Illustrated Key to the Female Anopheles of Southwestern Asia and Egypt (Diptera: Culicidae)’

JULY 1992

ILLUSTRATED KEY TO THE FEMALE ANOPHELES OF SOUTHWESTERN ASIA AND EGYPT (DIPTERA: CULICIDAE)’

JAYSON I. GLICK~

Walter Reed BiosystematicsUnit, Department of Entomology, Walter Reed Army Institute of Research, Washington,DC 2030 7-5 100

ABSTRACT. An illustrated key for the identification of the female Anophelesmosquitoes of southwesternAsia and Egypt is presented. Thirty-nine speciesand three subspeciesare treated, including 25 speciesand one subspeciesof Anopheles(Cellia) and 14 speciesand two subspeciesof Anopheles(Anopheles). A new speciesfrom Egypt of the subgenusCellia closely related to Anophelesstephgnsi Liston is left unnamed. Anopheles(Anopheles) pseudopictus Grassiis removed from synonomy with Anopheles(Anopheles) hyrcanus (Pallas), and Anoph- eles (Anopheles)habibi Mulligan and Puri is recognized as a junior synonym of Anopheles (Anopheles) claviger (Meigen). Tables providing important taxonomic references and the geographicdistribution for each speciesare included.

INTRODUCTION 1949, Senevet and Andarelli 1955a, Russell et al. 1963). An identification key for the Anopheles This work beganas a study of the Anopheles mosquitoesof the entire SouthwestAsian Re- mosquitoesof the Arabian Peninsula,empha- gion has long been a necessityfor entomolo- sizing the fauna of Saudi Arabia and Kuwait, gistsdealing with malaria vectors. Published and was expanded to include all of south- keys and speciesdescriptions for the region western Asia as defined by Harbach (1988), are scattered throughout the literature, and and modified to include all land south of the are often limited in scope to the Anopheles Russianrepublics between the Mediterranean species of a single country (Salem 1938, Seaand the Indus River of Pakistan,including Egypt; Pringle 1954, Iraq; Abdel-Malek 1958, all of Turkey and Egypt. The material exam- Syria; Shahgudian 1960, Iran; Postiglione et ined came largely from the collections of the al. 1973, Turkey; Danilov 1985, Afghanistan) National Museum of Natural History, Smith- or limited geographical region such as the sonian Institution, and the British Museum Arabian Peninsula (Mattingly and Knight (Natural History). The Anophelesfauna of the 1956, Shidrawi and Gillies 1987) and the SouthwestAsian Region presently consistsof Indian Subregion(Christophers 1933). Many 39 speciesand three subspecies,representing are now of limited value due to numerous two subgenera.The majority of the species nomenclatural changesand additions, and re- have Palaearctic affinities, while a smaller finementsin our ability to differentiate sibling number are clearly more Ethiopian or Ori- species,and are ineffective for identification ental in their distribution. of Anopheleson a region-widebasis. Available Two nomenclatural changes have been keys for the Anophelesof the Palaearctic Re- made for the An. (Anopheles)of the Southwest gion are similarly ineffective (Bates et al. Asian Region. Anopheles(Ano.) pseudopictus Grassi is removed from synonomy with An. ’ ’ The viewsof the authordo not purportto reflect (Ano.) hyrcanus(Pallas) based on the appar- the viewsof the Departmentof the Army or the ent absenceof evidence for its hybridization Departmentof Defense. with An. hyrcanusin any part of its distribu- ’ Reprint requests:Walter Reed Biosystematics Unit, MuseumSupport Center, Smithsonian Insti- tion, and the distinctnessof material studied tution,Washington, DC 20560. of both An. hyrcanus and An. pseudopictus 126 MOSQUITO SYSTEMATICS VOL. 24, No. 2 from Turkey, Iran and Afghanistan. Bruce A. (Colbourne and Smith 1964, Sebai 1988, Za- Harrison (personalcommunication) provided har 1985) An. (Cel.) pharoensis in Egypt characterswhich clearly show An. (Ano.) ha- (Zahar 1974), and An. (Cel.) pulcherrimusin bibi Mulligan and Puri to be a synonym of Afghanistan (Zahar 1974). An. (Ano.) claviger (Meigen). In particular, Indigenousmalaria has been eliminated for the lower proepisternalsetae (PeSL) are found the most part from Bahrain and Kuwait, only in An. habibi and An. claviger, and a where imported malaria is now the primary closely related western Mediterranean species problem (Amin 1989, Hira et al. 1985). An. (Ano.) petragnani Del Vecchio. Compar- Anopheles(Gel. ) stephensiand An. (Gel. ) pul- ison of the type female of An. habibi in the cherrimus are present in both countries and British Museum (Natural History) with An. are known vectors in neighboring countries. clavigerfrom Iraq and Israel showedno mor- In Iraq, primary malaria vectors presently phological differencesbetween the two; com- include An. (Ano.) sacharovi,An. (Gel. ) ste- parison with An. claviger from France, phensi and An. (Gel. ) superpictus(Abul-Hab Greece, Italy, Spain, Israel and Russiashowed and Al-Kassal 1986). Malaria eradication pro- no statisticaldifference in the length of the grams have reduced transmission in many wing petiole to the anterior forked cell be- areas of the Southwest Asian Region, while tween the two speciesas was stated in the there has been a resurgence of malaria in correction to the original description of An. others. Ramsdale and Haas (1978) reviewed habibi (Mulligan and Puri 1936b); and com- the problems of resurgentmalaria in southern parison of the genitalia of An. habibi males and southeastern Turkey where An. (Ano.) from Quetta, Baluchistan (BM 1938-663/ sacharovi, An. (Gel. ) superpictusand other 14 13) with An. claviger males from Israel, speciesmay be playing a role in transmission. Greece and England showedno salient differences. Anopheles habibi is therefore recog- METHODS AND PRESENTATION nized as a junior synonym of An. claviger (NEW SYNONOMY). The scopeof this study includesthe Anoph- Morphological characters used here are eles fauna from portions of the North Eura- based predominantly on previous usage in sian, Mediterranean, Afro-Arabian (Desert), published literature. Harbach and Knight Afrotropical and Indo-Iranian malarial epi- ( 1980) are followed for morphological terms demiological zones as defined by Macdonald and abbreviations, and wing spot characters (1957). Primary malaria vectorsin the South- and abbreviationsare taken from the nomen- west Asian Region include An. (Cellia) ara- clature usedby Wilkerson and Peyton (1990). biensis Patton, An. (Gel.) culicifaciesGiles, In the key, morphological featuresare writ- An. (Cel.) fluviatilis James, An. (Cel.) phar- ten out, followed by their abbreviation, to oensis Theobald, An. (Gel. ) pulcherrimus assistusers. Specimens were examined at 20- Theobald, An. (Ano.) sacharovi Favre, An. 120x magnification under blue-filtered tung- (Cel. ) sergentii (Theobald), An. (Gel. ) ste- sten light. Pure white was used as a reference phensi Liston and An. (Gel.) superpictus for determining other colors according to the Grassi. Secondary vectors include An. (Gel.) method of Peyton and Ramalingam (1988). annularis Van der Wulp, An. (Cel.) cinereus Taxonomic notes are indicated in the key for Theobald, An. (Ano.) claviger and An. (Gel.) certain speciesand presented in an “Expla- multicolor Cambouliu (White 1989, Zahar nation of Notes” section immediately follow- 1974). Although many of the primary vectors ing the key. are important in malaria transmissionover a Table 1 is a taxonomic index to the Anoph- widespreadarea of the region, several are of eles mosquitoes of southwestern Asia and concern in more limited areas,including An. Egypt, including a list of important taxo- (Gel.) arabiensis in the Arabian Peninsula nomic references for each species.Tables 2 JULY 1992 127

Table 1. Taxonomic index and referencesfor the Anophelesmosquitoes of southwesternAsia and Egypt.

Key Taxon couplet Taxonomic references Genus Anopheles SubgenusAnopheles 2,27 algeriensisTheobald 28 Marshall (1938) Senevet and An- darelli (1955a) claviger(Meigen) 32 Mulligan and Puri ( 1936a, 1936b), Marshall (1938) Rossand Rob- erts (1943), Torres Canamares ( 1945), Senevet and Andarelli (1955a, 1955b) coustaniLaveran 35 Evans ( 1938), de Meillon ( 1947) Senevet and Andarelli (1955a), Gillies and de Meillon (1968) gigas simlensis(James) 39 Christophers(1933), Reid ( 1968) hyrcanus (Pallas) 37 Christopher-s(1933), Ross and Roberts ( 1943), Reid ( 1953), Gutsevich ( 1976) lindesayi Giles 39 Christophers( 1933), Reid ( 1968) maculipennisMeigen 30 Hackett and Missiroli ( 1935) Marshall (1938), Bates (1940), Senevet and Andarelli (1955a), Rioux (1958) Gutsevich et al. (1974) White ( 1978) marteri sogdianusKeshishian 31 Keshishian ( 1938) Shahgudian (1956) martinius Shingarev 30 Hackett and Missiroli ( 1935) Bates (1940), White (1978) nigerrimusGiles 38 Christophers(1933), Reid (1953, 1968) Harrison ( 1972) Hani- son and Scanlon (1975) peditaeniatus(Leicester) 38 Reid (1953, 1968) Harrison (1972), Harrison and Scanlon (1975) plumbeus Stephens 32 Marshall (1938) Senevet and An- darelli (1955a) pseudopictusGrassi 37 Dow ( 1953), Senevet and Anda- relli (1955a) sacharoviFavre 30 Hackett and Missiroli ( 1935), Bates (1940), Ross and Roberts (1943), Rioux (1958) White (1978) subalpinusHackett and Lewis 30 Hackett and Lewis (1935), Bates (1940) Rioux (1958) White (1978) Cianci et al. (1987) tenebrosusDoenitz 35 Evans (1938), de Meillon (1947) Gillies and de Meillon (1968) SubgenusCellia 2,28 annularis Van der Wulp 5 Christophers( 1933), Ross and Roberts ( 1943), Bonne-Wepster and Swellengrebel( 1953), Hara (1959) Reid (1968) apoci Marsh 26 Marsh (1933) Table 1 continues. 128 MOSQUITO SYSTEMATICS VOL. 24, No. 2

Table 1. Continued.

Key Taxon couplet Taxonomic references arabiensisPatton 22 Evans (1938), de Meillon (1947), Senevet and Andarelli ( 1955a), Coluzzi (1964), Gillies and de Meillon (1968), Zahar et al. (1970), White (1975, 1985), Mat- tingly (1977) azaniae Bailly-Choumara 9 Bailly-Choumara ( 1960) Gillies and de Meillon (1968) cinereusTheobald 16 Evans (1938) de Meillon (1947), Senevet and Andarelli (1955a), Gillies and de Meillon (1968) Culicifacies Complex 23 Christophers( 193 3), Evans (1938), Ross and Roberts (1943), de Meillon ( 1947), Bonne-Wepster and Swellengre- be1( 1953), Gillies and de Meil- lon ( 1968), Harrison (1980) demeilloni Evans 25 Evans (1938), de Meillon (I 947), Gillies and de Meillon ( 1968) dthali Patton 13 Christophers(1933), Evans ( 1938), de Meillon ( 1947), Se- nevet and Andarelli (I 955a), Gillies and de Meillon (1968) fluviatilis James 25 Christophers( 1933), Ross and Roberts (1943), Bonne-Wepster and Swellengrebel( 1953) maculatusTheobald 12 Christophers( 1933), Ross and Roberts ( 1943), Bonne-Wepster and Swellengrebel( 1953) Hara (1959), Reid (1968) moghulensisChristophers 20 Christophers( 1933) multicolor Cambouliu 15 Christophers( 193 3), Evans (1938), Ross and Roberts (1943), de Meillon (1947), Se- nevet and Andarelli (1955a), Gillies and de Meillon (1968) paltrinierii Shidrawi and Gillies 26 Shidrawi and Gillies ( 1987) pharoensisTheobald Evans (1938), Ross and Roberts (1943), de Meillon (1947), Se- nevet and Andarelli (1955a), Gillies and de Meillon (1968) pretoriensis(Theobald) 11 Evans (1938), de Meillon (1947) Gillies and de Meillon (1968) pulcherrimusTheobald 4 Christophers( 1933) Gutsevich et al. (1974) rhodesiensisrupicola Lewis 9,28 Evans ( 1938), de Meillon ( 1947), Senevet and Andarelli ( 1955a), Mattingly and Knight (1956), Gillies and de Meillon (1968) sergentii(Theobald) 24 Christophers( 1933), Senevet and Andarelli (1955a), Mattingly and Knight (1956) splendidusKoidzumi 5 Christophers(1933), Bonne-Weps- ter and Swellengrebel( 1953), Hara (1959) Table I continues. JULY 1992 129

Table 1. Continued. Key Taxon couplet Taxonomic references squamosusTheobald 7 Evans (1938), de Meillon (1947) Gillies and de Meillon (1968) stephensiListon 19 Christophers (1933), Ross and Roberts ( 1943) subpictusGrassi 22 Christophers (1933), Ross and Roberts ( 1943) Bonne-Wepster and Swellengrebel ( 1953) Hara (1959) Reid (1968) superpictusGrassi 20 Christophers (1933), Ross and Roberts (1943) Senevet and An- darelli (1955a), Gutsevich et al. (1974) turkhudi Liston 16 Christophers(1933), Evans (1938), Saliternik and Theodor ( 1942), de Meillon ( 1947), Gillies and de Meillon ( 1968) willmori (James) 12 Christophers(1933) Reid (1968) n. sp. 19 B. A. Harrison, personal communication

Table 2. Distribution ofA@zeZes (Arzoplzeles) in southwestern Asia and Eapt.

algeizensis’ claviaer coustaui Jigas simlerzsis

hYK~IIlLS lirldesavi maculipermis marteri sogdiarw martinius rzifenimus peditaelziatus plum beus pseudopictus sacharovi subalpirw tenebroms 130 MOSQUITO SYSTEMATKS VOL. 24. No. 2

Table 3. Distribution ofAnqheles (Cellia) in southwesternAsia and Egypt.

annularis apoci ara biensis azaniae l cinereus l a l 0 l a culicifacies a l a l 0 l a demeilloni 0

dthali l 0.0.. 0 0 0 0 0 0 0

fluvia tilis 0 l a 0 l 0 a 0 maculatus 0 l moghulensis 0 a 0 multicolor a 0.00.. .ooooo 0 paltriniehi a a pharoensis a a l a a 0 pretonensis’ a a pulcherrimus 0 a 0 0 0 l a l 0 0 0 r/zodesiensis rupicola 0 l l 0 a l a l seren tii 0.0.. l .o@oo 0 0 splefldidus 0 0

sauamosus l stephensi a 0 a a a 0 l 0 a subpictus l 0 l superpictus 0 0.00.. a l 0.0 turkhudi l 0.0.. 0 0 0 0 willmon ’ a 0 n. sp. a and 3 include the distribution for each species terms and abbreviations used in the key within the region. Figures 1 and 2 provide a (taken from Wilkerson and Strickman 1990, summary of the majority of morphological with modifications). JULY 1992 131

Thorax

Fig. 1 (above). Female ilno$ze/es mosquito, lateral view. Ap, antepronotum; C-I. forecoxa: C-II. midcoxa: C-III, hindcoxa: Clp, clypeus; Fe-I, forefemur: Fe-II. midfemur: Fe-III. hindfemur; Hl, halter: La. labellum: Mks. meso- katepisternum: Mm, mesepimeron; MP~P,_~, maxillary palpus, palpomeres l-5; Mpn. mesopostnotum; MS, meso- thoracic spiracle: Mts, metepisternum: P, proboscis: Pa, paratergite; PA, postspiracular area: Ppn. postpronotum: Ps, proepisternum; S-I-VIII, sterna I-VIII: Scu, scutum; Stm. scutellum; Ta-III,_5, hindtarsomeres 1-5: Te-I-VIII, terga I- VIII; Ti-III. hindtibia; Tr-I, foretrochanter: Tr-II, midtrochanter; Tr-III, hindtrochanter.

Fig. 2 (below). Wing veins and crossveins of a female ilnupheles mosquito. C, costa: CuA. cubitus anterior; h. humeral crossvein: M, media: M,, media-one: Ml+?. media-one-plus-two: Mz. media-two: M.1+4. media-three-plus-four: mcu, mediocubital crossvein; R, radius: R,, radius-one: rl-r,. radial crossvein: RZ, radius-two: Rz+~, radius-two-plus-three: R3, radius-three: R4+5r radius-four-plus-five; R,, radial sector: Re. remigium: SC, subcosta: sc-r. subcostal crossvein: IA. anal vein. 132 MOSQUITO SYSI-EMATICS VOL. 24. No. 2 KEY TO THE FEMALE ANOPHELES OF SOUTHWESTERN ASIA AND EGYPT

1. Wings with contrasting pale and dark spots, at least on costa (C), radius (R) and radius-one (R,) (Fig. 3) . 2

Wing entirely dark-scaled (Fig. 4) . . 26

An. (Gel.) multicolor

Fig. 4. An. (Ano.) algeriensis

2( 1). Anterior margin of wing with at least 4 separate dark areas involving the costa (C), radius (R) and radius- one (R,) (Fig. 5). Anopheles(Cellia) (in part) 3

Anterior margin of wing with fewer than 4 separate dark areas involving the costa, radius and radius-one (Fig. 6) Anopheles(Anopheles) (in part) 33

Fig. 5. An. (Cel. ) pulcherrimus

3(Z). Hindtarsomeres 3-5 (Ta-II13_<) pale (Fig. 7) ...... 4

- Hindtarsomeres 3-5 not entirely pale (Fig. 8) ...... 6

Fig. 7. An. (Gel. ) splendidus JULY 1992 133

4(3). Maxillary palpus (MPlp) with 4 pale bands (Fig. 9): abdominal terga densely covered with broad pale scales. and prominent posterolateral dark scale-tufts on all segments (Fig. 10) pulcherrimusTheobald

Maxillary palpus with 3 pale bands (spots of pale scales may also be present) (Fig. 11): abdominal terga with narrow pale scales. and dark posterolateral or apical scales on distal segments only (Fig. 12) 5

An. (Gel. ) pulcherrimus An. (Cel. ) splendidus

Fig. 12.

5(4). Maxillary palpus (MPlp) with 2 most apical pale bands broad and basal band more narrow (Fig. 13): palpomere 4 ( MPlp4) pale at base and apex (Fig. 13): femora (Fe) and tibiae (Ti) with spots of pale scales (Fig. 14) ..__ ..___...,,...._...... ____.. splendidusKoidzumi

- Maxillary palpus with apical pale band broad and 2 most basal pale bands narrow (Fig. 15): palpomere 4 usually pale at apex only. occasionally with a few pale scalesat base (Fig. 15): femora and tibiae without pale spots(Fig. 16) unnulurisVan der Wulp

Fig. 13.

An. (Cel. ) splendidus

Fig. 15.

An. (Gel. ) annularis 134 MOSQUITO SYSTEMATICS VOL. 24, No. 2 b(3). Maxillary palpus (MPlp) with 4 pale bands (Fig. 17): abdominal terga II-VII with posterolateral dark scale tufts(Fig.l8)...... ___...... _...... 7

Maxillary palpus dark. or with at most 3 distinct pale bands (pale spots may also be present) (Fig. 19): abdominal terga II-VII without dark scale-tufts. although some posterolateral dark scales may be present on distal segments (Fig. 20) . 8

Fig. 18.

“\ An. (Cd. ) pharoensis Fig. 20.

7(b). Hindtarsomeres 3 and 4 (Ta-II13.,) pale over apical half. hindtarsomere 5 (Ta-IIIs) entirely pale (Fig. 21); abdominal terga densely covered with broad pale scales (Fig. 22) pharoensisTheobald

- Hindtarsomeres 3 and 4 pale at apex only, hindtarsomere 5 dark (Fig. 23); abdominal terga II-VII covered with moderately narrow dark scales (less dense than in pharoensis),and varying amounts of pale scales mesally and posteriorly. pale scales often confined to tergum II and some distal segments (Fig. 24); tergum VIII densely covered with broad pale scales, and with some broad dark scales posterolateraliy and mesally (Fig. 24) sqzfamosusTheobald

Fig. 22.

An. (Gel. ) pharoensis

An. (Cel. ) squamosus Te

-t JULY 1992 135 g(6). Maxillary palpus (MPlp) dark (Fig. 25) 9 - Maxillary palpus with pale bands (pale spots may also be present) (Fig. 26) 10

ig. 25. An. (Cel.) azaniae Fig. 26. An. (Cel.) stephensi

9(g). Erect head scales broad, white on vertex (V) and dark brown laterally and posteriorly (Fig. 27) ,,,,,, ..,, .._...... rhodesiensismpicola Lewis (in part) (I&te 1)

Erect head scalesnarrow. straw-yellow throughout (Fig. 28) azaniaeBailly-Choumara

Fig. 27. An. (Gel. ) rhodesiensisrupicola Fig. 28. An. (Cel.) azaniae

lO(8). Hindtarsomere 5 (Ta-1115) pale (Fig. 29) ...... 11

Hindtarsomere 5 dark (Fig. 30) ...... 13

Fig. 29. An. (Gel. ) pretoriensis

Fig. 30. An. (Cel.) multicolor 136 MOSQUITO SYSTEMATICS VOL. 24, No. 2

1l( 10). Hindtarsomere4 (Ta-I&) entirely pale (Fig. 31); abdominal terga without pale scales(Fig. 32) ...... pvetoriensis(Theobald)

- Hindtarsomere 4 pale only at base and apex (Fig. 33); abdominal terga with pale scaleson at least some distal segments(Fig. 34) 12

Fig. 32.

I\ ‘i An. (Cel. ) pretoriensis \ \ An. (Gel. ) willmori

12(11). Abdominal terga II-VIII largelycovered with pale scales(Fig. 35); dark scaleson posterolateral corners of terga VII and VIII, occasionally also on IV-VI (Fig. 35) . willmori James - Abdominal terga usuallywith pale scalesat most on segmentsV-VIII, occasionallytergum IV with a few pale scalesposteriorly (Fig. 36); dark scaleson posterolateralcorners of terga VII and/or VIII. rarely also on VI (Fig. 36) ...... rnaculatzlsTheobald

Te-II Te-VIII

Fig. 35. An. (Cel.) willmori

Fig. 36. An. (Cel.) maculatus L‘ b

13(10). Wing with pale spots confined to costa (C), radius (R) and radius-one (R,) (Fig. 37); erect head scales narrow, straw-yellowthroughout (Fig. 38); scutum (Scu) with setaeonly, no scales(Fig. 39)...... _.,..,...,..._.,._...._...__...__.._...... __,.._.._...... dthali Patton - Wing with pale spotspresent on nearly all veins (Fig. 40); erect head scalesbroad, white on vertex (V) and dark brown laterally and posteriorly(Fig. 41); scutum with obvious pale scalesin addition to setae(Fig. 42) . ..___...__.___.,.__,.__..._.,,,.,.. .,._.,,,_..,__..._..._..._...... __.._...._ 14 137 JULY 1992

Fig. 37.

An. (CM.) dthali

ig. 38.

Fig. 39.

15 14(13). Palpomere5 (MPlp,) dark at apex (Fig. 43) '7 Palpomere 5 entirely pale (Fig. 44) . . . . . I I VOL. 24, No. 2 138 MOSQUITO SYSTEMAIJCS

15( 14). Scutal fossa (SF) covered with scattered pale scales (Fig. 45): base of costa (C) pale-scaled (Fig. rnzrlticolorCambouliu 46)

Scutal fossa without scales, or at most a few scales present at extreme upper margin (Fig. 47): base of costa dark(Fig.48)...... ,.._. _.. ., __ .,...... ___._..., ._____...... _._ 16

Fig. 46.

Fig. 47.

I An. (Cel.) cinereus

16( 15). Wing with well defined pale- and dark-scaled areas on all veins (Fig. 49): anal vein (IA) with 3 dark spots cinereusTheobald (Fig. 49)

Wing spots indistinct posterior to radius (R) and radius-one (R,) (Fig. 50): anal vein with at most 2 indistinct turkhudi Liston (Note 2) dark spots, often appearing mostly dark-scaled (Fig. 50).

’ ’ 1A An. (Cel. ) cinereus

An. (Cel.) turkhudi JULY 1992 139 17( 14). Scutum (SW) with broad pale scales on median area (Fig. 51): upper proepisternal setae (PeSU) (Note 3) absent (Fig. 52) ..__...... 18 - Scutum with narrow pale scales on median area (Fig. 53): upper proepisternal setae present (Fig. 54) 2 1

An. (Cel. ) stephensi

An. (Gel. ) arabiensis

1X( 17). Femora (Fe) and tibiae (Ti) spotted with pale scales (Fig. 55): abdominal terga II-VIII largely covered with pale scales (Fig. 56) --* ., ,,,...... 19 - Femora and tibiae not spotted (Fig. 57): abdominal terga without pale scales (Fig. 58) 20

Fig. Fig.

n. (eel.) superpictus VOL. 24, No. 2 140 MOSQUITO SYSTEMATICS

19( 18). Anal vein (IA) with 3 dark spots (Fig. 59); scutal fossa (SF) covered with scattered pale scales (Fig. 60): abdominal sterna V-VIII usually with pale scales (Fig. 6 1) stephensiListon

Anal vein with at most 2 small poorly defined dark spots, one just past midlength and the other near apex. or appearing entirely pale-scaled (Fig. 62); scutal fossa with pale scales only at upper margin (Fig. 63): abdominal sterna usually without pale scales, or at most with pale scales on sternum VIII, rarely a few scales on VII (Fig. 64) , n. sp. (Note 4)

An. (Cel. ) stephensi

Fig. 60. Fig.

An. (Gel.) n. sp.

Fig. 64.

Fig. S-VI

20( 18). Anal vein ( 1A) with 3 dark spots (Fig. 65) moghdensisChristophers

- Anal vein with 2 dark spots, distal spot long (Fig. 66) superpictusGrassi 141 JULY 1992

Fig. 65. An. (Gel.) moghulensis 1~

Fig. 66 . An. (Cel.) superpictus 1A

2 l( 17). Scutalfossa (SF) coveredwith scatteredpale scales(Fig. 67); hindtarsomeres. . . .3 . and 4 (Ta-II13.4). . . pale. . . . at apex22 (Fig. 68) 23 Scutalfossa without scales(Fig. 69): hindtarsomeres3 and 4 entirely dark (Fig. 70) VOL. 24. No. 2 142 MOSQUITOSYSTEMATICS

22(21). Femora (Fe) and tibiae (Ti) with spots of pale scales(Fig. 71); anal vein (1A) usually with 3 dark spots (Fig. 72); radius (R) usuallywith distinct preaccessorysector dark (PASD) spot (Note 5) (Fig. 72) ,_.____...... _. .,,,,..,..______....._...... ,,..._.___...... ____.,,. aruhiensisPatton - Femora and tibiae not spotted (Fig. 73): anal vein with 2 dark spots (Fig. 74); radius usually without preaccessorysector dark spot (Fig. 74) szthpictzwGrassi

R PASD

CC:el. ) arabiensis 1A

23(21). Radius (R) with a dark spot just distal to humeral crossvein(h) (Fig. 75); wing fringe usually with l-2 inconspicuouspale spotson posteriormargin, rarely more (Fig. 75) CulicifaciesComplex (Note 6) Radiuswithout basaldark spotjust distal to humeral crossvein(Fig. 76): wing fringe usuallywith at least 4 pale spotson posteriormargin (Fig. 76) . . . . , 24

An. (Cel.) culicifacies

An. (Cel.) sergentii

24(23). Radius-four-plus-five(R4+5) dark-scaled except at baseand apex, occasionallywith somepale scalesin distal area (Fig.77). ~ . sergentii(Theobald) - Radius-four-plus-fivewith a distinct, large median pale area (Fig. 78). . 25 JULY 1992 143

Fig. 77. An. (Cel.) sergentii

Fig. 78. An. (Cel.)Jluviatilis

25(24). Radius (R) with preaccessorysector dark (PASD) spot (Fig. 79) demeilloniEvans Radius without preaccessorysector dark spot (Fig. 80). jkviatilis James

Fig. 79. An. (Cel.) demeilloni

Fig. 80. An. (Gel.) fluviatilis

26( 1). Erect head scalesnarrow, straw-yellowthroughout (Fig.8 1) ...... Anopheles(Cellia) (in part) ...... apoci Marsh and paltrinierii Shidrawi and Gillies (Note 7) Erect head scalesbroad, white on vertex (V) and dark brown laterally and posteriorly (Fig. 82). or dark brownthroughout(Fig.83) ...... 27

Fig. 82. An. (Cel.) paltrinierii An. (Cel. ) rhodesiensisrupicola An. (Ano.) algeriensis 144 MOSQUITO SYSTEMATICS VOL. 24, No. 2

27(26). Scutum (Scu) without pale scales on median area (Fig. 84) ...... 28 - Scutum with narrow (Fig. 85) to moderately broad (Fig. 86) pale scales on median area ...... __...._..,,.....,...... Anopheles(Anopheles) (in part) 29

Fig. 84. An. (Ano.) algeriensis

Fig. 85. An. (Ano.) maculipennis Fig. 86. An. (Ano.) plumbeus

28(27). Erect head scales white on vertex (V), dark brown laterally and posteriorly (Fig. 87) ...... Anopheles(CeNia) rhodesiensisrupicola Lewis (in part)

Erect head scales dark brown throughout (Fig. 88) ...... ilnopheles(Anopheles) algeriensis Theobald

Fig. 87. An. (Cel. ) rhodesiensisrupicola Fig. 88. An. (Ano.) algeriensis

29(27). Wing scales darker and more dense at crossveins and furcations, forming dark spots (Fig. 89) ...... Maculipennis Complex (Note 8) 30 - Wing scales uniformly distributed. dark spots inapparent (Fig. 90) ...... 3 1 An. (Ano.) claviger

30(29). Wing with distinct dark spots(Fig. 91); scutum (Scu) dark brown (Fig. 92);.. scutal ...... fossa .... (SF) .... usuallywith narrow, pihform pale scales,atleast on extreme upper margin (Fig. 92) ...... mucdipennisMeigen and subalpinusHackett and Lewis (Note 9) Wing spotsusually less distinct (Fig. 93): scutum pale brown (Fig. 94); scutalfossa without pale scales(Fig. 94) martinius Shingarevand sacharoviFavre (Note 10)

An. (Ano.) maculipennis

An. (Ano.) sacharovi 146 MOSQUITO SYSTEMATICS VOL. 24, No. 2

3 l(29). Labellum (La) distinctly paler than remainder of proboscis(P) (Fig. 95)...... marteri sogdianusKeshishian (Note 11)

Labellum not paler than remainder of proboscis(Fig. -_ 96) ...... 32

An. (Ano.) marteri sogdianus

32(31). Scutum (Scu) with very narrow, piliform pale scaleson median area (Fig. 97); lower proepisternalsetae (PeSL) (Note 12) present (Fig. 98); palpomere 5 (MPlp,) not longer than 0.50 length of palpomere 4 (MPlp,) (Fig. 99) claviger(Meigen) Scutum with narrow to moderately broad, spatulate pale scales on median area (Fig. 100); lower proepisternal setae absent (Fig. 101); palpomere 5 longer than 0.50 length of palpomere 4 (Fig. 102)...... plumbeusStephens

Fig

’ Fig. 99.

An. (Ano.) claviger

’ Fig. 102.

An. (Ano.) plumbeus JUI.Y 1992 147

33(Z). Basolateral area of clypeus (Clp) with a patch of dark laterally projecting scales (Fig. 103) 34

Clypeus without scales (Fig. 104) 39

An. (ho.) lindesayi

34(33). Hindtarsomeres 4 and 5 (Ta-III4 ,) entirely pale (Fig. 105) 35

Hindtarsomeres 4 and 5 not entirely pale (Fig. 106) _,._____.,_____.______,.___,.._ 36

Fig. 105. An. (Ano.) coustani Fig. 106. An. (ho.) gigassimlensis

35(34). Hindtarsomere 1 (Ta-III,) broadly pale at base and apex. hindtarsomere 2 (Ta-1112) pale over approximately apical half. hindtarsomere 3 (Ta-IIIj) dark at base only or entirely pale (Fig. 107); abdominal sternum VII with a group of posteromedian dark scales (Fig. 108) coustuniLaveran

- Hindtarsomeres 1 and 2 narrowly pale at apex only. hindtarsomere 3 pale over apical third to two-thirds (Fig. 109): abdominal sternum VII with or without posteromedian dark scales (Fig. 110) ...... trnehroszoDoenitz (Note 13)

VII

Fig. 108.

An. (Ano.) coustani An. (Ano.) tenebrosus 148 MOSQUITO SYSTEMATICS VOL. 24. No. 2

36(34). Hindtarsomere 4 (Ta-I&) pale at apex only or entirely pale, hindtarsomere 5 (Ta-1115) dark (Fig. 11I) 37

- Hindtarsomere 4 usually pale at base and apex, hindtarsomere 5 entirely dark or pale at base only (Fig. 112).... ,,...... ,._. . .._...... ,,...... 38

___~ ___--_- _-y ’ < _--l_--- s - -- To-Ill,

Fig. 111. An. (Am.) pseudopictus Fig. I 12. An. (Ano.) peditaeniatus

37(36). Hindtarsomere 4 (Ta-IIIJ) pale at apex only (Fig. 113) hyrcanzrs(Pallas)

Hindtarsomere 4 entirely pale (Fig. 1 14) pseudopictusGrassi

.- _?--_-___ _--- 5 _-__ _._ TCI-Ill,

Fig. 113. An. (Ano.) hyrcanus Fig. 114. An. (Ano.) pseudopictus

38(36). Humeral crossvein (h) with patch of dark scales (Fig. 115); remigium (Re) mostly dark-scaled (Fig. 115); pale markings on hindtarsomeres 4 and 5 (Ta-IIIJ.S) variable. often without basal pale bands (Fig. 116) _, _, _. _. _. _. .nigerrimusGiles (Note 14)

- Humeral crossvein without scales (Fig. 117);remigium mostly pale-scaled (Fig. 117): hindtarsomere 4 and usually 5 with basal pale bands (Fig. 1 18) peditaeniatzrs(Leicester)

R.? h

Fig. 115. Fig. 117. / /

Fig. 116. Fig. 118. An. (Ano.) nigerrimus An. (Ano.) peditaeniatus 149 JL I.Y 1992

39(33). Costa (C) dark-scaled except at apex (Fig. 119); hindfemur (Fe-III) with broad subapical pale band (Fig. 120); hindtarsomeres4 and 5 (Ta-IILi5) entirely dark (Fig. 12 1) iin&~u~YGiles

- Costa with several pale-scaled areas prior to apex (Fig. l-_77): hindfemur narrowly pale at base and apex only (Fig. 123): hindtarsomeres 4 and 5 narrowly pale at tarsal joints (Fig. 124). g&m simiensis(James)

Fig. 119.

Fig. 121. An. (Ano.) lindesayi

Fig. 122.

Fig. 124. An. (Ano.) gigassimlensis

EXPLANATION OF NOTES

I. Anopheles(Gel. ) rhodesiensisrupicola can from “Palestine” was not seen during this be found with (couplet 9) and without (cou- study. Anophelesturkhudi was found during plet 28) contrasting pale and dark spots on a faunistic survey of the Sinai Peninsula(Mar- the costa(C), radius (R) and radius-one (Ri). galit and Tahori 1973), but it was not listed This variation of the wing scalingis often seen by Pener and Kitron ( 1985) in a survey of in desert speciesand may not be confined northern Israel. In an annotated list of the solely to the C, R and R1. mosquitoes of Israel (Margalit and Tahori 2. The variety An. (Gel.) turkhuditelamali 1974), they record only the subspeciesAn. described by Saliternik and Theodor (1942) twkhudi telamali;however, J. Margalit (per- 150 MOSQUITO SYSTEMATIC-S VOL. 24. No. 2

sonal communication) feels that the statusof species,including An. maculipennis,An. mar- An. turkhudi telamali as a valid subspecies tinius, An. sacharovi and An. subalpinus. may be in doubt. Identifications of the speciesare best accom- 3. The upper proepisternal setae(PeSU) oc- plished by charactersof the eggsor by differ- cur in a group above the forecoxae on the ences in chromosomal inversions (White upper part of the proepisternum. 1978). 4. The dark spots on the anal vein (IA) of 9. Anopheles (Ano.) maculipennis and An. An. (Ccl. ) n. sp. may be reduced to the point subalpinuscan be distinguishedby the inter- that the entire vein appearspale-scaled. How- costal membrane of the egg float, which is ever, the speciescan be separated from An. rough in An. maculipennisand smooth in An. stephensiby the combination of the reduction subalpinus, and by chromosomal inversion in pale scaleson the scutal fossa(SF), and the differences.Anopheles subalpinus is known to reduction or absence of pale scales on the occur with certainty only in Iran, Iraq, Syria abdominal sterna. and Turkey. Postiglione et al. (1970) found 5. The preaccessorysector dark (PASD) spot An. subalpinusin Turkey, suggestingthat pre- is defined as the isolated group of dark scales vious records of An. messeaeFalleroni may occurringon the radius(R) before the splitting also be An. subalpinus. White (1978) shows of radius-one (R,) and the radial sector (R,), the geographic distribution of An. messeae and located between the sector pale (SP) and falling short of the Southwest Asian Region. accessorysector pale (ASP) spots.The PASD The presence of An. melanoon Hackett in is equivalent to the sector dark (SD) spot of Turkey requires confirmation. Wilkerson and Peyton (1990) as illustrated 10. Anopheles martinius and An. sacharovi in their Figure 1 for the condition where the can be distinguishedby the fixed paracentric ASP is present on the costa(C), subcosta(SC) inversions of their polytene chromosomes and R, thereby producing SD spots on all (White 1978). three veins. The typical condition for the pres- 11. Ribeiro et al. (1985) collectedAn. (Ano.) ence of the isolated PASD spot on the R in marteri Senevet and Prunelle in northeastern SouthwestAsian Cellia is the absenceof the Portugal, and after reviewing its geographical ASP spotsand the fusing of the SD spotson distribution, bioecology and taxonomy from the C and SC. the literature, determined that An. (Ano.) 6. Anopheles (Gel. ) culicifacies has been marteri sogdianusis a junior synonym of An. found to be a complex of four sibling species marteri, and that An. marteri is a polymor- (designatedspecies A, B, C and D), distin- phic, monotypic species.However, based on guishableonly by fixed chromosomal inver- their conclusionsconcerning the distribution sions. SpeciesA has been identified from an of subspeciesand expected morphological di- urban area on the border between Oman and vergenceof subspecies,An. marteri sogdianus the United Arab Emirates by Akoh et al. will be treated as a valid subspeciesin this (1984) and from Sistan and Baluchistan study. Province, Iran by Zaim and Javaherian 12. The lower proepisternalsetae (PeSL) oc- (199 1). SpeciesA and B have been detected cur individually or as a group mesad of the in Punjab Province, Pakistan (Mahmood et forecoxae and below the upper proepisternal al. 1984). setae(PeSU) on the lower part of the proepis- 7. Adult females of An. (Gel. ) apoci and An. ternum. (Gel. ) paltrinierii cannot be distinguishedex- 13. The posteromedian dark scales of ab- cept for the morphology of the pharyngeal dominal sternum VII in An. (Ano.) tenebrosus armature. Males of An. pahrinierii can be were occasionally absent in specimensfrom separated by the absence of leaflets on the Egypt, Israel and Saudi Arabia. However, the aedeagus(Shidrawi and Gillies 1987). pale banding of hindtarsomeres l-3 is a reli- 8. In southwestern Asia the Maculipennis able character for distinguishingAn. tenebro- Complex is representedby at leastfour sibling sus from An. coustani. JULY 1992 151 14. The adults of An. (Ano.) nigervimusare Bates,M., W.N. Beklemishevand L. La Face. generally similar to An. peditaeniatus, and 1949. Anophelines of the Palearctic Re- apparently the extent of pale banding on the gion, pp. 4 19-442. In: M.F. Boyd (ed.). hindtarsomeresis often variable. SeeHarrison Malariology. A comprehensive survey of and Scanlon (1975) for a discussionof char- all aspectsof this group of diseasesfrom a acterswhich help to separatethe two species. global standpoint. Vol. I. W.B. Saunders Early recordsof An. nigerrimusfrom Pakistan Co., Philadelphia. may be An. peditaeniatus(B.A. Harrison, per- Bonne-Wepster, J. and N.H. Swellengrebel. sonal communication). 1953. The anopheline mosquitoes of the Indo-Australian Region. J.H. de Bussy, ACKNOWLEDGMENTS Amsterdam. Christophers,S.R. 1933. The fauna of British The author is grateful to Ralph E. Harbach, India, including Ceylon and Burma. Dip- E.L. Peyton, Ronald A. Ward and Richard C. tera. Vol. IV. Family Culicidae. Tribe An- Wilkerson, Walter Reed Army Institute of ophelini. Taylor and Francis, London. Research (WRAIR), for their helpful com- Cianci, R., A. Sabatini, D. Boccolini, L. Bul- ments and review of the manuscript; Bruce lini and M. Coluzzi. 1987. Electrophoretic A. Harrison, National ResearchCouncil, for evidence of reproductive isolation between accessto his extensivenotes and reprint files, sympatric populations of Anophelesmela- and for his helpful comments and review of noon and Anophelessubalpinus, p. 156. In: the manuscript; Deborah Feher for preparing Proceedings of the Third International the illustrations;and Taina Litwak, WRAIR, Congressof Malaria and Babesiosis,An- for assistancein preparing the manuscript for nency. publication. Colbourne, M.J. and S.A. Smith. 1964. Prob- REFERENCES CITED lems of malaria in the Aden Protectorate (report on a visit). WHO/Ma1/442. Abdel-Malek, A.A. 1958. The anopheline Coluzzi, M. 1964. Morphological divergences mosquitoes of northern Syria. 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