1Also collaborator, Walter Reed Biosystematics Unit (WRBU), Smithsonian Institution, Washington DC, USA.

scales mostly broad and flat. Male coxite with strong apical lobe; basal lobe usually with a number of flattened hairs; claspette unusually large, with stout stem. Larva (except in A. stampari) with several hairs on siphon in addition to the usual pair of tufts." Most subsequent authors followed Edwards’ (1932) group classification including Natvig (1948:381) and Gutsevich et al. 1970 (English translation, 1974). Stone et al. (1959) included within Ochlerotatus the species listed by Edwards (1932) without a segregation by group and they included Ae. pseudorusticus Callot as a synonym of Ae. refiki. Stone (1963) listed Ae. stampari in synonymy with Ae. refiki. Mohrig (1969:95) follows the rusticus group classification of Edwards (1932) while treating the two species (i.e. Ae. refiki and Ae. rusticus) occurring in Germany. His studies were focused on an investigation comparing the female genitalia of subgenera in the genus Aedes occurring in that country and showed these two species (rusticus group) occupied a certain unusual position within the subgenus Ochlerotatus. Knight & Stone (1977), in the current "Catalog of the Mosquitoes of the World," listed subgenus Rusticoidus and included only Ae. refiki with the synonym Ae. pseudorusticus, however, the other species of Edwards’ (1932) rusticus-group were retained in Ochlerotatus. Dahl & White (1978:393) again synonymized Ae. stampari with Ae. refiki, following Martini (1931); this was also reported by Ward (1984:269). Ward (1987:94) stated "One species, Ae. refiki Medschid, was transferred to the monotypic subgenus Rusticoidus by Shevchenko and Prudkina (1973). Although this subgenus is well recognized (Knight & Stone 1977), no mention was made of the current status of Ae. refiki." Alekseev (1989) described the new species, Aedes (Rusticoidus) krymmontanus, and provided on page 177 the following information. The subgenus Rusticoidus at present in the USSR and adjacent countries comprised 5 species (Gutsevich et al., 1970, Gutsevich & Pendikova, 1972). On the basis of our material, as well as on the figures and descriptions of the larvae and adults of species of the subgenus (Gutsevich et al., 1970; Dubitzky, 1970; Martini, 1931; Medschid, 1928; Rioux, 1965; Roman, 1958; Gilot, 1968; Gilot et al., 1971; Mohrig, 1969) a table of characters diagnostic for the species has been constructed. Because of insufficient description and figures, as well as citation of range Ae.(R.) albescens has been omitted. Alekseev (page 176) provided a table of diagnostic characters of comparable species of mosquitoes of subgenus Rusticoidus (i.e. Ae. krymmontanus, Ae. rusticus, Ae. refiki, Ae. lepidonotus and Ae. subdiversus). This table included one male genitalic and 12 larval characters. A translation from Russian to English of the article was provided by the late George C. Steyskal, Gainesville, Florida. Ward (1992:189), for subgenus Rusticoidus, included citations of Ae. krymmontanus and Ae. refiki. Schaffner (1992:80, 188) raised the question whether Ae. rusticus and the other members of Edwards’ (1932) rusticus-group should be included in the subgenus Rusticoidus along with Ae. refiki. Later, Schaffner (1998:4) stated "The sub-genus Rusticoidus was described by Shevchenko & Prudkina (1973) following a study of the male genitalia in the genus Aedes. They proposed Ae. refiki as haplotype, and included in the sub-genus the related species previously incorporated in the rusticus group. Even though the catalogue of mosquitoes (Knight & Stone, 1977 and its additions) classifies Ae. refiki as being the only European species in this sub-genus, we share the opinion of other authors (Shevchenko & Prudkina, 1973; Encinas Grandes, 1982) that Ae. rusticus is also a member of that subgenus." In a series of articles Stojanovich & Scott (1993:16; 1995a:17; 1995b:15; 1995c:14; 1995d:19; 1996:6; 1997:18) included Ae. refiki, Ae. rusticus and Ae. subdiversus (one or more species per article) in subgenus Rusticoidus but indicated that some authors placed Ae. rusticus and Ae. subdiversus in subgenus Ochlerotatus. Snow & Ramsdale (1999:16) provided a chart giving the distribution for the species Ae. krymmontanus, Ae. lepidonotus, Ae. quasirusticus, Ae. refiki, Ae. rusticus and Ae. subdiversus, and even though it was not stated that the species belonged to the same subgenus, they were grouped together in the chart.

Because of the uncertainty of which species were assigned to subgenus Rusticoidus by Shevchenko & Prudkina (1973), several articles published since have retained one or more of the species of the "rusticus group" in subgenus Ochlerotatus.

From the above chronology, and my examination of specimens and species descriptions, I concur with Alekseev (1989) in the inclusion of the following European species in the subgenus Rusticoidus: Ae. krymmontanus, Ae. lepidonotus, Ae. refiki, Ae. rusticus, and Ae. subdiversus. Aedes quasirusticus Torres Canamares (1951) also should be included in Rusticoidus. The western Asian Aedes albescens should be questionably included in the subgenus pending a redescription and illustrations of the species to include adults, male genitalia, pupa and larva. Gutsevich & Pendikova (1972) and Gutsevich & Dubitzky (1981; 1987) should be examined for supplemental information concerning Edwards’ (1921) description of Ae. albescens and its separation from Ae. subdiversus. The related North American species will be treated by the author in a separate article.

The following is an expanded definition of subgenus Rusticoidus and is provided for comparison with the other subgenera of Aedes, especially subgenus Ochlerotatus. This definition is based on the examination of specimens of Ae. lepidonotus (C , C g, X , X g, L), Ae. refiki (C , C g, X , X g, L), Ae. rusticus (C , C g, X , X g, P, L), Ae. quasirusticus (C , C g, X , X g, P, L) and Ae. subdiversus (X , X g), and the published descriptions and illustrations of all included species. Symbols and abbreviations used above are: C = female, X = male, P = pupa, L = fourth-instar larva, and g = genitalia. Nomenclature used follows Harbach & Knight (1980).

Adults. Relatively large-sized mosquitoes with somewhat shaggy appearance. Head: Eyes separated in front; vertex and occiput covered with curved narrow scales; numerous erect forked scales on occiput and lateral areas of vertex; ocular line with curved narrow scales and numerous well developed setae; maxillary palpus of female brown with scattered pale scales; maxillary palpus of male noticeably longer than proboscis, palpomere 3 long with distal portion somewhat swollen, upturned, with numerous long (longer than palpomere 4 length), forward-directed setae lateroventrally, palpomeres 4,5 somewhat downturned, palpomere 4 with numerous moderately long setae ventrolaterally, palpomere 5 with several short to moderately long setae lateroventrally and more numerous on proximal portion (maxillary palpus developed similar to many Ochlerotatus). Antenna: Pedicel of female with 2 large patches of partly overlapping broad scales, one covering mesal surface and other covering lateral surface, patches maybe contiguous dorsally in some species. Thorax: Scutum covered with narrow scales, including most of prescutal area, 1-6 on parascutellar area, antealar and supraalar areas usually with curved narrow scales somewhat broader, no broad scales present; scutal setae (most on anterior half relatively short)--few anterior promontory, numerous anterior and posterior dorsocentral and acrostichal, scutal fossal (anterior, lateral, median and posterior), few posterior medial scutal, several prescutellar, several antealar, numerous supraalar, and 1-3 parascutellar; scutellum with curved narrow scales and numerous long setae on all lobes; pleural areas with scales broad and relatively long (occasionally few narrow scales on paratergite, hypostigmal and lower metepisternal areas in some species), most pale, occurring on following areas--large patch on antepronotum, large patch on upper proepisternum, large patch covering postpronotum, large elongate patch on subspiracular area, patch on hypostigmal area, patch on postspiracular area, elongate patch on lateral and ventral margins of paratergite, large patch covering upper and posterior areas of mesokatepisternum, patch on lower prealar area and often extending onto upper area, large patch covering mesepimeron from near ventral margin dorsally to anterodorsal setal patch and also posterodorsally to setae, patch covering metameron, patch on membrane posteromesad of metapostnotum; most pleural setae long, and as follow--numerous (30+) antepronotal, numerous (40+) upper proepisternal, several posterior postpronotal, few postspiracular, few upper and several posterior mesokatepisternal, numerous (25+) prealar, numerous upper and 1-9 (usually 2-5) long lower mesepimeral; mesomeron with dorsal margin well above base of hindcoxa. Legs: Patches of broad scales on both antecoxal and postcoxal membranes of foreleg; posttarsi I-III of female and male each toothed, larger unguis of I,II with 2 teeth in male. Wing: Costa with pale scales at base; remigium with 1-6 (usually 2,3) long setae. Abdomen: Terga with brown scales but with extensive pale-scaled areas.

Female genitalia. Tergum VIII: With basal 0.4-0.5 retracted into segment VII; base slightly concave; apex straight and wide; numerous setae and usually numerous moderately broad scales on most of area; 4-8 moderately long setae on apical margin. Sternum VIII: Base slightly convex to straight; apex with very small median indentation and broadly rounded lobe on each side of midline; numerous short and moderately long setae and moderately broad scales on most of area; apical margin with row of setae, short mesally and increasing in length to moderately long laterally. Tergum IX: Moderately wide; apex with moderately deep median indentation; several setae apically. Insula: Liplike; with several moderately long setae. Lower vaginal lip: Covered with short to moderately long spicules; narrow; lightly pigmented; lower vaginal sclerite absent. Upper vaginal lip: Covered with short to moderately long spicules; narrow; moderately to heavily pigmented; caudal margin curved; upper vaginal sclerite absent. Spermathecal eminence: Membranous; nonpigmented; ill-defined; large patch of short thin simple spicules on basomesal area. Postgenital lobe: Moderately long; moderately wide; with small median apical indentation; ventral surface narrowly attached to upper vaginal lip; with number of setae. Cercus: Moderately long; moderately wide; apex sharply rounded with few long stout setae; dorsal surface with number of short and few moderately long setae and numerous moderately broad scales; ventral surface with number of short setae on apical and lateral areas. Spermathecal capsules: 1 large- and 2 medium-sized ones; heavily pigmented; spherical.

Male genitalia. Prerotation position. Tergum IX: With narrow lobe on each side of midline bearing several short, stout, flattened setae apically. Gonocoxite: Long; moderately broad; dorsal surface with basal lobe bearing specialized setae mesally. Gonostylus: Long; narrow; curved mesally; with 2-4 short setae subapically on lateral margin; gonostylar claw moderately long, narrow, heavily pigmented, with apex truncate. Claspette: With long, moderately thick, curved stem, distal portion somewhat broader; filament short, with transverse annulations. Phallosome: Aedeagus simple, more or less troughlike, relatively long, narrow, median area narrower than basal and apical portions, base with more or less circular opening (bearing short triangular flap laterally on each side in some species), apex with median small lobe separating short flattened area with few tiny, stout spicules on each side; paramere moderately long to long, narrow; basal piece moderately long. Proctiger: Moderately long; shorter than aedeagus; paraproct heavily pigmented, apex curved into blunt point; few short cercal setae. Sternum IX: Relatively short; with heavily pigmented, median, caudal, small, rounded area bearing few long and short setae; heavily pigmented, short, triangular strip laterally on each side.

Pupae. Cephalothorax: Setae 4,5-CT with short thin branches, approximately equally developed; 7-CT with moderately long thin branches; 11-CT moderately stout, closer to 10-CT than to 12-CT. Trumpet: Moderately long; pinna moderately wide. Abdomen: Seta 1-II-VII with thin branches, 1-II, III moderately long, 1-IV-VII moderately long to long; 2-II, VII laterad of 1-II, VII; 2-III-VI mesad of 1-III-VI; 5-IV-VI very long, stout; 6-VII with short fine branches; 9-VII, VIII relatively short, with moderately stout branches, 9-VII noticeably cephalad and slightly laterad of 6-VII, 9-VIII very lightly aciculate. Paddle: Outer width from midrib greater than inner width; length noticeably longer than width; basal portion of outer margin with minute serrations, apical portions of outer and inner margins with minute spicules; seta 1-Pa short, usually single (occasionally 2-forked or branched).

Fourth-instar larvae. Head: Seta 1-C long, stout, curved mesad, simple; 4-C short, with fine branches, mesad and slightly cephalad of 6-C; 5-C long, stout, aciculate, caudad of 4,6-C, slightly mesad of 6-C and slightly laterad of 4-C; 6-C long, stout, aciculate; 7-C moderately long to long, stout, fanlike, with multiple aciculate branches, laterad of 4-6-C and slightly cephalad of 4,6-C, shorter than 5,6-C; 8-10-C moderately long, fine; 11,12-C short, with multiple fine branches; 13-C long, single; 14-C short, relatively stout; 15-C short, with thin branches. Antenna: Relatively short, heavily pigmented, shaft with numerous short stout spicules over entire length; seta 1-A short, stout, with aciculate branches; 2-6-A inserted at apex of shaft. Thorax: Seta 1-P long, aciculate; 12-P very long, aciculate, single. Abdomen: Seta 12-I short; 1-VII long, stout, aciculate; 1,3,5-VIII with multiple, stout, aciculate branches, 1-VIII noticeably shorter than 3,5-VIII; 4-VIII shorter than 3-VIII; comb on segment VIII consisting of several stout scales in 1 or 2 irregular rows, scales with long stout median spine and few short denticles basolaterally; saddle incompletely rings segment X, without acus, without stout posterior spines; 1-X long, stout, aciculate; 4-X consisting of several long, multiple-branched setae on well developed grid and at least 2 short, branched, precratal setae. Siphon: With well developed attached acus; seta 1-S long, stout, with multiple aciculate branches; 1 accessory seta laterally before midlength of siphon, inserted slightly above pecten proximally to 1-S, short, usually branched; at least 1 additional accessory seta (often several accessory setae that are stout and aciculate) inserted dorsally or slightly subdorsally on siphon; 6-S long, single; 8-S short, branched; 9-S developed as moderately stout hook; pecten consisting of several spines with at least 1 distal spine wider spaced than the remainder.

Eggs. The following are pertinent features that I feel may be of subgeneric importance and are extracted from the description of the egg of Ae. rusticus given by Service et al. (1997). Shape: Short but very wide; in profile almost subtriangular with rounded corners; ventral surface arched in middle; about equally rounded at both ends. Outer chorion: Dorsal, lateral, ventral, anterior and posterior areas more or less similarly developed; chorionic cells mainly roundish or elliptical, few quadrilateral or pentagonal with straighter sides; each cell encloses single variously sized, but usually large, tubercle with finely granular sloping sides and domed tuberculate top. Micropyle: Collar not very prominent; orifice not obvious.

Discussion.

Adults of the subgenus Rusticoidus are similar to some Holoarctic species of Ochlerotatus in habitus but are distinguished from all (?) species of this subgenus and the other subgenera of Aedes by possessing the following combination of six characters (1) both antecoxal and postcoxal membranes of foreleg have a patch of broad white scales, (2) antennal pedicel of female has two large patches of partially overlapping broad white scales, one patch covering the mesal surface and the other covering the lateral surface (patches maybe contiguous dorsally in some species), (3) postpronotum is covered with only broad flat scales, (4) parascutellar area has 1-6 scales in addition to 1-3 long setae, (5) hypostigmal area has a patch of broad scales, and (6) membrane posteromesad of metapostnotum has a patch of broad scales. Some species of Ochlerotatus possess one or more of the above characters but not all six. Other adult features that are useful in separating Rusticoidus from the other subgenera of Aedes are (1) scutum and scutellum having only narrow scales, (2) antepronotum, upper proepisternum and prealar area each has a large number of long setae, (3) metameron is covered with broad scales, (4) mesepimeron has 1-9 long setae on anteroventral area, (5) subspiracular area has an elongate patch of broad scales, and (6) male posttarsi having each unguis with a tooth and the larger unguis of posttarsi I and II has two teeth.

Adults of Ae. lepidonotus are interesting in that they possess a patch of golden narrow scales and 2 short golden or white setae on the median caudal area of the mesopostnotum. Species of the subgenus have 1-7 scales on the lower portion of the metepisternum (seen in specimens of Ae. lepidonotus, Ae. quasirusticus and Ae. rusticus but these scales may have been rubbed off in other species of the subgenus).

Male genitalia of Rusticoidus are distinctive in the development of the aedeagus (see above) and the claspette being moderately thick and having a short, transversely annulated filament. Other features, used in combination, that are useful in separating Rusticoidus from many other subgenera of Aedes are: tergum IX having a narrow lobe on each side of the midline and bearing several short, stout, flattened setae; sternum IX relatively short, having heavily pigmented areas restricted to a small, median, caudal, rounded area bearing a few long and a few short setae, and narrow basal strip connected to the lateral margins that have a short, triangular strip; and the gonocoxite with a basomesal lobe on the dorsal surface bearing specialized setae. Each species of the subgenus has distinctive features of the male genitalia (e.g., Ae. rusticus having the gonocoxite with the dorsal surface expanded into a narrow lobe bearing several moderately long, broad, fusiform scales ventrally, and the gonostylar claw is sinuous).

The pupal stage of Rusticoidus is known only for Ae. quasirusticus and Ae. rusticus and is imperfectly described and illustrated. Unfortunately, the specimens of these two species available for examination were in poor condition and no features of subgeneric rank could be observed for this stage.

Fourth-instar larvae of Rusticoidus are generally similar to many species of Ochlerotatus but are easily distinguished from these and all subgenera of Aedes by having the siphon with a short (usually branched) accessory seta laterally before the midlength and inserted slightly above the pecten and proximal to seta 1-S. Other features that are useful in separating Rusticoidus larvae from those of other subgenera of Aedes are (1) seta 12-P is very long, stout, aciculate and single, (2) siphon has at least a second accessory seta (often with several setae that are stout and aciculate) inserted dorsally or slightly subdorsally, (3) positions and development of setae 4-7-C are as described above, (4) saddle does not ring segment X, is incomplete ventrally and is without an acus or stout spines on the posterior margin, (5) seta 9-S is developed as a moderately stout hook, and (6) antenna is relatively short, heavily pigmented, and with numerous stout spicules over the entire length of the shaft.

Larvae with accessory setae on the siphon also are found in the subgenus Aedes and a few species of subgenus Ochlerotatus (e.g., Ae. crinifer (Theobald) (see Arnell 1976); Ae. hexodontus Dyar, Ae. hexodontus hokkaidensis Tanaka, Mizusawa & Saugstad and Ae. punctor (Kirby) (see Tanaka et al. 1979) but none of these species have an accessory seta inserted laterally on the siphon before midlength, slightly above the pecten and proximal to seta 1-S. Seta 12-P developed as a very long, aciculate, single seta is also infrequently found in some Ochlerotatus species (e.g., Ae. punctor). Other features mentioned above also are found in some species of Ochlerotatus.

In subgenus Rusticoidus the egg of the European Ae. rusticus has been described and/or illustrated (e.g., Marshall, 1938:85; Natvig, 1948:39; Hinton, 1981:425; Service et al., 1997:61). Service et al. (1997) provided a detailed description of this species and included several scanning electron microphotographs of the entire egg and other features. This article should be consulted for details of the egg. The chorionic pattern of Ae. rusticus eggs is similar to several species of Ochlerotatus (e.g., Ae. cantans (Meigen), Ae. punctor, Ae. vittiger (Skuse)) and, even though the shape is characteristic, it also resembles some species of Ochlerotatus (e.g., Ae. aculeatus (Theobald), Ae. vittiger).

Appreciation is expressed to D.R. Barnard and H. Oberlander (CMAVE) for providing facilities for conducting this study; to T.M. Howard (The Natural History Museum (NHM), London, UK) and T.V. Gaffigan (WRBU) for providing type and other specimens; and to R.E. Harbach (NHM), R.C. Wilkerson and M.A.M. Sallum (WRBU), and J.A. Seawright (CMAVE) for reviewing the manuscript.

______________________________________________________________________________________________

Ralph E. Harbach

Department of Entomology, The Natural History Museum, Cromwell Road, London SW7 5BD, U.K.

E-mail: r.harbach@nhm.ac.uk

The purpose of this note is to clarify the specific name of the species of Culex in southern Europe commonly known as Culex univittatus. It appears that continued use of the name univittatus Theobald, 1901 for this species stems from Minar (1991), who listed perexiguus Theobald, 1903 as a junior synonym of this epithet. Contrary to the evidence, I am surprised that univittatus is still in use for this taxon. It is obvious that Minar (1991) prepared his contribution to the Catalogue of Palaearctic Diptera without knowledge of my monograph on the subgenus Culex in southwestern Asia (Harbach, 1988). It seems he only had access to literature that was 10-12 years out of date, and simply repeated what was listed in the world catalogue of mosquitoes by Knight & Stone (1977), which he incorrectly cited as 1978. He did not reference the supplement to the world catalogue by Knight (1978) which listed Cx. perexiguus as a valid species (after White, 1975). Furthermore, Minar apparently had no knowledge of the crossmating experiments performed by Jupp (1971) (Cx. univittatus x Cx. neavei Theobald) and Jupp & Harbach (1990) (Cx. perexiguus x Cx. neavei). The available data indicate that Cx. univittatus is restricted to the temperate highlands in the East African Subregion of the Afrotropical Region, while Cx. perexiguus inhabits more arid areas in eastern Africa and the Mediterranean Subregion of the Palaearctic. I have examined only a few specimens from southern Europe (Greece, Italy and Turkey), but these appear to be Cx. perexiguus based on characters of the male genitalia and larvae. Hence, based on the available evidence, the species of the Univittatus Complex that occurs in southern Europe should be regarded as Cx. perexiguus rather than Cx. univittatus.