord 
et du Bassin Mediterranean; les genres C u 1 e x, Uranotaenia, 
Theobaldia, Orthopodomyia et Ma n s o n i a. — Ibid., 
Vol. 37: 1-384. 1959. 
Snodgrass,R.E. The Anatomical Life of the Mosquito. — Smithson. 

Misc. Collns., 139(8): 1-87. 1959. 
Stone, A., K. L. Knight, and H. Starcke. A Synoptic Catalog of the 
Mosquitoes of the World (Diptera, Culicidae). — The Thomas Say 
Foundation, 6. Washington. 1959. 358 pp. 
Wesenberg -Lund, C. Contributions to the Biology of the Danish 

Culicidae. - K. danske Vidensk. Selsk. Skr., Vol.7: 1-210. 1921. 



B. SYSTEMATICS, FAUNISTICS, MORPHOLOGY, BIOLOGY 
Publications in Russian 

A 1 e kt o r o v , A. A. K faune komarov Culicidae Dal'nevostochnogo kraya 

SSSR (On the Fauna of the Family Culicidae of the Soviet Far East).- 
Parazitologicheskii Sbornik Zoologicheskogo Instituta SSSR, Vol. 2 : 
229-248. 1931. 

Bashkareva, A. L. Nekotorye dannye o malyariinykh i drugikh 

komarakh Sochinskogo raiona (Some Data on Anophelines and Other 
Mosquitoes of the Sochi Region). — Ibid., pp. 55—58. 

Blagoveshchenskii, D. I. Materialy po faune naruzhnykh parazitov 

(Arthropoda) zhivotnykh Kazalinskogo i nekotorykh drugikh raionov 
yuzhnogo Kazakhstana (Data on the Fauna of External Parasites 
(Arthropoda) of Animals of the Kazalinsk Region and Other Regions 
in Southern Kazakhstan). — In Sbornik: "O vreditelyakh zhivotno- 
vodstva v Kazakhstane," pp. 11—84. Moskva— Leningrad, Izd. AN 
SSSR. 1937. 

Bozhkov, D. K. Krovososushchie komari (Diptera, Culicidae) Bolgarii 

(Mosquitoes (Diptera, Culicidae) in Bulgaria). — Entomologicheskoe 
Obozrenie, 45(3): 570-574. 1966. 



610144 60 



Bregetova.N.G. Aktivnost' napadeniya na cheloveka i ee sutochnyi 
ritm u komarov A e d e s caspius caspius (Fall.) Edw. i 
Aedes vexans Meig. v prirodnykh usloviyakh yuzhnogo 
Tadzhikistana (Activity of Attack on Man and its Diurnal Rhythm 
in Mosquitoes Aedes caspius caspius (Pall.) Edw. and 
Aedes vexans Meig. under Natural Conditions of Southern 
Tadzhikistan). — Izvestiya AN SSSR, Seriya Eiologicheskaya, 
Nos. 2-3: 251-279. 1946. 

Breev.K.A. O primenenii lovushek ul'trafioletovogo sveta dlya opredele - 
niya vidovogo sostava i chislennosti populyatsii komarov (Use of 
Ultraviolet Light Traps for Determination of the Specific Composi- 
tion and Population of Mosquitoes). — Parazitologicheskii Sbornik 
Zoologicheskogo Instituta AN SSSR, Vol. 18: 219-238. 1958. 

Breev,K. A. Vliyanie istochnikov sveta na chislennost' i vidovoi sostav 
komarov, sobiraemykh v svetovye lovushki (The Effect of Light 
on the Population and Specific Composition of Mosquitoes Collected 
in Light Traps). — Entomologicheskoe Obozrenie, 42 (2): 280—303. 
1963. 

Chagin.K. P. Nablyudeniya nad tsiklom razvitiya A e d e s (F.)togoi v 
laboratornykh i prirodnykh usloviyakh (Observations on the Cycle 
of Development of Aedes (F.)togoi in the Laboratory and in 
Nature). — Meditsinskaya Parazitologiya i Parazitarnye Bolezni, 
12 (2): 44-52. 1943. 

Chinaev, P. P. Metody kolichestvennogo ucheta krovososushchikh 

komarov (Diptera, Culicidae) (Methods of a Quantitative Census of 
Mosquitoes (Diptera, Culicidae)). — Entomologicheskoe Obozrenie, 
38 (4): 757-765. 1959. 

Denisova.Z.M. Izmenchivost' gipopigiev Anopheles maculipennis 
messeae Fall, v populyatsii Marbumstroya (Mariiskaya ASSR) 
(Variability of the Hypopygium of Anopheles maculipennis 
messeae Fall, in Marbumstroi (Mari ASSR)). — In Sbornik: 
"Voprosy fiziologii i ekologii malyariinogo komara," Vol. 1: 
113-119. 1940. 

Denisova,Z. M. Materialy po izucheniyu variatsii gipopigiev samtsov 

Anopheles maculipennis atroparvus Thiel i A n o p h e 1 e s 
maculipennis sacharovi Favre (A Study of the Variation of 
the Hypopygium of Males of Anopheles maculipennis 
atroparvus Thiel and A n o ph e 1 e s maculipennis sachar- 
ovi Favre). — Meditsinskaya Parazitologiya i Parazitarnye 
Bolezni, 3(17): 221-227. 1948. 

Den i sova , Z. M. Novyi vid A e d e s ( O c h 1 e r o t a t u s ) (A New Species of 
Aedes (Ochlerotatus)). -Ibid., 24(1): 58-61. 1955. 

Denisova, Z. M. Variatsii gipopigiev samtsov nekotorykh podvidov 

malyariinogo komara A no ph e le s maculipennis Mg. (Diptera, 
Culicidae) (Variation of the Hypopygium in Males of Some Subspecies 
of Anopheles maculipennis Mg. (Diptera, Culicidae)).— 
Entomologicheskoe Obozrenie, 43 (4) : 815-822. 1964. 

Enikolopov,S. K. Ob ekologii Anopheles algeriensis Theob. 
(On the Ecology of A n op he 1 e s algeriensis Theob.). — 
Meditsinskaya Parazitologiya i parazitarnye Bolezni, 6 (3): 
354-359. 1937. 



61 



Fedorov,V.G. K obnaruzheniyu Culex molestus Forskal v 

Leningrade (On the Finding of Culex molestus Forskal in 
Leningrad). -Ibid., 15(2): 58-67. 1946. 

F r i do 1 in , V. Yu. Zhivotno-rastitel'noe soobshchestvo gornoi strany 

Khibin (Animal and Plant Communities in the Khibiny Mountains).— 
Trudy Kol'skoi Bazy AN SSSR, Vol. 3: 1-295. 1936. 

Glagoleva, E. M. Nekotorye dannye po ekologii lichinok Anopheles 
v Tadzhikistane. I. Biotopy Anopheles algeriensis Theob. 

II. Biotopy An ophe le s lindesayi Giles i Anopheles 
marteri Sen. Prun. III. Biotopy A n op h e 1 e s bifurcatus. 
IV. Biotopy A nop he le s pulcherrimus Theob. (Data on the 
Ecology of A n op h e le s Larvae in Tadzhikistan. I. Biotopes of 
Anopheles algeriensis Theob. II. Biotopes of A n op he 1 e s 
lindesayi Giles and A n op h e 1 e s marteri Sen. Prun. 

III. Biotopes of Anopheles bifurcatus. IV. Biotopes of 
Anopheles pulcherrimus Theob.). — Meditsinskaya 
Parazitologiya i Parazitarnye Bolezni, 13 (4): 58—64 and 65—67; 
13(5): 47-52; 13(6): 41-44. 1944. 

Glagoleva, E.M. Ob ekologii lichinok Anopheles v Tadzhikistane 
(On the Ecology of Anopheles Larvae in Tadzhikistan). — 
Ibid., 15 (5): 60-67. 1946. 

Gutsevich, A. V. Novye i malo izvestnye formy komarov (New and 
Little -Known Forms of Mosquitoes). — Trudy Zoologicheskogo 
Instituta AN SSSR, Vol. 18: 320-324. 1955. 

G ut s e v i c h , A. V. Novyi vid krovososushchego komara roda Aedes 
(Diptera, Culicidae) iz Kazakhstana (A New Species of Aedes 
(Diptera, Culicidae) from Kazakhstan). — Entomologicheskoe 
Obozrenie, 41(4): 886-888. 1962. 

Gutsevich, A. V. Novyi vid komara iz Uzbekistana — Aedes 

(Ochlerotatus) simanini (Culicidae) (A New Species of 
Mosquito from Uzbekistan — Aedes (Ochlerotatus) simanini 
(Culicidae)). - Zoologicheskii Zhurnal, 45(3): 457-459. 1966. 

Gutsevich, A. V., Z. S. Donets, G.G. Ezhova, and A. M. Popov. 

Krovososushchie komary (Diptera, Culicidae) Chernovitskoi oblasti 
(Mosquitoes (Diptera, Culicidae) of the Chernovtsy Region). — 
Entomologicheskoe Obozrenie, 41 (2): 355-358. 1962. 

Kalmykov, E. S. Izmeneniya v rasprostranenii malyariinykh komarov 
v svyazi s khozyaistvennoi deyatel'nost'yu cheloveka (Changes of 
the Distribution of Anopheline Mosquitoes in Relation to the 
Economic Activity of Man). — Nauchnye Trudy po Parazitologii, 
pp. 55-61. Dushanbe, Izd. "irfon." 1965. 

Kazantsev,B.N. K voprosu ob izmenchivosti risunka i okraski kryl'ev 
komarov roda Anopheles Srednei Azii (Variation of the Pattern 
and Coloration of the Wings of the Genus Anopheles in Middle 
Asia). — Parazitologicheskii Sbornik Zoologicheskogo Instituta 
AN SSSR, Vol.1: 229-235. 1930. 

Kazantsev,B. N. Tsvetovye variatsii bukharskikh Aedes caspius 
(Color Variation of A e d e s caspius in Bukhara). —Ibid., 
Vol. 2: 85-90. 1931. 



62 



Keshish'yan, M. N. Culicidae Tadzhikistana (Culicidae of Tadzhikistan). 
Meditsinskaya Parazitologiya i Parazitarnye Bolezni, 10(1): 
77-80. 1941. 

Khelevin, N. V. VHyanie vneshnikh uslovii na vozniknovenie embrional' - 
noi diapauzy i na kolichestvo pokolenii u Aedes caspius 
dorsalis Mg. (Diptera, Culicidae) v techenie sezona (influence 
of the Environment on the Embryonic Diapause and Number of 
Generations in Aedes caspius dorsalis Mg. (Diptera, 
Culicidae) during a Season). — Entomologicheskoe Obozrenie, 
37(1): 24-47, 1958. 

Kovrov.B.G. and A. S. Monchadskii. O vozmozhnosti primeneniya 
polyarizovannogo sveta dlya privlecheniya nasekomykh (On the Use 
of Polarized Light to Attract Insects). - Ibid., 42 (l): 49-55. 1963. 

Lis ova, A. I. Materialy po faune komarov sem. Culicidae goroda 

Tashkenta i ego okrestnostei (Data on the Fauna of Culicidae of 
Tashkent and its Vicinity). — Parazitologicheskii Sbornik Zoologi- 
cheskogo Instituta AN SSSR, Vol. 5: 27-54. 1935. 

Lobkova, M. P. Nekotorye dannye po rasprostraneniyu komarov 

(podsemeistvo Culicinae) v Karelii (Data on the Distribution of 
Mosquitoes (Subfamily Culicinae) in Karelia). — In Sbornik: M K 
prirodnoi ochagovosti parazitarnykh i transmissivnykh zabolevanii 
v Karelii," pp. 108-119. Moskva-Leningrad, Izd. "Nauka." 1964. 

L ' v o v , D. K. O vidovoi samostoyatel'nosti komara-perenoschika 
yaponskogo entsefalita Aedes esoensis Yam. (Diptera, 
Culicidae) (On the Specific Status of the Vector of Japanese 
Encephalitis, Aedes esoensis Yam. (Diptera, Culicidae)). — 
Entomologicheskoe Obozrenie, 35(4): 929-934. 1956. 

Markovich, N. Ya. Novye dannye po biologii Anopheles bifurcatus 
(nablyudeniya na Severnom Kavkaze (New Data on the Biology of 
Anopheles bifurcatus in the Northern Caucasus). — 
Meditsinskaya Parazitologiya i Parazitarnye Bolezni, 10 (3/4): 
410-413. 1941. 

Martini, E. Komary Srednego Povolzh'ya (Mosquitoes of the Middle 
Volga Region). - Izvestiya AN SSSR, VI Seriya, 20(9): 607-618. 
1926. 

Maslov,A. V. Izmenchivost' i sistematika Pallasova malyariinogo 

komara (Variability and the Systematics of Pallas Anophelines). — 
Meditsinskaya Parazitologiya i Parazitarnye Bolezni, 15(6): 
39-46. 1946. 

Maslov, A. V. O sistematike krovososushchikh komarov gruppy C u 1 i s e - 
ta (Diptera, Culicidae) (On the Systematics of Mosquitoes of the 
Group Culiseta (Diptera, Culicidae)). —Entomologicheskoe 
Obozrenie, 48(1): 193-216. 1964. 

Monchadskii, A. S. K poznaniyu lichinok komarov (Diptera, Culicidae). 
II. Novye i maloizvestnye lichinki komarov Dal'nego Vostoka (On 
the Larvae of Mosquitoes (Diptera, Culicidae). II. New and Little - 
Known Larvae of Mosquitoes of the Far East). — Parazitologi- 
cheskii Sbornik Zoologicheskogo Instituta AN SSSR, Vol. 9.: 
167-180. 1947. 

Monchadskii, A. S. Lichinki Finlaya (Diptera, Culicidae) iz dupel 
derev'ev (Larvae of Finlaya (Diptera, Culicidae) from Tree 
Holes). -Ibid., Vol.11: 253-261. 1949. 



63 



Monchadskii,A. S. Letayushchie krovososushchie dvukrylye -gnus 

(Bloodsucking Insects). — Moskva— Leningrad, Izd. AN SSSR. 1952. 

Monchadskii,A. S. and Z. A. Radzivilovskaya. Novyi metod 

kolichestvennogo ucheta aktivnosti napadeniya krovososov (A New 
Method for a Quantitative Calculation of the Rate of Attack of 
Bloodsucking Insects). — Parazitologicheskii Sbornik Zoologi- 
cheskogo Instituta AN SSSR, Vol.9: 147-166. 1947. 

Olsuf'ev, N. G. Vidovoi sostav i sezonnaya dinamika chislennosti 

krovososushchikh dvukrylykh v del'te Volgi i ikh vozmozhnaya rol' 
v epidemiologii tulyaremii (Specific Composition and Seasonal 
Dynamics of Bloodsucking Insect Populations in the Volga Delta and 
their Possible Role in the Epidemiology of Tularemia). — 
Zoologicheskii Zhurnal, 18(5): 786-798. 1939. 

Pavlovskii.E. N. Metody izucheniya krovososushchikh komarov 

(Culicidae) (Methods of Studying Mosquitoes (Culicidae)). 2nd edition, 
pp. 1-176. - Moskva-Leningrad, Izd. AN SSSR. 1935. 

Pavlovskii, E. N. (editor). Laboratornyi praktikum meditsinskoi 

parazitologii (Komary) (Laboratory Course of Medical Parasitology 
(Mosquitoes)), pp. 200-249. - Leningrad, Medgiz. 1959. 

Petrishcheva,P.A. Fauna, ekologiya i biologiya Culicidae Turkmenii 
(Fauna, Ecology and Biology of the Culicidae of Turkmenia). — 
Parazitologicheskii Sbornik Zoologicheskogo Instituta AN SSSR, 
Vol. 6: 49-115. 1936. 

Petrishcheva,P.A. O krovososushchikh komarakh Primor'ya 

(Mosquitoes of the Maritime Territory). — In: Parazitologiya 
Dal'nego Vostoka, pp. 144—159. Leningrad, Medgiz. 1947. 

Petrishcheva,P.A. Metody izucheniya prirodnykh ochagov boleznei 
cheloveka (komary) (Methods of Studying Natural Foci of Human 
Diseases (Mosquitoes)), pp. 12 — 35. — Moskva, Izd. "Meditsina." 
1964. 

Prendel', A. R. Proiskhozhdenie i kharakteristika fauny krovososushchikh 
komarov UkrSSR (Origin and Characteristics of the Fauna of 
Mosquitoes in the Ukraine). — In Sbornik: "Kraevaya parazitologiya 
i prirodnaya ochagovost' transmissivnykh boleznei, " pp. 91—96. 
Kiev. 1966. 

R emm, Kh. Ya. Mater ialy po faune i ekologii komarov (Diptera, Culicidae) 
Estonskoi SSR (Data on the Fauna and Ecology of Mosquitoes 
(Diptera, Culicidae) in Estonia). — Entomologicheskoe Obozrenie, 
36 (1): 148-160. 1957. 

Saf'yanova, V.M. Materialy po ekologii krovososushchikh komarov 
Vladimirskoi oblasti (Data on the Ecology of Mosquitoes of the 
Vladimir Region). - Zoologicheskii Zhurnal, 39(2): 236-243. 1960. 

Sazonova,0. N. Novyi vid krovososushchego komara roda Aedes Mg. 
(Diptera, Culicidae) iz evropeiskoi chasti SSSR (A New Species of 
Aedes Mg. (Diptera, Culicidae) from the European USSR). — 
Parazitologicheskii Sbornik Zoologicheskogo Instituta AN SSSR, 
Vol. 16: 148-151. 1956. 

Sazonova, O. N. Tablitsa dlya opredeleniya samok komarov roda Aedes 
Mg. (Diptera, Culicidae) lesnoi zony SSSR (Key for the Determination 
of Females of Aedes Mg. (Diptera, Culicidae) of the Forest 
Zone of the USSR). — Entomologicheskoe Obozrenie, 37(3): 
741-752. 1958. 



64 



Shingarev,N. I. Zametki po Culicidae III. (Notes on the Family 

Culicidae. III). — Russkii Zhurnal Tropicheskoi Meditsiny, Vol. 6: 
47-53. 1928. 

Shlenova,M.F. Biologiya vazhneishikh vidov Aedes v srednei polose 
evropeiskoi chasti SSSR (Biology of the Important Species of 
Aedes in the Central European USSR). — Meditsinskaya 
Parazitologiya i Parazitarnye Bolezni, 28(2): 193 — 198. 1959. 

Solovei, V. Ya. and V.G. Likhoded. K faune i ekologii krovososu- 

shchikh dvukrylykh (Diptera) severo-zapadnoi chasti Murmanskoi 
oblasti (On the Fauna and Ecology of Bloodsucking Insects (Diptera) 
in the Northwestern Murmansk Region). — Entomologicheskoe 
Obozrenie, 45(3): 565-569. 1966. 

Val'kh.S.B. K poznaniyu fauny Culicidae vostoka Ukrainy (On the Fauna 
of Culicidae in the Eastern Ukraine). — Meditsinskaya Parazito- 
logiya i Parazitarnye Bolezni, Vol. 28(6): 687-695. 1959. 

Velichkevich, A.I. K faune i ekologii komarov Yuzhnogo berega Kryma 
(On the Fauna and Ecology of Mosquitoes of the Southern Coast of 
the Crimea). — Parazitologicheskii Sbornik Zoologicheskogo 
Instituta AN SSSR, Vol.6: 137-145. 1936. 

Vinogradova, E. B. Morfologicheskaya i biologicheskaya kharakteristika 
nekotorykh prirodnykh populyatsii krovososushchikh komarov 
Culex pipiens L. (Diptera, Culicidae) na territorii SSSR 
(Morphological and Biological Characteristics of Some Natural 
Populations of Culex pipiens L. (Diptera, Culicidae) in the 
USSR). — In Sbornik: "Ekologiya vrednykh nasekomykh i 
entomofagov"; Trudy Zoologicheskogo Instituta AN SSSR, Vol. 36: 
31-57. 1965. 

Vinogradova, E.B. Krovososushchie komary kompleksa Culex 

pipiens L. (Diptera, Culicidae), ikh prakticheskoe znachenie, 
sistematika i biologiya (Mosquitoes of the Culex pipiens L. 
Complex (Diptera, Culicidae), their Practical Importance, 
Systematics and Biology). — Entomologicheskoe Obozrenie, 
45 (2): 241-257. 1966. 

Vinogradskaya,O.N. Stigmal'nyi apparat A n op h e 1 e s i ego vido- 
izmeneniya v zavisimosti ot sukhoustoichivosti i vlagolyubivosti 
vidov (The Spiracular Apparatus of Anopheles and its Variation 
in Relation to the Tolerance of Aridity and Hygrophily of Species). — 
Ibid., 31(1/2): 151-154. 1950. 

Vinogradskaya, O. N. Uchastie trakheinoi sistemy v isparenii vody u 
Anopheles maculipennis messeae Fall, i sezonnaya 
izmenchivost' dykhal'tsevogo indeksa u vidov podsemeistva 
Culicinae (Diptera, Culicidae) (Participation of the Tracheal System 
in the Evaporation of Water in Anopheles maculipennis 
messeae Fall, and the Seasonal Variation of the Spiracular 
Index in Species of Culicinae (Diptera, Culicidae)). — Ibid., Vol.33: 
157-160. 1953. 

Volkova,M. I. Fauna krovososushchikh komarov (sem. Culicidae) doliny 
srednei Volgi v predelakh Tatarskoi ASSR (The Fauna of Mosquitoes 
(Culicidae) of the Middle Volga Valley in the Tatar ASSR). - 
Uchenye Zapiski Kazanskogo Gosudarstvennogo Universiteta, 
116 (5): 153-156. 1956. 



65 



V o 1 ' f t r u b , A. M. K voprosu o vidovom sostave i dinamike aktivnosti 

krovososushchikh dvukrylykh nasekomykh v Taimyrskom natsional'- 
nom okruge (On the Specific Composition and Dynamics of Activity 
of Bloodsucking Insects in the Taimyr Region). - Trudy Nauchno- 
Issledovatel'skogo Instituta Sel'skogo Khozyaistva Krainego Severa, 
Vol. 11: 113-116. 1963. 

Zdrodovskii,P.F. Malyariya na Mugani. Epidemiologiya i plan bor'by 
(Malaria in the Mugan Steppe. Epidemiology and Control). Moskva- 
Baku. 1926. 174 pp. 

Z hogole v , D. T. Svetovye lovushki kak metod sobiraniya i izucheniya 

nasekomykh -perenoschikov vozbuditelei boleznei (Light Traps for 
the Collection and Study of Insects - Vectors of Pathogenic Agents). 
Entomologicheskoe Obozrenie, 38(4): 766 — 773. 1959. 

Zhogolev.D. T. and V. P. Shcherbina. Novaya model' portativnoi 
svetovoi lovushki dlya sbora krovososushchikh dvukrylykh 
nasekomykh (A New Model of Portable Light Trap for the Collection 
of Bloodsucking Insects). - Meditsinskaya Parazitologiya i 
Parazitarnye Bolezni, 35(5): 619-621. 1966. 



Publications in Other Languages 

Bates, M. The Nomenclature and Taxonomic Status of the Mosquitoes of 
the Anopheles maculipennis Complex. - Ann. Ent. Soc. Am., 

Vol. 33: 343-356. 1940. 
Buonomini.G. and M. Mariani. World Anophelines Belonging to the 

Subgenus Maculipennia Buonomini e Mariani, 1946. — Riv. 

Malar., Vol.32: 173-188. 1953. 
C a 1 1 o t , J. and J. A. R i o u x . Liste sommaire des Culicides de France. - 

Annls. Parasit.hum. comp., 40(2): 242-245. 1965. 
Colless,D. H. Notes on the Culicine Mosquitoes of Singapore. IV. The 

Aedes niveus Subgroup (Diptera, Culicidae) : Introduction and 

Description of Five New Species and of One New Subspecies. - 

Ann.Trop. Med. Parasit., 52(4): 468-483. 1958. 
C o 1 u z z i , M. and C. C o n t i n i . The Larva and Pupa of Mansonia 

(Coquillettidia) buxtoni (Edwards), 192 3 (Diptera, Culicidae). - 

Bull. Ent. Res., 53 (2) : 2 15-2 18. 1962. 
Corbet, P. S. Reproduction in Mosquitoes of the High Arctic. - Proc. 

XII Intern. Congress of Entomology, pp. 817—818. 1965. 
Dabrowska,P. E. Communities of Mosquitoes in Three Types of Forest 

Land. - Ekol.polska.A, 12(36): 737-783. 1964. 
Dodge.H.R. Supergeneric Groups of Mosquitoes. - Mosquito News, 

22(4): 365-368. 1962. 
Edwards, F.W. Oligocene Mosquitoes in the British Museum, with a 

Summary of Our Present Knowledge Concerning Fossil Culicidae. - 

Q. Jl.Geol. Soc. Lond., Vol. 79: 139-155. 1923. 
Feng, L. C. The Tree Hole Species of Mosquitoes of Peiping, China. - 

' China Med. J., Suppl., Vol.2: 503-525. 1938. 
Gor don, R. M. and W. H. R. Lumsden . A Study of the Behavior of the 

Mouthparts of Mosquitoes When Taking up Blood from Living Tissue 

Together with Some Observations on the Ingestion of Microfilariae. - 

Ann. Trop. Med. Parasit., Vol. 33 : 259-278. 1939. 



66 



Jobling,B. On Two Subspecies of Cule x pipiens. - Trans. R. Ent. 

Soc. Lond., 87(8): 246-248. 1938. 
Krama'r,J. Culex territans v CSR. - Cslka. Parasit., Vol.4: 

201-204. 1957. 
Mariani.M., C. Eruno-Smiraglia, and N. Caravaglios. 

Ricerche sulla interspecificita labranchiae -atroparvus. - 

Riv. Malar., 43(1/3): 37-50. 1964. 
Martini.E. Beitraige zur medizinischen Entomologie und zur Malaria - 

Epidemiologie des unteren Wolgagebiets. — Abh. Geb. Auslandsk., 

(Die Stechmuckenfauna), Vol.29: 21-69. 1928. 
Mattingly,P. F. A Change of Name among the British Mosquitoes 

(Diptera, Culicidae). - Proc. R. Ent. Soc. Lond., B, Vol. 22 : 106-108. 

1953. 
Mat t ing ly , P. F. Le sous-genre Neoculex (Diptera, Culicidae) dans 

la sous-region mediterraneenne. I. Espece, sous-espece et 

synonymie nouvelles. — Annls. Parasit. hum. comp., Vol. 30: 

375-388. 1955. 
Matt ingly, P. F., L. E. Rozeboom, K. L. Knight, P. G. Shute, 

F. H. Drummond, and S. R. Christophers. The Culex 

pipiens Complex. - Trans. R. Ent. Soc. Lond., 102(7): 331-342. 

1951. 
P e u s , F. Zur Kenntnis der A e d e s -Arten des deutschen Faunengebietes 

(Diptera, Culicidae). Die Weibchen der A e d e s communis- 

Gruppe. - Konowia, Vol.12: 145-159. 1933. 
R e i d , J. A. The Anopheles hyrcanus Group in South-East Asia 

(Diptera, Culicidae). - Bull. Ent. Res., 44(1): 5-76. 1953. 
Reid,J.A. and K. L. Knight. Classification within the Subgenus 

Anopheles (Diptera, Culicidae). — Ann. Trop. Med. Parasit., 

55 (4): 474-488. 1961. 
Rioux, J.-A. and M. Arnold. Les culicides de Camargue (Etude 

systematique et e'cologique). — Terre Vie, Vol.4: 244—286. 
Risler,H. Das Gehororgan der Mannchen von C u 1 e x pipiens L., 

Aedes aegypti L. und Anopheles stephensi Liston 

(Culicidae). - Zool. Jber. Neapel., Sec. 2, Vol. 74: 478-490. 1955. 
Roubaud,E. and S. Ghelovitch. Observations sur le moustique 

anthropophile mediterraneen du groupe pipiens, Culex 

berbericus Roub. — C. r. hebd. Seanc. Acad. Sci., Paris, 

242 (25): 2900-2903. 1956. 
Seminar on the Ecology, Biology and Control of the Culex pipiens 

Complex. — World Health Organisation (Vector Control), Vol. 125: 

1-217. 1965. 
Senevet,G. and L. Andarelli. Les moustiques de l'Afrique du Nord et 

du bassin me'diterraneen. — Arch. Inst. Pasteur Alger., Vol. 41 : 

115-172. 1963; Vol.42: 94-192. 1964. 
Senior-White, R. Notes on Ceylon Mosquitoes. II. The Larvae of the 

Commoner non-Anopheline Mosquitoes. — Spolia zeylan. , (b), 

Vol. 14: 61-76. 1927. 
Skierska,B. Przegla.d pis'miennictwa dotyczacego komarow (Culicidae) z 

obszardw Polski oraz rejestracja i rejonizacja tych owaddw na 

terenie naszego kraju. — Wiad. parazyt. , 9(6): 579—597. 1963. 



67 



S 1 i f e r , E. H. and S. S. Sekhom. The Fine Structure of the Sense Organs 

of the Antennal Flagellum of the Yellow Fever Mosquito A e d e s 

aegypti (Linnaeus). - J. Morph., Ill ( 1 ) : 49-66. 1962. 
Stackelberg, A. A. A New Species of F inl ay a Theob. (Diptera, 

Culicidae) from Ussuri Land. - Bull. Ent. Res., 34(4): 311. 1943. 
Trpis, M. Typologie der Biotope in der Ostslowakei gemass der 

Gesellschaften von Larven und Imagines der Stechmucken. — 

Entomologicke Problemy, Vol.5: 9—69. Bratislava. 1965a. 
Trpis, M. Verbreitung der Stechmucken (Diptera, Culicidae) in der 

Ostslowakei. - Ibid., pp. 71-100. 1965b. 
Vockeroth, J.R. Specific Characters in Tarsal Claws of Some Species 

of Aedes (Diptera, Culicidae). - Can. Ent., Vol.82: 160-162. 1950. 
Vockeroth, J. R. The Specific Status of Aedes pionips Dyar 

(Diptera, Culicidae). -Ibid., Vol.84: 243-247. 1952. 
Vockeroth, J. R. Notes on Northern Species of Aedes, with 

Descriptions of Two New Species (Diptera, Culicidae). — Ibid., 

Vol 86: 109-116. 1954. 



C. MOSQUITOES AS VECTORS 
Publications in Russian 

Ananyan, S. A. Novye dlya SSSR, peredayushchiesya komarami, vozbudi- 
teli virusnykh infektsii (New Causative Agents of Infections 
Transmitted by Mosquitoes in the USSR). — In Sbornik: "Aktual'nye 
problemy virusnykh infektsii"; Materialy XII nauchnoi sessii 
Instituta poliomielita i virusnykh entsefalitov, pp. 194—195. 
Moskva. 1965. 

Bardos, V. and E. Cupkova. Virus Calovo — vtoroi arbovirus, 

. V 

vydelennyi iz komarov v Chekhoslovakii (Calovo Virus — the 
Second Arbovirus Isolated from Mosquitoes in Czechoslovakia). — 
Zhurnal Gigieny, Epidemiologii, Mikrobiologii, Immunologii, 
Vol. 6: 255-260. Praha. 1962. 

Bardos, V. and V. Danielova. Virus Tahyna — virus, vydelennyi iz 

komarov v Chekhoslovakii (Tahyna Virus, Isolated from Mosquitoes 
in Czechoslovakia). -Ibid., Vol. 3 (2): 182-193. Praha. 1959. 

Eeklemishev, V.N. O faktorakh, opredelyayushchikh znachenie 
razlichnykh vidov Anopheles kak perenoschikov malyarii 
(Factors Determining the Importance of Different Species of 
Anopheles as Vectors of Malaria). — Meditsinskaya Para- 
zitologiya i Parazitarnye Bolezni, 10(1): 5 — 8. 1941. 

Beklemishev, V. N. Ekologiya malyariinogo komara (Anopheles 

maculipennis Mgn. ) (The Ecology of Anopheles maculi- 
pennis Mgn.). -Moskva, Medgiz. 1944. 299pp. 

Detinova,T.S. Metody ustanovleniya vozrastnogo sostava dvukrylykh 
nasekomykh, imeyushchikh meditsinskoe znachenie (Methods for 
the Determination of the Age Composition of Medically Important 
Diptera). — World Health Organisation. Geneva. 1962. 220 pp. 



68 



Glushchenko.P.A., A. V. Gutsevich, and M. S. Dud kin a. 

Issledovanie komarov kak perenoschikov virusa limfotsitarnogo 
khoriomeningita na zapade Ukrainy (A Study of Mosquitoes as 
Vectors of the Virus of Lymphocytic Choriomeningitis in the 
Western Ukraine). - DAN SSSR,113(5): 1181-1183. 1957. 

Gutsevich, A. V. Nasekomye kak perenoschiki virusov — parazitov 

cheloveka i zhivotnykh (insects as Vectors of Viruses — Parasites 
of Man and Animals). - Zoologicheskii Zhurnal, 43(3): 429-442. 
1964. 

Il'enko, V.I., A.N. Sterkhova, N.M. Mirzoeva, K.A. Dyunina, 
and M. G. Akhundov. Materialy po izucheniyu transmissivnykh 
virusnykh infektsii v nekotorykh raionakh Azerbaidzhanskoi SSR 
(A Study of Transmissible Virus Infections in Some Regions of 
Azerbaidzhan). — In Sbornik: "Voprosy epidemiologii, bakteriologii, 
gigieny, parazitologii i virusologii, " pp. 182 — 184. Synopses of 
Reports. Dushanbe. 1962. 

Mirzoeva , N. M., V. I. Il'enko, K. A. Dyunina, and M. G. 

Akhundov. Dannye o serologicheskoi i virusologicheskoi 
razvedke arbovirusov v Kyzylagachskom zapovednike im. S. M. 
Kirova Azerbaidzhanskoi SSR (Data on the Serological and Viro- 
logical Cultivation of Arboviruses in the Kirov Kyzyl -Aga 
Reserve in Azerbaidzhan). — In: Kleshchevoi entsefalit, 
Kemerovskaya kleshchevaya likhoradka, gemorragicheskie 
likhoradki i drugie arbovirusnye infektsii, pp. 205—206. Moskva. 
1964. 

Olsuf ev,N. G. Novye eksperimental'nye dannye po voprosu o roli 
komarov v peredache i khranenii tulyaremiinoi infektsii (New- 
Experimental Data on the Role of Mosquitoes in the Transmission 
and Preservation of Tularemia Infection). — Sbornik rabot, 
posvyashchennyi 30-letiyu nauchnoi deyatel'nosti akademika E. N. 
Pavlovskogo, pp. 176 — 189. Leningrad— Moskva. 1941a. 

Olsuf'ev.N. G. Rol' komarov v peredache tulyaremiinoi infektsii dikim 
gryzunam, ptitsam i domashnim zhivotnym (The Role of Mosquitoes 
in the Transmission of Tularemia to Wild Rodents, Birds and 
Domestic Animals). - Ibid., pp. 190-197. 1941b. 

Olsuf ev, N. G. and D. A. G o 1 o v . Rol' komarov v peredache i 

khranenii tulyaremii (The Role of Mosquitoes in the Transmission 
and Preservation of Tularemia). — Trudy Otdela Parazitologii 
VIEM, Vol. 3: 213-246. 1938. 

Olsuf ev, N. G. and G. P. R u d n e v (editors). Tulyaremiya (Tularemia). — 
Moskva, Medgiz. 196 0. 45 9 pp. 

Orekhov.M. D. Dipetalonematoz verblyudov v Turkmenskoi SSR i mery 
bor'by s nim (Dipetalonemiasis of Camels in Turkmenia and its 
Control). - Veterinariya, Vol.4: 28-30. 1952. 

Osipov, A.N. K vyyavleniyu promezhutochnykh khozyaev vozbuditelya 
setarioza krupnogo rogatogo skota v SSSR (On the Discovery of 
Intermediate Hosts of Setariasis of Cattle in the USSR). — Materialy 
k Nauchnoi Konferentsii Vsesoyuznogo Obshchestva Gel' mint ologov, 
Vol. 4: 181-184. 1965. 



69 



Pavlovskii,E. N., A.K. Sht e in , and P. P. Perfil'ev. Eksperi- 
mental'nyeissledovaniya nad vliyaniem deistvuyushchikh nachal 
komara Culex pipiens na kozhu cheloveka (Experimental 
Study of the Effect of Active Principles of Culex pipiens on 
the Skin of Man). - DAN SSSR, A, Vol. 13: 251-254. 1928. 

Petrishcheva, P. A. Komary — perenoschiki yaponskogo entsefalita 
v Yuzhnom Primor'e (Mosquitoes as Vectors of Japanese 
Encephalitis in the Southern Maritime Territory). — In: 
Parazitologiya Dal'nego Vostoka, pp. 106 — 143. Leningrad, Medgiz. 
1947. 

Reha'cek, J. Kul'tivirovanie razlichnykh virusov v kleshchevykh 
tkanevykh kul'turakh (Cultivation of Viruses in Cultures of 
Tick Tissues). -Actavirol., Prague, 9(4): 332-337. 1965. 

Sazonova,0. N. Krovososushchie komary (Diptera, Culicinae) (Mosquitoes 
(Diptera, Culicinae)). —In: Perenoschiki vozbuditelei prirodno- 
ochagovykh boleznei, pp. 9-63. Moskva, Izd. "Meditsina." 1962. 

Simanin.P. I. K voprosu ob eksperimental'nom zarazhenii komarov 
tropicheskoi malyariei (Experimental Infection of Mosquitoes 
with Tropical Malaria). — Parazitologicheskii Sbornik Zoologiches- 
kogo Muzeya AN SSSR, pp. 37-44. 1930. 

Simkova,A., V. Danielova, and V. Bardos. Eksperimental'nyi 

perenos virusa Tahyna komarami A e d e s vexans (Experimental 
Transmission of Tahyna Virus by A e d e s vexans). —Actavirol., 
Prague, 4(6): 341-347. 1960. 

V ig ov ski i , A. I. and A. V. G u t s e v i c h . Predvaritel'nye rezul'taty 

izucheniya prirodnykh ochagov limfotsitarnogo khoriomeningita na 
zapade Ukrainy (Preliminary Results of the Study of Natural Foci 
of Lymphocytic Choriomeningitis in the Western Ukraine). — 
DAN SSSR, 140(5): 1223-1225. 1961. 



Publications in Other Languages 

B a r d o s , V. and E. Cupkova. The Calovo Virus — the Second Virus 

Isolated from Mosquitoes in Czechoslovakia. — J. Hyg. (Prague), 

6 (2): 186-192. 1962. 
Bates, M. The Natural History of Mosquitoes. New York. 1962. 379 pp. 
Chamberlain, R. W. and W. D. Sudia. Mechanism of Transmission 

of Viruses by Mosquitoes. — Ann. Rev. Entomol., Vol.6: 371—390. 

1961. 
Committee of Experts on Malaria. 10th Report. — World Health 

Organisation, Ser. Tech. Rep., 272: 1-62. Geneva. 1965. 
Cont acos, P. G., J. S. Lunn, G. R. Coatney, J. W. Kilpatrick, 

and F. E. Jones. Quartan-Type Malaria Parasite of New World 

Monkeys Transmissible to Man. — Science, Vol. 142: 676. 1963. 
Danielova', V. Experimentalni studie vztahu viru Tahyna k nekterym 

druhum komaru. — Cslka. Epidem. Mikrobiol. Immunol., ll(3): 

171-174. 1962. 
Devine,T.L., C. E. V e n a r d , and W. C. My s e r . Measurement of 

Salivation by Aedes a e gy p t i (L. ) Feeding on a Living Host. — 

J. Insect Physiol., 11 (3): 347-353. 1965. 



70 



E d e s on, J. F. B. and T.Wilson. The Epidemiology of Filariasis Due 

toWuchereria bancrofti and Erugia malayi. —Ann. 

Rev. Entomol., Vol.9: 245-268. 1964. 
Ferreira.E. Distribuicao geografica dos anofelinos no Brazil e sua 

relapao com o estado atual de erradicacao de malaria. — Revta. 

bras. Malar. Doenc. trop., 16(3): 329-348. 1964. 
Foote.R.H. and D.R.Cook. Mosquitoes of Medical Importance. - 

Agric. Handb. (U.S. Dept. Agric. ), Vol.152: 1-158. Washington. 

1959. 
Gordon Smith, C. E. Factors Influencing the Behavior of Viruses in 

their Arthropodan Hosts. "Host - Parasite Relationships in 

Invertebrate Hosts." Oxford. 1964. 31 pp. 
Hannoun.C, R. Panthier, J. Mouchet, and J. P. Eouzan. 

Isolement en France du virus West-Nile a partir de malades 

et du vecteur Culex m o d e s t u s Ficalbi. - C. r. hebd. Seanc. 

Acad. Sci., Paris, Vol.259: 4170-4172. 1964. 
Hawking, F. and M. J. Worms. Transmission of Filaroid Nematodes. — 

Ann. Rev. Entomol., Vol.6: 413-432. 1961. 
Hecht.O. Die Hautreaktionen auf Insektenstiche als allergische 

Erscheinungen. — Arch. Schiffs. - u. Tropenhyg., Vol. 33: 

364-366. 1929. 
H or s f al 1, W. R. Mosquitoes. Their Bionomics and Relation to Disease. 

New York. 1955. 723 pp. 
Huf f, C. G. Susceptibility of Mosquitoes to Avian Malaria. — Expl. Parasit., 

16 (1): 107-132. 1965. 
J ay ewardene, L. G. Larval Development of Erugia ceylonensis 

Jayewardene, 1962, in Aedes ae g y pt i, with a Brief Comparison 

of the Infective Larva with those of B r ug i a spp., Dirofilaria 

repens and Artionema digitata. — Ann. Trop. Med. Parasit., 

57 (3): 359-370. 1963. 
Jordan, H. B. Lizard Malaria in Georgia. — J. Protozool., 11(4): 

562-566. 1964. 
Lavoipierre, M. M. J. Studies on the Host -Parasite Relationships of 

Filarial Nematodes and their Arthropod Hosts. II. The Arthropod 

as a Host to the Nematode: A Brief Appraisal of Our Present 

Knowledge Based on a Study of the More Important Literature from 

1878 to 1957. -Ann. Trop. Med. Parasit. ,52 (3): 326-345. 1958. 
Mattingly, P. F. Ecological Aspects of the Evolution of Mosquito-Borne 

Virus Diseases. - Trans. R. Soc. Trop. Med. Hyg., 54(2): 97-112; 

Discuss., pp. 130-134. 1960. 
Mattingly,P.F. Some Considerations Relating to the Role of Culex 

pipiens fatigans Wiedemann in the Transmission of Human 

Filariasis. - Bull. Wld. Hlth. Org., Vol.27: 569-578. 1962. 
Mattingly, P. F. The Evolution of Parasite — Arthropod Vector 

Systems. — Evolution of Parasites, pp. 29—46. Oxford. 1965. 
Mims,C. A., M. F. Day , and J. D. Ma r s hall . Cytopathic Effect of 

Semliki Forest Virus in the Mosquito Aedes aegypti. — 

Am. J. Trop. Med. Hyg., 15 (5) : 775-784. 1966. 
N e 1 s o n , G. S. Factors Influencing the Development and Behavior of 

Filarial Nematodes in their Arthropodan Hosts. "Host-Parasite 

Relationships in Invertebrate Hosts," pp. 75 — 119. Oxford. 1964. 



71 



Pratt, H. D. Virus Diseases Transmitted by Mosquitoes and Other 

Arthropods. - Mosquito News, 24(2): 91-103. 1964. 
Raffaele, G. Adattamento alio sviluppo nel polio di Plasmodium 

berghei. - Riv. Malar., 44(1/3): 1-8. 1965. 
Reeves, H. C. Mosquitoes and Virus Diseases. — Biological transmission 

of disease agents, pp. 75—82. New York. 1962. 
Reeves.W. C. Ecology of Mosquitoes in Relation to Arboviruses. — Ann. 

Rev. Entomol., Vol. 10 : 25-46. 1965. 
Roy, D.N. Entomology (Medical and Veterinary). Calcutta. 1946. 358 pp. 
R us sell, P. F., L. S. West, and R. D. Ma n w e 1 1 . Practical Malariology. 

Philadelphia. 1946. 684 pp. 
Sandosham,A.A. Malariology with Special Reference to Malaya. 

Singapore. 1959. 327 pp. 
Serie, C, L. Andral, A. Lindres, and P. Neri. Epidemie de 

fievre jaune en Ethiopie (1960-1962). - Bull. Wld. Hlth. Org., 

30(3): 299-319. 1964. 
Singh, K. R. and Kh. M. P a v r i . Isolation of Chittoor Virus from 

Mosquitoes and Demonstration of Serological Conversions in Sera 

of Domestic Animals at Manjri, Poona, India. — Indian J. Med. 

Res., 54 (3): 220-224. 1966. 
Vincke, I.H. Le paludisme des rongeurs en Afrique. — Annls. Soc. beige 

Med. trop., 44(3): 579-586. 1964. 
Viruses Transmitted by Arthropods. — World Health Organization, 

Ser. Tech. Rep., 219: 1-87. Geneva. 1962. 
Warren, M., D. E. E y 1 e s , R. H. W h a r t on , and C. K. Ow Yang. 

The Susceptibility of Malayan Anophelines to Plasmodium 

cynomolgi b a s t i a n e 1 1 i i . — Indian J. Malar., 17 (l ): 

85-105. 1963/ 
W eb be r , W. A. F. Dirofilaria aethiops Webber, 1955, a Filarial 

Parasite of Monkeys. III. The Larval Development in Mosquitoes. — 

Parasitology, Vol. 45 : 388. 1955. 
Wha rt on , R. H., D.E. Eyles, M. W a r r e n , D. E. M o o r h o u s e , and 

A. A. San do sham. Investigations Leading to the Identification 

of Members of the Anopheles umbrosus Group as the 

Probable Vectors of Mouse Deer Malaria. — Bull. Wld. Hlth. Org., 

29(3): 357-374. 1963. 
Wharton, R. H., D.E. Eyles, M. Warren, and W. H. Cheong. 

Studies to Determine the Vectors of Monkey Malaria in Malaya. — 

Ann. Trop. Med. Parasit., 58(1): 56-77. 1964. 

Y oe 1 i , M. Studies on Plasmodium berghei in Nature and under 

Experimental Conditions. — Trans. R. Soc. Trop. Med. Hyg., 59 (3): 
255-271; Discuss., pp. 272-276. 1965. 

Y u c e 1 , A. and R. Deschiens. Depistage d'un foyer de filariose a. 

Wuchereria bancrofti en Turquie Orientale. — Bull. Soc. 
Path, exot., 53(5): 885-891. 1960. 



72 



SYSTEMATIC PART 



The family Culicidae was formerly considered to include the subfamilies 
Culicinae, Dixinae and Chaoborinae. They are at present considered as 
separate families of the Nematocera. Each family is distinctly character- 
ized by important morphological characters of the adult insects and the 
larvae. 



Family CULICIDAE 

Mouth parts forming a proboscis several times longer than the width of 
the head (Figures 1, 2). Antennae with 15 segments, with dense, long hairs 
in the male and sparser, shorter hairs in the female. Wings narrow (Fig- 
ure 6), veins with scales. Costa present at anterior and posterior margin 
of wing; subcosta long, extending to the costa; radius 4-branched, r2+3 
forked, r4+5 simple; cross-vein between ri and r2 absent; media with 3 
branches; cross veins rm and mcu present; cu 1 -branched;* anal vein 
long, extending to margin of wing. 

The larvae differ markedly in external appearance. All thoracic seg- 
ments fused, thorax broader than abdomen. Abdomen consisting of 9 seg- 
ments, segment 8 formed by the fusion of segments 8 and 9; segment 9 is 
morphologically segment 10. There is a posterior pair of spiracles on the 
dorsal side of segment 8 or at the end of the siphon. Some larvae are 
predaceous but the antennae are not modified for capturing the prey. 
Vesicular widenings of the main tracheal trunks, if at all present, only in 
the thorax. 

The family Culicidae is divided into three subfamilies: Anophelinae, 
Toxorhynchitinae and Culicinae. Each of the first two subfamilies is re- 
presented by a single species in the Palaearctic region and the diagnosis 
of the subfamily is therefore the same as that of the genus in the USSR 
(Anopheles Mg. and Toxorhynchites Theob. (Megarhinus Rob.- 
Des.)). They are included with the 6 genera of the subfamily Culicinae in 
the following key. 



Key to genera 

1 (2). Palps of female as long as the proboscis (Figure 28). Last two 
segments of the palps of the male club-shaped, (Figure 29). Ab- 
domen usually without scales, or at least sternites without scales 
over a more or less large area 1. Anopheles Mg. (p. 78 

* [Media 2-branched, cubitus 2-branched according to the usual interpretation.] 



73 




FIGURE 28 . Mouth parts of the female of Anopheles: 

lbr — labrum; mand — mandibles; h — hypopharynx; max — maxillae; 
lb — labium; m.p. — maxillary palps. 

[l). Palps of female short, several times shorter than the proboscis 

(Figure 1). Last two segments of the palps of the male different. 

Tergites and sternites of abdomen covered with scales. 
'4). Proboscis thick, distinctly tapering and curved ventrally in the 

apical half. Scutellum uniformly curved. Large mosquitoes, with 

tufts of bright hairs at the end of the abdomen 

2. Toxorhynchites Theob. (p. 119] 



:?3 




FIGURE 29. Palps of males (from E.N. Pavlovskii): 

A— Anopheles; B — Aedes; C— Culex; D — Culiseta. 



74 



4 (3). Proboscis thin, more or less uniformly thick its entire length or 

slightly tapering and slightly curved in the apical part. Scutellum 
3-lobed (Figure 30). (Subfamily Culicinae.) 



(73) 




FIGURE 30. Scutellum: 

A — Culicinae; B — Anophelinae. 

5 (6). Anal vein ending in posterior margin of wing at level of branching 

of common stem of r2+3 + 1-4+5 (Figure 31) 

3. Uranotaenia Arr. (p. 127) 



(73; 




FIGURE 31. Wing of Uranotaenia 



73 



(5). Anal vein ending in posterior margin of wing at a more or less 

great distance beyond the point of branching of the common stem 

of T2+3 + r4+5, near middle of wing. 
(8). Spiracular setae present (Figure 4) 

5.Culiseta Felt (Theobaldia N.-Lem.) (p.135) 

(7). Spiracular setae absent (Figure 4). 
) 1 0). Postspiracular setae present. Claws of female usually with 

denticles. Cerci of female more or less projecting (Figure 11). . 

7. Aedes Mg. (p. 174). 

(9). Postspiracular setae absent. Claws of female simple, without 

denticles. Cerci of females not projecting. 
[12). First segment of fore tarsi longer than the next 4 tarsal segments 

together; 4th segment of fore tarsi reduced in both sexes 

4. Orthopodomyia Theob. (p. 131). 

12 (11). First segment of fore tarsi not longer than next 4 tarsal segments 

together; 4th segment of fore tarsi of females not reduced. 

13 (14). Pulvilli present (Figure 5). Wing scales narrow 

8. Culex L. (p. 338). 

14 (13). Pulvilli absent. Wing scales usually broad 

6. Mansonia Blanch, (p. 16 7). 



10 



11 



75 



Remarks. Determination is facilitated by the fact that the 3 genera 
are represented by only one species in Russia. These species are dis- 
tinguished by their characteristic color : Uranotaenia unguiculata: 
light blue lateral stripes on the thorax, Orthopodomyia pulchri- 
p alp is: narrow, white longitudinal stripes on a background of dark, brown 
scales on the mesonotum, and Toxorhynchites christophi: tufts of 
long, brightly colored hairs in posterior part of abdomen. 



74 Males 

The structure of the hypopygium of males is characteristic for each 
species; the generic differences are less distinct. The key to genera 
usually does not include the structure of the hypopygium. The present key 
is only provisional. The genus Toxorhynchites is omitted in the key 
because the hypopygium of T. christophi has not been described. 

1 (2). Coxite without lobes. Style longer than coxite, rarely as long as 

coxite 1. Anopheles Mg. 

2 (l ). Coxite of most species (but not of all species) with 1 or 2 lobes. 

Style shorter than coxite, rarely nearly as long. 

3 (4). Coxites situated close together. Coxite with one subapical lobe 

with spines and setae, rarely closely beyond the middle. Tenth 
sternite with numerous spines or a transverse row of denticles at 
the apex 8. Culex L. 

4 (3). Coxites usually slightly separated. Coxite with 2 lobes (basal and 

apical), if with one lobe, it is situated at the base or in the middle 
of the coxite; lobe rarely absent. Tenth sternite without spines 
or a row of denticles at the apex (often with 1 — 3 large, sclerotized 
denticles). 

5 (6). Apical appendage of style longer than width of style (Figure 7); 

if the appendage is not longer than the width of the style, the ap- 
pendage is situated before the apex of the style; if an appendage 
is absent, the style is divided at the base 7. Aedes Mg. 

6 (5). Appendage of style short, usually shorter than the width of the 

style. 

7 (8). Lobe of coxite with one thick, strongly sclerotized, blunt spine 

(Figure 93) 6. Mansonia Blanch. 

8 (7). Lobe of coxite, if present, at least with 2 pointed spines or strong 

setae. 

9 (10). Coxite markedly thickened, short, with a small, flat lobe. Style 

broad, flattened (Figure 64) 3. Uranotaenia Arr. 

10 (9). Coxite not markedly thickened, more or less oblong, with a conical 

lobe, rarely without such a lobe. Style narrow. 

11 (12). Appendage of style about as long as the width of the style in its 

widest part (Figure 68) 4. Orthopodomyia Theob. 

12 (ll). Appendage shorter than the width of the style in its widest part 

(Figures 71 and 74); there may be 2 appendages at the apex of the 
style (C.longiareolata.) 5. Culiseta Felt. 



76 



Larvae 

1 (2). Spiracles situated on dorsal side of 8th abdominal segment, sur- 

rounded by a stigmal plate (subfamily Anophelinae) 

Anopheles Mg. (p. 78). 

2 (l). Spiracles and stigmal plate situated at end of siphon on dorsal side 

of 8th abdominal segment; siphon at least as long as wide at the 
base. 

3 (4). Lateral lobes of labrum adapted to capture prey, each lobe with 

about 10 strong setae with curved and serrated end (subfamily 
Toxorhynchites Theob. (p. 118). 

4 (3). Lateral lobes of labrum only rarely adapted to capture prey, each 

lobe with at least 30 hairs of different form (subfamily Culicinae). 

5 (6). Siphon and valves of stigmal plate transformed into a piercing 

apparatus Mansonia Blanch, (p. 167). 

6 (5). Siphon and valves of stigmal plate normal. 

7 (14). Siphon with one pair of hair tufts on the posterior surface. 

8 (9). Hair tufts situated at base of siphon Culiseta Felt (p. 135). 

9 (8). Hair tufts situated in middle or near apex of siphon. 

10 (11). Eighth abdominal segment with a chitinized plate at the sides with 

spines at the posterior margin. Denticles of pecten with fine 
spines at the margin Uranotaenia Arr. (p. 12 7). 

11 (10). Only scales in one row or in a spot on sides of 8th abdominal seg- 

ment. If a plate is present, the scales are isolated. Denticles of 
pecten, if present, with awl-shaped, pointed end. 

12 (13). Pecten on siphon absent. On the dorsal side of abdominal seg- 

ments 6—8 are chitinized plates which are larger on the posterior 
segments Orthopodomyia Theob. (p. 131). 

13 (12). Pecten on siphon present. Plates on abdominal segments 6—8 

absent Aedes Mg. (p. 174). 

14 (7). Siphon with several pairs of hair tufts on the posterior and lateral 

surface Culex L. (p. 338). 



I. Subfamily ANOPHELINAE 

Clypeus usually longer than wide. Mandibles and maxillae of females 
well developed. Palps usually as long as proboscis in both sexes, rarely 
slightly shorter in females (in some exotic species). Mesonotum weakly 
convex; scutellum simple, curved (exeept in the tropical genus Chagasia). 
Legs very long and thin, with small, dense scales. Pulvilli absent. Wings 
relatively narrow, often with dark or light spots of scales; basal part of 
r4+5 extending slightly beyond cross-vein r-m toward base of wing, alula 
with narrow scales at the margin. Abdomen with hairs or rarely with hairs 
and scales. Hypopygium simple (Figure 9); coxite short, usually without 
basal lobe; at its position are spines or strong setae of varying number 
(l — 6); anal segment completely membranous or weakly sclerotized; 
aedeagus simple, tubular, with or without terminal leaf-shaped appendages; 
females with one spermatheca. 



77 



The larvae have strongly developed lateral hairs on the body parallel to 
the surface of the water. A feathered type of branching is character- 
istic, a fanlike type absent. Head oblong. There are two pairs of clypeal 
76 hairs at the anterior margin, outer and inner, and behind them a pair of post- 
clypeal hairs. A row of 3 pairs of usually long, feathered hairs is situated 
behind the antennae on the frontoclypeus. There are 2 pairs of frontal hairs 
in the larvae of the tropical genus Chagasia. Antennae short, rod-shaped, 
with a branched, rarely simple hair, and two large, flattened spines and 
sensory hairs at the end. Because of feeding at the surface of the water, 
the head can be rotated through 180° toward both sides, so that the ventral 
side is upward. At the sides of the anterior margin of the thorax are two- 
lobed shoulder organs which can be retracted and hold the larvae at the 
surface film. The hairs on the thorax are arranged in three bands cor- 
responding to the thoracic segments, with 13—14 pairs of hairs in each band. 
The abdomen bears characteristic palmate hairs, one pair on the 7 anterior 
segments. These hairs may be rudimentary on segments 1 — 3. The palmate 
hairs and shoulder organs attach the larvae to the surface film. The 
spiracles are situated on the dorsal side of the 8th abdominal segment and 
are surrounded by the stigmal plate. This is supported by two lateral plates 
with a comb of spines which are connected posteriorly by a chitinized arc. 

The pupae have short, broadly open respiratory funnels; the lateral hairs 
of the abdominal segments are situated at the anterior lateral corner of the 
segment. Caudal paddles with 2 hairs, one at the apex of the medial rib, 
the other, smaller hair, anterior to it. 

Eggs usually with a fringe and air floats. 

The subfamily contains three genera. Only species of the main genus 
Anopheles are represented in the Palaearctic. 



1. Genus Anopheles Meigen 

With the characters of the subfamily; scutellum curved; stem of medial 
fork mi + mj straight. The larvae of Palaearctic species of Anopheles 
are divided into three groups: 1 ) all species of the subgenus Anopheles, 
2) A. (A.) plumb e us and 3) species of the subgenus Myzomyia. The 
characters of these groups are given in the key. The genus Anopheles 
is divided into six subgenera of which four are mainly neotropical. Species 
of the subgenera Anopheles Mg. and Myzomyia Blanch. (Cellia 
Theob.) are represented in the Palaearctic region. 

Key to Species 
Females 

1 (14). Cross -veins and base of forms r 2 + r 3 , va x + m 2 and m 3 + 4 + 

cui covered with dark scales; costa with uniformly dark scales 
or with one or two light spots at the anterior margin. (Subgenus 
Anopheles Mg . ) 

2 (11). Wings with dark scales (yellowish- brown or brown), often with 

denser spots of darker scales but without spots of white scales; 
fore femora thin, cylindrical. 



78 



3 (10). 



4 (5). 



5 (4). 



6 

a 


(?) 

(b) 


b 
7 
8 


(a) 
(6) 
(9) 


9 


(8) 


10 


(3) 


a 


(b) 



b a 





11 


(2 




12 


(13 




a 


(b 




b 


(a 


78 


13 


(12 



14 1 



Wings with more or less uniformly colored scales, without spots 

of darker scales; base of anterior fork of wing (r2+3) usually 

situated slightly closer to base of wing than base of medial fork 

(mi+m 2 ). 

Tuft of narrow, white scales absent on the vertex. Mesonotum 

uniformly brownish-yellow, with moderately long, dark brown 

hairs 1. A. (A.) algeriensis Theob. 

White tuft on vertex distinct. Lateral parts dark; hairs on meso- 
notum light in some parts. 

Apex of proboscis whitish 4. A. (A.) marteri Sen.-Prun. 

Fringe of wing white at the apex 

A. (A.) marteri marteri Sen.-Prun. 

Fringe of wing dark at the apex . . . A. (A.) marteri sogdianus Kesh. 
Proboscis dark at the apex. 

Smaller and darker; body mainly blackish-gray with a leaden tinge. 
Tuft of scales at anterior margin of mesonotum distinct, white . . 

2. A. (A.) plumbeus Steph. 

Larger and paler; body mainly yellowish brown or brown. Tuft 
of scales at anterior margin of mesonotum more weakly developed 

and with a yellowish tinge 3. A. (A.) claviger Mg. 

Wings with spots formed by groups of darker scales; base of 
forks of vein r2+3 and vein mi+2 situated at the same distance from 

base of wing 5. A. (A.) maculipennis Mg. 

Body dark brown; mesonotum with a broad, gray median longi- 
tudinal stripe and dark brown lateral parts; dark spots on wings 

distinct A. (A.) maculipennis maculipennis Mg. 

A. (A.) maculipennis messeae Fall. 

A. (A.) maculipennis melanoon Hack. 

A. (A.) maculipennis labranchiae Fall. 
A. (A.) maculipennis atroparvus van Thiel. 
Body pale yellowish brown; mesonotum more or less pale brown; 

dark spots on wings weak, especially in males 

A. (A.) maculipennis sacharovi Favre. 

Wings with white spots at anterior margin formed by white scales 
or with one white spot; fore femora thickened at the base. 
Costa with one white spot near the apex. Hind femora with dis- 
tinct, subapical white ring. Fore femora slightly thickened at the 

base 6. A. (A.) lindesayi Giles. 

White scales on lower surface of hind femora present in jz of 
length of femur from the base. . . A. (A.) lindesayi lindesayi Giles. 
White scales on lower surface of hind femora present only in /e of 
length of femur from the base ... A. (A.) lindesayi japonicus Yam. 
Costa with 2 pale spots. Hind femora without white, subapical 

ring. Fore femora distinctly thickened at the base 

7. A. (A.) hyrcanus Pall. 

Cross-veins and base of forks r2+ 3 and mi+2 with white scales; 
costa with four or more light spots at the anterior margin. (Sub- 
genus Myzomyia Blanch.). 



79 



15 (16). Abdomen covered with scales on both sides, forming projecting 

tufts. Fifth segment of hind tarsi white. Palps of females with 
upright scales in basal part .... 8. A. (M.) pulcherrimus Theob. 

16 (15). Abdomen with hairs, without scales. Fifth segment of hind tarsi 

dark. Palps of females with adpressed scales 

9. A. (M.) superpictus Grassi. 



Males 

1 (14 

2 (3 

3 (2 

4 (5 

5 (4 



79 



6 


(9 


7 


(8 


8 


(7 


9 


(6 


10 


(13 


11 


(12 


12 


(11 


13 


(10 


14 


(1 


15 


(16 


16 


(15 



4th- s 



1 (2 



Coxite with 2 (rarely 1 or 3) large setae on inner side of base, at 
least one of them situated on a teat-shaped tubercle. (Subgenus 
Anopheles Mg.) 

Coxite with one thick spine on inner side of base 

1. A. (A.) algeriensis Theob. 

Coxite with 2—3 large setae on inner side of base. 

Coxite with 3 large setae, 2 of them branched, on inner side of 

base 3. A. (A.) claviger Mg. 

Coxite with 2 simple (not branched) strong setae on inner side of 

base. 

Setae (spines) of claspettes situated close together but not fused. 

Aedeagus short and broad, without leaf-shaped appendages at the 

apex 2. A. (A.) plumbeus Steph. 

Aedeagus long and narrow, with leaf-shaped appendages at the apex 

5. A. (A.) maculipennis Mg. 

At least some setae (spines) of claspettes fused, forming a trans- 
parent plate. 

Lobes of tergite 9 conical or trapezoidal, short (not high), about as 
high as wide. 
Coxite with thick setae near middle of inner margin. (Wings with 

uniformly dark scales) 4. A. (A.) marteri Sen.-Prun. 

Coxite with a relatively thin seta near middle of inner margin. 
(Wings with spots of white scales) .... 6. A. (A.) lindesayi Giles. 
Lobes of tergite 9 long, 3-4 times longer (higher) than wide .... 

7. A. (A.) hyrcanus Pall. 

Coxite with 4-6 large setae on inner side of base, situated directly 
on surface of coxite, not on a tubercle. (Subgenus Myzomyia 
Blanch.) 

Aedeagus with short, leaf-shaped appendages at the apex 

9. A. (M.) superpictus Grassi. 

Aedeagus without leaf-shaped appendages at the apex 

8. A. (M.) pulcherrimus Theob. 



age larvae 

Frontal hairs short, simple. Lateral hairs on abdominal seg- 
ments 4-6 long, distinctly feathered 

2. A. (A.) plumbeus Steph. 



80 



2 (1). Frontal hairs long, feathered. Lateral hairs on abdominal seg- 

ments 4—6 not feathered, divided into a few branches from near 
the base. 

3 (16). Inner clypeal hairs situated close together. 

4 (ll). Outer clypeal hairs simpie or slightly branched at the end or from 

the middle. 

5 (6). Clypeal hairs with fine secondary feathering from the middle or last 

third. Leaflets of palmate hairs long and narrow. Comb at 

sides of base of stigma plate with 20-25 teeth 

1. A. (A.) algeriensis Theob. 

6 (5). Clypeal hairs smooth, without fine secondary feathering. 

7 (8). Postclypeal hairs with 2-5 branches. Leaflets of palmate hairs 

without terminal filaments, broad, deeply denticulate at the lateral 
margins. Spiracles not connected with central plate. Comb at 

sides of base of stigmal plate with 12—16 teeth 

3. A. (A.) claviger Meig. 

8 (7). Postclypeal hairs simple. Spiracles connected with central plate, 

often by a special process. 

9 (10). Palmate hairs on metathorax weakly developed, with 10—11 leaf- 

lets. Palmate hairs of abdomen ending in a long filament 

4. A. (A.) marteri Sen.-Prun. 

10 (9). These palmate hairs well developed, with 15-20 leaflets which 

become gradually pointed, without a long terminal filament .... 
6. A. (A.) lindesayi Giles. 

11 (4). Outer clypeal hairs branched, dendriform. 

12 (13). Antennal hair long, about half as long as antenna, situated in its 

middle or slightly proximal to middle. . . 7. A. (A.) hyrcanus Pall. 

13 (12). Antennal hair very short, not longer than the width of the antenna, 

situated nearer to the base. 

14 (15). Outer clypeal hairs reaching to anterior margin of lateral lobes 

of labrum or beyond. Stigmal plate smaller (0.38-0. 50mm long, 
width between ends of lateral lobes 0.39-0.52 mm), slightly pig- 
mented; middle of posterior part of central plate pale, alveoli 

and folds of anterior part weakly developed 

A. (A.) maculipennis sacharovi Favre. 

15 (14). Outer clypeal hairs different. Stigmal plate large (0.50-0.57 mm 

long, width between ends of lateral lobes 0.56-0.59 mm), darkly 
pigmented. Middle of anterior part of central plate, alveoli and 
folds of anterior part dark, distinct. . 5. A. (A.) maculipennis Meig. 

16 (3). Inner clypeal hairs widely separated. 

17 (8). Outer clypeal hairs finely branched, with 6—15 branches. Post- 
10 clypeal hairs with 2 — 4 branches, rarely simple. Central plate of 

stigmal plate distinctly tapering in anterior part of median part, 
its middle weakly pigmented with distinct outline. Anterior part 
of central plate in the form of an arrow-head, its lateral pro- 
cesses oblong, processes to spiracles absent 

8. A. (M.) pulcherrimus Theob. 

18 (17). Outer clypeal and postclypeal hairs simple. Central plate broad, 

not tapering in anterior part of median part; sharp outline in 
median part absent. Anterior part of central plate without lateral 



81 



processes, processes to spiracles present, rarely absent 

9. A. (M.) superpictus Grassi. 



1. Subgenus Anopheles Meigen 

Cross-veins and base of forks r2+ 3 and mi+ 2 + m3 with dark scales; 
costa usually with 1 — 2 light spots or without spots. Coxite with 2 (rarely 
1 or 3) large setae on inner side of base, at least one of them situated on a 
teat-shaped tubercle. 

Larvae of the subgenus Anopheles, except A. plumbeus, with inner 
clypeal hairs situated close together, with large, feathered frontal hairs, a 
branched hair on the antenna, and long lateral hairs (No. 6) on abdominal 
segments 4—5, branching from near the base into 2—5 branches, and short 
branches on segment 6. The larvae of A. plumbeus have more widely 
separated inner clypeal hairs, short simple frontal hairs, a simple hair on 
the antennae and long, feathered hairs on abdominal segments 4—6. 

The subgenus Anopheles is widely distributed (more than 100 species, 
10 species in the Palaearctic region). The species occur in all zoogeo- 
graphical regions. The Palaearctic species of Anopheles form two 
groups according to their distribution: the group related to A.maculi- 
pennis is mainly distributed in the temperate zone, far to the north; 
2 ) the other group, related to A.hyrcanus,is mainly distributed in the 
Oriental region and is represented by a number of species. It does not 
extend so far north. 



1. Anopheles (Anopheles) algeriensis Theobald, 1903 

This species differs from the other species of the genus in the absence 
of spots on the wings and the tuft of white scales on the vertex. Thorax 
and abdomen mainly pale brown; mesonotum with dark hairs. Hind tarsi 
sometimes with indistinct, pale rings. 

Hypopygium (Figure 32) very characteristic: at the base of the coxite 
is a large spine situated on a distinct tubercle. The outer spines of the 
basal part of the coxite, which are present in A. claviger, are 1 — 2 not 
constant, weak setae. Near the apex of the inner side of the coxite is a 
large seta. Claspettes 3-lobed, outer lobe with 2—3 large, pointed spines 
situated close together, median lobe covered with hairs, inner lobe with 
3 more or less equal setae. Aedaegus with 2 — 3 pairs of long, thin, leaf- 
shaped appendages at the apex. 

The 4th-stage larva (Figure 33) resembles that of A. claviger. 
Head: inner clypeal hairs long, simple, situated close together or rarely 
slightly branched at the ends, with thin secondary feathering which begins 
from the middle or near the apex; outer clypeal hairs half as long as inner 
hairs, also with secondary feathering, simple or with 2—3 branches at the 
ends. Postclypeal hairs smooth, simple, rarely 2-branched at the end, 
usually reaching to the base of the anterior clypeal hairs. Antennae dark, 
with more numerous, dark spines than in A. claviger. 



82 




FIGURE 32. Hypopygium of Anopheles algeriensis Theob. 




FIGURE 33. Anopheles a 1 g er i en s i s Theob. Fourth- stage larva: 

1 - anterior part of frontoclypeus; 2 - left antenna; 3 - median hairs of prothorax (right side); 4 - stigmal 
plate. 



83 



The palmate hairs of abdominal segments 3—7 consist of 16—18 slightly 
pigmented, lanceolate leaflets with a weakly developed terminal filament 
and distinctly denticulate margin. 

Stigmal plate not pigmented, without distinct alveoli in the middle of the 
central plate, its median part not different from the lateral parts. The 
median part is even usually lighter than the surrounding parts. The an- 
terior process usually does not extend posteriorly beyond the anterior and 
median part of the central plate. The comb on the 8th abdominal segment 
has 7—11 large teeth (the marginal teeth are the longest) and there are 
1 — 3 small spines between them. The spines bear a row of small spines at 
the base of the apical margin. 

Eggs oblong- oval, markedly widened in the middle and with distinct air 
floats but without a fringe. 
82 Distribution. Mainly in the Mediterranean to England in the north. 

The discovery ofA.algeriensis on Sarema Island at the west coast of 
Estonia is interesting (Remm, 1957). In the USSR it occurs in Trans- 
caucasia, the Northern Caucasus and in Middle Asia. It is usually rare. 
A mass appearance has been observed in the lower reaches of rivers, e. g. 
in the lower reaches of the Sulak River in Dagestan (Enikolopov, 1937). 

Biology. The larvae are found in large swamps densely overgrown with 
vegetation (a typical submerged plant in such water bodies is Hippuris 
vulgaris) in small, well shaded water bodies, mainly in stagnant, markedly 
saline water. In Tadzhikistan (Glagoleva, 1944), the larvae are found only in 
springs in swamps with hard, alkaline water. The larvae often occur to- 
gether with larvae of A. hyrcanus and Culex territans. They 
hibernate. The mosquitoes attack man and animals. In Middle Asia 
(Tadzhikistan) they appear mainly in spring and autumn. The mosquitoes 
are usually found in thickets of bulrushes; they rarely enter stables and 
still more rarely human habitations. 



2. Anopheles (Anopheles) plumbeus Stephens, 1828 
(nigripes Staeger, 1839) 

Distinguished by its dark color and slightly smaller size than 
A. claviger. 
83 Occiput with black scales laterally, with a tuft of narrow, white scales on 

the vertex, which is directed anteriorly. Proboscis and palps black. 
Thorax blackish-brown; mesonotum with a broad, median longitudinal stripe 
js as wide as the mesonotum; a tuft of narrow white scales in middle of 
anterior margin of mesonotum; lateral parts of mesonotum and sides of 
thorax blackish-brown; scutellum blackish. Legs black or blackish- brown, 
coxae and base of femora pale. Wings without spots, densely covered with 
dark brown scales; halteres with black head. Abdomen black, shiny 
dorsally, slightly paler ventrally. Hairs of thorax and abdomen pale brown 
with a golden tinge. 

Hypopygium (Figure 34): coxite with 2 strong, simple setae at the base; 
a large seta near middle of inner side of coxite; spines at apex of claspettes 
situated close together but not fused; aedeagus short, broad, without 
appendages. 



84 




FIGURE 34. Hypopygium of A nophel es plumbeus 
Steph. 

Fourth- stage larva (Figure 35): dark gray to dark brown. In 
recently molted specimens, the abdomen is ringed because the pigmentation 
between the segments is weak. 




FIGURE 35. Anopheles plumbeus Steph. Fourth- stage larva: 
1 - head, dorsal; 2 — left antenna; 3 — stigmal plate. 

!4 Head uniformly dark brown, with weakly developed compound eyes. 

Clypeal hairs thin, slightly branched at the ends. They are simple, smooth 
and unbranched in larvae of A. plumbeus var . barianensis. The 
hairs are situated at about the same distance from each other. Postclypeal 



85 



hairs, frontal and other hairs thin, short and simple. Antennae dark, 
straight, without spines, with a short, simple hair near the base. Abdomen: 
palmate hairs present on segments 2—7, short and simple on 1st segment; 
with about 15 leaflets of uniform color, lanceolate, with pointed end but 
without terminal filament; their margin may be slightly denticulate from 
lz from the base. Lateral hairs on abdominal segments 4—6 long and 
feathered. Stigmal plate with dark anterior lobe. Spiracles connected by 
a process, rarely directly connected with anterior part of central plate. 
This has the form of an onion, with a narrow dark anterior process. The 
granulation of the median part of the central plate is dark and sharply 
defined from the lighter lateral parts. 

Eggs oblong- oval, with a wide fringe but without air floats. 

Distribution. It is mainly distributed in the Mediterranean; in 
Western Europe to England and southern Sweden in the north and to Estonia 
in the USSR (Remm, 1957). It occurs in the Ukraine (rare), Northern 
Caucasus, Transcaucasia, Turkmenia (forests of Kopet-Dagh) and 
Tadzhikistan. 

The Anopheles in India and Pakistan is considered by most authors 
as the species A. barianensis James, 1911, but there are no distinct 
differences between A. plumbeus and A. barianensis. The adults 
and larvae from Tadzhikistan examined are typical A. plumbeus. 

In the southern parts of its range the species occurs only in broadleaved 
forests in mountains and valleys to an altitude of 1,600— 2,000 m. 

Biology. The larvae live in water in tree holes (of birch, elm, oak, 
ash in the Ukraine; hornbeam, oak, chestnut, maple, walnut, ash and fruit 
trees in the Crimea; beech, oak, chestnut, maple, walnut, Tartar maple, 
Oriental plane, maple, apple in the Caucasus; walnut, mulberry, Oriental 
plane tree in Tadzhikistan), mainly in shaded areas. Larvae were found in 
Abkhazia in shaded, artificial water bodies (abandoned pits, basins of sewage 
systems, etc . ). They were found together with larvae ofAedes geni- 
culatus, A. pulchritarsis, Orthopodomyia pulchripalpis, 
and also with larvae of My i at r op a f lore a L. (Syrphidae). The water 
in the breeding places of A. plumbeus, according to studies in Tad- 
zhikistan (Glagoleva, 1946), is distinctly alkaline, with a high concentration 
of salts of alkaline metals (mainly potassium and sodium), in addition to 
humic and ulmic acid, tannin, pigments, etc. extracted from the tree holes; 
the water is brown. The eggs and larvae of all stages hibernate. They 
!5 survive freezing of the water and the eggs are able to withstand long periods 
of frost. This is a typical forest insect which rarely enters buildings or 
bites man. 

Plasmodia of malaria may complete a cycle of sporogony, but the 
importance of this species as a vector is very small because of its ecology. 
It was of epidemiological importance in the past in forests in Caucasian resorts. 

56 3. Anopheles (Anopheles )claviger Meigen, 1804 
(bifurcatus auct.)* (Figure 36) 

Yellowish brown in its greater part. Head with narrow dark brown 
scales laterally, with narrow white scales on the occiput. Vertex with an 

* Linnaeus described the male of Culex pipiens as "Culex bifurcatu s." The name Anopheles 
bifurcatus L. should therefore be replaced with A.claviger Mg. 



86 




FIGURE 36. Anopheles claviger Mg. 



87 



anteriorly directed tuft of cream-colored scales and hairs; proboscis and 
palps uniformly dark brown; antennae brown, basal segments lighter; hair 
on the antennae branched. Mesonotum brown, with a broad, median, an- 
teriorly tapering, whitish gray stripe about /s as wide as the mesonotum, 
with a tuft of yellowish white scales in the middle anteriorly; hairs on 
mesonotum whitish yellow in the middle, brownish laterally; scutellum 
brown, often pale brown, with brownish setae at the posterior margin; 
pleurae of thorax brown, with weak tomentum. Legs brown or dark brown, 
paler on ventral side of femora and tibiae, and at the articulations; tarsi 
dark brown. Wings transparent, veins with brown scales; halteres with 
dark brown head. Abdomen brown, darker at posterior margin of tergites, 
with long, pale brown hairs. 




FIGURE 37. Hypopygium of Anopheles clavigerMg. 



Hypopygium (Figure 3 7) with one strong seta near the apex on the inner 
side of the coxite. Three large strong setae at base of coxite, the outer 2 
setae situated close together and branched in their apical half; inner seta 
simple, situated on a tubercle. 

Fourth-stage larva (Figure 38) large and dark, with strongly pig- 
mented head and stigmal plate. 

Inner clypeal hairs long, nearly as long as the antenna, simple or with 
2 — 4 branches at the apex, without secondary feathering, situated close 
together. Outer clypeal hairs shorter than inner hairs, smooth, rarely 
simple, often with 2—4 branches from the middle or in the last third. 



Postclypeal hairs short, thin, with 2—5 branches nearly from the base. 
Antenna half as long as the head, with sparse spines on the inside, hair 
short, with 4—7 branches, situated near the base. 




FIGURE 38. Anopheles claviger Mg. Fourth-stage larva: 

1 — anterior part of frontoclypeus (outer clypeal hair branched at the end at left); 2 — left antenna; 3 — 
stigmal plate. 



Palmate hairs well developed on abdominal segments 3—7; leaflets 
lanceolate, with slightly elongate apex, but without terminal filament, with 
smooth or slightly denticulate, weakly but uniformly pigmented. Lateral 
hairs of segments 1 — 3 feathered, branched on segments 4—6. 

Stigmal plate with entirely dark anterior lobe, without connection with 
the spiracles in anterior part of central plate, triangular with the apex 
anteriorly and distinctly pigmented in the middle, entirely dark in the 
anterior and posterior parts and alveolate, with dark outline in the median 
part. The stigmal plate of 2nd- and 3rd-stage larvae of A., algeriensis 
and A. hyrcanus and of 2nd-stage A.maculipennis is similar. 

Comb on 8th abdominal segment with 12—14 large and medium-sized 
spines alternating with smaller spines. 

Eggs oblong- oval, with a large number of air floats and a simple fringe, 
not transversely striated, present only at the ends of the egg. 

Note on systematics. Several subspecies of A. claviger have 
been described; one of them in the Mediterranean, A. petragnani Del 
Vecchio, 1939, should be considered as a distinct species. It differs in 
details of the chaetotaxy of the larva, and in the structure of the egg and pupa. 



89 



A. claviger and A. petragnani do not cross or produce viable off- 
spring but further information is necessary to determine the status of 
A. petragnani. 

Distribution. European USSR to the Leningrad Region (60°N). Wide- 
spread in the Crimea and Caucasus (Northern Caucasus, Dagestan, Georgia, 
Armenia, Azerbaidzhan); Middle Asia (Turkmenia, Uzbekistan, Tadzhikistan, 
Kirghizia); to 2,000m altitude in mountains of Middle Asia; Kazakhstan, and 
West Siberia to Tomsk in the east. Europe, from central Sweden 
and Norway to Italy; Southwest Asia (Palestine, Asia Minor, Iraq, Iran); 
North Africa. 

Biology. The larvae breed in water slightly warmed by the sun, in 
moderately or weakly mineralized water fed by springs, in stagnant (swamps) 
and flowing water, shaded by trees, shrubs, rocks, etc. They breed in the 
mountains in backwaters of streams or in springs, in irrigation canals fed 
by springs. In some mountainous and foothill localities, they are some- 
times found in or around populated areas. The mosquitoes often enter 
houses and stables. Larvae of the 3rd and 4th stage hibernate in water that 
does not freeze at the bottom. In southern parts of their area of distribution 
their development does not cease in winter but it becomes very slow. The 
larvae are very agile and descend to the bottom at the slightest disturbance. 
The adults spend their days among plants and enter buildings in spring. 
They usually bite in the open air. The females of some populations of 
A. claviger may have an autogenous first batch of eggs without sucking 
blood (Markovich, 1941). 

A. claviger is a potential vector of malaria of man. Its epidemio- 
logical importance in most of its range is not large, but in localities where 
the larvae breed in populated areas it may be an important carrier. Natural 
infection of A. claviger with tularemia has been established. 



4. Anopheles (Anopheles) marteri Senevet 
and Prunelle, 1927 

This species resembles A. claviger, but differs from it in the whitish 
apex of the proboscis (female) and in the hypopygium, which has two strong, 
simple, not feathered setae at the base of the coxite. The structure of the 
hypopygium resembles that of A. maculipennis but differs in the 
appendages of the claspettes, which form narrow plates. There are only 
setae on the claspettes in A. maculipennis or the setae are fused into 
a narrow plate only in the dorsal part of the claspettes. 

Occiput with whitish scales in the anterior part; long, hair like scales 
between the eyes. Proboscis with whitish apex, without pale rings; white 
scales at the base of the palps. Palps of male about as long as proboscis. 
Mesonotum with a whitish median stripe only in its anterior part ( /2 — lz) 
and two dark stripes laterally, with numerous narrow, brown or golden 
scales which are lighter and wider on the median longitudinal stripe. The 
dark stripes of the mesonotum are without scales or hairs in the anterior 
part. Scutellum with longer setae in the lateral parts. Legs: coxae 
yellowish white with pale yellow hairs; femora, tibiae and tarsi blackish 
brown with a bluish tinge; tarsi without rings. Wings without spots, with 
dense, dark scales; fringe at apex of wing white. 



90 




FIGURE 39. Hypopygium of Anopheles marteri Sen.-Prun. 



Hypopygium (Figure 3 9): coxite with 2 strong, spinelike setae in the basal 
part, medium seta thick, curved at the end and one strong seta in the apical 
third of the inner side and numerous small hairs on the dorsal surface. 
Claspettes with 2 lobes which are sometimes indistinctly divided. Dorsal 
lobe of claspettes with 3 narrow, plate-shaped processes. Ventral lobe with 
a plate which is wider toward the apex. Aedeagus with long, leaf-shaped 
appendages. 

Fourth-stage 1 a r v a resembling that of A . 1 i n d e s ay i. Clypeal 
hairs simple, inner hairs situated close together, more than twice as long 
as the outer hairs; clypeal hairs as long as outer clypeal hairs or longer. 
Frontal hairs feathered, with 6-7 pairs of lateral branches, slightly shorter 
than the inner clypeal hairs; ends of inner frontal hairs hardly reaching 
base of postclypeal hairs. Antennae with dark apex in / 5 of their length. 
Hair of antenna with 3-4 moderately long branches near the middle, slightly 
proximal to it. 

Hair No. 1 on metathorax forming a palmate hair with 10-11 leaflets 
which are wider at the base and taper apically with sharp steps to a 
terminal filament. Palmate hairs well developed on abdominal segments 
2-7, with 15-19 elongate, narrow leaflets. They pass from the middle in 
2-3 irregular steps into the terminal filament, which is fa as long as the 
leaflet. 

Stigmal plate with an anterior lobe which is darkly pigmented at the base 
and lighter at the anterior margin; spiracles connected with anterior part 
of central plate by processes. Central plate very broad in the middle, 
granulation in middle lighter. 

Distribution. Mediterranean: Corsica, Balkans, Algeria, Palestine. 
The subspecies A. marteri sogdianus Kesh. occurs in Middle Asia 
and Northern Iran. The species occurs in the USSR only in the mountains 
of Tadzhikistan (Keshish'yan, 1938). 



91 



Anopheles marteri sogdianus Keshishian, 1938 

It differs from the nominate subspecies in the absence of a white spot 
at the apex of the fringe of the wing. It was at first described as a distinct 
species from Tadzhikistan (Keshish'yan, 1938). 

Distribution. USSR: mountain regions of Tadzhikistan at altitudes 
above 1,000m. xVorthwest Iran. 

Biology. Larvae are found in large numbers in spring (April- June) 
in shaded rock pools in mountain rivers, streams and springs, often together 
with larvae of A. claviger and A.lindesayi. Its ecology resembles 
that of larvae of A. marteri. 



5. Anopheles maculipennis Meigen, 1818 (Figure 40) 

The classification of the "maculipennis complex" is one of the most 
difficult and controversial in the systematics of mosquitoes. A number of 
"races" of maculipennis have been described and later considered as 
subspecies. The following six forms occur in the Palaearctic: maculi- 
pennis Mg., messeae Fall., melanoon Hack, (subalpinus Hackett 
and Lewis), labranchiae Fall., atroparvus van Thiel, sacharovi 
Favre. The last was described as a distinct species. In a monograph of 
A. maculipennis (Beklemishev, 1944), all these forms are considered 
as subspecies. 

It was later suggested that these subspecies be raised to the rank of 
species (Bates, 1940). This was accepted by some authors, but not generally. It 
was stressed that some forms of the maculipennis complex were more 
closely related to each other than to other forms. The groups of the 
complex were suggested as species. This "compromise" was accepted by 
many authors, e.g. Stone et al. (1959). Instead of the one species 
A. maculipennis in the Palaearctic, three species have been accepted: 
maculipennis (with the subspecies messeae and melanoon), 
labranchiae (with the subspecies a t r op a r vu s ) and sacharovi. 
The North American A.occidentalis D.K. is also considered as a 
distinct species. These four species plus several related species form the 
group maculipennis, which is considered as one of the six groups into 
which the subgenus Anopheles was divided (Reid and Knight, 1961; this 

91 division of Anopheles into six groups was previously suggested by 
Edwards, 1932). It was also suggested that this group should form the sub- 
genus Maculipennia (Buonomini and Mariani, 1953), but this has not been 
accepted. 

Those in favor of dividing maculipennis into several species state 
that they can be recognized by the structure of the dorsal surface of the 
eggs and by important biological differences (see below). It has been 

92 emphasized that the forms do not cross and that there is therefore repro- 
ductive isolation in their common habitat (2-3 forms often occur in the 
same locality). 

In our opinion the following considerations should be taken into account 
in the classification of the maculipennis complex. 



610144 92 



91 




FIGURE 40. Anopheles maculipennis Mg 



The classification of mosquitoes is based mainly on the aggregate of 
characters of the adult mosquitoes (female and male) and the larvae. We 
therefore give three keys for each species. All Palaearctic species of 
mosquitoes are determined by at least one of these three "phases." This 
principle gives a firm support to classification, and we see no reason to 
reject it. 



93 



No reliable differences have been found so far between forms of the 
complex, between the larvae or the imago or, particularly, the structure of 
the hypopygium. Determination is possible only according to oviposition. 
The difference between sacharovi and the other forms of the complex 
are very small and at most of the rank of subspecies. Differences in the 
structure of the hypopygium, mainly in the form of the spines on the 
claspettes, have also been described. A study of the armature of the 
claspettes of different forms of A. maculipennis has shown that in- 
dividuals cannot be identified by this character and that there is variation 
of the character in all subspecies (Denisova, 1940, 1948, 1964). 

As to biological characters, important intraspecific differences are 
present in many species. Thus, in many species autogenous and anauto- 
genous populations have been found. These are apparently intraspecific 
groups, biotypes, or perhaps subspecies. If other polymorphic, widely 
distributed species had been studied as extensively as A. maculipennis, 
the same heterogeneity would probably have been found and there would be 
the problem of the division of many species; but this would not be justfied 
in our opinion. 

Strict reproductive isolation between species of the maculipennis 
complex is based mainly on experiments with atroparvus, which is 
easily bred in the laboratory. It is not known how complete and universal 
is the reproductive isolation of the forms of the complex. Experimental 
crossing of subspecies has not always given the same results. The pro- 
duction of fertile progeny depends on many factors. Thus, in experiments 
in hybridization of labranchiae X atroparvus, the result depended 
on whether the mosquitoes were captured wild or bred in the laboratory 
(Mariani et al., 1964). In other species, e. g. Culex p i p i e n s, at least 
partial crossing between different subspecies and even populations is known. 

The partial or even complete reproductive isolation of subspecies proves 
only the developing divergence which, however, has not advanced sufficiently 
to result in the formation of new species. Only sacharovi stands on the 
verge of transformation into a distinct species. 

We consider, therefore, A. maculipennis as a single, polytypic 
93 species with 6 Palaearctic subspecies: maculipennis, messeae, 

melanoon, labranchiae, atroparvus and sacharovi. The latter 
is more differentiated in some respects. Only this species is listed 
separately in the keys of females and larvae. We think that species status 
of these forms disturbs the continuity and proportion in the classification 
of the subgenus Anopheles since the differences between the forms are 
of a different character and smaller than the differences between other 
species of the subgenus Anopheles of the Palaearctic fauna. The 
taxonomic rank of the varieties of A. maculipennis is of no basic 
importance since, as Darwin stated, a variety is a developing species. 

We give below the description of A. maculipennis including all 
Palaearctic subspecies except A. m. sacharovi, followed by the dis- 
tinctive characters of the subspecies. The more important morphological 
and biological characters of the subspecies are compared in the table (see 
P. 98). 

Anopheles maculipennis varies widely in coloration and size . 
Specimens from southern localities are usually lighter and smaller. 



94 



Head: Occiput with upright, dark brown scales, with a tuft of long, whitish 
scales and hairs directed anteriorly on the vertex. Antennae brown. Palps 
and proboscis brown, mainly with adpressed scales. Mesonotum with a 
broad, median, grayish longitudinal stripe which tapers anteriorly and there 
are usually 2—3 indistinct brownish stripes in its anterior half. At the 
anterior margin of the mesonotum is a tuft of whitish scales. Lateral parts 
of mesonotum brown anteriorly, blackish brown posteriorly. Hairs in median 
part of mesonotum and on the scutellum pale brown, otherwise dark brown, 
especially above the base of the wings. Legs brown on ventral side of 
femora, lighter on inner side of tibiae. Tarsi dark. Wings with brownish 
scales, with 4 spots of blackish brown scales at base of radial and medial 
fork, at cross -veins and in basal part of stem of r2 + r3; scales at margin 
of wing usually pale, yellowish at the apex. Halteres yellow, with brownish 
head. Abdomen brown, sometimes blackish brown, often with lighter spots 
in lateral parts of tergites; abdomen with long, golden yellow or brownish 
hairs. 



94) 




FIGURE 41. Hypopygium of Anopheles maculipennis Mg. 



Hypopygium (Figure 41) with one strong seta near middle of inner side of 
coxite; two large setae situated on tubercles at base of coxite. Spines on 
claspettes situated close together but not fused. The spines vary in form 
(blunt or pointed), and their number also varies. Aedeagus long, narrow, 
with leaf-shaped appendages at the apex. The differences between the 
larvae of the subspecies of A. maculipennis are so small that no 



95 



(95) 




FIGURE 42. Anopheles maculipennis Mg. Fourth- stage larva: 

1 — anterior margin of frontoclypeus; 2 — base of outer clypeal hair (dichotomous branching); 
3 — strong branching of inner clypeal hairs; 4 — stigmal plate. 



96 


















15 



Eggs of subspecies of Anopheles mac uli pe nnis Mg. (after Missiroli, Hackett and Martini): 

1 — A.m.subalpinus (dark form, m el a no on); 2—6 — A.m.messeae; 7—8 - A.m.maculipennis; 
9— 12 — A . m .atroparvus; 13, 14 — A . m .labranchia e; 15 — A . m . sa charo vi. 



97 



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98 



characters have been found which would permit their reliable determination. 
A study of extensive material with variation- statistical methods indicated 
differences in the average number and relative development of the branches 
of some hairs (Nos. 1 — 6, after Martini) on the dorsal side of the abdominal 
segments with marked overlapping of extreme variations in all subspecies, 
so that the determination of single larvae is impossible. We therefore give 
only the description of the larvae of A. maculipennis and of A . m a c u - 
lipennis s a c h a r o vi, in which the differences are sufficiently distinct. 

Fourth-stage larvae (Figure 42 ) are very variable in appearance, 
color and size because of their different habitats and conditions. Larvae 
from northern latitudes are usually larger, with darker chitinized structures 
(head, stigmal plate, etc.). The color varies from grayish yellowish green 
to blackish green, according to the color of the environment and the illumina- 
tion of the water. 

Head with variable pigmentation, from the usual light brown spots to 
blackish brown stripes against a dark background. Clypeal hairs: inner 
hairs situated close together, with 3—8 branches at the end; outer hairs 
dendriform. The branching is variable: the stem is often divided into 
2 branches, which branch dichotomously or dendriform in the horizontal 
plane; the stem has rarely three or more branches which branch further. 
Hairs directed obliquely anteriorly. Outer clypeal hairs not reaching 
anterior margin of lateral lobes of labrum, situated on its base. Postclypeal 
hairs short, slightly branched, with 2-8 branches, extending slightly beyond 
base of anterior hairs. Antennae nearly straight, fa as long as the head, 
97 with darker end; hair short (Figure 48), usually as long as width of antennae 
at its position, situated near the base, with 4-6 thin branches. 

Median hairs of prothorax: outer hairs short and simple, median hairs 
feathered, with 3—7 lateral branches on each side, inner branches nearly 
half as long, slightly branched at the end. Palmate hairs on metathorax 
rudimentary. 

Palmate hairs present on segments 3— 7, with 16—24 leaflets which are 
slightly wider in the middle, which is more strongly pigmented. 
Terminal filament fa of the length of the leaflet. It is often pointed at the 
end, its base passing in irregular steps into the widest part of the leaflet. 

Stigmal plate large: 0.5-0. 57mm long, width between ends of lateral 
valves 0.56— 0.59mm. The chitinized areas are usually strongly pigmented. 
Anterior lobe with dark base, distinctly lighter otherwise. Spiracles with- 
out processes, situated some distance from the central plate. The base of 
the anterior process of the central plate does not reach the border between 
its anterior and median parts, passing into the folds and outlines of the 
alveoli in the median part. This is darkly pigmented from the posterior 
margin to the middle of the posterior part and then passes anteriorly into 
the granulated part which covers the entire median part and the posterior 
half of anterior part. It may be complete or the alveoli diverge to the sides 
in the median part leaving a lighter median area. If the granulation is 
complete, the alveoli of the median part may form a darker stripe than in 
A. claviger. 

The comb on segment 8 consists of 6—10 (usually 7—9) large, awl-shaped 
denticles and between them are 1 — 4 much smaller denticles. 

Lateral hair on saddle long, usually simple. Outer caudal hairs asym- 
metrically branched, their ends curved, inner hairs branched more strongly 



99 



on the dorsal side. Fin with 18-20 tufts. Gills usually as long as the 
saddle, both pairs of the same length. 

The eggs of A. maculip ennis, except those of A.m. sacharovi, 
have air floats which occupy the middle third and a fringe. The structure 
and color of the exochorion differ but are characteristic for the subspecies, 
This depends on changes of the structure of the exochorion and consists of 
dark and light spots and stripes. The differences of this character in the 
various subspecies are given in the table on p. 98 and in the figure on p. 97. 

Distribution. Europe, North Africa, North Asia (we are not includ- 
ing the North American form, which many authors now consider a distinct 
species, A . o c c i d ent ali s D.K.). The most northern record of 
A.maculipennis is from southwest of Murmansk, 69 °N (Solovei and 
Likhoded, 1966). The distribution then extends southeast (Karelia at 66°) 
and east along the Lena River at about the same latitude in river valleys and 
then further south along water divides and especially in mountains. From 
Yakutsk the border of distribution, which is not well known in this area, extends 
south and crosses the Amur valley about 150 km east of Blagoveshchensk. 
98 The area of distribution also includes Northeast China, Mongolia, Sinkiang, 
Northern Afghanistan and Iran near Transbaikalia, Southwest Asia and some 
parts of North Africa. 

Biology. There is a large literature on the ecology of A. maculi- 
p e n nis, including a monograph by Beklemishev (1944). We give below only 
a few of the more important ecological aspects which are valid for all 
subspecies. 

The larvae breed in permanent and temporary stagnant water bodies of 
varying type, depth and size, with full sunshine (partly shaded in the south) 
and with filamentous algae and other aquatic vegetation. They occur in 
permanent water bodies among aquatic plants which form a "second bottom" 
at the surface. When the water bodies dry up in summer, they are found in 
shallow water and in the backwaters of slowly flowing streams. They occur 
in slightly brackish water in the south. 

The females hibernate in empty buildings, often in stables, rarely in 
caves or burrows. In the northern and eastern parts of the range the 
mosquitoes hibernate in empty buildings for many months at low tempera- 
tures (A. m. messeae). 

The phenology of the species has been thoroughly studied according to 
the conditions in different landscape zones (Beklemishev and Shipitsina, 
1957). In warm weather, when the temperature is favorable for reproduction 
(minimum of development at about 10°), the species produces a number of 
generations: 2 generations in the north, 4—6 in the south. The maximum 
number of adults appears in midsummer, but in regions with a very hot 
summer there is a decrease in numbers in July— August. 

Most mosquitoes are concentrated in populated areas and their vicinity. 
They feed mainly on farm animals (cows, horses, pigs, etc.) and to a lesser 
degree on people. If there are few or no animals they attack mainly people. 
They may also feed on domestic and wild birds. 

A.maculipennis is an obligatory bloodsucker. A blood meal is 
necessary for reproduction. Autogenous development of the eggs has not 
been observed in' A . maculipennis. Gonotrophic harmony in the summer 
is characteristic for this species (probably for the whole genus). One 
complete blood meal is sufficient for the development of eggs. 



100 



Epidemiological importance. A.maculipennis was a 
dangerous carrier of the three main types of malaria in the past. In most 
of Europe, Kazakhstan, Siberia and Transbaikalia it was virtually the only 
vector of malaria. In the southern parts of its area of distribution it also 
carried malaria together with other species (and still does in some places' 

A.maculipennis is an intermediate host of the filariae of dogs 
Dirofilaria immitis and D . r e p e n s . 

It transmits Calova virus in Czechoslovakia. 

A. maculipennis may also have a part in spreading tularemia. 
Natural infections have been found. 



99 Anopheles maculipennis sacharovi Favre, 1930 
(elutus Edwards, 1921) 

It differs from the other subspecies in its lighter coloration. Lateral parts 
of mesonotum yellowish brown, more or less as in the middle. Dark spots 
on the wings, especially in males, weakly defined and not distinguishable in 
old specimens. The scales at the margin of the wings are of uniform color, 
without a light spot at the apex. 

These characters are not certain and determination of adults is not 
always possible. 



100. 




FIGURE 43. Anopheles maculipennis sacharovi 
Favre. Stigmal plate of 4th-stage larva. 



101 



Fourth-stage larva (Figure 43 ) closely resembling that of 
A. maculipennis but usually smaller. The differences are only 
quantitative, small and variable. Light yellowish green larvae, with weakly 
pigmented head and stigmal plate are common. The head pattern is not 
sharp and resembles that of the newly molted larvae of A. maculipennis. 
The hair on the antenna and the outer clypeal hairs are relatively longer 
and extend to the anterior margin of the extended lateral lobes of the labrum, 
or even slightly beyond them. However, there are many exceptions. This 
also refers to the variation of the number of leaflets of the palmate hairs 
which are usually fewer in A. m. sacharovi than in maculipennis 
but their form is identical. 

The stigmal plate differs from that of A. maculipennis in size and 
in the intensity of pigmentation of the central plate. It varies markedly 
in size but is distinctly smaller and more weakly pigmented. It is 0.38— 
0.5 mm long and the width between the ends of the lateral lobes is 0.39— 
0.52 mm. The central plate has a very weak, light, median granulation. 
The dark area in the posterior part reaches to the posterior margin It 
is interrupted anteriorly and is distinct only in the middle. The base of 
the anterior process barely reaches the anterior part, where it forms folds 
which only rarely, in less pigmented larvae, reach to the middle section. 

Eggs deposited in spring, summer and autumn have no floats, only a 
fringe. However, in the south of the range, where development in winter 
continues, eggs deposited in winter have small air floats, but the number and 
size of the cells are very variable. 

Distribution. Mediterranean, mainly the eastern part : Italy, southern 
Balkan, Southwest Asia. In the USSR: Middle Asia to the Aral Sea and 
Kirghiz Range, Eastern Transcaucasia, Dagestan. It also occurs in Iran, 
Afghanistan and the part of Sinkiang near to Middle Asia. 

Biology. It occurs mainly in arid zones, where it is common. The 
larvae breed in open, sunny, shallow water rich in algae and with sparse higher 
aquatic vegetation. They tolerate distinctly saline water. They are often 
found together with the larvae of Culex modestus Fie. They are little 
mobile, rarely leaving the surface. 
100 A.m. sacharovi is a serious vector of malaria everywhere. 



6. Anopheles (Anopheles) lindesayi Giles, 1900 

Coloration dark brown. Head black, with dense, black, upright scales on 
sides of occiput, middle of occiput white, with upright scales; vertex with a 
tuft of yellowish white hairs between the eyes, directed anteriorly. Antennae 
as in A. maculipennis, palps and proboscis black. Mesonotum light 
ash gray in the middle, dark brown at the sides, with a wide longitudinal 
stripe bordered by a narrow, grayish stripe. Sides of thorax mainly dark, 
scutellum gray, hairs on thorax pale yellowish. Legs blackish brown, coxae 
and trochanters pale, hind femora with a broad, white ring in 3 / 4 of its length. 
Wings mainly with dark scales; there is a large white spot on the costa 
near the apex of the wing which extends to n and r2; white scales present 
in middle of r3, in basal part of Ti+s, in middle of mi, in basal and apical 
thirds of m3+4 and at ends of veins r 4 + 5 , m2, cui and an; halteres pale, with 



102 



(101) 



black head. Abdomen blackish brown, grayish ventrally, with long, brownish 
hairs. 




FIGURE 44. Hypopygium of Anopheles lindesayi Giles (after La Casse and Yamaguti) 

Hypopygium (Figure 44) with two setae at the base of the coxite; spines 
at end of claspettes fused, forming a long, narrow plate; lobes of tergite 9 
short, conical. 

Fourth-stage larva (Figure 45) to 6 mm long, gray or grayish 
green. Head yellowish brown or dark brown. A dark, oval spot in the 
middle of the frontoclypeus near its posterior margin that extends anteriorly 
beyond the base of the frontal hairs; near the eyes are two small, oval 
spots. Clypeal hairs simple, postclypeal hairs rarely 2 -branched at the 
end; inner clypeal hairs situated close together, very long, outer hairs 
V2 — V3 shorter, postclypeal hairs as long as the outer clypeal hairs. Frontal 
hairs short, the longest inner hair hardly reaching beyond the base of the 
postclypeal hairs. Antennae more strongly pigmented at the apex, hair 
short, situated near the base, with 6—9 branches. 

Hair No. 1 on metathorax palmate, distinct, with 15—22 leaflets without 
terminal filament, sometimes rudimentary. Palmate hairs of abdomen well 



103 



(102) 










FIGURE 45. Anopheles lindesayi Giles. Fourth-stage larva (after Puri): 

1 - anterior margin of frontoclypeus; 2 - right antenna, inner side; 3 - median hairs of prothorax; 

4-6 - bases of lateral hairs of prothorax, mesothorax and metathorax; 7 - palmate hair of 4th abdominal 

segment; 8 - stigmal plate. 



104 



developed on segments 2 — 7, with 13- 17 leaflets on segment 2, 19— 24 on seg- 
ments 3—6 and 15—19 leaflets on segment 7. Leaflets dark, with a long 
terminal filament. At the base of the filament are long denticles, which pass 
gradually into the filament. 

Stigmal plate with an anterior lobe which is dark at the base, lighter 
103 anteriorly. Processes of spiracles usually distinct; if they are absent, the 
anterior part of the central plate is adjacent to the spiracles; its margins 
are distinctly striated, granulation in the middle distinct but only slightly 
darker than the lateral parts. 

Distribution. USSR: mountain regions of Tadzhikistan, 800— 1,600 m 
above sea level (Keshish'yan, 1938). Mountain regions of India, Malaya, 
China, Philippines. The subspecies A.lindesayi japonicus Yam. 

» occurs in Japan. 
Biology. The larvae occur in shaded mountain streams with little 
vegetation and in streams with pure, cold water (11 — 15°). They are found 
mainly in weaker currents and in winding areas near the bank. The larvae 
hibernate. The mosquitoes bite man at dusk, also outdoors. 



Anopheles lindesayi lindesayi Giles 

The strongly developed white scales on the ventral surface of the hind 
femora cover about x / 3 of the femur from the base. 



Anopheles lindesayi japonicus Yamada, 1918 

It differs from the nominate subspecies in the weak development of white 
scales on the ventral surface of the hind femora which cover only about % 
of the femur from the base. The larvae do not differ from those of the 
nominate subspecies. 

Distribution. Japan (Hokkaido, Honshu, Kyushu), Korea, China. 

Biology. It occurs in northern regions. The larvae are found to- 
gether with those of Anopheles koreicus Yam., A. hyrcanus, 
Culex hayashii and C. infantulus Edw. They prefer pure water. 

7. Anopheles (Anopheles) hyrcanus Pallas, 17 71 
(? sinensis Wiedemann, 1828) (Figure 46) 

Markedly varying in size and coloration. 

Head dark brown, with blackish brown upright scales at sides of occiput, 
white scales dorsally; a tuft of white hairs directed anteriorly situated on 
the vertex between the eyes; proboscis and palps dark brown, palps of 
females with 3 white rings and white apex; there are sometimes also white 
scales on the palps; the palps of the female, especially in the basal half, 
are covered with upright scales. Antennae of female dark brown, with white 
pubescence, with long, brown hairs; the first 5—7 antennal segments bear 
some white scales. Mesonotum brown, with a more or less distinct, broad, 
gray stripe in the middle, often divided by dark longitudinal stripes into 



105 



104 




FIGURE 46. Anopheles hyrcanus Pall. 



106 



two or four narrow gray stripes; mesonotum, except for the narrow stripes, 
sometimes covered with gray tomentum. At the anterior margin of the 
mesonotum is a tuft of white scales. Hairs on mesonotum yellowish brown. 
Sides of thorax brown, with grayish tomentum. Legs brown, lighter at 
ventral side of femora and inner side of tibiae. Base of fore femora 
distinctly thickened. Tarsi dark brown, with white rings at the apex of the 
first 3 or 4 tarsal segments, particularly on the hind legs, or whole 4th 
tarsal segment of hind legs white; the white color sometimes also extends 
to part of the 5th segment. Wings mainly with dark scales; there are two 
large white spots at the anterior margin, one at the distal third, the other 
near the apex. The first spot extends to ri and the second to r2. Constant 
areas of white scales are present in the basal third of ri, at the forks of 
r2+r3 and mi + m2, m3+ 4 and particularly on the cubitus and anal vein which 
are sometimes mainly covered with white scales. There are sometimes 
also scattered white scales between the brown scales. Fringe of wing white 
at the apex, otherwise brownish; halteres pale, with brown head. Abdomen 
brown, with long, dense, brown or golden hairs. 




FIGURE 47. Hypopygium of A n o p h e 1 e s hyrcanus Pall, (after La Casse 
and Yamaguti) 



107 



Hypopygium (Figure 47): coxite with 2 strong, simple setae on the inner 
side near the base. Spines of claspettes fused, forming a narrow plate; 
tergite 9 with long, narrow processes. 

Fourth- stage larva (Figure 48) very variable in color and pigment- 
ation of chitinized formations, particularly of the head and stigmal plate. 



106 




FIGURE 48. Antennae of larvae: 

1— Anopheles maculipennis Mg.; 2 — Anopheles hy re a nus Pall.; 3 — stigmal plate of 
4th- stage larva of A .hyrcanus. 

Inner clypeal hairs situated close together, simple or often slightly 
branched at the ends; outer hairs about fa as long as the inner hairs; they 
may reach to the anterior margin of the lateral lobes of the labrum, dendri- 
form. From the main stem extend 3 — 6 branches which are repeatedly 
branched, the inner branches longer than the outer branches; postclypeal 
hairs weakly developed, with 2—3 branches, reaching to the base of the 
anterior clypeal hairs. 

Antenna with a strongly developed hair near the middle, with 7—8 
branches, at least half as long as the antenna or longer. 

Hair No. 1 on metathorax weakly developed, with long leaflets without a 
terminal filament. Palmate hairs rudimentary on segments 1 — 2, well 
developed on segments 3— 7, with 17—24 leaflets with indistinct terminal 
filament passing in steps into the leaflet, which is dark in the basal half 
and light at the apex. 



108 



Stigmal plate dark only at base of anterior lobe, spiracles not connected 
with anterior part of central plate; the dark base of the central plate 
extends beyond the border between the anterior and middle part; median 
dark pigmentation of posterior part of central plate weakly developed only 
at the posterior margin; granulation light, present only in the middle near 
the posterior margin, without dark outline. 

Eggs with medium-sized air floats and a broad, smooth (not striated) 
fringe. 

Note on systematics. This species has been treated differently. 
The Oriental forms which were described as subspecies of A. hyrcanus 
(nigerrimus Giles, 1900; p s e u d o s i n e n s i s Baisas, 1935; lesteri 
Baisas and Hu, 1936) are now considered as distinct species. The Oriental 
sinensis Wied., described from Canton, possibly differs from the 
Palaearctic A. hyrcanus (see Reid, 1953). 

Intraspecific variations (aberrations) of A. hyrcanus have been 
described from the Palaearctic, particularly ab. pseudopictus Grassi 
(with a white 4th s egment of the hind tarsus ) and ab. mesopotamiae 
(with a light, diffuse pattern of the wing). However, in view of the wide 
variation of A. hyrcanus, there is no reason to consider these forms as 
subspecies (Maslov, 1946). 

"Ab . mesopotamiae" is probably a seasonal form, as the light, 
diffuse pattern of the wings is found mainly among hibernating females 
(Kazantsev, 1930). A.chodukini Mart., 1929, described from Tashkent, 
is probably an albinistic form. 

Distribution. It occurs from the Atlantic to the Pacific, including 
South and Southeast Europe, West, South and East Asia, and some regions of 
Central Asia (Mongolia, Sinkiang). This species and closely related groups 
are common in Southeast Asia (India, Sunda Islands, and Philippines), from 
where it extends far to the north and west in the Palaearctic. The eastern 
area of distribution is represented by China, Japan, Korea and the southern 
Maritime Territory (especially common near Lake Khanka and extreme 
southern coast of the territory). The northern border crosses the Amur 
and Aman' and then passes into Mongolia and China. From the upper 
reaches of the Irtysh (Lake Norzaisan) the northern border passes to 
Central Kazakhstan and to the lower reaches of the Ural, Volga and Don 
rivers, south of Kharkov and Kiev, and slightly north of Kishinev. It also 
occurs in South and Southeast Europe and Southwest Asia. It is widespread 
in Southern Kazakhstan, Middle Asia and the Caucasus, in river valleys and 
deltas. It occurs to 1,500— 1,600m in the mountains. 

Biology. The larvae breed in stagnant water warmed by the sun and 
overgrown with vegetation, in small and large swamps and lower reaches 
of rivers covered with bulrushes. They also occur in rice fields and 
irrigation canals. In the north and northeastern parts of the range (Far 
East), especially in winter, they breed mainly in open water. In the south, 
and other parts of the area of distribution, during the hot season, they inhabit 
shaded, muddy water with rich vegetation and medium mineralization and 
high oxidation. They occur together with larvae of Anopheles pulcher- 
rimus, A.maculipennis sacharovi, Culex modestus and 
others. The females hibernate among reeds in riparian forests, shrubs, 
etc. They are active from spring to November and their numbers increase 
in autumn. There are two (in the north) to four generations per year. 



109 



This species has little contact with man. The females rarely enter 
108 houses but do enter cattle sheds. After bloodsucking they fly away. They 
pass the day mainly in thickets of twinflower. They bite mainly in the open 
air, at dusk and in the night, but also in sunlight. Their attacks in riparian 
forests are sometimes in large numbers and persistent. In the southern 
Maritime Territory their breeding places are close to human habitations 
and they commonly enter houses. 

The epidemiological importance of the species depends on the conditions 
of the locality. In the Caucasus and Middle Asia it is of minor importance 
as a vector of malaria in the valleys of large rivers in the plains. A . hyr- 
canus did not become infected experimentally with tropical malaria but 
was infected with tertian malaria (Simanin, 1930). In the Far East (southern 
Maritime Territory) it was the only vector of malaria but it did not become 
widely distributed. In East Asia (Korea, some parts of China and Japan) it 
is the main vector of malaria. 

A. hyrcanus probably plays a part in the circulation of tularemia in 
some river foci. Natural infections have been found. 



2. Subgenus Myzomyia Blanchard 
(Cellia Theobald) 

Cross-veins and base of forks of wing with white scales; costa with 4 
or more white spots. Hypopygium with 4—6 large setae at base of coxite. 

The subgenus Myzomyia is mainly distributed in the Oriental and 
Ethiopian regions. About 80 species have been recorded from the 
Ethiopian region and 45 from the Oriental region. Fourteen species occur 
in Australia, but none in America. The West Palaearctic species are 
closely connected with the Ethiopian region (Mediterranean) and the eastern 
species, particularly A. . pulcherrimus, with the Indian part of the 
Oriental region. 

The larvae of Myzomyia have widely separated inner clypeal hairs, 
the distance between them larger than the distance between the outer hairs 
and the inner hairs. Frontal hairs feathered. Antenna with a short simple 
hair. Long, feathered, lateral hairs are present on abdominal segments 4—6. 

8. Anopheles (Myzomyia) pulcherrimus Theobald. 1902 

The species differs from the other species, except from the African 
A. pharoensis.by the distinct tufts of scales at the sides of the abdominal 
tergites and the white segments 4 and 5 of the hind tarsi. 

Head with light brown, upright scales laterally, white scales dorsally, 
and with an anteriorly directed tuft of white hairs on the vertex between the 
eyes. Antennae of female dark brown, with short, delicate, white pubescence. 
Each segment bears long, white hairs. The first 7—8 segments also bear 
109 white scales which cover the whole segment, sometimes only part of it (apex, 
inner or outer surface). Antennae of male brown, with long golden hairs. 
Proboscis with light brown, adpressed scales; palps brown, with light brown 



110 



scales, upright in the basal half of the palp, mainly adpressed in the apical 
half. All segments of palps with white rings at the apex; these rings are 
l /s to V4 as l° n g as the segment, except on the last segment, on which the 
white ring is about l fa as long as the segment. Mesonotum brown or gray- 
ish brown, densely covered with broad, white scales. The narrow stripes 
in the posterior half of the mesonotum and sometimes also the narrow 
stripes in the anterior half are the only parts without scales. Scutellum 
of the same color as the mesonotum and covered with white scales. Pleurae 
of thorax brownish gray, with grayish tomentum, densely covered with white 
scales, particularly in the middle. Legs brown, covered in a large part 
with white scales. Fore femora distinctly thickened in the basal third, 
covered with white scales often mixed with some brown scales. Fore 
tibiae covered with white scales; there are 2 sometimes fused, narrow, 
longitudinal stripes of brown scales on the outside. Fore tarsi blackish 
brown, with white rings at apex of the 1st, 2nd, 3rd and sometimes 4th seg- 
ment; mid-femora anteriorly with brown scales mixed with white scales 
which sometimes form a longitudinal stripe, with white spots in the apical 
third and at the apex and with white scales posteriorly, with a white ring 
near the apex; tarsi of mid-legs brown, with white rings at the apex of the 
first 2 — 3 segments; hind femora like femora of mid-legs. Scales on hind 
tibiae mainly white; tarsi of hind legs with a pale brown 1st segment with 
a white ring at the apex. Second segment with a dark brown or blackish 
brown ring at the base, j$— fa as long as the segment; apical part of 2nd 
segment and segments 3—5 white. 

Wings with white scales in the greater part. There are usually 6 black 
spots at the anterior margin; the first two spots are in the basal fourth of 
the wing, are small and present only on the costa; the 3rd black spot is 
much longer (about fa as long as its distance from the base) and broad and 
extends to the subcosta and ri; the 4th spot is 1.5 times as long as the 
3rd, and extends to the subcosta, n and also to the stem r2+3 + r4+s; the last 
spot is sometimes covered with white scales in the part adjacent to the 4th 
spot. The 4th spot is often divided into three small spots with white scales 
on ri. The 5th spot resembles the 3rd spot in form and distance from the 
apex and its distance from the base. The 6th spot is usually short and forms 
a curve toward the apex, occupying the costa, and ri and V2- There are also 
more or less constant spots of dark scales in the basal third of one or both 
branches of the fork r2+r3, in the middle third of r3, in the basal third of 
the fork mi + ni2, and also in the apical parts of all longitudinal veins from 
110 r3 to anal vein. There are also less constant spots of dark scales in other 
parts of the wing, especially in the apical part of the stem r2+r3 and mi + m2 
and on the anal vein, and in other parts. The fringe of the posterior margin 
consists in its apical half mainly of alternating parts of dark and white 
scales, only of white scales in the basal half. Halteres light, with dark 
brown head. Abdomen brownish black, with long, golden hairs on the 
posterior margin and at sides of tergites, covered nearly completely (except 
for a stripe at the anterior margin of the tergite) with broad, white scales. 
The white scales of the lateral parts of the abdomen sometimes have a 
golden or yellowish sheen. Posterior corners of tergites 2-7 with a tuft 
of projecting, white or yellowish brown scales. Abdomen dark brown 
ventrally, with white scales. 



11 




FIGURE 49. Hypopygium of Anopheles p ul cher rimus Theob. 

Hypopygium (Figure 49): coxite with 5 strong setae near the base on the 
inner side; claspettes with plates which are wider at the apex; aedeagus 
without leaf- shaped appendages. 

Fourth-stage larva (Figure 50) 5— 7 mm long, yellowish green. 
Inner clypeal hairs widely separated: distance between them twice the 
distance between the outer hairs and inner hairs. Inner clypeal hairs with 
6—15 sparse, thin lateral branches from the basal third, so that they appear 
feathered. Outer hairs nearly half as long as inner hairs; they branch 
into 4—12 (average 7) thin branches, usually of the same length, from the 
middle or from the apical third. Postclypeal hairs half as long as the 
outer clypeal hairs, with 2—4 branches from near the base. 

Hair No. 1 of metathorax well developed, palmate, with a long stem and 
8—13 long, narrow leaflets without terminal filament, forming a fan and not 
111 a rosette. Palmate hairs of abdominal segment 1 weakly developed 
and with 5—9 narrow leaflets without terminal filament, forming a fan. 
Leaflets distinct on segments 2-7, with 13-18 leaflets on the 2nd segment, 
with 18—23 leaflets on segments 3—6 and with 16—22 on segment 7. They 
are of uniform color except for the darker apex, which bears a long, pointed 
terminal filament with well defined, stepped denticles at the base. 

Stigmal plate (Figure 51) very characteristic; anterior lobe irregularly 
dark; spiracles adjacent to anterior part of central plate, in the form of 
an arrowhead with distinct, posteriorly directed lateral processes the posterior 
margin of which forms a distinct concavity at the border with the middle 
part; medial part of central plate weakly pigmented and forming a striated, 
light medallion. 

Eggs with lateral air floats and a broad, transversely striated fringe. 

Distribution. USSR: plains and valleys of Middle Asia (Turkmenia, 
Uzbekistan, Tadzhikistan, southern Kirghizia, south of Kazakhstan (Syr- Darya 
valley)); not higher than 900-1,000 m in mountains. The species was 



112 



formerly present in Azerbaidzhan, especially in the steppes in the lower 
reaches of the Kura River, Iran, Iraq, Afghanistan, Pakistan and India. 









FIGURE 50. Anopheles pul c he rrimus Theob. Fourth- stage larva (after Puri): 

1 — anterior margin of frontoclypeus; 2 — variation of branching of outer clypeal hair; 3 — median 
hairs of prothorax; 4,5 — base of lateral hairs of mesothorax and metathorax; 6 — palmate hair of 
4th abdominal segment. 



112 Biology. The larvae inhabit mainly shallow lakes, riparian forests 

and large swamps with rich aquatic vegetation, Potamogeton, Cera- 
t ophy llu m, etc. floating on or near the surface. In contrast to larvae 
of other mosquitoes which occur mainly near the shore. A. pulcher- 
rimus occurs mainly in the open parts where the dense thickets of 
Potamogeton replace the bottom. The larvae also are found in 
permanent pools, fed by underground water or by irrigation ditches in ponds, 
rarely in rice fields. They avoid small pools (less than 1— 2m ) or water 
of high salinity. In the USSR, they hibernate as larvae of the middle stages 
(adults apparently hibernate in India); there is normally a high mortality in 
winter, so that the first generation is small. Their numbers increase in autumn. 



113 



A sharp fluctuation of the size of populations in the same locality in 
different years is characteristic for A. pulcherrimus. Especially 
damaging are unusually low temperatures in winter and a long cold spring 
The mortality is very low in warm winters and springs, and numbers in- 
crease in summer. These fluctuations are often due to local factors. 




FIGURE 51. Anopheles pulcherrimus Theob. Stigmal 
plate of 4th-stage larva. 



Females enter houses and cattle sheds. They also bite outside, mainly at 
dusk and in the night but there have also been mass attacks on man during 
the day. In contrast to the other species, A. pulcherrimus likes light 
and aridity. 

It is usually considered as a minor vector of malaria. Experimental 
infection with tertian and tropical malaria was successful. Naturally in- 
fected mosquites have been found. In parts of Middle Asia, where 
A. pulcherrimus was common, it was considered as an important 
carrier of malaria, but its populations have now been reduced. 



113 



9. Anopheles (Myzomyia) superpictus Grassi, 
1899 (Figure 52) 



This is a very variable species, particularly in the color of the legs and 
wings. The hind tarsi are uniformly brown or have indistinct light rings. 
114 Head with brown, upright scales laterally and upright white scales 

dorsally. Antennae with brown hairs; proboscis long, thin, with light apex; 
palps black, with distinct white rings and white apex; scales on palps ad- 
pressed. Mesonotum light brown, slightly darker laterally, with sparse, 



14 



(us; 




FIGURE 52. Anopheles superpictus Grassi 



115 



pale brown hairs and dense, narrow, whitish scales, especially in the 
anterior part; sides light brown, with a grayish tinge. Legs dark brown, 
coxae light, with white narrow rings at the knees, and at the articulations 
of tibiae and tarsi; tarsi mainly brown, rarely slightly ringed. Wings of 
markedly varying color; there are usually 4 spots of white scales at the 
anterior margin (the 4th spot at the apex of the wing in the male), and there 
are more or less numerous white scales in the basal part of the costa; the 
other parts of the anterior margin are covered with blackish brown scales; 
the light and dark spots of the anterior margin extend to the subcosta and 
ri; the veins of the other parts of the wing (posterior to ri) are mainly 
covered with white scales; more or less constant spots of dark scales are 
present on the stem of r2+ r3, in the basal part of r^+ r 5 , on the media (2 
spots on the stem of mi + m2 and one spot on each branch of the fork on 
m3+4, and on the anal vein. Halteres brown, with dark head. Abdomen 
brown, with long, yellowish hairs. 




FIGURE 53. Hypopygium of A nopheles s u p er pi c tu s Grassi 

115 Hypopygium (Figure 53): coxite relatively short and thick, with 5 strong 

setae at the base; claspettes with a plate which is widened at the apex; 
aedeagus with short, leaf-shaped appendages. 



116 








FIGURE 54. Anopheles superpictus Grassi. Fourth-stage larva (after Puri): 

1 - anterior margin of frontoclypeus; 2 - median hairs of prothorax; 3—5 - base of lateral hair of pro- 
thorax, mesothorax and metathorax; 6 — palmate hair (No.l) of metathorax; 7 — palmate hair of 4th 
abdominal segment. 

Fourth-stage larva (Figure 54) small, about 5 mm long, gray or 
grayish green. Inner clypeal hairs widely separated, the distance between 
them greater than the distance between the outer and inner hairs. Inner 
hairs thin, simple, with sparse secondary feathering, the outer hairs half as lon| 
simple, smooth; postclypeal hairs simple, as long as the outer clypeal 
hairs. 

Palmate hair on metathorax weakly developed, consisting of 6—8 thin, 
narrow, long, lanceolate leaflets, folded in a fan. Palmate hairs present on 
abdominal segments 2—7. On the 1st segment they have 5—7 thin branches 
often folded together; they have 12—15 leaflets on the 2nd segment, 14-18 
on the third, 15—19 on segments 4—6 and 14—16 on segment 7. Leaflets 
with well developed terminal filament fa as long as the leaflet, striated and 
pointed, with narrow base and variable denticles. 
116 Stigmal plate (Figure 55): anterior lobe dark but irregularly pigmented; 
spiracles adjacent to anterior part of central plate or connected with it by 



117 



processes; central plate broad, not tapering in anterior part of middle 
part the lateral margins of which are parallel; anterior part without 
lateral processes, forming an angle; granulation in middle of central 
plate weakly developed, diffuse but not forming a striated, light medallion. 




FIGURE 55. Anopheles superpictus Grassi. Stigmal 
plate of 4th-stage larva. 



The eggs vary markedly, usually with a broad, transversely striated 
fringe but without air floats. Eggs deposited in the cold season, often with 
a few small air floats grouped more or less in an oval area on the flat- 
tened side at one end. 

Distribution. This species is characteristic for the Mediterranean, 
where it is common in countries with a dry climate and is rare or absent 
where the humidity is high. Its distribution in the USSR includes Ciscaucasia 
(Dagestan), Transcaucasia (Georgia, Armenia, Azerbaidzhan), Middle Asia 
(Turkmenia, Uzbekistan, Tadzhikistan, Southern Kirghizia, southern Kazakh- 
stan). Southern Europe: Spain, southern half of the Apennines, and Balkans. 
North Africa, from Algeria to Egypt. Asia Minor, Iraq, Iran, Afghanistan, 
Pakistan, northwest India. 

Biology. The larvae inhabit flowing, slowly flowing or stagnant springs 
of ground origin (surface filtration) with waters transparent and well 
warmed water. They also occur in shallow pools in dry river beds (hoof- 
marks). In rapid currents they occur among algae between stones or at the 
bank. They have been found in rice fields with water of stream or ground 



118 



origin. The larvae have a high resistance to mineralization of the water 
and the salts in them, but avoid muddy water polluted by decomposition of 
organic substances with a high oxidation. They are very agile. At the 
slightest disturbance they descend to the bottom where they lie under 
117 stones, etc. Stirring brings them to the surface. Fertilized females 
and only a few larvae hibernate. 

Females usually remain active in winter; some of them suck blood 
during this time. The spring generation of larvae (usually in April in 
Middle Asia) is not large; the larvae develop relatively slowly. Develop- 
ment lasts 24 days at 21.5°, 10 days at 27°. The average duration of develop- 
ment is 12—13 days. Their numbers increase rapidly at the end of May and 
in early June (in Middle Asia) after the recession of floods and reach a 
maximum in the middle of summer (July in Middle Asia). The numbers 
decrease in August and increase again in the second half of September. 
This may sometimes be connected with the drainage of water from rice 
fields. During this time there is also an intensified flight of females into 
houses because of the colder nights. They begin to hibernate in October. 

The females of A. superpictus feed on man and animals . They 
often enter houses during the day, between sunrise and dusk, and feed on the 
inhabitants. Many of them also enter cattle sheds during the day. They 
rest in dark corners, empty dishes, folds of clothes, etc. They also rest in 
tree holes, crevices in cliffs and clay structures, burrows of rodents 
(Citellus, Gerbillus, etc . ), caves , etc . Activity reaches a peak at sun- 
set and declines after nightfall. 

The epidemiological importance of the species as a vector of all forms 
of malaria is very high. In many parts of Middle Asia it is a more im- 
portant vector than other species of Anopheles because of its greatest 
numbers in the hot months (July— September), when the development of the 
Plasmodia is very short. This is intensified by the anthropophilic character 
of the females and the rapid digestion which increases the need for blood 
more than in other species of Anopheles. There are also the ecological 
characters of the larvae that sustain high numbers of the species. In the 
Near and Middle East, A. superpictus is the most important vector of 
malaria. 



II. Subfamily TOXORHYNCHITINAE 

Large, brightly colored mosquitoes. The clypeus is wider than long. 
Proboscis thick, thinner in the apical half, usually curved downward. 
Maxillae and mandibles extending to about the middle of the proboscis, 
without denticles. Spiracular setae present, postspiracular setae absent. 
Scutellum uniformly curved. Wings with short radial fork (r2+3); stem 
118 of fork several times longer than fork, between ni3+4 and cui is a thickened 
membrane in the form of a narrow triangular plate; the posterior margin 
of the wing has a small incision at the apex of cui. Abdomen covered with 
broad scales and often with tufts of bright hairs at the end. 

The larvae are large, about 20 mm long, dark wine red with a violet 
tinge; head rectangular, hairs weakly developed and situated mainly in 



19 



2 lateral groups in its anterior third. Antennae short, rod-shaped. Mouth 
parts predaceous. The lateral lobes of the labrum catch the prey; they 
consist of about 10 strong, hook-shaped, curved setae arranged in a row and 
serrated at the end. The mandibles have strongly developed denticles. 

Thorax relatively narrow. A pair of tracheal air sacs on the main trunks 
of the trachea with hydrostatic function in its posterior part. Hairs of the 
body partly shortened and forming coarse spines with secondary spines. 
The lateral hairs are strongly developed, the dorsal hairs weakly developed 
because the larvae live under the surface film. At the base of the hairs and 
spines are strongly sclerotized plates, usually 3 pairs on each segment on 
each side in 3 longitudinal rows. 

Larger plates are present on the sides of segment 8. On its dorsal side 
is a short siphon with a pair of hair tufts at its base; auricles and pecten 
absent. The short, posterior segment is surrounded by a chitinized ring 
with spines at the posterior margin. 

Biology. The larvae inhabit small accumulations of water in the axils 
of leaves of Bromeliaceae and in bamboo and tree holes, feeding on larvae 
of other mosquitoes. They are predaceous, killing even each other. They 
are used to kill the larvae of other mosquitoes in tree holes on tropical 
islands. 

The females deposit the eggs singly on the surface of the water. They 
are mainly found at the upper border of forests, where they feed on nectar 
and do not suck blood. 

The subfamily contains only one genus with three subgenera. 



2. Genus Toxorhynchites Theobald 
(M eg a r hi n u s Ronineau-Desvoidy)* 

With the characters of the subfamily. 

This is a tropical genus; only a few species occur in the southern 
temperate zone. Over 50 species have been described. The subgenera 
Ankylorhynchus and Lynchiella occur in the tropics of the western 
hemisphere while the species of the subgenus Toxorhynchites are 
mainly distributed in the Oriental region. The Palaearctic species are 
related to the Oriental group. 



1. Toxorhynchites christophi Portschinsky, 1884 
(?towadensis Matsumura, 1916) (Figure 56) 

119 These are large mosquitoes, 10—13 mm long without the proboscis. 

Nearly all scales are contiguous, with a green, blue, violet or silvery white 
metallic sheen. 

Female. Occiput with broad, dark brown scales with a blue metallic 
sheen. Posterior margin of eyes with bluish white scales. Proboscis very 
long ( lz — U as long as the body), curved downward, with dark brown scales 

* The name Meg arhinus is now considered as a synonym of the subgenus Lynchiella. 



120 



with a steely sheen. There is an indistinct ring of lighter scales in the 
middle of the proboscis and long, black hairs at its base. Palps short with 
blackish brown scales with a metallic sheen. Antennae brown, torus with 
white scales. 




FIGURE 56. Toxorhynchites christophi Portsch. 

Mesonotum with reddish brown, narrow scales, the anterior margin and 
lateral parts with broader, pale blue scales; above the base of the wings 
120 are tufts of yellow hairs. The pleurae are covered with broad, white, shining 
silvery scales in their greater part. Scutellum with shiny, broad scales, 
greenish in the middle and golden laterally, the golden scales more numerous, 
The setae on the margin of the scutellum form a complete row. 



121 



Legs: femora and tibiae with black, steel-colored scales. Basal part 
of femora with white scales ventrally. Apical half of 1st segment, the 
whole 2nd segment, sometimes also base of 3rd segment of fore tarsus, 
base (sometimes also apex) of 1st and other segments of the mid-tarsus, 
and 2nd segment of hind tarsus white. The other tarsal segments are 
black with a blue sheen. Claws simple. 

Wings transparent, slightly dark; veins with brown scales. Cross-vein 
between r2+3 and r4+5 situated more distally than cross-vein r— m, which is 
situated in a line with the cross-vein m— cu. 




FIGURE 57. Toxorhynchites christophi Portsch. 
Fourth-stage larva. Head, dorsal. 



Abdomen covered with black scales, with a greenish, metallic sheen in 
the anterior half and with a bluish sheen in the posterior half. In the 
anterior half of the 5th (and sometimes also the third) tergite is a more or 
less distinct transverse stripe of white scales; sides of tergites also white. 
Fifth tergite with white stripes laterally, 6th tergite with tufts of longer, 
dense black hairs, tergites 7 and 8 with tufts of bright orange-yellow hairs. 

Male. (We have no males in the collection and give the description of 
the male of T . t ow a de n s i s after La Casse and Yamaguti, 1955. ) Palps 
slightly shorter than proboscis, with dark scales and with 2 light rings; 
apex of palps curved dorsally, pointed. Coloration as in the female. 
Tergites 6 and 7 with thick tufts of dark hairs laterally, tergite 8 with tufts 
of golden brown hairs. 

There is no description of the hypopygium. 



122 



Fourth- stage larva (Figure 57) about 16— 20 mm long, wine-red- 
violet, resembling that of T. splendens Wied. 

Head rectangular, with rounded corners, frontoclypeus distinctly concave 
laterally behind the eyes. Anterolateral corners of frontoclypeus project- 
ing anteriorly, anterior margin concave. Clypeus strongly developed, 
especially its lateral parts with the lateral lobes of the labrum, which 
catch the prey. Each lobe bears 10 thick, curved, dark hooklike setae with 
2—3 short denticles at the end. The other mouthparts are situated at the 
anterior margin, so that the gular region is very large. Mandibles with 2 
large and 3—4 smaller denticles at the end. Antennae short, less than half 
121 as long as the head, their ends at the level of the ends of the setae of the 
labrum when extended anteriorly. Antennae without spines. Three hairs 
at the terminal quarter, one of them, the one farthest from the apex, simple 
and directed inward, the other two shorter and branched. 




FIGURE 58. Toxorhy nchit es christophi Portsch. Fourth- stage larva. 
Chaetotaxy of thorax (shown in a plane). Roman numerals designate thoracic 
segments, Arabic numerals numbers of hairs. 



123 



Clypeus with three simple hairs on each side, the groups widely 
separated. The two outer pairs correspond to the clypeal hairs of larvae 
of Anopheles and the longer inner hairs to the median frontal hairs . 
There are four pairs of hairs on the frontoclypeus in the anterior third which 
form two lateral rows. The two middle pairs are simple and the longer 
hairs correspond to the outer and inner frontal hairs, the shortest branched 
inner pair corresponds to the postclypeal hairs and the outer pair to the 
sutural hairs; near these and lateral to the frontal suture are 2 branched, 
transsutural hairs. 
123 The hairs on the body vary in structure; there are 3 types: l) large 

spines, with coarse secondary feathering, simple or 2-branched; 2) long, 
thin, smooth and simple hairs, sometimes slightly branched at the end; 
3) short hairs, simple or slightly branched some distance from the base. 
The groups of hairs are situated on sclerotized plates. 

Each thoracic segment bears 14 pairs of hairs which form several groups 
in 3 transverse stripes in several longitudinal rows: dorsal, dorsolateral, 
lateral, ventrolateral and ventral (Figure 58). The arrangement of the 
hairs and chitinized plates on the abdominal segments is shown in Figures 
59 and 60. 



(122) 




FIGURE 59. Toxorhy nchites christophi Portsch. Fourth-stage larva. Chaetotaxy of abdominal 
segments 1,2,4-6. Rows of dorsolateral (dl) .lateral (1) and ventrolateral (vl) hairs of left side. 

At the sides of the 8th segment are large plates and at their posterior 
margin are two long, secondarily feathered setae and above them two thin, 
short hairs and below the base of the siphon a longer, thin hair. These 
five pairs of setae and hairs correspond to the hairs behind the comb of 
larvae of Culicinae. A comb is absent. Siphon short, in the form of a 
truncate cone. It is slightly longer than wide at the base. Auricles and 
pecten absent. Two short hairs with 5—6 secondarily feathered branches 
situated on the posterolateral side of the siphon near the base. 



610144 



124 






122) 







FIGURE 60. Toxorhy nchites christophi Portsch. Fourth-stage larva. Posterior end, lateral. 

Last segment short and broad. The saddle surrounds it completely 
and has a row of closely situated spines of different length on the sides of 
the posterior margin. The lateral hair has the form of a spine and is 
shorter than the segment. The outer and inner caudal hairs have 6—7 
branches. The outer hairs are slightly longer than the inner. The fin is 
situated posteriorly, with 18 — 20 long, secondarily feathered hairs which 
form an asymmetrical fan. Gills short, spherical. 

Biology. The larvae live in water in the holes of elm and linden trees. 
They are found from the end of June to late August or early September. 
Remains of larvae of A. (St.) galloisi Yam. and A . ( F . ) alektorovi 
were found in the intestine. Very rare. 

Note on systematics. Shtakel'berg (1937) stated that T . t o w a - 
d e n s i s Mats . is apparently a synonym of T. christophi. There are no 
distinct differences between them. However, the description of T. towa- 
d e n s i s (La Casse and Yamaguti, 1955) mentions golden brown hairs on 
abdominal segment 8 of the female, and does not mention the bright hairs 
on tergites 5—7. These tufts of hairs, white, black and yellow, give the 
insect its characteristic appearance. They may have been lost on the 
specimen of M.towadensis described. 



125 



Distribution. Southern Maritime Territory (Alektorov, 1931; 
Monchadskii, 1951). Japan (Honshu). 



2. Toxorhynchites sp. (Figures 61, 62) 

Only the larvae are known. 

The 4th- stage larva resembles that of T. christophi but differs 
as follows: l) head wider; 2) spines on the body more strongly developed; 
simple spine number 8 on lateral plate of mesothorax not 2-branched; 
125 siphon nearly cylindrical, not truncate-conical; tuft of hairs on siphon 

longer than width of siphon at its position; lateral spine on saddle longer 
than last segment; ends of caudal hairs slightly curved upward, the outer 
hairs weakly branched; fin with about 15 hairs on which the secondary 
feathering begins at the base; gills larger, oval. 

(124) 




FIGURE 61. Toxorhynchites sp. Fourth- stage larva. Anterior end, dorsal. 



126 



;i24) 




FIGURE 62. Toxorhy nchites sp. Fourth- stage larva. Posterior end, lateral. 

Biology. The larvae were found in water in holes of trunks of Korean 
pine and in root holes of maple in July and August together with larvae of 
A. (St.) galloisi, A . ( F . ) alektorovi and A . ( F . ) nipponicus. 

Distribution. Southern Maritime Territory, Barabash area, 
"Kedrovaya pad'" Reserve (Monchadskii, 1940; Amosova, 1951). 



III. Subfamily CULICINAE 

Clypeus longer than wide, its anterior margin rounded. Proboscis thin, 
rarely slightly thicker apically, not curved; mandibles and maxillae of 
females long, rarely (some exotic species) extending to end of labium, with 
denticles in the apical part. Mesonotum distinctly convex. Scutellum dis- 
tinctly 3-lobed, with 3 tufts of setalike hairs; space between lobes bare. 
Wings with long radial fork, which is rarely shorter than its stem; thickened 
plate between ni3+4 and cui absent. Abdomen with scales, which are usually 
contiguous. Hypopygium of varying structure; anal segment always with 
well developed sternite; aedeagus without leaf-shaped appendages. Females 
usually with 3 spermathecae . 

The head of the larva is often wide, rarely rounded or oblong, not rotating 
through 180°. Frontoclypeus with 3 pairs of frontal hairs in its anterior 
part, arranged in a curved row or forming 2 symmetrical triangles, and a 
pair of postclypeal hairs which are branched in a fan but are not feathered. 
Antennae of varying form and length; they are longer in larvae with the 
filtration type of feeding and shorter, rod- shaped with different types of hairs 
and sensory setae at the end in larvae feeding on periphyton and on predators. 
The mouth parts differ according to the type of feeding. 



127 



Hairs of the body branched or fan- shaped, situated in one plane or 
palmate: their branches arranged like the ribs of a polyhedral pyramid; 
the branches may be secondarily feathered. 

Thorax broad, without shoulder organs. Abdomen without palmate hairs. 
Abdominal segment 8 with a varying number of scales at the sides forming 
the comb behind which are 5 hairs. At the posterior margin of the dorsal 
side of segment 8 is the siphon with a pecten and from 1 to 5-6 pairs of 
variously arranged hair tufts. The last segment, bears the saddle, which 
sometimes forms a complete ring, and two pairs of caudal hairs and one 
pair of lateral hairs. Fin present, rarely absent. 

The large subfamily Culicinae has been divided into a different number 
of tribes (2-12). Dodge,(l962) suggests dividing the subfamily into four 
tribes: Sabethini, Aedini, Uranotaeniini, Culicini. We prefer the simpler 
126 classification of Stone et al. (1959), in which the subfamily Culicinae is 

divided into the tribe Sabethini with 8 genera (tropics of both hemispheres, 
mainly in South America) and the tribe Culicini with 19 genera. Eight 
genera of Culicini occur in the Palaearctic : Uranotaenia, Culiseta, 
Orthopodomyia, Mansonia, Aedes, Culex, Heizmannia and 
Armigeres. The first six genera occur in the USSR. Only one species 
of Tripter oides, T. bambusa Yam. (R a c hi o n o t o m y i a bambusa^ 
occurs in the Palaearctic in Japan and China. 



3. Genus Uranotaenia Arribalzaga 

Small, dark species. Eyes contiguous in dorsal part of head above the 
antennae and ventrally (behind the proboscis). Palps of both sexes short, 
usually not longer than Ve of the length of the proboscis. Mesonotum with 
well developed acrostichal and dorsocentral setae. Lobes of pronotum 
widely separated, usually with 3 setae; one spiracular seta which is rarely 
absent (in some exotic species); postspiracular setae absent; there are 
also 2 prealar, several upper sternopleural, usually 2 upper mesepimeral 
and one lower mesepimeral setae. Legs slightly widening at base of mid- 
femora rarely also fore femora. Tibiae and tarsi of males of many exotic forms 
with distinct secondary sexual characters . Claws of both sexes simple, without 
denticles, pul villi absent. Microtrichia very small, wings appearing bare 
without high magnification; forks short, fork mi+ 2 distinctly shorter than 
its stem. Anal vein ending in posterior margin at level or before base of 
stem of r 2 +3 and r 4 + 5 ; postalar sclerites without scales. Abdomen short, 
with blunt apex in females. Hypopygium with a short coxite with a moder- 
ately large basal lobe which is covered with setae. Style simple, with an 
apical appendage; anal segment membranous, without sclerotized denticles; 
phallosome divided into plates, sometimes with denticles. 

Larvae small, with dark, slightly oblong head and short antennae. At the 
sides of abdominal segment 8 are chitinized plates with a row of a few 
spines at the posterior margin. Denticles of pecten of siphon with 
fringed margin, not pointed. One pair of hair tufts near middle of the 
siphon. The larvae keep horizontal under the surface of the water. 

The genus contains about 140 species, most of them in the tropics of 
both hemispheres. It is represented in the Palaearctic by one species in 



128 



(127) 




FIGURE 63. Uranotaenia unguiculata Edw. 



129 



the Mediterranean area in the wide sense (including the Caucasus and Middle 
Asia) and by one Oriental species in Japan. 



1. Uranotaenia unguiculata Edwards, 1913 (Figure 63) 



Small mosquitoes with narrow, silvery stripes at sides of mesonotum and 
on the pleurae. General color dark brown. 

Head covered with dark brown scales in its greater part, usually with a 
longitudinal stripe of silvery scales dorsally. Proboscis and palps of both 

127 sexes blackish brown. Palps short in both sexes. Antennae and antennal 
hairs brown in both sexes. Basal segments of antennae with silvery scales. 
Mesonotum with dark brown scales, sometimes with a reddish tinge, and 
with a narrow longitudinal stripe of silvery scales at the sides from the 
anterior margin to the base of the wings. Scutellum with brown scales; 
pleurae with a longitudinal stripe of silvery scales in the middle. Legs: 
coxae of fore and mid-legs with silvery scales; femora with brown scales 
anteriorly, usually with a sharply defined longitudinal stripe of white scales 

128 which is complete on the fore and mid-legs and fused with the white scales 
on the posterior surface on the hind legs; posterior surface of fore and mid- 
femora also with white scales; tibiae anteriorly with blackish brown 
scales, often with a diffuse white spot in the middle; tibiae with white 
scales posteriorly; apex of femora and tibiae with white scales; tarsi black; 

claws of fore tarsus of male with one 
denticle. Wings transparent, slightly 
grayish, veins covered with broad, brown 
scales; stem of radial veins with white 
scales in the basal part. Abdomen with 
dark brown scales dorsally, with tri- 
angular spots of silvery scales at the 
sides, sometimes only on the last ter- 
gites, with white scales ventrally. 

Hypopygium (Figure 64): coxite 
thick and short, irregularly conical, 
with a small, flat lobe with 3 setae on 
the inside in the basal third; style 
broad, slightly tapering in the basal 
third, with a spine -shaped appendage at 
the apex. 

Fourth-stage larva (Figure 65) 
small, yellowish green or yellowish 
brown, with dark head. 

Head oblong. Mouth parts situated 
anteriorly, so that the gula is 1.5—2 times 
as large as in larvae of other genera 
and the base of the labrum is situated 
more anteriorly. This is connected with 
the feeding on the lower side of the 
surface film and bending the head 
posteriorly when the body is horizontal. 




FIGURE 64. Hypopygium of Urano- 
taenia unguicul at a Edw. 



130 




FIGURE 65. Uranotaenia ungu i c ul a t a Edw. Fourth- stage larva: 
1 — head .dorsal; 2 — posterior end, lateral. 

Outer frontal hairs with 4-7 short branches, median hairs longer, simple, 
thick. The inner hairs are situated behind the median hairs, close together, 
simple, rarely with 2—3 branches. Postclypeal hairs short, simple or 
slightly branched, situated before the inner frontal hairs. Antennae short, 
straight, slightly tapering apically, dark, almost or completely without 
spines. Hair of antenna thin, short, situated in the middle of the dorsal 
surface. 

Abdomen: segment 8 with a chitinized plate at the sides, the upper 
margin of which may extend to the dorsal side. Plates indistinct in recently 
molted larvae, darker toward the end of the stage. In the ventral part of 
the posterior margin are the spines of the comb in a row of 5-8, usually 
6, dark, conical, pointed spines which are covered with thin, very short hairs 
in a row. 

Siphon with well developed auricles at the base, moderately long (index 
3.2-3.8, usually 3.5-3.6), with straight anterior side, slightly tapering 
apically. Pecten with 13-18 (usually 15-16) weakly pigmented denticles 
129 which are regularly spaced (except the 2—4 basal denticles). Each denticle 
forms a scale with short, thin spines at the margin. One pair of hairs with 
8—12 secondarily feathered branches in the middle of the siphon near the 
distal denticles of the pecten. 



131 




FIGURE 66. Uranotaenia unguiculata Edw. Fourth- stage 
larva. Stigmal plate. 

The stigmal plate (Figure 66) has a posterior process of the "stirrup," 
which ends in two pointed clawlike, inward directed appendages, and a thick, 
lateral margin of the spiracles and a slightly twisted, anteriorly directed 
hair of the chaetoid type with a wide base. 

The last segment is surrounded by a dark chitinized ring with a thin, 
3-branched lateral hair. Outer caudal hairs 2-branched, one hair as long 
130 as the siphon, the inner hairs with 4 shorter branches. Fin with about 10 
tufts. Gills lanceolate, with rounded end, much shorter than the last 
segment. 

Distribution. Mediterranean. USSR: southern Ukraine, Volga delta, 
Caucasus, Middle Asia, to the basin of the Hi River in the Alma-Ata Region 
in the northeast; Southern Europe, North Africa, Southwest Asia, Iran, 
Pakistan. 

Biology. The larvae are found in shaded parts of small, stagnant pools 
overgrown with vegetation (sometimes Lemna). They prefer fresh or 
nearly fresh water and avoid water with a salinity higher than 0.1 — 0.2°7oo. 
They are found together with larvae of Anopheles maculipennis 
sacharovi, A. hyrcanus, Culex pipiens and Aedes caspius 
caspius. They rarely bite man. 



4. Genus O r t h o p o d o my i a Theobald 

The genus is characterized by the absence of pulvilli and the presence 
of spiracular setae. Two proepimeral setae. First segment of fore tarsi 
longer than the other segments. Large insects, mainly black with a white 
pattern. 



132 



131) 




FIGURE 67. O rthopodomy i a p ul c hr ip al p is Rond. 



133 



The larvae are reddish or pinkish and become bluish violet before 
pupation; the hairs of the thorax and abdomen are very long, particularly 
the lateral hairs. There are plates on abdominal segments 6—8, a pecten 
on the siphon is absent, and the comb consists of 2 straight rows of long 
scales of characteristic form. The main tracheal trunks are widened in 
the metathorax The larvae develop in tree holes. 

This is a small genus with about 20 species in the Oriental region and 
in the southern part of North America and in Central and South America. 
One Mediterranean species in the Palaearctic. 



1. Orthopodomyia pulchripalpis Rondani, 1872 
(a 1 b i o n e n s i s MacGregor, 1919) 

This species differs from all other Palaearctic mosquitoes in the 
characteristic pattern on the mesonotum (Figure 67). 

Head black, with black scales mixed with white scales. Posterior 
margin of eyes with white scales dorsally. Proboscis with black scales 
and with a more or less broad white ring near the middle. Palps of male 
distinctly longer than proboscis, black, with white rings at the articulations 
and with white terminal segment. Palps of female nearly half as long as 
the proboscis, black, with a white ring in the middle and with white apex. 
Antennae black, basal segment with white scales. Mesonotum black, with 
narrow black scales and with narrow, white longitudinal stripes; the 2 
median stripes extend from the anterior margin to the scutellum, parallel 
in the anterior half, slightly curved in the posterior half; the two lateral 
stripes are present only in the posterior half above the base of the wings; 
lateral margin of mesonotum with a narrow white stripe. Legs with black 
132 scales with a steel sheen; base of femora with yellowish white scales; 

dorsal surface of femora with speckles of white scales; tibiae with a white 
spot; tarsi black with white rings covering the apex of the preceding seg- 
ment and the base of the following segment; white rings on fore and mid- 
tarsi weakly developed, present usually only on the basal segments; white 
rings distinct on all segments of the hind tarsi; segment 5 of hind tarsi 
white. Wings transparent; veins with brown scales, base of stem of radial 
veins with silvery white scales. Abdomen blackish brown, covered with 
scales of the same color, with moderately broad transverse stripes of white 
scales at the anterior margin of the tergites. 

Hypopygium (Figure 68): coxite with a conical lobe near the base; lobe 
with 4—5 large setae; style relatively narrow, slightly thicker in the basal 
half, with an appendage about as long as the width of the style in its widest 
part at the apex. 

Fourth-stage larva (Figure 69 ) reddish, becoming violet before 
pupation. Head dark, rounded. Frontal hairs long, with secondary feather- 
ing; outer hairs shorter than the others, median hairs (9— 10-branched) 
and inner hairs (5— 7-branched) reaching far beyond the anterior margin of 
the head. Slightly before and between the inner frontal hairs are the post- 
clypeal hairs, which resemble the outer frontal hairs in length and branch- 
ing. Antennae straight, without spines. A hair with 7 short branches form- 
ing a fan at fa of the length from the base. Mouth parts transitional to the 
filtration type of feeding. 



134 




FIGURE 68. Hypopygium of Or t ho po do my i a 
pulchripalpis Rond. 

Thorax wide, especially posteriorly, with very long lateral hairs. Ab- 
dominal segments distinctly narrower posteriorly Lateral hairs also very 
long. In older larvae, there are dark plates on the dorsal side of segments 
6-8. The plates are small and narrow on segment 6, extend to the sides on 
segment 7, are narrower but also extend to the sides on segment 8. There 
are also narrow lateral plates between the 8th and last segment. 

Siphon (index 3.5-4) without a pecten and auricles. There is a hair with 
8-10 branches forming a fan on the posterior surface near the middle. 
133 The comb consists of two rows of long denticles with a pointed spine; some 
denticles of the posterior row (16-20) are situated between the denticles 
of the anterior row (7-10). Only the median hair of the hairs behind the 
comb is strongly developed, resembling the hair on the siphon. 

The last abdominal segment has a ring-shaped saddle, its ventral margin 
much shorter than the dorsal-margin. Outer caudal hairs nearly as long 
as the abdomen, simple, inner caudal hairs forming a short, asymmetrical 
fan with 9-14 branches. Fin with 14 tufts. Gills lanceolate; dorsal pair 
as long as the saddle, ventral pair half as long. 

Distribution. Mainly Mediterranean. USSR: southern coast of the 
Crimea, Transcaucasia from Sochi to Lenkoran'. Southern Europe, North 
Africa (Algeria, Tunisia), Turkey. It extends to Belgium and the south of 
England in the north. 



135 




FIGURE 69. Orthopodomy ia pulchrip al pis Rond. Fourth- stage larva: 

1 — posterior end, lateral; 2—4 — scales of comb (2 — large, 3 — small, 4 — lateral). 

Biology. The larvae breed in water in tree holes (beech, horse chest- 
nut, and others) or below the roots, often together with larvae of Anopheles 
plumbeus, Aedes pulchritarsis and Aedes geniculatus. 
They hibernate in the 4th stage. They rarely bite man during the day in 
shaded places . 



5. Genus Culiseta Felt* (Theobaldia , 

Neveu-Lemaire) 

The name The obaldia Neveu-Lemaire, 1902 is preoccupied by a 
genus of molluscs (1885). 
134 Usually medium-sized or large mosquitoes. Spiracular setae present, 
postspiracular setae absent. Claws of female simple, pul villi absent. 
Stem of radial vein with hairs in the basal part which are more numerous 
on the ventral surface. Scales on veins narrow. Cerci of female not 
projecting. Hypopygium: coxite usually with a lobe, style with a short 
appendage; claspettes absent. 



A.V.Maslov (1964,1967) was of great help in the preparation of this chapter. 



136 



Larvae large to very large, with a broad head (except in Allotheo- 
ba 1 d ia), mouth parts adapted to feeding on the substrate (Allotheo- 
b a 1 di a) or to mixed Culiseta s. str. ) or filtration feeding. Antennae 
short, with a weakly developed tuft (Allotheobaldia) or longer (Culi- 
seta) or very long, with a tuft in the form of a broad fan (Culic e 1 1 a); 
variations of frontal hairs similar. Scales of comb on abdominal segment 8 
numerous, without a main spine. Siphon of varying length, with a short 
pecten, often continuing toward the distal end as spines (Allotheobaldia, 
partly Culicella) or as a row of hairs (Culiseta s. str.). Tuft at base 
of siphon always present. Stigmal plate with a posterior process of the 
"stirrup" in the form of a more or less large plate. Main tracheal trunks 
either wide, ribbon-shaped, with oval cross section (Allotheobaldia, 
Culiseta s. str.) or narrow, with rounded cross section. Saddle either 
weakly developed (Allotheobaldia) or surrounding the last abdominal 
segment as a ring. Fin strongly developed, some of the tufts situated before 
the common base on the saddle, Gills of varying length. 

The genus contains only about 30 species of which 12—13 occur only in 
the Palaearctic. Most of the species occur in the temperate zone of the 
Palaearctic and Nearctic. 



Key to species 
Females 

1 (2). Mesonotum with distinct longitudinal white stripes forming a lyre- 

shaped pattern (Figure 70). Femora and tibiae with distinct white 
spots anteriorly. Costa covered with white scales in a large part. 
(Palps of males slightly shorter than proboscis.) (Subgenus 

Allotheobaldia Brol.) 

; . . l.C. (Allotheobaldia) longiareolata Macq. 

2 (l). Mesonotum without white stripes. Femora and tibiae with uni- 

formly dark scales or more or less numerous, scattered, light 
scales anteriorly. Costa covered mainly or completely with dark 
scales. (Palps of male longer than or as long as proboscis.) 

3 (10). Cross-veins c— m and m— cu situated in a line; if they are slightly 

separated, the distance between them is usually not longer than 
the cross-vein m— cu. Wings often with dark spots. (Subgenus 
Culiseta Felt.) 

4 (7). Tarsi dark. 

5 (6). Wings with more or less indistinct dark spots 

5. C. (Culiseta) bergrothi Edw.* 

135 6 (5). Spots on wings absent or indistinct 

2. C. (Culiseta) glaphyroptera Schin* 

7 (4). Tarsi with white rings. 

8 (9). First segment of hind tarsi without a light ring in the middle; 

femora without subapical pale ring 

3. C. (Culiseta) alakaensis Ludl. 

" The females of C.bergrothi and C. glaphyroptera are difficult to distinguish, but the 
hypopygium of males is distinctly different. 



137 



a (b). Dark scales on the body and legs and dark brown or black 
and white scales form the contrast of the pattern. Light scales 
on wings absent or few. Dark spots on wings distinct. A stripe 
of white scales at the base of the abdominal tergites which is 
narrower in the middle. Light scales absent in posterior half of 

tergites C.(Culiseta) alaskaensis alaskaensis Ludl. 

b (a). Dark scales on body and legs and light brown or brown or 
light ocher-colored scales form a diffuse pattern on legs and body. 
Scattered light scales on the wings. Dark spots on wings indis- 
tinct. Stripes of pale scales on tergites indistinct; a few light 

scales are scattered all over the tergites 

C. (Culiseta) alaskaensis indica Edw. 

9 (8). First segment of hind tarsi with a white ring in the middle; femora 

with a subapical, light ring 4. C. (Culiseta) annulata Schr. 

a (b). Costa with dark (black) scales. Dark spots on wings dis- 
tinct. Distinct anterior stripes of light (white) scales on the 

tergites; light scales absent in posterior half of tergites 

C. (Culiseta) annulata annulata Schr. 

b (a). Costa with dark and light scales. Dark spots on wings 
diffuse. Light stripes indistinct at base of abdominal tergites and 
formed by yellowish (not white) scales; such scales are also 
present among the dark scales in the posterior half of the tergites 
C. (Culiseta) annulata subochrea Edw. 

10 (3). Cross-vein m— cusituated distinctly nearer to the base of the 

wing than r— m, the distance between them usually at least the 
length of m— cu. Wings without dark spots. (Subgenus Culicella 
Felt.) 

11 (14). Sternites of abdomen usually with a pattern of dark scales in the 

form of a V, its apex anteriorly. 

12 (13). Light rings present on all segments of the fore, mid- and hind 

tarsi 9. C. (Culicella) fumipennis Steph. 

13 (12). Light rings not present on all tarsal segments 

9. C. (Culicella) setivalva Masl. 

14 (ll). Sternites of abdomen without a pattern of dark scales in the form 

of a V. 

15 (16). Narrow stripes of light scales present only at base of tergites. 

Scales not forming spots on wings 

6. C. (Culicella) morsitans Theob. 

16 (15). Narrow, indistinct stripes of light scales at the base, sometimes 

also at the apex of the tergites, or else absent; tergites some- 
times completely covered with light scales. There may be an 
indistinct dark spot in the middle of the wings, at the base of r4+5 
7. C. (Culicella) ochroptera Peus. 



136 Males 



1 (2). Tergite 9 with long lateral processes; aedeagus thick 

l.C. (Allotheobaldia) longiareolata Macq. 

2 (l). Tergite 9 simple, without lateral processes; aedeagus normal. 



138 



3 (4). Coxite with an apical lobe which has a tuft of long, scalelike 

plates 2. C. (Culiseta) glaphyroptera Schin. 

4 (3). Apical lobe of coxite with hairs or lobe absent. 

5 (6). Coxite with slightly convex apical lobe densely covered with short 

hairs 3. C. (Culiseta) alaskaensis Ludl. 

6 (5). Coxite without apical lobe. 

7 (8). Usually an area with dense hairs on the inner surface of the coxite 

before the apex. Phallosome strongly sclerotized 

4. C. (Culiseta) annulata Schr. 

8 (7). Area with dense hairs before apex of coxite absent. Phallosome 

usually weakly sclerotized. 

9 (10). Coxite with long, strong setae, 2—3 setae on inner surface of 

coxite 9. C. (Culicella) setivalva Masl. 

10 (9). Coxite with moderately long, thin setae. 

11 (12). Basal lobe of coxite with 2 setae. Sternite 8 with a row of 

usually at least 10 spines in the middle of the posterior margin 
5. C. (Culiseta) bergrothi Edw. 

12 (ll). Basal lobe of coxite with 3 large setae. Posterior margin of 

sternite 8 with fewer spines (at most 10) or without spines. 

13 (14). Sternite 8 without spines at the posterior margin 

7. C. (Culicella) ochroptera Peus (part). 

14 (13). Posterior margin of sternite 8 with a comb consisting of densely 

placed spines or with a few scattered spines. 

15 (16). Distal segments of palps of male (at least one segment) distinctly 

thickened 6. C. (Culicella) morsitans Theob. 

16 1 1 ) Distal segments of palps of male not thickened, as thick as the 

preceding segments. 

17 (18). Basal lobe of coxite with 3—4 large setae 

8. C. (Culicella) fumipennis Steph. 

18 (17). Basal lobe of coxite with 5—8 large setae (rarely 3—4) 

7. C. (Culicella) ochroptera Peus (part). 



Larvae 

1 (12). Siphon relatively short and thick (index at most 4). Main tracheal 

trunks broad, ribbonlike. Antennae shorter than the head, with a 
weakly developed tuft. 

2 (3). Pecten of siphon with a few large denticles, widely separated, on the 

entire siphon. Spines on antennae absent; tuft short, situated in 
middle of antenna, with at most 3 branches (subgenus Allo- 
theobaldia Brol.)... 1. C. (Allotheobaldia) longiareolata Macq. 

3 (2). Pecten of siphon with thinner denticles at the base, continuing 

distally in a row of long hairs. Antennae with sparse spines; 
tuft in middle of antenna short, slightly nearer the base, with more 
than 5 branches (subgenus Culiseta Felt). 

4 (7). Antennae about half as long as the head. Median frontal hairs 

with 4—8 branches. Comb usually with more than 60 scales. 

/ \ 2/ 

5 (,6). Antennae /3 as long as the head. Pecten of siphon occupying about 

/4 of length of siphon, with more hairs than scales. Before the 



139 



fin are 5 tufts, 3 of them situated on the saddle 

2. C. (Culiseta) glaphyroptera Schin. 

6 (5). Antennae half as long as the head. Pecten of siphon occupying 

/a of the length of the siphon, with as many scales as hairs or 
slightly more. Before the fin are 3—4 tufts, 2 of them situated on 
the saddle 5. C. (Culiseta) bergrothi Edw. 

7 (4). Antennae less than half as long as the head. Median frontal hairs 

with 3 branches. Comb usually with less than 50 scales. 

8 (9). Siphon shorter and thicker, slightly tapering apically, ratio of 

length to its width at the base 2.4—3.0 (2.7), and to its width at the 
apex 3.4—3.9(3.7). Scales of comb narrow, elongate, with parallel 

lateral margins, not distinctly wider at the base 

3. C. (Culiseta) alaskaensis Ludl. 

9 (8). Siphon longer and thinner, markedly tapering apically, ratio of 

length to its width at the base 3—4, to its width at the apex 5—6. 
Scales of comb slightly narrower in the middle, distinctly wider 
at the base. 

10 (ll). Distance between postclypeal hairs as large as distance between 

inner frontal hairs 4. C. (Culiseta) annulata Schr. 

11 (10). Distance between postclypeal hairs shorter than distance between 

inner frontal hairs C. (Culiseta) annulata subochrea Edw. 

12 (l). Siphon long and thin (index more than 5). Main tracheal trunks 

narrow, rounded. Antennae longer than the head, with a well 
developed tuft (subgenus Culicella Felt). 

13 (16). In addition to the pecten, the siphon bears spines which are longer 

than the scales and irregularly scattered on the posterolateral 
surface. Hair at the base of the lateral valves of the stigmal 
plate forming a half- open fan on a chitinized base. 

14 (15). About 50 scales in the comb. Median hair behind the comb as 

long as the last abdominal segment. Outer caudal hairs forming 
an asymmetrical fan, branched dorsally, with about 6 branches. 
Anterior tufts of the fin not situated on the common base, just 

more than half as long as the posterior tufts 

8. C. (Culicella) fumipennis Steph. 

15 (14). About 100 or more scales in the comb. Median hair behind the 

comb much shorter than last abdominal segment. Outer caudal 
hairs with 3 branches. Anterior tufts of fin not situated on the 
common base, as long as or only slightly shorter than the posterior 
tufts 9. C. (Culicella) setivalva Masl. 

16 (13). Additional spines apart from the pecten on the siphon absent. 

,00 Hair at base of lateral valves of stigmal plate of the usual structure. 

17 (18). Inner frontal hairs with 5—9 branches. Scales at posterior margin 

of comb with a dark, longitudinal ridge, pointed at the end. Hair 
at posterior valves of stigmal plate curved only at the end. Gills 

1.5—2 times as long as the saddle 

7. C. (Culicella) ochroptera Peus. 

18 (17). Inner frontal hairs with 2-3 branches. Scales at posterior margin 

of comb without dark longitudinal ridge. Hair at posterior valves 

of stigmal plate curved their entire length. Gills shorter than 

the saddle 6. C. (Culicella) morsitans Theob. 



140 




FIGURE 70. Culiseta 1 ongi a re ola t a Macq. 



141 



1. Subgenus Allotheobaldia Brolemann 

The males of this subgenus have short palps which are about 
/i as long as the proboscis. Coxite without apical lobe, tergite 9 with 2 
long processes, aedeagus thick. 

Larvae with a small head, which is less than 1.5 times wider than long; 
mouth parts of the mixed type of feeding, substrate feeding type predominat- 
ing over filtration type. Antennae short, without spines, with a short, 
slightly branched hair. The short siphon is usually not sclerotized to the 
base and has no auricles. There is a pair of hair tufts at the base and a 
pecten consisting of a few spine-shaped, widely separated denticles on the 
entire length of the siphon. Main tracheal trunks broad, ribbon-shaped. 
Last abdominal segment with a weakly developed saddle, markedly branched 
tufts of the fin and short gills. 

The only species of the subgenus is C. (All.) longiareolata. 

It was recently suggested (Maslov, 1964) that the subgenus Allotheo- 
baldia Brol. be removed from the genus Culiseta and considered as 
a distinct genus. C. (All.) longiareolata Macq. indeed differs from 
all other species of the genus in the structure of the hypopygium, the length 
of the palps of the male and other characters, but the same standard should 
be maintained in different groups of the family. In some genera (e.g. in 
Aedes, Culex) the differences between the species are at least as sub- 
stantial as in Culiseta. The differences ofC. (All.) longiareolata 
are sufficiently recognized if the species is placed in a different subgenus. 

1. Culiseta (Allotheobaldia) longiareolata Macquart, 1838 

This species differs sharply from all other Palaearctic species of the 
genus in the white, lyre-shaped pattern on the mesonotum (Figure 70) and 
in the white scales on the anterior margin of the costa. 

Head with broad, white, contiguous scales dorsally in the middle, with 
sparse brown setae laterally. There are also white scales at the sides of 
the head and on the occiput and dense white scales along the margin of the 
eyes. Palps of male brown, shorter than the proboscis, long segment of 
palps white at the base and in the middle; penultimate segment of palps 
with white scales at the base, last segment with white scales also at the 
apex; last segment markedly thickened and short. Palps of female short, 
140 with dark brown and white scales; apex of palps of female with white scales. 
Proboscis blackish brown. Antennae of male blackish brown, with white 
rings at the base of the segments and long brown hairs; antennae of female 
blackish brown, the basal segments with white scales. Mesonotum with 
brown or yellowish brown, narrow scales; a median stripe of white scales 
from the anterior margin to the scutellum; similar white stripes in the 
anterior half of the mesonotum at the lateral margin; they turn inward 
along the transverse suture and extend to the scutellum lateral to the dorso- 
central setae; mesonotum with white scales at the base of the wings, on the 
scutellum and on the pleurae. Legs black, with numerous white spots which 
often form longitudinal stripes; femora and tibiae covered with white scales 
on a large part of the posterior surface; tarsi black, with scattered spots 



142 



of white scales and narrow, white rings at the base of the first 2, 3 or 4 
segments (on the hind tarsi); claws of female simple, claws of male with 
a denticle on the fore and mid-tarsi. Wings with brown scales on the veins; 
costa with white scales on the entire anterior margin. Abdomen markedly- 
varying in color of scales, usually with brown scales dorsally, with broad 
transverse stripes of white scales at the base of the tergites; posterior 
third of tergites with yellowish scales which often form a process to the 
white scales at the anterior margin or cover the part of the tergite without 
white scales (often on the posterior tergites), replacing the brown scales; 
sternites with white scales. Sternite 8 of female with 2 broad lobes with 
an incision between them. 

Hypopygium (Figure 71) with strongly developed, finger-shaped process 
of tergite 9 and a thick, sclerotized aedeagus. 




FIGURE 71. Hypopygium of Culi- 
seta longia reol at a Macq. 



Fourth-stage larva (Figure 72 ) large, gray, with small dark head 
and short siphon. Head less than 1.5 times wider than long. Frontal hairs 
weakly developed: outer hairs with 3—4 branches, medianhairs situated before 
the inner, simple; inner hairs simple, rarely 2-branched. Postclypeal hairs 
simple, situated between the median frontal hairs. Antennae short, without 
spines, with a short hair with 1—3 branches which is not longer than twice 
as long as the width of the antenna. 

Comb with 40—75 scales, very variable in size and form, with a distinct 
spine or only with a row of denticles. The median hair behind the comb is 
strongly developed. 

Siphon (index 1.5—2.1) dark, but usually not sclerotized to the base; 
auricles absent. Pecten with 4—7 large, smooth spines, widely separated 
on the entire siphon, and with 3—7 smaller, rudimentary denticles in its 
basal, not sclerotized part. There are all transitions between these 
denticles and the scales of the comb. A hair tuft with 10—15 secondarily 
feathered branches, thinner and shorter than on the median hair behind the 
comb, at the base of the siphon near the border of the sclerotized part. 



143 




FIGURE 72. Culiseta longiareolata Macq. Fourth-stage larva: 
1 — head, dorsal; 2 — posterior end, lateral; 3 — scales of comb. 

Last segment of abdomen with a weakly developed saddle which covers 
at most fa of its dorsal side and extehds little to the sides; its posterior 
margin with dense, short spines. Outer caudal hairs as long as the siphon, 
with 1 — 2 branches, inner caudal hairs forming a fan with 10—16 branches, 
as long as the fin, which consists of 16—18 strongly branched tufts. Length 
of gills very variable, according to the salinity of the water, 0.5—1.5 times 
as long as the last segment. 

Distribution. Mainly Mediterranean. USSR: steppe and desert 
zones, Central and Southern Ukraine to Kharkov, Lower Volga area> Northern 
Caucasus, Transcaucasia (Georgia, Armenia, Azerbaidzhan), Kazakhstan and 
some southern regions of West Siberia (Omsk), Middle Asia (Turkmenia, 
Uzbekistan, Tadzhikistan, Kirghizia). Western Europe to southern England; 
Africa; Southwest Asia, to Iran, India and Pakistan. 

Biology. The larvae inhabit shallow pools, wells, prospecting pits, 
142 etc. and natural waters, fresh or slightly saline. They tolerate a high 

degree of pollution. Most of their life is spent at the surface; they rarely 
descend to the bottom. They often occur together with larvae of Culex 
p i p i e n s, rarely with larvae of Aedes caspius. The larvae hibernate. 
In Middle Asia they continue to develop even in cold water so that adults 
are found even in winter. They rarely bite man but transmit blood para- 
sites of birds. Several Plasmodia of birds may complete the cycle of 



144 



sporogony in this species. Autogenous deposition of eggs has been found 
in some populations. 




FIGURE 73. Culiseta longiareolata Macq. Fourth-stage 
larva. Stigmal plate. 



2. Subgenus Culiseta Felt 

Palps of male longer than or as long as the proboscis, with 2 thickened, 
apical segments. Coxite often with a subapical lobe or a sharply defined 
area of dense hairs in the upper third of the inner surface; tergite 9 without 
long appendages; aedeagus normal: phallosome markedly sclerotized, with 
curved apical parts in most species. 

Head of larva usually 1.5 times wider than long; mouth parts usually 
of the mixed type of feeding, with short antennae which are only rarely 
(C . glaphyroptera) more than half as long as the head, with a relatively 
short tuft and with subapical and apical setae in two groups. Comb with 
numerous densely arranged scales. Siphon moderately long (index less 
than 4), with a pair of hair tufts at the base and pecten continuing distally 
in a row of long, thin, dense hairs. Main tracheal trunks broad, ribbon- 
shaped. Saddle surrounding the last segment completely like a ring. 
Two to four tufts of the fin situated on the saddle. Outer caudal hairs with 
a few branches. 

The subgenus is represented in the Palaearctic by six species, one of 
them Oriental. 



145 



2. Culiseta (Culiseta) glaphyroptera Schiner, 1864 




FIGURE 74. Hypopygium of Culi' 
seta glaphyroptera Schin. 



The species is characterized by the structure of the hypopygium and by 
the dark brown tarsi without light rings and the absence or weak development 

of dark spots on the wings. 

Head laterally and dorsally with whitish 
scales at margin of eyes, dorsal part of 
occiput with upright, brown scales. Pro- 
\ \\ \ / boscis and palps dark brown in both sexes, 

palps of male distinctly longer than the 
proboscis, with widened but flattened 
terminal segment. Antennae of male brown, 
with light rings on the basal and middle 
segments and uniformly brown terminal 
segments; antennae of female brown. 
Mesonotum with brown or golden brown 
scales which form indistinct stripes and 
spots; pleurae of thorax with contiguous 
white scales in some places. Legs with 
brown scales; posterior surface of femora 
and tibiae with yellowish white scales; tarsi 
dark brown, without light rings. Wings with 
dark brown scales, sometimes with indistinct 
spots of darker scales at the base of the 
radial and medial forks and at the cross- 
veins. Abdomen dark brown dorsally, with transverse stripes of white 
scales at the anterior margin of the tergites which occupy about fa of the 
length of the tergite; sternites with white scales in a large part and stripes 
of brown scales at the posterior margin. 

Hypopygium (Figure 74): tergite 9 without lateral processes; coxite 
with a subapical process with a group of long, scalelike plates. 
144 Fourth- stage larva (Figure 75) large, yellowish brown, semitrans- 
parent, with a broad head. Frontal hairs strongly developed, secondarily 
feathered, outer hairs with 8—12, median hairs with 5—7, inner hairs with 
7—9 branches, reaching to the labrum. Postclypeal hairs situated before 
the inner frontal hairs, close together. Antennae long, distinctly more than 
half as long as the head, with a hair tuft in the middle slightly nearer to the 
base, about half as long as the antenna, usually with 10 branches. Mouth 
parts of the filtration type, without comblike ends on the inner hairs of the 
labrum, but with short outer lobes. 

Comb with 60—70 or more elongate scales with wide base, narrower in 
the middle and with broad, rounded end; margin of scales with a row of 
variously developed spines. Siphon (index 3.4—3.5) weakly tapering in 
apical third. Pecten with 12- 14(usually 15—21) spinelike denticles, con- 
tinuing apically in a row of 17—29 (usually 19 — 22) longer hairs to the apical 
quarter of the siphon. Siphonal hair tuft situated near the base, between 
denticles 9—12 of the pecten, with 8 branches that are distinctly longer than 
the width of the siphon at the base. Stigmal plate resembling that of 
C. alaskaensis but distinctly smaller. 



146 



(143) 




FIGURE 75. Culiseta glaphyroptera Schin. Fourth -stage larva (after Peus): 
1 — head, dorsal; 2 — posterior end, lateral; 3 — scales of comb. 

Last abdominal segment completely surrounded by the saddle, which has 
a deep incision on the ventral side. Outer caudal hairs with 3—4 branches, 
inner caudal hairs forming a fan, usually with 14 branches. Lateral hair 
short, 2-branched. Fin with 13-16 tufts on the common base and 5 shorter 
tufts before it. The anterior 3 tufts are situated on the saddle. Gills 1.5— 
2.5 times as long as saddle, transparent, pointed at the end. 

Distribution. Mountains of Central and Southeastern Europe. 
USSR: Transcarpathians; records from other regions of the USSR (Lenin- 
grad Region and others) apparently refer to C. bergrothi. 

Biology. The species is restricted to mountain regions. The larvae 
occur in partly shaded water with a stony bottom in small mountain rivers 
and streams. They are not found in pure, transparent water with a clean, 
stony bottom with poor microflora and fauna; often common in polluted 
water with fallen leaves and rich detritus. 



3. Culiseta (Culiseta) alaskaensis Ludlow, 1906 

The species differs from other Palaearctic species of the subgenus with 
dark spots on the wings in the absence of white rings before the apex of the 
femora and in the middle of the first segment of the hind tarsi. 



147 



Head dorsally and laterally with yellowish white, contiguous scales and 
with dark brown, upright scales. Antennae of male as in C. glaphyrop- 
tera; antennae of female dark brown, 2 — 3 basal segments, especially on 
the inner side, with white scales. Proboscis and palps of both sexes with 
brown scales mixed with white scales; palps of male distinctly longer than 
the proboscis. Mesonotum with dark brown and golden scales which form 
spots or are irregularly distributed; lateral parts of mesonotum usually 
145 lighter; pleurae of thorax with spots of white scales. Legs: femora and 
tibiae anteriorly with black scales mixed with numerous white scales; 
posterior surface and apex also with white scales; tarsi black, with white 
rings at the base of segments 2—3 or segments 2—4 (hind tarsi). Wings 
with dark brown scales mixed with isolated white scales; black scales 
forming dark spots at the cross-veins and forks r2+r3, mi+m2 and m.3+4+ cui. 




FIGURE 76. Hypopygium of Culiseta a la s ka e ns is Ludl. 

Abdomen with black scales dorsally, with stripes of white scales at the 
base of the tergites which are about V3 as long as the tergite and slightly 
narrower in the middle; ventral side nearly completely with white scales. 

Hypopygium (Figure 76): sternite 8 with a comb of short spines at the 
posterior margin near the middle; coxite with a slightly convex subapical 
lobe with dense hairs; basal lobe of coxite with 2, rarely 3, strong setae, 
which are sometimes bent at an angle. Phallosome sclerotized. 

Fourth-stage larva (Figure 77) very large, yellowish brown to 
nearly black. The abdomen of young larvae often appears transversely 



148 



striated by pigment in the outer layer of the fat body, which is absent 
between the segments. 



(146) 




FIGURE 77. Culiseta a la ska ens is Ludl. Fourth -stage larva. Posterior end, 
lateral. 



Head broad. Frontal hairs secondarily feathered, outer hairs shorter, 
with 9—11 branches, median hairs with 2—3, inner hairs with 5—6 branches. 
Postclypeal hairs short, thin, 3-branched, the distance between them as long 
as the distance between the inner frontal hairs. Antennae distinctly less 
than half as long as the head; hair tuft situated at /s of the antenna from 
the base, with 8—11 branches which reach to the apex. Mouth parts of the 
mixed type. 

Comb with 35—55(46) scales of varying form but not narrower in the 
middle and with longer, dense spines at the rounded end. Siphon short, 
broad, slightly tapering apically, index 2.5—3.0, ratio of length to width at 
the apex 3.4—3.9. Pecten with 6—8 long, spinelike denticles and 3—6 rudi- 
mentary denticles at the base, continuing apically in a row of 16—18 hairs 
which are usually of the same length except for the 2—3 shorter distal hairs. 
146 Hairs not reaching apical third or quarter of siphon. A hair tuft with 7—10 



149 



weakly secondarily feathered branches at the base which are slightly more 
than half as long as the siphon. Stigmal plate very large (Figure 78). Last 
segment about half as long as the siphon, saddle surrounding it completely. 
Outer caudal hairs with 3—5 branches, middle branch as long as the siphon, 
inner hairs forming a large fan with 22—27 branches. Lateral hairs short, 
with 2—3 branches. Fin with 16—18 strongly developed tufts and 3—4 shorter 
tufts before the common base. Gills 1.5 times as long as the saddle or 
shorter, lanceolate, pointed at the end. 



(147) 




FIGURE 78. Culiseta alaskaensis Luld. Fourth-stage 
larva. Stigmal plate. 



Distribution. Boreal zone of the Palaearctic and Nearctic. The 
species occurs in the tundra, taiga and forest zone of the USSR. It also 
occurs in the northern regions of the European and Asian parts of the USSR 
(Kola Peninsula, Cape Chelyuskin). Western Europe. North America. 
South of the USSR, in mountain regions in Iran and Pakistan; the subspecies 
C.(C. ) alaskaensis indica occurs in Northern India. 



150 



Biology. The larvae are usually found in partly shaded water which 
does not dry up in summer and which has little aquatic vegetation and with 
fallen leaves at the bottom, also in clearings of broadleaved forests or 
shrubs or in open areas near them; they inhabit swamps in the tundra. 
147 Often found together with larvae of Aedes cinereus, A. caspius 

dorsalis, A. cantans, A. excrucians and others, rarely with larvae 
of Anopheles maculipennis or Culex territans. They hibernate 
as adults in tree holes, caves, cellars, often together with Culex pipiens, 
Anopheles maculipennis, Helomyzidae (Acalyptrata), Petauristidae 
(Nematocera) and Mycetophilidae. They usually leave their winter places 
earlier than other species of mosquitoes (in mid- or late April near Lenin- 
grad). They bite man rarely, usually at dusk and in bright sunlight. They 
attack reindeer in the tundra and forest-tundra. 



Culiseta alaskaensis alaskaensis Ludlow 

It occurs to the southern border of the forest zone. Specimens from 
southern regions are lighter. 



Culiseta alaskaensis indica Edwards, 1920 

Much lighter than the nominate subspecies. Palps of male often with 
light scales. Mesonotum with golden yellow scales. Abdomen often with 
broader stripes of white scales at the base of the tergites. Hypopygium 
mainly as in the nominate subspecies, but the spines at the posterior margin 
of sternite 8 are often weakly developed (only 3 spines in some specimens). 

The larvae do not differ from those of the nominate. subspecies. 

Distribution. Mountain regions of the southern USSR: Caucasus, 
Middle Asia, Uzbekistan, Tadzhikistan, Kirghizia, Kazakhstan. 

4. Culiseta (Culiseta) annulata Schrank, 1776 

The species is closely related to C. alaskaensis, but differs from it 
in the presence of a white ring in the apical third of the femora and in the 
middle of the first segment of the hind tarsi. 

Head with yellowish white, contiguous scales at the sides which are 
especially dense along the posterior margin of the eyes; head with yellowish 
brown setae dorsally, with upright scales of the same color posteriorly. 
Proboscis of male light brown, with yellowish white scales in a large part; 
148 palps of male distinctly longer than the proboscis, light brown, with broad, 
light rings at the apex of the penultimate and base of the last segment; 
hairs on palps brown, with yellowish white scales in some places; proboscis 
of female brown, with brown and white scales; palps short, brown, apical 
segments with white scales, otherwise brown scales. Antennae as in 
C. alaskaensis. Mesonotum with brown, golden and white scales which 
are broader in the lateral parts; pleurae of thorax with spots of broad, 
white scales; scutellum with white scales. Legs blackish brown, with dark 



151 



brown scales, with spots of white scales on femora and tibiae; posterior 
side of femora and tibiae with white scales in a large part; femora with a 
distinct white ring in the apical third, knees and articulation of tibia and 
tarsus with white scales, tarsi with white rings in middle of first segment 
and at base of 2nd, 3rd and 4th segment. Wings with brown scales, with 
scattered white scales in the anterior part; cross-veins and forks r2 + r3, 
mi + m.2 and 1113+4+ cui with distinct spots of blackish brown scales. Abdomen 
dark brown, with white anterior transverse stripes about ji of the length 
of the tergite; tergites with dark brown scales posteriorly; tergite 2 
usually with a narrow longitudinal stripe of white scales in the middle; 
sternites with yellowish white scales. 




FIGURE 79. Hypopygium of Culiseta annulata Schr. 



Hypopygium (Figure 79): basal lobe of coxite with 2-5 curved, strong 
setae; there is usually a subapical area densely covered with scales on the 
inner surface of the coxite. Phallosome strongly sclerotized. Spines at 
posterior margin of sternite 8 usually absent. 

Fourth- stage larvae (Figure 80) large, yellowish brown or green- 
ish brown. Head broad. Frontal hairs as in C.alaskaensis. Post- 
clypeal hairs and inner frontal hairs equally widely separated. Antennae 
distinctly less than half as long as the head, with a hair tuft with 10-15 
branches in the middle, not reaching the apex. Mouth parts of the mixed 
type, with indistinct combs at the ends of the inner hairs of the labrum. 
Comb with 40—50 or more scales which are slightly narrower in the middle. 
Siphon relatively longer and more strongly tapering apically than in 
149C. alaskaensis; index 3.6-4.0, ratio of length :width at the apex 6. 
Pecten with 11-18(13-15) spinelike denticles, the basal denticles often 



152 



rudimentary. The denticles pass distally into a row of 11 — 21(14—17) hairs 
which reach to half, rarely 2 /3,of the length of the siphon. Hair tuft situated 
slightly away from the base, usually with 9—10 weakly secondarily feathered 
branches, not half as long as the siphon. Stigmal plate as in C.alaskaen- 
s i s, but distinctly smaller. Last abdominal segment less than half as long 
as the siphon, the saddle surrounding it completely like a ring. Outer caudal 
hairs with 3 branches, median hairs half as long as the siphon, inner hairs 
with 13—19 branches, lateral hair thin, with 3 branches. Fin usually with 
18 tufts, the anterior tufts short, passing without distinct border into the 
tufts before the common base. Gills lanceolate, as long as the saddle. 




FIGURE 80. Culiseta annulata Schr. Fourth-stage larva (after Peus): 
1 — head, dorsal; 2 — posterior end, lateral; 3 — scales of comb. 



Distribution. European USSR, to the Leningrad Region and Estonia 
in the north; Caucasus and Transcaucasia (mountains); Middle Asia, 
Kazakhstan. Widely distributed in Western and Central Europe, Sweden 
and Norway; Mediterranean; North Africa; Southwest Asia. 

Biology. The habitats of the larvae are very varied, artificial and 
natural water bodies, puddles, ditches, reservoirs, barrels, and marshes. 
They tolerate moderate pollution. Sometimes found in water with duckweed. 
Hibernation normally in the adult stage; larvae may hibernate in southern 
latitudes (southern coast of the Crimea). The mosquitoes feed mainly on 
mammals but also bite man and birds to whom they transmit some plasmodia 
of birds. In the southern part of the European USSR and in the Caucasus the 
species is very common; it is rare in the north (Leningrad Region, Estonia). 



153 



150 Culiseta annulata annulata Schrank 

With the characters of the species: Hypopygium: basal lobe of coxite 
usually with 2 large setae; spines at posterior margin of sternite 8 often 
absent, if present, at most 4 spines. 



Culiseta annulata subochrea Edwards, 1921 

It differs from the nominate subspecies in the more weakly developed 
spots on the wings and the more numerous white scales, especially in the 
anterior part of the wings. Except for the narrow white stripes at the base 
of the tergites, the abdomen is nearly entirely covered with ocher-yellow 
scales, usually with some isolated brown scales; white rings on tarsi much 
wider than in the nominate subspecies. Hypopygium: basal lobe of coxite 
usually with 3— 4 strong setae; spines often present (4—8) at posterior margin 
of sternite 8. 

Fourth-stage larva large, light to dark brown, with lighter head 
and siphon. Head broad. Frontal hairs secondarily feathered, outer hairs 
with 7—12 branches, median hairs with 3 (rarely 2), inner hairs with 5—8 
branches; postclypeal hairs with 3—4 thin branches, the distance between 
them less than the distance between the inner frontal hairs. Antenna about 
half as long as the head; hair tuft situated in the middle, slightly basal to 
the middle, with 9—14 thin branches nearly reaching to the end of the 
antenna. Mouth parts of the mixed type of feeding. Comb with 30—50 
scales resembling those of C. annulata but with more densely arranged 
scales and with narrower base. Siphon as in C. annulata (index 3.3— 3.6; 
ratio of length to width at the apex 4.5—5.3). Pecten with 10—15 thin, spine- 
like denticles and 2—5 small, rudimentary spines at the base. Pecten pass- 
ing into a row of hairs (18— 26), the basal hairs 0.3—0.4 as long as the siphon. 
Hair tuft with 7—11 secondarily feathered hairs situated slightly away from 
the base, usually less than half as long as the siphon. Stigmal plate as in 
C. annulata. Last segment less than half as long as the siphon, saddle 
surrounding it completely. Outer caudal hairs with 3—5 branches; the 
ventral hairs are the longest, longer than the siphon, the inner hairs forming 
a fan with 16—19 branches. Lateral hair thin, 3-branched. Fin with 16—18 
tufts, 2—3 tufts before the common base. Development of tufts as in 
C. annulata. Gills lanceolate, with pointed end, slightly shorter than the 
saddle. 

Distribution. Widely distributed in Middle Asia. Occurs in southern 
Norway, Finland, Denmark, England, France and other countries. North 
Africa, Southwest Asia (Palestine, Syria, Iran, Iraq). The areas of distribu- 
tion of the two subspecies overlap in Western and Central Europe. 

Biology. The subspecies is typically halophilous. The larvae are 
found in pools with rich aquatic vegetation. In southern latitudes (Middle 
Asia), they occur mainly in water well shaded by trees or reeds, or in deep 
wells. The females hibernate. The larvae may apparently also hibernate, 
so that adults appear early. There are 4—5 generations per year. Rarely 
bites man. 



154 



151 5. Culiseta (Culiseta) bergrothi Edwards, 1921 
(borealis Shingarev, 1927) 

The species is characterized by uniformly black tarsi without white 
rings and indistinct dark spots on the wings; r — m situated slightly more 
distally than m— cu. 

Head with pale golden contiguous scales and blackish brown upright 
scales and setae. Proboscis with black scales with a metallic sheen. 
Palps of female with black scales and a few light scales. Palps of male 
long, with blackish brown scales and with long, yellowish brown hairs in 
the apical half (from apical part of long segment). Mesonotum markedly 
varying in color, either more or less uniformly but not densely covered with 
narrow, golden scales or with dark brown, nearly black scales with longi- 
tudinal stripes of golden scales in the posterior half of the mesonotum along 
the dorsocentral setae of the same color and with broader stripes along the 
lateral margins of the mesonotum; lateral margins of the bare space before 
the scutellum and scutellum covered with narrow, white scales; posterior 
margin of scutellum with long, blackish brown setae; pleurae of thorax with 
spots of white, lanceolate scales. Legs: femora and tibiae with blackish 
brown scales anteriorly and scattered white scales, particularly in basal 
part of femora; femora and tibiae posteriorly and hind femora also an- 
teriorly, except on the dorsal surface and an indistinct subapical ring, and 
also the apex of all femora and tibiae, with white scales. Tarsi black, with- 
out light rings; first segment of fore tarsi of male very long, usually longer 
than the other tarsal segments together. Wings with narrow, dark brown 
scales on the veins and indistinct dark spots at the base of the radial and 
medial forks, and on the cross-veins. 




FIGURE 81. Hypopygium of Culiseta bergrothi Edw. 



155 



Abdomen with black scales, with moderately broad stripes of white 
scales at the base of the tergites; dark scales on dorsal surface sometimes 
with a bluish or metallic sheen. 



(152) 




FIGURE 82. Culiseta bergrothi Edw. Fourth -stage larva: 
1 - head, dorsal; 2 — posterior end, lateral. 



Hypopygium (Figure 81): coxite long; basal lobe well developed, with 
2 strong, long setae; apical lobe absent; sternite 8 with a row of spines at 
the posterior margin. 



610144 



156 



Fourth-stage larva (Figure 82 ) large, grayish brown or yellowish 

152 brown. Head broad. Frontal hairs secondarily feathered, outer hairs with 
8— 16(10— 11 ) branches, median hairs with 5—9(6—7), inner hairs with 9—13 
(10— ll) branches; postclypeal hairs with 2—6(3—5) branches situated close 
together before the inner frontal hairs. Antennae half as long as the head; 
hair tuft with 5—14(9—11) secondarily feathered branches which reach to 
the apex of the antenna, situated slightly basal to the middle. Mouth parts 
of the filtration type. 

PComb with 50—80(60—70) narrow, long scales, narrower basally; margins 
with a row of spines, situated closer together at the distal end. Siphon 
distinctly tapering apically, of varying length. In larvae from the Far East 
the index is 3.1-4.1(3.6-3.9) and in larvae from Karelia 2.8-3.6(3.2). The 
pecten consists of 14(9—22) moderately long denticles with 2—3 accessory 
denticles at the base and 2—8(3 — 4) rudimentary denticles at the basal ring 

153 of the siphon; it continues in a row of 7—15(10—12) hairs, the 3—4 basal 
hairs shorter; the hairs extend to jz of the siphon from the base. Tuft 
with 4—9(6—7) secondarily feathered branches, as long as the last segment, 
situated near the base. Stigmal plate as in C . annul at a, more strongly 
pigmented, much smaller than in C.alaskaensis. Saddle incised on the 
ventral side, surrounding the segment completely. Outer caudal hairs with 
2—4 branches, median hairs longer than the siphon, inner hairs forming a 
fan with 13—16 branches, lateral hair with 2—6 short branches. Fin with 
13—19 strongly developed tufts and 3—4 shorter and less branched tufts 
before the common base; 1 — 2 tufts situated in the ventral incision and 2 
tufts on the saddle. Gills narrow, lanceolate, with pointed ends, usually 
twice as long as the saddle. 

Distribution. Widely distributed in the southern part of the tundra in 
the USSR (Kola Peninsula, Nenets National District), in the taiga and the 
northern part of the forest zone. In the Far East it spreads to the broad- 
leaved forest zone. Northern Europe (Sweden, Norway, Finland). 

Biology. The larvae occur in stagnant water, swamps in the tundra, 
pools in forests and ponds in inhabited areas. In the southern part of its 
distribution, the species inhabits shaded water bodies and tolerates a high 
degree of pollution. There is one generation per year in the north, 2—3 in 
the south. Females hibernate. They feed on mammals, mainly cattle, and 
sometimes enter homes and cattle sheds. 



3. Subgenus Culicella Felt 

Palps of male as long as or longer than the proboscis; last two segments 
of palps not thickened or more or less thickened, with distinct hairs; femora 
and tibiae without sharply defined light spots; wings without dark spots; 
cross-vein m— cu situated distinctly before r— m; coxite without subapical 
lobe or with a sharply defined area of dense hairs. Tergite 9 without pro- 
cesses; phallosome weakly sclerotized, its terminal part slightly curved, 
but not hook-shaped. 

Larvae large, with broad head and mouth parts of the filtration type, with 
long, curved antennae, the strongly developed tuft on the antennae situated 



157 



far distal to the middle; two long setae situated near the apex, which bears 
only one long seta. Siphon (index 5—7) with a pair of tufts at the base and a 
pecten with a few spinelike denticles. Last abdominal segment long, with a 
ring-shaped saddle surrounding it completely. Outer caudal hairs branched. 
Gills moderately long, with pointed ends. Main tracheal trunks thin, rounded. 
The subgenus contains 8 species of which 5—6 are Palaearctic. 



6. Culiseta (Culicella) morsitans Theobald, 1901 

Head with yellowish white scales and with upright, brown scales dorsally 
which gradually pass into long, brown setae anteriorly. Proboscis of male 
154 light brown, slightly darker at base and apex, with pale yellowish scales; 
palps of male distinctly longer than the proboscis, brown, with broad, light 
rings at the base and narrow light rings on the two terminal segments; 
terminal segment distinctly thickened; proboscis and palps of female dark 
brown, with blackish brown scales. Mesonotum with narrow, brown and 
golden scales (golden scales in the midline, near the dorsocentral setae and 
at the lateral margin); pleurae of thorax with spots of white scales; scutel- 
lum with golden scales. Legs brown; femora and tibiae with brown scales 
anteriorly, with yellowish white scales posteriorly; knees and articulation 
of tibia and tarsus with white spots; tarsi with narrow, white rings at the 
base of the 2nd, 3rd and sometimes 4th segment. Wings with brown scales, 
without spots. Abdomen with brownish black integument and scales, with 
narrow stripes of white scales at the base of the tergites and with yellowish 
white scales ventrally. 




FIGURE 83. Hypopygium of Culiseta morsitans Theob. 
(after Natvig). 



158 



Hypopygium (Figure 83): coxite at most 3 times as long as wide at the 
base; basal lobe of coxite with 3 — 6 strong setae; sternite 8 with an ir- 
regular row of spines or with 3—10 long, strong setae at the posterior 
margin. 



(155) 




FIGURE 84. 1 — Culiseta ochropte r a Peus, posterior end, lateral; 2 — Culiseta morsitans 
Theob., posterior end, lateral (after Peus). 



Fourth-stage larva (Figure 84) large, varying in color from semi- 
transparent to yellowish green and brown, with a pale head which is more 
than 1.5 times wider than long. Frontal hairs longer than the head, weakly 
secondarily feathered, outer hairs with 6—7 branches, median and inner 
hairs with 2—3 branches. Postclypeal hairs situated before the inner frontal 
hairs, situated closer together, short, simple. Antennae longer than the 
head, S-shaped, curved, with dark, narrow apical end and base; hair tuft 
forming a dense fan with 20—25 secondarily feathered branches, situated at 
lz of the length of the antenna from the base; subapical setae situated near 
the apex, one of them long. Comb with more than 100 closely arranged, 
long, narrow scales with slightly widened base and a row of thin spines 
distally. Siphon (index 5—7) straight, slightly tapering apically, with a 
pecten of 5—8 long, thin denticles, the distal denticle slightly separated, 
situated at about x /q of the length of the siphon from the base. Tuft with 
4—6 branches, fa as long as the siphon, situated at the base. Stigmal plate 
very characteristic (Figure 85). Main tracheal trunks thin, with round 
cross section. Last abdominal segment long, the ring-shaped saddle surround- 
155 ing it; posterior margin with spines on the dorsal side which are absent 



159 



anteriorly and laterally. Outer caudal hairs with 3 branches, the median 
hairs as long as the siphon, the inner hairs forming a fan with 14—17 branches 
branches. Lateral hair simple, half as long as the saddle. Fin with 13-14 
tufts on the common base and about 6 tufts before it, one of them situated 
in the incision of the saddle, the others on the saddle. Gills lanceolate, 
nearly half as long as the saddle; the upper pair may be slightly shorter 
than the lower. 




FIGURE 85. Culiseta morsitans Theob. Fourth-stage larva. 
Stigmal plate. 



156 Distribution. European USSR from the Baltic area and Leningrad 

Region to the Crimea and Northern Caucasus; Central Urals; West 
Siberia. Europe from Norway, Sweden and Finland to the Mediterranean. 
North Africa. Southwest Asia. 

Biology. The larvae develop in different water bodies, including 
stagnant and slightly flowing, more or less shaded water bodies with fallen 
leaves on the bottom. They often occur in temporary water bodies in 
forests or at their edge. The larvae hibernate in permanent water bodies 
but die when the water freezes. They occur together with larvae of Ano- 
pheles claviger and in spring with larvae of A e d e s . The adults are 
common in forests, mainly of the broadleaved type. During the day they 
rest among dense grass, on building sites, etc. In the north (Baltic area, 
Leningrad Region), they are common in late summer and early autumn 
(second half of July to September). They bite farm animals and man, but 
feed apparently also on wild animals and probably birds. 



7. Culiseta (Culicella) ochroptera Peus, 1935 

Head with contiguous whitish scales and with upright black scales; 
long, dark setae at the margin of the eyes. Palps of female dark brown, 
sometimes with light scales at apex of 3rd and 4th segment. Palps of male 



160 



projecting beyond apex of the proboscis for 1.5 times the length of the 
apical segment; terminal segment not thickened; palps covered mainly with 
yellowish scales, only the 1st segment, base of the 2nd and a ring at base of 
3rd segment dark. 

Mesonotum with rust brown or golden brown scales; laterally and at 
the posterior margin are narrow, crescent-shaped, whitish-brass-colored 
scales, mainly before the scutellum and also on the scutellum. Such scales 
sometimes form indistinct longitudinal stripes on the mesonotum. Legs: 
femora dark anteriorly, pale yellow posteriorly. Tarsi with narrow, light 
rings at the base of the first 3 segments and sometimes a few light scales 
at the apex of the segments; light rings on the tarsi often indistinct. Wings 
with brown scales on the veins, scales sometimes more densely grouped 
at the base of r4+5, forming a dark spot. 

The color of the scales on the abdomen is very variable. The tergites 
are usually brown with light yellow scales forming transverse stripes at 

the base and sometimes narrower stripes at the 
apex. Last tergite completely light. Sternites 
with light and dark scales arranged as on the 
tergites. Far Eastern populations have uniformly 
yellowish (sometimes brown) scales on the 
tergites. There are also specimens with narrow, 
basal, light stripes on the tergites or with isolated 
light scales scattered on the dark background. 

Hypopygium (Figure 86): lobe of coxite with 
3—9 slightly curved spines (strong setae). 
Posterior margin of 8th sternite with about 8 
spines, sometimes without spines. Phallosome 
usually ovoid, weakly sclerotized. 

Fourth-stage larva (Figures 84 and 87) 
large, light or dark brown, with yellowish brown 
head and siphon (Monchadskii, 1947). Frontal 
hairs secondarily feathered; outer hairs with 
7—13 branches, 2 /3 as long as the antenna, median 
hairs longer than the antenna, always 2-branched, 
inner hairs as long as the outer hairs, 5—9- 
branched, Postclypeal hairs with 2—4 thin 
branches. Antennae slightly shorter than the head, darkly pigmented at the 
base and in the tapering apical part, curved where the long subapical setae 
are situated, distinctly away from the apex; tuft with 20—37 branches. 
Comb with 60—95 scales, some of them with a longitudinal, dark ridge; the 
narrowing in the middle, if present, is less distinct than in C.morsi- 
tans. Siphon long (index 5.7-7.0), slightly tapering apically. Pecten with 
2—7 denticles, the 2—3 distal denticles spinelike, more widely separated and 
with 2—4 rudimentary denticles at the base. Tuft with 5—10 branches, 
situated at the base, /$— l /z as long as the siphon. Branches forming a 
divergent fascicle, not a fan. Stigmal plate characteristic (Figure 87). 
Last segment long, saddle surrounding it like a ring. Outer caudal hairs 
with 2—3 branches, less than % as long as the siphon; inner hairs forming 
a fan with 12—23 branches; lateral hair long and simple. Fin with 10—22 
tufts on the common base and 6—8 shorter tufts before it, some of them, 
situated on the saddle. Gills narrow, lanceolate, long (1 — 2.6 times as long 
as the saddle ). 




FIGURE 86. Hypopygium of 
Culiseta ochroptera Peus 



161 



157! 




FIGURE 87. Culiseta ochroptera Peus. Fourth -stage 
larva: 

1 — head, dorsal; 2 — stigmal plate. 



Note on systematics. A. V. Maslov (1964) considered A. ochro- 
ptera as a synonym of C. silvestris Shingarev, 1928, but to judge from 
Shingarev's incomplete description, "it is certain that the two species are 
not identical" (Shtakel'berg, 1937). The holotype of C. silvestris is 
lost and Maslov examined only paratypes. Because of the incomplete 
original description of C. silvestris, it seems advisable to retain the 
name C. ochroptera. 

Maslov (1964) suggests that the species be divided into four subspecies: 
C. silvestris silvestris Shing. (Eastern Europe, Urals, West 
Siberia), C. s. ochroptera Peus (Central Europe, Baltic area), 



162 



C. s. amurensis Masl. (Amur area and Maritime Territory), and 
C. s. minnesotae Barr. (North America). He gives a key to the sub- 
species based on females, males and larvae; however, this division into 
subspecies is provisional because of the inadequate study of the species. 




FIGURE 88. Culiseta ochroptera Peus. Fourth-stage larva: 

1 — posterior end, lateral; 2 — denticles of pecten; 3 — scales of comb. 



159 



Distribution. Forest zone of the Palaearctic, from Western Europe 
to the Maritime Territory and Northeast China (the Japanese species 
Culiseta (Culicella) nipponica La Casse and Yamaguti may also 
refer to C. ochroptera; the species was described from larvae from 
Hokkaido. The northern and southern boundaries of its distribution have 
not been determined. In the USSR, C. ochroptera occurs in the 
Vladimir Region (Saf 'yanova, ^SO), Urals, West Siberia, and some other 
areas in addition to the Far East. 

Biology. The larvae occur only in large, shallow marshes, in forest 
clearings, and on the muddy shores of lakes; they apparently hibernate. 
But in the eastern Ukraine females hibernate (Val'kh, 1959). The mosquitoes 



163 



are found mainly in wild nature and feed on birds, rarely mammals; they 
sometimes bite man. 



8. Culiseta (Culicella) fumipennis Stephens, 1825 

It is closely related to C.morsitans, but differs in the presence of 
white rings on all tarsal segments, a V-shaped pattern formed by dark 
scales on most abdominal sternites, and pale scales in the middle of the 
proboscis. 




FIGURE 89. Hypopygium of C ulise ta fumipennis 
Steph. (after Natvig) 



160 Hypopygium (Figure 89): coxite 3—3.5 times longer than wide at the 
base. Basal lobe of coxite with 3—4 1 strong setae; posterior margin of 
sternite 8 with a few spines. 

Fourth-stage larva (Figure 90) large, semitransparent, greenish 
or yellowish brown, with a broad head. Frontal hairs secondarily feathered, 
outer hairs with 5—6 branches, median hairs with 2, inner hairs with 2—4 
branches. Antennae longer than the head, with a tuft with 10—13 secondarily 
feathered branches at the base of the tapering apical third of the antenna. 
Subapical setae situated some distance from the apex. Comb with 120—160 
long, narrow scales which are wider distally and have thin spines at the 
margin. Siphon wider at the base, tapering at first distinctly and then 
gradually apically; the length : width index (about 5 ) of the base does not give a 
correct picture of the length of the siphon. Pecten with 7—8 well developed 
and 4—6 rudimentary, densely grouped spaced denticles in an oblique row arour 
the base of the hair tuft; the pecten continues in a row of 2—5(3 — 4) large, 



164 



smooth, strongly pigmented spines, which are irregularly distributed and 
absent in the apical third. Hair tuft situated near the base, with 4-5 smooth 
branches, half as long as the siphon. Stigmal plate with strongly developed, 
hook-shaped setae on the posterior valves and large hairs on the lateral 
valves in the form of a partly spread fan with 8-12 branches. Main 
tracheal trunks narrow, with round cross section. 



(159; 




FIGURE 90. Culiseta fumipennis Steph. Fourth- 
stage larva. Posterior end, lateral (after Seguy). 

Last segment long, surrounded by the ring-shaped saddle. Outer caudal 
hairs with 3 branches, median hairs thick, longer than the siphon, inner 



165 



caudal hairs forming a fan with 14—15 branches. Fin with 14 tufts and 6 
shorter, less branched tufts before it. Gills lanceolate, more than half as 
long as the last segment. 

Distribution. European USSR (Estonia, Leningrad Region, central 
Urals, Ukrainian Polesie, Northern Caucasus). All records of adult 
mosquitoes from the USSR need confirmation. Larvae of C. fumipennis 
have not been found in the USSR. Western Europe to southern Sweden and 
Norway. Mediterranean, including Algeria. 

Biology. The larvae develop in the spring in shallow water with green 
vegetation and also in water covered with Lemna minor and L.tri- 
s u 1 c a, often together with larvae of Culiseta morsitans, Culex 
territans and Culex hortensis (southern parts of the range), rarely 
with larvae of A n o p he 1 e s claviger. They feed on plankton and spend 
most of their time submerged. 



9. Culiseta (Culicella) setivalva Maslov, 1937 

Closely resembling C. fumipennis in coloration, differing from it in 
the structure of the hypopygium and particularly in the absence of spines in 
the middle of the posterior margin of sternite 8 and in the presence of long, 
strong setae on the coxite. 

Head with contiguous whitish scales and upright blackish- brown scales 
and setae. Proboscis of male light brown, slightly darker toward the base 
and at the apex, with light yellow scales; palps long, projecting beyond the 
proboscis for about the length of the apical segment, with brown scales 
mixed with light scales which form more or less broad rings near the middle 
161 of the long segment and at the base of the subapical and apical segment. 
Hairs on palps of male brownish, especially long in the apical half of the 
long segment; hairs at base of subapical and apical segment light yellow. 
Proboscis and palps of female with dark brown scales, with only a few 
scattered light scales. Mesonotum and scutellum with sparse, narrow, light 
golden scales; pleurae of thorax with small spots of broad, white scales. 
Legs (femora and tibiae) dark brown anteriorly, without or with a few light 
scales, white posteriorly; hind femora sometimes with an anterior longi- 
tudinal stripe of light scales; apex of femora and tibiae with yellowish 
scales; narrow white rings at the base of tarsal segments 1 — 3 or 1—5. 
Tarsi of fore legs of male short, distinctly shorter than the tibia; first 
segment of fore tarsi about as long as or slightly shorter than the other 
tarsal segments together. Wings with dark brown scales on the veins. 
Abdomen with dark scales dorsally and stripes of yellowish white scales 
at the anterior margin of the tergites, about fa— fa as long as the tergite; 
venter with yellowish white scales with more or less numerous dark scales 
which sometimes form an inverted V as in C . f u m ip e n n i s. 

Hypopygium (Figure 91): sternite 8 without spines at the posterior 
162 margin, with 2—4 long, strong setae lateral to the median process (one at 
each side) and similar setae (about 8), behind the middle of the sternite; 
coxite with very long, strong setae, 2—3 on the inner side of the coxite; 
basal lobe small but distinct, with 2 strong setae at the apex; style long and 
thin. 



166 



(161) 




FIGURE 91. Hypopygium of Culiseta setivalva Masl. 

Fourth-stage larva resembling that of C. fumipennis, but 
differing in a number of characters, mainly in the more weakly developed 
hairs of the posterior (and, possibly, the anterior) end. Comb with over 
100 scales. The median hair behind the comb is distinctly shorter than 
the last abdominal segment. The additional, widely separated distal 
denticles of the pecten on the siphon are weakly developed; the tuft at the 
base of the siphon is more strongly developed. On the dorsal side of the 
posterior margin of the 8th abdominal segment (rudiment of 9th segment) 
is a narrow, transverse, chitinized arc. The 7—8 tufts before the fin are 
slightly shorter than the anterior tufts on the common base. Outer caudal 
hairs with only 3 branches, one hair longer than the others. Gills more than 
half as long as the last segment. 

Distribution. Southern European USSR (south coast of the Crimea), 
Black Sea coast of the Caucasus, Asia Minor (Anatolia). 

Biology. According to Velichkevich (1936), the larvae breed in shallow, 
silty water bodies overgrown with vegetation, rich in detritus, stagnant or 
slowly flowing and with a high concentration of calcium (101.2 mg per liter). 
These water bodies are shaded by trees and shrubs on their banks in summer; 
they are warmed by the sun in autumn and winter, after leaves have fallen. 



167 



One generation per year. Larvae of the 3rd and 4th stage hibernate and 
pupate at the end of April. Flight begins in early May. The water bodies 
dry up nearly completely in summer. Eggs deposited in the water or on 
moist ground are in diapause. In autumn, when rain fills the water bodies, 
the larvae hatch arid reach the 2nd to 4th stage in December, and then enter 
hibernation. The larvae are found together with larvae of Culiseta 
morsitans, C.annulata, Culex pipiens, C.territans, C.hor- 
tensis, and Anopheles maculipennis. The females bite man and 
animals during the day and at dusk, particularly in shaded places and among 
shrubs. They do not fly far from their breeding places, resting in thickets 
of reeds, Sparganium, in shrubs and tree holes. They do not enter 
buildings. 



6. Genus Mansonia Blanchard 

Eyes contiguous or nearly contiguous. Proboscis moderately long, more 
or less uniformly thick. Palps of male as long as or longer than the 
proboscis; palps of female short, not more than ,4 as long as the proboscis. 
Mesonotum with well developed acrostichal and dorsocentral setae. 
Spiracular setae absent, postspiracular setae present or absent; upper 
sternopleural setae well developed; upper mesepimeral setae usually 
present. Legs: 1st segment of hind tarsi shorter than hind tibiae; fore 
tarsi of male with one larger claw and usually with 2 denticles, the other 
claw smaller and usually without denticles; claws of female simple; pulvilli 
163 absent. Wings with relatively broad scales on the veins. Forks of wings 
comparatively long. Postalar sclerites with scales. Coxite with well 
developed basal lobe. Eighth segment of female short and broad; cerci 
short, not projecting. 

The larvae have a Very broad head, long antennae with a whiplike apical 
part, subapical setae situated far from the end and with a strongly developed 
tuft. Mouth parts of the filtration type. Thorax with strongly developed 
lateral hair tufts, with chitinized bases on the mesothorax and metathorax. 
The lateral branches of the main tracheal trunks are widened in the meta- 
thorax. The short siphon has valves which are transformed into a piercing 
apparatus which receives air from the air spaces of aquatic plants. Last 
segment long, with a ring-shaped saddle surrounding it and with strongly 
developed fin and caudal hairs. 

There are 60 species of Ma ns o ni a, most of them from the Tropics of 
the eastern and western hemispheres. Only three species occur in the 
Palaearctic: one of them (M . richiardii Fie.) is widely distributed in the 
USSR, another (M . buxtoni Edw.) is rare in the Mediterranean and occurs 
in the southwestern USSR, and a tropical species (M.uniformis Theob.) 
reaches the Palaearctic in the southern islands of Japan (Kyushu, Honshu). 
The genus is divided into 4 subgenera. The two species in the USSR belong 
to the subgenus Coquillettidia. 



168 



Subgenus Coquillettidia Dyar 

Postspiracular setae absent. Seventh abdominal segment of male large, 
8th segment not large but distinct, without chitinized hooks. Hypopygium: 
basal lobe of coxite with a rod- shaped or other appendage that is as long 
or longer than the lobe; style with a short terminal appendage. 

The subgenus contains about 35 species, most of them in tropical Africa, 
5 in the Oriental region, 5 in Australia and New Zealand, one in America, 
and 2 in the Palaearctic. 



Females 

1 (2). Tarsi with light rings. Proboscis usually with a light ring in the 

middle. Scales on wings relatively broad 

1. M. (C.) richiardii Fie. 

2 (l). Tarsi without light rings. Proboscis without light ring. Scales on 

wings relatively narrow 2. M. (C.) buxtoni Edw. 



Males 

1 (2). Style broad at the base, with a constriction in the middle 

1. M. (C.) richiardii Fie. 

2 (l). Style narrower at the base, without a constriction 

2. M. (C.) buxtoni Edw. 



Larvae 

1 (2). The dorsal hair of the hairs behind the comb on the 8th abdominal 

segment with 2—4 branches. Saddle covered with isolated chaetoids, 
especially on the dorsal side 1. M. (C. ) richiardii Fie. 

2 (l). The dorsal hair of the hairs behind the comb on the 8th abdominal 

segment with 5—7 branches. Saddle with shorter chaetoids, form- 
ing groups of 2—8 in a row 2. M. (C.) buxtoni Edw. 



1. Mansonia richardii Ficalbi, 1889 

The species is characterized by the broad wing scales and by a white rin£ 
on the proboscis and in the middle of the first segment of the hind tarsi in 
most specimens (Figure 92). 

Head with yellowish white contiguous and brown upright scales. Pro- 
boscis with brown scales, with a ring of whitish scales in the middle which 
is broader in the female. Palps of female with brown and yellowish white 
scales; they are usually grouped in the male on the basal part of the seg- 
ments and form a light ring. Mesonotum with brown and golden scales; 
scutellum with golden scales; pleurae of thorax with spots of broad, yellow- 
ish white scales. Legs: femora and tibiae with brown and yellow scales 
anteriorly; tarsi brown, usually with broad white rings at the base of the 



169 



(164) 




FIGURE 92. Mansonia richiardii Fi 



170 



segments (particularly on the hind tarsi); first segment of hind tarsi with 
a white ring in the middle, ring rarely indistinct or absent or legs mainly 
with light scales. Such variation (var. nikolskyi Shing.) are distributed 
in Middle Asia and Kazakhstan. Claws of male with denticles (2.0; 2.0; 
0.0), claws of female simple. Wings slightly gray, veins with relatively broad, 
white and brown scales. Abdomen with brown scales and triangular spots 
of yellowish white scales at the sides of the tergites. 




FIGURE 93. Hypopygium of Mansonia richiardii Fie. (after Coe, Freeman and Mattingly). 
Style of M.richiardii (A) and M.buxtoni Edw. (B) (after Aitken). 



Hypopygium (Figure 93): coxite thick, short, with a lobe bearing a strong 
sclerotized spine with blunt end. Style irregularly widened at the base and 
166 in the apical third. Tenth sternite with a few denticles at the apex. Phallo- 
some markedly sclerotized, with denticulate margin. Processes of tergite 
9 with 8—10 thin setae. Sternite 8 with a group of 10—15 strong, straight 
setae situated close together. 

Fourth- stage larva (Figure 94) yellowish or yellowish brown. 
Head broad but not narrowing near the base of the antennae. Outer frontal 
hairs with 9 branches, the median hairs with 4—5 branches, long and 



171 



secondarily feathered, inner hairs short, resembling the postclypeal hairs, 
situated anterior to them; both pairs usually with 5—6 branches. Antennae 
1.2—1.8 times longer than the head because of the long terminal part. 
The subapical setae are situated far from the end; tuft with 15—20 sec- 
ondarily feathered branches. 




2SS3T 



FIGURE 94. Hypopygium of Mansonia 
richiar d ii Fie. Fourth-stage larva. 
Head, dorsal. 




FIGURE 95. Mansonia richiardii Fie. Fourth-stage larva. 
Siphon, lateral (after Wesenberg-Lund). 



172 



Abdomen tapering posteriorly. Comb consisting of an irregular row of 
scales (more than 25), with a well developed main spine. Hairs behind the 
comb forming two groups: a dorsal group of 2 hairs situated near the dorsal 
scales of the comb and a ventral group of 3 hairs situated near its middle; 
dorsal hair with 2—4 long branches, median hair with 2—3 branches. 

Siphon very short, in the form of a truncate cone. Auricles at the base, 
pecten and hair tufts absent. Valves and "stirrup" of stigmal plate forming 
a piercing apparatus, the hairs of the plate forming long spines. 

Last segment elongate, surrounded by the ring-shaped saddle which is 
wider posteriorly. It is covered, especially on the dorsal side near the 
posterior margin, with short, isolated chaetoids. Caudal hairs forming a 
partly spread fan; outer hairs longer than inner hairs. Lateral hair with 
2—3 branches. Fin displaced posteriorly, with 10—14 long tufts. Gills 
lanceolate, with pointed ends, not longer than widest part of last segment. 

Distribution. Widely distributed in the western Palaearctic. Re- 
corded from the Leningrad Region (north to Priozersk), Estonia, Latvia, 
Moscow Region, Urals, Ukraine, Northern Caucasus, Transcaucasia, Middle 
Asia (Turkmenia, Uzbekistan, Tadzhikistan, Kirghizia), Kazakhstan and the 
southern part of West Siberia. Widely distributed in Europe, from 
southern Sweden, Finland, Denmark and England to Italy, Hungary, Bulgaria 
and Rumania. Southwest Asia (Palestine). 

Biology. The larvae occur in different permanent water bodies with 
rich vegetation (Acorus, Carex, Glyceria, Ranunculus, Spar- 
ganium, Ty ph a, etc.) from the air spaces of which they obtain air. They 
live submerged and move very little. Their specific gravity is higher than 
that of the water. They remain horizontal during swimming because of the 
tracheal air sacs. There is one generation per year in the northern part 
of their distribution, 2—3 generations in the south. The larvae hatch in 
autumn and usually hibernate in the 3rd or 4th stage. Pupation takes place 
in May— June. The pupae have modified breathing horns and, like the 
larvae, pierce the stems and roots of plants at the bottom of the water. 
Before the mosquito hatches, the end of the horns breaks off and the pupa 
comes to the surface. In the south (Middle Asia) they breed in large num- 
bers in permanent water bodies in the floodplains of the middle and lower 
reaches of large rivers, in fresh and slightly saline water, often together 
with larvae of Anopheles pulcherrimus. The mosquito is an active 
bloodsucker in Middle Asia and in some areas (floodplains of large rivers) 
distinctly more numerous than other species of mosquitoes. They bite 
mainly in wild nature, rarely entering buildings. 



2. Mansonia buxtoni Edwards, 1923 

It differs from M. richiardii in the absence of rings on the tarsi 
and the dark proboscis and palps. 

Head with yellowish white contiguous scales and brown upright scales. 
Proboscis and palps with brown scales, which are partly upright. Proboscis 
about as long as the fore femora. Mesonotum with golden brown scales, 
lighter before the scutellum and at the base of the wings. Legs: femora 
and tibiae with dark scales mixed with light scales which form indistinct 



173 



longitudinal stripes; apex of femora and tibiae with white scales; tarsi 
brown; hind tibia distinctly longer than first tarsal segment; claws of 
female simple. Wings with dark scales, which are narrower than in 
M.richiardii (but broader than in Culex). Abdomen: tergites with 
168 brown scales and distinct, whitish-cream spots at the sides; the spots are 
connected by narrow stripes at the anterior margin of the tergites on some 
segments; sternites with light stripes at the base. Hypopygium resembling 
that of M . r i c hi a rdi i, differing only in the form of the style (Figure 93); 
it is comparatively broad at the base, without a constriction in the middle, 
the apical half markedly swollen. 




FIGURE 96. Mansonia buxtoni Edw. Fourth-stage larva: 

1 — head, dorsal; 2 — posterior end, lateral; 3 - groups of chaetoids on last segment . 



174 



Fourth-stage larva (Figure 96) closely resembling that of 
M.richiardii. It is light green, 5— 7 mm long, head 1.4 times wider than 
long. Frontal hairs: outer hairs moderately long, with 9 secondarily 
feathered branches, median hairs nearly twice as long, with 5—7 secondarily 
feathered branches; inner hairs short, with 8 simple branches re- 
sembling the postclypeal hairs situated before them. Antennae as in 
M. richiardii. 

Comb of 8th abdominal segment with 16-22 scales in an irregular row 
(partly double dorsally) and with a pointed main spine. The dorsal hair of 
the hairs behind the comb has 5—7 (average 6) branches, the median hair 
and intermediate hairs with 2 branches, the latter half as long as the median 
hair. 

The last segment resembles that of M. richiardii but it is covered 
with rows of short chaetoids in groups of 2—8. Lateral hair with 4 branches. 

Distribution. Israel, Syria, Sardinia, Corsica, Morocco. It occurs 
in the USSR only in the Chernovtsy Region (Gutsevich, Donets, Ezhova and 
Popov, 1962). 

Biology. Little known. The mosquitoes bite man in the open. The 
larvae occur in water between the roots of A c o r u s and Typha. 



7. Genus Aedes Meigen, 1818 

Spiracular setae absent, postspiracular setae present. Claws of female 
usually with denticles on all legs. Denticles present either on two pairs or 
on one pair of legs in some species. Pulvilli absent. The first segment 
of the hind tarsi is shorter than the tibia. Posterior end of abdomen of 
female more or less pointed, cerci more or less projecting. 

Larvae with mouth parts for feeding on plankton or periphyton. An ex- 
ception are the predaceous larvae of the tropical subgenus Mucidus. 
Antennae rod-shaped, usually short, with a hair tuft near the middle or basal 
to it. Median and lateral hairs of prothorax moderately developed. 
Tracheal air sacs and main tracheal trunks with round cross section absent. 
Dorsal chitinized plates absent, except the saddle on the last segment. 
Siphon relatively short (index at most 4), always with a pecten and a pair of 
hair tufts on the posterior surface, rarely with additional hairs on the 
anterior side and on the sides of the siphon. Stigmal plate usually with a 
2-branched posterior process of the "stirrup." 

About 700 species of this large genus have been described. Forty-two 
species occur in the USSR and form half of the species of mosquitoes in 
the Soviet Union. Aedes predominates in number of species and speci- 
mens in the tundra and in forest belts. 

The genus is divided into more than 20 subgenera of which 5 occur in the 
Palaearctic: Ochlerotatus, Aedimorphus, Finlaya, Stegomyia, 
and Aedes. The subgenera are distinguished by the structure of the 
hypopygium, less distinctly by the females. 

The larvae are grouped differently than the adults. The larvae of the 
subgenera Aedes, Ochlerotatus and Aedimorphus are very 
similar. On the other hand, the larvae of some species of the subgenus 
Ochlerotatus resemble those of Stegomyia and many larvae of 



175 



Stegomyia those of F inlay a. Only a general key of the larvae is there- 
fore given and no special key for the larvae of the different subgenera. 

The species of Aedes cause great harm as bloodsuckers and vectors 
of diseases of man and animals. 



170 Key to Species 
Females 

1 (50). Tarsi with light rings, which are sometimes very narrow, more 

distinct on the hind tarsi (light rings better visible against a dark 
background). 

2 (13). Each light ring of the tarsi extends to 2 segments: apex of one 

and base of following segment (Figure 97, A). 

3 (6). Wing veins with light and dark scales. 

4 (5). Abdomen with a dorsal longitudinal light stripe, sometimes nearly 

completely covered with light scales 

1. A. (Ochlerotatus) caspius Pall. 

a (b). Mesonotum with golden or rust brown scales, with 2 longi- 
tudinal light stripes. Wing veins with light and dark scales .... 

A. caspius caspius Pall. 

b (a). Mesonotum with a brown longitudinal stripe; lateral parts 
with silvery gray or cream- colored scales. Light scales on the 
wing forming groups on the veins A. caspius dorsalis Mg. 

5 (4). Abdomen without longitudinal light stripes. (The species should 

be determined by the larvae and the males for greater certainty) 
2. A. (Ochlerotatus) mariae Serg. 

6 (3). Wing veins with uniformly dark scales. 

7 (8). Light rings on tarsi very narrow, present only on hind legs, some- 

times also on midlegs. Palps of both sexes with dark scales . . . 
38. A. (Finlaya) alektorovi Stack. 

8 (7). Light rings broader, present on all tarsi. Palps of female dark, 

with white apex, with white rings in the male. 

9 (l0). Last tarsal segment white. (Mediterranean) 

3. (A. (Ochlerotatus) pulchritarsis Rond. 

a (b). Mesonotum without longitudinal light stripe 

A. pulchritarsis pulchritarsis Rond. 

b (a). Mesonotum with a narrow longitudinal stripe of white 

scales A. pulchritarsis asiaticus Edw. 

Last tarsal segment entirely dark or dark with a light ring at the 
base. (Far East.) 

Mesonotum with a large white spot 

37. A. (Finlaya) seoulensis Yam. 

Mesonotum without a white spot . . . 39. A. (Finlaya) togoi Theob. 
Light rings present only on base of tarsal segments (Figure 97, B). 
White rings on tarsi very broad, only apex of segments dark. In addi- 
tion to contiguous scales there are also dark upright scales on the 

hind tibiae 32. A. (Ochlerotatus) kasachstanicus Guts. 

15 (14). Light rings on tarsi less broad, not wider than fa of the length of 

the segment, usually narrower. Hind tibiae without upright scales. 



176 



10 


(9) 


11 


(12) 


12 


(11) 


13 


(2) 


14 


(15) 



16 (37). Proboscis distinctly longer than fore femora. Scutellum with 

yellowish or whitish, narrow, curved scales. 

17 (20). Cerci short, little projecting. Light spots at sides of abdomen 

formed by silvery white scales. 

18 (19). White rings present only on first 3 segments of hind tarsi 

40. A. (Finlaya) japonicus Theob. 

19 (18). White rings present on 4 segments of hind tarsi 

41. A. (Finlaya) koreicus Edw. 

20 (17). Cerci longer, distinctly projecting. Light abdominal scales with- 

out silvery sheen. 

21 (22). White rings on tarsi very narrow, usually not more than /$ of the 

length of the segment (Figure 97, C) ................... 

33. A. (Aedimorphus) vexans Mg. 





FIGURE 97. Tarsi of Aedes: 

A — A. c asp i us Pall.; B- 
A.cantans Mg.; C — 
A .vexans Mg. 



FIGURE 98. Spots of scales on pleurae of thorax of Aedes: 

pe — proepimeral; lis - hypostigmal; ps - parastigmal; 
ms - metastigmal; pc - postcoxal; me - mesepisternal 
(sternopleural); msp — mesepimeral. 



177 



(172) 




FIGURE 99. Arrangement of spots of scales on pleurae of thorax of A edes (after 
O.N.Sazonova): 

A -A.cataphylla Dyar; B— A.pullatus Coq.; C - A . intrudens Dyar; 
D — A.punctor Kirby; E — A.nigrinus Eck.; F — A . diantaeus H.D.K. 



178 



172 



173 



a (b). Posterior half of abdominal tergites with uniformly dark 

scales A.vexans vexans Mg. 

b (a). Tergites with light spots in posterior part of segments, 

sometimes forming an interrupted longitudinal stripe 

A. vexans nipponii Theob. 

22 (21). Light rings on tarsi broader, not less than /a of the length of the 
middle segment of the hind tarsi (Figure 97, B). The species of 
the group cantans cannot always be determined with certainty 
by the females.* 

23 (26). Abdomen with light scales dorsally, sometimes mixed with isolated 
dark scales. 

24 (25). Females usually ocher yellow. Mesonotum with silverj yellow 

scales. Pleurae of thorax with cream- colored scales, not dif- 

»fering distinctly in color from the scales on the mesonotum. 
Lower mesepimeral setae present 

1.1. A. (Ochlerotatus) cyprius Ludl. 

25 (24). Females yellowish gray. Mesonotum with small, rust brown 

scales. Pleurae of thorax covered with grayish white scales 
which differ sharply in color from the dark scales on the meso- 
notum. Lower mesepimeral setae absent 

10. A. (Ochlerotatus) flavescens Mull. 

26 (33). More or less numerous (not isolated) dark scales on the abdominal 
tergites, dark scales sometimes predominating. 

27 (28). Abdomen almost completely with dark scales dorsally, without 
light transverse stripes; light scales (females) usually forming 
diffuse spots in the midline of the body. Mesonotum usually with 

small, bronze or rust-colored scales 

6. A. (Ochlerotatus) behningi Mart. 

28 (27). Abdomen with more or less light transverse stripes dorsally, if 

stripes absent, light scales scattered, not forming spots. Meso- 
notum of different color, with golden scales and light scales 
and with a dark longitudinal stripe or with dark brown scales and 
with indistinct light spots. 

29 (32). General coloration comparatively dark. Mesonotum mainly with 

dark brown scales or with lighter scales and with a dark longi- 
175 tudinal stripe. Pleurae of thorax with white scales. Abdomen 

with light transverse stripes dorsally which are usually distinct 
on the anterior segments. 

30 (31). Mesonotum mainly with chocolate brown scales in small, indistinct 

spots, sometimes with a longitudinal dark stripe. White rings on 
middle segments of hind tarsi usually less than half as long as the 

segment. Claws sharply curved 

4. A. (Ochlerotatus) cantans Mg. 

31 (30). Mesonotum with a more or less distinct dark longitudinal stripe, 

lateral parts with golden scales. White rings on middle segments 
of hind tarsi at least half as long as the segment. Claws slightly 
curved 5. A. (Ochlerotatus) riparius D. K. 

* In the determination of the species of the cantans group and particularly of the communis group, 
the arrangement of spots of scales on the pleurae of the thorax (Figures 98—101) and the form of the 
claws (Figure 102) are important. 



179 



174 



(173) 




FIGURE 100. Arrangement of spots of scales on pleurae of thorax of A e d e s (after 
O.N.Sazonova): 

A — A .behningi Mart.; B — A . bekle mishev i Den.; C - A .excrucians 
Walk.; D— A.ripariusD.K.; E — A.cyprius Ludl.; F - A.flavescens Mull. 



a (b). Light scales among the dark scales on the wings, proboscis 

and palps A. riparius riparius D. K. 

b (a). Wings, proboscis and palps with dark scales 

A. riparius ater Guts. 

32 (2 9). General coloration lighter. Mesonotum usually with mainly golden 
or cream- colored scales, sometimes with a dark longitudinal 



180 



stripe. Pleurae of thorax usually with yellowish scales. Abdomen 
often without light stripes dorsally, if stripes are present they are 
formed by yellowish scales. 




FIGURE 101. Arrangement of spots of scales on pleurae of thorax of Aedes (after 

O.N.Sazonova): 

A - A. impiger Walk.; B-A.caspius dorsalis Mg.; C- A.vexans Mg. 

D-A.cinereusMg.; E-A.geniculatus 01. 



181 



^ c*£ ^^ ^ 5h 





^ ^ 



^^^ 



L — # c — is 



38 


(41 


39 


(40 


40 


(39 


41 


(38 




176 33 (36). 

34 (35). 

35 (34). 

36 (33). 

37 (16). 



FIGURE 102. Claws of species of A edes (after O.N. Sazonova): 

1— A.cinereus Mg.; 2 — A.impiger Walk.; 3 — A .nigripes Zett.; 4 — 
A.cataphylla Dyar; 5— A.leucomelas Mg.; 6— A.pullatus Coq.; 7 — 
A .intrudens Dyar; 8 - A . punc tor Kirby; 9 — A .h ex od on t us Dyar; 10 — 
A .pionips Dyar; 11— A. communis Deg.; 12 — A.nigrinusEck.; 13 — 
A.diantaeus H.D.K.; 14 — A.sticticus Mg.; 15 — A.caspius dorsalis Mg.; 
16 — A.vexansMg.; 17 — A.behningi Mart.; 18 — A.cantansMg.; 19 — 
A .riparius D.K.; 20 - A . bekle m i sh e v i Den.; 21 — A . ex cru ci ans Walk.; 
22 — A.cyprius Ludl.; 23 — A .flavescens Mull-; 24 — A.geniculatus 01. 

. Mesonotum covered mainly with golden scales, without distinct 

longitudinal dark stripe. 
. Claws sharply curved, large; denticle nearly parallel to apical 

part of claw 7. A. (Ochlerotatus) excrucians Walk. 

Claws less sharply curved, with a large, widely diverging denticle. 

(The species can be reliably determined only by the larvae) .... 

8. A. (Ochlerotatus) beklemishevi Don. 

Mesonotum with a distinct longitudinal dark stripe, lateral parts with 

grayish yellow scales 9. A. (Ochlerotatus) annulipes Mg. 

Proboscis not longer than fore femora. Scales of scutellum 

broad, straight, silvery white. 

Mesonotum with snow white spots. 

Mesonotum with 4 round, white spots in the middle 

44. A. (Stegomyia) vittatus Big. 

Mesonotum with 2 triangular or semicircular white spots at sides 

of anterior half 48. A. (Stegomyia) chemulpoensis Yam. 

Mesonotum with one or several longitudinal white stripes. 



182 



42 (43). Mesonotum with 4 narrow longitudinal stripes, the lateral stripes 

more distinct, curved in the anterior half 

42. A. (Stegomyia) aegypti L. 

[42). Mesonotum with a silvery white median longitudinal stripe. 
[45). Last segment of hind tarsi moderately or completely dark at the 

apex 45. A. (Stegomyia) galloisi Yam. 

,44). Last segment of hind tarsi white. 

[47). Claws of fore and mid-tarsi of female with a denticle 

(Mediterranean) 43. A. (Stegomyia) cretinus Edw. 

[46). Claws of fore and mid-tarsi of female without a denticle (Far 

East). 
[49). A small, light spot at the margin of the mesonotum above the base 

of the wing, formed by broad, straight, silvery white scales .... 

46. A. (Stegomyia) albopictus Sk. 

[48). An indistinct, light spot above the base of the wing, formed by 

narrow, curved, whitish or yellowish scales 

47. A. (Stegomyia) flavopictus Yam. 

(l). Tarsi without light rings. 
,96). Proboscis distinctly longer than fore femora. 
[57). Cerci of female short, little projecting. Light spots on abdomen 

shining, silvery. 
[54). Mesonotum with 2 large, silvery white spots, sometimes fused into 

one spot 36. A. (Finlaya) nipponicus La Casse and Yamag. 

[53). Mesonotum with abroad, longitudinal dark stripe, without silvery 

white spots. 
[56). Scutellum with narrow yellowish scales 

34. A. (Finlaya) geniculatus Ol. 

,55). Scutellum with broad white scales 

35. A. (Finlaya) echinus Edw. 

[52). Cerci long, distinctly projecting. Light spots on abdomen without 

silvery sheen. 
[69). Light scales present or predominating in apical half of tergites. 
v.62). Scales in dorsal part of proepimeron broad, straight, black. 

Mesonotum with a broad, longitudinal, dark stripe or with 2 dark 

stripes close together. 
,61). Light stripes on abdominal tergites not sharply defined, and not 

widened in the middle; abdomen sometimes with light and dark 

scales dorsally, light scales usually predominating 

13. A. (Ochlerotatus) refiki Med. 

,60). Light stripes on abdominal tergites often distinct, usually with a 

process in the midline of the body 

12. A. (Ochlerotatus) rusticus Rossi. 

,59). Scales in upper part of proepimeron often narrow, curved, but if 

straight, yellowish or light brown, not black. Mesonotum usually 

without dark longitudinal stripe. 
,66). Light scales on abdominal tergites not forming transverse stripes; 

abdomen completely covered with light scales dorsally or among 

them are more or less numerous dark scales, sometimes forming 

indistinct spots. 



183 



64 


(65) 


65 


(64) 


66 


(63) 


67 


68) 


68 


67) 


69 


58) 


70 


73) 


71 


72) 


72 


71) 


73 ( 


70) 


74 ( 


77) 


75 


76). 


76 ( 


75). 


77 


74) 


78 ( 


81). 


79 ( 


80). 


80 ( 


79) 


81 ( 


78). 


82 ( 


83). 


83 ( 


82) 


84 ( 


91). 



Postnotum (the tongue-like process behind the scutellum with a 

group of scales 14. A. (Ochlerotatus) lepidonotus Edw. 

Postnotum without a group of scales 

15. A. (Ochlerotatus) subdiversus Mart. 

Abdominal tergites usually with basal stripes of light scales (more 
or less numerous light scales also in posterior half of tergites). 
Postcoxal spot of scales absent. Thorax with brown integument. 
Light scales mixed with dark scales in posterior half of tergites 

30. A. (Ochlerotatus) detritus Hal. 

Postcoxal spot of scales present. Thorax with blackish brown 
integument. Posterior half of tergites with dark scales and more 
or less numerous light scales (sometimes only a few light scales) 

31. A. (Ochlerotatus) simanini Guts. 

Posterior part of tergites with dark scales; light scales forming 

stripes or spots at base or sides of tergites. 

Mesonotum with long, dense, black setae. Setae present on entire 

posterior half of proepimeron. 

Spot of light scales on sternopleuron extending to anterior angle 

of sclerite (at base of fore legs). Claws slightly curved, long . . . 

26. A. (Ochlerotatus) nigripes Zett. 

Spot of light scales on sternopleuron not extending to the anterior 

angle. Claws sharply curved 

27. A. (Ochlerotatus) impiger Walk. 

Setae of mesonotum less long or dense, often brown or golden. 
Setae on proepimeron present only at the posterior margin. 
Fore femora variegated anteriorly, with numerous light and dark 
scales. Light scales mixed with dark scales in different parts of 
the wing, especially on the costa and ri. 

Proboscis with more or less numerous light scales, especially in 
the middle. Numerous light scales scattered on the veins of the 

wing 29. A. (Ochlerotatus) leucomelas Mg. 

Proboscis with uniformly dark scales. A few light scales only in 

anterior part of wing .... 28. A. (Ochlerotatus) cataphylla Dyar. 

Fore femora usually not variegated anteriorly, mainly with dark 

scales or with a few light scales. Light scales present only at the 

base of the wing or absent. 

A small spot of light scales below the anterior spiracle (hypo- 

stigmal spot). 

Spot of white scales on mesoepimeron extending to its lower 

margin. Lower mesepimeral setae present 

25. A. (Ochlerotatus) pullatus Coq. 

Spot of white scales on mesepimeron not extending to its lower 

margin. Lower mesepimeral setae usually absent 

24. A. (Ochlerotatus) intrudens Dyar. 

Hypostigmal spot absent. 

Spot of white scales on sternopleuron not extending to its anterior 

angle 23. A. (Ochlerotatus) diantaeus H. D. K. 

Spot of white scales on sternopleuron extending to its anterior 
angle (at base of fore legs). 

Spot of white scales on mesepimeron extending to its lower margin. 
Lower mesepimeral setae present. 



184 



85 (86) Postcoxal scales absent (i. e. scales absent on membranous part 

between sternopleuron and articulation of prothorax with coxae of 
fore legs 16. A. (Ochlerotatus) communis Deg. 

86 (85). Postcoxal scales present. 

87(88). Base of costa with dark scales. Light stripes of abdominal ter- 
gites 2-5 sharply narrowed in the middle 

6 18. A. (Ochlerotatus) punctor Kirby. 

88 (87) More or less numerous light scales usually at base of costa. 

Light stripes of abdominal tergites 2-5 of uniform width or only 
slightly narrower in the middle, (it is difficult to distinguish the 
adults of the last two species, but they can be reliably distinguished 
by the larvae.) . 

89 (90). Mesonotum with light, grayish yellow scales, with 2 indistinct 

longitudinal brown stripes close together 

17. A. (Ochlerotatus) pionips Dyar. 

90 (89). Mesonotum with dark scales, often more or less uniformly rust 

brown 19. A. (Ochlerotatus) hexodontus Dyar. 

91 (84). Spot of white scales on mesepimeron ending distinctly before its 

lower margin. Lower mesepimeral setae absent. 

92 (95). Hind tibiae usually covered mainly with light scales on the outside. 
93(94) Wing veins with dark scales. Light stripes on abdomen markedly 

narrower in the middle. First segment of flagellum of antennae 

yellow at the base 20. A. (Ochlerotatus) sticticus Mg. 

94(93) Light scales present at base of wing. Light stripes on abdomen 
of more or less uniform width. First segment of flagellum of 

antennae entirely black 21. A. (Ochlerotatus) nigrinus Eck. 

1 95 (92). Hind tibiae with dark scales on the outside ....... . . 

v ; 22. A. (Ochlerotatus) hungancus Mih. 

96 (51). Proboscis "not longer than fore femora or slightly longer than fore 

femora (A . aureus). 

97 (98). Mesonotum with golden scales and a longitudinal dark stripe. 

Wings with a large admixture of light scales. Bright golden 

scales form a large spot on the occiput • ■ • 

51. A. (? subgenus) aureus Guts. 

98 (97). Mesonotum without dark stripe. Wings with dark scales. Occiput 

without spot of golden scales. 

99 (100). Abdominal tergites with white spots at the sides, without light 

V stripes • 50. A. (?Aedes) nobukonis Yam. 

100 (99). Abdominal tergites dark or dark with light stripes • 

v 49. A. (Aedes) cmereus Mg. 

a (d). Abdomen without light stripes dorsally. 

b (c). Scales of mesonotum dark, reddish brown, sometimes with 

a golden or bronze sheen. Head mainly with dark scales •-•••• 

... A. cinereus cinereus Mg. 



c '(b). Scales of mesonotum lighter, golden brown or yellowish. 
Head with light, whitish gray scales 



A. cinereus rossicus D.G. M. 



Abdomen with light stripes at base of tergites 

... A. cinereus esoensis Yam. 



185 



Males * 

Key to Subgenera 

1 (2). Style divided into two branches, a median and a lateral. Palps 

several times shorter than the proboscis, as in females 

subgenus Aedes Mg. (p. 189). 

2 (l). Style simple, not divided into two branches. Palps about as long 

as the proboscis, sometimes slightly longer or slightly shorter. 

3 (6). Claspettes present. 

4 (5). Coxite with more or less large basal and apical lobes or at least 

with one of them subgenus Ochlerotatus Arr. (p. 185). 

5 (4). Coxite without lobes (sometimes rudimentary, basal lobe in the 

form of a small tubercle) subgenus Finlaya Theob. (p. 188). 

6 (3). Claspettes absent. 

7 (8). Appendage of style situated some distance from the apex. Style 

broadly cylindrical, slightly wider at the apex. Basal lobe forming 
a flask-shaped formation with hairs at the end. The only species 
of the subgenus in the USSR— .... A. (Aedimorphus) vexans Mg. 

8 (7). Appendage of style situated at the apex or slightly subapical. 

Style of different form. Lobe situated in middle of coxite 

subgenus Stegomyia Theob. (p. 188). 



180 Key to Species 

Subgenus Ochlerotatus Arribalzaga 

1 (8). Basal lobe of coxite with one row of long, lanceolate scales. 

2 (3). Style with S-shaped appendage 12. A. (O.) rusticus Rossi. 

3 (2). Style with straight or nearly straight appendage. 

4 (5). Wing of claspette elongate, transversely striated 

13. A. (O.) refiki Med. 

5 (4). Wing of claspette shorter, not striated. 

6 (7). Only one lobe in basal part of coxite, bearing a group of lanceolate 

scales 14. A. (O.) lepidonotus Edw. 

7 (6). In addition to the lobe with long, lanceolate scales, there are small 

lobes with 1—3 long and several slightly shorter setae 

15. A. (O.) subdi versus Mart. 

8 (l). Lanceolate scales on basal lobe of coxite absent. 

9 (14). Coxite with 3 large spines on the inside, 2 spines usually situated 

on the basal lobe. 

10 (ll). Coxite with a dense tuft of hairs directed inward 

23. A. (O.) diantaeus H. D. K. 

11 (10). Coxite different. 

12 (13). Stem of claspette with a finger-shaped process near the middle. 

Coxite with a distally directed tuft of hairs at the apex 

24. A. (O.) intrudens Dyar. 

13 (12). Stem of claspette without a process. Coxite without a tuft of 

hairs „ 25. A. (O.) pullatus Coq. 

* The males of A . nobukonis and A .aureus are not known. 



186 



NOTE 

Footnote on page 187 should read: 

The hypopygium of A. hungaricus Mih(p. 256) is similar. This species differs 
from A.caspius in the absence of white rings on the tarsi, dark scales on the 
wings and other characters. 



14 o: 

15 (ie; 



Coxite with 1—2 spines on the inside (on the basal lobe) or without 

spines. 

Basal lobe of coxite with 2 spines .... 1. A. (O.) caspius Pall. * 

(Structure of the hypopygium very similar in both subspecies, 

intermediate forms are common.) 

a (b). Basal lobe slightly convex, spines situated close together; 

anterior spine strongly curved at the apex 

o A. caspius caspius Pall. 

b (a). Basal lobe markedly convex, spines widely separated; 
anterior spine slightly curved ........ A. caspius dorsalis Mg. 

Basal lobe with one spine or without a spine. 

Apical lobe of coxite weakly developed, sometimes indistinct. 

Basal lobe very weakly developed, with short, thin setae and hairs, but 

without spines or large setae 2. A. (O.) mariae Serg. 

Basal lobe more or less developed, with long setae, usually one of 
them distinctly thicker. 

Apical lobe absent, Basal lobe slightly convex 

3. A. (O.) pulchritarsis Rond. 

Apical lobe present, small. Basal lobe convex, conical. 

Coxite with long hairs. Basal lobe without distinct spine. Phallo- 

some strongly sclerotized, bifurcate. Appendages of tergite 9 

with 14-20 setae 26. A. (O.) nigripes Zett. 

Coxite with shorter hairs. Basal lobe with a spine, which is more 
or less distinct among the setae. Phallosome weakly sclerotized, 

with 2 denticles. Appendages of tergite 9 with 4—12 setae 

27. A. (O.) impiger Walk. 

Apical lobe of coxite well developed. 

Basal lobe of coxite with one spine or with a large seta, which is 

distinct between the thinner setae and hairs. 

Wing of claspette narrow, without transparent, plate-shaped 

widening. 

Stem of claspette short; wing of claspette usually sclerotized, 

slightly widened in the middle 18. A. (O.) punctor Kirby. 

19. A. (O.) hexodontus Dyar. 
Stem of claspette long; wing of claspette very narrow, slightly 

sclerotized 16. A. (O.) communis Deg. 

17. A. (O.) pionips Dyar. 
Wing of claspette with a transparent, plate- shaped widening, which 
is sometimes distinct only if the position of the claspette is 
changed. 

Coxite with very long hairs directed medially; ends of hairs of 
both coxites overlapping. Stem of claspette sharply curved. 

Lobes of tergite 9 usually with 6—10 short, straight setae 

28. A. (O.) cataphylla Dyar. 

Lobes of tergite 9 usually with 10—15 long setae directed slightly 
outward 29. A. (O.) leucomelas Mg. 



The hypopygium of A .hungaricus Mih (p. 256) is similar. This species differs from A caspius in 
the white rings on the tarsi, dark scales on the wings and other characters. 



16 


(15) 


17 


(24) 


18 


(19) 


19 


(18) 


20 


(21) 


21 


(20) 


22 


(23) 


23 


(22) 


24 


(17) 


25 


(48) 


26 


(29) 


27 


(28) 


28 


(27) 


29 


(26) 


30 


(33) 


31 


(32) 


32 


(31) 



187 



182 



33 


(30) 


34 


(37) 


35 


(36) 


36 


(35) 


37 


(34) 


38 


(39) 


39 


(38) 


40 


(41) 


41 


(40) 


42 


(45) 


43 


(44) 


44 


(43) 



45 (42). 

46 (47). 

47 (46). 



48 (25). 

49 (50). 

50 (49). 

51 (52). 



52 (51). 



Hairs on inside of coxites usually shorter, their ends usually not 

overlapping (except in A. detritus). Stem of claspette straight 

or slightly curved. 

Apical lobe with broad base (extending proximally to middle of 

coxite), with short hairs. Wing of claspette short. 

Distal part of basal lobe tapering, thin . . . 20. A. (O.) sticticus Mg. 

Distal part of basal lobe broader, rounded 

21. A. (O.) nigrinus Eck. 

Apical lobe reaching to distal third of coxite, with long hairs. 

Wing of claspette usually long. 

Basal lobe narrow, markedly higher than wide at the base. Wing 

of claspette very broad, not wider than long 

4. A. (O.) cantans Mg. 

Basal lobe not higher than wide at the base. Wing of claspette 

narrower, longer than wide. 

Basal lobe flattened, with short, dense hairs and a thick, markedly 

sclerotized spine 10. A. (O.) flavescens Mull. 

Basal lobe conical, with a spine or a large, moderately thick seta 
and long hairs. 

Spine of basal lobe well developed, distinct between the setae and 
hairs. (Tarsi with light rings.) 

Stem of claspette long, curved 11. A. (O.) cyprius Ludl. 

Stem of claspette short, straight, but if curved, the transparent widen- 
ing of the wing begins at a distance from the base of the wing 

("manubrium" present) 5. A. (O.) riparius D. K. 

a (b). Manubrium of wing of claspette short or absent, a trans- 
parent widening in the basal and distal part of the wing 

A. riparius riparius D. K. 

b (a). Manubrium of wing of claspette about half as long as the 
wing; a transparent widening present only in apical half of wing 

of claspette A. riparius ater Guts. 

Spine (large seta) of basal lobe not distinct between the smaller 
setae. (Tarsi without light rings.) 

Inner surface of coxite with thick, long hairs. Transparent widen- 
ing of wing of claspette present in middle and at apex of wing. 

Phallosome ovoid, without a constriction 

30. A. (O.) detritus Hal. 

Inner surface of coxite with sparse, shorter hairs. Transparent 
widening of wing of claspette present on a short part of its middle 

third. Phallosome oblong, with a slight constriction 

31. A. (O.) simanini Guts. 

Basal lobe without a spine (large seta). 

Basal lobe conical, about as high as wide at the base 

6. A. (O.) behningi Mart. 

Basal lobe flattened, markedly wider than high. 

Basal lobe slightly convex, with long, very dense hairs. Wing of 

claspette with a small, hook- shaped appendage on the concave side 

at the base 32. A. (O.) kasachstanicus Guts. 

Basal lobe indistinct, flattened, with short hairs. Wing of claspette 
without hook-shaked appendage at the base. 



188 



53 (54). Stem of claspette thin, long, distinctly tapering apically 

7. A. (O.) excrucians Walk. 

54 (53). Stem of claspette thicker, not tapering apically 

9. A. (O.) annulipes Mg. 



Subgenus Finlaya Theobald 

1 (2). Coxite with a tuft of very large scales in distal part of inner 

surface 36. A. (F.) nipponicus La Casse and Yamag. 

2 (l). Coxite with small scales mainly on the outside. 

3 (6). Wing of claspette longer than stem. 

4 (5). Wing of claspette much longer than stem. Appendage of style 

short, l fa- l fa as lon g as the st Y le 39. A. (F.) togoi Theob. 

5 (4). Wing of claspette slightly longer than stem. Appendage of style 

much longer, about fa as long as the style 

37. A. (F.) seoulensis Yam. 

Wing of claspette not longer than stem. 

Coxite with a small tubercle with 2 strong setae and a thinner 

seta on the inner surface at the base , 

38. A. (F.) alektorovi Stack. 

Tubercle with strong setae on inner surface of coxite absent. 

Lobes of tergite 9 with 2—6 strong setae. 

Coxite with very long, dense hairs .... 35. A. (F.) echinus Edw. 

Hairs of coxite not long or dense ..... 34. A. (F.) geniculatus Ol. 

Lobes of tergite 9 with 5—10 thin hairs. 

Lobes of tergite 9 convex, hemispherical. Claw of sternite 10 

simple at the apex 41. A. (F.) koreicus Edw. 

14 (13). Lobes of tergite 9 flattened. Claw of sternite 10 bifurcate at the 

apex 40. A. (F.) japonicus Theob. 



Subgenus Stegomyia Theobald 

1 (2). Style flask- shaped, markedly widened at the apex 

44. A. (S.) vittatus Big. 

2 (l). Style cylindrical or spindle-shaped. 

3 (6). Appendage situated at apex of style. Posterior margin of tergite 9 

with an incision. 

4 (5). Lobe of coxite rounded, distinctly outlined. Posterior margin of 

tergite 9 with a shallow, curved incision 

48. A. (S.) chemulpoensis Yam. 

5 (4). Lobe of coxite not distinctly outlined. Posterior margin of 

tergite 9 with a deep incision 42. A. (S.) aegypti L. 

6 (3). Appendage of style situated a small distance from the apex. 

Posterior margin of tergite 9 convex or with a process. 

7 (8). Lobe of coxite transversely elongate . . . 45. A. (S.) galloisi Yam. 

8 (7). Lobe of coxite longitudinally elongate. 



189 



6 


(3) 


7 


(8) 


8 


(?) 


9 


(12) 


10 


(11) 


11 


(10) 


12 


0). 


13 


(14) 



9 (10). Posterior margin of tergite 9 serrated, curved (Far East) 

47. A. (S.) fiavopictus Yam. 

(Mediterranean) ... 43. A. (S.) cretinus Edw. 
10 (9). Posterior margin of tergite 9 smooth, with a process in the middle 
46. A. (S.) albopictus Sk. 



Subgenus Aedes Meigen 

Subspecies of A. (Aedes) cinereus Meigen 

1 (2). Lateral (longer) branch of style bifurcate at the apex 

A. cinereus cinereus Mg. 

2 (l). Lateral branch of style simple or serrated but not bifurcate. 

3 (4). Lobe of coxite with double apex .... A. cinereus rossicus D.G.M. 

4 (3). Lobe of coxite simple, stepped A. cinereus esoensis Yam. 



184 Fourth- stage larvae* 

1 (2). Antenna distinctly longer than the head 

A. (Ochlerotatus) diantaeus H. D. K. 

2 (l). Antenna shorter than the head. 

3 (10). Frontal hairs situated in one curved row. 

4 (5). Frontal hairs not displaced toward anterior margin of head, 

situated in middle of frontoclypeus. Comb on 8th abdominal seg- 
ment with a few scales (to 20) with a pointed main spine. Distal 
denticles of pecten more widely separated, reaching beyond middle 
of the siphon; hair tuft with 3—5 short branches situated near the 
apex Subgenus Aedes Meig. 

a. Typical larvae: with the characters of the subgenus 

A. (Aedes) cinereus cinereus Meig. 

b. Typical larvae: in addition to the subapical setae, there are 2 
pairs of thin, short hairs with 2—5 branches on the anterior side 

of the siphon A. (Aedes) cinereus rossicus D.G.M. 

c. Typical larvae with 4 pairs of additional, very short and thin 

hairs at lateral surfaces at apex of siphon 

A. (Aedes) cinereus esoensis Yam. 

5 (4). Frontal hairs displaced toward anterior margin of frontoclypeus. 

Comb on 8th abdominal segment with 40-50 scales without a main 
spine. 

6 (7). Siphon short (index about 2), as long as the last segment; hair tuft 

situated near apex of siphon. Gills short, bluntly conical 

A. (Finlaya) togoi Theob. 

7 (6). Siphon longer (index about 3), distinctly longer than the last seg- 

ment; hair tuft situated near the middle. Gills as long as the 
last abdominal segment or longer, lanceolate. 

* The larvae of A.simanini and A.kasachstanicus are described on pp. 279 and 283, those of 
A. aureus and A.nobukonis are unknown. 



190 



8 (9). Denticles of pecten evenly spaced, not more than 2 distal denticles 
extending beyond the hair tuft A. (Finlaya) koreicus Edw. 

9 (8). Distal denticles of pecten (1—4) more widely spaced, forming 
large spines' with a more acute angle to the longitudinal axis of 
the siphon A. (Finlaya) japonicus Theob. 

10 (3). Frontal hairs forming a triangle on each side, median hairs 
situated anterior to the inner hairs. 

11 (12). Median and inner frontal hairs displaced toward anterior margin 
of head; postclypeal hairs as long as the frontal hairs, situated 
between the median hairs; all hairs strongly branched. Comb 
with 8—15 scales arranged in one row, with a pointed main spine 
A. (Finlaya) nipponicus La Casse and Yamag. 

[ll). Median and inner frontal hairs not displaced toward anterior 

margin of head; postclypeal hairs much shorter than frontal hairs, 
if not shorter, frontal hairs weakly branched. 

[26). Auricles absent at base of siphon. Antennae with a small, simple 
hair, short, without spines or, rarely (A. vittatus) spines very 
weakly developed subgenus Stegomyia Thecb. 

[19). Outer frontal hairs simple. 

16). Hairs of body, including lateral hair of last segment, stellate. 

Gills sausage- shaped, 5 times as long as the saddle 

A. (Stegomyia) chemulpoensis Yam. 

.15). Hairs of body branched in one plane. Gills at most 3 times as 
long as the saddle. 

[18). Scales of comb, in addition to the main spine, with 2—6 or more 
spines at the sides of the base, 1—2 spines large, sometimes as 
large as the main spine. Additional hairs at sides of siphon 
absent A. (Stegomyia) aegypti L. 

[17). Scales of comb without large spines at sides of base of main 

spine, with only a row of thin, short setae. One hair at sides of 

siphon, in addition to the tuft, which is as long as the tuft 

A. (Stegomyia) cretinus Edw. 

,14). Outer frontal hairs with at least 2—3 branches. 

[21). Pecten about fa as long as the siphon. In additionto the pectenthere 
is a large, smooth spine near the apex. The hair on the siphon is 
situated between the spine and the distal denticle of the pecten . . 
A. (Stegomyia) vittatus Big. 

[20). Pecten not more than half as long as the siphon, without a larger 
spine near the apex. 
-3). Gills 1.2—1.6 times as long as the siphon. Postclypeal hairs only 
slightly shorter than the frontal hairs, strongly branched, with 
9—10 branches. Lateral hair on last segment with 2—4 short 
branches A. (Stegomyia) galloisi Yam. 

[22). Gills shorter chan siphon. Postclypeal hairs distinctly shorter 

than the frontal hairs, weakly branched, with 2—5 branches. Lateral 
hair with 2 longer branches of different length. 

[25). Hairs of body stellate .... A. (Stegomyia) flavopictus Yam. 

i24). Hairs of body branched in one plane 

A. (Stegomyia) albopictus Skuse. 



191 



26 


13) 


27 


28) 


28 


27) 


29 


28) 


30 


,35) 


31 


32) 


32 


31) 


33 


34) 


34 


33) 


35 


30) 


36 


37) 


37 


36) 


38 


29) 


39 


46) 


40 


43) 


41 


'42) 


42 


,41) 


43 


[40) 



Auricles well developed at base of siphon. Antennae different, 
usually with distinct spines, rarely smooth, with a simple or, 
usually, a branched hair* 

Median frontal hairs very long, longer than the head, twice as long 
as the branched inner hairs. Gills long, pointed at the end, the 

shorter lower pair as long as the siphon 

• A. (Finlaya) alektorovi Stack. 

Median frontal hairs about as long as the inner hairs. Gills of 

varying length. 

Antennae smooth, without spines. Gills longer than the saddle. 

Hair tuft on antenna in the form of a simple hair. Denticles of 

pecten in form of long, pointed, equally spaced spines. 

Comb with 55—60 scales, forming an irregularly triangular spot. 

All frontal hairs strongly branched, with 5 branches. Gills 

sausage- shaped, upper pair longer than the siphon, lower pair 

shorter A. (Finlaya) seoulensis Yam. 

Comb with 11—18 scales in one, curved row. Frontal hairs weakly 
branched, with 1 — 4 branches. Upper pair of gills not as long as 
siphon, lower pair distinctly shorter. 
Pecten /$— /$ as long as the siphon. Hairs of body strongly 

developed, stellate, with a few, thinner branches 

A. (Finlaya) geniculatus Ol. 

Pecten at least half as long as the siphon. Hairs of body strongly 

developed, stellate, with more numerous thicker branches 

A. (Finlaya) echinus Edw. 

Hair tuft of antenna with 2—4 short branches. Denticles of pecten 
short, not spinelike, with a broad base. 

Siphon 4—5 times longer than wide at the base, base 1.4—1.5 times 
wider than apex, slightly but uniformly tapering toward the apex. 
Gills sausage- shaped, with rounded ends; usually as long as the 

siphon A. (Ochlerotatus) pulchritarsis pulchritarsis Rond. 

Siphon 3.0—3.5 times longer than wide at the base, base 1.5 times 
wider than apex, strongly tapering toward the apex. Gills sausage- 
shaped, usually twice as long as the siphon 

A. (Ochlerotatus) pulchritarsis asiaticus Edw. 

Antennae with sparse spines. If the spines are weakly developed 

(A. m a r i a e), thegills are very short and spherical. 

On the anterior surface of the siphon, in addition to the subapical 

setae, are 2—3 pairs of coarse, well developed hairs and on the 

lateral surface, in addition to the tuft, is a pair of thin lateral hairs 

near the distal denticles of the pecten. 

Denticles of pecten extending beyond hair tuft of siphon. 

Hair tuft of siphon with 6—8 branches. Distal denticle of pecten 

not extending to the apical third of the siphon 

A. (Ochlerotatus) rusticus Rossi. 

Hair tuft of siphon in the form of a long, simple hair. Distal 
denticle of pecten situated in apical third of siphon, nearly at its 

end A. (Ochlerotatus) subdiversus Mart. 

Distal denticles of pecten not extending beyond the hair tuft of the 
siphon. 



192 



Three pairs of hairs on anterior surface of siphon. The ventral 

of the hairs behind the comb is the longest 

A. (Ochlerotatus) lepidonotus Edw. 

Hairs on anterior surface of siphon, except the subapical setae, 
absent, only the tuft present on the sides. 

Distal denticles of pecten widely spaced, extending beyond hair 
tuft of siphon. 

Saddle surrounding last abdominal segment like a ring 

A. (Ochlerotatus) nigripes Zett. 

Saddle of last abdominal segment extending only beyond middle of 

sides A. (Ochlerotatus) cataphylla Dyar. 

Distal denticles of pecten not extending beyond hair tuft. 

Four to six shorter tufts before the common base of the fin. Hair 

tuft in middle of siphon (except in A . m a r i a e, which has a very 

short siphon), at least as long as the width of the siphon where 

the tuft is situated. 

Whole surface of body with small dark spines in dense rows. 

Head and long siphon (index 4—5) ocher yellow; pecten with 2—3 

larger, widely separated denticles 

A. (Ochlerotatus) cyprius Ludl. 

Surface of body bare, without small spines. 
Comb on 8th abdominal segment usually with 6—12 scales. 
Scales of comb arranged in one irregular, often curved row. 
Median and inner frontal hairs with 2—3 branches. Index of siphon 

3.5-4.0 A. (Ochlerotatus) riparius Dyar and Knab. 

Scales of comb arranged in 2 (rarely 3) irregular rows. Median 

and inner frontal hairs simple. Index of siphon 2.0 

■ A. (Ochlerotatus) nigrinus Eck. 

Comb on 8th abdominal segment with 15—45 scales. 

Saddle of last abdominal segment weakly developed, not completely 

covering it dorsally and not extending beyond middle of sides. 

Gills short, spherical. Index of siphon at most 2 

A. (Ochlerotatus) mariae Ed. and Et. Serg. 

Saddle well developed. Gills at least half as long as the saddle. 

Index of siphon at least 3. 

Hairs, at apex of posterior valves of stigmal plate hook- shaped 

and thickened. 

Comb with 30-40 (usually 32-36) scales. Postclypeal hairs with 

2—3 thin, short branches. Hairs at apex of posterior valves of 

stigmal plate strongly thickened 

A. (Ochlerotatus) excrucians Walk. 

Comb with 20-28 (average 24) scales. Postclypeal hairs with 
6—8 thin, short branches. Hairs at apex of posterior valves of 

stigmal plate less thick A. (Ochlerotatus) behningi Mart. 

Hairs at apex of posterior valves of stigmal plate of the usual 

form: slightly curved, not thickened. 

Distal denticles of pecten widely spaced. 

The distal 1 — 3 denticles of the pecten situated beyond the middle 

of the siphon. Hair tuft with 2-4 branches 

A. (Ochlerotatus) beklemishevi Den. 



193 



66 (65). Distal denticles of pecten distinctly not reaching middle of siphon. 

Hair tuft with 4—7 (usually 5—6) branches 

A. (Ochlerotatus) flavescens Mull. (part). 

67 (64). Distal denticles of pecten not widely spaced, space between the last 

2 denticles usually as long as the space between the preceding 2 
denticles. 

68 (69). Six or seven shorter tufts before the fin, not situated on the com- 

mon base A. (Ochlerotatus) flavescens Mull. (part). 

69 (68). Four or five shorter tufts before the fin,not situated on the com- 

mon base. 

70 (71). Fin with 18—19 tufts situated on the common base. Posterior 

appendage of stigmal plate, in addition to the usual 2 branches, with 

a different median branch between them 

A. (Ochlerotatus) cantans Meig. 

71 (70). Fin with at most 16 tufts situated on the common base. Posterior 

appendage of stigmal plate of the form normal for the subgenus . . 
A. (Ochlerotatus) annulipes Meig. 

72 (51). At most 3 tufts not situated on the common base before the fin. 

If there are 4 tufts before the fin (A . v e x a n s ), the hair tuft on the 
siphon is short, shorter than the width of the siphon at its position, 
situated at the beginning of the apical third. 

73 (78). Gills usually shorter than saddle, rarely only l /z of its length, not 

pigmented. 

74 (75). All scales of the comb without a long main spine, with a row of 

spines at the margin, the apical spine the longest. "Stirrup" of 
stigmal plate without processes to the spiracles, markedly dis- 
placed posteriorly to between the posterior valves. Median frontal 
hairs with 1—2 branches, inner hairs with 2—3 branches. Gills 
spherical A. (Ochlerotatus) detritus Hal. 

75 (74). At least some scales of the comb with a long main spine. "Stirrup" 

of stigmal plate with processes to the spiracles, situated in the 
middle. Median and inner frontal hairs usually simple. Gills 
oblong, at least l /z as long as the saddle. 

76 (77). Tufts of fin branched far from the base: length of single stem of 

median tufts of fin 1.5—2 times as long as the lateral processes 
of the base. Some scales of the comb near the ventral side have 
a long, main spine, the others have 2—3 longer spines at the apex 

and 2—3 shorter spines at the sides of the base 

A. (Ochlerotatus) leucomelas Meig. 

77 (76). Tufts of fin branched from near the base: single stem not longer than 

the lateral processes of the base. Scales of comb with a more or 

less distinct main spine, isolated in any part of the comb 

A. (Ochlerotatus) caspius Pall. 

a. Hair tuft on siphon situated beyond the middle, nearer to the 
apex, with 5—10 branches. Inner caudal hairs with 12—15 branches. 

Median hair behind the comb with 7—14 branches 

A. (Ochlerotatus) caspius caspius Pall. 

b. Hair tuft on siphon situated near the middle, with 3—5 (rarely 
more) branches. Inner caudal hairs with 4—12 branches. Median 

hair behind the comb with 5—8 branches 

A. (Ochlerotatus) caspius dorsalis Meig. 



194 



94 



93 



Gills at most 1.5 times as long as the saddle, often pigmented. 
More than 40 scales in the comb; all or nearly all scales without 
a main spine, with a row of small spines at the margin. 
Median and inner frontal hairs simple, rarely one or two hairs 

with 2—3 branches A. (Ochlerotatus) communis De Geer. 

Median and inner hairs with 3—7 branches. 

Antennae thin, long, about % as long as the head or slightly longer. 

All scales of comb without a main spine 

A. (Ochlerotatus) pionips Dyar. 

Antennae thicker and shorter, about half as long as the head. 

Some scales of comb with a weak main spine 

A. (Ochlerotatus) pullatus Coq. 

Comb with less than 30 scales. 

Saddle ring-shaped, surrounding the last segment, or there is a 

narrow slit on the ventral side between the margins of the saddle. 

Comb with 10-20 scales, scales small, 0.07-0.08 mm 

A. (Ochlerotatus) punctor Kirby. 

Comb with 5—9 scales; scales larger, 0.11— 0.13mm 

A. (Ochlerotatus) hexodontus Dyar. 

Saddle not surrounding the last abdominal segment completely, 

only extending to the lower margin of the sides. 

Inner frontal hairs simple. Distal denticles of pecten not widely 

spaced. 

Median frontal hairs simple. Comb with 7—16 (average 13—14) 

scales. Gills pigmented A. (Ochlerotatus) impiger Walk. 

Median frontal hairs with 2 branches. Comb with 16—24 (usually 

20) scales. Gills not pigmented 

A. (Ochlerotatus) hungaricus Mih. 

Inner frontal hairs with 2—5 branches. 

Distal denticles of pecten (1—3) always widely spaced, distinctly 

larger than the others. 

Hair tuft situated beyond the middle of the siphon, always distal 

to the distal denticle of the pecten, with 3—8 (usually 4—6) thin 

branches, shorter than the width of the siphon at its position .... 

A. (Aedimorphus) vexans Meig. 

Hair tuft situated in the middle or slightly beyond the middle of 
the siphon, with 3—8 (usually 4—6) branches, as long as the width 
of the siphon at its position or longer. If there are more than 2 

widely spaced denticles, the tuft is situated between them 

A. (Ochlerotatus) intrudens Dyar. 

Denticles of pecten equally spaced; one of the distal denticles 
may be rarely slightly separated from the others. Comb with 
16-27 (usually 22-24) scales .... A. (Ochlerotatus) sticticus Meig. 



1. Subgenus Ochlerotatus L. Arribalzaga* 

Hypopygium: coxite with a basal and an apical lobe, or at least with 
one lobe; style simple, not bifurcate; appendage of style situated at the 

* See also p. 283. 



195 



190 



apex; claspettes present; phallosome simple (not divided into 2 plates). 
Palps of male slightly longer than or as long as the proboscis, rarely slightly 
shorter, usually slightly thickened on the 3rd and 4th or 3rd and 5th 
segments. Female: abdomen tapering posteriorly, cerci distinctly pro- 
jecting; proboscis distinctly longer than fore femora. 

The females are very difficult to determine. The specimens have to be 
well preserved and undamaged. Determination is sometimes possible only 
by the males or the larvae. An important systematic character is the 
arrangement of the spots of scales on the pleurae of the thorax (Peus, 1933). 
The form of the claws is important in some species (Vockeroth, 1950). 
Sazonova (1958) mentions many useful characters. She gave a key to the 
females of species of Aedes of the forest zone. 




FIGURE 103. Aedes caspius caspius Pall. 



196 






The subgenus contains about 140 species distributed mainly in the 
Palaearctic and Nearctic. Twenty-nine species occur in the USSR. The 
subgenus Ochlerotatus is characteristic for the forest zone and tundra. 
The distribution of some species extends to 75— 78°N. The species of 
Ochlerotatus are active bloodsuckers and have an important role in the 
spread of tularemia and virus infections. 

The Palaearctic species of Ochlerotatus may be divided into 4 
groups according to the adults, named after one of the series : caspius, 
cantans, rusticus, communis. The first two groups have light rings 
on the tarsi (Figure 97), the other two groups do not. 



caspius group 

Tarsi with white rings on the base and apex of the segments; the borders 
of the segments are visible on the white rings (Figure 103). The hypo- 
pygium is characterized by the weak development or absence of an apical 
lobe on the coxite, which, however, is also the case in some species of the 
other groups. . 

Three Palaearctic species belong to the caspius group. 



1. Aedes (Ochlerotatus) caspius Pallas, 1771 

This species is very variable in coloration and is represented in the 
Palaearctic by two subspecies, one of which, A. c. caspius, occurs in 
more southern and dry regions, and the other, A. c. dorsalis, in more 
northern and humid regions. Intermediate forms occur in the Far East, 
Siberia, Kazakhstan and in some parts of the European USSR. 

Head with white and yellowish brown scales dorsally. Proboscis and 
palps with brown and white scales in varying numbers. The coloration 
of the thorax and wings of the two subspecies differs. Tarsi with white or 
cream-colored rings at the base and apex of the segments; light rings 
sometimes indistinct. Abdomen with indistinct light transverse stripes and 
a light longitudinal stripe which may not be present on all segments. Light 
scales usually predominate on the abdomen. There are sometimes only 
dark spots at the sides of the tergites; the abdomen is rarely completely 
covered with light scales. 

Hypopygium (Figure 104): basal lobe of coxite with 2 spines, one of them 
longer and curved, the other shorter and straight. Wings of claspette 
narrow, without transparent widening. 

The larvae of the two subspecies differ only in quantitative characters: 
size of body, siphon, stigmal plate, gills, degree of branching of hairs, etc. 
The quantitative importance of these differences even in distinctly sepa- 
rated populations of the subspecies, is contiguous or transitional in their 
extreme variations. This becomes clear in the descriptions below. 

Distribution. From the Atlantic to the Pacific and from 64°N 
(Finland) and Yakutsk to the southern boundary of the Palaearctic (North 
Africa, Iran), widely distributed in North America (only A. c. dorsalis). 



197 



192 Biology. Their most characteristic habitat are river valleys, where 

they often predominate among mosquitoes attacking man and animals. In 
mass flights, they enter inhabited areas, houses and cattle sheds. They 
breed in large numbers in floodplains after the floods. A new generation 
appears after the summer rain which is smaller than the spring generation. 



(191) 




FIGURE 104. Hypopygium of A e d e s caspius dorsalisMg. 



Note on systematic s. Many authors consider A. c. caspius 
Pall, and A. c. dorsalis Mg. as different species. Their distribution 
overlaps in a large part. In Western and Central Europe the two forms are 
easily distinguished by the coloration of the adults, less by the structure of 
the hypopygium. Specimens with intermediate characters are rarely found. 
Further north, specimens become more common the subspecies of which is 
difficult or impossible to determine. Intermediate forms are also found in 
some parts of the European USSR (Volga area, Northern Caucasus), 
Kazakhstan and Siberia. 

The characters of the Far Eastern A. caspius are mixed and inter- 
mediate. In Transbaikalia, the Maritime Territory and Northeast China 
specimens predominate, in which the mesonotum has two longitudinal, broad 
stripes of white or cream- colored scales with a narrow, dark, chestnut- 
colored or brown stripe between them. The lateral margins of the meso- 
notum are also covered with chestnut- colored scales. Deviations from 
this intermediate variation between A. c. caspius and A. c. dorsalis 
have been observed, e. g. instead of longitudinal, light stripes there may be 
light spots of irregular form, cream- colored scales may predominate and 
the dark scales form only a longitudinal stripe in the anterior part of the 
mesonotum. Specimens from the Maritime Territory are also intermediate 



198 



in the structure of the hypopygium. They resemble A. c.dorsalis in 
the coloration of the wings. Southern populations (Kwangtung) resemble 
A. c. caspius more closely. 

A.c. caspius occurs in Middle Asia but there are also intermediate 
forms. A. c. dorsalis inhabits mainly high mountain regions (Pamirs). 

Although A.c. caspius and A. c. dorsalis (at least in the eastern 
part of their distribution) are connected by transitional characters, it seems 
justified to consider them as subspecies. 

A detailed study of the geographical variation of A. caspius may make 
it possible to establish further subspecies, but some variations in coloration 
and other characters definitely depend on ecological factors, mainly on the 
conditions of development of the larvae. A description of new subspecies 
without a study of their ecology is therefore not justified. 

Aedes duplex Mart, was described from two males from the 
Saratov Region (Martini, 1926, 1928). It resembles A. caspius but differs 
in the light color of the scales on the mesonotum, the weakly developed 
light rings on the tarsi, and the double number of spines on the basal lobe 
of the coxite. On each lobe are two straight spines situated close together 
and two curved spines, but similar characters of coloration are also often 
observed in A. caspius in different parts of its distribution. A double 
number of spines on the coxite is sometimes considered as an aberration. 
In the collections of the Zoological Institute there are males of A. caspius 
from Sarepta in the Volgograd Region with one, two or several additional 
spines in addition to the two spines on the basal lobe which are of nearly 
the same size and structure as the "normal" spines. There are transitions 
between the normal and aberrant specimens. 

The status of A. duplex is thus doubtful in our opinion. 



Aedes caspius caspius Pallas 

In addition to the characters given in the key, we give the following. 
Proboscis and palps of male mainly with light scales. In the basal quarter 
of the costa there are dark scales among the light scales or the dark scales 
predominate. The dark spots on the abdominal tergites are brown, the 
light spots are yellowish or white in some places. The proximal spine of 
the basal lobe is sharply curved, with thin, hooked end. 

The coloration of the mesonotum is very variable. The light longitudinal 
stripes may be narrow, distinct (the most common coloration) or wider and 
diffuse. They are sometimes yellowish and indistinct against the background 
of brown scales. The mesonotum is often also covered with yellowish brown 
or sand-colored scales. The size of the dark spots at the sides of the 
abdominal tergites varies to the complete absence of dark spots. 
B.N.Kazantsev (1931) found that the light, "sand-colored" A. caspius 
developed in Bukhara in water with a high salinity, while mosquitoes breed- 
ing in fresh water had a contrasting color. These color variants are 
characteristic not for the area but are found in different parts of the dis- 
tribution of the subspecies. 

Fourth-stage larva (Figure 105) medium-sized, yellowish, with 
lighter head and siphon. Frontal hairs slightly secondarily feathered, 



199 



outer hairs with 7—10 branches, median hairs situated before the inner 
hairs, both pairs simple, rarely 2-branched, very rarely one of the hairs 
194 3- branched; postclypeal hairs with 3—5 short, thin branches. Antenna about 
half as long as the head, with sparse, small spines; hair tuft situated basal 
to the middle, with about 9 branches which are half as long as the antenna. 



(193) 




FIGURE 105. Aedes caspius caspius Pall. Fourth-stage larva. Posterior end, 
lateral. 



Comb with 18—28 (usually 20-25) scales in 2-3 irregular rows. The 
scales vary markedly, from short scales without a main spine, with a row 
of small spines at the apex and margin, and narrower at the base to longer 
scales with one (or more) distinct spine and shorter scales at the base. 
Hairs behind the comb: dorsal hair with 5—10 branches, ventral hair with 
6—7, median hair with 7—14 branches, intermediate hairs simple. Siphon 
straight, slightly tapering from the middle to the apex (index 1.8—2.6), of varying 
length. Pecten with 17—26 (usually 20—22) denticles, 1 — 4 denticles near 



200 



the base rudimentary; denticles equally spaced; pecten slightly extending 
beyond middle of siphon. Hair tuft with 5—10 secondarily feathered, short 
branches, situated beyond middle of siphon. Stigmal plate typical for larvae 
of Ochlerotatus: 0.30— 0.36 mm long, 0.2 7—0.32 mm wide at the posterior 
valves, 0.28— 0.34 mm wide between the ends of the lateral valves. 

Saddle extending to middle of sides of last segment; lateral hair simple, 
short, less than half as long as the saddle; outer caudal hairs simple, 
longer than the siphon, inner hair with 12—15 branches which are half as 
long. Fin with 14—17 tufts on the common base and 2-3 shorter tufts 
before it; 9—12 thin branches in each of the middle tufts. Gills short, 0.3— 
0.9 of length of saddle, leaf-shaped or lanceolate, lower pair shorter than 
the upper. 

Distribution. The subspecies is characteristic for the steppe; it 
occurs in dry steppes with saline waters. It is distributed in Europe to 
England, southwest Finland, and the central parts of the European USSR. It 
also occurs in the southern regions of Siberia, Kazakhstan and Middle Asia. 
It is distributed in Mongolia, North and West China, West Asia and North 
Africa. In the steppes of the Ukraine, the Lower Volga area, east of Cis- 
caucasia and Transcaucasia, Kazakhstan and Middle Asia, A. c. caspius 
is the predominant species which attacks people and animals in masses, e.g., 
in the Mugan steppe (Zdrodovskii, 1926)^ in many parts of Turkmenia 
(Petrishcheva, 1936) and in the lower reaches of the Syr Darya (Blago- 
veshchenskii, 1937). 

Biology. The larvae develop in open or shaded waters (south), per- 
manent or temporary water bodies formed by the snowmelt, flooded rivers 
or overflows of irrigation, with little vegetation and with muddy bottom, 
often with a high concentration of salt (to 5% and higher). The eggs are 
deposited in moist soil at the edge of water bodies which do not dry up. 
There are several generations per year and they are sometimes very large 
(Middle Asia). The females bite man and animals, especially at dusk. 

The species has a high resistance to heat and drought. In Tadzhikistan, 
activity is highest at temperatures of 19-31° (Bregetova, 1946). They often 
bite during the day. They may migrate for long distances, sometimes to 
10km. 

They have a role in the spread of tularaemia and may be infected for 
a month (Olsuf'ev, 1938). They transmit Tahyna virus in Czechoslovakia 
(Bardos and Danielova, 1959). 



Aedes caspius dorsalis Meigen, 1830 

The main characters are given in the key. Proboscis and palps of fe- 
male mainly with dark scales. Costa with white scales in the basal quarter, 
without dark scales. Dark spots on the abdominal tergites blackish brown, light 
spots with a whitish gray tone. Proximal spine of basal lobe of coxite 
slightly curved, its apex usually not hook-shaped. 

Fourth-stage larva (Figure 106) slightly larger than in A . c . 
caspius and darker, to brownish black. Frontal hairs secondarily 
feathered, outer hairs with 4—8 branches (usually 5—6), median hairs 
situated before the inner, both pairs simple, rarely 2-branched; postclypeal 



201 



hairs short, with 2—5 thin branches. Antenna about half as long as the head, 
with sparse spines and a tuft with 4—7 branches in the middle, not more 
than half as long as the antenna. 

Comb with 13—34 (usually 20—25) scales in 2—3 irregular rows; scales 
196 very variable, as in A . c . c a s piu s. Hairs behind the comb: dorsal hair 
with 3—8 (usually 4—5) branches, ventral hair with 4—6, median hair with 
5-8 branches, intermediate hair simple. Siphon with an index of 2.5-3.0, 
rarely less. Pecten with 14—23 denticles, the first 4 denticles at the base 
rudimentary; pecten not reaching middle of siphon; denticles irregularly 
spaced, the distal denticles rarely slightly outside the row. Hair tuft with 
3—8 (usually 4—5) secondarily feathered branches, situated in the middle of 
the siphon, branches not longer than the width of the siphon at the base. 
Stigmal plate (Figure 107) slightly larger and more darkly pigmented than 
in A. c. caspius. 



(195) 




FIGURE 106. Aedes caspius dorsalis Mg. Fourth-stage larva. Posterior end, lateral. 



Saddle as in A. c. caspius but more strongly pigmented and with 
sharply defined lower margin. Inner caudal hair with 4—12 (usually 8—9) 



202 



branches, more than half as long as the outer hair. Number of tufts of the 
fin as in A . c . caspius, but each tuft is less branched, usually with 7—9 
branches. Gills slightly longer than in A. c. c a s p iu s, about 1.3 times as 
long as the saddle in fresh water, not more than 0.3—0.4 of the length of the 
saddle in saline water. 




FIGURE 107. Aedes caspius dorsalis Mg. Fourth- 
stage larva. Stigmal plate. 



Distribution. Characteristic for open habitats in mixed forests and 
forest-steppe. In the north it extends to the northern boundary of the 
distribution of the species, in the south to Hungary, Bulgaria, steppes in the 
Ukraine, Northern Caucasus, Lower Volga area and mountains of Middle 
Asia, including the Pamirs. Also Siberia, Far East (see above) and North 
America. 

Biology. The larvae develop mainly in small, open water bodies and 
swamps. They are found in permanent or temporary water bodies formed 
by thawing snow, floods, rainfall or groundwater fresh or saline. The 
eggs hibernate. Several generations per year, especially during periodic 
floodings. They are very numerous in some localities. They are mass 
bloodsuckers in river valleys, e. g. the Volga and Dnieper. They bite at 
temperatures of 16—22° in the Moscow Region. The seasonal numerical 
curve has two peaks (Shlenova, 1959). There are 2—3 generations per year 
in the central European USSR (Khelevin, 1958). 

They transmit western encephalitis (equine encephalomyelitis) in the 
U.S.A. 



203 



2. Aedes (Ochlerotatus) mariae Ed. and Et. Sergent, 1902 

The species differs from A. caspius in the absence of a longitudinal 
light stripe on the abdomen and the absence of spines on the basal lobe of 
197 the coxite. The description is based mainly on that of Senevet and Andarelli 
(1964), who noted great variation in the characters of this species. 






FIGURE 108. Abdomen of Aedes mariae Serg., 
dorsal (variation of coloration) (after Senevet and 
Andarelli) 



198 



Proboscis with reddish brown scales, sometimes mixed with whitish 
scales in the middle of the proboscis. Mesonotum with rust brown scales 
usually with a few whitish scales, which sometimes form an indistinct, 
narrow longitudinal stripe ("var. zamittii" Theob.). Tarsi with white 
rings, each ring extending to two segments; last segment of hind tarsi 
completely white. Wings with dark and light scales, more numerous on the 
costa, subcosta and ri. Abdomen markedly varying in coloration (Figure 108] 
At the base of the tergites are narrow stripes of light scales which usually 
widen laterally into spots. A large part of each tergite is covered with 
dark scales; a longitudinal stripe of light scales is absent. 

Hypopygium (Figure 109): apical lobe of coxite indistinct. The moder- 
ately convex basal lobe bears hairs and 4—5 larger setae. Claspettes with 
short, straight stem and narrow, curved wing, without transparent widening. 
Phallosome more or less rectangular. Lobes of tergite 9 with long setae 
(compared with those of A. caspius), 4— 6 setae at each side. 

Fourth- stage larva (Figure 110) of varying coloration. Frontal 
hairs: outer hairs usually with 7 branches, both inner pairs simple, a third 
as long as the outer, the median hairs situated before the inner hairs; 
postclypeal hairs very thin and short, branched. Antennae thin, slightly 
curved, with weak spines; hair tuft situated in the middle, with 6—9 branches, 
half as long as the antenna. 

Comb with 16—25 scales in 2—3 rows; scales with distinct main spine 
with a varying number of denticles of different size at the base or with 



204 



1-2 denticles as large as the main spine. Siphon short (index 1.4-2.0), 
slightly tapering, with a varying degree of chitinization at the base; pecten 
reaching to middle of siphon, with 15 or more thin denticles which are 
longer distally and with accessory denticles in the basal third or half of 
the denticles; hair tuft with 6 branches, as long as the width of the siphon, 
situated slightly distal to the middle of the siphon. 



197' 





FIGURE 109. Hypopygium of Aedes mariae Serg. (after Sene vet 
and Andarelli): 

A — basal lobe; B — claspette; C — lobes of tergite 9. 




FIGURE 110. Aedes mariae Serg. Fourth-stage larva (after Edwards): 
1 — head, dorsal; 2 — posterior end, lateral. 



205 



Saddle on last segment weakly developed, not completely covering the 
dorsal side and only extending slightly to the sides; lateral hair simple. 
Outer caudal hairs simple, long, inner hairs fa — fa as long as the outer, 
with 11—14 branches. Fin strongly developed, with 12—13 tufts on the com- 
mon base and 4—5 tufts before it. Gills short, spherical. 

Distribution. Europe, Africa and Asia, mainly on the Mediterranean 
coast from Morocco to Syria. Not recorded with certainty from the USSR. 

Biology. The larvae are found in rock pools on the sea shore, often 
in the surf zone. The usual concentration of salt in such pools is 2—4%, but 
the larvae tolerate a much higher concentration, to 18%. 

The species transmits the parasite of bird malaria, Plasmodium 
relic turn. 

Note on systematics. The occurrenc eofA.mariae in the 
USSR is not certain. There is a record from the Ukraine (Prendel 1 , 1966), 
199 with the note that this needs confirmation. In the collections of the Zoo- 
logical Institute is a preparation (possibly of A. mariae) which was found 
near Dnepropetrovsk in May 1928 by V. V. Goritskaya. The hypopygium 
resembles that of A. caspius, with the following differences: basal lobe 
of coxite, in addition to numerous hairs, with an irregular row of 7—8 more 
or less equal setae; spines or large setae absent; the weakly developed 
apical lobe bears a few hairs which are longer than those in A. caspius. 

A . mariae possibly occurs in the south of the USSR but has not been 
differentiated from A. caspius. 



3. Aedes (Ochlerotatus) pulchritarsis Rondani, 1872 

The species is characterized by the absence of a light longitudinal stripe 
on the abdomen and the brown scales of the wings. It is divided into two 
subspecies. 

Head with yellowish and brown scales . Palps of female brown, with white 
apex; palps of male brown, with white rings on the last segments. The coloration 
of the mesonotum differs in the two subspecies. Scutellum with yellowish 
scales. Pleurae of thorax with spots of white scales. Tarsi with contrast- 
ing white rings on the apex of the preceding and the base of the next seg- 
200 ment. Last tarsal segment white. Wings with dark scales. Abdomen 
brown, with stripes of white scales at the base of the tergites. 

Hypopygium (Figure 111): an apical lobe on the coxite absent; the 
weakly convex basal lobe has a long spine curved at the end and several 
slightly thinner setae; wings of claspettes narrow, long; lobes of tergite 9 
slightly convex, with 4—6 short setae. 

The white scales on the tarsi and abdomen of A. pulchritarsis give 
it some resemblance to species of the subgenus Stegomyia, but 
A. pulchritarsis has no lyre- shaped pattern on the mesonotum, which 
is characteristic for A. (Stegomyia) aegypti. In species of Stego- 
myia, the white rings on the tarsi are situated at the base of the segments 
while in A. pulchritarsis each ring extends to the apex of one segment 
and the base of the next. 

Larvae semitransparent, milk white to grayish, with yellowish brown head 
and darker siphon. 



206 



(199) 







FIGURE 111. Hypopygium of A ed es p ul chri t arsi s Rond. 

The subspecies differ mainly in the coloration of the mesonotum and 
the length of the gills in the larvae. 

Distribution. Mediterranean, South Asia. USSR: southern coast of 
the Crimea and other parts of the Ukraine, to Uzhgorod and Kharkov in the 
north, the Caucasus and Middle Asia. 

Biology. It develops in tree holes. It occurs in humid forests and 
on irrigated farm lands. It is not an active bloodsucker. 



Aedes pulchritarsis pulchritarsis Rondani 

Mesonotum with golden scales and brown spots, the dark color some- 
times predominating. Femora and tibiae dark anteriorly, with small spots 
of dark scales or with scattered light scales, often also completely dark 
anteriorly, with white apex. Tarsi with white rings. 



207 



Fourth-stage larva with nearly rounded head and broad anterior 
margin of the frontoclypeus. Size varying according to the size of the tree 
hole. 

Frontal hairs situated near anterior margin of frontoclypeus, outer hairs 
with 6—13 (usually 10—12) branches, median hairs with 4—7 (usually 6—7), 
situated before the longer inner hairs, with 8—10 (rarely fewer) branches 
situated on one line with the outer hairs; postclypeal hairs with 11 — 18 
branches, situated between the median and inner frontal hairs. Antennae 
thin, slightly curved, without spines; hair tuft with 1 — 4 (usually 2—3) thin, 
short branches, situated in the middle. 

Comb on sides of 8th segment with 6—10 (usually 8) scales in one, some' 
times irregular row; scales with a pointed terminal spine, with short, thin 
spines at the base. Siphon thin, straight, index 4—5; pecten with 17—24 
closely spaced denticles; they are short, with a broad base and few spines; 
hair tuft situated in the middle, with 3—4 branches. Stigmal plate with a 
horseshoe-shaped posterior appendage to the "stirrup"; the formations on 
the lateral valves are more strongly connected by their base with the 
appendages of the "stirrup" than with the spiracles. 

Last segment with a weakly developed saddle, which often covers the 
201 segment only dorsally; lateral hair simple, long; outer caudal hairs simple, 
long; inner hairs with 3—5 branches, longer than the gills. Fin weakly 
developed, with 6—8 (rarely 9) slightly branched tufts (with 2— 4, usually 3 
branches), as long as the inner caudal hairs. Gills as long as the siphon, 
sausage-shaped, with rounded ends. 

Distribution. Western part of the distribution of the species, in the 
Ukraine and Caucasus in the USSR. 

Biology. The larvae develop in tree holes (maple, oak, elm, etc.), in 
tree stumps and among roots together with the larvae of A. geniculatus, 
A. plumbeus and O. pulchripalpis. They hibernate in the egg stage. 
The larvae hatch when the tree holes become filled with water . A few generations 
per year from March— April. The females are not active bloodsuckers. 

Note on systematics. A.pulchritarsis var . berlandi 
Se'guy, 1921 was first described as a variety, and later as a different species 
(Rioux and Arnold, 1955; Callot and Rioux, 1965). In A. b e r 1 a nd i, light 
spots on the femora and tibiae are absent or indistinct, the mesonotum is 
covered mainly with golden scales of different tones, sometimes with an 
indistinct longitudinal, white stripe; there are no differences in the structure 
of the hypopygium. A. berlandi is distributed in the Mediterranean, 
from Algeria to Iran, and often occurs together with "typical" A. pulchri- 
tarsis. We do not think that A. berlandi should be considered as a 
species as the differences are very variable. Specimens from the Crimea 
and Transcaucasia show variation in the coloration of the mesonotum and 
similar light spots on the femora and tibiae. 

Aedes pulchritarsis asiaticus Edwards, 1926 

(var. stegomyina Stackelberg and Montchadsky, 1926) 

The subspecies differs mainly in the coloration of the mesonotum. Head 
with white and black scales which form spots. Mesonotum with blackish 



208 




FIGURE 112. Aedes pulchrit arsis asiaticus 
Edw. Fourth -stage larva. Head, dorsal. 



brown or dark brown scales, with a 
longitudinal median stripe of white 
scales and white transverse stripes, 
stripes on the mesonotum some- 
times formed by yellowish scales 
which are indistinct against the 
background; lateral margins of 
mesonotum with white scales. 
Rings on tarsi cream-colored. 

The larva (Figures 112 and 
113) differs from that of the 
nominate subspecies in the shorter 
siphon (index 3.0—3.3), fewer 
denticles in the pecten (16—18), gills 
twice as long as the siphon and 
nearly as thick, and fin with the same 
number of tufts but more weakly- 
branched, with at most 2 branches 



because of the stronger development of the gills. 

Distribution. Eastern part of the distribution of the species. 
USSR: Middle Asia ("Uzbekistan, Turkmenia, Tadzhikistan), to 2,100m above 
sea level. 



202 




FIGURE 113. Aedes pu 1 c hri t a r s is as i a t i c us Edw. Fourth- stage larva: 
1 — stigmal plate; 2 - posterior end, lateral. 



209 



Biology. The larvae breed only in water in tree holes (poplar, walnut, 
Phellodendron, elm, etc.). among roots flooded by water from irrigation 
and in tree trunks at a height of 0.5— 2.0m, fromMarch to October. They 
hibernate and pass the dry period in the egg stage; the eggs are very 
resistant to drought. In Tadzhikistan it occurs together with A. plumbeus 
but mainly in tree holes with less alkaline and saline water. The females 
are not active bloodsuckers. Several generations per year. 



can tans group 

Tarsi with broad, light rings at base of segments (Figure 97, B). Large, 
rarely medium-sized mosquitoes. The hypopygium does not show char- 
acters present in all the species of the group. The larvae have 4—7 shorter 
tufts of the fin before the common base. 

The species bite nearly always in the open, rarely entering houses. 

Eight Palaearctic species belong to the cantans group which are 
characteristic for the forest zone of the Palaearctic and Nearctic; all of 
them occur in the USSR. Identification of the females is very difficult and 
not always certain. The boundaries of their geographical distribution need 
more exact definition. 



4. Aedes (Ochlerotatus) cantans Meigen, 1818* 

Large mosquitoes of dark coloration. Many contrasting white or yellow- 
ish scales among the dark scales on the proboscis, wings, femora and 
abdomen. 

203 Head with yellowish white and brown scales, the brown scales usually 
forming two large spots. Palps of females dark, with scattered white 
scales and white apex; palps of males brown, with indistinct white rings at 
the base of the segments. Mesonotum markedly varying in coloration, with 
chocolate brown scales; lateral parts with whitish gray or cream- colored 
scales forming irregular spots or covering the parts nearly completely. 
Scutellum with white or yellowish scales. Pleurae with spots of white 
scales. Tarsi with white rings at the base of the segments; each ring 
occupies nearly half the length of the segment on the middle segments of 
the hind tarsi. Claws curved (Figure 102). Wings with dark scales, more 
or less mixed with light scales. Abdomen with weakly defined white or 
cream-colored stripes at the base of the tergites, stripes sometimes very 
narrow or nearly absent. More or less numerous light scales among the 
dark scales on the tergites. 

Hypopygium (Figure 114) characteristic: basal lobe of coxite with one 
thick spine, lobe oblong, much higher than wide; wing of claspettes broad. 
Fourth- stage larva (Figure 1 15) large, brown, with paler head and 

204 siphon. Frontal hairs: outer hairs with 6—11 (usually 7—8) branches, 
median hairs with 2—3 (usually 2) branches, situated before the inner hairs 
with 3—4 branches; postclypeal hairs with 2 (rarely 3) short, thin branches, 
situated between the median hairs. Antenna half as long as the head or 
longer, slightly curved, with distinct spines and a short tuft with 6—9 
branches. 

* Many authors name this species A.maculatus Meigen, 1804 , but this name is a synonym ot 
A.rusticus Rossi, 1790. 

210 



(203) 




FIGURE 114. Hypopygium of Aedes cantans Mg. 

Comb on 8th abdominal segment with 28—40 (usually 35—36) scales in 
2 — 4 irregular rows, each scale with a pointed main spine and with small 
spines at the sides of the base, 1—2 spines sometimes distinctly larger. 
Siphon straight, tapering from /s from the base (index 3.0 or slightly less); 
pecten with 21 — 33 (average 26—27) denticles, not reaching middle of siphon; 
hair tuft situated in the middle, with 5—7 branches, not longer than the width 
of the siphon at the base. Stigmal plate (Figure 116) with characteristic 
structure of the posterior appendage of the "stirrup": between the lateral 
205 branches is a variously developed (often weak) median branch in the form 
of an oblong leaf, and the anterior valve is much longer than in the other 
species. 

Saddle of last segment reaching to middle of sides; lateral hair long, 
simple; outer caudal hairs long and simple, inner hairs with 7—9 branches 
half as long as the outer hairs. Fin with 18—19 tufts on the common base 
and with 4 (rarely 5) shorter tufts before it. Gills of varying length, rarely 
as long as the saddle, usually 1.5 times as long. 

Distribution. Western Europe to the Far East. It is very common 
in the western parts of its distribution area but rare in the east. Charac- 
teristic for mixed forests and also enters the taiga. Records from the 
forest-tundra and tundra probably refer to other species. It extends to the 



211 



Northern Caucasus, Crimea and Southern Europe (Italy and Spain) in the 
south. 



(204) 




FIGURE 115. Aedes cantansMg. Fourth- stage larva. Posterior end, lateral. 

Biology. The larvae develop in the spring in puddles, pits and ditches 
with numerous fallen leaves, without vegetation or with poor vegetation on 
the edge of forests or in thinned parts of forests, often together with 
A. communis and A.punctor. A. mid-spring species which appears 
later than the preceding species and has one generation per year. There 
is sometimes a second generation after the summer rains but this is small. 
They hibernate in the egg stage; the eggs are deposited on moist soil near 
the water. It is one of the most widespread mass bloodsuckers which bite 
man and animals. In many parts of the forest belt of the European USSR 
it is one of the most numerous species. It bites almost exclusively in the 
open. 



212 




FIGURE 116. Aedes cantans Mg. Fourth-stage larva. 
Stigmal plate. 

5. Aedes (Ochlerotatus) riparius Dyar and Knab, 1907 
(semicantans Martini, 1920) 

It is closely related to A. cantans, but differs distinctly in males and 
larvae. Medium-sized, rarely large. 

Mesonotum usually with a distinct longitudinal stripe of dark brown or 
bronze scales, lateral parts with lighter golden or cream-colored scales, 
sometimes with narrow, bronze stripes, of the same color as the broad 
median stripe. 

Anterior side of hind femora often light. Claws slightly curved. White 
stripes at base of abdominal tergites relatively broad and distinct; there 
are sometimes narrow, light stripes at the apex of the tergites. 

The coloration is very variable. The North American populations have no 
dark stripe on the mesonotum. Because of the variation, the females cannot 
always be determined with certainty. The structure of the hypopygium 
gives more reliable characters (Figure 117). The basal lobe of the coxite 
is shorter than in A. cantans, about as high as wide at the base; wing of 
claspette narrower. 

Fourth -stage larva (Figure 118) medium-sized, dark. Frontal 
hairs secondarily feathered, outer hairs with 4—9 (average 5—6) branches, 
the median hairs situated before the inner hairs, both pairs with 2 (rarely 3) 
branches; postclypeal hairs situated between the median hairs, thin, short, 
with 3—4 branches. Antenna slightly longer than the head, nearly straight, 
with distinct spines and a tuft with 3—5 branches in the middle. 

Comb of 8th abdominal segment with 6—9 large scales in an irregular 
row, scales with a strongly developed, pointed main spine and smaller 



213 



spines near the base. Siphon straight, slightly tapering from the basal 
third, index 3.5—4.0; pecten with 14—21 (average 16) thin denticles occupying 
slightly more than fa of the siphon, 2—3 distal denticles more widely spaced; 
tuft situated in the middle, slightly basal to it, with 3—5 branches, with slight 
secondary feathering, about fa as long as the siphon. 




FIGURE 117. Hypopygium of A e d e s riparius riparius D.K. 
(after Carpenter and La Casse) 



Saddle of last segment covering A fa of the sides or more; lateral hair 
simple, nearly as long as the saddle; outer caudal hairs simple, longer 
than the siphon, inner hairs forming a fan with 5—8 branches, half as long 
as the outer hairs. Fin with 15-18 tufts on the common base and 4—6 
shorter tufts before it. Gills to 1.5 times as long as the saddle, thin, pig- 
mented, with pointed ends. 



214 




FIGURE 118. Aedes riparius D.K. Fourth-stage larva (after Carpenter 
and La Casse): 

1 — anterior end, dorsal; 2 — posterior end, lateral. 



215 



The differences between the larvae of the two subspecies of A. riparius 
have not been determined. 
208 Distribution. Forests of Europe and North Asia, North America. 

It extends to the boundary of the forest zone in the north and also apparently 
occurs in the tundra. It enters the forest-steppe in the south. The 
boundaries of its distribution have not been exactly determined. This is 
a relatively rare species. 

Biology. The larvae breed in ponds in spring in open landscapes, 
mainly in peat bogs with a bottom of Sphagnu m, together with A . punc- 
tor. They hibernate in the egg stage. Usually one generation per year. 
The females are active bloodsuckers but do not occur in large numbers. 



Aedes riparius riparius Dyar and Knab 

Proboscis dark, with an admixture of light scales which form an in- 
distinct ring in the middle. Palps of female with dark scales and a few 
light scales'. Last segment of hind tarsi with a white ring at the base. 
Wings with dark and light scales. Second abdominal tergite without a 
longitudinal light stripe or with an indistinct stripe not extending on the 
whole tergite. 

Hypopygium (Figure 117) resembling that of A. cantans; stem of 
claspette straight, wing with a short manubrium, plate- shaped widening 
distinct in the middle or near the base. 

Europe, Siberia. The North American form is apparently more closely 
related to the European form. 



Aedes riparius ater Gutsevich, 1955 

Proboscis, palps of female and wings with brown scales; slightly lighter 
scales may be present at the base of the costa. Last segment of hind tarsi 
without a white ring. Second abdominal tergite with a distinct longitudinal 
white stripe extending to the posterior margin. 

Hypopygium (Figure 119) different from that of A. cantans; stem of 
claspette curved, wing with long manubrium, plate- shaped widening present 
only in the distal half, sometimes only in the distal third of the wing. 

Larvae unknown. 

Distribution. Maritime Territory, Khabarovsk Territory, Siberia, 
northeastern part of the European USSR. 

Note on systematics. There are distinct differences in coloration 
and especially in the structure of the hypopygium between the two forms 
(subspecies?). A. r. ater should possibly be considered as a different 
species closely related to the North American A. squamiger Coquillett, 
1902. 



6. Aedes (Ochlerotatus) behningi Martini, 1926 

Smaller than the other species of the cantans group, mainly dark. 



216 



(209) 




209 



FIGURE 119. Hypopygium and claspette of Aedes riparius ater Guts. 

Proboscis and palps brown, with a small admixture of light scales. 
Mesonotum with narrow, bronze or rust-colored scales, sometimes (often 
in males) with an indistinct dark brown longitudinal stripe. Mesonotum 
blackish brown. Tarsi with light rings on the middle segments of the hind 
tarsi about half as long as the segment or shorter. Wings with brown 
scales and a few light scales. Abdomen with brown and light scales, usually 
forming indistinct spots in the midline, one spot sometimes extending to two 
tergites. Distinct, light stripes absent. In males, light scales are usually 
scattered irregularly on the tergites without forming spots. 

Exact determination of the females is not always possible. The hypo- 
pygium (Figure 120) differs distinctly from that of the other species. The 
conical basal lobe of the coxite is without spines and large setae. The 
basal lobe bears a spine in the other species of the cantans group, but if 
a spine is absent the lobe is flat and indistinct. 



217 



(210) 




FIGURE 120. Hypopygium of Aedes behningi Mart. 



Fourth -stage larva (Figure 121) closely resembling that of 
A. excrucians and. often practically indistinguishable from it. Frontal 
hairs coarse, secondarily feathered, outer hairs with 6—8 branches, median 
210 hairs situated before the inner hairs, with 2—3 branches; inner hairs with 
2—4 (usually 2) branches; postclypeal hairs with 6—8 thin, short branches. 
Antenna about half as long as the head, with distinct spines and a hair tuft 
with 8—9 branches in the middle. 

Comb with 20—28 (usually 24) scales in 2—3 irregular rows; scales 
with well developed pointed main spine and several spines at the sides of 
the base. Siphon tapering apically but less strongly than in A. excruc- 
ians (index 3.0— 4.0, average 3.5); pecten with 18— 28 (usually 22— 24) 
denticles, 1—3 denticles larger and more widely spaced; however, the 
distance between them is quite often not greater than between the adjacent 
denticles; tuft with 5 branches, less than half as long as the siphon, situated 
in the middle, slightly distal to it. Hairs on posterior valves of stigmal 
plate curved like a hook but slightly thinner than in A. excrucians. 
The stigmal plate has not been described. 

Saddle reaching nearly to the lower margin of the segment; lateral 
hair simple, as long as the saddle; outer caudal hairs simple, as long as 



218 



the siphon, inner hairs forming a fan with 14—17 branches. Fin with 14—17 
tufts on the common base and 5—6 shorter tufts before it. Gills as long as 
the saddle or slightly longer, not pigmented. 

Distribution. USSR and Siberia, to Krasnoyarsk and Barnaul in the 
east, to the southern boundary of the taiga in the north, to the Ukrainian 
steppe in the south. The species also occurs in Czechoslovakia (Trpis, 
1965b); the distribution in the west has not been determined. 

Biology. The larvae are found in large numbers in floodplains of 
rivers. One generation per year in the spring. 



7. Aedes (Ochlerotatus) excrucians Walker, 1856 

It differs from the three preceding species in its lighter color and the 
structure of the hypopygium. Markedly varying in coloration. 




FIGURE 121. Aedes behningi Mart. Fourth-stage larva. Posterior 
end, lateral (after Martini). 



Proboscis and palps with brown and yellowish brown scales. Mesonotum 
with rust yellow or reddish brown scales, scales at sides usually lighter, 
sometimes with an indistinct broad longitudinal stripe of darker scales. 
Integument of mesonotum brown, rarely blackish brown in the middle, brown 
laterally. Scales on pleurae of thorax yellowish. Anterior side of fore 
femora variegated. Tarsi with light rings which are about half as long as 
the segment on the middle segments of the hind tarsi. Claws large, sharply 
curved; the large denticle nearly parallel to the claw (Figure 122). There 



219 



are specimens with a less developed denticle directed toward the claw 
Wings with dark brown and yellowish white scales, dark scales usually 'pre- 
dominating. Abdomen with brown and yellowish white scales which usually 
form indistinct stripes at the base and sometimes also at the apex of the 
tergites. Tergites often completely variegated. The coloration of the 
abdomen is very variable. For example, in some of the specimens from 
Transbaikalia and the Far East, there are few or no dark scales on the 
abdomen, and light scales predominate on the wings. These specimens 
resemble A.cyprius and A.flavescens. 



(212) 







FIGURE 122. Hypopygiumof Aedes excrucians Walk, (after Carpenter and La Casse): 
1 - claw; 2 - hypopygium. 



610144 



220 



Hypopygium (Figure 122): basal lobe low, densely covered with short 
hairs, without large setae or spines; stem, of claspette long, slightly curved, 
slightly tapering apically; wings of claspette 3 times longer than their 
widest part. 



212 




213 



FIGURE 123. Aedes excrucians Walk. Fourth -stage larva: 
1 — head, dorsal; 2 — posterior end, lateral. 

Fourth-stage larva (Figure 123) large, yellowish brown. Frontal 
hairs with varying secondary feathering: outer hairs with 4—9 (usually 7—8) 
branches, median hairs with 1 — 3 (usually 2), inner hairs situated behind the 
median hairs, with 2—6 (usually 2—3) branches; postclypeal hairs with 2—3 
thin, short branches. Antenna about half as long as the head, covered with 
spines, hair tuft with 3—8 (usually 7—8) secondarily feathered hairs half as 
long as the antenna, situated in the middle, slightly basal to it. 

Comb of 8th abdominal segment with 30-40 (usually 32-36) scales. The 
number of scales varies markedly, there may be about 20 or rarely even 
fewer scales. The larvae cannot be determined by this character. Each 
scale has a distinct, long main spine and smaller, thinner spines at the 



221 



sides of the base. Siphon straight, tapering distinctly from / 5 from the 
base (index 3.2 to 4.4, sometimes 5); pecten with 16-27 spinelike denticles 
which are longer distally, 1-3 distal denticles more widely separated and larger; 
tuft situated distal to the middle, with 5-6 broadly divergent, secondarily 
feathered branches. 

Stigmal plate (Figure 124) characteristic; it differs from that of related 
species in that the distance between the apex of the posterior valves is 
much greater than the length of the plate, in the hook-shaped, thick hairs 
at the apex of the posterior valves and in the narrow, dark, chitinized 
anterior valve, which is slightly wider apically. 

Saddle extending beyond middle of sides of last segment; lateral hair 
simple, as long as the saddle; outer caudal hairs very long, longer than the 
siphon, inner hairs forming a fan with 7-9 branches, half as long as the 
214 outer hairs. Fin with 16—20 tufts on the common base and of 4—6 shorter 
tufts before it. Gills long, not longer than the saddle, but sometimes as long 
as the siphon, not pigmented. 




FIGURE 124. Aedes excrucians Walk. Fourth-stage larva. Stigmal plate. 

Distribution. Europe, Kazakhstan, Siberia, Far East of the USSR, 
Mongolia, Northwest China, Japan, North America. It extends to the tundra 
in the north and sometimes enters the tundra, particularly in river flood- 
plains (north of Norway, to 70°). To the Crimea, Caucasus and mountains 
of Asia Minor in the south. It is distributed throughout the zone of the 
forest and forest-steppe, and sometimes occurs in large numbers. 

Biology. The larvae breed in temporary water bodies in spring, rarely 
in permanent water bodies with vegetation, in open pools in meadows and 
floodplains, at the edge of forests, in thinned out forests, sometimes in 
slightly saline water at the sea shore. There is normally one generation 
in spring from hibernating eggs. Flight begins a week to ten days after the 
first appearance of Aedes. The larvae descend to the bottom if disturbed 



222 



and often remain there for a long time. Stirring brings them to the surface. 
The females are mass bloodsuckers. Flight continues for a long time, 
sometimes also in summer. 

The species transmits tularemia. Natural infections have been found. 



(215) 





FIGURE 125. Aedes b eklemishe vi Den. Fourth-stage larva (after O.N. Sazonova): 
1 — head, dorsal; 2 — posterior end.lateral. 

8. Aedes (Ochlerotatus) beklemishevi Denisova, 1955 
(grandilarva Sazonova, 1956) 

Closely related to A. excrucians. Determination is possible only 
by the larvae. 



223 



Mesonotum with golden or bronze scales. Fore femora light anteriorly, 
with a few dark scales. Claws (Figure 102, 20) large, with sharply diverging 
denticle ( Sazonova, 1958). Wings mainly with dark scales mixed with yellow 
scales. Anterior tergites with distinct light stripes at the base, usually 

215 with narrow stripes at the apex of the tergites. There are no differences 
from A. excrucians in the hypopygium 

Fourth- stage larva (Figure 125) very large, usually more than 

1 cm long, dark. Frontal hairs secondarily feathered, coarse, outer hairs 
with 5-9 (usually 7-8) branches, median hairs with 1-3 (2 in 90%), situated 
before the inner hairs with 2-4 (usually 3) branches; postclypeal hairs thin, 
short, situated between the median hairs, with 2-7 (usually 3-6) branches. An- 
tenna short, less than half as long as the head, slightly curved and with small 
spines, and with a tuft with 2-7 (usually 3-5) branches in the middle. 

Comb on 8th abdominal segment with 14-20 (usually 16-18) scales in 

2 — 3 irregular rows, with a long pointed main spine and small, thin spines 

216 laterally at the base. Siphon long, straight, distinctly tapering in the apical 
third (index 3.1-3.5); pecten with 16-28 (usually 22-26) denticles, 2-5 
rudimentary denticles at the base and 2—6 (usually 3—4) distal denticles 
which are larger and widely separated, the distal denticle in the apical 
third of the siphon. Near it is the tuft with 2-4 (usually 3) thin, slightly 
secondarily feathered branches, not longer than the width of the siphon at 
the base. 

Hairs at apex of posterior valves of stigmal plate not hook- shaped and 
thick as in A. excrucians, but as usual for larvae of Ochlerotatus. 

Saddle of last segment extending to 
nearly /± of the sides; lateral hair 
simple, about /s as long as the saddle. 
Outer caudal hairs simple, slightly 
longer than the siphon, inner hairs form- 
ing a fan with 11—16 (usually 14—16) hairs 
which are half as long as the outer hairs. 
Fin with 17-22 (usually 19-20) tufts on 
the common base and 4—7 shorter tufts 
before it. Gills long, nearly as long as 
the siphon, slightly pigmented. 

Distribution. Described from 
the Kursk, Yaroslavl and Moscow regions 
(Denisova, 1955). Recorded from many 
localities in the European USSR and 
Siberia, to the Sea of Okhotsk in the 
north. Boundaries of distribution not 
known. It often occurs together with 
A. excrucians but is usually less 
numerous. 

Biology. The larvae develop in 
temporary, open water bodies on mea- 
dows, felled areas, rarely in forests. 
One generation in late spring, but 
sometimes also in autum if there is 
rain. Hibernation in the egg stage. 
Widely distributed, but not numerous. 
The females are active bloodsuckers and 
bite during the day and at night. 




FIGURE 126. Hypopygium of A edes 
annulipes Mg. (after Martini) 



224 



9. Aedes (Ochlerotatus) annulipes Meigen, 
1830 (quartus Martini, 1920) 

Closely related to the preceding species, but differs in the coloration of 
thorax and abdomen. There is a small difference in the structure of the 
hypopygium. Because of the large variation of coloration, this character 
is not reliable for determination. 

Mesonotum with a broad, dark longitudinal stripe of chocolate brown 
scales, lateral parts with cream-colored or grayish scales. Abdomen with 
distinct yellowish stripes at the base of the tergites; light scales may also 
be present at the apex of the tergites. 

Hypopygium (Figure 126) mainly as in A. excrucians, but stem of 
claspette relatively thick, not tapering apically, sometimes even thicker 
apically than at the base. 



(217) 




FIGURE 127. Aedes annulipes Mg. Fourth-stage larva. Posterior end, lateral 
(after Marshall). 



225 



Fourth-stage larva (Figure 127) resembling that of A. c ant an s. 
Some larvae of cant an s differing slightly from typical larvae have been 
identified as A. annulipes. 

217 Outer frontal hairs usually with 7 branches, median hairs situated before 
the inner hairs, the median hairs with 1-4 (usually 2), inner hairs with 3-6 
(usually 4) coarse, secondarily feathered branches; postclypeal hairs 
situated between the median hairs, short, thin, with 2—3 branches. Antenna 
slightly curved, darker apically, with distinct spines and a tuft with 4—7 
branches in the middle. 

Comb on 8th abdominal segment with 29—45 (usually 33—34) scales in 
2—4 irregular rows, scales with a pointed main spine and a row of spines at 
the sides of the base. One or two spines near the base may be larger but 
not as large as the main spine. Siphon moderately tapering from the 
middle (index 3.0 to 3.5). Pecten with 17-26 (usually 21-23) densely 
situated denticles and 1—4 rudimentary denticles at the base, extending to 
2 / 5 of the siphon from the base. Tuft with 3-7 (usually 5) branches, situated 
in the middle, slightly secondarily feathered, longer than the width of the 
siphon at its position. Stigmal plate with bifurcate posterior appendage of 
the "stirrup," as usual in the subgenus. 

Saddle of last segment extending beyond the middle of the sides; lateral 

218 hair simple; outer caudal hairs simple, slightly longer than the siphon, 
inner hairs half as long, usually with 8 branches. Fin usually with 16 tufts 
on the common base and 4-5 shorter tufts before it. Gills usually slightly 
shorter than the saddle, lanceolate. 

Distribution. West and Central Europe, from southern Sweden to 
Yugoslavia. In some parts of Poland and Czechoslovakia the species appears 
in large numbers (Skierska, 1963; Dabrowska-Prot, 1964; Trpis, 1965a). 
There are many records from the European and Asian USSR, mainly from 
the forest zone. Most of these records probably refer to other species of 
the cantans group. Without males, identification by females is not 
reliable. This was noted also by Sazonova (1958). A. annulipes is 
probably distributed only in the European USSR: Estonia (Remm, 1957), the 
Ukraine (Val'kh, 1959). 

Biology. The larvae are found in open ponds in spring or in water 
bodies partly shaded by shrubs. In the plains, they occur on meadows at 
the edge of forests, often together with A. cantans but in smaller numbers. 
A mid-spring species with one generation. Hibernation in the egg stage. 
Behavior of the larvae as in A. excrucians. The females are active 
bloodsuckers. 



10. Aedes (Ochlerotatus) flavescens Muller, 
1764 (lutescens Fabricius, 1775) 

Large, rust brown mosquitoes of mainly dark coloration, differing from 
the preceding species in the light abdomen and structure of the hypopygium. 
219 Proboscis and palps of female with brown and yellowish brown scales. 
Palps of male with light scales forming a broad ring. Mesonotum with 
small, dark rust brown or reddish brown scales. Pleurae of thorax with 
grayish white scales covering the entire dorsal mesepisterna. Lower 



226 



mesepimeral setae absent. Legs with light rings about half (or slightly 
more) as long as the segment on the middle segments of the hind tarsi. 
22 Wings with brown and yellowish white scales. Abdomen dorsally with 
yellowish gray scales, sometimes with a few dark scales. 



(218) 




FIGURE 128. Hypopygium of A e d e s flavescens Mull. 

Hypopygium (Figure 128): basal lobe of coxite with a very thick, dark 
spine; the lobe is slightly flattened, and densely covered with short hairs 
on small tubercles. Claspette with a short, straight stem, wing short, with 
a manubrium and a distinct, plate -shaped widening in the apical half. 

Fourth-stage larva (Figure 129) large, semitransparent, yellowish 
brown. Frontal hairs secondarily feathered, outer hairs with 6-9 branches, 
median hairs with 2-4 (usually 3) branches, situated before the inner hairs 
with 2-4 branches; postclypeal hairs very small, with 2-3 branches, 
situated between the median hairs. Antenna short, less than half as long as 
the head, with distinct spines and a tuft with 5-8 branches, with slight 
secondary feathering, situate- 1 , in the middle, slightly more basally, and 
slightly more than half as long as the antenna. 

Comb on 8th abdominal segment with 17—36 (usually 2 0—27) scales in 
3 rows; scales with a long, pointed main spine and with narrow basal part 
and distinct spines at the margin at the base. Siphon (Figure 130) straight, 



227 



(219) 




FIGURE 129. Aedes f 1 a v esc e ns Mull. Fourth-stage larva (after Carpenter 
and La Casse): 

1 — anterior end, dorsal; 2 — posterior end, lateral. 



228 



tapering from the middle, index 3.2—4.0. Pecten 
not reaching middle of siphon, consisting of 
17—28 (usually 19—22) denticles, the distal 3 
denticles more widely spaced, or not; exact 
determination is difficult. Hair tuft situated in 
the middle, with 4—7 (usually 5—6) secondarily 
feathered branches, X k— l /z as long as the siphon. 

Saddle of last segment extending beyond middle 
of sides; lateral hair simple, as long as the saddle, 
outer caudal hairs simple, longer than the siphon, 
inner hairs forming a fan with 12—13 branches, 
half as long as the outer hairs. Fin with 18—19 
tufts on the common base and 6—7 shorter tufts 
before the base. Gills of varying length, shorter 
than the saddle to twice as long, not pigmented, 
the upper pair longer than the lower. 

Distribution. From the Atlantic to the 
Pacific; to Karelia and Yakutsk in the north, 
to Transcaucasia, Middle Asia, North China 
(Tsaidam) and Mongolia. North America, nearly 
to the Arctic Circle in Alaska. 

Biology. The larvae are found in small, 
open or large water bodies on meadows or flood - 
plains in swamps formed by snowmelt or by the 
flooding of rivers and lakes in spring, overgrown 
with vegetation. Their development is slow and 
the flight of females takes place at the end of 
spring and continues to the end of summer. One 

generation per year. Hibernation in the egg stage. 
221 In many parts of the zone of mixed forests and forest- steppe they appear 

in large numbers although they are not usually considered as mass species. 

They enter inhabited areas but rarely enter houses. They bite people and 

farm animals. 

Bacteria of the tularemia were found in A. flavescens during 20 days 

in the laboratory, infectivity by bite during 16 days, and bacteria in the feces 

14 days after experimental infection (Olsuf'ev, 1941a). 




FIGURE 130. Aedes flav- 
escens Mull. Fourth-stage 
larva. Siphon. 



11. Aedes (Ochlerotatus) cyprius Ludlow, 1920 
(freyi Edwards, 1921) 

Large, rust brown mosquitoes, differing from A. flavescens in the 
lighter color of the females; males of darker coloration. 

Proboscis and palps with brown and yellowish scales. Mesonotum with 
light golden or ocher yellow scales, sometimes with a weakly marked 
darker longitudinal stripe. Integument of mesonotum light brown in fe- 
males, dark brown or brown in males. Pleurae of thorax with yellowish 
white scales. Scales not covering dorsal mesepisterna completely, forming 
two more or less separated spots. Lower mesepimeral setae present. 
Tarsi with broad, light yellow rings, contrasting with the dark color of the 



229 



distal part of each segment. On the middle segments of the hind tarsi the 
ring is about 2 / 3 as long as the segment. Wings with brown and yellow 
222 scales, yellow scales usually predominating. The color of the wings and 

legs is not as variegated in males as in females. Dark scales predominate 
on the wings. Abdominal tergites with ocher yellow scales and a few dark 
scales. 




FIGURE 131. Hypopygium of Aedes cyprius Ludl. 



Hypopygium (Figure 131): basal lobe of coxite conical, with one spine 
and thin setae; stem of claspette long, slightly curved; wing of claspette 
without manubrium, plate -shaped widening most distinct in the middle. 

Fourth-stage larva (Figure 132) large, dark, not transparent, with 
bright yellow head and siphon. Entire body of larvae from the 2nd stage 
onward, densely covered with dark, small chaetoids in dense transverse 
rows. The larvae differ sharply from other species of Aedes in this 
character. 

Outer frontal hairs with 2—3 branches, median hairs situated before the 
inner hairs, both pairs with 2 branches, rarely simple; postclypeal hairs 
thin, often simple. Antenna short, slightly curved, thicker at the base, with 
small spines and a tuft with 1—3 branches in the middle. 



230 




""( /// i , 
"in tut 1 1 
\ 1 I //I I \/\'\ J ) 

ii '//'////// 

n ///1U11//1 




" u !itji" u ifi'iint'ii'iii nmi 
"i 1 1 ln m / / // /'Hii'/!,//// / ' / 




/ 3 

FIGURE 132. Aedes cypriusLudl. Fourth-stage larva: 

1 - siphon, lateral; chaetoids on cuticle of 3rd-stage larva (2) and 4th-stage 

larva (3). 

Comb of 8th abdominal segment 9-15 (usually 10-12) scales in two 
irregular rows, each scale with a large, pointed main spine and several 
shorter spines at the sides of the base. Siphon long, gradually tapering 
(index 4.0-4.6); pecten with 17-27 (usually 20-21) denticles, the 2-3 distal 
denticles larger and more widely spaced; tuft situated in the middle, 
slightly distal to it, with 3-4 thin branches which are longer than the width 
of the siphon at the base. 

Saddle of last segment extending to middle of sides; lateral hair simple; 
22 3 outer caudal hairs simple, distinctly longer than the siphon, inner hairs 

forming a fan with 9-10 branches, half as long as the outer hairs. Fin with 
16-18 tufts on the common base and with 3-6 (usually 4-5) short tufts before 
it. Gills as long as the saddle, or at most 1.5 times longer, lanceolate, 
pigmented. 

Distribution. Forest and forest-steppe of the Palaearctic from 
Western Europe to Khabarovsk and the Maritime Territory. It extends to 
the southern taiga (to 63° in Karelia) to the Ukrainian steppe and Central 
Kazakhstan in the south. 

Biology. The larvae develop in large, 40- 80- cm-deep water bodies formed 
by snowmelt or by floods of rivers, with banks overgrown with shrubs and 
herbaceous vegetation. They occur together with the larvae of A.flaves- 
cens, A. excrucians, A. cantans, A. riparius and other species 
but do not adhere to the banks, preferring greater depths with lower tem- 
peratures. A mid -spring species with one generation. Hibernation in the 
egg stage. The females attack man and animals and sometimes are vicious 
bloodsuckers, especially in Siberia. 



231 



rusticus group 

This group contains four relatively rare species, A. rusticus, 
A. refiki, A. lepidonotus and A. subdiversus with a character- 
istic hypopygium: basal lobe of coxite with a group of elongate, transparent, 
lanceolate scales. The females do not have such a distinct character. 
The rusticus group differs from the two preceding groups in the absence 
of light rings on the tarsi. The scales on the proepimera are usually broad 
and straight (differing from the communis group). 

The larvae differ in the presence of additional hairs on the siphon 
besides the pair of tufts in its middle which is usual for Aedes: 2—3 pairs 
of thick hairs on the anterior surface and one pair of thinner and shorter 
hairs on the sides near the distal denticles of the pecten. 



12. Aedes (Ochlerotatus) rusticus Rossi, 1790 
(diversus Theobald, 1901) 

Large mosquitoes, characterized by the dark color of the broad scales on 
the dorsal part of the proepimera. 



(224) 





FIGURE 133. Hypopygium of Aedes rusticus Rossi (after Natvig): 

A - style; B - claspette; C - basal lobe of coxite. High magnification. 



232 



Proboscis and palps with black scales. Mesonotum with golden bronze 
scales, with 2 longitudinal, dark brown or black stripes, which are more 
distinct in the anterior half; there may also be 2 more lateral, narrow dark 
stripes in the posterior half of the mesonotum. Tarsi mainly with dark 
scales. Dark scales predominate on the wings but there are also some 
light scales. Abdominal tergites with light basal stripes which are widened 
in the middle, sometimes forming a nearly continuous longitudinal light 
stripe. 

Hypopygium (Figure 133): basal lobe of coxite with a group of lanceolate 
scales, without spines or large setae; appendage of style twisted; stem 
of claspette long, wing short, irregular, without plate -shaped widening. 



(22 5) 




FIGURE 134. Aedes rusticus Rossi. Fourth-stage larva. Posterior 
end .lateral (after Marshall). 



Fourth-stage larva (Figure 134) large, brown to nearly black. 
224 Outer frontal hairs usually with 8 branches, median hairs with 2 (rarely 3) 
branches, situated before the inner hairs, which have 3 (rarely 2) branches. 



233 



Antennae short, about half as long as the head, thin, nearly straight, covered 
with spines; hair tuft situated near the middle, slightly basal to it, with 
5—6 secondarily feathered branches, half as long as the antenna. 

Comb with 12—18 (usually 14—15) scales in 2 irregular rows; scales 
with a long, pointed main spine, 1—2 shorter lateral spines and thin spines 
at the base. Siphon straight, long (index 3— 3.5), tapering from the basal 
half. Pecten extending in 2 /s °f siphon, consisting of 15—25 densely 
arranged denticles which are longer distally; the pecten continues in 1—3 
longer, widely spaced spines which do not extend to the apical third of the 
siphon. Hair tuft situated in middle of siphon between the distal denticles, 
usually with 6—8 secondarily feathered branches. On the anterior surface 
of the siphon are 3 (rarely 4) pairs of long, thick, secondarily feathered, 
sometimes branched hairs; a pair of thinner hairs with 1—2 branches on 
the sides beyond the middle of the pecten. 

Saddle extending beyond middle of sides of last segment; lateral hair 
simple, nearly as long as the saddle; outer caudal hairs long and thick, 
inner hairs forming a fan with 7—9 branches, nearly half as long as the 
outer hairs. Fin strongly developed, usually with 16 tufts on the common 
base and 3—4 shorter tufts before it. Gills nearly half as long as the saddle, 
elongate -leaf -shaped, the upper pair longer than the lower. 

Distribution. Western, Central and Southern Europe, to Scotland 
and Denmark in the north. North Africa, Asia Minor (var. subtrichurus 
225 Martini, 1927, in which the scales of the mesonotum are usually golden 
brown, whitish on the sides). In the USSR it occurs in the Carpathians. 
Records from the Leningrad Region need confirmation. 

Biology. The larvae occur in puddles and temporary pools with 
herbaceous vegetation in forests, partly shaded by shrubs and trees, rarely in 
open, permanent ponds. They are found together with the larvae of 
A. leucomelas and A. cantans. The annual cycle change s in the 
south. Eggs, larvae and fertilized females may hibernate according to the 
conditions. If the water dries up, there is one generation per year. The 
females are vicious biters but are not mass bloodsuckers. 



13. Aedes (Ochlerotatus) refiki Medschid, 1928 

It differs from A.rusticus in the coloration of the abdominal tergites 
and the structure of the hypopygium. 

Proboscis and palps with black scales, sometimes with some light 
scales. Mesonotum with yellowish scales and a broad, dark, longitudinal 
stripe or with two stripes close together. Pleurae of thorax with spots of 
white scales; scales of proepimera straight, 2 -colored, whitish in the 
lower part, dark brown or black in the upper part. Postcoxal and hypo- 
stigmal spots of scales present. Fore and mid-femora and tibiae with 
brown and light scales anteriorly. Tarsi dark. Wings mainly with dark 
scales, base of radius with light scales. The coloration of the abdomen varies 
markedly. The tergites are often covered with black scales and with an 
indistinct white stripe at the base; the white stripes are often reduced, 
sometimes to small spots at the sides; there may also be narrow, light 
stripes at the apex of the tergites; sometimes the light scales predominate 
and there are only a few dark scales. 



234 



Hypopygium (Figure 135): basal lobe flattened, situated nearly in the 
226 middle of the inner surface of the coxite, covered with lanceolate, trans- 
versely striated scales; near the lobe is another tubercle with 2—3 long 
setae. Stem of claspette long, wing elongate, transversely striated. The 
elongate appendages of tergite 9 bear 7—10 short, strong setae. Very long 
setae at the posterior margin of sternite 8. 




FIGURE 135. Hypopygium of A e d e s refiki Med. 

Fourth-stage larva (Figure 136) small and less strongly pigmented 
than that of A.lepidonotus. Outer frontal hairs with 6— 9 branches, 
median hairs simple, rarely with 2—3 branches; inner hairs situated behind 
the median hairs, with 2—5 (usually 3) thick, dark, secondarily feathered 
branches; postclypeal hairs short, thin, with 3 branches. Antenna short, 
slightly curved, with spines; hair tuft with 12—15 short, thin branches. 

Comb on 8th abdominal segment with 6—11 scales in an irregular row 
with a well developed, pointed main spine and several shorter lateral spines 



235 



at the base. Siphon straight, slightly tapering apically (index 3.0—4.0, 
rarely more); pecten with 12—21 (usually 13—16) densely arranged denticles 
and 1—2 widely spaced, large, smooth spines slightly extending beyond basal 
third of siphon. Hair tuft with 6—9 strongly developed, secondarily 
227 feathered branches situated in middle of siphon but slightly basal to it; on the 
anterior surface are 3 pairs of thick, secondarily feathered hairs and on 
the lateral surfaces a pair of thin hairs with 2—5 branches near the distal 
spines of the pecten. 

Saddle of last segment extending nearly to lower margin of sides; lateral 
hair with 1—3 long, secondarily feathered branches; outer caudal hairs 
long, simple, inner hairs 3 / 4 as long, usually with 9 branches. Fin well 
developed, with 15 tufts on the common base and 2—3 tufts before it. Gills 
leaf -shaped, with rounded ends, as long as the saddle, slightly pigmented. 




FIGURE 136. Aedes refiki Med. Fourth-stage larva (after Medschid): 
1 — head, dorsal; 2— posterior end, lateral. 

Distribution. Southern and Central Europe (Spain, France, East 
Germany, West Germany, Czechoslovakia, Hungary, Yugoslavia); Asia 
Minor. In the USSR it has been recorded only from Simferopol on the 
southern coast of the Crimea. A rare species. 

Biology. The larvae have been found in the Crimea in early spring 
in water with turf holes with grass or clay bottom and with fallen leaves. 
Hibernation in the egg stage. Development begins at the end of February 
and flight at the end of April. Summer populations develop only if water 
is present where the eggs are deposited. Bites man (Velichkevich, 1936). 



236 



14. Aedes (Ochlerotatus) lepidonotus Edwards, 1920 

The species differs in the presence of scales on the postnotum. 



(228) 




FIGURE 137. Hypopygium of Aedes lepi- 
donotus Edw. 



FIGURE 138. Aedes lepidonotus Edw. 
larva (after Medschid): 

1 — head, dorsal; 2 — posterior end, lateral. 



Fourth-stac 



Head dorsally with yellowish scales and black, upright scales lateral to 
them; lateral parts of head with white scales. Proboscis black. Palps 
of female a third as long as the proboscis, black, with a few white scales. 
Palps of male mainly covered with white scales; apex and, sometimes, base 
of segments with black scales; last segment distinctly thickened, with tufts 



237 



of long hairs .yellow hairs with black apex on the penultimate segment and 
228 black hairs on the last segment. Mesonotum with light yellow scales; 
pleurae of thorax densely covered with white scales; postnotum with a 
group of light yellow scales. Legs: femora and tibiae with yellowish white 
scales, except at the apex, which is covered with black scales; tarsi with 
black scales and with some white scales at the base of the first segment. 
Wings mainly covered with yellowish white scales except on the costa and ri, 
which are covered with dark scales. Abdomen usually covered with light 
grayish scales. The middle of the tergites is sometimes nearly completely 
covered with black scales, especially in males. 

Hypopygium (Figure 137): coxite with weakly developed basal lobe with 
a tuft of narrow scales; a process with long setae near the basal lobe 
absent; apical lobe hardly distinguishable; cla^pette with straight, thick 
stem and a short wing of irregular form. 
229 Fourth-stage larva (Figure 138) medium-sized. Outer frontal 

hairs with 8—10, median hairs with 1—3, inner hairs with 3—4 branches, 
situated behind the median hairs; all hairs secondarily feathered; post- 
clypeal hairs short, thin, with 2—3 branches. Antennae less than half as 
long as the head, with spines; hair tuft with 5—6 secondarily feathered 
branches, half as long as the antenna, situated slightly basal to the middle. 

Comb with 6—11 large scales in an irregular row, scales with a pointed 
main spine and lateral spines at the base or lateral spines as large as the 
main spine, which may be rudimentary or absent. Siphon straight, of varying 
form (index 2.5— 4.0), tapering from the basal third. Pecten with 9—21 
(average 14—16) denticles of varying number according to the length of the 
siphon; 1—2 distal denticles more widely spaced but not extending beyond 
basal half; hair tuft with 8—13 secondarily feathered branches situated in 
the middle of the siphon, usually before the distal denticles of the pecten. 
On the anterior surface of the siphon are 2 pairs of thick, long, secondarily 
feathered hairs. There is usually a simple, smooth hair, longer than the 
width of the siphon, on the sides, near the distal denticle of the pecten. 

Saddle of last segment extending to the lower margin of the sides; lateral 
hair well developed, with 3—5 long, secondarily feathered branches; outer 
caudal hairs simple, longer than the siphon, inner hairs two thirds as long, 
usually with 9 branches. Fin with 16 tufts on the common base and 2 much 
shorter tufts before it. Gills shorter than the saddle, leaf-shaped, upper 
pair slightly longer than the lower. 

Distribution. Greece, Turkey (Anatolia). 

Biology. The larvae occur in waters formed by spring floods, slightly 
saline, often together with A. detritus. One generation per year in early 
spring. Hibernation in the egg stage. 

Note on systematics. A. albescens Edwards, 1921 was 
described from Siberia and synonymized by Martini (1931) with A. lepi - 
d o n o t u s . Edwards writes that A. albescens is related to A . 1 e p i - 
donotus,but the postnotum (postscutellum) is without scales. Nearly all 
the scales on the body are whitish but brown on the greater part of the 
proboscis and on the palps . The scales on the proepimera are whitish, narrow, 
some of them curved. Palps of female long, a third as long as the proboscis. 
Wing membrane whitish; all scales at anterior margin of wing whitish 
yellow, some scales on the radius dark, whitish yellow on the legs, brown 



238 



on the last tarsal segments; tarsi without white rings. Claws: formula 
1:1, 1:1, 1:1. Siberia: Omsk (Grand); one female in the museum in 
Helsinki. Martini (1931), who examined this specimen, stated that one scale 
was distinctly preserved on the postnotum and that there were several 
dots, probably the points where the scales had fallen off. 

Because of this description of A. albescens and the synonymy of 
albescens and lep id onotus, we identified the mosquitoes of the 
group rusticus from North Kazakhstan as A . lep idonotus, but as we 
have not found larvae, males or mosquitoes with scales on the postnotum 
which would correspond to the description of this species, we think that the 
mosquitoes of the group rusticus from Siberia and Kazakhstan should 
be considered as A.subdiversus (see below). The status of A. al- 
bescens Edw. is not clear: it is either identical with a species of the 
rusticus -lepidonotus group or with subdiversus. We suggest 
that A. lepidonotus has not been found in the USSR and that A . a 1 - 
bescens is a doubtful species. 



230 15. Aedes (Ochlerotatus) subdiversus Martini, 1926 

The species differs from the three preceding species of the rusticus 
group in the structure of the hypopygium. Head with light and dark scales 
dorsally. Proboscis and palps (female) with black scales, often also with 
some light scales. Mesonotum with a broad, sometimes indistinct, longi- 
tudinal stripe of bronze or yellowish brown scales which consist apparently 
of three small stripes, lighter in the middle and darker laterally. Pleurae 
of thorax with spots of silvery or yellowish cream-colored scales. Upper 
half of proepimera with yellowish or darker, bronze, straight or slightly 
curved scales. Postcoxal and hypostigmal spots of scales present, the 
latter forming part of a longitudinal stripe below the anterior spiracle. 
Legs: femora and tibiae of fore legs variegated anteriorly; tarsi brown, 
without white rings but with a large admixture of light scales. Wings mainly 
with dark scales, light scales mainly at the base and on some veins, mainly in 
the anterior part of the wing. Abdomen: tergites with whitish gray or 
cream-colored scales, often with some dark scales, which sometimes form 
indistinct spots; proportion of light and dark scales on abdomen strongly 
varying. 

Hypopygium (Figure 139): style straight or slightly sickle -shaped (not 
undulant); basal lobe of coxite with narrow, lanceolate scales, and near it a 
small process with 2—3 long and some shorter setae; stem of claspette 

231 long, with short wing, not transversely striated; phallosome sclerotized, 
serrated at the apex, with a constriction in the middle; appendages of 
tergite 9 with 8—10 short, thick setae; posterior margin of sternite 8 densely 
covered with long, thin hairs in the middle. 

Fourth-stage larva large, yellowish brown, with slightly paler 
head and siphon. 

Head 1.5 times wider than long. Frontal hairs slightly secondarily 
feathered, outer hairs with 5, median hairs with 2—3, inner hairs with 2 
branches; median hairs situated before the inner hairs, slightly more widely 
spaced; between them are the 2 -branched postclypeal hairs. Antennae 



239 



half as long as the head, covered with spines, hair tuft situated in the middle, 
with 3 branches, about a third as long as the antenna. Mouth parts of the 
mixed type, substrate and planktonic, as in A. rusticus. 

Comb with 12—17 (usually 14—15) scales with a long, pointed main spine 
and small denticles at the base, usually in 2 irregular rows. Hairs behind 
comb as in A. rusticus. Siphon (index 3—3.3) with pecten occupying 
about % °f ^s length and consisting of 11—16 denticles, 2—5 near the base 
rudimentary; 1—2 distal denticles larger and more widely spaced. Each 
denticle in the form of a spine with accessory spines at the base. Between 
the distal denticle of the pecten and the apex of the siphon are 3—4 smooth, 
longer spines. Hair tuft in the form of a long, simple hair near the distal 
denticle of the pecten or slightly apical to it. Lateral to the median 
denticles of the pecten is a thin, 2 -branched hair. On the anterior surface 
of the siphon are 3—4 pairs of long, slightly secondarily feathered hairs, 
sometimes arranged in a zigzag row. 



(230) 




FIGURE 139. Hypopygium of A e d e s subdiversus Mart. 



240 



Saddle of last segment covering about % of the sides. Outer caudal hairs 
simple, as long as the siphon, inner hairs forming a fan with 11 branches. 
Fin with 14—15 tufts on the common base and 6—7 shorter tufts before it. 
Gills and lateral hairs were damaged in the specimens available. 

Note on systematics. The species varies widely in coloration. 
The light variants which we identified as A.lepidonotus in the past 
(see above) have light scales on the tergites, a large admixture of light 
scales on the palps and proboscis of the female, and lighter scales in the 
dorsal half of the proepimera. There are transitions between the "typical" 
forms and the light variants. 

Distribution. From the Volga (the species was described from the 
Saratov Region; Martini, 1926, 1928) to Transbaikalia. Common in 
Kazakhstan and southern Siberia. The boundaries of the distribution area 
have not been determined. 

Biology. One generation per year in spring. Mass attacks on people 
and farm animals have been observed in some localities, e. g. in the 
Kustanai Region. 



communis group 

Usually medium -sized mosquitoes. They differ from the species of 
the caspius and cantans groups in the dark tarsi without light rings 
and from those of the rusticus group in the structure of the hypopygium. 
This large heterogeneous group contains 16 Palaearctic species, some of 
them mass bloodsuckers. They usually predominate among mosquitoes 
232 inhabiting forests and tundra and attack people and farm animals. They 
usually bite in the open, but when they are numerous, they enter houses and 
cattle sheds. 

Most species of this group have only one spring generation. After the 
rains, there may be a small summer generation, much smaller than the 
spring generation. The seasonal curve of activity of this group has usually 
one peak, with the maximum at the end of spring and in early summer. In 
the northern taiga and tundra, where most of the mosquitoes of this group 
are found, they appear in small numbers in the middle or at the end of 
summer. 

An important systematic character is the arrangement of the spots of 
scales on the pleurae of the thorax (Peus, 1933; Figure 99). The scales 
are easily lost and the females of the communis group can therefore be 
determined reliably only from well preserved specimens. Determination 
can be certain only by the hypopygium of the males and by the larvae. 



16. Aedes (Ochlerotatus) communis De Geer, 1776 
(nemorosus Meigen, 1818) 

Medium-sized, differing from related species in the absence of a post- 
coxal spot of scales and the structure of the hypopygium. 

Palps of female and proboscis with brown scales. Last segment of 
palps of male thickened. Mesonotum with golden yellow or bronze scales, 



241 



usually with 2 approximate longitudinal dark stripes which may be fused, 
or sometimes indistinct or absent. Pleurae of thorax with white scales 
without silvery sheen. Postcoxal spot absent, i. e. scales absent on the 
small membranous part between the base of the fore coxae and the anterior 
233 margin of the mesepisternum. Spot of scales on the mesepisterna extend- 
ing to the anterior corner. Spot of light scales below the anterior spiracle 
(hypostigmal spot) absent. Scales on the mesepimera extending to the 
lower margin. Tarsi with blackish brown scales, sometimes with a few 
gray or whitish scales. Wings with dark scales; light scales only for a 
short distance on the base of the costa. Abdomen with transverse white 
stripes dorsally near the anterior margin of the tergites, other part of 
tergites with dark scales; stripes on abdominal segments 2—5 more or 
less of the same width, not distinctly narrower in the middle. 



(232) 




FIGURE 140. Hypopygium of Aedes c omm unis Deg. (after Natvig): 

A - basal lobe of coxite; B - lobes of tergite 9; C - wing of claspette. 
High magnification. 

Hypopygium (Figure 140): coxite with long hairs and well developed 
basal and apical lobes. Basal lobe half -rounded, with a long spine; a row 
of short hairs curved to one side at the inner margin of the lobe. Stem of 
claspette long, straight or slightly curved, wing very narrow, long, not 
widened, slightly sclerotized. 

Fourth-stage larva (Figures 141— 143) large, of varying color, 
usually dark. Frontal hairs secondarily feathered, outer hairs with 4—8 
234 branches, median hairs situated before the inner hairs, both pairs simple, 
rarely one of the hairs with 2, exceptionally with 3, branches; postclypeal 
hairs thin, short, with 3—4 branches. Antenna about half as long as the 
head, covered with spines and slightly curved, with a hair tuft with 6—7 



242 



;233) 



branches in the middle, not half as long 
as the antenna. 

Comb usually with 60—80 scales 
forming an irregular triangular spot, 
scales without a main spine, their 
distal end with a row of pointed spines 
which sometimes extend to the lateral 
margin. Siphon tapering from the 
middle, index 2—3 (average 2—7). 
Pecten with 17—26 (usually 21—22) 
closely spaced denticles, the 1— 4 basal 
denticles rudimentary, the others be- 
coming longer apically; pecten not ex- 
tending to middle of siphon. Hair tuft with 
5—9 (usually 6—7) secondarily feathered 
branches situated in the middle, about a third as long as the siphon. 

Saddle extending to middle or lower margin of last segment; lateral 
hair short, simple; outer caudal hairs simple, longer than the siphon, inner 
hairs forming a fan with 6—8 shorter branches. Fin with 16—19 tufts on the 
common base and 2 shorter tufts before it. Gills long, 1.5—2 times as long 
as the saddle, pigmented, with pointed ends. 




FIGURE 141. Aedes communis Deg. 
stage larva. Head, dorsal. 



Fourth- 



(233) 




FIGURE 142. Aedes communis Deg. Fourth -stage larva. Posterior end, lateral. 



243 




FIGURE 143. Aedes communis Deg. Fourth-stage 
larva: 

1 — scales of comb; 2 — stigmal plate. 

Distribution. Forest and tundra zone of both hemispheres : Europe, 
Siberia, Far East (USSR), North America. It extends in the north to the 
Arctic Ocean (Murmansk, mouth of the Kolyma), in the south to East Kazakh- 
stan, the Northern Caucasus, Ukrainian steppes and Bulgaria. 

Biology. The larvae are found in permanent and temporary water 
bodies formed by melting snow. Their bottom covered with fallen leaves. 
They are found in the south in shaded forest habitats, in open areas in the 
north. They may hatch from hibernating eggs while still under the ice. 
One generation per year, but there may be another generation in a wet 
autumn. After a dry spring, numerous larvae may appear with the summer 
rains. The forest zone is one of their most common habitats. They take 
235 first place among mosquitoes attacking people and animals in many 

localities. Flight begins earlier than in most other species of Aedes. 
Their numbers decrease sharply in midsummer. 

They apparently take part in the transmission of tularemia (natural 
infections have been found). 



244 



17. Aedes (Ochlerotatus) pionips Dyar, 1919 

Resembling A. coramunis,but differing in the presence of a postcoxal 
spot of scales and in the structure of the larvae. 




\ \ 

1M w 






FIGURE 144. Hypopygium of Aedes pionips Dyar (after Carpenter and 
La Casse) 



Head with light yellowish brown scales dorsally. Proboscis and palps 
with brown scales. Last segment of palps of male thin. Mesonotum with 
yellowish gray or golden bronze scales with an indistinct broad longitudinal 
brown stripe or 2 approximate stripes divided by a narrow stripe of golden 
scales. Lateral longitudinal dark stripes usually present in the posterior 
half of the mesonotum. Spots of scales on pleurae of thorax as in A. c om - 
munis, except that there are also scales on the postcoxal membrane. 
Wings with dark scales and usually some light scales at the base. Abdomen 



245 



236 




FIGURE 145. Aedes pionipsDyar. Fourth-stage larva (after Carpenter and 
La Casse): 

1 - anterior end, dorsal; 2 - posterior end, lateral. 



246 



with brown scales dorsally and light transverse stripes of more or less the 
same width. 

Hypopygium (Figure 144) as in A. communis; the differences of the 
hypopygium of the two species ("Vockeroth, 1952) are small, apparently in 
the range of intraspecific variation. The hypopygia of the two species are 
indistinguishable for practical purposes. 

Fourth- stage larva (Figure 145) large, dark brown, head 1.5 times 
wider than long. Antennae thin, slightly curved, more than fa as long as the 
head, covered with spines; hair tuft situated near the middle, slightly more 
basally, with 7—13 (average 8—9) slightly secondarily feathered branches, 
about half as long as the antenna. Frontal hairs: outer hairs with 5—9 
(average 7—8) secondarily feathered branches, median hairs with 3—5 
branches (with 3 branches in 50% of the specimens), inner hairs with 3—5 
(average 4, rarely 2) branches; postclypeal hairs with 3—5 short, thin 
branches, situated between the median hairs. 

Comb on 8th abdominal segment with 61—78 scales (average 68, rarely 
fewer than 60), scales oblong, slightly wider distally, their margin with a 
row of sometimes asymmetrical, short spines which are smaller toward the 
base. Siphon straight, slightly tapering, index about 3.0. Pecten with 18—24 
closely spaced denticles which are longer distally and with 1—6 rudimentary 
denticles at the base, pecten fa — fa as long as the siphon. Hair tuft situated 
in the middle, with 4—9 (average 6) secondarily feathered branches. Saddle 
of last segment extending to fa of the sides; lateral hair thin, simple; outer 
caudal hairs distinctly longer than the siphon, simple, inner hairs with 9—14 
(average 12) smooth branches, as long as the last segment. Fin with 17—21 
(average 19) tufts on the common base and with 2—3 short tufts before the 
base. Gills lanceolate, pointed, pigmented, of varying length, as long as the 
saddle or twice as long. 

Distribution. Alaska, Canada, Northwestern U. S. A. Until recently it 
has not been found in the Palaearctic (Sazonova, 1958) and has probably not 
been distinguished from A. com munis. It occurs in the northern half of 
the European USSR, West and East Siberia, Khabarovsk Territory and Kamchat- 
ka. The boundaries of distribution have not been determined. Records of the 
wide distribution and large numbers of A. comm unis in northern regions 
refer perhaps to A. pionips and A. hexodontus (see below). 

Biology. The larvae hatch from hibernating eggs in early spring in 
pools and other small water bodies formed by snowmelt in open and forest 
localities, in plains and mountain landscapes to about 1,000 m above sea 
level. 



18. Aedes (Ochlerotatus) punctor Kirby, 1837 
(meigenanus Dyar, 1921) 

Closely resembling the two preceding species in coloration, differing 
from them in the structure of the hypopygium, the absence of light scales 
on the wings, and the form of the light stripes on the abdomen (with an 
indentation in the middle). 

Proboscis and palps with blackish-brown scales. Mesonotum with golden 
bronze or reddish bronze scales, usually with a dark brown longitudinal 



247 



238 stripe or with 2 approximate dark stripes. Specimens with a uniformly- 
colored mesonotum have been found. Postcoxal spot present (in contrast 
to A. communis). Spot of white scales extending to the anterior corner 
on the mesepisterna and to the lower margin on the mesepimera. Hypo- 
stigmal scales absent. Tibiae and tarsi with dark scales. Wing with dark, 
nearly black scales; there may be rarely a few yellowish scales at the 
base of the costa. The light transverse stripes on the abdominal tergites 
are distinctly narrower in the middle; there are sometimes only white spots 
at the sides of the segment connected by a narrow, light stripe at the base 
of the tergites. 




FIGURE 146. Hypopygium ofAedes punctor Kirby (after Carpenter and La Casse) 



248 



Hypopygium (Figure 146): apical lobe of coxite flattened, covered with 
short, curved hairs, extending proximally nearly to the middle of the coxite, 
239 basal lobe with one spine, strongly convex, densely covered with short hairs 
on small tubercles. Claspettes with short, straight stem, wings slightly 
widened in the middle, long, lanceolate, 2—3 times longer than wide. The 
wing of the claspette is usually brown in preparations, without a trans- 
parent widening as in most species of Ochlerotatus. 




FIGURE 147. Aedes punctor Kirby. Fourth- stage larva. Posterior end. lateral. 

Fourth-stage larva (Figure 147) medium-sized to large, older 
larvae dark brown. Frontal hairs: outer hairs with 2—8 (average 5) 
branches, median and inner hairs with 1—3 (usually 2) branches with slight 
secondary feathering; postclypeal hairs situated between median hairs, 
thin, short, with 2—4 branches. Antennae less than half as long as the head, 
slightly curved, hair tuft situated in the middle, with 4—7 branches, less than 
half as long as the antenna. 

Comb with 11-24 (usually 14-15) small scales (0.07-0.08 mm), with a 
main spine and small spines at the base, arranged in 2—3 irregular rows. 
The number of scales is very variable even on the left and right side of a 
specimen. Siphon straight, slightly tapering (index about 3.0); pecten 
distinctly not reaching middle of siphon, with 14-26 (usually 19-22) denticles, 
1—6 rudimentary denticles at the base; denticles closely spaced, becoming 



249 



longer apically; hair tuft with 3—9 (average 5) branches in the middle of 
the siphon, often basal to it. 

Saddle surrounding last segment like a ring; on the ventral side is a 
deep indentation in which the anterior tufts of the fin are situated; the 
240 lower margins of the ring are rarely not fused, but form a narrow median 
slit. Lateral hair simple, as long as the saddle; outer caudal hairs long, 
simple, inner hairs with 5—8 branches. Fin with 16—19 tufts on the common 
base and 1—2 tufts before the base. Gills long, tapering, pigmented, usually 
1.5—2 times as long as the saddle. 

Distribution. Forest and tundra zones of both hemispheres: Europe, 
Siberia, Far East, North America. Extending in the north to the Arctic 
Ocean and in the south to the Crimea, Northern Caucasus and Eastern 
Kazakhstan. It occurs in the mountains of Algeria. Its distribution to the 
north should be determined more exactly because the related A . hexo- 
dontus has been found there. 

Biology. The larvae are found in small, open or partly shaded water 
bodies formed by snowmelt in turf or forest swamps, their bottom covered 
with rotting leaves, needles of conifers, moss or herbaceous vegetation. 
In the taiga and tundra and also in the forests of the Maritime Territory, 
A.punctor is an important bloodsucker, biting man and farm animals. 
It is an early spring species which appears together with A. communis 
or a little later. The mosquitoes sometimes breed again after the summer 
rains, but not in large numbers. Natural infections of A. punctor with 
tularemia have been found. 



19. Aedes (Ochlerotatus) hexodontus Dyar, 1916 

Closely resembling A. punctor, but differing in the light scales at 
the base of the wing, and from A. communis in the presence of a post- 
coxal spot of scales. It closely resembles A. pionips, but differs from 
it in its darker coloration, especially of the mesonotum. It can be reliably 
determined only by the larvae. 

Head mainly with dark brownish bronze scales dorsally. Proboscis and 
palps with black scales. Mesonotum usually with rust brown scales and 
there is sometimes an indistinct dark longitudinal stripe. Wing with dark 
scales, light scales present only at the base. Light scales are sometimes 
few, but they often cover the base of the costa and radius to the humeral 
cross-vein. Abdomen with dark scales and light stripes which are usually 
slightly narrower in the middle, rarely of the same width. The brown 
scales of the tergites sometimes have a slight bronze tone and do not differ 
much from the grayish white scales of the stripes at the base of the 
segments. 

Hypopygium (Figure 148). The males of A. hexodontus and A . pu n c 
tor are practically indistinguishable. 

Fourth- stage larva (Figure 149) dark or yellowish brown, head 
1.3—1.4 times wider than long. Frontal hairs thick, with sparse secondary 
feathering; outer hairs with 4 (3—6) branches, median hairs situated before 
the inner, both pairs usually simple, rarely with 2 branches; 2-branched 
hairs are often found in the central and southern parts of the area of 



250 



distribution. Postclypeal hairs thin, with 2-3 branches. Antennae about 
half as long as the head, with spines, hair tuft with 3-4 branches, not reaching 
the apex. Mouth parts of the mixed type, substrate feeding predominating. 



241 




/ / \ Vt - 






FIGURE 148. Hypopygium of Aedes hexodontus Dyar (after Carpenter and La Casse) 



251 



(242) 




FIGURE 149. Aedes h e x o d o n t us Dyar. Fourth- stage larva (after Carpenter 
and La Casse): 

1 - anterior end, dorsal; 2 - posterior end, lateral. 



610144 



252 



Comb usually with 7 scales (6-9), rarelylO-ll), with a distinct main 
spine and small, thin spines at the base. Siphon (index about 3) straight, 
tapering. Pecten with 12—17 closely spaced denticles, occupying /$— / 5 
of the length of the siphon from the base; the distal denticle is sometimes 
larger than the others and rarely more widely separated. Hair tuft 
situated distal to the pecten, with 2—7 (4—5) branches. 

Last segment of abdomen with a ring-shaped saddle surrounding it. 
243 Outer caudal hairs long, simple, inner hairs with 4—10 (6—8) branches 

forming a fan; lateral hair simple. Fin with 16-18 tufts on the common 
base, 2 (rarely l) shorter tufts before the base. Gills lanceolate, slightly 
pigmented, usually twice as long as the saddle. 

Distribution. The species was until recently confused with other 
species of the communis group so that its distribution was not clear. 
In the USSR it occurs to the Kola Peninsula, Karelia in the north 
(Prionezhskii District), Leningrad area (Petrodvorets) and in the Nenets 
National District (Nizhnepecherskii District) and in West and East Siberia. 
It is apparently widely distributed in the subarctic tundra and taiga, and 
in mountain regions in the south. Northern Sweden (Vockeroth, 1954), 
Canada, Alaska and mountainous regions of the U. S. A. 

Biology. An early spring species; the larvae appear when the snow 
begins to melt in various types of small water bodies; they occur in the 
mountains in water with a stony bottom. The adults apparently fly earlier 
than A. punctor and are vicious bloodsuckers. 



20. Aedes (Ochlerotatus) sticticus Meigen, 1838 

It differs from the preceding species in the smaller scales on the 
mesepimera, coloration of mesonotum and tibiae, and in the structure of the 
hypopygium. 

Proboscis, palps and wings with dark scales. Mesonotum with a broad, 
distinct longitudinal stripe (or with 2 approximate stripes) of dark brown 
scales. There may also be 2 longitudinal dark lateral stripes. Lateral 
parts of mesonotum with silvery gray or cream-colored scales. Spot of 
white scales on mesepimera not reaching their lower margin. Fore and 
mid-femora dark anteriorly. Light scales forming a longitudinal stripe on 
the outer surface of the hind tibiae which is sometimes indistinct. Abdomen 
dorsally with black scales with a bronze sheen and with white, triangular 
spots on the sides connected by transverse stripes at the base of the seg- 
ments; stripes narrower in the middle, sometimes interrupted. 

Hypopygium (Figure 150) resembling that of A. punctor, but differing 
in the form of the basal lobe of the coxite and wing of the claspette: basal 
lobe separated from the coxite in the distal part; wing of claspette with a 
small, transparent widening, wing short, about twice longer than wide. 

Fourth-stage larva (Figure 151) large, brown, with lighter head 
and siphon. Frontal hairs secondarily feathered: outer hairs usually with 
5 branches, median hairs situated before the inner, both pairs with 2—3 
branches, rarely more; one of the hairs may be rarely simple. Postclypeal 
hairs situated between the median hairs, short, with 1—4 branches. Antenna 
short, 0.4—0.5 times as long as the head, with sparse spines; hair tuft with 
4—5 short branches, situated in the middle, slightly basal to it. 



253 



(2 43) 




FIGURE 150. Hypopygium of Aedes sticticus Mg. (after 
Carpenter and La Casse) 

Comb on 8th abdominal segment with 16—27 (usually 20—24) scales in 
2—3 irregular rows, scales with a moderately large main spine and a few 
shorter spines at the sides of the base, spines near the main spine nearly 
as large as this. Siphon relatively short, straight (index 2—2.5, rarely 3.0), 
tapering from the basal third. Pecten extending beyond the middle, usually 
with 20—25 equally spaced denticles, the distal denticle sometimes slightly 
more widely separated than the others. Hair tuft situated beyond the middle, 
with 5—6 secondarily feathered branches, as long as the width of the siphon 
at the base. 



254 




FIGURE 151. Aedes sticticus Mg. Fourth-stage larva (after Peus): 
1 — 8th abdominal segment, lateral; 2 - head, dorsal. 

Saddle of last segment extending to middle of sides or to its lower margin; 
lateral hair short, simple; outer caudal hairs simple, much longer than the 
siphon; inner hairs with 6—7 branches. Fin with about 20 tufts on the 
common base and 1 — 2 tufts before it. Gills longer than the saddle, pointed, 
not pigmented. 

Distribution. In Europe to Finland and Karelia in the north, to 
Yugoslavia, the southern Ukraine and Northern Caucasus in the south; 
Siberia, Transbaikalia, Khabarovsk Territory, Maritime Territory; Japan, 
North America. 

Biology. The larvae occur in water bodies formed by snowmelt or 
flooded rivers, with partly shaded trees, their bottom covered with fallen 
245 leaves, often together with larvae of A . vexans. Hibernation in the egg 
stage. The eggs may remain viable for 2—3 years in the soil. Character- 
istic for the zone of mixed forests and forest-steppe. The records of 
A. sticticus from the northern taiga and tundra are probably not correct. 
In some places (Western Ukraine, Transbaikalia) they bite in large numbers. 
One generation per year. They appear much later than A. communis 
and A. punctor. This is more a summer than a spring species . 



21. Aedes (Ochlerotatus) nigrinus Eckstein, 1918 

Closely related to A. sticticus, but differing from it in details of 
coloration of the antennae, legs and wings (Peus, 1933). 

First segment of flagellum of antennae thick, black (not thick, yellow at 
the base in A. sticticus). Base of costa, subcosta and base of media 
with white scales. Fore and mid-femora variegated anteriorly. Light 
stripes on the abdomen not narrowing in the middle or narrowing only 
slightly. 

Hypopygium: basal lobe of coxite shorter and wider than in A. sticti 
cus, its outer margin uniformly convex; wing of claspette with a small, 
plate- shaped widening, twice as long as wide. 



255 




246 



FIGURE 152. Aedes nigrinus Eck. Fourth-stage larva (after Peus): 
1 — head, dorsal; 2 - 8th abdominal segment, lateral. 

Fourth-stage larva (Figure 152) resembling that of A. sticti- 
cus. Frontal hairs slightly secondarily feathered, outer hairs with 
5 branches, median hairs situated before the inner, both pair's simple, rarely 
one hair 2-branched; postclypeal hairs short, with few branches. Antenna 
slightly less than half as long as the head, with sparse spines, hair tuft with 
3-5 short branches situated at 2 / 5 of the length of the antenna from the base. 

Comb usually with 10-12 (rarely 15-17) scales, which are more elongate 
and have a longer main spine than in A. sticticus. Siphon short, straight, 
slightly tapering (index 2.0 or slightly more). Pecten nearly reaching 
middle of siphon, usually with 17 denticles and several rudimentary denticles 
at the base. Tuft with 4-6 branches situated in the middle or slightly 
beyond it. 

Saddle of last segment extending nearly to the lower margin of the sides; 
lateral and caudal hairs as in A. sticticus. Fin usually with 13-14 
tufts on the common base and 3 — 4 tufts before the base. Gills of varying 
length, sometimes as long as the siphon. The upper pair is longer than the 
lower in specimens with shorter gills; the two pairs are of the same length 
in specimens with long gills. 

Distribution. Europe: West and East Germany, Denmark, southern 
Scandinavia, Finland, Czechoslovakia. Also found in some parts of the 
European USSR: Estonia, Leningrad and Vologda regions, Northern Urals, 
West Siberia. A rare species which inhabits mainly open meadows. 

Biology. The larvae occur in open pools on meadows with rich 
vegetation, formed by snowmelt or flooding. Hibernation in the egg stage. 
Records of several generations in a summer need confirmation. 



22. Aedes (Ochlerotatus) hungaricus Mihalyi, 1955 

This is a recently described species and its systematic position is not 
quite clear. 

Small mosquitoes. Proboscis and palps of females with dark scales. 
Mesonotum with whitish gray scales and a distinct, dark brown longitudinal 
stripe, which is divided into two stripes in the posterior half. Spot of light 
scales nearly reaching the anterior margin on the mesepisterna and not 



256 



reaching the lower margin on the mesepimera as in A. sticticus and 
A. nigrinus. Hypostigmal and postcoxal spots absent. Tarsi dark, 
without light stripe on the outer side of the hind tibiae. Wings with dark 
scales, which are especially dense on the costa and r-i-. Abdomen with dark 
scales dorsally and light stripes at the base of the tergites which are 
slightly narrower in the middle; spots of white scales at sides of tergites. 




FIGURE 153. Hypopygium of A e d e s hungaricus Mih. 
(after Mihalyi) 



Hypopygium (Figure 153): coxite narrow, with dense hairs; ends of 
hairs of left and right coxite slightly overlapping. Basal lobe of coxite 
moderately convex, with 2 strong setae, one long and curved, the other 
thinner, short and straight. Apical lobe weakly developed. Claspettes with 
nearly straight stem and long wing which is distinctly wider in the middle; 
the narrow basal part of the wing ("manubrium") is distinct. Phallosome 
short, ovoid. Appendages of tergite 9 with 3—4 long, curved setae. 

Fourth-stage larva yellowish- brown, with darker head and light 
spots at points of attachment of muscles. Frontal hairs: outer hairs with 
6—8, median hairs with 2 branches, inner hairs simple, situated behind the 
median hairs, all hairs slightly secondarily feathered. Postclypeal hairs 
indistinct between the median frontal hairs. Antennae slightly more than 
half as long as the head, with sparse spines, hair tuft with 8—10 branches, 
situated at j$ of the length of the antenna from the base. 

Comb on 8th abdominal segment with 16—24 (average 20) scales with a 
long, pointed main spine and shorter spines at the base; the marginal 
spines near the main spine may be as long as this. Siphon cylindrical, 
247 widest at a third from the base; index 2.0—2.4 (average 2.2); pecten with 
19—24 (average 20) closely spaced denticles which are longer distally and 
extend beyond the middle of the siphon, each denticle in the form of a thin, 
slightly curved spine with several accessory denticles at the base; tuft 



257 



with 3—5 branches with slight secondary feathering, situated beyond the 
middle of the siphon, distinctly separated from the distal denticle. 

Saddle of last segment covering the sides to the middle, lateral hair 
simple, half as long as the saddle; outer caudal hairs simple, inner hairs 
forming a fan with about 10 branches. Fin with 15 tufts on the common 
base and 3, rarely 4 (young larvae), shorter tufts before the base. Gills 
lanceolate, not pigmented, upper pair longer than the lower, distinctly longer 
than the saddle. 

Distribution. Hungary, Danube valley (Mihalyi and Gulyas, 1963). 
Apparently characteristic for floodplains. Not found in the USSR. 

Note on systematics. A.hungaricus resembles the other 
species of the communis group in coloration, particularly A.sticticus 
and A. nigrinus, but differs from them in the absence of a light stripe on 
the outer side of the hind tibiae; the structure of the hypopygium, the 
presence of two strong setae on the basal lobe of the coxite and the weak 
development of the apical lobe are more characteristic for the species of 
the caspius group. 



23. Aedes (Ochlerotatus) diantaeus Howard, 
Dyar and Knab, 1917 

They are usually slightly larger than specimens of the communis 
group. Fresh specimens are distinguished by the steel sheen of the black 
scales on the proboscis, legs and abdomen. 




FIGURE 154. Hypopygium of A e d e s diantaeus H.D.K. (after Natvig): 

A — basal lobe of coxite; B — wing of claspette; C — lobes of tergite 9. 
High magnification. 



Proboscis and palps with black scales. Mesonotum with golden or 
silvery gray scales, with a broad longitudinal dark stripe or 2 approximate 
stripes. Spot of white scales on the mesepisterna not extending to their 



258 



(248) 











259 



anterior margin (a difference from all preceding species of the communis 
group). Postcoxal and hypostigmal scales absent. Wings, tibiae and tarsi 
249 with dark scales. i\.bdomen with black scales dorsally and triangular, white 
spots at the sides, sometimes connected by narrow stripes at the anterior 
margin. 

Hypopygium very characteristic (Figure 154). Inner surface of coxite 
with a large, dense tuft of hairs directed inward. Basal lobe with 3 spines, 
two of them close together on a tubercle. Claspettes with angularly bent 
stem and broad wing. 

Fourth- stage larva (Figures 155— 157) large, light to dark brown; 
the very long antennae distinguish the larvae of this species and are visible 
with the naked eye. 




FIGURE 157. Aedes diantaeus H.D.K. Fourth-stage 
larva. Stigmal plate. 



Frontal hairs secondarily feathered, long: outer hairs with 3—6 (usually 
4) branches, median hairs with 2—5 (usually 3—4) branches, situated before 
inner hairs with 2—6 (usually 3—4) branches; postclypeal hairs situated 
between the median hairs, short, very thin. Antennae 1.1—1.3 times as long 
as the head, nearly straight, thin, with distinct spines and a tuft with 3—7 
(usually 4—5) secondarily feathered branches situated near the middle. 

Comb of 8th abdominal segment with 6—13 (11 — 12) large scales in 2 
irregular rows, scales with a long, pointed main spine and short, thin spines 
at the sides of the base. Siphon straight, tapering from the basal third; 
pecten with 15 — 22 (usually 16—18) long, spinelike denticles, which are 
shorter toward the base and with 1—2 distal, more widely spaced denticles; 
tuft situated in the middle, slightly basal to it, with 6—10 (usually 7—8) 
slightly secondarily feathered branches, as long as the width of the siphon 
at the base. 



260 



Saddle extending to lower margin of sides of last segment; lateral hair 
simple, shorter than the saddle; outer caudal hairs longer than the siphon, 
simple, inner hairs nearly half as long, with 10—13 branches. Fin with 
21 — 22 tufts on the common base and 2—4 tufts before the base. Gills narrow, 
lanceolate, slightly longer than the saddle. 

Distribution. Western Europe to the Maritime and Khabarovsk 
territories and from Norway and Karelia in the north to the Southern 
Ukraine and Northern Caucasus. It also occurs in part of North America. 

Biology. The larvae develop in different types of temporary water 
bodies in forests, formed by snowmelt. In pits, ditches, puddles, etc., 
shaded or in open localities, with cold water. Hibernation in the egg stage; 
usually one generation per year. A late spring species with delayed larval 
development; it is characteristic for mixed forests but also occurs in the 
250 taiga; often in thinned forests in glades and at the edges. Flight at the end 
of spring, later than many other species of Aedes. They are active blood- 
suckers but usually appear in small or moderate numbers. They are the 
predominant species in some parts of the Carpathians. 



24. Aedes (Ochlerotatus) intrudens Dyar, 1919 

The species differs from the preceding species in the presence of a 
hypostigmal spot, i. e. a few scales situated below the anterior spiracle. 
Proboscis, wings, legs and abdomen covered with dark scales without a 
steel sheen as in A. diantaeus. 




FIGURE 158. Hypopygium of Aedes intrudens Dyar 



261 



Proboscis and palps with brown scales and with grayish white scales 
scattered on the palps. Mesonotum with golden bronze scales, lighter 
(whitish) scales at the lateral margin, sometimes with an indistinct longitudinal 
dark stripe or with two narrow stripes. Postcoxal spot absent, hypostigmal 
spot present. Scales on mesepimera not reaching lower margin of sclerite. 
Fore femora with dark scales, with more or less white scales, mainly 
anteriorly. Tarsi dark. Wings with dark scales, sometimes a few light 
scales at the base of the costa. Abdomen dark dorsally, with white stripes 
at the base of the segments; stripes of uniform width, rarely narrower in 
the middle. 



(251) 




FIGURE 159. Aedes intrudens Dyar. Fourth-stage larva (after Carpenter and La Casse): 
1 — head, dorsal; 2 — posterior end, lateral. 



262 



Hypopygium (Figure 158) as in the preceding species, with 3 spines on 
the basal lobe and a dense tuft of hairs on the inner surface of the coxite. 
This tuft is smaller than in A., diantaeus and situated more distally, on 
the apical lobe, the hairs directed inward and posteriorly. Stem of claspette 
with a finger-shaped process in the middle, with a hair at the apex. 

Fourth-stage larva (Figure 159) medium-sized to large, yellowish 
brown to dark brown. Frontal hairs secondarily feathered: outer hairs with 
5-9 (usually 6-7) branches, median hairs with 3-4 (usually 3) branches, 
situated before the inner hairs with 3-5 (usually 4) branches; postclypeal 
hairs situated between the median hairs, short, with a few, thin branches. 
Antenna slightly more than half as long as the head, with sparse spines, hair 
tuft with 6-8 branches situated in the middle, slightly basal to it. 

Comb with 10-18 (usually 12-16) scales in two irregular rows, scales 
251 large, dark, with a long, pointed main spine and short, thin spines at the 
sides of the base. Siphon straight, distinctly tapering from the middle, 
index 2.8-3.5 (average about 3.0). Pecten with 14-22 (usually 18-19) 
denticles which are longer distally, 1-4 (usually 2-3) more widely spaced 
and forming larger, smooth spines while the others have 1—2 accessory 
denticles near the base; the most distal spine is situated distinctly beyond 
the middle of the siphon. Hair tuft with 4-10 (average 7-8) thin branches 
with weak secondary feathering, as long as the width of the siphon at its 
position. Hair tuft situated in the middle or distal to it; if there are 1—2 
widely spaced denticles, the tuft is situated distal to them, if there are more 
widely spaced denticles, the tuft is situated between them. 

Saddle of last segment extending to middle or lower margin of sides, of 
irregular form; lateral hair simple, short; outer caudal hairs simple, 
2 distinctly longer than the siphon; inner hairs usually with 6—8 branches, 
about 2 /z as long as the siphon. Fin with 16—18 tufts on the common base 
and 1—3 shorter tufts before the base; the branching of the tufts, especially 
of the largest, posterior tufts of the fin, begins far from the base, and the 
stem is distinctly longer than the lateral processes of the base. Gills 
longer than the saddle, markedly varying in length, pointed, not pigmented. 

Distribution. Western Europe to the Khabarovsk Territory and 
Kamchatka. Extending in the north to the forest-tundra, but apparently not 
to the tundra. It occurs in the steppes of the Ukraine. Northern and 
Central Europe, northern part of the U.S.A. and southern Canada. 

Biology. The larvae occur in shallow pools and swamps in forests 
formed by snowmelt or filtration during rise of the groundwater. They live 
in water without vegetation with a bottom covered with fallen leaves and 
are found together with larvae of A. excrucians, A. cata phy 11 a, 
A. communis, A. diantaeus and A. cinereus. The females attack 
man in large numbers mainly in forests, primarily alder forests. This is 
a forest species. It has one generation in spring which appears earlier 
than most other species. Their numbers decrease sharply in summer. 



2 5. Aedes (Ochlerotatus) pullatus Coquillett, 1904 

Closely related to A. intrudens, differing in details of coloration and 
in the structure of the hypopygium. 



(2 53) 




FIGURE 160. Hypopygium of Aedes pullatus Coq. 

Proboscis and palps with dark scales, sometimes with some light scales 
on the palps. Mesonotum with golden or whitish scales, with 2 dark narrow 
longitudinal stripes, with dark brown scales or without scales. Integument 
of mesonotum usually black. Hypostigmal spot present, postcoxal scales 
absent. Scales on mesepimera extending to lower margin of sclerite. 
Lower mesepimeral setae present (1 — 5). Legs: fore femora dark an- 
teriorly, tarsi with brown scales. Wing with dark scales, a few light scales 
at the base. Abdomen with dark scales dorsally, with light stripes at the 
base of the tergites, stripes sometimes narrower in the middle. 

Hypopygium (Figure 160): coxite with long, dense hairs, with 3 spines 
at the base as in the preceding two species, but a tuft of dense hairs on the 
coxite is absent; one of the two spines flattened, forming a narrow plate, 
sometimes with 2 delicate small branches at the end; stem of claspette 
angularly bent, without finger-shaped process. 

Fourth- stage larva (Figures 161 and 162) medium-sized, light 
brown or darker. Frontal hairs strongly branched, secondarily feathered; 
outer hairs with 8—13, median hairs with 3—8 (average 6), inner hairs with 
3—7 (average 5) branches; postclypeal hairs short, thin, with 4—5 branches. 
Antennae short, only slightly more than half as long as the head, with spines 
and a tuft with 5 secondarily feathered branches in the middle. 



264 



253' 




FIGURE 161. Aedes pullatus Coq. Fourth- stage larva. Head, dorsal (after 
Carpenter and La Casse). 



Comb with 50—60 scales of varying form: a main spine may be absent 
and there may be spines at the margin which are shorter at the base, or the 
main spine is more Or less distinct with shorter lateral spines. Siphon 
uniformly tapering from the basal third; index 3.0-3.5. Pecten with 15—25 
254 closely spaced denticles which are longer distally, and 2—3 rudimentary 
denticles, not reaching the middle of the siphon; hair tuft situated in the 
middle, distinctly beyond the distal denticle, with 5-8 secondarily feathered 
branches, longer than the width of the siphon at the position of the tuft. 

Saddle of last segment covering more than half of the sides; lateral hair 
simple, slightly more than half as long as the saddle; outer caudal hairs 
simple, long, inner hairs forming a symmetrical fan with 6—10 smooth 
branches. Fin usually with 15 tufts on the common base and 1—3 shorter 
tUfts before the base. Gills pigmented, lanceolate, slightly curved; upper 
pair slightly longer than lower pair, 1.5—2 times as long as the saddle or 
longer, sometimes as long as the siphon. 

Distribution. The distribution is disjunct. In Europe and the 
south of the USSR, it occurs in the mountains, outside mountain areas 
in the north. Pyrenees, Alps, Black Forest, Harz, Tatra, Carpathians, 
Caucasus, mountains of Tien Shan, southeast Kazakhstan, Leningrad, Moscow 
and Saratov regions, Kola Peninsula, Tuva region, Transbaikalia, Maritime 
Territory. The collections at the Zoological Institute contain specimens 
of A. pullatus with the label: western foothills of Otkhon- Tengri, 2, 300 m, 
Kozlov, July 1924. Widespread also in North America: Alaska, Canada, 
U.S.A., Rocky Mountains to altitudes of 3, 000-4,000m. 

Biology. The species occurs in the north in lowlands and plains: it 
is restricted to mountains of 2,000 m and higher in the south. The larvae 
occur in different types of small water bodies in spring, shaded (in the south) 



255 



265 



or open (in the north and in mountains), without vegetation. The species 
occurs in the mountains in small lakes with sandy or rocky bottom. In the 
Khibiny Mountains it occurs in turf waters and marshes along rivers with 
silt or stony bottom or with a bottom covered with fallen leaves of dwarf 
birch. One generation per year from hibernating eggs, later than in other 
species of Aedes; mainly a summer species. They are usually found in 
small numbers, but they appear in large numbers on the Kola Peninsula 
and in the Khibiny Mountains (Fridolin, 1936). 






(254) 




FIGURE 162. Aedes pullatusCoq. Fourth-stage larva. Posterior end, lateral (after Carpenter and 
La Casse). 



26. Aedes (Ochlerotatus) nigripes Zetterstedt, 
1838 (alpinus Edwards, 1921) 

Dark, medium- sized mosquitoes. The species differs from the other 
species of the communis group, except A. impiger, in the hairs on 
the thorax and in the position of the proepimeral setae. 



266 



Proboscis and palps with dark brown scales. Palps of male distinctly- 
shorter than the proboscis, terminal segment of palps not thickened. Meso- 
notum densely covered with long, brownish black setae. Integument of 
mesonotum black, scales brown or reddish brown, with a golden sheen 
laterally. Pleurae of thorax with creamy white scales. Postcoxal spot 
present, hypostigmal spot absent. Setae on proepimera irregularly 
scattered in the posterior half and at the dorsal margin. Scales on mes- 
episterna extending to anterior margin of sclerite. Femora variegated 
anteriorly with white and brown scales, sometimes mainly dark, with a few 
light scales. Claws long, slightly curved, with a short denticle which is 
less than half as long as the claw. Wings with dark scales, light scales 
present only at the base. Abdomen with dark scales dorsally (often with a 
slight reddish tone), with white stripes at the base of the tergites; stripes, 
especially on the posterior segments, sometimes indistinct, with diffuse 
margin. 



(256) 




FIGURE 163. Hypopygium of Aedes nigripes Zett. 



267 



(257) 




FIGURE 164. Aedes nigripes Zett. Fourth-stage larva (after Carpenter and 
La Casse): 

1 — anterior end, dorsal; 2 — posterior end, lateral. 



268 



Hypopygium (Figure 163): coxite with dense, long hairs; ends of hairs 
of left and right coxite usually slightly overlapping. Apical lobe of coxite 
weakly developed; basal lobe with setae but without distinct spine or a 
distinct strong seta; marginal seta usually larger than the others. Stem 
of claspette curved, rarely straight. Wing of claspette with weakly marked 
plate-shaped widening in the middle third. Processes of tergite 9 large, 
with 14—20 large, short setae. Phallosome strongly sclerotized, dark, 
bifurcate. 

Fourth- stage larva (Figure 164) medium-sized to large, dark or 
yellowish brown, with moderately broad head. Frontal hairs with slight 
secondary feathering; outer hairs usually with 5 branches, median hairs 
and inner hairs situated behind them, simple; postclypeal hairs with 3—5 
thin, short branches. Antennae short, less than half as long as the head; 
hair tuft with 1 — 3 short branches situated nearer to the apex. Mouth parts 
of the mixed type, substrate feeding predominating. 

Comb with 10—20 pointed scales, with very small spines, at the base. 
Siphon (index 2.5-3) straight, slightly tapering. Pecten with 12—17 denticles, 
256 the 1-4 distal denticles in the form of longer, widely spaced, smooth spines. 
Hair tuft with 2—6 thin, short branches about as long as the width of the 
siphon at the base. 

Last segment with ring-shaped saddle surrounding it completely or with 
a narrow slit in the middle of the ventral side. Outer caudal hairs long, 
simple, inner hairs usually with 5—6 branches; lateral hair with 1—2 
branches. Fin with 16—18 tufts on the common base. Gills 1.5—2.5 times 
as long as the saddle, lanceolate, both pairs of the same length. 

Distribution. A circumpolar species characteristic for the tundra. 
Alaska, Canada (including Arctic islands), Greenland, northern part of 
Scandinavia and Finland. In the USSR, it occurs on the Kola Peninsula, in 
the Arkhangelsk Region, Nenets National District, Kara River basin, Noril'sk, 
Obdorsk, Srednekolym.sk, Khatanga basin, mouth of the Lena, Kamchatka and 
the Komandorski Islands. 

Biology. An early spring species developing in pools and other 
temporary water bodies formed by snowmelt, in the tundra in marshes over- 
grown with grass and with dwarf willow and birch. They are mass blood- 
suckers in some parts of the tundra. Their ecology was studied on 
Ellesmere Island (west of Greenland) at 81°49'N (Corbet, 1965). The 
larvae appeared in early June in water bodies formed by thawing snow. 
258 Flight begins in 3-4 weeks. The mosquitoes feed at first on the nectar of 
flowers, particularly Dryas i n t e g r i f o 1 i a. Copulation takes place 
during swarming. The females may suck blood on the second day after 
taking flight. Autogenous deposition of eggs has been observed. The eggs 
are deposited at the edge of the water, 5—10 cm above the surface. 



27. Aedes (Ochlerotatus) impiger Walker, 1848 
(nearcticus Dyar, 1919; parvulus Edwards, 1921) 

A. nearcticus Dyar and A. parvulus Edw. were considered as 
different species in the past, distinguished mainly by the structure of the 
hypopygium, the presence (A. nearcticus) or absence (A. parvulus, 



269 



of a spine on the basal lobe of the coxite. However, this character varies 
markedly. In some specimens the basal lobe bears a few, nearly equal 
setae, in others one seta is slightly larger than the others and in still others 
there is a spine which differs distinctly from the thinner setae. A . ne- 
ar c t i c u s and A. parvulus should therefore be considered as the same 
species (Natvig, 1948) and both names are synonyms of A. impiger Walk. 
A study of the types of Culex i m pi g e r Walk, and A e d e s nearcticus 
Dyar showed that they are identical (Vockeroth, 1952). 




FIGURE 165. Hypopygium of Aedes impiger Walk. 



259 



Aedes impiger, like A. nigripes, differs from the other species 
of the communis group in the thoracic setae and arrangement of the 
proepimeral setae. The two species differ in the structure of the hypo- 
pygium, the form of the claws, arrangement of the scales on the mese- 
pisterna and in size - A. impiger is smaller. 



270 



Proboscis and palps with dark scales. Palps of male as long as or 
slightly shorter than the proboscis. Mesonotum densely covered with long, 
black setae. Integument of mesonotum black, scales reddish brown, some- 
times with lighter, yellowish brown scales, forming small spots. Setae on 
the proepimera scattered in posterior half of sclerite. Scales on mese- 
pisterna not reaching their anterior margin. Postcoxal spot present, 
hypostigmal spot absent. 

Femora variegated anteriorly or mainly with dark scales and some light 
scales . Claws more strongly curved than in A. nigripes, with a long 
denticle (more than half as long as the claw). Wing with dark scales and 
with a few light scales at the base. Abdomen dark dorsally, with light 
stripes at the base of the tergites. 

Hypopygium (Figure 165): coxite with moderately long hairs; ends of 
hairs of left and right coxite usually not overlapping. Apical lobe weakly 
developed; basal lobe with setae, the marginal seta curved at the end and 
distinctly thicker and longer. Claspette: stem curved from the middle, 
wing with distinct plate- shaped widening to the second third. Processes 
of tergite 9 small, with 4—10 setae. Phallosome weakly sclerotized, with 2 
denticles at the end. 




FIGURE 166. Aedes impiger Walk. Fourth-stage larva. 
Head, dorsal (after Carpenter and La Casse). 



Fourth-stage larva (Figures 166 and 167) medium- sized, light to 
dark brown, with brown head, which is V4 wider than long. Frontal hairs: 
outer hairs with 2—4 secondarily feathered branches, median hairs smooth 
and simple, inner hairs simple, often secondarily feathered; postclypeal 
hairs thin, short, with 2—4 branches, situated between the median hairs. 
Antennae short, less than half as long as the head, with sparse spines; hair 
tuft with 2-4 secondarily feathered branches situated in the middle, slightly 
basal to it. 

Comb of 8th abdominal segment with 7—16 (average 13—14) scales in 2 
irregular rows, scales with a well developed, pointed main spine and small 
spines at the base. Siphon tapering from the basal third; index about 3.0. 



271 



Pecten with 10—20 (average 12—14) closely spaced denticles which are 
longer distally and occupy the basal third of the siphon. Hair tuft situated 
260 in the middle, slightly basal to it, with 2-6 (average 4) secondarily feathered 
branches which are longer than the width of the siphon at its position. 




FIGURE 167. Aedes impigerWalk. Fourth -stage larva. Posterior end, lateral (after Carpenter 
and La Casse). 



Saddle of last segment extending slightly beyond dorsal half of sides, 
lateral hair short, simple. Outer caudal hairs simple, lorig, inner hairs with 
3-4 branches. Fin with 14-16 tufts on the common base arid 1-3 short 
tufts before the base. Gills 1.5 times longer than the saddle, lanceolate, 
pointed, pigmented. 

Distribution. Tundra of both hemispheres. The species usually 
extends further to the north together with A. nigripes than other 
species of mosquitoes. They have been found on Ellesmere Island at 
82°30'N, which is the most northern locality where mosquitoes have been 
found. In the south, the species occurs in the forest zone, mainly in 
mountain regions. It is distributed in Alaska, Canada, Greenland, northern 
Scandinavia and Finland. In the USSR, it has been found on the Kola 
Peninsula, in the Arkhangelsk Region, Nenets National District, Taimyr, 
mouth of the Kolyma, and the Novosibirskie Islands. It usually occurs 
together with A. nigripes. A.impiger predominates in Noril *sk 
(70°N) (Vol'ftrub, 1963). 



272 



Biology. The larvae develop in water bodies formed by snowmelt in 
the tundra and forest-tundra. They are numerous (lower reaches of the 
Yenisei) and attack man and animals, especially reindeer. They appear 
earlier in the spring than the other species. 



261 28. Aedes (Ochlerotatus) cataphylla Dyar, 1916 
(rostochiensis Martini, 1920) 

The species differs from the preceding species of the communis 
group in the presence of light scales not only at the base but also on other 
parts of the wing. Also characteristic are the variegated scales on the 
femora. 




-\ /I T\^ T ;l|--H...-/ / N 



FIGURE 168. Hypopygium of Aedes c a ta phy 1 1 a Dyar (after Carpenter and 
La Casse). 

Lobes of tergite 9 of A. cataphylla Dyar (A) and A.leucomelas Mg. (B). 



273 



262 




Proboscis with uniformly black scales. Palps 
with dark scales and some light scales. Meso- 
notum with silvery or light gray scales laterally 
and with a broad, usually indistinct median stripe 
of reddish brown or dark brown scales. The 
entire mesonotum is sometimes covered with 
chestnut brown scales and with lighter scales at 
the margin. Postcoxal spot and hypostigmal spot 
present. The spots are connected by a stripe of 
scales with a round, white spot on the parastigma, 
a small sclerite between the anterior spiracle and 
the anterodorsal margin of the mesepisternum. 
Femora variegated anteriorly, with white and 
brown scales. Tibiae and tarsi with dark scales 
and some light scales. Wings with brown scales 
with a few whitish scales near the base and a few 
whitish scales on the costa, subcosta and ri. 
Abdomen with blackish brown scales dorsally, 
broad, white stripes of more or less the same 
width at the base of the tergites; the last tergites 
are sometimes covered mainly with light scales. 
Hypopygium (Figure 168) with long, thick hairs 
on the inner surface of the coxites; apex of hairs 
of left and right coxite overlapping. Basal lobe 
of coxite small, with one spine or a strong seta. 
Apical lobe strongly convex. Claspette with 
sharply curved stem, wing with a plate -shaped 
widening. Processes of tergite 9 narrow, with 4—13 (usually 6—8) short, 
strong spines directed posteriorly. 

Fourth- stage larva (Figure 169) medium-sized, dark. Frontal 
hairs: outer hairs with 3—6 secondarily feathered branches, median hairs 
situated before the inner hairs, both pairs smooth, simple; postclypeal 
hairs situated between the median hairs, with 2-3 short, thin branches. 
Antennae less than half as long as the head, straight, with moderately distinct 
spines and a hair tuft in the middle or slightly more apically, with 3—5 
branches, less than half as long as the antennae. 

Comb with 10 (rarely) to 28 scales (average 25) in 2—3 irregular rows, 
scales with a long, pointed main spine and smaller spines near the base. 
Siphon straight, tapering in apical half (index 2.5—3.0, rarely more); pecten 
with 11 — 21 closely spaced denticles and 2—4 larger, smooth pointed spines 
which are widely separated and extend to % of the length of the siphon from 
the base; hair tuft situated in the middle, slightly basal to it, where the pecten 
ends, with 3—5 branches. 

Saddle of last segment extending beyond middle of sides; lateral hair 
short, simple; outer caudal hairs simple, usually as long as the siphon, 
inner hairs half as long, with 5—8 branches. Fin with 18 tufts on the com- 
mon base and 1—2 shorter tufts before the base. Gills pigmented, 
lanceolate, pointed, of varying length, as long as the saddle or twice as long. 

Distribution. Europe, Kazakhstan, Siberia, Far East and part of 
North America. It extends in the south to the Ukrainian steppe, Northern 



FIGURE 169. Aedes cat a- 
phylla Dyar. Fourth-stage 
larva. Siphon. 



274 



Caucasus and Kirghizia (mountains). It occurs in the north in the tundra 
according to the literature. The northern boundary of distribution needs 
more exact determination. In Scandinavia it occurs at 64°N, i. e. it does 
not reach the northern forest boundary (Natvig, 1948). It is found at 65°N 
in Alaska (Gjullin et al., 1961). 

Biology. Hibernation in the egg stage. The larvae occur in puddles 
formed by snowmelt and spring floods, mainly in open water or at the banks. 
They are very common in the forest zone and attack in large numbers but 
they have not been reported as a predominant species. They are more 
characteristic for open habitats with a thin forest cover. One generation 
per year. They appear in spring a few days earlier than the other species 
or together with A. communis and A. punctor. Flight ends in mid- 
summer. 



29. Aedes (Ochlerotatus) leucomelas Meigen, 1804 
(salinellus Edwards, 1921) 

This species is closely related to A. cataphylla in coloration and the 
structure of the hypopygium, but differs in the presence of light scales on 
the proboscis and the numerous light scales on the wings. 

Proboscis with dark scales, with more or less numerous yellowish brown 
scales which usually predominate in the middle of the proboscis. Meso- 
notum with bronze brown scales, without dark longitudinal stripe, scales at 
the sides lighter, golden or grayish white. Wings with dark scales, with 
many light scales which are more numerous on the costa, subcosta and 
radius. Pleurae of thorax and legs as in A. cataphylla. Abdomen with 
broad white transverse stripes on the tergites, which are usually distinct. 
Some specimens have diffuse margins of the stripes and isolated light scales 
in the posterior half of the tergites (sometimes numerous on the posterior 
tergites). Such specimens resemble A. detritus to some extent. 




FIGURE 170. Base of tufts of fin of 4th-stage larvae (after Fedorov): 
1— Aedes leucomelas Mg.; 2 — Aedes caspius Pall. 

Fourth- stage larva (Figure 170) medium- sized and of varying 
shades of brown. Frontal hairs secondarily feathered: outer hairs with 
3—6 branches median, hairs situated before the inner hairs; both pairs 



275 



simple, inner hairs rarely 2-branched; postclypeal hairs situated between 
the median hairs, thin, short, with 4 branches. Antenna nearly half as long 
as the head, nearly straight, with weakly developed spines and a hair tuft 
with 3—6 branches in the middle which are half as long as the antenna. 

Comb on 8th segment with 18—29 (average 24) scales in 2—3 irregular 
rows, of markedly varying form: most scales near the dorsal side are 
shorter, without a main spine, and bear 2—3 longer spines and some shorter 
spines at the sides; the scales near the ventral side have a distinct main 
spine and smaller spines at the base. Siphon straight, tapering from the 
second third, index 2.5—3 (average 2.6). Pecten occupying basal third of 
siphon, with 15—24 (usually 20) closely spaced denticles and several rudi- 
mentary denticles at the base. Hair tuft situated in middle of siphon 
slightly basal to it, with 3—8 branches which are secondarily feathered. 
264 Saddle extending to middle of sides of last segment; lateral hair simple, 

as long as the saddle; outer caudal hairs simple, long, inner hairs with 
5—9 branches, half as long. Fin with 15—18 tufts on the common base and 
1—3 shorter tufts before the base. The structure of the tufts is very 
characteristic: the branching begins far from the base and the long basal, 
single part of the tuft is 1.5—2 times as long as the lateral process of the 
common base. Gills as long as the saddle or shorter, leaf- shaped, the 
lower pair shorter than the upper. 

Distribution. The species occurs in Europe from southern Scandin- 
avia and Karelia, where it is distributed to 65°N (Lobkova, 1946) to Hungary, 
steppes of the Ukraine and the Northern Caucasus. It occurs in Kazakhstan, 
Siberia, Transbaikalia (Ulan-Ude) and Mongolia (Ulan- Bator ). The 
boundaries of its distribution, especially in Asia, need more exact 
determination. 

Biology. The larvae occur in open or partly shaded water bodies in 
spring, mainly in lowlands, on the edge of forests or in thinned parts, their 
bottom covered with fallen leaves, often near isolated trees. They are 
found in slightly saline water together with larvae of A. detritus, 
A. cataphylla and A. punctor. This is a widely distributed but not a 
mass species in the forest and forest-steppe. One generation per year. 
The mosquitoes appear in spring with the earliest species of Aedes. 
Flight ends in midsummer. 

30. Aedes (Ochlerotatus) detritus Haliday, 1833 
(salinus Ficalbi, 18 96) 

The species differs distinctly from the other species of the communis 
group, except A. s im a n i n i, in the coloration of the abdomen (Figure 171). 

Proboscis and palps with brown scales with more or less numerous 
light scales. Mesonotum with yellowish brown and white scales. Integu- 
ment of thorax brown. Pleurae of thorax with broad, white scales. Post- 
coxal spot absent, hypostigmal spot present. Femora variegated anteriorly; 
tibiae and tarsi with dark scales and with more or less numerous light 
scales. Wings with broad, dark and light scales. Abdomen dorsally with 
indistinct light transverse stripes at the base of the segments. Posterior 
part of tergites with brown and yellowish white scales, dark scales pre- 
dominating on the anterior segments and light scales on the posterior segments . 



276 



(265) 




FIGURE 171. Aedes detritus Hal. 

Hypopygium (Figure 172): hairs of coxite thick, long; ends of hairs of 
left and right coxite slightly overlapping. Basal lobe with one spine (strong 
seta) and shorter setae. Stem of claspette short, curved, wing with a 
manubrium; a plate- shaped widening present in the middle and apical part 
of the wing. Phallosome short, ovoid, without a constriction. 

Fourth-stage larva (Figures 173 and 174) medium-sized, yellowish 
to brown; frontal hairs: outer hairs with 7—12 branches, median hairs with 
1—3 branches (usually simple), situated before the inner hairs with 2—3 



277 



(266) 



265 



branches; postclypeal hairs with 2—3 
branches, thin, short. Antenna about 0.6 of 
the length of the head, thin, slightly curved, 
with sparse spines, hair tuft situated in the 
middle, with 5—8 (average 6) branches, half 
as long as the antenna. 

Comb with a varying number of scales, 
25—60 or more, scales without a main spine, 
short, tapering from the base, with a row of 
spines at the margin, the apical spines 
longer. Siphon short, index 2.2—2.5, slightly 
tapering from the middle. Pecten with 
18—27 denticles, occupying half of the siphon, 
denticles regularly spaced, only the distal 
denticle sometimes more widely separated. 
Hair tuft situated in the middle or nearer 
to the base, with 6—10 (usually 8) branches. 
Stigmal plate differing in some characters 
of the structure of the "stirrup": it is not 
situated as usual in the middle of the plate 
between the spiracles, but displaced toward 
the posterior valves; the posterior arc is 
therefore absent and the posterior process 
has a very jagged posterior margin and thin, small branches; shoulders 
and processes of spiracles absent; anterior arc of "stirrup" strongly 
developed, strongly projecting into the space between the spiracles. 




FIGURE 172. Hypopygium of Aedes 
detritus Hal. 



266 




FIGURE 173. Aedes detritus Hal. Fourth-stage larva: 
1 - head, dorsal; 2 - stigmal plate. 



278 



(267) 




FIGURE 174. Aedes detritus Hal. Fourth-stage larva. Posterior end, 
lateral (after Marshall). 

Saddle of last segment extending to middle of sides, lateral hair simple, 
as long as the saddle; outer caudal hairs long, longer than the siphon, inner 
hairs forming a fan with 8—11 branches. Fin with 16—18 tufts on the com- 
mon base and 2—3 shorter tufts before the base. Gills very short, spherical. 

Distribution. Western and Eastern Europe; to England and southern 
Scandinavia in the north; North Africa, Southwest Asia. USSR: Middle Asia 
and Kazakhstan. There are also records from the Leningrad region, 
Ukraine, Lower Volga area and West Siberia. Some of these records may 
be incorrect. It has also been recorded from Kashgar (Sinkiang) and from 
northwest Mongolia (Natvig, 1948). 

Biology. The larvae occur in small, temporary, stagnant water bodies 
and pools in spring, mainly with markedly saline, rarely with fresh, water. 
In the south the water is predominantly saline, with 0.83 — 5.2% salts; often 
inmarshes onthe seashore. In Tadzhikistan and Kirghizia they are found in 
the mountains at 1,500— 2,000m above sea level and in water bodies formed 
by snowmelt. They occur sometimes together with A.caspius caspius, 
C. longiareolata and other species. Intemperate latitudes there are 
267 two or more generations per year after the summer rains; in southern 

latitudes there is one generation per year, possibly due to the early drying 
of the water bodies. An early spring species which hibernates in the egg 



279 



stage. The females attack man, sometimes in large numbers. The species 
is characteristic for areas near the sea and continental regions with 
abundant saline water bodies. In the USSR it is numerous in deserts and 
steppes in Turkmenia, Tadzhikistan and Uzbekistan. In a large part of its 
distribution (which has not been satisfactorily determined) it is considered 
as a relatively rare species. 

A. detritus is probably an intermediate host of the filaria D i - 
petalonema evansi, which causes a serious disease of camels 
(Orekhov, 1952). 



31. Aedes (Ochlerotatus) simanini Gutsevich, 1966 

Closely related to A. detritus but differing in its darker color, the 
presence of a postcoxal spot of scales and the structure of the hypopygium. 

Proboscis and palps with dark and light scales, light scales usually more 
numerous. Thoracic integument blackish brown. Mesonotum with yellow- 
ish bronze and whitish scales, the latter predominating laterally and before 
and on the scutellum. Pleurae of thorax with white scales except in the 
upper half of the proepimera, which is covered with straight, relatively 
broad, grayish scales. Postcoxal membrane densely covered with scales. 
268 Hypostigmal spot continuing posteriorly in a stripe. Legs: fore and mid- 
femora and tibiae variegated anteriorly; tarsi without light rings but with 
numerous white scales on all segments, white scales sometimes predominat- 
ing; articulations of tarsi dark. Wings variegated, with large, brown and 
white scales on all veins. Abdominal tergites with blackish brown scales, 
and with broad, white, transverse stripes at the base of the segments. A 
few white scales on the dark part which covers most of the tergites; the 
light scales sometimes form an indistinct stripe also at the posterior 
margin of some segments. 

Hypopygium (Figure 175): coxite with moderately long hairs, ends of 
hairs of left and right coxite not overlapping. Basal lobe of coxite with 
one large seta and smaller setae and hairs. Apical lobe present. Stem 
of claspette straight, wing with a plate- shaped widening, present only for a 
short distance near the base. Phallosome elongate -oval, with a slight 
constriction. 

Fourth- stage larva medium- sized, yellowish brown, with darker 
head and lighter siphon (after A. M. Dubitskii). 

Head nearly 1.5 times wider than long. Frontal hairs: outer hairs 
with 5 (3—8) branches, median and inner hairs simple (in 60% of specimens) 
or 2-branched (40%), all hairs with indistinct secondary feathering. Post- 
clypeal hairs situated between median frontal hairs, with 2, rarely 3—4, 
short, thin branches. Sutural and transsutural hairs longer than the 
postclypeal hairs, with 1 or 2 branches. Antennae short, about 0.4 times 
as long as the head, with sparse spines; hair tuft situated in the middle 
slightly basal to it, with 4—5 (3—6) branches, not reaching the end of the antenna. 

Median hairs of prothorax with slight secondary feathering: anterior 
and median hairs simple or 2-branched, posterior hair with 2—3 longer 
branches extending to posterior margin of eyes. 

Comb usually with 22 (16—36) scales in 2, rarely 3, rows, forming a 
spot projecting posteriorly, half-moon shaped. Scales with a distinct 



280 



main spine and a few spines at the sides of the base. Base of dorsal and 
ventral scales in the spot distinctly broader than the others. Hairs behind 
comb: dorsal hair with 5 (4—7), median hairs with 7 (6—9), ventral hair with 
5 (4—8) secondarily feathered branches; intermediate hair smooth and 
simple. Darker, sclerotized plates of irregular form around the base of the 
dorsal and median hairs. Siphon (index about 3.4) slightly tapering apically. 
Pecten usually with 13 (12—17) equally spaced denticles and 2—4 smaller 
denticles at the base, occupying about /§, at most /i, of the basal part of the 
siphon. Denticles moderately long, with wider base and usually with one 
larger and 2—3 smaller accessory denticles at the base. Hair tuft with 5 
(4—7) secondarily feathered branches, slightly longer than the width of the 
siphon at its position, situated in basal part of siphon but distinctly before 
the distal denticle of the pecten. 




FIGURE 175. Hypopygium of A edes simanini Guts: 

A — basal lobe of coxite; B — lobe of tergite 9; C — wing of claspette. High 
magnification. 



281 



Last segment 1.5 times longer than wide. Saddle extending to middle 
of sides of segment, beyond it only in its anterior half. Lateral hair simple, 
rarely 2-branched, at most half as long as the saddle. Caudal hairs: outer 
hairs simple, usually as long as the siphon, inner hairs half as long, with 
5 (4—8) branches. Fin with 1.7 (14—18) tufts on the common base and one, 
rarely 2, shorter tufts before the base. Gills short, leaf- shaped, at most 
U as long as the saddle. 

Biology. The larvae occur in temporary shallow water bodies, 
strongly polluted by organic matter, with a varying degree of mineraliza- 
tion, open and shaded, in the tugai zone and near inhabited areas. The 
larvae are very adaptable ecologically. They tolerate temperatures to 
-5, -7° for 3—4 days. Development continues at temperatures of 4—5°. 
An early spring species with one generation per year. The larvae are 
found at the end of March and the adults in early April. Flight and attacks 
at the end of April. 
269 The species was described (Gutsevich, 1966) from material collected by 

P.I. Simanin in Kokand (Uzbekistan) in March-April 1929. It was also 
found in Kazakhstan. Iliisk village, Alma-Ata Region, vicinity of Alma-Ata, 
1,800 m above sea level. It probably also occurs in Middle Asia but has 
not been distinguished from A. detritus. 



32. Aedes (Ochlerotatus) kasachstanicus Gutsevich, 1962 

The species is characterized by the white tarsi and upright scales on 
the hind tibiae. It cannot be placed in any of the groups of Palaearctic 
Ochlerotatus (Gutsevich, 1962). A . ka s a c h s t a ni c u s differs from 
the other species of Ochlerotatus in its shorter proboscis, which is 
about as long as the fore femora. 
2 70 Palps and proboscis of female with dark and light scales, the latter pre- 

dominating in the middle of the proboscis. Palps of male with light rings. 
Mesonotum with dark chestnut scales and some whitish scales which form 
an indistinct border anteriorly at the sides of the mesonotum and also cover 
the scutellum and the area before the scutellum. Pleurae of thorax with 
broad white scales. Postcoxal spot of scales present, hypostigmal spot 
absent. 

Femora and tibiae with light and dark scales anteriorly. On the hind 
tibiae, in addition to contiguous scales, are also upright, mainly dark scales; 
the upright scales on the tibiae are larger in the males. Tarsi light, dark 
only at apex of segments. Wings with relatively broad, white scales; white 
scales present on all veins, but brown scales predominate. On the abdomen, 
light scales predominate dorsally, dark scales are present mainly in the 
middle of the tergites. Cerci distinctly projecting. 

Characters like the broad scales on the pleurae of the thorax and the 
upright scales on the hind tibiae stress the resemblance of A. kasach- 
stanicus to species of the subgenus Mu ci du s, which occurs in the 
Ethiopian, Oriental and Australian regions. 

Hypopygium (Figure 176): coxite with a basal and an apical lobe, apical 
lobe moderately large, basal lobe large, projecting, densely covered with 
long, seta-like hairs in several rows. Distinct spines or large setae absent 



282 



on the basal lobe. Claspettes: stem straight, wing with a small, plate- 
shaped widening and with a hook- shaped denticle near the base. 



(269) 




FIGURE 176. Aedes kasachstanicus Guts: 

A - hypopygium; B - wing of claspette; C - area of cross veins of wing; D - part of hind tibia. 



Distribution. Many localities along the Hi River (Alma-Ata Region, 
Kazakhstan). 

Biology. Forests and shrub thickets in floodplains. It appears in May 
and attacks in large numbers in June. Flight ending in August. Apparently 
only one generation per year. Active bloodsuckers. 



283 



Fourth- stage larva medium-sized, dark brown with dark siphon 
and lighter head (after A. M. Dubitskii). 

Head 1.4 times wider than long. Frontal hairs: outer hairs usually with 
8 (6—11) branches, median hairs with 3 (2—4), inner hairs with 5 (4—7) 
branches with sparse secondary feathering hairs, postclypeal hairs situated 
between the median frontal hairs, with 3 (2—7) very short branches. Sutural 
hairs with 2 (1—3), transsutural hairs with 3 (2— 4) thin branches. Antennae 
short, about half as long as the head, covered with sparse, small spines and 
with a tuft with 7—8 (6—11) branches situated in the middle of the antenna, 
slightly nearer to the base. 

Inner median hairs of prothorax relatively short: anterior and median 
hairs with 1—2 branches, posterior hairs with 2— 3, rarely with 4, longer, 
sparsely secondarily feathered branches, not reaching to posterior margin 
of eyes. 

Comb usually with 31 (27—34) scales in 2—3 rows, forming a posteriorly 
projecting, half-moon-shaped spot. Scales with a distinct main spine and 
a row of thinner and shorter spines at the sides of the base. Hairs behind 
brush: dorsal hair with 4 (3—6), medianhair with 6 (5—10), ventral hair with 
5 (4—7) branches with distinct secondary feathering; intermediate hair 
smooth and simple. Siphon long, cylindrical in the basal third and tapering 
apically, index 3.9—4.0. Pecten usually with 20 (16—21) equally spaced 
denticles and with 2—6 small denticles at the base. Pecten occupying basal 
third of siphon. Each denticle with 2—4 accessory denticles at the slightly 
widened base. Hair tuft usually with 4 (3—5) secondarily feathered branches, 
slightly longer than the width of the siphon at its position, situated distal to 
the end of the pecten at a distance of the length of one denticle. 

Saddle of last segment twice as long as wide, extending beyond middle of 
sides of segment. Lateral hair simple, slightly shorter than the saddle. 
Caudal hairs: outer hairs simple, slightly longer than siphon, inner hairs 
half as long, with 9 (8-10) branches. Fin with 16 (12-18) tufts on the com- 
mon base and with 2—3 shorter tufts before the base. Gills short, about 
one-third as long as saddle, ovoid or leaf- shaped. 

Biology. Breeding places are only water bodies formed by flooding 
of the rivers in spring or thawing snow, mainly in the tugai* zone with 
abundant fallen leaves and slightly saline water. A late spring species; 
one generation per year. 

!7 0a) 

SUPPLEMENT 

While the manuscript was in press, new data on mosquitoes of the USSR 
have been obtained: two new species, A e d e s (Ochl.) montchadskyi 
Dub., 1968 and Aedes (Ochl.) stramineus Dub., 1970, have been 
described. Two species new for the fauna of the USSR have been found. 
They are Aedes (Ochl.) fitchii Felt and Young and A . (Ochl. ) 
rempeli Vocker. The authors are especially grateful to A. M. Dubitskii 
(Alma-Ata) and P. E. Polyakova (Novosibirsk) for providing material of 
larvae and adults of these species and data on their biology. 

* [Formation with forest shrubs and meadows in river valleys (Middle Asia).] 



610144 284 



Aedes (Ochlerotatus) montchadskyi Dubitzky, 1968 

Female. Proboscis and palps with dark and light scales; apex of 
proboscis dark. Scales of mesonotum mainly golden yellow; whitish 
scales at the margins of the mesonotum and also forming 2 small spots in 
its middle. Pleurae of thorax with white scales that extend to the anterior 
corner of the sternopleura and to the lower margin of the mesepimera. 
Hypostigmal spot present, postcoxal spot absent. Femora and tibiae mainly 
with dark and light scales anteriorly. Tarsi without light rings; anterior 
surface of first 2 or 3 segments of tarsi with dark scales, posterior surface 
with light scales; lateral surface of tarsi with light scales, mainly in basal 
part of segments, forming indistinct light rings; terminal segments of tarsi 
dark. Wings with dark scales, with numerous light scales on the costa and 
ri. Abdomen dark dorsally and with light stripes at the base of the tergites, 
which are wider laterally and occupy about /± of the length of the tergite; 
light scales on the dark background absent. 

Male. Coloration as in the female but the light abdominal stripes are 
wider, occupying about fa of the length of the tergite. Hypopygium: coxite 
with moderately long hairs; basal lobe large, with a curved spine, a row of 
setae and several rows of hairs; apical lobe moderately large. Stem of 
claspette slightly curved, wing narrow, with a small, plate- shaped widening. 
Processes of tergite 9 with 5—8 setae. 

Fourth-stage larva grayish brown, with distinct secondary feather- 
ing on the hairs of the body. 

Head slightly wider than long. Frontal hairs: outer hairs with 8—10 
branches, median hairs with 2—3, inner hairs with 3—4 branches; inner 
hairs longer than the median hairs. Postclypeal hairs short, with 2—3 
branches. Sutural hairs simple, transsutural hairs shorter, with 2 branches. 
Antennae at most half as long as the head, with sparse spines; hair tuft 
short, with 6 — 8 branches, situated in the middle, slightly nearer to the apex. 

Median hairs of prothorax: anterior hairs thinner and shorter, extending 
to middle of compound eyes, posterior hairs thick, distinctly longer; both 
pairs 3-branched from the base; intermediate hairs simple, as long as the 
anterior hairs; all with sparse secondary feathering. 

Comb usually with 27 (22—23) scales in 2—3 rows, forming a half-moon- 
shaped spot which is convex posteriorly; scales of varying size and form, 
the shorter scales with a row of spines at the distal end, the longer scales 
with a main spine and sparse, shorter spines at the sides of the base. 
Hairs behind the comb strongly developed: dorsal hair with 4—7 branches, 
median hair with 5—7, ventral hair with 3 branches; intermediate hairs 
smooth and simple. 

Siphon tapering toward the apex, index 3.3—3.5. Hair tuft situated in the 
middle, slightly nearer to the base, with 5—8 branches, 1.5—2 times longer 
than the width of the siphon at its position. Pecten with 10—20 thin, 
contiguous denticles with 2—5 accessory denticles at the base, the distal 
denticle not reaching to the hair tuft. 

Saddle distinctly longer than wide, nearly extending to ventral margin 
of segment. Lateral hair characteristic, with 3 (rarely 2 or 4) secondarily 
feathered branches, distinctly longer than the saddle. Outer caudal hairs 
simple, 1.5 times as long as the siphon, inner hairs with 8—10 secondarily 



285 



feathered branches, more than half as long as the outer hairs. Fin with 
17—18 tufts on the common base and one shorter tuft before the base. Gills 
small, oval, l /&— /e as long as the saddle. 

Distribution and biology. Southeast Kazakhstan. Floodplains 
of the upper and middle reaches of the Hi River. In small temporary ponds 
in spring, in tugai thickets or near them, in water with a varying degree of 
mineralization. A late spring species, one generation per year. 

Note on systematics. It belongs to the communis group; the 
arrangement of the light scales on the tarsi resembles that in species of 
the cantans group. The variegated coloration of the femora and tibiae 
and the white scales on the wing place this species near A. leucomelas, 
but it differs from it in the coloration of the thorax and structure of the 
hypopygium. 

270b 

Aedes (Ochlerotatus) stramineus Dubitzky, 1970 

Female. Proboscis mainly with white scales, apex dark. Palps with 
dark and light scales. Mesonotum without a stripe, with yellowish scales 
which are darker at the margin, lighter in the posterior third. Pleurae of 
thorax with white scales to the anterior corner of the sternopleura and to 
the lower margin of the mesepimera. Hypostigmal and postcoxal spots of 
scales present. Femora and tibiae with dark and light scales anteriorly, 
tarsi with indistinct light rings at the apex of one segment and base of the 
next segment. Wings mainly with light scales and more or less numerous 
dark scales on the costa and other veins. Wing membrane yellowish. 
Abdomen with uniformly yellowish white scales dor sally. 

Male. Coloration as in the female. Hypopygium: coxite with short 
hairs and a few, thick, long hairs at the apex; basal lobe of coxite convex, 
with two spines in the distal part, a larger, hook- shaped spine in the apical 
half, and a smaller, straight spine; the lobe also bears a row of setae which 
are nearly as thick as the spines and several rows of hairs; apical lobe 
weakly developed, indistinct. Stem of claspette straight, short; wing of 
claspette as long as the stem, moderately sclerotized, slightly wider in the 
distal half. Processes of tergite 9 with 6—8 short setae. Phallosome 
broad, with a slight constriction. 

Fourth- stage larva medium-sized (to 1 cm), muddy brown to dark 
brown, with dark head and lighter siphon. 

Head slightly wider than long. A dark spot in the middle of the fronto- 
clypeus which is distinct against the lighter lateral sclerites. If the pig- 
mentation of the frontoclypens is less strong, it is wider anteriorly, then 
becomes narrower and widens again, attaining its greatest width between 
the inner frontal hairs; it then narrows again and becomes wider at the 
median frontal hairs. The pigmentation increases with age and its outline 
becomes less distinct because of the general darkening of the head capsule. 
Frontal hairs: outer hairs with 9—11 secondarily feathered branches, median 
hairs simple, inner hairs with 5, rarely with 3—6 branches. Postclypeal 
hairs situated between the median frontal hairs, short, with 3—4 branches. 
Sutural hairs short, simple, transsutural hairs much longer, with 2 branches. 
Antenna /$ as long as the head, with small spines; hair tuft situated in the 
middle, with 5—8 branches. 



286 



Inner median hairs on prothorax: anterior hairs shorter, reaching to 
posterior margin of eyes, with 3—4 branches, posterior hairs slightly longer, 
with 3—6 branches; both pairs branched from the base; intermediate hairs 
simple, as long as the posterior hairs. None of the hairs feathered. 

Comb of 8th segment usually with 30 (24—36) scales which form a tri- 
angular spot. Scales of varying size and form, shorter and wider with a 
row of spines at the distal end to narrower and longer, pointed but without 
a distinct main spine, with a row of spines which become shorter toward 
the base at the lateral margin. Hairs behind the comb: dorsal hair with 
7—9, median hair with 9—10, ventral hair with 6—7 secondarily feathered 
branches; intermediate hair smooth and simple. 

Siphon tapering from the middle, index 3.1—3.3. Hair tuft with 7—9 
secondarily feathered branches, at most 1.5 times longer than the width 
of the siphon at its position, situated beyond the middle, slightly distal to it. 
Pecten with 29—33 regularly spaced denticles, occupying the basal half of 
the siphon, not reaching to the tuft for the length of one denticle; denticles 
thin, long, with a row of weak denticles at the base. 

Saddle nearly twice as long as wide, reaching to the middle of the sides 
of the segment. Lateral hair simple, thin, slightly more than half as long 
270c as the saddle. Caudal hairs: outer hairs simple, nearly 1.5 times as long 
as the siphon, inner hairs with 12—14 branches, a third as long as the outer 
hairs. Fin with 14—16 tufts on the common base and 1—2 (rarely 3) shorter 
tufts before the base. Gills short, /s— /e as long as the saddle, oval, the 
lower pair slightly shorter than the upper. 

Distribution and biology. Described from Kazakhstan (Dubitskii, 
1970). Mosquitoes which probably belong to this species occur in many 
parts of Middle Asia, in Kazakhstan and West Siberia. It develops in 
spring in large, open, saline temporary ponds together with A. detritus 
and A . mo nt ch ad s ky i. One generation per year. 

Note on systematics. It is closely related to A. caspius from 
which it was not distinguished in the past. 



Aedes (Ochlerotatus) fitchii Felt and Young, 1904 

Large, dark mosquitoes. 

Female. Proboscis dark, palps mainly with dark scales and with a 
few light scales. Mesonotum with yellowish bronze scales and with a 
longitudinal dark brown stripe and small, indistinct, whitish spots. Pleurae 
of thorax with white scales which extend to the anterior corner of the 
sternopleura and do not reach the lower margin of the mesepimera. Hypo- 
stigmal spot of scales absent, postcoxal spot present. Femora and tibiae 
variegated anteriorly. Tarsi with white rings at the base of the segments; 
last two segments of fore tarsi and one segment of the mid-tarsi dark; 
rings on median segments of hind tarsi about half as long as the segment. 
Wings mainly with dark scales and more or less numerous light scales. 
Abdomen with light stripes dorsally which are broad at the base of the 
tergites and narrow at the apex; light scales predominate on the posterior 
tergites. 



287 



Male. Coloration as in the female. Hypopygium: coxite with relatively 
short hairs; basal lobe large, rounded or blunt- conical, slightly asym- 
metrical; lobe with one spine and numerous thick, dense hairs; apical lobe 
strongly convex. Stem of claspette slightly S- shaped or nearly straight; 
wing of claspette shorter than stem, slightly sclerotized, narrow, with a 
small incision at the base. Processes of tergite 9 with 7-9 long setae. 
Phallosome cylindrical, elongate, with a deep incision at the apex. 

Fourth-stage lar v a medium- sized. Head at least 1.5 times wider 
than long. Frontal hairs secondarily feathered, inner hairs with 3-4, 
rarely 2, branches, median hairs situated before the inner hairs, with 2-3 
shorter branches, outer hairs forming a tuft with many branches. Post- 
clypeal hairs situated between the median frontal hairs, short, with 3-4 
branches. Sutural and transsutural hairs simple. Antennae about half as 
long as the head, with spines, hair tuft situated near the middle, but not 
reaching the end of the antenna. 

Median hairs of prothorax secondarily feathered, inner hairs simple or 
with 2-3 branches, median hairs long and simple, outer hairs long, with 
2 branches or simple. 

Comb with 12-28 scales with a distinct main spine and a row of spines 
Vt- Va a s long as the main spine at the base. Hairs behind comb long, the 
median, the dorsal, and the ventral hair with 3-4 secondarily feathered 
branches, intermediate hair smooth and simple. 

Siphon (index 4-5) tapering from near the base, cylindrical in the apical 
fourth. Pecten with 15-24 regularly spaced denticles, not quite reaching 
middle of siphon; denticles longer apically. Tuft with 3-6 branches with 
slight secondary feathering, situated near the middle, beyond the distal 
denticle of the pecten. Hairs on posterior valves well developed, curved. 

Last segment long, saddle extending slightly beyond middle of sides. 
Lateral hair simple, smooth, as long as the saddle. Outer caudal hairs 
long, simple, inner hairs forming a fan with 6 branches. Fin with 1 or 2 
tufts before the common base. Gills pointed, lanceolate, 1.5-2 times as 
long as the saddle. 

Distribution and biology. It was found in the Magadan Region 
on the Anadyr' River and in the middle reaches of the Kolyma River at 
about 64°N (Polyakova, 1970). It appears in June, flies to the end of August 
and attacks people in large numbers. Widespread in the western hemisphere: 
Alaska, Canada, U.S.A. 

Note on systematics. It belongs to the c a nt a n s group. The 
species cannot always be determined with certainty by the females because 
of the resemblance to other species of the group, mainly A. riparius. 

270d 

Aedes (Ochlerotatus) rempeli Vockeroth, 1954 

The description of the adults is taken from the literature (Vockeroth, 
1954; Carpenter and La Casse, 1955) because we had no material. Only 
the larvae have been found in the Soviet Union to date. 

Female. Proboscis dark, palps with dark scales and with a few light 
scales. Mesonotum with grayish yellow scales and with a dark brown longi- 
tudinal stripe. Pleurae of thorax with whitish scales which reach the lower 
margin of the mesepimera. Hypostigmal and postcoxal spots absent. 



288 



Abdomen dark dor sally, with light stripes at the base of the tergites. 
Femora and tibiae dark anteriorly with a small admixture of light scales. 
Tarsi dark. Wings with dark scales, only a few light scales at the base of 
the costa. 

Male. Coloration as in the female. Hypopygium: coxite with long 
hairs, basal lobe large, conical, without a spine, with a few setae in a row in 
which they gradually become shorter; apical lobe small, with only a few 
short hairs. Stem of claspette curved, wing slightly shorter than the stem, 
with a widening in the basal third, curved and tapering at the apex. Phallo- 
some about as wide as long. 

Fourth -stage larva dark brown to black, medium- sized, with dark 
head and siphon. 

Head slightly wider than long. Frontal hairs with sparse secondary 
feathering: median hairs situated before the inner hairs, both pairs simple; 
outer hairs with 4—6 thinner branches with weak secondary feathering. 
Postclypeal hairs situated between the median frontal hairs, short, with 
4—5 branches. Sutural hairs thin, simple, transsutural hairs shorter, with 
2 branches. Antennae short, 0.5—0.6 times as long as the head, covered 
with spines; hair tuft with 5—7 thin branches, not reaching the end of the 
antenna, situated in the middle, slightly nearer to the base. Inner median 
hairs of prothorax smooth, the 2 anterior hairs simple and thinner, 
posterior hairs 2-branched from the base. 

Comb with 32—46 (average 38) scales forming a spot which is slightly 
curved posteriorly. Scales of varying size and form, long and narrow to 
broad and short, mainly at the ventral margin. Scales without main spine 
or rarely with a weak spine, their apical margin with a row of spines. 
Hairs behind the comb; dorsal hair with 2—4, ventral hair with 2—6, median 
hair with 4—7 secondarily feathered branches, intermediate hairs smooth 
and simple. 

Siphon tapering in the terminal third; index 3.5—4. Hair tuft with 8—12 
secondarily feathered branches, situated near the middle of the siphon, 
distinctly nearer to the base. Pecten with 20—23 adjacent denticles, occupy- 
ing only slightly more than the basal fourth of the siphon. Subapical spines 
well developed. Hairs on posterior valves of stigmal plate hook- shaped. 

Last segment short, slightly longer than high. Saddle surrounding the 
segment like a ring, saddle much longer dorsallythan ventrally. Lateral 
hair simple, distinctly longer than the saddle. Outer caudal hairs simple, 
smooth, 1.5 times as long as the siphon, inner hairs with 4—5 shorter 
branches. Fin with 14—16 tufts on the common base and 1—2 shorter tufts 
before the base; the unbranched base of each tuft is about 2—3 times longer 
than the process of the base, the length of the unbranched base increases 
from the anterior to the posterior tufts. Gills narrow-lanceolate, 1.7—2 
times as long as the saddle, weakly pigmented. 

Distribution and biology. The adults were described from Canada 
(Vockeroth, 1954). In the USSR, it occurs in Yakutia (Aksenova and 
Anufrieva, 1969), in parts of Siberia (Baraba Steppe, Kolyma basin (material 
of P. E. Polyakova and L. P. Kukharchuk), and also in the Altai in East 
Kazakhstan (Dubitskii and Tupitsin, 1970). The record of A . (O.) ni gripes 
from Saratov probably refers to A.(0.) rempeli. 



289 



Bibliography to the supplement 

Akse nova, A. S. and V. N. A nuf r i e va. Fauna i nekotorye voprosy 

biologii gnusa v raione almaznykh razrabotok v Yakutskoi ASSR 
(Fauna and Some Aspects of the Biology of Bloodsucking Flies in 
Diamond Mining in Yakutia).- Meditsinskaya parazitologiya i 
parazitarnye bolezni, 47, 1:8—16. 1969. 

Dubit skii, A. M. Novyi vid komara Aedes montchadskyi sp. n. 
iz basseina 111 (A New Species of Mosquito, Aedes mont- 
chadskyi sp. n., in the Basin of the Hi River). - Parazitologiya, 
2, 3:218-224. 1968. 

Dubitskii,A.M. Novyi vid komara Aedes stramineus sp. n. 
(Diptera, Culicidae) (A New Species of Mosquito, Aedes 
stramineus sp. n. (Diptera, Culicidae)). - Ibid., 4, 5:408-413. 
1970. 

Dubitskii,A.M. and Yu. N. Tu p i t s i n. Opisanie lichinok novogo dlya 
fauny SSSR komara Aedes (O.) rempeli Vockeroth (Des- 
cription of the Larvae of A e d e s (O.) rempeli Vockeroth, 
a New Species of Mosquito for the Fauna of the USSR).- Ibid., 
4,2:171-174. 1970. 

Polyakova.P.E. Novyi dlya fauny SSSR vid komara (Diptera, Culicidae) 
Aedes fitchii Felt and Young (A e d e s fitchii Felt and Youn| 
a New Species of Mosquito for the Fauna of the USSR (Diptera, 
Culicidae)). - Ibid., 4, 4:389-391. 1970. 



270 2. Subgenus Aedimorphus Theobald, 1930 
(Ecculex Felt) 

Closely related to the subgenus Ochlerotatus, but differing in the 
structure of the hypopygium; claspettes absent, basal lobe of coxite usually 
strongly developed, sometimes flask- shaped; rod- shaped appendage (if 
present) situated before apex of style; aedeagus divided into two serrated 
plates . The larvae do not differ from larvae of Ochlerotatus. 

The subgenus is distributed in the Ethiopian and Oriental regions and 
contains over 90 species. It is represented in the Palaearctic by two 
species, one widely distributed, the other, A. alboscutellatus Theob. , 
occurring only in the south of Japan. 



33. Aedes (Aedimorphus) vexans Meigen, 1830 

The narrow, light rings at the base of the tarsal segments (Figure 97) 
are characteristic. 



290 




FIGURE 177. Hvpopygium of Aedes vexans Mg. (after 
Carpenter and La Casse) 



291 



Proboscis and palps with brown scales with a few white scales. Meso- 
notum with reddish brown or bronze colored scales. A small area before 
the scutellum and the scutellum covered with golden scales. Femora and 
tibiae with dark scales anteriorly and with some scattered white scales. 
Tarsi dark, with white rings, which usually occupy not more than / 5 - / 4 of 
the length of the segment. This distinguishes it from species of the 
cantans group in which the rings are wider. The width of the light rings 
varies markedly. Specimens with relatively wider rings resemble 
A. cantans, but this species differs in the coloration of the mesonotum 
and abdomen and the presence of white scales on various veins of the wing. 
The wings of A. vexans are covered with brown scales, isolated light 
scales are present only at the base of the wing. The coloration of the 
abdomen varies in the two subspecies. 

Hypopygium very characteristic (Figure 177): basal lobe nearly divided 
from the coxite in the form of a flask- shaped process with wider end which 
is densely covered with hairs; style broad, plate- shaped, with a rod-shaped 
appendage, situated far from the apex of the style. 

The larvae of A. vexans are very variable because of the distribution 
of the species and different conditions of development and habitats of the 
adults, especially in the habitat characters, so that there are apparently 
ecological and geographical intraspecific differences. However, a compari- 
son of the descriptions of larvae from Europe, Asia, Africa and North 
America and examination of the material available in the USSR show that it 
is not possible to distinguish the larvae from different localities. 

Fourth-stage larva (Figure 178) medium- si zed (south) to large 
2 72 (north), yellowish in the south to dark brown, nearly black in the north. 
Frontal hairs secondarily feathered, outer hairs with 6—12 (usually 7—9), 
median hairs situated before the inner hairs, slightly more widely separated, 
with 1 — 2 (rarely 3) branches, inner hairs with 2—5 (usually 3—4) branches. 
Postclypeal hairs with 2—5 short, thin branches, situated between the median 
hairs. Antenna about half as long as the head, with moderately distinct 
spines which are more distinct in the northern forms, hair tuft with 5—12 
branches (half as long as the antenna), situated at lz of its length from the 
base. 

Comb with 7—13 (usually 8—10) scales with a pointed main spine and a 
row of thin spines at the sides of the base. Siphon tapering from the second 
third, its length very variable: index 2.5—4.3 (average 3.0— 3.5), less in 
southern forms. Pecten extending to middle of siphon or beyond it (in 
larger larvae), consisting of 12—25 (usually 18-20) denticles, 1—3 denticles 
at the base rudimentary and 1—3 distal denticles more widely separated and 
larger than the others. Hair tuft situated in the apical part, with 3—8 
(usually 4—6) short branches. 

Saddle on last segment extending to middle or lower margin of the sides; 
lateral hair simple, rarely with 2—3 branches; outer caudal hairs simple, 
longer than the siphon, inner hairs with 4—9 branches, half as long. Fin 
with 11—13 tufts on the common base and 3—4 shorter tufts before the base. 
Gills 1.5 times as long as the saddle or longer, lanceolate, not pigmented, 
both pairs of the same length or the lower pair slightly shorter. 

Distribution. Europe, Asia, Africa, North America. It extends in 
Europe to about 60— 62°N in river valleys and outside them. It is especially 



292 



widely distributed in Asia: Middle Asia, Kazakhstan, Siberia, Far East, 
Japan, China, Indo- China, India, Iran, Asia Minor. It is distributed from the 
Mediterranean coast of Africa to South Africa. In America, from the south 
of Canada to the Gulf of Mexico. 




FIGURE 178. Aedes vexans Mg. Fourth- stage larva. Posterior 
end, lateral (after Marshall). 

Biology. The larvae are restricted in the south to fresh or saline 
water bodies in floodplains, floodplains of deltas after the spring floods, 
with little vegetation. It occurs in the north in various water bodies in 
273 spring (puddles, ditches, pools) in open water or at the edge, but in fresh 
water. Its numbers are especially large in floodplains and river deltas. 
Flight begins in spring, slightly later than in most other species of Aedes. 
The spring generation is usually small. After the summer rains or summer 
floods of rivers from the south there is a second generation and flight may 
continue to the autumn. The period of mass activity lasts 2—4 months. 



293 



The seasonal numerical curve often has 2 or several peaks and the maxi- 
mum may occur in different months depending on the conditions in the 
rivers and the distribution of rainfall in summer. 

Mass breeding of A. vexans has been observed in many parts of the 
distribution, including the Maritime and Khabarovsk territories, lower 
reaches of the Danube, Dnieper, Don, Volga and Terek, some parts of Trans- 
caucasia, the Rhine valley and many parts of the U.S.A. The species often 
predominates among insects attacking people and animals. The mosquitoes 
may fly 10-20 km and further from their breeding places. They attack 
mainly in the open but also enter houses and cattle sheds in large numbers. 

Epidemiological importance. In foci of tularemia A . vexans 
may play a part in spreading the disease. Natural infections have been 
found and it also transmits tularemia experimentally. A virus of the group 
of lymphocytic choriomeningitis was isolated in Transcarpathia from 
A . vexans, and Tahyria virus in Slovakia. The species transmits eastern 
equine encephalitis in the U.S.A. The larvae of Di r o f i 1 ar i a immitis 
may develop in A. vexans to the infective stage. 



Aedes vexans vexans Meigen 

Abdomen with brown scales dorsally and white transverse stripes at the 
anterior margin of the tergites which are slightly narrower in the middle, 
the stripes resembling a horizontal letter B. Posterior margin of last 
tergites often with white scales. Specimens from Transcaucasia and Middle 
Asia usually differ in the light color of the scales and integument of the 
mesonotum. 

Description of the larvae on p. 291. 

Distribution. Europe, America, Africa, West Asia, Kazakhstan, 
Siberia. 



Aedes vexans nipponii Theobald, 190 7 

The subspecies differs in its slightly larger size and the numerous light 
scales on abdomen, proboscis, palps, femora and base of the wings. In 
addition to the broad transverse stripes, light scales form spots on the 
abdomen in the form of a longitudinal stripe (usually on the anterior seg- 
ments) or in the form of a triangle with the apex anteriorly, usually on the 
posterior segments. Light scales sometimes predominate on the posterior 
tergites. Light scales are present at the base of the wing but there are 
very few on other parts of the wing. 

The larvae do not differ from those of the nominate subspecies. 

Distribution. The subspecies is distributed mainly in the Far East: 
2 74 Maritime and Khabarovsk territories, Transbaikalia, Mongolia, China, Japan, 
also in Siberia and Kazakhstan. The two subspecies are connected by 
transitions. There may be "typical forms" and transitional forms in the 
same locality. 



294 



3. Subgenus F inlay a Theobald 

Differs from the other subgenera of Aedes mainly in the structure of 
the hypopygium. Claspettes present. Basal lobe of coxite absent or 
weakly developed, apical lobe absent. Appendage of style situated at the 
apex. Aedeagus simple, not divided. Palps of male usually slightly shorter 
than the proboscis. 

The posterior segments of the abdomen of the female are of character- 
istic structure: 8th sternite broad, not retracted into the preceding seg- 
ment; cerci short (in contrast to the two preceding subgenera). Proboscis 
distinctly longer than fore femora. Coloration: white spots at sides of 
abdomen. The light scales on the abdomen usually not white in species of 
Ochlerotatus. Proboscis and wings of Palaearctic species of F inlay a 
with dark scales. 

The larvae of Finlaya show no differences from those of the other 
Palaearctic subgenera of Aedes. The antennae are usually without spines, 
they are rarely weakly developed; hair on the antenna simple or 
2-branched. The hairs on the body are stellate or form strongly developed 
spines. Siphon short, with distinct auricles at the base. Denticles of pecten 
often forming a curved or slightly S- shaped row. Gills of different form; 
fin developed in inverse proportion to their size. 

The larvae of most species occur in small water bodies (tree holes, leaf- 
axils, holes in bamboo stems, rock pools, etc.). The biotopes of the species 
are specific, so that the distribution of most species is sporadic and often 
narrow. 

About 190 species of this subgenus are known, most of them from the 
Oriental region. Most of the species in the Palaearctic region are con- 
nected with the Oriental groups. Five species occur in the USSR, only one 
in Europe, the others only in the Far East. 

The species of Finlaya are usually not mass bloodsuckers but some 
transmit Japanese encephalitis and filariasis and a few species apparently 
transmit yellow fever (Aedes leucocelaenus Dyar and Shannon in 
South America). 



34. Aedes (Finlaya) geniculatus Olivier, 1791 
(ornatus Meigen, 1818) 

Large mosquitoes with a broad, dark longitudinal stripe on the mesonotum 
and white spots at the apex of the femora (Figure 179). Proboscis with 
dark scales, legs and abdomen with a violet tone. 

Proboscis and palps with black scales. Mesonotum with a broad, sharply 
defined longitudinal stripe of blackish brown or chocolate brown scales, or 
with 2 approximate dark stripes; two other dark stripes at the sides of 
the posterior half of the mesonotum. Lateral parts of mesonotum with 
silvery gray or cream-colored scales. Scutellum with narrow, yellowish 
scales. Pleurae of thorax with spots of broad, white scales. Tibiae and 
tarsi with black scales. At the apex of the femora is a small spot of white 
scales, distinct against the black tibiae and distal part of the femora. 
Claws of fore and mid-tarsi of females with one denticle, simple on the hind 



295 



tarsi. Wings with brown scales. Abdomen with black scales dorsally and 
triangular, white spots at the sides of the segments. 




FIGURE 179. Aedes geniculatus 01. 



296 



276 




FIGURE 180. Hypopygium of Aedes geni 
culatus Ol. (after Natvig) 




FIGURE 181. Aedes geniculatus Ol. Fourth- stage larva: 
1 — posterior end, lateral (after Marshall); 2 — stigmal plate. 



297 



Hypopygium (Figure 180): coxite densely covered with scales on the 
outside and with hairs; basal lobe forming a small tubercle with hairs and 
without spines or setae. Stem of claspette straight, longer than wing; 
hair on stem situated nearer to the base; wing of claspette narrow, curved. 
Lobes of tergite 9 with 3—6 setae. 

Fourth-stage larva (Figure 181) large, weakly pigmented, grayish 
or yellowish, with darker head and siphon; the marked stellate branching 
of the hairs gives the larvae a spinose appearance. Frontal hairs weakly 
developed: outer hairs short and thin, with 2—4 branches, median hairs 
displaced anteriorly, with 1—2 (usually) thicker branches, situated before 
the inner hairs, simple, but twice as long as the inner hairs; postclypeal 
hairs situated between the median hairs, with 7—8 short, thin branches 
forming a fan. Antenna thin, about half as long as the head, without spines, 
with a simple, rarely 2-branched, short hair. 

277 Comb on 8th segment with 11—15 scales in a posteriorly curved row, 
scales with a long main spine and with short, thin setae at the margin near 
the base. Siphon straight, short, index about 3.0. Pecten with 14—20 
denticles which are longer distally, closely spaced in a slightly curved row, 
extending beyond the basal third of the siphon; hair tuft with 4—5 branches 
situated distal to the pecten. 

Saddle of last segment extending beyond middle of sides, its posterior 
margin with small spines; lateral hair much longer than the saddle, with 
2—3 branches; outer caudal hairs simple, thicker, longer than the siphon, 
inner hairs usually with 3 branches, nearly half as long. Fin with 7—10 
tufts with 2—4 branches on the common base and 1—2 shorter tufts before 
the base. Gills sausage-shaped, slightly tapering at the ends, longer than 
the saddle, the lower pair shorter. 

278 Larvae from northern Iran (Figure 182) differ in their smaller size, 
longer and more strongly branched but thinner hairs on the body, more 
distinct secondary feathering in their basal part, longer tufts of the fin, 
and shorter gills. 

Distribution. Europe, to England and southern Scandinavia in the 
north (58°), North Africa, Southwest Asia to Iran. West and south of the 
USSR: Lithuania, Belorussia, Ukraine, Caucasus. Single specimens have 
been recorded from Turkmenia, Chuvash, Kuibyshev Region, Voronezh. 

Biology. The larvae develop in tree holes (beech, oak, walnut, poplar, 
ash) and in strongly shaded (tree holes) and well illuminated waters (tree 
stumps). They sometimes develop in pools, puddles, etc. together with 
larvae of A. plumbeus, A. pulchritarsis and Orthopodomyia. 
Hibernation in the egg stage in the north, as larvae in the south, so that 
adult mosquitoes may appear in warm winters. Large numbers are found 
in broadleaved forests with abundant summer rains and warm winters. 
They are rarely found in mixed forests and still more rarely in conifer 
forests, in forests in the Carpathians, Caucasus and Talysh they are mass 
bloodsuckers and the predominant species. Rare in other areas of the 
USSR. They attack mainly in the open, rarely entering inhabited areas. 
They appear later in spring than most other species of A e d e s, as they 
depend on summer rains. They are usually most numerous in midsummer. 

Transmission of yellow fever by A. geniculatus has been proved 
experimentally, but the species does not occur in foci of yellow fever. 



298 



(277) 




FIGURE 182. Aedes geniculatus 01. from Northern Iran. Fourth-stage larva. Posterior end, lateral. 



35. Aedes (Finlaya) echinus Edwards, 192 

Closely related to A. geniculatus, but differs in the coloration of the 
thorax and particularly in the coloration and form of the scales on the 
scutellum. 

Proboscis and palps with black scales. Mesonotum with 2 approximate 
chestnut-brown longitudinal stripes, distinctly bordered laterally by white 
scales. Scutellum with broad, white scales. Pleurae of thorax with spots 
of white scales, distinct against the dark background of the integument. 
Femora with a small white spot at the apex; tibiae and tarsi black. Wings 
with dark scales. Abdomen dark dorsally, with narrow, whitish stripes at 
the base of the tergites which widen laterally into triangular, white spots. 



299 



There are apparently no distinct differences from A. geniculatus 
in the structure of the hypopygium. However, the coxite is covered with 
very long, dense hairs in A. echinus. 



'279) 




FIGURE 183. First abdominal segment, dorsal (after Edwards): 
1— Aedes geniculatus Ol.; 2— Aedes echinus Edw. 



Fourth- stage larva. (Figures 183 and 184) distinguished by the 
unusually strong development of the thoracic and abdominal hairs, which 
are stellately branched from the base into thick spines with secondary 
feathering, so that the larvae have the appearance of a hedghog. The description 
is based on material from Bulgaria which was kindly provided by 
D. K. Bozhkov. 

Frontal hairs long, without secondary feathering: outer hairs with 2—4 
(usually 3) branches, median hairs with 1 — 2 branches, situated before the 
inner hairs, which are always simple; postclypeal hairs with 7—10 (usually 
279 7) thinner and shorter branches divided from the base. Antenna long 

(0.60—0.75 of the length of the head), distinctly curved, without spines, with 
a simple, rarely 2-branched hair situated at / 5 of the length of the antenna 
from the base; the hair is sometimes divided into 2—3 branches at the apex. 

Comb with 12—18 (usually 14) elongate scales in a slightly posteriorly 
curved row; scales with a strongly developed main spine and with small 
spines from the apex or from below it at the margin which are shorter 
basally. Siphon nearly cylindrical, index 3.0—3.5 or slightly more, with 
distinct auricles at the base. Pecten with 15— 27 (usually 18—22) closely 
spaced, long, spiniform denticles which are shorter toward the base and 2 — 4 
rudimentary denticles near the base; at the base of each denticle is a larger 
spine and several smaller spines; the pecten extends beyond the middle of 



300 



the siphon. Hair tuft situated distal to the pecten, with 2 — 4 (usually 3) 
branches which are distinctly longer than the width of the siphon. 



(280) 




FIGURE 184. Aedes echinus Edw. Fourth-stage larva (after Edwards): 
1 — head, dorsal; 2 — posterior end, lateral. 

Saddle of last segment nearly reaching lower margin of sides; lateral 
hair with 2 thick branches, longer than the saddle; outer caudal hairs simple, 
more than twice as long as the siphon, inner hairs with 3 (rarely 2) shorter 
branches. Fin with 9—13 tufts with 2—3 branches on the common base and 
1 — 2 tufts before the base. Gills strongly developed, upper pair 3—4 times 
and lower pair 1.5 times as long as the saddle. 

Distribution. Mediterranean: North Africa, Asia Minor, Southern 
Europe including the Balkans. It occurs in Bulgaria (Bozhkov, 1966) but 
not in the USSR. 

Biology. The biology of the larvae has been little studied, but it 
apparently resembles that of A. geniculatus. The larvae have been 
280 found in Anatolia in root holes of olive trees which fill with water during 
spring floods. They are numerous on the Black Sea coast of Bulgaria. 



301 



36. Aedes (Finlaya) nipponicus La Casse and 
Yamaguti, 1948 

Resembling the two preceding species, but differing in the absence of 
white rings on the tarsi. It differs from the other Palaearctic species with 
dark tarsi in the presence of white spots on the mesonotum. Previously 
known as A.niveus Ludlow, 1903. The new subspecies A.niveus 
nipponicus La Casse and Yamaguti, 1 948 was described from Japan and 
the authors considered it as a different species. It belongs to the geni- 
culatus group, but the subgroup niveus is represented by a large 
number of species in the Oriental region (Colless, 1958). The differences 
between the species are apparently very small. In particular, the characters 
distinguishing A. niveus and A. nipponicus (see below) do not justify 
considering A. nipponicus as a distinct species. 



281 




FIGURE 185. Hypopygium of A edes nipponicus La Casse 
and Yamag. (after La Casse and Yamaguti) 



302 



Proboscis and palps with brown scales. Mesonoturn with 2 large spots 
of silvery white scales divided in the middle by a dark longitudinal stripe 
which is wider posteriorly. Scutellum with light scales. Wings and tarsi 
with brown scales. Abdomen with brown scales dorsally and white spots 
at the sides of the segments, sometimes connected by narrow white stripes 
at the base of the tergites; the stripes are usually present on the posterior 
segments. Spermatheca single (La Casse and Yamaguti, 1955), which is an 
exception; there are usually 3 spermathecae in species of Aedes. 

Aedes nipponicus differs from A. niveus in that "the white spots 
of the mesonoturn are nearly always (but there are exceptions) bordered on 
the medial side by a concave line" (La Casse and Yamaguti, 1955). The 
coloration of the mesonoturn in specimens from the Maritime Territory is 
very variable. The dark, longitudinal stripe dividing the white spots usually 
extends to the anterior margin of the mesonoturn while the line separating 
the light and dark parts may be straight or (usually) slightly concave. The 
dark stripe often does not reach the anterior margin of the mesonoturn and 
the white spots are connected. Sometimes, especially in males, the entire 
anterior half of the mesonoturn is covered with silvery scales. 




FIGURE 186. Aedes nipponicus La Casse and Yamag. 
Fourth-stage larva. Head, dorsal. 



Hypopygium very characteristic (Figure 185). Not only is the outer 
surface of the coxite covered with scales, as in the other species of 
F inlay a, but there is also a group of large scales on the inner surface, 
Basal lobe weakly developed, with 2 short setae. At the base of the coxite 
is a tuft of dense, short hairs. Style with a long appendage about j± as long 
as the style. Lobes of tergite 9 situated close together, with several thin, 
long setae. 

Fourth-stage larva (Figures 186 and 187), as in all species living 
in tree holes, of varying size, greenish gray or grayish brown, with dark head and 
siphon. 



303 




FIGURE 187. Aedes nipponicus La Casse and Yamag. Fourth-stage larva. Posterior end, lateral. 



283 



Frontal hairs displaced toward anterior margin of head, secondarily 
feathered, central branches longer than lateral branches; the outer hairs 
are the longest, with 6—16 (usually 9—12) branches, median hairs with 7—12 
(usually 9—10) and inner hairs with 8—18 (usually 9-12) branches. Post- 
clypeal hairs strongly developed, resembling the median hairs, situated 
between them and with 6-12 (usually 9—11) branches. Antenna long (0.6— 
0.6.5 of the length of the head), thin, covered with small spines, hair tuft with 
5—11 (usually 6—9) branches, half as long as the antenna. 

Comb with 8—15 (usually 10-12) scales in one row, scales with a pointed 
main spine and with a few small setae at the margin at the base. Siphon 
slightly S- curved, index 2.5—3.8, usually about 3, with weakly developed 
auricles at the base. Pecten with 10—24 (usually 16—18) denticles and 1 — 5 
(usually 2) small denticles at the base, all denticles equally spaced, the 
distal denticle not larger than the others, denticles with a large accessory 
denticle and several small setae at the wide base. Hair tuft with 3—7 
(usually 4—6) secondarily feathered branches, stellate. 

Saddle of last segment extending beyond middle of sides, with dark 
anterior margin and with spines at the posterior margin; lateral hair 
with 1—5 (usually 2—4) branches; outer caudal hairs thick, simple, inner 
hairs with 2—5 (usually 3—4) branches, the ventral branch the longest. Fin 



304 



with 5—8 (usually 6) tufts with 2—3 branches and 1—2 shorter anterior tufts, 
the tufts not situated on a common base. Gills lanceolate, upper pair 
usually longer than the saddle, lower pair less than half as long. 

Distribution. Japan, Korea, North China. Only in the extreme south 
of the Maritime Territory in the USSR. 

Biology. The larvae develop almost always in tree holes and in 
stumps, in small water bodies, in stems of bamboo in the south. They are 
often found in holes of linden (Tilia amurensis) and maple (Acer 
mono and A. manshuricu m), rarely in birch (Betula ermani and 
B. dahurica), holes of Phellodendron, Manchurian walnut and Siberian 
stone pine. Larvae have been found in roots and tree holes to 8 m high 
together with larvae of A. galloisi, A. alektorovi and Toxo- 
rhynchites sp. Hibernation in the egg stage. The larvae appear in 
early June. The larval stages last 20—25 days, pupation 4 days. Flight 
of the first summer generation begins in early July. There may be 4 
generations per year. 

The females are bloodsuckers and attack man in large numbers. They 
fly from the -breeding place for a distance of 1 km. They are active through- 
out the day in forests. Their numbers markedly decrease toward autumn. 
They also bite inside buildings. 



37. Aedes (Finlaya) seoulensis Yamada, 1921 

The species is characterized by a white spot on the mesonotum and 
white rings on the tarsi. 

Proboscis and palps with black scales. Mesonotum with narrow, white 
scales in the anterior half, with dark brown scales in the posterior half. 
Posterior border of part with white scales curved. The area without scales 
before the scutellum is bordered by white scales. Scutellum with white 
scales. Tarsi with white rings on the first three segments, rings extending 
to apex of one segment and base of next. Rings on fore tarsi present only 
on the first two segments. Wings with brown scales, with a spot of light 
scales at the base of the costa. Abdomen with black scales with a steel 
sheen dorsally; narrow whitish stripes at anterior margin of tergites, white 
spots at the sides; light scales at posterior margin of tergites on the last 
segments. 

Hypopygium: coxite with scales on the outer surface. Appendage of 
style about /± as long as the style. Claspettes with short stem and narrow 
wing, wing slightly longer than the stem. Lobes of tergite 9 with 6—8 hairs. 

The fourth-stage larva has not been described in detail (Feng, 
1938). Frontal hairs not displaced toward the anterior margin, median 
hairs situated before the inner, all three pairs, especially the outer and 
median, long, with about 5 branches. Postclypeal hairs not shorter than 
the median hairs, branched, situated between the median and inner hairs 
but closer together. Sutural hair simple, long, not shorter than the frontal 
hairs. Antenna slightly more than half as long as the head, with sparse 
spines and a simple or 2-branched hair in the middle, slightly distal to it, 
its end reaching the apex. 



305 



Comb with 55—60 closely spaced scales with a main spine and with thin 
setae from the base to 2 j% of its length. Siphon straight, index 3.0. Pecten 
with 20—22 long, equally spaced denticles, the distal denticle situated beyond 
the middle of the siphon. Hair tuft with 5—6 branches situated in the apical 
part. 

Last segment very long, saddle reaching to middle of sides; lateral hair 
simple; outer caudal hairs simple, more than 1.5 times as long as the 
siphon, inner hairs with 6 shorter branches. Fin with 10 tufts on the com- 
mon base. Gills sausage-shaped, upper pair 1.3 times as long as the siphon, 
lower pair 0.75 times as long. 

Distribution. Korea, North and Northwest China. Not recorded 
from the USSR. 

Biology. The larvae usually develop in tree holes (Ulmu s pumila, 
Quercus acutissima and Soph or a j ap on i c a) during the summer. 



38. Aedes (Finlaya) alektorovi Stackelberg, 1943 

The species differs from the two preceding species in the absence of 
a white spot on the mesonotum; tarsi with narrow white rings but not on 
all segments (Stackelberg, 1943). 




FIGURE 188. Hypopygium of Aedes alektorovi Stack. 



Proboscis and palps with dark scales. Palps of male as long as the 
proboscis or slightly shorter. Mesonotum with scales of 2 colors: chest- 
nut brown scales provide the background for the golden scales, which form 



306 



stripes and spots. The golden scales sometimes predominate, especially 
in males. Tarsi of fore and mid-legs (or only fore tarsi) dark, except for 
285 a small white spot at the articulation of tibia and tarsus. Hind tarsi (some- 
times also mid-tarsi) with 2, 3 or 4 narrow white rings, which extend to 
the apex of one segment and the base of the next. Wings with dark scales. 
Abdomen dark, with white spots at the sides, sometimes connected by narrow, 
white stripes at the base of the tergites. Cerci hardly visible. 

Hypopygium (Figure 188): basal lobe of coxite forming a small tubercle 
with 2 large, curved setae and 1 (or 2) thinner seta. Appendage of style 
about l± as long as the style. Stem of claspette longer than the wing, slightly 
which is slightly curved in the middle, slightly wider in the apical half. 
Lobes of tergite 9 with 3—6 setae. 



(286) 




FIGURE 189. Aedes alektorovi Stack. Fourth- stage 
larva. Head, dorsal. 



Fourth-stage larva (Figures 189— 191) of varying size, depending 
on the size of the tree hole and its population, light brown with darker head 
and especially dark siphon (Monchadskii, 1949). Frontal hairs: outer hairs 
with 4—8 (usually 6) secondarily feathered branches, median hairs simple, 
smooth, very long, about 1.25 times as long as the head, inner hairs situated 
behind the median hairs, with 3—5 secondarily feathered branches, half as long 
as the median hairs. Postclypeal hairs situated before the median hairs 



307 



(286) 




<u 2 E 




ri <-> 




308 




FIGURE 192. Aedes togoi Theob. 



309 



closer together, with 10—12 short, thin branches. Antenna about half as 
long as the head, with a few spines, hair tuft with 1 — 2 branches situated in 
the middle. 

Hairs of body very long, especially the lateral abdominal hairs. Comb 
with 20—41 (usually 25—30) scales without a main spine, in 2—3 rows, with a 
row of spines which are shorter toward the base at the margin. Siphon thin, 
slightly curved, index 3.9—4.5. Pecten not reaching middle of siphon, with 
19—27 (usually 21 — 24) denticles and 1—3 rudimentary, closely spaced 
denticles near the base; denticles longer distally. Hair tuft with 3—4 
branches situated in the middle, as long as the width of the siphon. 

Saddle of last segment not reaching middle of sides; lateral hair short, 
with 2—4 (usually 3) thin branches; outer caudal hairs nearly 1.5 times as 
long as the siphon, simple, inner hairs with 2—3 shorter branches. Fin 
weakly developed, lateral processes of base not forming a common base, 
with 4—7 (usually 5—6) tufts, with lateral processes at the base and 3—4 
shorter tufts anteriorly. Gills 1.5 times as long as the siphon, sausage- 
shaped, slightly pointed, lower pair 15—25% shorter than upper pair. 

Distribution. It occurs in several localities in the Far East, south 
of Khabarovsk. 

Biology. A rare species characteristic for the broadleaved forests 
of the Maritime Territory. The larvae develop only in tree holes (Tilia 
amurensis; Acer mono, A.manshuricum and A. pseudo- 
si e b ol di a n um; Betula ermani and B.dahurica; Phello- 
dendron amurense, and others) together with larvae of A. galloisi, 
A.nipponicus and Toxorhynchites christophi. They usually 
develop in tree holes with narrow openings and choose the darkest places, 
avoiding roots in the open. Hibernation in the egg stage. Development in 
July lasts about 15 days. 



288 39. Aedes (Finlaya) togoi Theobald, 1907 

The species is characterized by white rings on the tarsi of all legs, the 
rings extending to two segments (Figure 192). 

Proboscis with dark scales. Palps of female with black scales and with 
white apex. Palps of male about /s as long as the proboscis, black, with 
white rings at the articulations. Mesonotum with blackish brown scales 
and weak, narrow longitudinal stripes of yellowish white scales in the mid- 
line along the dorsocentral setae at the lateral margins of the mesonotum. 
Scutellum and a small area before it with yellowish white scales. Tarsi 
black, with narrow white rings on the apex of one segment and base of the 
next. Wings with brown scales. Abdomen with black scales, with tri- 
angular white spots on the sides, connected by narrow stripes at the base 
of the tergites. 

Hypopygium (Figure 193) with a characteristic structure of the 
claspettes: the narrow wings are distinctly longer than the stem. Coxite 
with long, slightly flattened, longitudinally striated setae dorsally and on 
the inside. Numerous scales on the outer surface of the coxite, sometimes 
isolated scales also on the inner surface. Appendage of style short. Lobes 
of tergite 9 with 6—9 thin hairs. 



310 




FIGURE 193. Hypopygium of A e d e s togoi Theob. 
(after La Casse and Yamaguti) 



Fourth- stage larva (Figure 194) medium-sized to large, with 
weakly colored body, dark head and lighter siphon. 

Frontal hairs displaced toward anterior margin of head and arranged 
in a curved row which is convex anteriorly; outer hairs with 6—7, the longer 
median hairs with 7-8 and the inner hairs with 11-13 branches forming fans. 
Postclypeal hairs situated between the inner hairs, usually with 7 shorter 
branches. Antenna nearly straight, short, irregularly covered with sparse 
spines and with a short hair with 2—3 branches. 

Comb with 80—125 closely arranged scales of varying size, with rounded 
end and with a row of spines at the margin which are shorter near the base. 



311 



289 Siphon short (index 1.8—2.3), slightly barrel- shaped. Pecten extending 
distinctly beyond middle of siphon, with 18—25 closely spaced, spinelike 
denticles which are shorter near the base; at the lower margin of each 
denticle are 2—5 widely spaced accessory denticles. Hair tuft with 7—10 
secondarily feathered branches, forming a fan, situated near the apex, longer 
than the width of the siphon. 




FIGURE 194. Aedes togoi Theob. Fourth- stage larva: 
1 - head, dorsal; 2 - posterior end, lateral. 



Last segment as long as the siphon, with a small saddle, which almost 
does not reach the sides; lateral hair short, simple; outer caudal hairs 
simple, usually 2.5 times as long as the siphon, inner hairs half as long, 
forming a fan with 9—11 branches. Fin with 1—14 tufts with 7—12 branches, 
and 1 — 2 shorter tufts before the common base. Gills irregularly leaf- 
shaped, fa— fa as long as the last segment. 

Distribution. Coastal belt of China, Korea and Japan, including 
Hokkaido. It occurs in the USSR in the southern part of the Maritime 
Territory, Sakhalin and on the southern Kurile Islands. 

Biology. The larvae occur in large numbers in small pools on rocky 
coasts near the surf zone where the water is often very salty, with marked 
fluctuations because of rain, breakers or drying, which they tolerate. They 
are sometimes found together with larvae of C.vagans and A . h y r c a nu s. 
Several generations per year, from May to October. Secondary biotopes 
are stone basins, reservoirs or other artificial water bodies in inhabited 
areas, sometimes far from the coast, for example in Peking. 



312 



They are mass bloodsuckers in some localities, attacking in the open 
290 and in inhabited areas. In sparsely populated areas they apparently feed 
mainly on sea birds. In the Maritime Territory adult mosquitoes are 
found from June to the end of October, mass flight from the middle of July 
to early September. 

Epidemiological importance. This species is of great 
importance because it transmits Japanese encephalitis. Transmission of 
the virus was proved experimentally and natural infections were found. 
In China, A . t o g o i is an intermediate host of Brugia malayi (an 
important vector ) and Wuchereria bancrofti (vector of minor 
importance). 



40. Aedes (Finlaya) japonicus Theobald, 1901 

In contrast to A. togoi, A. japonicus has white rings only at the 
base of the tarsal segments. 

Proboscis and palps with black scales. Mesonotum with blackish brown 
scales, with 3 longitudinal stripes of light yellowish scales in the anterior 
half in the midline and along the dorsocentral setae; at the sides of the 
posterior half of the mesonotum are two curved, light stripes. Scutellum 
and area before it with yellowish white scales. Area below the anterior 
spiracle without scales. Tarsi with narrow, white rings at the base of the 
first two segments (fore tarsi) or first three segments (mid- and hind tarsi) 
Wings with dark scales. Abdomen with black scales dorsally, with small, 
white, triangular spots at the sides; usually narrow, transverse, white 
stripes at the base of the tergites which are usually not connected with the 
lateral spots. 

Hypopygium: coxite with scales on the outer surface and moderately 
long hairs. Stem of claspette longer than wing or as long as this. Lobes 
of tergite 9 small, flattened, with 4—9 thin hairs, sclerotized claws at apex 
of sternite 10 bifurcate. 

Fourth- stage larva medium- sized to large. Frontal hairs smooth, 
displaced toward anterior margin of head and arranged in a curved row 
which is convex anteriorly; outer hairs with 4—7, median hairs with 3—6, 
inner hairs with 4—7 branches. Postclypeal hairs very short, situated 
between the inner hairs, with 2—5 thin branches. Antenna less than half as 
long as the head, with sparse spines, hair tuft with 2—3 short branches 
situated beyond the middle. 

Comb with 43—85 closely spaced scales without a main spine, with a row 
of marginal spines which are longest at the rounded distal margin. Siphon 
straight, tapering from the tuft, index 2.3—3.8. Pecten with 14—29 denticles 
extending distinctly beyond middle of siphon, 1—4 of the distal denticles 
forming a more pointed angle to the longitudinal axis of the siphon in the 
form of larger, more widely spaced spines. Hair tuft with 4—7 secondarily 
feathered branches, as long as the width of the siphon, situated distal to its 
middle. 

Saddle of last segment covering the sides to the middle, its posterior 
margin with spines of different size; lateral hair with 1 — 2 branches, about 
as long as the last segment; outer caudal hairs more than 1.5 times as 



313 



long as the siphon, inner hairs with 2—4 branches about jz shorter. Fin 
291 with tufts with 2—5 branches on the common base. Gills lanceolate, usually 
1.5 times as long as the saddle. 

Distribution. China, Japan (Hokkaido). Not recorded from the USSR. 
Biology. This is a common and widespread species in Japan which 
occurs in mountains to an altitude of 1,300 m. The larvae occur in 
artificial water bodies with abundant decomposing organic matter in sun- 
light and in the shade. They occur in large numbers in spring and autumn. 
Their numbers remain the same from May to October in the south of Japan. 
They apparently rarely attack people. 




FIGURE 195. Hypopygium ofAedes koreicus Edw. (after 
La Casse and Yamaguti) 



314 



41. Aedes (Finlaya) koreicus Edwards, 1917 
(japonicus var. koreicus Edwards) 

Closely related to A. japonicus, and was previously not distinguished 
from it or regarded as a subspecies. In the Soviet literature on mosquitoes 
of the Maritime Territory and on vectors of Japanese encephalitis, it was 
named A. japonicus. 

Proboscis and palps with black scales. Mesonotum with dark scales 
and light stripes, as in A . japonicus, but stripes narrower . Area below 
anterior spiracle with a stripe of broad, white scales. Tarsi with narrow, 
white rings at the base of the segments. Rings on fore tarsi present on the 
first two segments, on the mid-tarsi on 3 or 4 segments, on the hind tarsi 
on 4 segments, last segment dark. Wings with dark scales. Abdomen dark 
dorsally, with narrow, transverse, light stripes at the base of the segments. 
Stripes not connected with the lateral white spots, which are usually not 
visible dorsally except on the 8th tergite. 

Hypopygium (Figure 195) resembling that of A. japonicus with the 
following differences: lobes of tergite 9 hemispherical; claws at apex of 
sternite 10 simple. 




FIGURE 196. Aedes koreicus Edw. Fourth-stage larva: 
1 — head, dorsal; 2 — ends of hairs of labrum. 

Fourth-stage larva (Figures 196 and 197) large, often darkly 
pigmented. Frontal hairs without secondary feathering, displaced toward 
anterior margin of head and arranged in an anteriorly curved row; outer 
hairs shorter, with 3-6 (usually 4-5) branches, median hairs with 2-6 
293 (usually 3-5), inner hairs with 4-6 (usually 5-6) branches. Postclypeal 
hairs short, with 2-4 thin branches from middle of stem. Antenna less 
than half as long as the head, irregularly covered with sparse spines and 
a short tuft with 1-4 (usually 2-3) short branches. 



315 



(292 




FIGURE 197. Aedes koreicus Edw. Fourth-stage larva. Posterior end, lateral. 

Comb with 49—70 (average 58) densely arranged scales of varying size 
and form, without a main spine, with a row of spines at the margin which 
are longer at the apex. Siphon distinctly tapering from the hair tuft, index 
2.8—3.4 (average 3.1). Pecten about half as long as the siphon or slightly 
longer, with 18—27 (often 20—25) densely arranged denticles, the distal 
denticles larger and forming a more pointed angle with the longitudinal axis 
of the siphon; the hair tuft is situated at the position of these denticles, 
forming a fan with 4—7 secondarily feathered branches, a third as long as 
the siphon. 

Saddle of last segment small, with irregularly jagged lateral margin, 
extending to dorsal third of sides and with small and larger spines at the 
posterior margin; lateral hair simple, rarely 2-branched; outer caudal 
hairs simple, 1.5 times as long as the siphon, inner hairs with 3—5 (usually 4) 
slightly shorter branches. Fin with 12, rarely 11, tufts on the common 
base and 1, rarely 2, tufts before the base. Gills lanceolate, narrow, usually 
1.75 times as long as the saddle. 

Distribution. Northeast China, Korea, Japan (Hokkaido). In the 
Maritime Territory in the USSR. 

Biology. The larvae occur in large numbers in rainwater barrels 
which become filled during the summer, always together with C. vagans. 
The species is very common in forests in the Southern Pvlaritime Territory 
and in inhabited areas or their vicinity. They bite in the open, sometimes 
in large numbers, but rarely enter houses. They bite people and farm 
animals and are often found in chicken coops. Adult mosquitoes are found 
in the Maritime Territory from June to September, mass flight from July 
to August. Probably two generations per year. 



610144 



316 



This species is apparently a potential vector of Japanese encephalitis. 
Experimental transmission of the virus has been reported. 



4. Subgenus Stegomyia Theobald, 1901 

Small, rarely medium- sized. Proboscis relatively thick and short, 
about as long as the fore femora. Palps of male slightly longer than the 
proboscis, with thin, usually upward curved apical segments, without hair 
tufts. Eighth sternite of female well developed, cerci short. 

Hypopygium: claspettes absent, basal (or subapical) lobe of coxite well 
developed, apical lobe absent, aedeagus divided into two plates, with small 
denticles at the apex in most species. 

Coloration characteristic: mesonotum with a pattern of light scales, 
tarsi with white rings at the base of the segments. Light spots on head, 
scutellum, pleurae of thorax and abdomen formed mainly by white scales 
with a silvery sheen. 

We give the characters of coloration common to the species of Stegomyia 
described below. Proboscis with dark scales. Palps of female dark in the 
basal half, white in the apical half; palps of male with white rings. Head 
with dark and light scales dorsally, usually forming a longitudinal stripe 
in the middle, round spots laterally and at the posterior margin of the eyes. 
Mesonotum with a light pattern which is characteristic for each species. 
Scutellum with 3 white spots. Tarsi with white rings at the base of the 
segments, which are more distinct on the hind legs. Wings with dark 
scales, sometimes with a small group of white scales at the base of the 
costa. Abdominal tergites with white stripes at the base and white spots 
at the sides. 

The frontal hairs of the larvae are not displaced toward the anterior 
margin of the head. Median hairs situated before the inner hairs. Antenna 
without spines, with a small, usually simple hair. Plate at base of lateral 
hairs of mesothorax and metathorax usually with a spine. Scales of comb 
few, usually arranged in one row. Siphon without auricles at the base. 

The subgenus contains over 90 species, mainly in the Ethiopian and 
Oriental regions. Some species reach the southwest Palaearctic (3 species) 
and the southeast Palaearctic (4 species). Four of these species occur 
(or occurred in the past) in the USSR. 

Some of these species are important vectors of virus infections, yellow 
fever, dengue and others. 



42. Aedes (Stegomyia) aegypti Linneus, 1762 
(argenteus Poiret, 1787; fasciatus Fabricius, 1805; 
calopus Meigen, 1818 (Figure 198) 

It differs from the other species of the subgenus in the pattern on the 
mesonotum (lyrelike) and structure of the hypopygium. 

Characters common to all species of Stegomyia are described above. 
Mesonotum with dark brown scales and with 2 distinct, lateral white stripes 



317 



curved in the anterior half and 2 less distinct, narrow, straight stripes 
near the midline. Apex of femora white, tibiae dark. Fore and mid-tarsi 
black with white rings at the base of the first two segments. Hind tarsi 
with white rings l fa — fa as long as the segment on the first 3 segments 
about fa as long on the 4th segment. Last segment of hind tarsi com- 
pletely white. The width of the white rings varies. Claws of fore and 
mid-tarsi of females with a denticle, claws of hind tarsi simple. Wings 
densely covered with brown scales on the veins. Abdomen with blackish- 
brown scales dorsally, usually with a reddish or steel sheen. Narrow white 
stripes in anterior part of the tergites, a silvery white spot at the sides. 



(295) 




FIGURE 198. Aedes aegypti L. 



318 



(296) 




FIGURE 199. Hypopygium of Aedes aegypti L. (after Carpenter and La Casse) 

Hypopygium (Figure 199): coxite short, thick, with an indistinct, flattened 
lobe in the middle of the inner surface, densely covered with short, seta- 
like hairs. Style markedly tapering apically, with a short appendage. 
Posterior margin of tergite 9 with a deep incision. 

Fourth- stage larva (Figures 200 and 201 ) very variable in size 
and coloration, according to the conditions. Frontal hairs long, simple; 
295 median hairs shorter, displaced anteriorly, inner hairs situated behind them, 
at the level of the outer hairs. Postclypeal hairs situated between the 
median hairs, with 4—6 short, thin branches. Antenna short, about half as 
long as the head, without spines and with a simple, short hair beyond the 
middle. 

Comb with 8—12 (usually 10, sometimes to 18) scales in a curved row; 
scales variable, often with a more or less distinct main spine and usually 
with shorter spines at the margin which are shorter near the base. Siphon 
short, tapering beyond the middle (index 1.5— 2.5), without auricles at the 
base. Pecten with 12—22 (usually 15—17) closely spaced denticles, the 



319 



296 distal denticle may be slightly more widely separated and situated beyond 
the middle of the siphon; denticles with broad base and a larger and 1—3 
smaller accessory denticles. Hair tuft situated near the distal denticle, 
with 2—5 short branches. 



(297) 




FIGURE 200. Aedes aegypti L. Fourth -stage larva (after Ma cfie): 

1 — posterior end, lateral; 2 - head, dorsal; 3 — 2nd abdominal segment, dorsal; 4 — base of lateral 
hairs of metathorax. 



Saddle of last segment reaching to lower margin of sides, lateral hair 
with 2—3 short branches; outer caudal hairs simple, twice as long as the 
siphon; inner hairs with 2—4 shorter branches. Fin with 8—10 tufts with 
2—3 branches. Gills sausage-shaped, with rounded end, slightly shorter 
than the siphon. 

Distribution. Tropics and subtropics of both hemispheres. Europe: 
France, Spain, Portugal, Italy, Yugoslavia (south of the Adriatic coast), 
Albania, Greece. Greater part of Africa. Asia: Southwest Asia, Iran 
(Shatt al-Arab), Pakistan, India, Southeast Asia, coastal parts of China (not 
in North and Northeast China), Japan (only in the far south). Western 
hemisphere: South, Central and North America, to Illinois and Indiana, i. e. 
2 97 nearly to 40°N. However, it has now been eradicated or has become rare 
in many countries where it was previously common. 



320 




This species used to be distributed on the 
Black Sea coast, from Sochi to Batumi, and 
often in localities far from the Black Sea, 
e. g. Kutaisi, Tbilisi, Baku in Transcaucasia. 
Because of the work instigated by 
N. P. Rukhadze, the incidence of A. aegypti 
was reduced, and it has not been found there 
in recent years. 

Biology. The larvae develop in 
artificial water bodies near and in inhabited 
areas and tolerate strongly polluted water. 
Development of the larvae at 2 7—30° lasts 
about 10 days and ceases at temperatures 

FIGURE 201. Aedes aegypti L. below 20°. 

Fourth-stage larva. Stigmal plate. The Qriginal habitat of ^ . a e g y p t i is 

tropical Africa, where it is still found in 
nature. Outside this area it occurs almost exclusively in inhabited areas 
and is more closely connected with man than any other species. It bites 
mainly in houses or near them. Copulation takes place in buildings. The 
298 mosquitoes copulate in small cages so that it is easy to breed them in the 
laboratory (at temperatures of 20—22°, optimum 25—30°). Several genera- 
tions per year in nature. 

The epidemiological importance of A. aegypti is very great . 
It is the most important or the only carrier of yellow fever outside natural 
foci of the disease and is therefore called the "yellow fever mosquito." 
Most urban foci of yellow fever have now been eradicated but if A. aegypti 
is present, outbreaks of the disease may occur transmitted from natural 
foci to inhabited areas. This species is also the main vector of dengue 
fever. Under experimental conditions and, possibly, under natural conditions 
it may transmit also other viruses infecting man, e.g. encephalitis and 
lymphocytic choriomeningitis. It also transmits the virus of myxomatosis 
of rabbits (mechanical transmission). The microfilariae of Brugia 
m a 1 a y i develop in A. aegypti, but microfilariae of Wuchereria 
bancrofti usually do not reach the infective stage. 

The literature on the biology of A. aegypti and its medical importance 
is enormous (see Horsfall, 1955). Christophers (i960) has written a large 
monograph on this species. 



43. Aedes (Stegomyia) cretinus Edwards, 1921 
(lindtropi Shingarev, 1927) 



299 



It differs from A. aegypti in the coloration of the mesonotum and the 
structure of the hypopygium. 

Mesonotum with blackish brown scales, with a longitudinal white stripe 
in the middle and 2 small, round, white spots laterally. Coloration and 
claws otherwise as in A. aegypti. 

Hypopygium (Figure 202): coxite oblong, the flattened process on the 
inner surface densely covered with hairs which are longer apically. Style 
narrower in the middle, with a thin, relatively long appendage situated 
slightly subapically. Posterior margin of tergite 9 with a curved process. 



321 



;298) 




FIGURE 202. Hypopygium of A e d e s cretinus Edw. 

The larvae have not been described in detail. They resemble those 
of A. aegypti but differ from them in the thinner antennae with a simple, 
thick hair in the middle; comb with 10 scales in one row, with a long, pointed 
spine and short, thin spines at the margin near the base; index of siphon 
about 2.0, pecten with 1 1 denticles which are narrower than in A. aegypti 
and with smaller denticles at the base; hair tuft with 3 branches, situated 
in the middle of the siphon and with a simple hair on the side in the apical 
third; gills at most % as long as the siphon, fin more strongly developed. 

Distribution. Crete, Macedonia. A few specimens have been found 
on the Black Sea coast of the Caucasus: Sukhumi, Gudauta, Sochi (Bash- 
kareva, 1931). The last record was by Gul'ripsh (near Sukhumi), who caught 
two females biting in a bamboo grove on 20 August 1939 (collection of 
T. Ya. Avdeeva). Two badly preserved specimens of Stegomyia from 
the southern coast of the Crimea probably belong to this species. 

Biology not studied. The larvae were found in tree holes together 
with larvae of A. plumbeus, O. pulchripalpis and A. geniculatus. 



44. Aedes (Stegomyia) vittatus Bigot, 1861 

It differs in the coloration of the mesonotum and in the presence of a 
white ring on the tibiae. 



322 




Head and its appendages as in 
^ . aegypti. Mesonotum with blackish 
brown scales and with 4 silvery white, 
round spots in the middle. Tibiae of 
all legs with a white ring in the middle. 
Tarsi with white rings at the base of 
the first 3 segments; rings on fore 
and mid-tarsi narrow, wide on the hind 
tarsi. Last 2 segments of hind tarsi 
white, sometimes dark at the apex. 
Hypopygium (Figure 203) with a 
very characteristic style, which is 
markedly widened apically and has a 
long, curved appendage in the middle 
of the inner side of the style. 
Fourth-stage larve 
(Figure 204): frontal hairs long, outer 
hairs with 3—6 branches, directed 
laterally, median hairs situated before 
the inner, both pairs simple. Post- 
clypeal hairs situated between the 
median hairs, small, branched. Antenna 
slighty more than half as long as the 
Bi g- head, with weakly developed spines and 

a small hair with 3 branches in the middle . 
Comb with 6—9 (usually 8) large scales with a long, pointed main spine 

and a few thin, short spines at the margin, near the base. Siphon short 

(index 2.0 or slightly more), slightly tapering, without auricles at the base. 

Pecten with 20—34 spinelike denticles, occupying about jz of the siphon; 

there is also a larger, smooth spine nearer to the apex. Between this spine 

and the most distal denticle of the pecten is a tuft with 3—6 branches as long 

as the width of the siphon at its position. 

Last segment longer than in other larvae of Stegomyia. Saddle weakly 

developed, extending only slightly to the sides; lateral hair short, simple; 

outer caudal hairs simple, long, inner hairs with 4—6 branches. Fin with 

5—7 two-branched tufts on the common base and 3—4 tufts before the base. 

Gills only slightly shorter than the siphon, wider at the base and pointed at 

the end. 

Distribution. Southern Europe, Africa, South and Southeast Asia . 

Not found in the USSR. 

Biology. The larvae occur in rock pools and also in artificial water 

bodies near inhabited areas. They sometimes attack people in large 

numbers. They are apparently vectors of yellow fever. Transmission of 

the virus of yellow fever has been proved experimentally. 



FIGURE 203. Hypopygium of Ae d e s vittatus 



45. Aedes (Stegomyia) galloisi Yamada, 1921 

It is characterized by the coloration of the thorax (mesonotum with a 
white longitudinal stripe and hind tarsi (last segment completely or partly 
dark) and by the structure of the hypopygium. 



323 




FIGURE 204. Aedes vittatusBig. Fourth-stage larva (after Barraud): 

1 - antenna; 2 - siphon, lateral; 3 - denticles of pecten; 4 - scales of comb. 

Mesonotum with chestnut brown scales, with a longitudinal light stripe 
which tapers slightly posteriorly and light lateral stripes which are curved 

301 in the anterior half of the mesonotum and straight in the posterior half; scales of 
the median and lateral stripes with a yellowish tone in the posterior half of the 
mesonotum. Tarsi of all legs with narrow, white rings on the first two 
segments. The next segments of the fore and mid-tarsi are dark; third 
segment of hind tarsi white in the basal half; 4th and 5th segments white, 
with dark apex. Claw of fore and mid-tarsi with a denticle, claws of hind 
tarsi simple. Abdomen with black scales and narrow white stripes at the 
base of the segments, and with white spots at the sides. 

In the specimens from Khabarovsk and the Maritime Territory, there 
are many variations in the coloration of the mid- and hind tarsi: last 
segment of hind tarsi completely dark, 2nd segment of mid-tarsi of female 
very variable, completely white, white with a dark apex, or dark with a white 
ring at the base (as in specimens from Japan); this segment is sometimes 
covered with white scales in the outside and dark scales on the inside. The 
coloration of the tarsi is apparently very variable. There may be two or 
several forms (subspecies?). 

Hypopygium (Figure 205): wart-shaped process on inner surface of 
coxite convex, transverse, densely covered with hairs at the outer margin. 
Style narrower in the middle, the rod- shaped appendage situated 
slightly subapically. Posterior margin of tergite 9 forming a curve which 
is convex posteriorly, sometimes with a triangular process in the middle. 

302 Fourth- stage larva (Figure 206) brown, with strongly developed, 
stellate hairs on the body. Their size varies according to the conditions 
(size of water body, density of population, food). 

Frontal hairs weakly developed, median hairs situated before inner 
hairs, displaced to the anterior margin, outer and median hairs with 2—3 



324 



(301) 




FIGURE 205. Hypopygium of Aedes galloisi Yam. 




FIGURE 206. Aedes galloisi Yam. Fourth-stage larva: 
1 - head, dorsal; 2 — posterior end, lateral. 



325 



branches, inner hairs simple. Postclypeal hairs situated between the 
median hairs, only slightly shorter than these, with 9—10 branches. Antenna 
short, about a third as long as the head, without spines, with a short simple 
hair in the middle. 

Comb with 8—12 (usually 9—10) scales in one row, with a large pointed 
main spine and short, thin setae at the sides of the base. Siphon without 
auricles, dark, distinctly tapering from the middle, index 2.9—3.1. Pecten 
with 10—16 small, thin, spine- shaped denticles, with broad base, and with 
1—3 basal accessory denticles; the distal denticle is situated before the 
middle of the siphon. Hair tuft situated in the middle, with 4—8 branches, 
not longer than the width of the siphon at the base. 

Saddle extending to the lower margin of the sides of the short last seg- 
ment, rarely surrounding it like a ring; lateral hair short, with 2—4 branches; 
outer caudal hairs simple, as long as the siphon, inner hairs with 2 branches, 
dorsal branch shorter. Fin weakly developed, with 6 simple (rarely 2- 
branched) hairs as long as the gills on the common base and 2 shorter and 
simple hairs before the base. Gills sausage- shaped, with rounded end, 
1.2—1.6 times as long as the siphon. 

Distribution. Northeast China, Korea, Japan, USSR : Khabarovsk and 
Maritime territory, Sakhalin, recorded also from Siberia in the Tomsk 
Region and the Kolyvan District in the Novosibirsk Region. The presence 
of this species in West Siberia is possibly due to accidental introduction by 
railroad. 
303 Biology. The larvae occur in large numbers in water in the root and 

trunk holes in linden, maple, birch, Phellodendron, Siberian stone pine, 
together with larvae of A. alektorovi, A., nipponicus and Toxo- 
rhynchites. They are also sometimes found in rain barrels and other 
water storage containers which are completely or partly dark, together with 
larvae of A. koreicus and C.vagans. Specimens from such biotopes 
are twice as large as specimens from tree holes. Several generations per 
year from May to October. Hibernation in the egg stage. A forest species 
which rarely attacks man. 

46. Aedes (Stegomyia) albopictus Skuse, 1895 

Mesonotum with a white longitudinal stripe. The species differs from 
other soecies with a similar pattern of the mesonotum in the structure of 
the hypopygium and coloration of the hind tarsi. 

Mesonotum mainly covered with chestnut brown scales; the white longi- 
tudinal stripe is narrower posteriorly. In the posterior half of the meso- 
notum there are lateral, longitudinal, white stripes, which are absent in the 
anterior half (in contrast to A.galloisi). At the margin of the mesonotum, 
before the base of the wings, is a small spot of irregular form with silvery, 
broad, straight scales. Tarsi of fore and mid-legs with narrow white rings 
at the base of the first two segments. The first four segments of the hind 
tarsi with wider white rings; last segment of hind tarsi completely white. 
Claws without denticles. Abdomen with white spots at the sides of the 
tergites, and sometimes with stripes at the base of the tergites which are 
narrower or interrupted in the middle. 



326 




FIGURE 207. Hypopygium of A e d es albopictus Sk. 
(after La Casse and Yamaguti) 

Hypopygium (Figure 207): lobe of coxite situated in the middle, longi- 
tudinally elongate, covered with long, simple hairs. Posterior margin of 
tergite 9 with a process in the middle. 

Fourth- stage larva (Figure 208) medium-sized. Frontal hairs 
weakly developed, outer hairs with 2, rarely 1 or 3, short branches, median 
304 hairs displaced anteriorly, with 1—2 branches, inner hairs situated behind 
the median hairs, longer, simple, with slight secondary feathering. Post- 
clypeal hairs stronger than the median hairs, displaced anteriorly, situated 
close together, with 6—15 thin branches and with a short stem. Antenna 
short, thin, without spines, hair situated in the middle, short and simple. 

Comb with 6—13 (usually 8—10) large scales in one row, with a large, 
pointed main spine and a row of short, thin setae at the sides of the base. 
Siphon without auricles, short (index 1.7—2.5), distinctly tapering from the 



327 



middle. Pecten with 7—23 (usually 9—16) denticles, with broad base and 
several accessory denticles, the distal denticle situated in the middle of 
the siphon. Distal to it is a tuft with 2 — 4 (rarely 5—6) branches as long as 
the width of the siphon. 




FIGURE 208. Aedes albopictus Sk. Fourth-stage larva: 

1 — antenna; 2 — siphon; 3 — scales of comb; 4 — denticles of pecten; 5 — 
head, dorsal; 6 — base of a lateral thoracic hair (1—4 after Barraud,5— 6 after 
Senior-White). 



Last segment short, saddle extending to ventral margin of sides; lateral 
hair with 2 (rarely more) branches, one of them longer than the saddle, the 
other half as long; outer caudal hairs simple, twice as long as the siphon 
or longer, inner hairs with 1—3 (usually 2—3) slightly shorter branches. 
Fin usually with 8 usually 2- branched tufts, the posterior branch as long as 
the outer caudal hair. Gills 1.5—2 times as long as the saddle, with slightly 
tapering, rounded end. 

Distribution. It is distributed mainly in the Oriental region and 
Oceania, also in Africa and Australia. In the Palaearctic it occurs in North 
305 China and Japan (except in Hokkaido). Not recorded in the USSR. 

Biology. The larvae develop in large numbers in artificial water 
bodies, containers filled with water and in houses, only rarely in small, 



328 



natural water bodies. Several generations per year (April-October). 
A. albopictus is a mass species in East Asia which occurs in large 
numbers in towns. Observations in Southeast Asia suggest that it feeds 
mainly on man. It is easily bred in the laboratory. 




FIGURE 209. Hypopygium of A e d e s f 1 a v o p i c tu s Yam. 
(after La Casse and Yamaguti) 

Epidemiological importance. A . a 1 b op i c t u s is an important 
vector of dengue in East Asia and Oceania. Some plasmodia of birds 
(Plasmodium lophurae, P. gallinaceum) develop in A. albo- 
pictus to the stage of sporozoites. It is not infected by filanae of man, 



329 



but Dirofilaria immitis of dogs develops to the infective stage in 
A. albopictus. 

47. Aedes (Stegomyia) flavopictus Yamada 1921 

Closely related to A. albopictus but differing in details of the colora- 
tion of the mesonotum and structure of the hypopygium. 

Mesonotum with chestnut brown scales, with a longitudinal white stripe 
which tapers posteriorly and is divided before the scutellum into two narrow 
stripes. There are also two indistinct, lateral longitudinal stripes in the 
posterior half of the mesonotum. Before the base of the wings, slightly 
above it is a weakly outlined spot of whitish or yellowish, narrow, slightly 
curved scales. Narrow, white rings at the base of the first two segments 
of the fore and mid-tarsi; the white rings are wider and present at the 
base of four segments on the hind tarsi; last segment of hind tarsi nearly 
completely white. Claws simple. Wings with dark scales and a small 
306 white spot at the base of the costa. Abdomen with white spots at the sides 
of the tergites and sometimes narrow, white stripes at the base of the 
segments. 

Hypopygium (Figure 209): lobe of coxite longitudinally elongate, extend- 
ing nearly to apex of coxite; hairs of two types present on the lobe: simple 
hairs, slightly shorter than in A. albopictus, and flattened, lanceolate 
hairs. Posterior margin of tergite 9 serrated, convex, curved. 

Fourth- stage larva with strongly developed stellate hairs on the 
body. Frontal hairs: outer hairs with 2—3 short branches, median hairs 
situated before the inner hairs, both pairs with 1 — 2 branches, the last hairs 
longer than the others; postclypeal hairs displaced anteriorly, situated 
close together, with 9—16 branches, with thin secondary feathering and with 
a short stem. Antenna about half as long as the head, without spines, with 
a simple, short hair in the middle. 

Comb with 6—12 scales in a row, with a main spine and with short, thin 
setae at the sides of the base. Siphon short, index 2.2—2.5, tapering from 
the middle, without auricles. Pecten with a varying number of denticles 
(4—16) with a row of accessory denticles at the broad base; pecten nearly 
reaching middle of siphon. Hair tuft with 3—6 branches situated in the 
middle, about as long as the width of the siphon at the base. 

Saddle of last segment extending to lower margin of sides, short, its 
posterior margin covered with spines; lateral hair with 2—7 branches 
which are longer than the saddle; outer caudal hairs simple, twice as 
long as the siphon, inner hairs with 2—3 slightly shorter branches. Fin 
with 6—8 tufts with 1—2 branches which are as long as the inner caudal 
hairs. Gills sausage- shaped, of varying length, as long as the saddle to as 
long as the siphon. 

Distribution. Japan (including Hokkaido), Korea, India, Pakistan. 
In the USSR found only in the Kedrovaya Pad' Reserve (Southern Maritime 
Territory , collected by A. S. Monchadskii). 

Biology. The larvae occur in water in stumps of bamboo, tree holes, 
and artificial water bodies in inhabited areas. The females are active 
bloodsuckers. 



330 



48. Aedes (Stegomyia) chemulpoensis Yamada, 1921 

It is characterized by the coloration of the legs and mesonotum (absence 
of a white longitudinal stripe) and the structure of the hypopygium. 

Mesonotum with dark brown scales and a white pattern consisting of the 
following parts: a) three small spots at the anterior margin, b) two large, 
triangular or semicircular spots at the sides of the anterior half of the 
mesonotum, c) small spots before the base of the wings, d) two longitudinal 
stripes and a small median spot (sometimes absent) in the posterior half 
of the mesonotum. Fore and mid-femora with a white stripe in the basal 
part anteriorly and several white spots in the apical half. Tibiae with a 
spot or ring of white scales in the first third. Tarsi with white rings at 
the base of the segments; last segment of hind tarsi completely white or 
white with dark apex. Claws without denticles. Abdominal tergites with 
white spots at the sides and narrow basal white stripes of uniform width. 

Hypopygium (Figure 210): a sharply defined, round, rugose process 
densely covered with hairs in the middle of the inner surface of the coxite. 
307 Style sword- shaped, without a constriction or a widening at the apex; ap- 
pendage situated at the apex. Posterior margin of tergite 9 with a shallow 
incision. 




FIGURE 210. Hypopygium of Aedes chemulpoensis Yam. 



331 



Fourth-stage larva not described in detail, resembling that of 
A. galloisi. Frontal hairs all simple, median hairs situated before the 
inner hairs, strongly displaced anteriorly, median hairs nearly twice as 
long as the outer hairs. Postclypeal hairs situated between the median 
hairs, with 4—5 branches. Antenna short, smooth, with a short, simple hair 
beyond the middle. 

Hairs on body stellate. Comb on 8th segment with 6—10 scales in a row 
and 2—4 accessory denticles at the sides of the base of the main spine. 
Siphon without auricles, index about 3.0. Pecten extending nearly to middle 
of siphon, with 15 denticles. Tuft with 4 branches, situated in the middle, 
slightly nearer to the apex. 

Saddle extending to lower margin of sides of last segment; lateral hair 
stellate, with many branches; outer caudal hairs long, simple, inner hairs 
with 2—3 branches, shorter. Fin with 6 weakly branched tufts. Gills 
sausage- shaped, 5 times as long as the saddle, both pairs of the same 
length. 

Distribution. Korea, North China. Not found in the USSR. 

Biology. Not known. The larvae occur in holes in trunks and roots 
of Salix matsudana, Quercus acutissima and Soph or a 
japonic a. 



308 5. Subgenus Aedes Meigen, 1818 

The most distinctive character of this subgenus are the short palps of 
the male, which, as in the females, are about /& as long as the proboscis. 
The structure of the hypopygium is also characteristic: style bifurcate at 
the base, without an appendage; claspettes absent. The differences are less 
distinct in the females: proboscis usually as long as the fore femora, spots 
of scales on pleurae of thorax weakly developed, cerci distinctly projecting. 
There are over 70 species in the subgenus, mainly in the Oriental region. 

The larvae closely resemble those of the subgenera Aedimorphus 
and Ochlerotatus. The only character which distinguishes the sub- 
genus Aedes from the other subgenera is the structure of the stigmal 
plate and some characters of the adult mosquitoes. 

49. Aedes (Aedes) cinereus Meigen, 1818 

Small. Proboscis and palps dark brown. Proboscis about as long as 
the fore femora. Pleurae of thorax with small spots of yellowish or white 
scales without a silvery sheen. Tarsi dark. Wings with dark scales, with- 
out light scales. Head, mesonotum and abdomen of varying coloration in 
the different subspecies. 

Hypopygium (Figure 211): coxite short, thick at the base, strongly 
tapering apically. Basal lobe of coxite well developed, densely covered 
with hairs; dorsal part of lobe differentiated into a dorsobasal process. 
Style articulated with coxite distinctly below the apex, bifurcate, the lateral 
branch longer and narrower than the median branch. 



332 




FIGURE 211. Hypopygium of A e des cinereus rossicus D.G.M. 

Fourth-stage larva (Figure 212) medium- sized, semitransparent, 
yellowish to brown or dark. Frontal hairs secondarily feathered, situated 
in a posteriorly concave, curved row; outer hairs with 6—14 (usually 7—8), 
median hairs with 3—7 (usually 5— 7), inner hairs with 3—7 (usually 4—6) 
branches. Postclypeal hairs short, thin, with 3—7 branches. Antenna long, 
about 0.7 times as long as the head, with sparse spines, hair tuft with 4—7 
branches situated at fa of the length of the antenna from the base, half as 
long as the antenna. 

Comb on 8th segment with 10—19 (usually 14—16) scales in 1 — 2 irregular 
rows, scales with a pointed main spine and with a row of small, thin spines 
at the sides of the base. Siphon thin, index 3.0—4.0. Pecten fa — fa as long 
as the siphon, with 13—21 denticles, 1 — 3 denticles near the base rudimentary 
and 1—3 distal denticles more widely spaced and larger. Hair tuft short, 
with 3—7 (usually 4—5) branches, situated nearer to the apex, distinctly 
beyond the distal denticle of the pecten. There are two pairs of short, thin, 
often hardly visible tufts with 2—5 branches on the anterior surface of the 
siphon in some larvae. At the anterior margin of the apical part of the 
siphon, in addition to the apical spines which are present in all larvae, 
there may be 4 pairs of short, thin, slightly branched hairs at the margin of 
the lateral surface; sometimes these hairs are fewer or they are absent. 

The stigmal plate (Figure 213) differs in the structure of the posterior 
process of the "stirrup"; the posterior bow forms a long, chitinized stem 
which is wider distally; at the sides of its posterior margin are two nail- 
shaped processes, curved toward each other. 



333 




FIGURE 212. Aedes cinereus cinereus Mg. Fourth-stage larva: 
1 — head, dorsal; 2 — posterior end, lateral. 

310 Saddle of the last segment extending to middle of sides or beyond this; 
lateral hair with 2, rarely 1 or 3, branches; outer caudal hairs longer than 
the siphon, simple, inner hairs less than half as long, with 4—8 (usually 7) 
branches. Fin with 9—10 tufts on the common base and 2—4 shorter tufts 
before the base. Gills long, at least twice as long as the saddle, narrow, 
pointed, not pigmented. 

Distribution. Europe, Siberia, Far East of the USSR (including 
Sakhalin and Kamchatka), Northeast China, Korea, Japan, Canada, U.S.A. 
It extends rarely to 69°N (Norway), to the Kola Peninsula and the lower 
reaches of the Pechora River. It reaches Transcaucasia and Middle Asia 
in the south. 



Aedes cinereus cinereus Meigen 

Head mainly with dark scales. Scales of mesonotum reddish brown, 
sometimes with a golden or bronze tone. Abdomen without light stripes 



334 



dorsally, with a longitudinal, yellowish white stripe at the sides of the 
tergites. 




FIGURE 213. Aedes cinereus Mg. 
Fourth-stage larva. Stigmal plate. 



FIGURE 214. Style (left) and basal lobe of coxite of Aedes 
cinereus Mg. (after D.K.L'vov): 



A— A. cinereus cinereus; B,C,D 
E — A. cinereus esoensisYam. 



transitional forms: 



311 Hypopygium (Figure 214): lateral branch of style bifurcate at the apex; 

median branch half or a third as long as the lateral branch. Dorsobasal 
process of lobe with two branches, the lateral branch longer than the median. 

Fourth-stage larva with the characters of the species. Typical 
forms do not have additional hairs on the anterior surface of the siphon or 
in its apical part. Transitional forms have one or two pairs of additional 
hairs on the anterior surface of the siphon. The gills become longer 
toward the eastern part of the area of distribution. 

Distribution. Europe, Caucasus, Middle Asia, Kazakhstan, Siberia, 
Far East of the USSR, North America. 



335 



Biology. The larvae develop in permanent, deep, shaded or partly 
shaded water bodies in thinned forests or at their edge and in meadow 
swamps with shrubs or in pools with little vegetation. A. c. cinereus 
is a mass species in many localities and often predominates among 
mosquitoes attacking man. Most of its distribution is in the forest zone, 
but it is more characteristic for open biotopes: meadows, felled forests, 
shrub thickets. The mosquitoes rest in the grass and bite the legs of 
passers by. They are active in the morning and evening and often also 
during the day. 

Aedes cinereus is an all-season species. The mosquitoes appear 
in spring later than the early spring species of Ochlerotatus and fly 
to the beginning of the autumn frosts. After the first generation in late 
spring, there is usually (but not always) a second generation after the 
summer rains. If the second generation is large, maximum numbers are 
observed in August— September (Remm, 1957). 

This is one of the important vectors (mechanical transmission) in some 
outbreaks of tularemia. 



Aedes cinereus rossicus Dolbeshkin, Gorickaja 
and Mitrofanova, 1930 (= A. tarnogradskii 
Martini, 1931) 

It differs in the structure of the hypopygium and in coloration. Head 
mainly with whitish gray scales. Mesonotum with golden brown scales. 
Mesonotum with golden brown scales. An indistinct stripe of yellowish 
white scales at the lateral margins of the mesonotum which is more 
distinct in the posterior half. Abdomen with blackish brown scales dorsally, 
without light, transverse stripes, with a light longitudinal stripe at the sides. 

Hypopygium (Figure 211): style not bifurcate at the apex; median branch 
of style lz— I2 as long as the lateral branch. Dorsobasal process of lobe 
with 2 branches. 

Typical fourth-stage larvae resemble those of the nominate sub- 
species in all characters, but always have two pairs of additio nal, short hairs on 
the anterior surface of the siphon which may be partly or completely absent 
in transitional forms (Figure 215). 
312 Distribution. Ukraine (Dnepropetrovsk, Kiev, Transcarpathian and 

Volyn regions), Caucasus (Ordzhonikidze, Teberda), Chuvashia, Urals 
(Perm, Orenburg) and Yakutia. Also Hungary, Czechoslovakia and Japan. 

Biology. The larvae occur in small, temporary, open water bodies 
with vegetation. Several generations per year. A. c. rossicus 
apparently occurs mainly in floodplains of rivers in large numbers. The 
mosquitoes are vicious biters, active during the day. 



Aedes cinereus esoensis Yamada, 1921 

It differs in the coloration of the abdomen and structure of the 
hypopygium. Head and thorax as in A. c. cinereus. Abdomen with 
blackish brown scales dorsally, sometimes with a bronze sheen. Stripes 



336 




j/~*\ of yellowish brown scales at the base of the tergites 

and spots of the same color at the sides. 

Hypopygium (Figure 214): lateral branch of style 
not bifurcate at the apex, median branch about j$ as 
long as the lateral branch. Dorsal process of lobe 
simple, consisting only of the median branch. 

Fourth-stage larvae of this subspecies 
resemble those of the nominate subspecies in all 
characters but differ from it in the presence of 4 
pairs of additional, short hairs at the apex of the 
siphon at the margins of the lateral surface. The 
hairs may not be fully developed on one side or may 
be absent in transitional forms. Gills longer than 
the siphon. 

Distribution. Far East of the USSR, Kazakh- 
stan, Northeast China, Korea, Japan. 

Biology. There are apparently no differences 
from A.c.cinereus. In many parts of the 
Maritime and Khabarovsk territories, A . c . e s o - 
ens is is a mass bloodsucker. It is considered as 
a possible vector of Japanese encephalitis 
(Petrishcheva, 1947). 

Note on systematics. The forms we con- 
sider as subspecies of A. cinereus are often 
considered as different species. They differ not 
only in coloration but also in important structural 
characters of the genitalia. However, there are 
transitions and intermediate variations in all 
characters. Thus, in the forms transitional between 
cinereus and esoensis, the light stripes on the 
tergites are not present on all segments, they are sometimes very narrow or 
incomplete, and sometimes replaced by a small spot consisting of a few 
scales. The fork at the end of the lateral branch of the style, which is 
characteristic for A. c. cinereus, is formed by two unequal processes, 
one of which may be very short and indistinct. The lateral branch of the 
dorsobasal process is sometimes shortened or forms a small tubercle, etc. 

A special study (L'vov, 1936) showed that of the two subspecies named 
above, only A. c. cinereus occurs in the European USSR. There are 
often transitional forms in Transbaikalia, which are very common in the 
Khabarovsk and Maritime territories where "typical" A. c. cinereus 
and A. c. esoensis also occur, the latter predominating. The transitions 
between A. c. cinereus and A. c. rossicus are similarly related. 
They differ less in coloration than A. c. esoensis from A. c. ciner- 
eus. The hypopygium of A. c. rossicus is intermediate in structure 
between that of A. c. cinereus and A . c . esoensis. 

The same is observed in the larvae. All the important larval characters, 
including the structure of the stigmal plate, are the same in all subspecies 
and do not exceed the variation of the nominate subspecies. Typical larvae 
of A. c. cinereus do not have the additional two pairs of hairs on the 



FIGURE 215. Aedes 
cinereus rossicus D.G.M. 
Fourth-stage larva. Siphon 
lateral. 



337 



anterior surface of the siphon, but larvae occur in the same sample with 
one or two pairs of these hairs but in smaller numbers. The reverse is 
313 the casein A. c.rossicus. Typical larvae of A. c. esoensis with 
four pairs of additional hairs in the apical part of the siphon are found 
rarely in the Far East. Most larvae have 1—2, rarely 3, pairs of hairs, 
frequently of different development, on both sides. One larva with com- 
pletely developed hairs at the apex had two pairs of hairs on the anterior 
surface of the siphon. Only the typical A. c. esoensis has been des- 
cribed from Japan (La Casse and Yamaguti, 1955) and no transitional 
forms or typical A. c. cinereus have been found. 

The data given speak against the species independence of A. eso- 
ensis and A. rossicus. 



50. Aedes (Aedes?) nobukonis Yamada, 1932 

This species is placed in the subgenus Aedes with reservation since 
the males are unknown. It differs from the other species of the subgenus 
in the presence of white spots at the sides of the abdominal segments and 
the absence of light stripes on the tergites. 

Head mainly with brown scales dorsally. Broad grayish white scales 
form spots at the sides of the head. Narrow light scales are scattered 
in the middle of the occiput and form a border at the posterior margin of 
the eyes. Proboscis and palps brown. Proboscis about as long as the 
fore femora. Integument of mesonotum brown, scales narrow, golden- 
bronze. Lighter golden scales form an indistinct longitudinal stripe which 
is divided before the scutellum and borders a small area without scales. 
Scutellum with narrow, golden scales. Pleurae of thorax with spots of 
white scales. Legs: femora dark anteriorly, light posteriorly; tibiae and 
tarsi completely covered with dark scales. Claws on fore and mid-legs 
with a denticle, claws on hind legs simple. Wings with narrow, brown 
scales. 

Abdomen covered with brown scales dorsally. A spot of darker scales 
with a faint metallic sheen on the first tergite. Basal white spots at the 
sides of the tergites; light stripes on tergites absent, so that the abdomen 
is completely dark dorsally. 

Males and larvae unknown. 

Distribution. Japan (Kyushu). The collection of the Zoological 
Institute contains a specimen with the label: Kedrovaya Pad', Maritime 
Territory, 26 July 1940, banks of the Kedrovka, pit under tree roots, 
collected by A. S. Monchadskii. This specimen shows only small dif- 
ferences from the original description, which states that light scales form 
two submedian and two lateral stripes on the head; mesonotum with a 
median and two submedian stripes. 



338 



Supplement to the genus Aede s 
51. Aedes aureus Gutsevich, 1955 

The males of this species are unknown, and its relation to the subgenera 
therefore remains open. 

Medium -sized mosquitoes, with bright golden scales on the lateral parts 
of the mesonotum, a golden spot on the head, and light scales on the wings. 

Head with a distinct median spot of upright, golden scales and hairs 
dorsally. Proboscis and palps with brown scales, and possibly some light 
scales in the middle of the proboscis, which is slightly longer than the fore 
femora. Mesonotum with a longitudinal stripe of chocolate brown scales 
which is wider posteriorly, lateral parts with golden scales. 
314 Tarsi dark. Wings: light, cream -colored scales predominate on the sub- 

costa and also at the base of radius, media and cubitus; veins otherwise 
completely or mainly covered with dark scales. Abdomen mainly with dark 
scales dorsally, lighter, yellowish gray scales forming spots in the middle 
and at the anterior margin of the tergites. Cerci very short. 

Male and larvae unknown. 

A character like the short proboscis (slightly longer than fore femora) 
suggests that the species is related to the subgenus Aedes. 

K.P.Chagin found a few specimens in the Khasan area in the southern 
part of the Maritime Territory (Gutsevich, 1955). 



8. Genus Culex Linnaeus 

Medium-sized or small forms. Palps of male usually longer than or as 
long as the proboscis, rarely shorter (C. hayashii and many exotic 
species); palps of female short. Spiracular setae absent; sternopleural 
and upper mesepimeral setae well developed; usually one lower mesepimeral, 
rarely absent, still more rarely 2—3 setae; at least 4 lower mesepimeral 
setae only in the subgenus Lutzia. A large part of pleurae of thorax 
without scales; there are only some spots of a few scales. Legs: first 
segment of hind tarsi as long as or longer than the hind tibiae, except in the sub- 
genus Barraudius. Claws simple in females. Wings with long radial 
fork. Hair absent at the base of the radius. Postalar plates with narrow 
scales. Cerci of female not projecting. 

Hypopygium: basal lobe of coxite and claspettes absent; subapical lobe 
usually well developed, with more or less modified setae (plates, finger- 
shaped processes, etc.); 10th sternite with a group of spines or with a 
transverse comb of denticles at the end; phallosome usually consisting of 
two pairs of plates, which are often serrated at the margin and with 
secondary denticles. 

The larvae have a large, broad head, long antennae with a well developed 
hair tuft in the form of a fan with secondarily feathered branches and long 
subapical setae often situated some distance from the apex. Mouth parts 
of the filtration type of feeding, with long tufts of simple hairs on the lateral 
lobes of the labrum and on the mandibles. An exception are the predacious 
larvae of L u t z i a, in which the labrum is adapted to capture prey. 



339 



Thoracic hairs usually long and branched. Chitinized plates and processes 
at their base moderately developed. Plates on abdominal segments absent 
except on the last segment, the saddle of which surrounds it like a ring. 
Comb at sides of 8th abdominal segment usually with numerous scales. 
Siphon, with rare exceptions (Lutzia, Barraudius), long or very long 
and thin, with well developed pecten and four or more pairs of hairs or 
hair tufts, situated at the lateral and posterior surface of the siphon or 
displaced posteriorly and forming a zigzag or straight row. Main tracheal 
trunks of two types: thin s with round cross section or broad, ribbon-shaped, 
with oblong-oval cross section. The structure of the tracheae of the first 
type corresponds to the stigmal plate ofC. territans, the tracheae of the 
second type to that of C.pipiens. 
315 The genus contains numerous species (over 400), most of them in the 

tropics of the western and eastern hemisphere. It is divided into 16 sub- 
genera of which those occurring in the USSR are described below. 

The larvae of the subgenera do not completely correspond to the subgeneric 
division of the adults. The species of Lutzia form a distinct group in a 
number of characters, combined with their predacious mode of life, but they 
have some characters which connect them with the larvae of Culex and 
Barraudius (arrangement and number of hair tufts on the siphon, details 
of structure of the stigmal plate). The larvae of the last two subgenera and 
of the subgenus Lophoceratomyia do not have characters which would 
make it possible to separate them. The tracheal trunks of the larvae of 
Neoculex are thin and with round cross section and their stigmal plate is 
of a special type, but the larvae of C.mimeticus and C. jacksoni, which 
belong to the subgenus Culex, also have such characters. The larvae of 
Culex form a distinct group, but this does not correspond to the adults. 



Key to Species 
Females 

1 (4). First segment of hind tarsi distinctly shorter than hind tibiae 

( / 5 — 6 / 7 as long as the tibia). Small species (subgenus 
Barraudius Edw.). 

2 (3). Abdomen with a longitudinal stripe of light scales at the sides 

which sometimes form more or less developed, triangular spots 

at the anterior margin of the tergites 

3. C. (Barraudius) modestus Fie. 

3 (2). Abdomen with more or less developed spots of light scales at the 

sides of the base 4. C. (Barraudius) pusillus Macq. 

4 (1). First segment of hind tarsi slightly longer than or as long as the 

hind tibiae. 

5 (8). Lower mesepimeral setae numerous, at least 4, usually 6—10. 

Large mosquitoes (subgenus Lutzia Theob.). 

6 (7). All abdominal tergites with light stripes of more or less the same 

width at the posterior margin. Basal half of anterior surface of 

hind femora with numerous dark scales . 

2. C. (Lutzia) vorax Edw . 



340 



7 (6). Abdominal tergites 2—4 dark or with narrow, light stripes at the 

posterior margin; tergites 5—8 completely covered with light 
scales or with wide stripes of light scales at the posterior margin. 
Basal half of anterior surface of hind femora with white scales, 
without dark scales 1. C. (Lutzia) fuscanus Wied. 

8 (5). Usually one lower mesepimeral seta which is rarely absent, still 

more rarely 2—3 setae. Usually medium-sized or smaller (sub- 
genera Cul ex L. and N e o c u 1 e x Dyar). 
9(20). Proboscis and tarsi with white rings which are sometimes narrow. 

1 0(1 3). Wings, especially on the costa, with large, light spots formed by 
white scales. 

11(12). Apical part of cui with light scales 

14. C. (Culex) orientalis Edw. 

316 12(11). Apical part of cui with dark scales 

12. C. (Culex) mimeticus Noe. 

13. C. (Culex) jacksoni Edw. 

13(10). Wings without spots, sometimes with more or less numerous, 
scattered white scales. 

14(15). Scales on wings brown, with more or less numerous white 

scales 9. C. (Culex) bitaeniorhynchus Giles. 

15(14). Wings with uniformly brown scales. 

16(19). Scales of mesonotum of 2 colors: anterior 2 / 3 of mesonotum with 

light yellow or whitish gray scales, posterior third with dark brown 
scales. 

17(18). Stripes of light scales at posterior margin of abdominal tergites. 
Anterior part of mesonotum with light yellow scales. Wings with 
narrow scales 10. C. (Culex) sinensis Theob . 

18(17). Stripes of light scales at anterior margin of abdominal tergites. 
Anterior part of mesonotum with white scales. Wing scales 
broader , 11. C. (Culex) whitmorei Giles. 

19(16). Mesonotum completely covered with more or less dark scales . . . 
scales 15. C. (Culex) tritaeniorhynchus Giles. 

20 (9). Proboscis and tarsi without light rings. 

21(24). Abdomen brown dorsally, without transverse stripes or lateral spots 
formed by light scales. 

22(23). Abdomen with blackish brown scales dorsally, sometimes with in- 
distinct stripes of white scales at posterior margin of tergites 

2 3(22). Abdomen with reddish brown scales dorsally. (Palps of male much 

shorter than the proboscis.) Far East 

8. C. (Neoculex) hayashii Yam. 

(Palps of male longer than the proboscis.). Mediterranean .... 
7. C. (Neoculex) martinii Med. 

24(21). Abdominal tergites with more or less developed transverse stripes 
formed by white or yellowish scales (some specimens of 
C. pipiens have rarely no light stripes on the tergites). 

25(28). Fore and mid-femora and tibiae dark anteriorly and with a light 

longitudinal stripe; rarely only the fore femora with a light stripe. 

26(2 7). Light transverse stripes on the abdomen usually triangularly pro- 
duced posteriorly. Hind femora with a dark stripe in the apical 
third on the ventral side 17. C. (Culex) theileri Theob. 

27(26). Light transverse stripes on the abdomen of more or less the same 
width, not produced posteriorly. Hind femora light ventrally their 
entire length 18. C. (Culex) vagans Wied. 



341 



28(25). Fore and mid-femora and tibiae without anterior light stripe. 

29(32). Light transverse stripes at posterior margin of abdominal tergites. 
Proepisterna with broad, light scales. 

30(31). Palps with dark and light scales. Light stripes on abdomen 

relatively broad, sometimes with an anterior process. Hind tibiae 
with a white spot at the apex. Last segments of palps of male bare 
or nearly bare 6. C. (Neoculex) hortensis Fie. 

31(30) Palps with dark scales. Light stripes on abdomen narrow, without 
anterior process. Hind tibiae without a white spot at the apex. 

Last segments of palps of male with long hairs 

317 5.C. (Neoculex) territans Walk. 

32(29). Light transverse stripes or light lateral spots situated at anterior 
margin of tergites. Proepisterna with narrow, light scales. 

33(34). Light transverse stripes on abdomen formed by white scales . . . 

16. C. (Culex) univittatus Theob. 

34(33). Light transverse stripes on abdomen formed by yellowish scales; 
the stripes are sometimes reduced or form spots at the sides of 

the abdomen 20. C. (Culex) pipiens L. 

19. C. (Culex) torrentium Mart. 

Males 

1 (4). Coxite with small scales on the outer surface. Subapical lobe 

situated nearly in middle of coxite. 

2 (3). Style relatively long and thin; phallosome short, its posterior 

margin not extending beyond apex of 10th sternite 

3. C. (Barraudius) modestus Fie. 

3 (2). Style short and thick; phallosome long, its posterior margin ex- 

tending beyond apex of 10th sternite 

4. C. (Barraudius) pusillus Macq . 

4 (1). Coxite without scales on the outer surface. Subapical lobe situated 

distinctly beyond middle of coxite. 
5(14). Subapical lobe of coxite without plate -shaped appendages; 10th 

sternite with a row of large denticles at the apex, or with several 
rows of short spines. 

6 (9). Tenth sternite with numerous spines at the apex. 

7 (8). Denticles of phallosome (sometimes indistinct) situated on body of 

phallosome 1. C. (Lutzia) fuscanus Wied. 

8 (7). Denticles usually well developed, situated on a triangular process 

on the outer margin of the phallosome 

2. C. (Lutzia) vorax Edw. 

9 (6). Tenth sternite with a transverse row of denticles at the apex. 
10(11). Coxite with a broad, flattened, sclerotized process at the apex, ex- 
tending distinctly beyond the base of the style 

6. C. (Neoculex) hortensis Fie. 

11(10). Coxite without a process at the apex. 

12(13). Style short, markedly widened beyond the middle 

7. C. (Neoculex) martinii Med. 

13(12). Style distinctly tapering apically 

5. C. (Neoculex) territans Walk. 



342 



a(b). Phallosome with numerous blunt denticles at the apex .... 

C. territans Walk. 

b(a). Phallosome nearly smooth, without denticles at the apex . . . 

C. territans judaicus Edw. 

14(5). Subapical lobe of coxite with one or several transparent, oval or 

lanceolate plates, plates rarely narrow, knife -shaped. Tenth 

sternite with several rows of spines at the apex. 
15(16). Subapical lobe of coxite with a few, knife -shaped plates 

8. C. (Neoculex) hayashii Yam. 

16(15). Subapical lobe of coxite with one plate. 

17(18). Phallosome with a simple plate at each side which is not denticulate 

or divided. Plate of subapical lobe of coxite lanceolate 

10. C. (Culex) sinensis Theob. 

18(17). Phallosome with to two sclerites on each side, often divided twice 

or denticulate. 
19(20). Phallosome with 2 simple (not divided) sclerites on each side, which 

are pointed at the end and sickle-shaped. Plate of subapical lobe 

of coxite narrow, knife -shaped 

9. C. (Culex) bitaeniorhynchus Giles. 

20(19). Phallosome with 3 sclerites on each side, if there are 2, one of 

them is divided or denticulate at the apex. Plate of subapical lobe 

of coxite broader, leaf-shaped. 
21(24). Tenth sternite without basal appendage; if an appendage is present/ 

it is rudimentary. 
22(23). Second part of phallosome with 4—5 long, finger-shaped 

processes 11. C. (Culex) whitmorei Giles. 

23(22). Second part of phallosome simple .... 20. C. (Culex) pipiens L. 

a(b). Second part of phallosome narrow, hook-shaped 

C. pipiens pipiens L. 

C. pipiens molestus Forsk. 

b(a). Second part of phallosome broad, lobe-shaped 

C. pipiens fatigans Wied. 

24(21). Tenth sternite with a distinct basal appendage which is usually 

very large. 
25(26). Coxite with a tuft of dense hairs at the apex, directed distally, near 

base of style 14. C. (Culex) orientalis Edw. 

26(25). Tuft of hairs at apex of coxite absent. 

27(32). Phallosome relatively simple, with a few denticles. 

28(31). Style tapering from base to apex. Plate of subapical lobe oval. 

29(30). Phallosome divided at the apex into 2 plates, one of them with 2—3 

laterally directed denticles 17. C. (Culex) theileri Theob. 

30(29). Phallosome divided into 3 plates at the apex, one of them directed 

distally 18. C. (Culex) vagans Wied. 

19. C. (Culex) torrentium Mart. 
31(28). Style widened in apical third. Plate of subapical lobe broad, 

asymmetrical 16. C. (Culex) univittatus Theob. 

32(27). Phallosome complicated, consisting of many parts. 
33(34). Second part of phallosome with 2—3 laterally directed denticles 

12. C. (Culex) mimeticus Noe. 

13. C. (Culex) jacksoni Edw. 



343 



34(33). Second part of phallosome with 4—5 denticles, the median denticles 
directed distally 15. C. (Culex) tritaeniorhynchus Giles. 



Fourth 



1 (2 



2 (1 



3 (4 



4 (3 
5(12 

6 (7 



7 (6 

8 (9 



9 (8 
10(11 

11(10 

12 (5 

13(14 



14(13 
15(28 



'i- stage larvae 

Lateral lobes of labrum with 30—40 large, S— curved setae with a 
comb of short spines in the distal half. Siphon short, not longer 
than last abdominal segment, sharply tapering apically, pecten 
occupying its entire length to the tapering part (subgenus 

Lutzia) .' C. (Lutzia) vorax Edw. 

C. (Lutzia) fuscanus Wied, 
Lateral lobes of labrum with normal setae. Siphon longer than last 
abdominal segment, not sharply tapering apically, pecten occupying 
at most half its length. 

Siphon short, usually 3 times longer than wide at the base. Hair 
tufts on siphon arranged in a zigzag row on the posterior surface 

C. (Barraudius) pusillus Macq. 

Siphon at least 4 times longer than wide at the base. 

Comb of 8th abdominal segment with 4—8 large scales with a 

pointed main spine. 

Mentum with numerous small denticles of equal size at the margin. 

Hairs of lateral lobes of labrum less than the usual number of thin, 

angularly bent or hook-shaped, slightly branched hairs 

C. (Culex) bitaeniorhynchus Giles. 

Mentum with a few denticles which are larger in the middle. 
Mentum with 10—15 small denticles at the base on each side and 
6—8 denticles which are larger in the middle. Suba^ical setae of 
antenna situated in the middle between end and hair tuft. Pecten 

with 2—4 small denticles at the base of the siphon 

C. (Culex) sinensis Theob. 

Mentum only with larger denticles. Pecten with 8—14 denticles 
occupying x / 4 of length of the siphon. 

Hair tufts on siphon short, as long as the width of the siphon at 
their position, each with 3 — 5 branches. Gills at most half as long 

as the siphon *C. (Culex) vishnui Theob.* 

Hair tufts on siphon very long, more than twice as long as the width 
of the siphon at their position, each with 2 branches. Gills more 

than half as long as the siphon C. (Culex) whitmorei Giles. 

Comb of 8th abdominal segment with at least 20 scales, without a 
main spine in most species, and with a row of denticles or spines 
of different size at the margin. 

Siphon, in addition to the pecten and subapical setae, with 2 — 9 addi- 
tional spines on the posterior side in the distal quarter 

C. (Culex) jacksoni Edw. 

Siphon without additional spines in the distal quarter. 
All hair tufts on the siphon (or except for one or two at the apex) 
situated on the posterior surface in a zigzag row or in pairs situ- 
ated close together in the midline. 



* An asterisk indicates species included only in the key to larvae; they do not occur in the USSR. 



344 



16(21). Main tracheal trunks narrow, less than half as wide as the siphon, 
with round cross section. Stigmal plate of the "C . territans" 
type. 

17(18). Inner and median frontal hairs short, at least half as long as the 
outer hairs. Denticles of pecten forming long, thin spines with a 
row of pointed, thin accessory denticles on the entire posterior 
margin. Fat body dark, so that the larvae appear transversely 
striated C. (Neoculex) hayashii Yam. 

18(17). Inner and median frontal hairs as long as or longer than the outer 
hairs. Fat body not dark, larvae not appearing transversely 
striated. 
320 19(20). The basal 2—3 hair tufts on the siphon situated between the more 

widely spaced distal denticles of the pecten, at least 3 times longer 
than the width of the siphon, the following tufts become shorter 
toward the apex C. (Neoculex) hortensis Fie. 

20(19). Basal hair tufts on siphon situated distal to the last denticle of the 
pecten; hair tufts usually twice as long as the width of the siphon 
C. (Culex) mimeticus Noe. 

21(16). Main tracheal trunks ribbon- shaped, broad, more than half as wide 
as the siphon, with oval cross section. Stigmal plate of the 
"C. pipiens" type. 

22(25). The first 3 basal hair tufts on the siphon situated between the more 
widely spaced distal denticles of the pecten. 

23(24). In addition to the hair tufts forming a zigzag row on the posterior- 
surface of the siphon, there are 2 pairs of shorter tufts in the 
apical quarter of the siphon, one pair farther from the apex and 
situated at the lateral surface, the other at the border of the lateral 
and posterior surface *C. (Culex) laticinctus Edw . 

24(23). All hair tufts on the siphon situated on the posterior surface . . . 

C. (Barraudius) modestus Fie. 

25(22). Basal hair tufts on siphon situated distal to the last denticle of 
the pecten. 

26(27). Subapical setae of antenna situated at the apex of the narrow 

terminal part of the antenna. Five pairs of hair tufts on the siphon, 
the penultimate tuft from the apex situated on the lateral surface 
C. (Culex) theileri Theob. 

27(26). Subapical setae on antenna situated at V 4 — V 3 of the length of the 
narrow apical part of the antenna before the apex. Six to 8 pairs 
of hair tufts on the siphon situated either in pairs or in a zigzag 
row. Two pairs of tufts near the apex situated on the lateral sur- 
face of the siphon C. (Culex) orientalis Edw. 

28(15). Hair tufts on siphon situated in pairs at the margins of the 

posterior surface, some tufts may be situated on the lateral surface. 

29(32). Siphon widest in the middle, more strongly tapering apically than 
toward the base. 

3 0(31). Four pairs of hair tufts situated at sides of posterior surface of 
siphon in a line, their length less than the width of the siphon at 
their position. Width of siphon at the apex half its width at the 
base *C. (Culex) gelidus Theob. 



345 



31(30). The penultimate pair from the apex of the 4 pairs of hair tufts on 
the siphon situated on the lateral surface; hair tufts at least as 
long as the width of the siphon at their position. Width of siphon 

at the apex more than half its width at the base 

C. (Culex) pipiens fatigans Wied. 

32(29). Siphon widest at the base. 

33(36). Main tracheal trunks narrow, less than half as wide as the siphon, 
with round cross section. Stigmal plate of the "C . territans" 
type. 

34(35). Siphon (index about 7) distinctly wider at the apex. Denticles of 

pecten with 1—2 accessory denticles at the posterior margin .... 
C. (Neoculex) territans Walk. 

35(34). Siphon (index 7.5—11) not wider at the apex or only slightly 

widened. Denticles of pecten with 3—5 long accessory denticles at 

the posterior margin C. (Neoculex) martinii Med. 

321 36(33). Main tracheal trunks ribbon-shaped, broad, more than half as wide 
as the siphon, with oval cross section. Stigmal plate of the 
"C . pipiens" type. 

37(38). Hair tufts on siphon very short, less than half as long as the width 

of the siphon at their position C. (Culex) laurenti Newst. 

38(37). Hair tufts on siphon more than half as long as its width at their 
position, or longer. 

3 9(42). All hair tufts on siphon of the same length. 

40(41). Six to 7 pairs of tufts on the siphon. Inner caudal hairs with 

3 branches. Subapical setae on antenna situated at Y 4 — Y 3 of the 

length of the narrow terminal part from the apex 

C. (Culex) tritaeniorhynchus Giles. 

41(40). Four to 5 pairs of hair tufts on the siphon. Inner caudal hairs with 
2 branches. Subapical setae on antenna situated near apex of 

narrow terminal part of antenna 

C. (Culex) univittatus Theob. 

42(3 9). At least the first two pairs of hair tufts from the base of the 
siphon longer than the others and longer than the width of the 
siphon at their position. 

43(44). A pair df hair tufts near the apex is the most lateral 

C. (Culex) torrentium Mart. 

44(43). The second pair of hair tufts from the apex of the siphon is the 
most lateral. 

45(48). Four pairs of hair tufts on the siphon. The basal pair of hair tufts 
of the siphon situated distinctly distal to the distal denticle of the 
pecten. Lateral hair on last segment ususally simple. 

46(47). Siphon longer (index more than 4.5). Gills distinctly longer than 
the saddle, both pairs of the same length. Ends of posterior tufts 

of fin at the same level as ends of gills 

C. (Culex) pipiens pipiens L. 

47(46). Siphon shorter (index at most 4.5). Gills not longer than the 

saddle, lower pair slightly shorter than the upper. Ends of posteri- 
or tufts of fin distinctly extending beyond ends of gills 

C. (Culex) pipiens molestus Forsk. 



346 



48(45). Five pairs of tufts on the siphon. The basal pair situated at the 
same level or between the 1st and 2nd distal denticles of the 
pecten. Lateral hair on last segment usually 2 -branched ...... 

• • • C. (Culex) vagans Wied, 



1. Subgenus Lut z ia Theobald 

The species of Lutzia differ from the species of the other subgenera 
of C u 1 e x in the presence of 4 or more lower mesepimeral setae. 

The larvae have a small, rounded head with mouth parts adapted for a 
predacious mode of life. The lateral lobes of the labrum catch the food; 
their hairs are modified to about 40 of large, curved setae on a strong, broad 
base. The teeth on the maxillae are large and long. Frontal hairs long, 
usually simple. Siphon short, with a median row of hair tufts along nearly 
the entire posterior surface; pecten extending distinctly beyond middle of 
siphon. Last abdominal segment very long, often longer than the siphon. 
The ring of the saddle of this segment is much wider on the dorsal side than 
on the ventral side. 

A small subgenus (7 species) in the tropics of the New World (2 species) 
and the Old World (4 species in the Oriental region, one in the Ethiopian 
region); two Oriental species occur in the Palaearctic. 

The larvae occur in small natural and artificial water bodies, often near 
inhabited areas, and together with larvae of other species of mosquitoes 
(A. koreicus, C. vagans, and others). They occur from June to 
autumn. The females are normally not bloodsuckers. 



1. Culex (Lutzia) fuscanus Wiedemann, 1820 

Large mosquitoes, with a distinct coloration of the abdomen. Head with 
narrow, whitish scales and numerous upright, dark scales. Proboscis with 
dark scales at the base and apex, with light scales in the middle which 
sometimes predominate. Palps of female with dark and light scales, palps 
of male with indistinct light rings and light apex. Mesonotum with golden 
brown, narrow scales and with light scales which form indistinct spots or 
stripes. Legs: anterior surface of fore and mid -femora with dark scales 
and numerous light scales; anterior surface of hind femora with white 
scales in the basal half, except at the dorsal margin. The light scales in the 
apical half of the dorsal surface of the hind femora form a line to the apex. 
Tarsi without light rings. Wings with brown scales. Abdomen: tergites 
5—8 with yellow scales or the scales form broad stripes at the posterior 
margin of the tergites; tergites 2—4 dark or with narrow stripes of light 
323 scales at the posterior margin. The abdomen thus appears distinctly 

2 -colored dorsally: the anterior half dark, the posterior half light. This 
contrast is less distinct in the males. 

Hypopygium (Figure 216): coxite moderately thick; subapical lobe of 
coxite with 3 thick spines and several hairs; style tapering apically. 



347 



;322) 




FIGURE 216. Hypopygium of Culex fuscanus Wied. 



Phallosome markedly sclerotized, without denticles on its narrow, curved 
part or with a few small denticles. Plates of 10th sternite with numerous 
spines at the apex; basal process of 10th sternite well developed but 
weakly sclerotized. Lobes of 9th tergite slightly convex, with a row of 
long spines. 

Fourth-stage larva* (Figures 217 and 218) large, with yellowish 
brown body and yellowish red or brownish red head and lighter siphon. 

Head slightly oblong. Labrum modified for catching prey, lateral lobes 
consisting of about 40 strong, slightly S -curved setae with a row of thin 
spines at the end. Teeth of mandibles large and long. Antennae short, 
with a short hair near the base and large, pointed setae at the end. Frontal 
hairs long, simple or 2 -branched. Siphon short, with distinctly tapering 
apex because of the curved posterior surface, on which are situated 14—15 
long hair tufts with 2—3 secondarily feathered branches, extending its 
entire length. Beyond the middle of the siphon is a pair of thin, shorter 
hairs on the lateral surface. Pecten with 8 — 11 denticles, occupying the 
siphon to the apical tapering. Comb with 3 5 or more scales, varying from 
elongate scales with a distinct main spine to short scales with a row of 
spines at the margin. Last segment distinctly longer than the siphon. 

* The species of Lutzia are not distinguishable with certainty by the larvae. This description may refer also 
to larvae of C ul ex (Lutzia) vorax. 



610144 



348 




FIGURE 217. Culex fuscanus Wied. Fourth-stage larva. Head, dorsal, and seta 
of labrum- 



Saddle surrounding the segment like a ring and more than twice as long on 
the dorsal side than on the ventral side; its posterior margin thus sharply 
sloping. Saddle covered with small spines. Caudal hairs long, simple. 
Gills short. 

Distribution. USSR : southern part of the Maritime Territory. 
Korea, Japan, China, Southeast and South Asia. 

Biology. The larvae are found in small, stagnant water bodies, natural 
and artificial, near inhabited areas. They feed on larvae of other Culicidae. 



2. Culex (Lutzia) vorax Edwards, 1921 

Closely related to C. fuscanus, differing as follows: femora and tibiae 
mainly with dark scales and with light scales which form spots which 
regularly alternate with the dark parts. Anterior surface of hind femora 
with numerous dark scales in the basal half of anterior surface, without a 
longitudinal line formed by white scales. Abdominal tergites 2—8 with 
narrow stripes of light scales at the posterior margin. Apex of palps of 
male dark. Denticles of phallosome situated on a distinct, triangular, lateral 
process (Figure 219). 



349 





FIGURE 218. Culex fuscanus Wied. 
Fourth-stage larva. Siphon, lateral. 



FIGURE 219. Hypopygium of C ulex vorax Edw. 
(after Barraud) 



Larvae, see C. ( L . ) fuscanus. 

Distribution : USSR : southern part of the Maritime Territory. 
Japan (Hokkaido, Honshu, Shikoku, Kyushu), Korea, China, Southeast and 
South Asia. 
325 Biology. The larvae occur in Japan in various water bodies, wooden 

or metal rain barrels, etc., rarely in natural water bodies. They tolerate 
a high degree of pollution. They often occur together with larvae of 
Culex pipiens and C. t r i t a e n i o rhy n c hu s, which they kill in 
large numbers. It rarely bites man. 



2. Subgenus B a rr au d iu s Edwards 

First segment of hind tarsi distinctly shorter than the tibia (about % as 
long as the tibia); palps of male longer than the proboscis, without long 
hairs; head with narrow scales at the margin of the eyes. Coxite with 
scales on the outside; subapical lobe situated slightly beyond the middle, 
without plate -shaped appendages. 

Larvae, see p. 339. 

The subgenus contains only two species. 



350 



3. Culex (Barraudius) modestus Ficalbi, 1889 

Small mosquitoes. Head with yellowish brown scales. Proboscis and 
palps brown. Palps of male very thin as they do not bear long hairs. 
Mesonotum with brown scales, scutellum with whitish scales, femora and 
tibiae with brown scales anteriorly and white scales posteriorly. First 
segment of hind tarsi distinctly shorter than hind tibia. Wings with brown 
scales; abdomen with brown scales, with a longitudinal stripe of yellowish 
white scales laterally. 



(32 6) 




FIGURE 220. Hypopygium of C ul e x modestus Fie. 



Hypopygium (Figure 220): coxite with more or less dense scales on the 
outside; subapical lobe situated slightly beyond middle of coxite, without 
plate -shaped appendage, lobe with 2—3 thick, blunt spines and a few hairs; 
style curved in apical half, thin and long, more than half as long as the 
coxite, rarely as long as this; phallosome forming a simple, nearly straight 



351 



plate on each side which does not project beyond the apex of the 10th 
sternite which is covered with spines. 



(32 6) 




FIGURE 221. Culex modestus Fie. Fourth-stage larva. Posterior end, lateral. 



327 



Fourth-stage larva (Figure 221) weakly pigmented, nearly trans- 
parent, with light head and siphon. Head nearly 1.5 times wider than long. 
Mouth parts of the filtration type. Antennae moderately spinose, light, 
with darker base, tapering from the hair tuft (at about % of the length of the 
antenna from the base), with 15—25 secondarily feathered branches, half as 
long as the antenna. Frontal hairs secondarily feathered, outer hairs with 
7—8, median hairs with 3—4 (rarely 2), inner hairs with 3 — 5 (rarely 2) 
branches. 

Comb of 8th abdominal segment with 50 and more adjacent, small, 
oblong scales with a row of spines at the apex and part of the sides. 
Siphon straight, tapering apically, 3.8 — 5 times longer than its width at the 
base. Pecten occupying about % of the length of the siphon, with 12—13 
relatively widely spaced, thin denticles with 3 — 5 accessory spines at the 
posterior margin. About 10 hair tufts in the middle of the posterior sur- 
face in a zigzag row; they begin at the 2 — 3 distal denticles of the pecten 
and extend nearly to the apex. The 2—4 apical tufts may be shorter than 
the others and less branched. The other tufts are as long as or longer 
than the width of the siphon at their position, with 5— 7 branches. Main tracheal 



610144 



352 



trunks broad, ribbonlike. Stigmal plate as in C. pi pi ens. Last segment 
surrounded by the saddle like a ring, slightly longer than wide. Lateral 
hair with 2 — 3 short branches. Outer caudal hairs long, simple, inner hairs 
shorter than the outer, with 3—4 branches, the lower branch longer. Fin 
with 11 — 13 tufts with 5—7 branches. Gills distinctly shorter than the 
saddle, both pairs of the same length. 

Distribution. USSR : European part to the Moscow and Sverdlovsk 
regions in the north, Caucasus, Transcaucasia, Middle Asia, Kazakhstan, 
southern Siberia to the Maritime Territory. Southern Europe, Southwest 
Asia, Northern India. 

Biology. The larvae occur from the end of spring to late autumn in 
fresh or slightly saline water bodies, in well illuminated marches, with rich 
vegetation, together with larvae of A. maculipennis, A. m. sacharovi, 
A. hyrcanus and rarely of A. pu 1 ch e r r i m u s. The mosquitoes are 
found on meadows and other open habitats. 

Natural infection of C. modestus with tularemia has been found. 



4. Culex (Barraudius) pusillus Macquart, 1850 

Small mosquitoes, differing, like C. modestus from other Palaearctic 
species of Culex in the shorter first segment of the hind tarsi, which are 
% as long as the tibia. It differs from C. modestus in its darker color- 
ation and white lateral spots at the base of the abdominal tergites. 

Hypopygium (Figure 222): style straight or slightly curved, relatively 
short (in distinction to C. modestus), less than half as long or half as 
long as the coxite; the markedly convex lobe of the coxite bears two thick, 
spines of different size, with thin, curved end and numerous setae and hairs; 
phallosome on each side with a separate narrow, long plate which is curved 
at the end; apex of phallosome usually projecting beyond apex of 10th 
sternite, densely covered with spines, nearly reaching to the subapical lobe. 

Fourth-stage larva (Figure 223) nearly transparent, yellowish or 
pale yellowish green, with slightly pigmented head and siphon. Head trans- 
verse-oval, 1.4 times wider than long. Antennae weakly spinose, long, 0.8 — 
0.9 times as long as the head, pale, except at the base and with tapering 
terminal part. Hair tuft with 15—2 5 secondarily feathered branches, more 
than half as long as the antenna; subapical setae situated near the apex. 
Frontal hairs long, outer hairs with 7, median and inner hairs with 2 — 3 
branches. Comb of 8th abdominal segment with 50 or more small, adjacent 
scales with a row of spines at the margin. Siphon short, 2.6—3.2 longer than 
its width at the base. Pecten about half as long as the siphon, with 11 — 13 
denticles. About 10 hair tufts are situated in a zigzag or straight line in 
the middle of the posterior surface of the siphon; about 3 basal tufts are 
situated between the distal denticles of the pecten. Tufts with 8-10 branches 
at least as long as the width of the siphon. Main tracheal trunks broad, 
ribbonlike. Stigmal plate as in C. pi pi ens. Last segment slightly shorter 
than the siphon. Lateral hair short, with 1-2 branches. Outer caudal hairs 
more than 1.5 times as long as the siphon, inner hairs shorter, with one, 
rarely 2, shorter branches. Fin with 12 well developed tufts. Gills about 
half as long as the saddle, upper pair slightly longer than the lower. 



353 




FIGURE 222. Hypopygium of C ulex pusillus Macq. 



Distribution. Mediterranean, mainly the eastern part. USSR: Middle 
Asia (Turkmenia, Uzbekistan, Tadzhikistan). North Africa (Algeria, Egypt), 
West Asia (Anatolia, Iran, Iraq). 

Biology. The larvae occur in stagnant water bodies and swamps with 
a high content of salt (to 3%), often together with larvae of A. c a s p i u s, 
A. pulcherrimus and others. Breeding places mainly near the coast, 
salt lakes and oases. Not a common species. 



3. Subgenus Neoculex Dyar 

Coxite without scales, subapical lobe with more or less large, spine - 
shaped appendages, usually without a leaf-shaped plate. Tenth sternite of 
most species without a basal process, with a group of spines or a row of 
blunt, plate -shaped processes at the apex; phallosome simple, formed by 
one pair of plates. 



354 




FIGURE 223. Culex p u sill u s Macq. Fourth-stage larva (after Edwards): 
1 — head, dorsal; 2 — posterior end, lateral. 



355 



330 The larvae have a long, thin siphon which is sometimes slightly wider 

apically, thin main tracheal trunks with round cross section, and stigmal 
plate of the C. (N.) territans type. The larvae of C. ( C ul e x ) 
mimeticus and C. (Culex) jacksoni closely resemble those of 
Neoculex. 




FIGURE 224. Hypopygium of C ul e x territans Walk, (after Carpenter and 
Lac 



356 



The subgenus contains about 60 species, 30 species in tropical Africa, 
9 in the Oriental region, 10 in Australia, 5 in Central America and 8 in the 
Palaearctic region (one of them also in the Nearctic region). C. terri- 
331 tans has a holarctic distribution, 5 species occur in the Mediterranean 

in the broad sense (2 species reach Middle Asia) and 2 occur in the Far East. 



5. Culex (Neoculex) territans Walker, 1856 (apicalis auct., nee Adams) 

Described from the United States of America. The related species 
C. apicalis Adams, 1903 was also described from the U. S. A. The 
European species of the subgenus Neoculex was first identified as 
C. apicalis and appeared under this name in many publications (Edwards, 
1921,1926; Martini, 1931; Monchadskii, 193 6, 1951; Shtakel'berg, 1937, and 
others). However, later study showed that the Palaearctic species is 
identical with C. territans and not with C. a p i c a li S, which is 
distributed in the southwestern part of the United States and in Mexico, but 
does not occur in the Palaearctic (Mattingly, 1953, 1955; Kramar, 1957). 
Thus all Russian records and mentions of C. apicalis refer to 
C. territans. 





FIGURE 225. Culex territans Walk. Fourth-stage larva: 
1 — siphon, lateral; 2 — stigmal plate. 



357 



C. rubensis Sasa and Takahashi, from Japan is probably a Far 
Eastern subspecies of C. territans. The two forms are indistinguishable 
in adults and larvae (La Casse and Yamaguti, 1955) and C. rubensis 
is probably a synonym of C. territans. 

Culex territans differs from related forms, particularly from 
C. hortensis, in its usually darker coloration, the long hairs on the 
palps of the male and structure of the hypopygium. 

Head with white and yellowish scales. Proboscis and palps dark brown, 
with dark brown scales, without light scales. Palps of male with long hairs 
on the last two segments. Mesonotum with brown scales; anterior and 
lateral margins of mesonotum and scutellum with whitish gray scales. 
Pleurae of thorax gray, with spots of white scales. Legs dark brown, 
femora and tibiae with dark brown scales anteriorly, tarsi entirely with 
dark brown scales; femora and tibia with whitish scales posteriorly; apex 
of hind tibiae without a white spot. Veins of wings with dark brown scales. 
Abdomen with blackish brown scales dor sally, with narrow, white trans- 
verse stripes of more or less the same width at the posterior margin of the 
tergites. Abdomen usually greenish ventrally, with brown and white 
scales, usually forming broad transverse stripes. 

Hypopygium (Figure 224): subapical lobe of coxite with 2 thick, trans- 
parent spines, curved at the end and several setae, straight or slightly 
332 curved or denticulate. Phallosome with 2 simple plates connected by a 
transverse bridge at the base and before the apex (there is no transverse 
bridge in the apical part of the phallosome in C. apicalis which is the 
main difference from C. territans). The development of this bridge 
varies markedly according to material from different parts of the USSR. 
It may be broad or narrow, sometimes very narrow. 

Fourth — stage larva (Figure 22 5) at first transparent, later 
yellowish green or bluish green. Head and siphon also become brown. 

Head 1.6 times wider than long. Mouth parts of the filtration type. 
Antennae nearly as long as the head, with a fan -shaped tuft of 25—32 
secondarily feathered branches, 3 / 4 as long as the antenna; subapical setae 
situated at the apex. Frontal hairs long, outer hairs secondarily feathered, 
with 6—8 median branches, hairs longer, simple, inner hairs with 2 branches. 

Comb of 8th abdominal segment with 50 or more oblong, small scales 
with a row of small spines at the margin. Siphon thin, 7 times longer than 
wide at the base; slightly but distinctly widening at the apex. Pecten with 
13—16 denticles occupying about % — % of the length of the siphon from the 
base. Four to 6 pairs of hair tufts which are longer than the width of the 
siphon, with 2—4 branches situated at the border between the sides and the 
posterior surface of the siphon. Basal pairs situated more closely 
together than the apical pairs. Main tracheal trunks thin, with round cross 
section. Stigmal plate (Figure 225) with strongly developed posterior 
valves, a long, thin, posterior process of the "stirrup" and small, deeply 
invaginated, rounded spiracles. Last abdominal segment long, with a ring- 
shaped saddle. Caudal hairs: outer hairs longer than the siphon, simple, 
inner hairs with 4 shorter branches. Fin with 13—14 tufts. Gills nearly 
as long as the saddle, lower pair shorter than the upper. 

Note on systematic s. The species is divided into two subspecies — 
C. 1. 1 e r r it an s and C.t. judaicus, which is sometimes considered as 
a different species (Mattingly, 1953), but more proof is necessary to confirm 
its status. 



358 



Distribution. USSR : European part from the Leningrad and 
Sverdlovsk regions in the north to the Ukraine, Crimea, Ciscaucasia and 
Lower Volga area in the south, Caucasus and Transcaucasia, Middle Asia, 
Kazakhstan, Siberia, Far East. Greater part of Europe outside the USSR, 
Japan, North Africa and North America. 

Biology. The larvae occur from June to the end of September in 
temperate latitudes, mainly in small water bodies which do not dry in 
summer, with strong sunlight and abundant vegetation. They are found with 
larvae of A. maculipennis, A. cinereus, C. alaskaensis and 
others. In the south they occur only in shaded water bodies, sometimes in 
springs and swamps. They feed on reptiles and amphibians. They 
usually do not bite people and farm animals. 



333 Culex territans judaicus Edwards, 1926 

It differs from the nominate subspecies in the shorter palps of the male, 
the length of the two apical segments, which are about half as long as the 
long 3rd segment (they are about % as long in the nominate subspecies); 
long segment of palps of male without long hairs, terminal segments with 
thinner hairs, phallosome with indistinct tubercles at the apex. Small 
mosquitoes, wings about 3 mm long (about 4 mm inC. t. territans). 

Distribution. Southwest Asia (Palestine) . 



6. Culex (Neoculex) hortensis Ficalbi, 1889 

Closely related to C. (N) territans, differing in the light spot at the 
apex of the hind tibiae, the bare or nearly bare apex of the segments of the 
palps of the male and the structure of the hypopygium. Palps of female 
with light scales dorsally. The light transverse stripes on the abdominal 
tergites are usually broader than in C. territans and the stripes are 
sometimes triangularly produced anteriorly into the area with dark scales. 
The coloration of body and scales is usually paler. 




FIGURE 226. Hypopygium of C ul ex hortensis Fie. and distal 
part of coxite, lateral 



359 



(334) 




FIGURE 227. Culex hortensis Fie. Fourth-stage larva. 
Siphon, lateral. 



610144 



360 



Hypopygium (Figure 22 6): coxite with a broad, sclerotized, flattened 
process at the apex; spine shaped appendages of subapical lobe of coxite 
relatively short; plate -shaped appendage of lobe absent. 

Fourth-stage larva (Figure 227) often not transparent, yellowish 
brown, cuticle covered with small chaetoids, which are denser on the 
mesothorax and metathorax. 

Head more than 1.5 times wider than long. Antennae as long as the head, 
densely covered with spines, often darkly pigmented. Hair tufts with 25— 
3 5 secondarily feathered branches, % as l° n g as the antenna. Subapical 
setae situated a short distance before the apex. Frontal hairs: outer hairs 
with 7—12 branches, median hairs situated before the inner, slightly more 
widely spaced, both pairs with 2—4 longer and thicker branches. 

Comb of 8th abdominal segment with 35—4 5 elongate scales with a row 
of spines at the margin of the slightly wider terminal part. Siphon long, 

334 about 8 times longer than the width of the distinctly widened base. Pecten 
about y 3 of the length of the siphon, with 11 — 14 denticles. Nine to 12 hair 
tufts on the posterior surface of the siphon in a zigzag row, 2—4 basal tufts 
situated between the distal denticles of the pecten. The first 6—8 tufts are 
very long, the next become gradually shorter. There are also 1 — 2 pairs of 
shorter lateral tufts on the lateral surface of the siphon near the apex. 

335 Main tracheal trunks thin, rounded; stigmal plate as in C.territans. 
Last segment long, with a ring-shaped saddle. Lateral hair with 2 branches. 
Fin with 12—14 tufts. Gills as long as the saddle or longer. 

Distribution. Mediterranean. USSR : Crimea, Caucasus and Trans- 
caucasia, Middle Asia (Turkmenia, Uzbekistan, Tadzhikistan), South Kazakh- 
stan; Europe to Berlin in the north; West Asia (Asia Minor, Palestine, 
Syria, Iran); North Africa. 

Biology. The larvae occur in fresh water with algae and vegetation, 
often together with larvae of Anopheles maculipennis, Culiseta 
longi ar e ol at a, C. annulata, Culex laticinctus Edw. and others. 
The mosquitoes rest during the day in caves and other dark places. They 
do not bite man often. 



7. Culex (Neoculex) martinii Medschid, 1930 

Very small mosquitoes. Coloration yellowish brown. Abdomen with 
uniformly colored scales dorsally. Proboscis and tarsi without light rings. 
Head pale; genae with white scales. Proboscis brown, slightly thickened 
in the apical third. Palps of female very short, brown; palps of male longer 
than the proboscis by about the length of the last segment, not thickened at 
the apex. Mesonotum yellowish, with small, golden brown scales and long, 
brown hairs. Femora and tibiae with dark scales anteriorly and light 
scales posteriorly. Wings with brown scales at the veins. Abdomen with 
brown scales dorsally and triangular whitish spots at the sides of the 
tergites which become wider toward the posterior margin of the tergites. 

Hypopygium (Figure 228): coxite short and thick; lobe situated beyond 
middle of coxite, near the apex, lobe with 2 moderately long but very thick 
spines. Style short and broad, distinctly wider in the middle. Tenth 
sternite with broad, lobe -shaped lateral processes with small hairs and a 
row of narrow, blunt denticles which form a comb or a fringe at the apex. 



361 




FIGURE 228. Hypopygium of C ul e x martiniiMed. 



336 




FIGURE 229. Culex martiniiMed. Fourth-stage larva. Posterior end, lateral. 



362 



Fourth -stage larva (Figure 229) : head 1.5 times broader than 
long. Antenna at least % as long as the head, hair tuft with 22— 26 secondarily 
feathered branches, 0.7— 0.8 times as long as the antenna. Subapical setae as 
long as the tuft, situated a short distance before the apex. Outer frontal 
hairs usually with 5 branches, median hairs simple, rarely 2 -branched, 
very long, reaching to the apex of the antenna or even beyond it, situated 
before the distinctly shorter, 2 -branched inner hairs and more widely 
spaced than these. Comb with 35—40 small, markedly elongate scales with 
a row of spines at the margin and at the apex. Siphon very thin, long, 
7.5—11 times longer than the width of the wide base, sometimes slightly 
widened at the apex. Pecten with 12—16 denticles in the basal % of the 
siphon and with 3—5 accessory denticles. Ten to 12 hair tufts in a zigzag 
row at the sides of the posterior surface of the siphon and then in irregular 
pairs toward the apex, some of them situated on the lateral surface. Hair 
tufts short, with 2—5 branches, as long as the width of the siphon or slightly 
longer, apical tufts shorter. Tracheae thin, with round cross section; 
stigmal plate of the C. territans type. Last segment long, with a ring- 
shaped saddle. Lateral hair 2 -branched. Outer caudal hairs simple, 
usually as long as the siphon, inner hairs 4-branched. Fin with 11 — 12 tufts. 
Gills about half as long as the saddle. 

Distribution. Mediterranean. USSR : Middle Asia, Uzbekistan 
(Lisova, 1935 — "C. sp."), Tadzhikistan, Kirghizia. Balkans (Yugoslavia), 
Asia Minor (Anatolia). 

Biology unknown. 

8. Culex (Neoculex) hayashi Yamada, 1917 

Small mosquitoes, differing from the other species of Culex in the 
relatively short palps of the male and the structure of the hypopygium. 
Proboscis and palps dark. Palps of male % — % as long as the proboscis. 
Mesonotum brown, with narrow, reddish brown scales with a bronze sheen. 
337 Tibiae and tarsi without light rings. Wings with dark scales. Abdomen 
with dark brown scales dorsally, without light stripes. 

Hypopygium (Figure 230): subapical lobe of coxite with 4—5 closely 
spaced, narrow plates and 2—3 longer, rod -shaped appendages with hooked 
end. Right and left plates of phallosome connected by abridge near the base, 
each plate broad and with denticles of different size on the surface and 
short, sclerotized processes directed anteriorly and inward. Tenth sternite 
with numerous spines in several rows at the apex. Lobes of 9th tergite 
weakly developed, with 4—6 hairs. 

Fourth-stage larva (Figures 231— 233) transparent but the brown 
fat body gives it an apparent transverse striation. 

Head 1.6—1.7 times wider than long. Antennae with spines, nearly as 
long as the head, with dark base and narrow apical part. Hair tuft with 

338 25—30 branches, subapical setae situated at the apex. Frontal hairs: outer 
hairs long, with 5—8 branches, median and inner hairs short, with 2—4 
branches, nearly as long as the postclypeal hairs. Comb with 34—46 narrow, 
elongate scales with a row of spines in the apical part. Siphon 5.5—6 times 

339 longer than wide at the base, slightly widened at the apex. Pecten with 
10—13 regularly spaced denticles in the form of thin spines with a row of 
thin, pointed accessory denticles their entire length. Eleven to 13 hair 



363 



tufts, situated in a zigzag row on the posterior surface, a paired arrange- 
ment is exceptional. Tufts with 3 — 6 smooth branches, longer than the width 
of the siphon at their position. Main tracheal trunks thin, with round cross 
340 section. Stigmal plate (Figure 233) resembling that of C. (N). territans 
but differing in some details (presence of a distally directed chaetoid at the 
end of the posterior process, structure of the organs on the lateral valves, 
etc.). Saddle of last segment ring-shaped. Lateral hair short, with 3—4 
branches. Outer caudal hairs as long as the siphon, inner hairs shorter, 
with 2—5 hairs which are shorter dorsally. Fin with 12—13 tufts. Gills 
lanceolate, usually as long as the saddle, lower pair longer than the upper. 



(33 7) 




FIGURE 230. Hypopygium of C ul ex hayashii Yam. (after La Casse and Yamaguti) 



364 



(338) 




FIGURE 231. Culex hayashiiYam. Fourth-stage larva. Distribution of pigment in the fat body: 

A - thorax and first two abdominal segments (d — dorsal and v — ventral); b - abdominal segments 1—8, lateral. 




FIGURE 232. Culex hayashii Yam. Fourth-stage larva. Head, dorsal. 



365 



(339) 




FIGURE 233. Culex hayashiiYam. Fourth-stage larva: 
1 - posterior end, lateral; 2 — stigmal plate. 



366 



Distribution. USSR : southern Maritime Territory. Korea, China, 
Japan (Hokkaido, Honshu, Kyushu). 

Biology. The larvae usually occur in small numbers from the end of 
spring to autumn, in shallow sunlit water bodies with decomposing leaves 
and with vegetation, also in reservoirs and cisterns together with A. hyr - 
canus, A. communis (spring), C. modestus, C. bit aeniorhyn - 
chus, C. t r it aenio rhyn chu s, C. infantulus Edw. and C. vishnui 
Theob. They are found only in thinned forests in the foothills. They are 
not important bloodsuckers. A rare species. 



4. Subgenus C u 1 e x Linnaeus 

Coxite without scales; style usually simple, without subapical row of 
spines; 10th sternite with a tuft of dense hairs or spines at the apex and 
with a more or less developed appendage at the base in most species; 
phallosome of complicated structure, often with denticles or processes. 
Ninth tergite forming a narrow transverse stripe. Tarsi usually dark, 
rarely with light rings, which sometimes extend to the apex of the segment 
and the base of the next. 

The larvae of the species of this subgenus vary in structure and can 
sometimes be characterized only by negative characters: absence of mouth 
parts adapted to predacious feeding, arrangement of the scales of the comb, 
which never form a regular row. The arrangement of the adults in groups 
does not correspond to the grouping of the larvae, which have not been 
sufficiently studied. The larvae of some species resemble those of species 
of Barraudius, and the larvae ofC. mimeticus resemble those of 
the subgenus Neoculex. 

The subgenus contains over 180 species which are divided into two 
groups according to the chaetotaxy of the pleurae of the thorax, and the 
structure of the hypopygium. The first group includes all Palaearctic 
species with light rings on the tarsi which are mainly Oriental and only a 
few species occur in Africa. This group is distributed mainly in the south- 
eastern Far East of the Palaearctic (Maritime Territory) and a few species 
occur in the south (C. mimeticus). The Palaearctic species of the 
second group have dark tarsi without rings and are more closely related 
to the Ethiopian species.* This group attains the highest specific diversity 
and many species are very closely related. C. p i p i e n s, which occurs in 
many parts of the USSR, belongs to this group. 



9. Culex (Culex) bitaeniorhynchus Giles, 1901 

Closely related to C. sinensis, but differs from it in the presence of 
light and dark scales on the wings and in the structure of the hypopygium. 

Head with narrow, light golden, contiguous and upright, forked scales 
in the middle of the vertex; lateral margins of eyes with broad, light, con- 
tiguous scales. Palps of female short, % as long as the proboscis, with 

* This group contains numerous neotropical species, which resemble the Ethiopian species of the group. 



367 



342 



dark scales; apex of palps of female with light scales. Palps of male 
distinctly longer than the proboscis (nearly by the length of the 2 terminal 
segments); terminal segments with a ring of light scales at the base; 
long segment of palps with 2 light rings; terminal part i 1 ^ — %) of apical 
segment light. Proboscis with a white ring beyond the middle. Mesonotum 
dark brown, with light yellow or golden, narrow scales in the anterior two 
thirds; two distinct, rounded spots of dark scales near the middle of the 
mesonotum with light scales; posterior third of mesonotum with dark 
scales". Pleurae of thorax with white scales on sternopleura and mesepi- 
mera. Legs brown; femora and tibiae with brown scales and with more or 
less numerous scattered white scales anteriorly; and mainly with white 
scales posteriorly; tarsi dark brown, with white rings at the base of tarsal 
segments 1—4. Wings with brown and yellowish scales. Abdomen brown, 
with more or less broad stripes of white scales at the apex of the tergites, 
sometimes also with indistinct, but much narrower stripes at the base of the 
tergites. 



'341' 




FIGURE 234. Hypopygium of C ul ex b i ta e nio rhy nchus Giles: 

A - inner sclerites; B - coxite and style; C — basal lobe of coxite; 
D — second part of phallosome. 



368 



Hypopygium (Figure 234) characteristic; plate on subapical lobe of 
coxite narrow, knife -shaped; phallosome with two, pointed, sickle -shaped 
plates at each side; at the sides of the phallosome is a process with 4 or 
5 finger-shaped denticles; basal process of 10th sternite weakly developed. 




FIGURE 235. Culex b i ta e ni orhy nchus Giles. Fourth-stage larva: 
1 — head; dorsal; 2 — mentum. 



Fourth-stage larva (Figures 235 and 236) at first transparent, 
later green or brownish green. Head small, relatively narrow. Antennae 
situated anteriorly, rod -shaped, narrower from the middle, from the hair 
tuft, which has 15—25 branches; subapical setae directed inward, situated 
at the apex. Mouth parts adapted to feeding on filamentous algae. Lateral 
lobes of labrum with fewer than usual, curved hairs; mandibles and 
maxillae also modified; mentum forming an isosceles triangle with very 
small, regular incisions at the margin. Frontal hairs secondarily feathered, 
outer hairs with 4—7 branches, median hairs longer, with 2 branches, 
inner hairs with 3 branches. Comb with 4—7 large, spine -shaped scales. 
Siphon long, thin, 6—8 times as long as wide at the base. Pecten with 7—9 
denticles at the base of the siphon. Four to 5 pairs of hair tufts on the 



369 



posterior surface arranged in a zigzag row, each with 2-5 short branches, 
344 shorter than the width of the siphon. Saddle surrounding last abdominal 
segment like a ring. Lateral hair short, with 1-3 branches. Outer caudal 
hairs simple, shorter than the siphon, inner hairs with 3-4 branches. Fin 
with 12 tufts. Gills slightly longer than the saddle, lanceolate. 



(343) 




FIGURE 236. Culex b i ta eniorhy nchus Giles. Fourth -stage larva. Posterior end, lateral. 



Distribution. USSR : southern Maritime Territory, to the Iman River 
in the north, where it is widespread in lowland and coastal areas. Japan, 
to Honshu, Korea, China, Southeast and South Asia, Africa, Northern Australia. 



370 



345 



Biology. The larvae occur in large numbers in open, sunlit, stagnant 
or slowly flowing, pure water with abundant vegetation and filamentous 
algae. They are found together with larvae of A. hy re anus, rarely 
with those ofC.tritaeniorhynchus, C. vishnui, C.hayashii and 
others. They are active bloodsuckers which bite mainly at night in 
rice fields. 

Epidemiological importance. According to Japanese authors 
(Yamaguti and La Casse, 1955) it is considered as a vector of Japanese 
encephalitis. Specimens of this species have been infected with this virus 
(Petrishcheva and Shubladze, 1940). 



10. Culex (Culex) sinensis Theobald, 1903 



(344) 




FIGURE 237. Hypopygium of C ul ex sinensis Theob. (after La Casse and Yamaguti) 



371 



It resembles C. bitaeniorhynchus in coloration but differs from it 
in the brown scales on the wings (without light scales) and in the structure 
of the hypopygium. 




FIGURE 238. Culex sinensis Theob. 
Mentum of larva (after Barraud) 



The white scales on the legs usually form small, slightly irregularly 
arranged spots (not a regular row of spots). Tarsi with narrow white 
rings at the base of the segments; abdomen with stripes of light scales at 
the base of the tergites. 

Hypopygium (Figure 237) resembling that of C. bitaeniorhynchus; 
plate of subapical lobe of coxite narrow, lanceolate; phallosome with a pair 
of simple (not segmented or serrated), slightly curved plates; basal pro- 
cesses of 10th sternite large, markedly sclerotized. 

Larva resembling that of C. bitaeniorhynchus, differing as 
follows: subapical setae of antenna situated distinctly before the apex; 
mentum (Figure 238) with larger denticles in the middle; comb with 4—6 
large scales with a pointed spine; pecten with 2—4 small denticles at the 
base of the siphon; a large, slightly curved, spine -shaped hair on the anterior 
surface near the apex of the siphon; gills 2.5 times longer than the saddle, 
inner caudal hairs with 2 branches. 

Distribution. Mainly in the Oriental region. USSR: southern 
Maritime Territory. Japan (Honshu, Kyushu), Korea, China, Southeast and 
South Asia. 

Biology. The larvae occur in rice fields and on swampy meadows. 



11. Culex (Culex) whitmorei Giles, 1904 

The species has a distinctly two-colored mesonotum which is covered 
with white scales in a large part, posterior third with dark scales; probos- 
cis and tarsi with white rings; wings with broad dark scales on the anterior 
veins and on the cubitus, with a spot of dark scales at the base of V4 45. 



372 



346 




FIGURE 239. Hypopygium ofCulex whitmerei Giles (after La Casse and Yamaguti) 



Greater part of head with white scales. Proboscis dark brown at the 
base and in the apical part, with a distinct yellowish white ring in the middle. 
Palps of female % as long as proboscis, with brown and white scales; base 
of palps and base of 3rd segment of palps with white scales; palps of male 
1.5 times longer than the proboscis, blackish brown; apex and base of 5th 
segment of palps and base of 4th segment with white scales; the more or 
less broad but indistinct ring in the middle of the 3rd segment and apex 
of 2nd segment light; hairs on palps numerous, long, dark brown. Mesono- 
tum brown, its anterior two -thirds with white scales, with a golden, kidney - 
shaped spot or 3 small spots near the middle; posterior third of mesonotum 
with dark golden scales and setae and with some white scales. Legs: 
femora and tibiae mainly with brown scales and a few white scales 
anteriorly; tarsi with narrow white rings at the base of segments 1—4. 
Wings with brown scales; costa, subcosta, r^ r 4 -+s and cubitus with broad, 



373 



dark scales. Abdomen with light stripes or triangular spots of white 
scales which extend posteriorly from the base of the tergites; tergites5— 8 
with more or less large spots of white scales at the sides; ventral side 
brown, with s tripes of light scales. 
347 Hypopygium (Figure 239): subapical lobe of coxite with a broad plate, 
2 rod -shaped appendages, one of them thin and pointed and the other thick 
and blunt, and with setae; 10th sternite with a short, narrow basal process; 
apical part of phallosome with well developed median part which forms a 
hook and a broad, triangular lateral plate; lower part of phallosome 
normally with 4 denticles of more or less the same size. 





FIGURE 240. Culex w hi tm orei Giles. Fourth -stage larva 
(after Barraud): 

1 — siphon, lateral; 2 — antenna. 



Fourth-stage larva (Figure 240): head nearly 1.5 times wider 
than long. Frontal hairs secondarily feathered, outer hairs with 6—10 
branches, median hairs situated slightly posterior to and inward of the 
outer hairs, both pairs long, 2 -branched, one of them rarely 3 -branched. 
Postclypeal hairs short, thin, 2 -branched, rarely simple. Sutural and 
transsutural hairs slightly longer, with 3 or more branches. Antennae as 
long as the head, covered with spines. Hair tuft with 10 or more branches, 
situated beyond the middle of the antenna, which is narrow distal to the 
tuft. Two subapical setae. 

Comb with 5—8 scales with a large main spine in an irregular, zigzag 
row, lateral spines at base of scales weakly developed, practically absent. 



374 



Hairs behind the comb: dorsal hair with 3—5, ventral hair with 2—3 branches, 
median hair with 6—8 branches, intermediate hair simple. Siphon wider 
at the base and then tapering, index 5.0—6.0. Pecten with 6—10 thin denticles, 
the 1—2 distal denticles larger and more widely spaced, occupying the basal 
part of the siphon, not reaching third of siphon from the base; 4—6 pairs of 
hair tufts in one row in the middle of the posterior surface of the siphon. 
Tufts with 2 long, slightly secondarily feathered branches and also with 
2 pairs of much shorter, 2 -branched tufts on the lateral surfaces, one in the 
middle, the other distal to the distal ventral tuft. Main tracheal trunks 
thin, with round cross section. 

Last segment distinctly longer than wide; saddle surrounding the seg- 
ment like a ring. Lateral hair simple, 2 -branched. Caudal hairs longer 
than the siphon, simple or 2 -branched. Fin with 12 tufts. Gills lanceolate, 
of varying length, as long as the saddle to as long as the siphon. 

Distribution. Southern Maritime Territory. Japan (Kyushu, Honshu — 
rare), Korea, East and Southeast Asia. 

Biology. Not studied in detail. The larvae occur in stagnant or slowly 
flowing water, in rice fields, etc. Mass attacks on man and domestic 
animals have not been observed. 



12. Culex (Culex) mimeticus Noe, 1899 

A very characteristic species, with spotted wings, slightly resembling 
A. superpictus (Figure 241) in habitus. 

Head with yellowish white and brown scales. Proboscis blackish brown, 
with a broad white ring near the middle in females which is slightly nar- 
rower in males. Palps of female with blackish brown scales, apex of palps usu- 
ally with white scales; palps of male blackish brown with blackish brown scales, 
usually with 3 white rings; proximal ring on the long segment of the palps 
distinctly wider than the other rings; apex of palps of male light. Antennae 
brown in both sexes, with light rings in the male. Mesonotum with light or 
golden brown scales in the middle, grayish or whitish scales at the margins; 
pleurae of thorax with spots of white scales. Legs: femora and tibiae with 
blackish brown scales anteriorly, often with some white scales which some- 
times form a narrow longitudinal stripe, with white scales posteriorly; 
tarsi with narrow white rings at the base of segments 1—4; rings partly 
extending to apex of preceding segment. Wings with blackish brown scales 
on the veins, with 3 large white spots at the anterior margin, situated at the 
beginning and end of the median third of the anterior margin and near the 
apex and occupying veins c, sc and ri; larger parts with white scales at the 
base of the forks r 2 + r 3 and mi + m 2 and on r 4 + 5 (this vein is covered with 
white scales in a large part), m 3 + 4 and an; cui only with blackish brown 
scales. Abdomen dark brown, with black scales dorsally; tergites with 
transverse stripes of white scales at the base, usually a third as wide as 
the tergite, ventral side with white scales. 

Hypopygium (Figure 242): coxite with relatively short hairs; subapical 
lobe of coxite with a broad, leaf-shaped, platelike appendage and usually 
with 5 large setae; style tapering apically; 10th sternite with a basal ap- 
pendage; lower part of phallosome with 3—5 laterally directed denticles. 



375 



(349) 




FIGURE 241. Culex mimeticus Noe 



610144 



376 



(350) 




FIGURE 242. Hypopygium of C ul ex mimeticus Noe 



FIGURE 243. Culex mi- 
meticus Noe. Fourth- 
stage larva. Siphon, lateral 



Larvae (Figure 243) resembling those of the subgenus Neoculex in 
many characters. 

Head very broad, often dark; median frontal hairs situated before the 
inner hairs, more widely spaced; both pairs with 2 secondarily feathered 
branches. Antennae light, apex darker, with spines for 2 / 3 from the base; 
hair tuft with numerous secondarily feathered branches, half as long as 
the antenna, situated in the middle or nearer to the apex. Subapical setae 
as long as the tuft, situated in middle of apical narrow part of antenna. 
Abdomen: comb with 20—35 scales, with a distinct, pointed main spine and 
a row of small spines at the sides of the base. Siphon straight, wider at 
the base and tapering apically; index 4.5— 7, varying according to the width 
of the base. Pecten about x / 3 as long as the siphon, with 12—13 denticles, 
the 1 — 2 distal denticles may be larger and more widely spaced. Five to 
6 hair tufts, the first 3—4 pairs longer, with 4—5 branches, often arranged 
in a zigzag row, irregularly spaced; the next pair of tufts with 2 thin, 
shorter branches, situated more laterally. 



377 



Main tracheal trunks thin, with round cross section. Stigmal plate of 
the C. territans type. Last segment with simple outer caudal hairs 
about as long as the siphon, inner hairs with 2 branches, the dorsal branch 
half as long as the ventral and with a short, 2 -branched lateral hair. Fin 
with 12 tufts. Gills 1.5—2 times as long as the saddle, pointed at the end. 

Distribution. South of the Palaearctic and Oriental region. USSR : 
Northern Caucasus, Dagestan, Transcaucasia, Middle Asia (Turkmenia, 
Tadzhikistan), Maritime Territory. Southern Europe, Asia Minor, Iran, India, 
China, Japan and other countries in Asia. 

Biology. The larvae occur in small pools in crevices in rocks and in 
backwaters of rapidly flowing mountain streams overgrown with vegetation, 
sometimes together with larvae of A. marteri, A. claviger, 
A. superpictus and Culiseta longiareolata. 



13. Culex (Culex) jacksoni Edwards, 1934 

The adults closely resemble C. mimeticus, but the larvae differ 
distinctly. 
351 According to the description, the mesonotum is covered with reddish 

brown scales, with lighter scales at the sides and in the area before the 
scutellum. Otherwise, including the structure of the hypopygium, as in 
C. mimeticus. 

Fourth-stage larva greenish (young) to yellowish brown (older), 
medium-sized (to 1 cm). 

Head about 1.5 times longer than wide. Frontal hairs with well devel- 
oped secondary feathering, outer hairs with 6—12 (average 8) branches, 
longer than the median and inner hairs, with 2 — 6 (average 3) and 3 — 7 
(average 5) branches respectively. Postclypeal hairs short and thin, with 

2 — 3 (1 — 5) branches; sutural and transsutural hairs slightly longer, with 

3 — 4 (1 — 5) thin branches. 

Antennae slightly shorter than the head, only slightly curved, weakly pig- 
mented at the base and with dense, thin, hairlike setae; distal part dark, 
narrower, with sparse, short, thick spines. Hair tuft forming a fan with 
11 — 26 (average 18) secondarily feathered branches reaching to the end of 
the antenna. Two long subapical setae, the apical setae as long as these 
but slightly thinner. 

Comb of 8th abdominal segment with 19—40 (average 32) pointed scales 
which form an irregular spot. Scales with a row of marginal spines at the 
sides of the base, spines near the main spine shorter than this. Hairs 
behind comb: dorsal hair with 5, ventral hair with 4, median hair with 7 
secondarily feathered branches; intermediate hair smooth and simple. 

Siphon long, slightly wider at the base (index about 6). Pecten with 
8—20 (average 14) denticles, 2 — 4 rudimentary denticles at the base. The 
denticles are thin spines with several accessory denticles at the slightly 
widened base; all denticles weakly pigmented. Pecten occupying 1 / 3 to 
half of the length of the siphon, rarely more. One to 5, often 3 of distal 
denticles longer and more widely spaced. At the posterolateral surface 
of the terminal quarter of the siphon are additional, larger, smooth spines, 
directed apically in the continuation of the pecten. This is a distinctive 
character of the larvae. The number of these spines varies from 9 to 2 



378 



(average 4, i. e., two pairs). If there are fewer spines, some may be 
rudimentary: a small, short spine on a broad base. 

The number of hair tufts on the siphon is relatively constant: 3 pairs on 
the posterior surface in an irregular zigzag row; 1-2 basal tufts are situ- 
ated near or between the 1-2 distal denticles and the apical tufts are 
situated between the additional spines. Tufts short, usually as long as the 
width of the siphon at their position, with 2-6 smooth branches. Two pairs 
of short, 2 -branched (rarely simple) hairs on the lateral surface of the 
terminal quarter of the siphon. 

Main tracheal trunks broad, ribbon-shaped. Stigmal plate of the 
"pi pi ens" type. 

Last segment short, only slightly longer than wide. Saddle surrounding 
the segment like a ring. Lateral hair thin, short 2-3 -branched. Caudal 
hairs: outer hairs simple, usually as long as the siphon, inner hairs % as 
long, with 2, rarely 3 branches, rarely simple. Fin situated toward 
posterior margin of segment, with 12 tufts. Gills lanceolate, pointed at 
352 the ends, both pairs of nearly the same length, 1.25-1.5 times as long as 
the saddle, rarely longer or shorter. 

Biology. The larvae were found in shallow swamps with filamentous 
algae (Suchans District; A. I. Lisova) and in small pools and backwaters of 
streams (Furugel'ma Island; V. K. Trasis), mainly in foothills, together with 
larvae of A. hyrcanus, A. vexans, C. b i t a e n i o rh y n ch u s, 
C. mimeticus, C. modestus, C. vagans and others. 




FIGURE 244. Hypopygium of C u 1 e x orientalis Edw. 



379 



14. Culex (Culex) orientalis Edwards, 1921 

Closely related to C. mimeticus, but differs from it as follows: 
wing scales longer and the wings therefore appear more densely covered 
with scales; cui with light scales at the apex; the part covered with light 
scales passes over to the stem of m 3+ 4 + cui; palps of male with numerous 
light scales at the apex which are sometimes more numerous than the dark 
scales. 



(353] 




FIGURE 245. Culex orientalis Edw. Fourth-stage larva. Head, dorsal. 



Hypopygium (Figure 244): coxite broad, slightly thickened, densely 
covered with hairs, especially in the apical part near the subapical lobe, 
which bears a transparent plate and a row of spines which are slightly 
denticulate or curved at the apex and surrounded by long hairs; style 



380 



353 



markedly widened in the middle, pointed apically; second part of phallosome 
with 3 large and several small denticles; the hypopygium differs from that 
of C. mimeticus in the presence of dense hairs in the apical part of the 
coxite. 

Fourth-stage larva (Figures 245— 247) semitransparent, becoming 
yellowish or green before pupation. 



(354) 




FIGURE 246. Culex orientalis Edw. Fourth-stage larva. Posterior end, lateral. 



Head broad, light yellow. Frontal hairs long, secondarily feathered, 
outer hairs with 5—10 branches, median and inner hairs displaced posteri- 
orly, the median hairs more widely spaced, with 2 branches, inner hairs 
with 3—4 branches. Antennae long, more than 3 / 4 as long as the head, 
pigmented at the base and in the apical part; tuft with 20—35 secondarily 
feathered branches, half as long as the antenna, situated distinctly beyond 
the middle; subapical setae longer than tuft, situated at % of the length of 
the terminal part from the apex, apical seta shorter. 



381 



354 



(355) Comb with 20-40 (average 30) scales, 

with a row of spines at the margin which 
are distinctly larger at the apex. Siphon 
long, thin (index 6.2—7.9). Number and 
arrangement of hair tufts varying; 10— 
13 tufts on the posterior side arranged 
in pairs or in a zigzag row; basal tufts 
longer, slightly longer than the width of 
the siphon, with 2—5 branches; at the 
lateral surfaces, near the apex, are 1— 
2 pairs of tufts; all tufts near the apex 
short. Pecten with 9—14 denticles, 1 — 
4 rudimentary denticles at the base. 
Main tracheal trunks with oval cross 
section. Stigmal plate of the C.pipiens 
type. Last segment with long, simple 
outer caudal hairs (1.3 — 1.5 times as 
long as the siphon), inner hairs %— x / 3 
shorter with 2—4 branches. Fin with 
11 — 13 tufts. Gills lanceolate, as long as 
or longer than the saddle (to 1.7 times 
its length); lower pair shorter. 

Distribution. East Asia. USSR : 
Khabarovsk and Maritime territories 
but not in the mountain -taiga. Japan, 
Korea, China, Philippines. 

Biology. The larvae occur mainly 
in sunlit water bodies with rich vegeta- 
tion and algae, in ditches and rice fields, 
often together with larvae of A. hyr- 
canus, C. b i t a e n i o r hy n chu s, 
C. t r i t a e n i o rh y n chu s, A.vexans 

and others. Two, rarely three, generations per year. They have not been 

reported to bite man or domestic animals. 




FIGURE 247. Culex or ie nt a li s Edw. 
Fourth-stage larva. Variation of arrangement 
of hairs on the siphon. 



15. Culex (Culex) tritaeniorhynchus Giles, 1901 

The species is characterized by a light ring on the proboscis, light rings 
on the tarsi and the absence of light scales on the wings (Figure 248). 
355 Head with reddish brown scales. Proboscis with a white ring which 

sometimes extends nearly to the base in the female, particularly on the ventral 
surface. Palps of female dark, palps of male with 2 rings on the long seg- 
ment, one ring distinctly wider than the other; apex of palps of male dark. Me - 
sonotum with dark reddish brown scales. Femora and tibiae with brown scales; 
ventral surface of femora with whitish scales. Tarsi brown, with narrow, 
white rings at the base of the segments; rings sometimes indistinct, more dis - 
tinct on the fore tarsi. [Abdomen with dark scales dorsally and with light 
stripes at the base of the tergites.] 

Hypopygium (Figure 249) resembling that of C. m i m e t i c u s. Subapical 
lobe of coxite with a leaf -shaped plate and 3—4 spines. Lower part of phallosome 
with 4—5 denticles on each side, some of them (usually the median) directed 



382 



(3 56 



posteriorly (in distinction to C.mimeticus), Basal appendages of 10th 
sternite well developed. 

Fourth-stage larva (Figure 250) nearly transparent, weakly 
pigmented. 

1 




FIGURE 248. Culex tr ita enio rhy nchus Giles 



383 



(3 57) 




FIGURE 249. Hypopygium of C u 1 e x t ri ta e ni orhy nchus Giles: 

A — inner sclerites; B — coxite and style; C — basal lobe of coxite; D — lobe 
of 9th tergite. 



(357 




FIGURE 250. Culex t r ita e nio rhy nchus Giles. Fourth -stage larva 
(after Kirkpatrick): 

1 — head, dorsal; 2 — posterior end, lateral. 



384 



Head moderately broad, 1.3 times wider than long. Frontal hairs long, 
secondarily feathered, outer hairs with 6—8, median hairs with 2, inner hairs 
with 3 branches, situated more closely together than the median hairs. 
Antennae about 0.7 times as long as the head, dark at the base and with narrow- 
ing terminal part; tuft situated at about % of the length of the antenna from 
the base, with about 30 secondarily feathered branches. Subapical setae 
longer than the tuft, situated at about % the length of the narrow apical part 
before the apex; apical seta shorter. 

Comb with about 50 short scales with a few spines at the margin which 
are longer at the apex. Siphon long (index 6.5—7.8) thin, with distinctly 
widened base; pecten with 9-15 denticles, the 2-3 distal denticles more 
widely spaced, with 5—9 thin, pointed accessory denticles on the entire 
ventral margin which are longer toward the apex. Five pairs of hair tufts 
at the sides of the posterior surface beginning at its second third from the 
base. Tufts with 3—4 branches (rarely 2, near the apex), not longer than the 
width of the siphon except the basal tufts; one pair of shorter, 2— 3 -branched 
lateral tufts in the apical third of the siphon. Tracheal trunks with oval 
cross section. Stigmal plate of the C. pipiens type. Last segment of 
abdomen with simple outer caudal hairs which are longer than the siphon, 
inner hairs with 3 branches, the lower as long as the siphon and each follow- 
ing branch half as long as the preceding, lateral hair short, 2— 3 -branched. 
Fin with 12 tufts. Gills short, half as long as the saddle, pointed at the end, 
upper pair longer than the lower. 

Distribution. The wide distribution includes the Oriental region, 
southern and southeastern parts of the Palaearctic and a large part of the 
Ethiopian region. It is found in the USSR in the southern part of Azer- 
baidzhan, in Turkmenia and Uzbekistan, and in the south of the Maritime 
Territory (to the Iman River in the north.). It also occurs in Turkey, Iran, 
China, North Korea and Japan. 

Biology. The larvae occur mainly in shallow, well warmed water 
bodies, which are sometimes only periodically flooded, and also on oxbow - 
lakes, where they are usually found only along the banks without vegetation. 
They usually inhabit meadow pools only after the hay harvest when the 
water is warmer. Their numbers are highest from the middle of July to 
the end of September. Active bloodsuckers, biting mainly in the open, 
usually from sunset to midnight or one in the morning and at 0600—0800 hr. 
In bad weather they may attack for days. They appear in inhabited areas 
near the breeding places and often enter houses and cattle sheds. 

Epidemiological importance. This species is one of the 
main vectors of Japanese encephalitis. Transmission of the virus by bite 
and natural infection have been proved. 



16. Culex (Culex) univittatus Theobald, 1901 (perexiguus Theobald, 1903) 

It differs from related species in its small size, narrow, white, trans- 
verse stripes on the abdomen and in the presence of a more or less distinct 
white stripe on the anterior (outer) surface of the hind tibiae. 

Head dorsally with brown and golden scales; white scales at the margin 
of the eyes. Proboscis brown, sometimes with white scales in thebasalhalf. 



385 



Palps with brown scales, sometimes with an admixture of white scales at 
the apex in females. Mesonotum with golden brown scales, its lateral 
parts, the area before the scutellum and the scutellum with gray scales; 
pleurae of thorax with spots of white scales. Legs with brown scales; 
femora ventrally, hind femora also anteriorly in the basal half with white 
scales; hind tibiae and sometimes also mid -tibiae with a more or less 
distinct white longitudinal stripe on the outside; tarsi blackish brown. 
Veins of wing with brown scales. Abdomen dark brown and with dark 
brown scales; narrow transverse stripes of white scales at the base of 
the tergites. 




FIGURE 251. Hypopygium of Culex univittatus Theob. 
A — coxite and style; B — inner sclerites. 



359 



360 



Hypopygium (Figure 251): subapical lobe with a broad, asymmetrical 
plate; style distinctly widened beyond the middle; basal appendage of 10th 
sternite well developed; second part of phallosome broad, simple. 

Fourth-stage larva (Figure 2 52) weakly pigmented, nearly trans- 
parent. Head 1.4 times wider than long. Frontal hairs secondarily 
feathered, outer hairs with 7—9, median hairs with 2 — 3, inner hairs with 
3—4 branches; median hairs situated before the inner hairs, more widely 
spaced. Antennae less than 3 / 4 as long as head, with a tuft of 24—28 second- 
arily feathered branches, situated at % — % of the length of the antenna 
from the base. Subapical setae situated near the apical seta. 

Comb with 30—40 narrow, elongate scales, their distal end with a row 
of dense spines at the margin. Siphon (index 5.9—7.6) of varying length, 
the larger index corresponds to larger specimens. Pecten with 10—12 
large denticles, occupying the basal x / 4 of the siphon; the distal denticles 
may be widely spaced. Five pairs of tufts at the posterolateral surface 
of the siphon, with 2—4 branches, not longer than the width of the siphon; 



386 



the length of the tufts and their branching decrease toward the apex. The 
tufts are situated gradually more laterally toward the apex. One of the two 
tufts near the apex is situated on the lateral surface of the siphon and the 
other on the posterior surface. Main tracheal trunks broad, with oval cross 
section. Stigmal plate of the C. pipiens type. Last abdominal segment 
with simple outer caudal hairs which are as long as the siphon, inner hairs 
with 2 branches, lateral hair with 2-3 branches. Fin with 12 tufts. Gills 
0.75—1.5 times as long as the saddle, oblong, ovoid. 



(3 59) 




FIGURE 252. Culex univittatus Theob. Fourth-stage larva (after Edwards): 
1 — head, dorsal; 2 - posterior end, lateral. 



387 



Distribution. Ethiopian region and Mediterranean, Iran, Pakistan, 
India. USSR: Turkmenia. 

Biology. The larvae occur in stagnant water bodies with fresh or 
slightly brackish water with up to 0.2% salt and a pH of 7.2—9.7. They 
are often found together with larvae of C. laurenti Newst., more rarely 
with those of C. pi pi e ns, A. caspius and A. pharoensis Theob. 
The females feed mainly on birds. 

Epidemiological importance. A vector of West Nile virus. 




361 



FIGURE 253. Hypopygium of Culex 
theileri Theob. Inner sclerites. 



17. Culex (Culex) theileri Theobald, 1903 

A large species with light longitudinal stripes on the anterior surface 
of femora and tibiae. 

Head with white or yellowish contiguous 
scales and brown upright scales. Probos- 
cis with brown scales and with more or less 
numerous white scales. Palps of female with 
brown scales, with a few white scales 
dorsally. Palps of male with brown scales, 
with more or less numerous spots of white 
scales which sometimes form rings. Meso- 
notum with brown scales, strongly varying 
in coloration; area before the scutellum with 
whitish scales. Legs: femora and tibiae 
with brown scales anteriorly and a distinct 
longitudinal stripe of white scales, with white 
scales posteriorly. Wings with brown scales. 
Abdomen with blackish brown scales, 
with a stripe of white scales at the base 
of the tergites usually triangularly 
produced posteriorly so that they form a 
light median longitudinal stripe with light scales ventrally. 

Hypopygium (Figure 253: subapical lobe of coxite with a relatively broad, 
elliptical plate; style sickle -shaped; 10th sternite with a well developed 
basal appendage; phallosome with numerous small denticles. 

Fourth -stage larva (Figure 254) transparent or yellowish, often 
with a dark head. Frontal hairs secondarily feathered, long, outer hairs 
with 7—9 branches, median hairs with 2—3 branches, situated before the 
inner hairs and more strongly branched, inner hairs with 3—5 branches. 
Antennae % as long as head, with darker base and apical part. Hair tuft 
with 25—30 branches; subapical setae situated near the apex, as long as the 
tuft and the apical seta. Comb with 30 or more scales, with weakly devel- 
oped apical spine. Siphon nearly straight, tapering apically, index 5.5—6. 
Pecten extending slightly beyond basal quarter of siphon, with 6—11 widely 
spaced, spine -shaped denticles, with 3—5 short accessory denticles at the 
base. Five pairs of hair tufts, beginning at the second third from the base: 
3 basal pairs longer, with 4—8 branches, situated on the posterior surface of 
the siphon, often in a zigzag row, 4th pair situated on the lateral surface, 
with 2—4 shorter branches, the 5th pair situated near the apex, on the 
posterolateral surface, with 2—4 short branches. Main tracheal trunks with 



388 



oval cross section. Stigmal plate of the C. pipiens type. Last abdominal 
segment with simple outer caudal hairs, which are longer than the siphon, 
inner hairs with 4 branches, lateral hair short, simple. Fin usually 
with 14 well developed tufts. Gills as long as or slightly longer than the 
saddle, with pointed end, both pairs of about the same length. 




FIGURE 254. Culex theileri Theob. Fourth-stage larva (after Edwards): 
1 — posterior end, lateral; 2 — head, dorsal. 



Distribution. Ethopian region, Mediterranean and western parts of 
the Oriental region (India). USSR: Ukrainian steppes, Crimea, Caucasus 
and Transcaucasia, Middle Asia. 

Biology. The larvae occur in stagnant water bodies and ditches 
overgrown with vegetation, often in strongly polluted water, usually in fresh 
or slightly saline water (to 0.2%), but the salinity is often 0.5— 1.0%, pH 9.5). 
They occur in spring on flooded meadows. The mosquitoes bite mainly in 
the open, sometimes in large numbers (for example, in some parts of 
Armenia), sometimes entering houses and other buildings. 



389 



18. Culex (Culex) vagans Wiedemann, 1828 (exilis Dyar, 1924) 

It resembles Ctheileri to some extent, specimens with a weakly- 
developed light stripe on the fore and mid -femora and tibiae resemble 
C. pipiens. It differs distinctly from both species in the structure of the 
hypopygium. C, exilis Dyar, 1924, described from Vladivostok, is a 
synonym of C. vagans Wied. 




FIGURE 255. Hypopygium of C u 1 e x vagans Wie. 
A — coxite and style; B — inner sclerites. 



Head with yellowish or light brownish scales. Proboscis brown, 
usually with some light scales in the middle; palps of female dark, with 
a few light scales dorsally; palps of male longer than the proboscis by 
about the length of their last segment; long segment of palps of male dark, 
with some light yellow scales; terminal segments of palps not thickened. 
Mesonotum brown, with brownish scales with a bronze or yellowish tone, 
lighter laterally. Legs dark on the dorsal surface, light on the ventral 
surface; fore and mid -femora and all tibiae with a white longitudinal 
363 stripe anteriorly, rarely with a white longitudinal stripe also on the fore 
femora. Abdomen with dark scales and distinct stripes of yellowish 
white scales at the base of the tergites dorsally; the stripes are more or 
less triangularly produced posteriorly in some specimens, so that they 
resemble C.theileri. 

Hypopygium (Figure 255) with a relatively broad plate on the subapical 
lobe of the coxite; phallosome divided into 2—3 plates in the apical part, 
one plate directed distally; 10th sternite with a basal appendage. 

Fourth -stage larva (Figure 256) yellowish brown, with dark head 
and siphon. Head usually 1.5 times longer than wide. Frontal hairs 



390 



secondarily feathered, outer hairs with 6—11, median hairs with 4—6, inner 
hairs with 5—7 branches, both last pairs situated close together, the inner 
hairs more closely. Antennae about % as l° n g as the head, brown, darker 
at the base. Hair tuft situated at % to % of the length of the antenna from 
the base, with 25—30 branches; branches turned inward more strongly 
feathered. Subapical setae situated at the apex, slightly shorter than the 
apical seta. 




FIGURE 256. Culex vagans Wie. Fourth -stage larva 
(after Barraud): 

1 — siphon, lateral; 2 — antenna; 3 — last segment, lateral. 



Comb with 37—40 (24—47) scales with thin spines at the margin which 
are longer at the apex. Siphon (index 5—6) tapering from second third of 
antenna, slightly more than half as wide at the apex than at the base. Pecten 
with 7—13 denticles, 2 — 4 rudimentary denticles at the base, , occupying 
about x /3 °f the siphon, the distal denticles more widely spaced. Five pairs of 
of hair tufts, the 3 basal tufts longer, at least 1.5 times longer than the width of 
the siphon, with 3—4 (2—6 ) branches, situated on the posterolateral surface, 



39] 



the first pair often situated at the level of the distal denticle of the pecten, 
the 4th pair situated on the lateral surface, with 3 (2—4) shorter branches, 
5th pair on the posterolateral surface shorter than the others, with 3—4 
branches. Main tracheal trunks broad, with oval cross section. Stigmal 
plate of the C. pipiens type. Last segment of abdomen with simple 
outer caudal hairs which are longer than the siphon, inner hairs with 
2 branches, lateral hair with 2 branches. Fin with 12 tufts, slightly longer 
than the gills. Gills twice as long as the saddle, pointed at the end. 
364 Distribution. USSR : Transbaikalia, Khabarovsk and Maritime 
territories. Japan (Hokkaido, Honshu, Shikoku, Kyushu), Korea, China, 
India (mainly mountains and foothills). 

Biology. The larvae occur in natural and artificial stagnant water 
bodies (puddles, ditches, rain barrels, etc.) mainly near inhabited areas, 
so that they are distributed in virgin parts of the taiga. They rarely bite 
man. 



19. Culex torrentium Martini, 1925 

Martini described this species, but it is often named C. exilis (Dyar, 
1924); however, C. exilis was described from Vladivostok, where 
C. torrentium does not occur, as far as is known. C. exilis Dyar, 
as stated above, is a synonym of C. vagans Wied. The relationship of 
the two forms and its relation to the C. pipiens complex (see below) is 
not clear. The hypopygia of C. torrentium and C. vagans are 
practically indistinguishable, but they differ distinctly in coloration: 
C. vagans has longitudinal stripes of white scales on the anterior side 
of the femora and tibiae which are absent in C. torrentium, but this 
development ofthe stripes is variable. There are specimens with indistinct 
stripes which are almost indistinguishable from C. torrentium. C. tor- 
rentium should possibly be considered as a subspecies of C. vagans, 
particularly as both forms are similar ecologically. 

However, there is the question of its relation to C. pipiens. C. tor- 
rentium and C. pipiens differ distinctly in the structure of the hypo - 
pygium: the basal processes of the 10th sternite are well developed in 
C. t o r r e nt iu m . There are rarely some specimens with weakly devel- 
oped processes which are apparently intermediate between C. torrentium 
and C. pipiens. Other differences described (coloration of mesonotum 
-and abdomen, length of the stem of the radial fork) do not give reliable 
differences, to judge from our material. 

There are 3 possibilities: a) to consider pi pi e n s — t o r re nt iu m — 
vagans as the same species; b) to consider torrentium as a western 
subspecies of vagans; c) to consider them, as is usual, as 3 different 
species: C. pipiens (cosmopolitan), C. torrentium (Europe, Siberia) 
and C. vagans (Siberia, Far East ofthe USSR, China, Japan). We are 
inclined to accept the last view until the problem has been studied in detail 
(the study should be made in East Siberia, where apparently all three forms 
occur). 

C. torrentium closely resembles C. pipiens (see below) from 
which it differs only in the structure of the hypopygium. The body and 
scales of C. torrentium are darker; the mesonotum is covered with 



392 



yellowish brown scales, usually without the golden tone characteristic for 
C. pipiens. The femora and tibiae are dark brown anteriorly, without 
light scales. The scales on the abdomen are nearly blackish brown. The 
scales on the light stripes on the abdomen are more distinctly white than 
in C. pipiens. 

The structure of the hypopygium closely resembles that of C. vagans 
(Figure 255). The 10th sternite has a long basal process, the phallosome 
has 3 plates at the apex, one directed distally and the other two laterally. 

Other differences described make it possible also to distinguish the 
females: coloration of sternites, position of cross -veins, and especially, 
length of the radial fork: the stem of the fork is longer in C.torrentium, 
the fork 3—4 times as long as the stem; the stem is shorter in C. pipiens, 
the fork 5—6 times as long as the stem. Examination of a large material 
from different localities shows that these differences overlap and are not 
constant because of the wide variation of C. pipiens. In a specimen in 
the collection of the Zoological Institute, identified by Martini ("C. tor r en - 
tium, Saratov, April 1925, E. Martini"), the stem of the radial fork is 
very short and the specimen should therefore be identified as C. pipiens. 

Fourth -stage larva yellowish brown, with moderately broad head. 
Median frontal hairs situated before the inner hairs, much more widely 
spaced, usually with 5 branches. Antennae with a tuft of 20 and more 
branches which are % as long as the antenna. Subapical setae situated near 
the apical seta. 

Comb with 35—40 scales with a row of spines at the margin that are 
longer at the apex. Siphon (index 6—7) tapering from the second third. 
Pecten with 12 denticles, occupying about % of the length of the siphon from 
the base. Four pairs of hair tufts with 2—3 branches are situated on the 
posterolateral surface of the siphon, the apical pair is the most lateral, 
the first 3 pairs 1.5 times as long as the width of the siphon at the base, 
the 4th pair distinctly shorter. Tracheal trunks broad, with oval cross 
section. Stigmal plate of the C. pipiens type. Last abdominal segment 
with long, simple outer caudal hairs and 2 -branched inner hairs. Fin with 
11 — 12 tufts. Gills lanceolate, twice as long as the saddle. 

Distribution. Widespread in the USSR: Leningrad, Perm, Sverdlovsk, 
Orenburg regions, Urals, Lower Volga area, Crimea, Northern Caucasus, 
West Siberia and. other regions. Central Europe, Asia Minor. 

Biology. The species was confused with C. pipiens in the past 
because its ecology had not been studied in detail. The larvae occur in 
different types of water bodies. In the Lower Volga area, they are found in 
small water bodies with rich vegetation, in floodplains of rivers; in Central 
Europe (Black Forest) they occur in rock pools along mountain streams 
together with larvae ofCuliseta glaphyroptera. It is not 
synanthropic. It apparently bites man very rarely. 



20. Culex (Culex) pipiens Linnaeus, 1758 (Figure 257) 

Head with brown scales; lateral parts of head and posterior margin of 
eyes usually with white scales. Proboscis brown. Palps of female with 
brown scales, sometimes with white scales in the middle and at the apex. 



393 



366 




FIGURE 257. Culex pipiens L. 



394 



Palps of male distinctly longer than the proboscis, with brown scales; two 
terminal segments with white scales ventrally; dark brown, bristlelike 
hairs at the apex of segments 3—5 of palps well developed; more or less 
distinct rings of white scales on long segment of palps. Mesonotum with 
reddish brown or yellowish brown scales: lateral margins of mesonotum and 
area before the scutellum with lighter scales; pleurae of thorax with in- 
distinct spots of white scales on the proepisterna, sternopleura, and mese- 
pimera. Legs with brown scales, femora with yellowish white scales 
ventrally. Wings with brown scales. Abdomen with dark brown scales and 
more or less broad stripes of light yellow or whitish scales at the anterior 
margin of the tergites. These stripes are rarely absent. 




FIGURE 258. Hypopygium of C u 1 e x pipiens L. (after Coe, Freeman and Mattingly): 
A — inner sclerites of C. p. pipiens L.; B — same ofC.p. fatigans Wied. 



Hypopygium (Figure 258): subapical lobe of coxite with a broad plate 
with rounded apex; second part of phallosome simple, not divided, without 
denticles; basal process of 10th sternite absent or forming a small, rod- 
shaped or triangular process; it is rarely larger, but in distinction to 
C. t o r r e nt iu m, it is not curved or sclerotized. 

Fourth-stage larva (Figure 259) very variable in coloration, 
grayish yellow to yellowish brown; with yellowish or brown head and siphon. 
Medium-sized, rarely large. 



395 



(368) 




FIGURE 259. Culex pipiens pipiens L. Fourth-stage larva: 
1 — posterior end, lateral; 2 — stLgmal plate. 



Head 1.3 times wider than long. Frontal hairs well developed, second- 
arily feathered; outer hairs with 6 — 10 branches, median hairs situated 
before the inner hairs, more widely spaced, both pairs with 3 — 7 (average 4) 
branches. Postclypeal hairs short, simple, situated between the median 
frontal hairs. Antennae about 0.6—0.7 times as long as the head, more 
densely covered with spines in the basal part and with isolated spines in 
the apical third where the tuft is situated; tuft with 20—30 secondarily 
feathered branches, x / 3 as long as the antenna. Subapical setae situated 
near the apex, shorter than the tuft. 

Comb with 30—55 densely arranged oblong scales of varying size which 
form a half-round spot; scales without a main spine, with a row of spines 
at the margin and apex. Hairs behind the comb: dorsal hair with 3 — 7, 
ventral hair with 3—6, the longest median hair with 6—10 secondarily 
feathered branches; intermediate hairs shorter, simple. Siphon of varying 
length and form (index 4— 6.5), nearly straight, widest at the base or slightly 
curved and wider in the middle. Pecten occupying about l / 5 — 74 of the length 
of the siphon, with 9 — 18 closely spaced denticles, 1—3 denticles near the 
base rudimentary. 



396 



Four pairs of hair tufts on the siphon (rarely 3 or 5 pairs), situated at 
the sides of the posterior surface. First pair near the base situated 
slightly distal to the distal denticle of the pecten, with 3—8 branches, first 
pair of tufts situated close together. Second pair in middle of siphon 
situated more laterally, with 2 — 7 branches, 3rd pair with 2 — 7 shorter 
branches situated most laterally, 4th pair the shortest, with 2 — 5 branches, 
situated like the 2nd pair. Tufts shorter than the width of the siphon at 
their position or distinctly longer. 

Main tracheal trunks broad, ribbon -shaped, at least half as wide as the 
siphon. Stigmal plate (Figure 259) slightly wider than long. "Stirrup" 
broad, with a large cavity, narrowing between the spiracles and forming an 
anterior arc which extends far beyond the middle of the spiracles. 
Posterior arc rounded, with a long chaetoid with a broad base. Behind it is 
a posterior appendage forming an irregularly oval, slightly pigmented plate, 
darker at the distal end where the posterior valves are attached. Spiracular 
processes of "stirrup" weakly developed. 

Spiracles with strongly inward narrowing openings and deeply invaginated 
inner margin, following the curve of the "stirrup" and with thickened and 
strongly projecting distal anterolateral margin. 

Anterior valves more strongly chitinized only in the middle in two 
stripes on their upper and lower surface. Organs on lateral valves with 
broad base ending in process a directed toward the spiracles. 

Last segment short, slightly longer than wide. Saddle surrounding the 
segment like a ring, with a short, simple lateral hair. Outer caudal hairs 
2 -branched (rarely simple), 1.5—2 times as long as the gills, inner hairs 
with 2—4 shorter branches, the ventral branch the longest. Fin usually 
with 12 tufts, the longer, posterior tufts longer than the gills or as long as 
these. The last tufts as long or longer than the saddle, sometimes twice 
as long. 

Note on s y s t e mat i c s .* The species of the Culex pipiens 
complex form a polytypical species, including some intraspecific forms 
which differ morphologically and biologically to a varying degree (Jobling, 
1938; Mattingly et al., 1951); Seminar on the Ecology, Biology and Control 
of the Culex pipiens complex, 1965). The complex included originally 
C. p. pipiens L., C.p. fatigans Wied. and C. p. molestus Forsk. 
More detailed studies in different parts of their distribution showed that 
there are populations with intermediate morphological characters and 
different combinations of biological characters which were given different 
names, e.g., C.p. pallens Coq. C.p.australicus Dobr. and C. glo - 
bocoxitus Dobr. were recently described from Australia. The taxon- 
omic status and systematic position of the latter are not clear. 
C.p.australicus occurs only in Australia, mainly in rural areas. It 
resembles C.p. pipiens biologically and the intermediate forms obtained 
by the crossing of C. p. fa t i g a n s and C.p. pipiens morphologically, 
although the latter species has not been recorded from Australia. 

C.p. fatigans (it is often named C.p. quinquefasciatus Say in 
the American literature) differs distinctly from the other forms in the width 
of the second part of the phallosome of the male and in the structure of the 
siphon of the larvae. The species copulates in a small space (stenogamous) 

* Written by E.B.Vinogradova; for details see Vinogradova (1965, 1966, 1969). 



397 



and there is no diapause (homodynamic). The morphological differences 
between C.p. pipiens and C.p. molestus are quantitative, but they 
are significant biologically: C.p. molestus is homodynamic, stenogamous 
and the first egg -batch may develop without a blood meal (autogenous); 
C.p. pipiens has a diapause (heterodynamic), a large space is required 
for copulation (eurygamous), and the eggs develop only after a blood meal. 
The morphological differences between these two forms are small in the 
adults (coloration, relative length of the proboscis and first 4 segments of the 
the palps of the male) and in the larvae (siphonal index, number of branches of 
the tufts of the siphon, branching of hairs, etc.). The best diagnostic charac- 
ter of the larvae, the siphonal index (at most 4.5 or less in C.p. molestus, 
more than 4.5 in C.p. pipiens) shows marked variation which is continu- 
ous but not clinal. There are often micropopulations in the same breeding 
place which differ distinctly in the mean of the siphonal index. The temper- 
atures during development may have a slight influence on the siphonal index. 
There may also be a combination of variations (natural hybridization) which 
has probably taken place in some intermediate populations. Such a high 
degree of variation in morphological characters makes the determination of 
the subspecies of a specimen often difficult or impossible. The reliability 
increases if a series of specimens is examined (mean values of quantitative 
characters supplemented by morphological and biological characteristics 
which are of decisive importance in distinguishing C.p. pipiens and 
C. p. molestus). 

During studies of the C. pipiens complex, the taxonomic status of its 
forms has been differently interpreted. C.p. fatigans was considered for 
a long time as a different species, but it is considered as a subspecies at 
present. A more complicated problem is C.p. molestus (some authors, 
e. g., French authors, name it C.p. autogenicus Roubaud). The 
370 numerous biological differences between C.p. pipiens and C. p. m o 1 e st u s 
were the reason that some authors considered C. molestus as a different 
species, but this is not generally accepted at present. C.p. pipiens and 
C.p. molestus are increasingly being considered as subspecies of 
C. pipiens, including the autogenous and anautogenous forms, because they 
cannot be distinguished by morphological characters and because of the 
existence of many intermediate forms (further details will be given below). 
We agree, but we think that it would be useful to give below the biological 
characteristics of the extreme typical forms C.p. pipiens and 
C.p. molestus. 

In addition to the subspecies mentioned above, there are intermediate 
forms of the complex in different parts of the distribution which have a 
varying stability. C.p.pallens combines the characters of C.p. pipiens 
and C.p. fatigans and was found in a wide zone adjacent to the area of 
the above subspecies in North America (between 39° and 36°N) and in the 
Far East. It shows intermediate morphological characters, particularly 
the moderately broad second part of the phallosome. Its Far Eastern 
populations form long lived, stable populations, but the North American forms 
are possibly seasonal. 

Hybrids between C.p. molestus and C.p. fatigans are known from 
Australia; their appearance is restricted in some localities to the summer- 
autumn period, and their elimination to the winter. 



398 



The forms intermediate between C.p. pipiens and C.p.molestus 
are widespread in nature, mainly in the southern parts of the Palaearctic, 
North Africa, France, Italy, USSR (Crimea, Northern Caucasus, Transcau- 
casia). They usually show intermediate morphological characters (mean 
values of the siphonal index of the larvae 4—5) and combinations of biological 
characters. Some authors, particularly Roubaud and Ghelovitch, 1956; 
Rioux, 1958, and others have named these biotypes: C.p. berbericus 
Roubaud (stenogamous, anautogenous, anthropophilic and homodynamic). 
We do not think that this is advisable because the possible number of 
combinations of biological characters is quite large and may result in 
complication of the terminology. The anthropophilic populations of C. pi pi - 
ens in the USSR are of practical and scientific interest. Mass attacks on 
people have been observed in Moscow, Rostov-on-Don, Odessa, Krasnodar, 
Stavropol, Novorossiisk and other localities. 

The development of populations with intermediate characters is of 
interest. Most authors consider some, if not all, populations as of hybrid 
origin. Evidence of this are the many experiments in crossing these forms 
which produce fertile progeny, e. g., an experimental generation of C. p. pal - 
lens. The tracing of morphological and biological characters (siphonal 
index of the larvae, structure of the hypopygium, autogenesis, stenogamy) 
also conforms with this view. 

During the development of C.p. pipiens and C.p.molestus, the 
adaptation of C. molest us to existence in the human environment was 
of first importance. In temperate climates and harsh winters, the existence 
of both forms, of the form with a diapause (C. p. p i p i e n s), is possible, or 
the existence of the homodynamic form (C.p.molestus), which is 
restricted to specific ecological niches like warm cellars and similar 
habitats in which they reproduce throughout the year. The existence in 
closed habitats also facilitates the selection of autogenous and stenogamous 
forms, which has been confirmed experimentally and in nature. The 
morphological and biological differentiation of the two subspecies increases 
further north. This is probably connected with climatic factors. 



Culex pipiens pipiens L. 

Mesonotum usually with reddish brown scales. Second part of phallo- 
some narrow, hook-shaped (Figure 258, A). 

Fourth -stage larva (Figure 259) with very variable characters in 
external habitus and branching and development of hairs. The branching 
of the hairs in the nominate subspecies is usually weaker but this cannot be 
determined on individuals but has to be determined by examination of a 
large material. The median and inner frontal hairs are 3 — 5 -branched 
(average 4 branches). Median hair behind the comb with 6—8 branches. 
Hair tufts on siphon (counted from the base): first tuft with 3 — 4, 2nd and 
3rd tufts with 2-3, 4th tuft with 2-4 branches. 

Siphon straight, slightly curved in the middle but not widened, tapering 
apically, longer than in the other subspecies (index 4.5—6.4, average 5.3 — 5.4). 

Last segment relatively short, slightly longer than wide. Outer caudal 
hairs usually 1.5 times longer than the gills, with 2 branches. Ends of the 



399 



longest posterior tufts of the fin reaching to the end of the gills but not 
projecting beyond them. Gills lanceolate, with pointed end, distinctly 
longer than the saddle; both pairs of the same length. 

Distribution. It occurs mainly in temperate climates. Palaearctic, 
northern half of North America, North Africa; rarely found in West Africa. 
The disturbance of the normal process of mating probably prevents its 
occurrence in the tropics. It is distributed throughout the USSR in river 
valleys and in inhabited areas, nearly to the Arctic Circle. It is rarely 
east of Lake Baikal. 

Biology. The larvae are found in the most diverse natural and arti- 
ficial stagnant water bodies, swamps, shallow parts of large water bodies, 
ditches, pools, puddles, irrigation ditches, rain barrels, reservoirs. They 
prefer clean water but sometimes develop in polluted water. The breeding 
places are not always connected with inhabited areas. They have a large 
vertical distribution, to 2,000—2,500 m in the Talysh and western Pamirs. 
They occur from May and in more southern regions from March to Septem- 
ber-November. They reach maximum numbers in late summer. Two or 
more generations per year in the South. Fertilized females hibernate in 
diapause in all possible natural shelters and in unheated cellars and 
basements. The numbers of hibernating females may be very large. 
Continuous development throughout the year probably takes place in the 
southern parts of the distribution. The adults are eurygamous, requiring 
a large space for copulation. The eggs develop only after a blood meal. 
This subspecies feeds only on birds. Many authors have observed that 
they are practically absent among mosquitoes biting man. Attacks on man 
are more frequent in southern regions, but reliable determination of the 
subspecies of biting mosquitoes is difficult. Laboratory observations 
confirm that the southern forms are more aggressive to man than the 
northern forms. ; 

Their role as vectors of diseases is not clear. C.p.pipiens has been 
considered as a vector of western equine encephalitis and St. Louis 
encephalitis in North America, but the determination of the subspecies is 
doubtful. 



372 Culex pipiens molestus Forskal, 1775 

Indistinguishable from C.p.pipiens by morphological characters 
and coloration. 

Fourth -stage larva (Figure 260), also very variable, but the 
branching of the hairs is usually stronger. Median and inner frontal 
hairs with 4—6 (average 5) branches. Median hair behind the comb with 
6—10 branches. Hair tufts of siphon (counted from the base): first tuft 
with 3-8, 2nd and 3rd tufts with 2-7, 4th tuft with 2-5 branches. 

Siphon relatively shorter than in the nominate subspecies (index at most 
4.5) and tapering slightly more strongly apically but not widened in the 
middle. 

Last segment longer than in the nominate subspecies. Outer caudal 
hairs twice as long as the gills or longer, although they are absolutely 
shorter than in the nominate subspecies. Gills as long as or shorter than 
the saddle, upper pair longer than the lower. 



400 



FIGURE 260. Culex pipiens molestus Forsk. Fourth-stage larva. 
Posterior end, lateral. 



Distribution. C. p. molestus is sympatric with C. p. pi p i e n s ; 
the only exception is Australia, where only C.p. molestus occurs 
(introduced). C. p. molestus is distributed at present in many cities in 
Europe, Asia, Africa and America; England, Sweden, Norway, Poland, 
France, Germany, Czechoslovakia, Italy, Egypt, Israel, Sudan, Japan. In the 
USSR it has been found in Leningrad (Fedorov, 1941), Vologda, Moscow, 
Kiev, Kharkov, Dnepropetrovsk, Donetsk, Krasnyi Liman, Nikolaev and Baku. 
It bites people in hot countries and in years of mass reproduction in tem- 
perate climates. 

Biology. The larvae occur in temperate climates in water in cellars 
and attics (mainly in localities where reproduction continues throughout the 
year), also in rain barrels and other artificial receptacles near houses 
3 73 (summer and early autumn). They may develop throughout the year in 

open water bodies in the tropics and subtropics. The larvae prefer strongly 
polluted water and stagnant water in sewers, but they may also develop in 
less polluted water. Several generations per year. The females do not 
undergo diapause. The adults are stenogamous, the minimum space neces- 
sary for copulation is 27 cm 3 . Often used in the study of stenogamy are the 
cages of Frizzi (5 dm), of Roubaud (26 dm) and of Bates (337 dm). The 
females are autogenous (the first egg batch is deposited without a blood meal 
at the expense of larval reserves). If the larvae are badly fed, autogeny is 
present only in a few females. The structure of the ovaries in young, 
recently hatched females is characteristic: the first follicles are usually 
in different stages of development, giving a mosaic picture. Autogeny is 
an inherited character. 

Epidemiological importance. C.p. molestus is a vector 
of West Nile virus. 



401 



Culex pipiens fatigans Wiedemann, 1828 (quinquefasciatus Say, 1823) 

Closely resembling C.p„ pipiens in coloration, differing in the 
yellowish brown, not reddish or dark brown scales on the mesonotum; palps 
of males less densely covered with hairs; transverse stripes on abdominal 
tergites yellowish, usually narrowing laterally, as a rule half -moon-shaped. 
It differs distinctly from C. p. pipiens in the structure of the phallosome 
(Figure 2 58, B), the second part of which is lobe -shaped and very wide 
(it is narrow, hook-shaped in C. p. p i p i e n s ) . The females of the two sub- 
species cannot be distinguished with certainty. 




FIGURE 261. Culex pipiens fatigans Wied. Fourth -stage larva 
(after Barraud): 

1 — siphon, lateral; 2 — antenna; 3 — last abdominal segment, lateral. 



374 Fourth -stage larva (Figure 261) differing from that of the other 

subspecies mainly in the form of the siphon, in which the anterior surface 
is distinctly convex and the posterior surface is convex in the middle and 
slightly concave (rarely straight) in the tapering part near base and apex. 
The maximum width of the siphon is therefore not at the base but in the 
middle. The width at the apex of the siphon is slightly more than half the 
width at the base. The tufts on the siphon are usually more strongly 
branched than in the two other subspecies: the 2 pairs near the base with 
6—7 branches, as long as or slightly longer than the width of the siphon at 
their position. The other hairs of the body, particularly the frontal hairs, 
are more strongly branched than in the larvae of the other subspecies 



402 



(median and inner hairs with 5—7 secondarily feathered branches). Gills 
as long as saddle or longer. 

Distribution. It occurs throughout the tropics and subtropics. 
Not recorded with certainty from the USSR. 

Biology. The larvae occur in artificial water bodies in towns and 
sparsely inhabited areas, rarely in natural water bodies. They may 
develop in strongly polluted water. This gives them a distinct advantage 
over other species of mosquitoes in a biocenosis. Their recent rapid 
increase in numbers in the tropics and subtropics is connected with 
urbanization, which has created favorable conditions for their reproduction. 
The adults are stenogamous. Several generations per year without 
diapause. 

A blood meal is necessary for the development of the eggs. The range 
of hosts for food is broad, but different populations vary in this respect. 
C.fatigans feeds mainly on man and is a vicious bloodsucker in some 
parts of its distribution. 

Epidemiological importance. It is an important vector of 
filariasis caused by Wuchereria bancrofti, a vector of Brugia 
malayi and D i r o f i 1 a r i a i mm it is and also of some Plasmodia of 
birds. It is a vector of St. Louis encephalitis and of western (equine) 
encephalitis in America. It also transmits Japanese encephalitis. 



403 



INDEX OF SCIENTIFIC NAMES* 



aconitus, Anopheles 49 

Aedes 9, 10, 11*, 12*, 14*, 15, 16*, 17, 

18, 23, 24*, 25, 33, 34, 39, 43, 44, 47, 

48, 51, 54, 57, 59, 72, 74, 75, 169—314 
Aedes (subgenus) 179, 183, 308 
Aedimorphus (subgenus) 43, 44, 171, 270 
aegypti, Aedes 9, 39, 48, 51, 53—56, 176, 

183, 185, 294, 295*, 296*, 297* 
africanus, Aedes 54 
alaskaensis, Culiseta 46, 135 — 137, 144, 

145*, 146*, 147* 
alaskaensis alaskaensis 135, 147 
alaskaensis indica 135, 147 
albescens, Aedes 229 
albimanus, Anopheles 50 
albionensis, Orthopodomyia 130 
albitarsis, Anopheles 50 
albopictus, Aedes 53—55, 176, 183, 185, 

303*, 304* 
alektorovi, Aedes 19, 46, 170, 183, 185, 

284*, 286* 
algeriensis, Anopheles 45, 50, 77, 78, 79, 

80, 81*, 82* 
Allotheobaldia (subgenus) 134, 136, 138 
alpinus, Aedes 255 
annulata, Culiseta 25*, 26*, 135—137, 

147, 148*, 149* 
annulata annulata 135, 150 
annulata subochrea 135, 137, 150 
annulipes, Aedes 176, 182, 188, 216, 217* 
annulirostris, Culex 54 
Anopheles 9, 10, 15*, 16*, 17, 18*, 19, 

20, 21*, 22*, 23—25, 26*, 27, 29*, 

30*, 31*, 32, 34*, 35*, 36—40, 42— 

44, 49—52, 54, 57, 59, 61, 62, 72, 

74, 76—117 
Anopheles (subgenus) 76, 78, 80 
Anophelinae 18, 42, 43, 75 
antennatus, Culex 54 
apicalis, Culex 331 
aquasalis, Anopheles 50 
Armigeres 44, 54 
aureus, Aedes 45, 179, 313 
argenteus, Aedes 294 

balabacensis, Anopheles 51 

bancrofti, Anopheles 53 

barbirostris, Anopheles 49, 56 

Barraudius (subgenus) 325 

behningi, Aedes 173, 182, 187, 208, 210*, 

211* 
beklemishevi, Aedes 176, 182, 187, 214, 

215* 



bergrothi, Culiseta 134, 136, 137, 151*. 

152* 
bifurcatus, Anopheles 86 
bitaeniorhynchus, Culex 25, 45, 56, 316, 

318, 319, 341*, 342*, 343* 
borealis, Culiseta 151 
buxtoni, Mansonia 163, 164, 167, 168* 

calopus, Aedes 294 

cantans, Aedes 56, 175, 181, 188, 202, 

203*, 204*, 205* 
caspius, Aedes 56, 57, 170, 180, 188, 191 
caspius caspius 170, 180, 188, 190*, 192, 

193* 
caspius dorsalis 170, 180, 188, 191, 195*, 

196* 
cataphylla, Aedes 178, 181, 186, 261*. 

262* 
Chagasia (subgenus) 43 
Chaoboridae 43 
chemulpoensis, Aedes 176, 183, 185, 306* 

307* 
chloropterus, Sabethes 54 
christophi, Toxorhynchites 45, 118, 119*, 

120, 121*, 122* 
cinereus, Aedes 19*, 34, 56, 57, 179, 183, 

308, 310* 
cinereus cinereus 179, 183, 184, 309*, 

310* 
cinereus esoensis 179, 183, 184, 312* 
cinereus rossicus 179, 183, 184, 308*, 311* 
claviger, Anopheles 35, 50, 53, 59, 60, 

77—79, 85*, 86*, 87* 
communis, Aedes 56, 178, 181, 188, 

232, 233, 234* 
Coquillettidia (subgenus) 163 
cretinus, Aedes 176, 183, 185, 298* 
Culex 9, 12*, 13*, 15*, 16*, 17 — 19, 

21—25, 33, 34, 37, 43—45, 48, 51, 52, 

54, 57, 59, 73—75, 314—374 
Culicella (subgenus) 17, 34, 37, 59, 153 
Culicidae 9, 13, 24, 25, 38, 42—46, 71 
culicifacies, Anopheles 49 
Culicinae 9, 10, 17, 18, 24*, 30, 32*, 

34—36, 39, 40, 42, 61, 62, 125 
Culicini 32, 125, 126 
Culiseta 11*, 15, 17—19, 23—25, 28, 

33, 34, 44, 57, 59, 72, 74, 75, 133—162 
cyprius, Aedes 20, 172, 182, 187, 221, 

222* 

darlingi, Anopheles 50 
Deinocerites 43 



* [Reproduced photographically from the Russian original. Page numbers refer to those of the original, which 
appear in the left-hand margin of this translation.] Bold face figures indicate pages with description, 
italics denote synonyms; an asterisk indicates a figure of the species. 



405 



detritus, Aedes 38, 177, 182, 188, 264, 

265*, 266*, 267* 
diantaeus, Aedes 23, 56, 178, 180, 184, 

247*, 248*, 249* 
diver sus, Aedes 223 
Dixidae 43 
duplex, Aedes 192 
dureni, Anopheles 51 

echinus. Aedes 19, 176, 183. 186, 278, 

279*, 280* 
Eretmapodites 43 
excrucians, Aedes 57, 60, 176, 182. 187, 

211, 212*, 213*, 214* 
exilis, Culex 362, 364 

farauti, Anopheles 49, 53 

fasciatus, Aedes 294 

Ficalbia 43 

Finlava (subgenus) 16, 17, 23. 44. 170, 

179, 274 
fitchii, Aedes 270c 
flavescens, Aedes 57, 172, 181, 187, 218, 

219* 220* 
flavopictus, Aedes 45, 176, 183, 185. 

305* 
fluviatilis, Anopheles 49 
freeborni, Anopheles 50, 51 
jreyi, Aedes 221 

fumipennis, Culiseta 135—137, 159* 
funeslus, Anopheles 50, 54. 56 
fuscanus, Culex 45. 315, 317, 318, 322*. 

323*, 324* 

galloisi, Aedes 19, 38, 39, 176, 183, 185, 

300, 301*, 302* 
gambiae, Anopheles 50, 53, 54, 56 
gelidus, Culex 54 — 56, 320 
geniculates, Aedes 19, 20, 45, 176, 183, 

186, 274, 275*, 276*, 277* 
glaphyroplera, Culiseta 135—137, 142, 

143* 
grandilarva, Aedes 214 

hackeri, Anopheles 51 

hayashii, Culex 45, 316, 317, 319, 336, 

337*, 338*, 339* 
Heizmannia 43 
hexodontus, Aedes 178, 181, 189, 240, 

241*, 242* 
hortensis, Culex 20, 316, 317, 320, 333*, 

334* 
hungaricus, Aedes 179, 189, 246* 
hyrcanus, Anopheles 35, 36, 49 — 51, 53, 

78, 79, 103, 104*, 105*, 106* 

impiger, Aedes 177, 181, 189, 258*, 259*, 

260* 
intrudens, Aedes 178, 180, 189, 250*, 

251* 

jacksoni, Culex 45, 316, 318, 319, 350 
japonicus, Aedes 170, 183, 184, 290 
jeyporiensis, Anopheles 49 

kasachstanicus, Aedes 170, 182, 269* 
Kerteszia (subgenus) 43 



kochi, Anopheles 51 

koreicus, Aedes 171, 183, 184, 291*, 292* 

laticinctus, Culex 320 

laurenti, Culex 321 

lepidonotus, Aedes 47, 177, 180, 186, 

227, 228* 
letifer, Anopheles 49, 51 
leucocelaenus, Aedes 54 
leucomelas, Aedes 178, 181, 188, 263* 
leucosphyrus, Anopheles 49, 51 
Limatus 43 
lindesayi, Anopheles 31, 50, 77, 78, 

100, 101*, 102* 
lindesayi lindesayi 77, 103 
lindesayi japonicus 77, 79, 103 
Undtropi, Aedes 298 
longiareolata, Culiseta 22, 34, 45, 134, 

136, 138, 139*, 140*, 141*, 142* 
Lophocoraomyia 43 
Lophopodomyia 43 
lutescens, Aedes 218 
Lutzia (subgenus) 25, 29, 321 

maculatus, Aedes 202 
maculatus, Anopheles 49, 51 
inaculipennis, Anopheles 11, 18, 35, 36, 

40*, 49, 50, 53, 56, 77—79. 90, 91*, 

94*, 95*, 96*, 97, 106* 
inaculipennis maculipennis 77, 90, 96* 
inaculipennis atroparvus 77, 90, 96* 
inaculipennis elutus 99 
maculipennis labranchiae 77, 90, 96* 
inaculipennis melanoon 77, 90, 96* 
inaculipennis messeae 77, 90, 96* 
maculipennis sacharovi 77, 79, 90, 98, 

99, 100* 
maculipennis subalpinus 90, 96* 
Mansonia 13, 17—19, 21, 23—25, 37, 

43, 53, 54, 73—75, 162—169 
mariae, Aedes 170, 180, 187, 196, 197*, 

198* 
marten, Anopheles 50, 77, 79, 88, 89* 
marteri marteri 77, 90 
marteri sogdianus 77, 90 
martinii, Culex 316, 317, 320, 335*. 336* 
Megarhinus (subgenus) 76, 118 
meigenanus, Aedes 237 
melanimon, Aedes 54 
Melanoconion (subgenus) 43 
melanura, Culiseta 54, 55 
mimeticus, Culex 316, 318, 320, 348, 

349*, 350* 
minimus, Anopheles 49, 53 
modestus, Culex 56, 315, 317, 320, 325, 

326* 
montchadskyi, Aedes 270a 
morsitans, Culiseta 24*, 25*, 26*, 60, 

135, 136, 138, 153, 154*, 155* 
multicolor, Anopheles 50 
Myzomyia (subgenus) 31, 43, 44. 108 

nearcticus, Aedes 258 
nemorosus, Aedes 232 
Neoculex (subgenus) 37, 328 
nigerrimus, Anopheles 53 
nigrinus, Aedes 178, 181, 187, 245* 



406 



nigripes, Aedes 177, 181, 186, 255, 256*, 

257* 
nigripes, Anopheles 82 
nili, Anopheles 50 
nipponicus, Aedes 45, 176, 182, 184, 

280, 281*, 282* 
niveus, Aedes 280 
nobukonis, Aedes 45, 179, 313 
Nyssorhynchus (subgenus) 43 

Ochlerotatus (subgenus) 16, 19, 23, 28, 

44—46, 170, 179 
ochroptera, Culiseta 135, 136, 138, 155*, 

156, 157*, 158* 
Opifex 44 
orientalis, Culex 315, 318, 320, 352, 

353*, 354*, 355* 
ornatus, Aedes 274 
Orthopodomyia 19, 20, 33, 34, 37, 44, 

46, 73—75, 130—133 

parvulus, Aedes 258 

pattoni, Anopheles 49 

perexiguus, Culex 358 

perturbans, Mansonia 54 

petragnani, Anopheles 88 

pharoensis, Anopheles 50 

philippinensis, Anopheles 49, 51 

Phoniomyia 43 

pionips, Aedes 178, 181, 188, 235*, 236 

pipiens, Culex 24, 46, 54, 317, 318, 365, 

366*, 367* 
pipiens pipiens 52, 318, 321, 368*, 370 
pipiens fatigans 52—54, 318, 320, 373* 
pipiens molestus 318, 321, 372* 
pipiens quinquefasciatus 369, 373 
plumbeus, Anopheles 23, 25, 27, 35, 

50, 77—79, 82, 83* 
polyiiesiensis, Aedes 52 
pseudopunctipennis, Anopheles 50 
Psorophora 43 
pulcherrimus, Anopheles 35, 49—51, 78, 

80, 108, 110*, 111*, 112* 
pulchripalpis, Orthopodomyia 19, 46, 130, 

131* 132* 133* 
pulchritarsis, Aedes 38, 40, 170, 180, 199* 
pulchritarsis pulchritarsis 170, 186, 200 
pulchritarsis asiaticus 170, 186, 201*, 202* 
pulchritarsis berlandi 201 
pullatus, Aedes 46, 47, 178, 180, 189, 

252 253* 254* 
punctor, Aedes 178, 181, 189, 237, 238*, 

239* 
punctulatus, Anopheles 49 
pusillus, Culex 315, 317, 319, 327, 328*, 

329* 

quadrimaculalus, Anopheles 50, 51 
quartus, Aedes 216 

refiki, Aedes 177, 180, 186, 225,226* 

rempeli, Aedes 270d 

restuans, Culex 54 

richiardii, Mansonia 57, 59, 163, 164*, 

165*, 166* 
riparius, Aedes 40, 175, 182, 187, 205, 

207* 



riparius riparius 175, 182, 206*, 208 
riparius ater 175, 182, 208, 209* 
rostochiensis, Aedes 261 
rufipes, Anopheles 50 
rusticus, Aedes 177, 180, 186, 223, 224* 
225* 

Sabethes 43 

Sabethini 43, 125, 126 

salinellus, Aedes 263 

salinus, Aedes 264 

semicantans, Aedes 205 

seoulensis, Aedes 170, 182, 185, 283 

sergenti, Anopheles 50 

setivalva, Culiseta, 46, 135, 136, 137, 

160, 161* 
silvestris, Culiseta 47, 158 
simanini, Aedes 177, 182, 267, 268* 
simpsoni, Aedes 54, 55 
sinensis, Anopheles 103 
sinensis, Culex 25, 45, 316, 318, 319, 

344*, 345* 
sollicitans, Aedes 54 
Stegomyia (subgenus) 15, 17, 23, 33, 43, 

44, 54, 55, 176, 179, 293 
stephensi, Anopheles 49 — 51 
Stethomyia 43 
sticticus, Aedes 56, 178, 181, 189, 243*, 

244* 
stramineus, Aedes 270b 
subdiversus, Aedes 177, 180, 186, 230* 
subpictus, Anopheles 56 
sundaicus, Anopheles 49, 51 
superpictus, Anopheles 35, 49, 50, 60, 

78, 80, 113*, 114*, 115*, 116* 

tarnogradskyi, Aedes 311 

tarsalis, Culex 54, 55 

territans, Culex 19, 33, 43, 52, 317, 320, 

330*, 331* 
territans territans 317, 332 
territans judaicus 317, 333 
tesselatus. Anopheles 56 
theileri, Culex 45, 316, 318, 320, 360, 

361* 
Theabaldia 72, 133 
titillans, Mansonia 54 
togoi, Aedes 53, 54, 170, 182, 184, 287*, 

288*, 289* 
Toporayia 43 

torrentium, Culex 317, 318, 321, 364 
Toxorhynchites 9, 18, 19—25, 29, 30, 

33, 34, 37, 38, 44, 72, 75, 118—125 
Toxorhynchitinae 117 
Toxorhynchitini 32 
Trichoprosopon 43 
trichurus, Aedes 46 
Tripteroides 44 
tritaeniorhynchus, Culex 45, 54, 55. 

316, 318, 321, 354, 356*, 357* 
turkhudi, Anopheles 31 

umbrosus, Anopheles 49 

unguiculata, Uranotaenia 126, 127*, 128*, 

129* 
univittatus, Culex 54, 317, 318, 321, 

358*, 359* 



407 



Uranotaenia 9, 17, 18, 20, 22 24 27 
31, 33, 43, 44, 72, 74, 75, ' 126-130 

vagans, Culex 316, 318, 321, 362*, 363* 
vexans, Aedes 46, 53, 54, 56 57 171 

179, 189, 270, 271*, 272* 
vexans vexans 171, 273 
vexans nipponii 171, 273 



vishnui, Culex 54, 319 

vittatus, Aedes 176, 183, 185, 299* 300* 

vorax, Culex 45, 315, 317, 318,' 324* 

whitmorei, Culex 45, 316, 318 319 
345, 346*, 347* ' ' 

Wyeomyia 43 



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v. 3 



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