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Two mosquito species, Culex (Culex) hepperi and Culex (Culex) maxi are reported for the first time for Bolivia, in Tarija department. Aedes (Ochlerotatus) stigmaticus and Mansonia (Mansonia) indubitans are the first records in Aniceto Arce province, in Tarija department, southern Bolivia. In addition, Aedes (Protomacleaya) alboapicus and Culex (Phytotelmatomyia) renatoi are reported for the first time in the northwestern region of Argentina. Anopheles (Nyssorrhynchus) nuneztovari s.l. is reported for the first time in Jujuy and Tucumán provinces, and Aedes (Ochlerotatus) raymundi and Anopheles (Anopheles) neomaculipalpus are the first reports in Tucumán province. The 3 species are extending their geographical distribution in these provinces. Data on the collection localities and comments about the medical relevance of some species are also presented.

Refereed/Peer-reviewed Japanese encephalitis virus (JEV) continues to cause significant numbers of human infections and fatalities despite the availability of efficacious vaccines. It is regarded as an emerging mosquito-borne pathogen with the potential of introduction into many countries. In 2022, JEV was detected in Australia on a hitherto unprecedented scale, with local transmission by indigenous mosquitoes to amplifying swine hosts and to humans. In this study, we review this recent disease activity, propose possible routes of virus movement, ecological drivers of activity, and consider possible future transmission scenarios. Measures to enhance current surveillance systems and potential strategies for health authorities to minimize future risks are discussed.

To calculate human biting rates for various mosquito species, we performed simultaneous collections for 15 wk at 6 ecologically variable sites in Hungary. Of the dominant species, the relative abundance of Aedes vexans, Ae. sticticus, and Coquillettidia richiardii showed a significant positive correlation between CO2 + Biogents lure and human landing catch (HLC). The relative abundance of Culex pipiens was significantly lower in the HLC samples than in the CO2 + BG lure samples. Of the invasive species, Aedes korecius was found more frequently in HLC, while Ae. japonicus was more common in CO2 + BG lure samples. Estimated human biting rates, determined with the 2 collection methods, showed no significant differences at high mosquito density (100–120 bites/h/person), but there was considerable variation at low mosquito biting rates. Therefore, correcting the CO2 + BG lure trapping data to include only species biting humans provides estimates approaching the values of the HLC. Our study confirmed that while HLC is the gold standard method for determining the human biting rate, provided appropriate data adjustments are made, trapping methods performing automated data collection can provide similar data while reducing the exposure of the data collector.

The emergence of West Nile virus and Usutu virus in Europe poses a significant risk to public health. In the absence of efficient antiviral therapy or vaccine candidates, the only strategy to control these arboviruses is to target the Culex (Diptera: Culicidae) mosquito vector. However, the selection pressure caused by exposure to insecticides for vector control or agricultural pest control can lead to insecticide resistance, thereby reducing the efficacy of insecticide-based vector control interventions. In Culex mosquitoes, two of the most common amino acid substitutions associated with insecticide resistance are the kdr L1014F in voltage gated sodium channels and G119S in acetylcholinesterase. In this study, Culex pipiens biotype pipiens, Culex torrentium, and Culex modestus were sampled from 2019 to 2021 in three distinct environmental habitats (urban, peri-urban, and agricultural) in and around the city of Leuven, Belgium. Individual mosquitoes were screened for two mutations resulting in L1014F and G119S amino acid substitutions. Both mutations were observed in Cx. pipiens and Cx. modestus but not in Cx. torrentium mosquitoes across the four collection sites. Furthermore, multi-resistance or cross-resistance in Cx. pipiens could be a threat in these areas, as both mutations were observed at low frequencies. These results provide the first report of kdr L1014F and ace-1 G119S resistance mutations in Cx. pipiens and Cx. modestus mosquitoes from Belgium, highlighting the importance of mosquito surveillance to design effective arbovirus outbreak control strategies.

In Mexico, the genus Uranotaenia includes 11 species distributed mainly in the tropical and subtropical regions in the southeast of the country. Uranotaenia sapphirina has been reported in 18 states in Mexico: Campeche, Coahuila, Colima, Chiapas, Guerrero, Hidalgo, Jalisco, Mexico City, Mexico State, Michoacán, Morelos, Oaxaca, Quintana Roo, Sinaloa, Tabasco, Tamaulipas, Veracruz, and Yucatán; whereas Ur. socialis has been reported in Chiapas and Quintana Roo. In recent surveillance studies of mosquito species in Tabasco, Ur. sapphirina and Ur. socialis were omitted due to the lack of recent collection records, but in historical records, the presence of Ur. sapphirina and one species consistent with the description of Ur. socialis were mentioned. During a mosquito survey collection, immature stages from ground-level natural habitats in conservation areas of Tabasco, Ur. sapphirina and Ur. socialis were collected in association with Anopheles albimanus, Culex erraticus, Mansonia titillans, and Ur. lowii. Additionally, 2 Mexican entomological collections were reviewed, searching additional records of those species. An identification key to separate larvae and adult females of Ur. sapphirina and Ur. socialis is provided. With the addition of Ur. sapphirina and Ur. socialis to the mosquito fauna of Tabasco, there are currently 107 species in the state, being the 3rd state in Mexico with the highest richness of mosquito species. Specimens collected during this study were deposited in the Collection of the Entomological and Bioassay Research Unit of Tabasco.

Culex (Culex) vansomereni Edwards subspecies draconis Ingram & de Meillon, 1927 —original combination: Culex draconis (subspecific status by Edwards 1941). Distribution: Republic of South Africa (Ingram & de Meillon 1927). subspecies elgonicus Edwards, 1941 —original combination: Culex vansomereni ssp. elgonicus. Distribution: Ethiopia, Uganda (Wilkerson et al. 2021). subspecies vansomereni Edwards, 1926a —original combination: Culex vansomereni. Distribution: Democratic Republic of the Congo, Ethiopia, Kenya, Liberia, Mozambique, Republic of South Africa, South Sudan [but not Sudan (Simsaa et al. 2021)], Tanzania, Uganda, Zimbabwe (Wilkerson et al. 2021). The record of Lewis (1956) is from Gilo in South Sudan. Subspecies draconis was originally described as a distinct species (Ingram & de Meillon 1927) and interpreted as a subspecies of vansomereni by Edwards (1941). This subspecies differs from the type form in having a pair of submedian yellow stripes on the posterior half of the scutum and sometimes an ill-defined pair of curved lines of yellow scales on the anterior half. In males, the lateral plates of the phallosome are of a slightly different shape with more numerous denticles, and more importantly, the subapical lobe of the gonocoxite is not divided, seta g is larger, as long as seta f, and seta h is noticeably flattened. Based on information provided by Jupp (1996), it seems likely that the two forms occurs in sympatry in South Africa. Subspecies elgonicus was described by Edwards (1941) from a female and two males (with dissected genitalia) collected by G. R. L. Hancock (Mattingly 1956) at high elevation (6,500 ft., about 1,980 m) on Mt Elgon in Uganda. Edwards indicated that elgonicus resembles the typical form “in nearly all respects” and distinguished it based on differences observed in the male genitalia, including the phallosome with slightly different lateral plates (with a longer and more incurved ventrolateral process and a larger number of marginal denticles), subapical lobe less distinctly divided, seta d present, setae d and e small and slender, seta f broader distally and noticeably separated from setae d and e and seta g larger, broader and as long as seta f. Catalogers, beginning with Stone et al. (1959), indicate that elgonicus has been recorded from Ethiopia without providing the source of the record. If elgonicus does in fact occur in Ethiopia, then it is more widely distributed and obviously distinguishable from the type form. Available evidence suggests that Cx. vansomereni is a complex of species. Hopkins (1952) observed that “The larvae of this species vary very considerably, particularly in the degree of sclerotization of the head and siphon, the siphonal index, and the number of comb-scales. Examination of a long series of specimens from Nairobi (the type-locality of the species), South Africa (ssp. draconis Ingram and de Meillon), and various localities in Uganda (ssp. vansomereni) has shown that though the majority of the specimens from South Africa and of those from high elevations in Uganda (6000–7000 ft.) are of the form with strongly-sclerotized and long siphon, and of those from Nairobi and from low elevations (5000 ft. and below) in Uganda of the form with short, weakly-sclerotized siphon, the differences are not constant; in at least one of the Uganda localities specimens of both forms and also intermediates have been found breeding in the same pool. According to Edwards (1941), larvae of vansomereni draconis are separable by their longer siphon (index about 6), the fact that the “coronet” of the subapical spines on the siphon is divided into dorsal and ventral groups, and by the subventral tufts of the siphon being double and scarcely longer than the diameter of the siphon.” This statement, however, pertains only to larvae, which have not been studied in detail. When other larval features, e.g. branching of the dorsal head setae and anal papillae, topographic and distributional data, with indication of sympatry, and particularly differences in structures of the male genitalia are considered, we conclude that the three described forms are likely to be different species; thus, subspecies draconis and elgonicus are herewith elevated to specific status: Culex (Culex) draconis Ingram & de Meillon, 1927 and Culex (Culex) elgonicus Edwards, 1926a . Both nominal forms are currently listed as species in the Encyclopedia of Life., Published as part of Harbach, Ralph E. & Wilkerson, Richard C., 2023, The insupportable validity of mosquito subspecies (Diptera: Culicidae) and their exclusion from culicid classification, pp. 1-184 in Zootaxa 5303 (1) on page 93, DOI: 10.11646/zootaxa.5303.1.1, http://zenodo.org/record/8043342, {"references":["Ingram, A. & de Meillon, B. (1927) A mosquito survey of certain parts of South Africa, with special reference to the carriers of malaria and their control. (Part I). Publications of the South African Institute for Medical Research, 4 (4), 1 - 81, 15 pls.","Edwards, F. W. (1941) Mosquitoes of the Ethiopian Region III. - Culicine adults and pupae. Printed by order of the Trustees, British Museum (Natural History), London, viii + 499 pp.","Wilkerson, R. C., Linton, Y. - M. & Strickman, D. (2021) Mosquitoes of the world. Vols. 1 & 2. Johns Hopkins University Press, Baltimore, Maryland, 1332 pp. https: // doi. org / 10.1186 / s 13071 - 021 - 04848 - 6","Edwards, F. W. (1926 a) Mosquito notes. - VI. Bulletin of Entomological Research, 17 (2), 101 - 131. https: // doi. org / 10.1017 / S 0007485300019143","Simsaa, M. A. A., Harbach, R. E., Almalik, A. M. A., Ahmed, E. M., Esia, A. A., Mohamed, A. H. & Azrag, R. S. (2021) Culex mosquitoes (Diptera: Culicidae) recorded along the Nile River in northern Sudan, with a key for the identification of all species of the genus known to occur in the country. Zootaxa, 4963 (3), 401 - 411. https: // doi. org / 10.11646 / zootaxa. 4963.3.1","Lewis, D. J. (1956) The Culex mosquitoes of the Sudan. Bulletin of Entomological Research, 47 (4), 703 - 721. https: // doi. org / 10.1017 / S 0007485300046940","Jupp, P. G. (1996) Mosquitoes of southern Africa: Culicinae and Toxorhynchitinae. Ekogilde Publishers, Hartebeespoort, 156 pp.","Stone, A., Knight, K. L. & Starcke, H. (1959) A synoptic catalog of the mosquitoes of the world (Diptera, Culicidae). The Thomas Say Foundation. Vol. VI. Entomological Society of America, College Park, Maryland, vi + 358 pp.","Hopkins, G. H. E. (1952) Mosquitoes of the Ethiopian Region I. - Larval bionomics of mosquitoes and taxonomy of culicine larvae. 2 nd Edition. British Museum (Natural History), London, viii + 355 pp."]}

Culex (Microculex) inimitabilis Dyar & Knab subspecies fuscatus Lane & Whitman, 1951 —original combination: Culex (Microculex) inimitabilis fuscatus. Distribution: Brazil, State of Rio de Janeiro (Lane & Whitman 1951; Lane 1953). subspecies inimitabilis Dyar & Knab, 1906b —original combination: Culex inimitabilis. Distribution: Brazil [(Middle Coastal States, São Paulo, Rio de Janeiro, Pará (Lane 1953)], Colombia, Ecuador, French Guiana, Guyana, Lesser Antilles (includes Grenada, Trinidad and Tobago), Panama, Suriname, Venezuela (Wilkerson et al. 2021). The subgenus Microculex has not been dealt with since Lane & Whitman (1951) and Lane (1953), and the species are generally very poorly known. Culex inimitabilis was described from larvae, apparently lost (Stone & Knight 1957a), that were collected in Trinidad. Howard et al. (1915) reproduced the very brief original description of the larva, provided a more detailed description of the larva and described the adult male and female, which established the currently accepted morphological concept of the species. They did not describe the pupa. Dyar (1928) provided a less detailed description of the female, male and larva, but he also did not describe the pupa Lane & Whitman (1951) established fuscatus as a subspecies of inimitabilis based on specimens reared from larvae collected in Rio de Janeiro, Brazil. The authors, and also Lane (1953), provided brief descriptions of the female, male, pupa and larva of both forms and distinguished fuscatus based on the following differences: Abdominal terga of the adult female without basolateral white spots (present in the type form), siphon 10–12 times longer than the basal width (8–10 times longer in the type form), abdominal terga II–IV of the pupa with somewhat triangular area of dark pigmentation, the broad areas on terga each with a pair of unpigmented spots (dark area of pigmentation on terga III and IV and a pair of unpigmented spots on III in the type form), pupal seta 5-II nearly twice as long as the tergum (slightly shorter than the tergum in the type form). It is interesting that Howard et al. (1915) described the siphon of the larva of inimitabilis as being about 14 times longer than the basal width, and yet, in agreement with Lane & Whitman, Dyar (1928) indicated that the siphon of the type form is 10 times longer than the basal width, which brings into question the value of the siphon index for distinguishing the two forms. That aside, the differences exhibited by adult females and pupae, coupled with the recorded occurrence of both forms in Rio de Janeiro State, suggests that fuscatus and inimitabilis are separate species that exist in sympatry. Therefore, until additional observations may prove otherwise, we believe it is prudent to recognize fuscatus as a separate species: Culex (Microculex) fuscatus Lane & Whitman, 1951 . Culex fuscatus is currently listed as a species in the Encyclopedia of Life., Published as part of Harbach, Ralph E. & Wilkerson, Richard C., 2023, The insupportable validity of mosquito subspecies (Diptera: Culicidae) and their exclusion from culicid classification, pp. 1-184 in Zootaxa 5303 (1) on page 102, DOI: 10.11646/zootaxa.5303.1.1, http://zenodo.org/record/8043342, {"references":["Lane, J. & Whitman, L. (1951) The subgenus \" Microculex \" in Brazil (Diptera, Culicidae). Revista Brasileira de Biologia, 11 (3), 341 - 366.","Lane, J. (1953) Neotropical Culicidae. Vols. I & II. University of Sao Paulo, Sao Paulo, 1112 pp.","Dyar, H. G. & Knab, F. (1906 b) The larvae of Culicidae classified as independent organisms. Journal of the New York Entomological Society, 14 (4), 169 - 230, 13 pls.","Wilkerson, R. C., Linton, Y. - M. & Strickman, D. (2021) Mosquitoes of the world. Vols. 1 & 2. Johns Hopkins University Press, Baltimore, Maryland, 1332 pp. https: // doi. org / 10.1186 / s 13071 - 021 - 04848 - 6","Stone, A. & Knight, K. L. (1957 a) Type specimens of mosquitoes in the United States National Museum: IV, The genus Culex (Diptera, Culicidae). Journal of the Washington Academy of Sciences, 47 (2), 42 - 59.","Howard, L. O., Dyar, H. G. & Knab, F. (1915) The mosquitoes of North and Central America and the West Indies. Vol. 3. Systematic description (in two parts). Part I. Publication No. 159. Carnegie Institution of Washington, Washington, D. C., vi + 523 pp.","Dyar, H. G. (1928) The mosquitoes of the Americas. Publication No. 387. Carnegie Institution of Washington, Washington, D. C., 616 pp."]}

Culex (Oculeomyia) aurantapex Edwards subspecies aurantapex Edwards, 1914 —original combination: Culex aurantapex. Distribution: Democratic Republic of the Congo, Kenya, Madagascar, Mozambique, South Africa, Zambia (Wilkerson et al. 2021), also Zimbabwe (Jupp 1996). subspecies ellinorae Ovazza, Hamon & Neri, 1956 —original combination: Culex (Culex) aurantapex var. ellinorae (although originally designated a variety, ellinorae is a replacement name for a recognized subspecies (abyssinicus van Someren, 1945) and has been recognized as a subspecies of aurantapex since Stone et al. 1959). Distribution: Ethiopia (Ovazza et al. 1956). subspecies jinjaensis Edwards, 1941 —original combination: Culex (Culex) aurantapex var. jinjaensis (subspecific status by Harbach & Howard 2007). Distribution: Uganda, Zambia (Wilkerson et al. 2021). Culex aurantapex was described from a single female collected in Nairobi, Kenya (Edwards 1914), a black species with distinctive abdominal scaling—“segments 2–4 black-scaled (dorsally), with a few scattered orange scales; segments 5–8 almost entirely orange-scaled both above and below [terga and sterna].” Edwards (1941) described jinjaensis (as a variety) from a series of seven males and seven females from Jinja, Uganda and a male from Kampala (Mattingly 1956), characterized as being blacker than the type form with “the abdomen lacking the conspicuous orange tip; tergites 5–8 [terga V–VIII] in both sexes with apical lateral yellow patches, which tend to unite to form rather irregular lateral yellow stripes on distal half of abdomen.” Jinja is located on the shore of Lake Victoria in southern Uganda. Edwards stated that the male genitalia of jinjaensis differ “little if at all from those of the typical form”, the genitalia of which Edwards described from specimens collected in Nairobi. Subspecies ellinorae, described by van Someren (1945), under the preoccupied name of abyssinicus, from females collected at Sciasciamanna and Lake Awasa in present-day Ethiopia that differ “from the type form by having more extensive orange markings on the abdominal tergites [terga]”—“2 with narrow black lateral borders and 2 large triangular orange spots the bases of which usually meet on the apical border of the segment to form a narrow orange band; 3 sometimes like 2 and sometimes the same as 4 which is mainly orange with narrow black lateral borders and 2 small median black triangles; 5 orange with narrow black lateral borders; 6–8 all orange. The black markings have a few scattered orange scales. Sternites 6–8 [sterna VI–VIII] all orange; 5 black with a few scattered orange scales and a narrow orange apical band; remainder black with scattered orange scales.” Culex aurantapex is very poorly known morphologically and taxonomically. The larva of the type form and that of subspecies jinjaensis are both only known from a single exuviae from their type localities (Hopkins 1952). Hopkins did not describe or illustrate these larvae because he was unable to find any differences between the exuviae and the larva of Cx. annulioris (see above). The pupa was described by Ingram & de Meillon (1927), which was apparently (not explicitly stated) identified from adults reared from larvae collected from a large swamp east of forest at Empangeni KwaZulu-Natal (as Zululand), South Africa. Edwards (1941) acknowledged the description but pointed out that “confirmation of the identity of this is desirable, but the paedotype pelt [exuviae] from Nairobi is similar in most respects.” Obviously, the immature stages of the nominal forms are essentially unknown. It is unfortunate that the male genitalia of the nominal forms have not been fully described and illustrated, and the immature stages have not been unequivocally associated with the type form. Because the variation observed in adults is suggestive of a species complex, with respect for the intuitive interpretation of morphological observations of the authors of the nominal forms, we are compelled to regard them as separate species pending morphological and molecular confirmation: Culex (Oculeomyia) ellinorae Ovazza, Hamon & Neri, 1956 and Culex (Oculeomyia) jinjaensis Edwards, 1941 . Culex ellinorae and Cx. jinjaensis are both currently listed as species in the Encyclopedia of Life., Published as part of Harbach, Ralph E. & Wilkerson, Richard C., 2023, The insupportable validity of mosquito subspecies (Diptera: Culicidae) and their exclusion from culicid classification, pp. 1-184 in Zootaxa 5303 (1) on page 106, DOI: 10.11646/zootaxa.5303.1.1, http://zenodo.org/record/8043342, {"references":["Edwards, F. W. (1914) New species of Culicidae in the British Museum, with notes on the genitalia of some African Culex. Bulletin of Entomological Research, 5 (1), 63 - 81. https: // doi. org / 10.1017 / S 0007485300034568","Wilkerson, R. C., Linton, Y. - M. & Strickman, D. (2021) Mosquitoes of the world. Vols. 1 & 2. Johns Hopkins University Press, Baltimore, Maryland, 1332 pp. https: // doi. org / 10.1186 / s 13071 - 021 - 04848 - 6","Jupp, P. G. (1996) Mosquitoes of southern Africa: Culicinae and Toxorhynchitinae. Ekogilde Publishers, Hartebeespoort, 156 pp.","Ovazza, M., Hamon, J. & Neri, P. (1956) Contribution a l'etude des dipteres vulnerants de l'empire d'Ethiopie. I. Culicidae. Bulletin de la Societe de Pathologie exotique, 49 (1), 151 - 182.","van Someren, E. C. C. (1945) Ethiopian Culicidae: Descriptions of two East African culicine larvae, not hitherto described, and a new subspecies from Abyssinia (Diptera). Proceedings of the Royal Entomological Society of London, Series B: Taxonomy, 14 (5 - 6), 77 - 80. https: // doi. org / 10.1111 / j. 1365 - 3113.1945. tb 00024. x","Stone, A., Knight, K. L. & Starcke, H. (1959) A synoptic catalog of the mosquitoes of the world (Diptera, Culicidae). The Thomas Say Foundation. Vol. VI. Entomological Society of America, College Park, Maryland, vi + 358 pp.","Edwards, F. W. (1941) Mosquitoes of the Ethiopian Region III. - Culicine adults and pupae. Printed by order of the Trustees, British Museum (Natural History), London, viii + 499 pp.","Harbach, R. E. & Howard, T. M. (2007) Corrections in the status and rank of names used to denote varietal forms of mosquitoes (Diptera: Culicidae). Zootaxa, 1542 (1), 35 - 48. https: // doi. org / 10.11646 / zootaxa. 1542.1.3","Hopkins, G. H. E. (1952) Mosquitoes of the Ethiopian Region I. - Larval bionomics of mosquitoes and taxonomy of culicine larvae. 2 nd Edition. British Museum (Natural History), London, viii + 355 pp.","Ingram, A. & de Meillon, B. (1927) A mosquito survey of certain parts of South Africa, with special reference to the carriers of malaria and their control. (Part I). Publications of the South African Institute for Medical Research, 4 (4), 1 - 81, 15 pls."]}

Culex (Culex) shoae Hamon & Ovazza subspecies shoae Hamon & Ovazza, 1954 —original combination: Culex shoae. Distribution: Ethiopia (Hamon & Ovazza 1954). subspecies ugandae van Someren, 1967 —original combination: Culex (Culex) shoae ugandae. Distribution: Kenya, Uganda (van Someren 1967). The type forms of Cx. shoae and subspecies ugandae are only known from the original descriptions. The type locality of the typical form is an unspecified locality in the Shewa region (Romanized as Shoa) of central Ethiopia. Shewa lies in the Ethiopian Highlands, with elevations mainly above 1,500 m. The type locality of ugandae is Ngogwe, Uganda. Ngogwe is a municipality with an elevation of 1,200 m in the Central Region of the country. Van Someren (1967) indicated that larvae of ugandae were also found at Kakamega (elevation 1,535 m) in western Kenya. The immature stages of both forms have been found in the axils of wild banana plants, but larvae of ugandae have also been found in the axils of Colocasia. Van Someren noted that although subspecies ugandae resembles the type form, all life stages exhibit “small but apparently constant differences”. The larva of ugandae is readily distinguished from the type form by having head seta 4-C with 2–4 branches, comb scales evenly fringed around the apex, the siphonal pecten with 8–10 spines, each with a coarse proximal denticle, and seta 1-X only slightly longer than the saddle. Seta 4-C is unusually large in the type form, with about 13 branches, the comb scales are unique in having lateral spicules that grade distally into blunt denticles and a blunt apex, the pecten consists of about 14 spines, the proximal spines have 1 or 2 denticles and the distal spines are simple, without denticles, and seta 1-X is 3 times as long as the saddle. Differences in the adult and pupa, which van Someren characterized as “only slight and perhaps unreliable”, include the following: In adults of subspecies ugandae the femora have small and inconspicuous knee spots (produced as narrow bands in the type form); in females the dark dorsal scaling of the hindfemur reaches the base of the femur (it does not reach the base in the type form); in males the outer division of the lateral plate of the phallosome has 6 denticles (3 in the type form) and seta d of the subapical lobe of the gonocoxite is stout and bristle-like (fine and hair-like in the type form); in the pupa seta 10-CT has 3 branches (6 branches in the type form), seta 11-CT is single or bifid (with 3 or 4 branches in the type form) and seta 5-VI is usually bifid but sometimes single (bifid in the type form). Differences in the male genitalia which van Someren did not notice include the differently shaped seta g of the subapical lobe (narrowed and slightly pointed distally in ugandae and broadly rounded apically in the type form), the ventrocaudal process of the outer division of the lateral plate is broad and somewhat duck-head shaped (narrow and more tooth-like in the type form) and the dorsal process of the outer division is broadly triangular in lateral view (more or less digiform in the type form). There is a wide gap of much lower terrain between the high elevations of Ethiopia and Uganda / Kenya that include the type localities of the typical form and subspecies ugandae, respectively, and that area of lower elevation could be a barrier to gene flow and explain the allopatric separation and morphological distinctions exhibited by the two forms. Because the morphological differences between the adults, larvae and pupae of typical shoae and subspecies ugandae, coupled with their occurrence in separate geographical areas, provide credible evidence for the recognition of ugandae as a separate species, it is hereby formally raised to specific status: Culex (Culex) ugandae van Someren, 1967 . Culex ugandae is currently listed as a species in the Encyclopedia of Life., Published as part of Harbach, Ralph E. & Wilkerson, Richard C., 2023, The insupportable validity of mosquito subspecies (Diptera: Culicidae) and their exclusion from culicid classification, pp. 1-184 in Zootaxa 5303 (1) on pages 89-90, DOI: 10.11646/zootaxa.5303.1.1, http://zenodo.org/record/8043342, {"references":["Hamon, J. & Ovazza, M. (1954) Une nouvelle espece de moustique (Culex shoae n. sp.) vivant sur les bananiers du haut plateau d'Ethiopie. Bulletin de la Societe de Pathologie exotique, 47 (3), 416 - 421.","van Someren, E. C. C. (1967) The female and early stages of Culex (Culex) nakuruensis Mattingly, with a description of a new subspecies of Culex (Culex) shoae Hamon & Ovazza. Proceedings of the Royal Entomological Society of London, Series B: Taxonomy, 36 (1 - 2), 11 - 16. https: // doi. org / 10.1111 / j. 1365 - 3113.1967. tb 00528. x"]}

Culex (Maillotia) salisburiensis Theobald subspecies capensis de Meillon, 1935 —original combination: Culex (Neoculex) salisburiensis var. capensis (subspecific status by White 1975). Distribution: South Africa (de Meillon 1935). subspecies coursi Doucet, 1949 —original combination: Culex coursi (subspecific status by White 1975). Distribution: Madagascar (Doucet 1949). subspecies salisburiensis Theobald, 1901b 1901c —original combination: Culex salisburiensis. Distribution: Democratic Republic of the Congo, Kenya, Lesotho, Madagascar, Nigeria, Saudi Arabia, South Africa, South Sudan [but not Sudan (Simsaa et al. 2021)], Uganda, Yemen, Zambia, Zimbabwe (Wilkerson et al. 2021). This is another of the many Afrotropical species of Culex that have been poorly collected, little studied and are imperfectly known taxonomically. Many nominal species placed in the genus have been reduced to subspecies or synonyms when they were not studied as part of revisionary works. This applies to White (1975), who recognized variety capensis de Meillon, 1935 and the species coursi Doucet, 1949 as subspecies of salisburiensis. The concept of capensis is clouded by the loss of the type material, the original description based only on the adult female and its synonymy with salisburiensis by Edwards (1941). The issue is further complicated by the description of Cx. salisburiensis naudeanus Muspratt, 1961 from Cape Province of South Africa, which includes the type locality of capensis. As surmized by White (1975), Muspratt “dismissed the availability of the name capensis because of the lack of type-material. He might also have wanted to allow for it being a species distinct from salisburiensis, since its identity cannot be completely resolved until further topotypic material is obtained. However, the published description of capensis quite supports its inclusion as a form [subspecies] of salisburiensis.” But how the description supports Mattingly’s conclusion is questionable as it only states that capensis is similar to salisburiensis but is darker and the proboscis is slightly longer than the forefemur. It is interesting to note that de Meillon (1935) questioned the recognition of capensis as a mere variety of salisburiensis: “Unfortunately, we did not collect any males, so for the present we have given this insect varietal rank only.” Unlike capensis, Muspratt described naudeanus from a series of males and females with associated larval and pupal exuviae and fourth-instar larvae, and found that “This form appears to differ chiefly from C. salisburiensis *) in the larval comb and pecten spines… which are quite distinct.”—the asterisk and parenthesis refer to a footnote that says “The variety capensis De Meill. could not be compared owing to lack of type specimens.” As for the male genitalia, Muspratt stated that “no differences [were] found from [the] salisburiensis typical form.” Knight (1953) identified a male and two females with associated larval and pupal exuviae collected in Yemen as specimens of Cx. salisburiensis, but questioned the identification because of many differences noted between the Yemen adults and the description of salisburiensis provided by Edwards (1941). To confirm the identification, Knight sent a full description of the specimens and a drawing of the male genitalia (fig. 2 in Knight 1953) to Peter Mattingly for comparison with specimens of salisburiensis deposited in the museum in London known at the time as the British Museum (Natural History). Comparisons were made with specimens from [South] Sudan, Kenya, Zambia (as N. Rhodesia), Zimbabwe (as S. Rhodesia) and many locations in South Africa, including Cape Province. Mattingly responded with the following assessment. I am afraid almost all of the differences you noted are due to errors in Edwards’ description. The only difference of any significance is that, while the Nairobi males have terminalia [genitalia] identical with the Yemen form, those from further south lack the longest of three accessory bristles [seta f] on the subapical lobe and one of the two small setae on the dististyle [gonostylus]. It is clear therefore that we have a northern and a southern form but it is impossible to say which is the type form since I have no males from Salisbury [Harare, Zimbabwe]…. It may be possible eventually to distinguish two subspecies. The southern forms are certainly much darker than yours, especially with respect to the scutal scaling but the Nairobi specimens are intermediate and, I should think, would probably intergrade in Eritrea. Also the two specimens from Chilanga [Zambia] suggest that there is considerable seasonal variation since one, collected in January, is very dark, while the other collected in November, is as pale as the Nairobi form. Edwards’ figure for the length of the female palps [maxillary palpi] seems to have been based on a single aberrant or shrunken specimen; and the tergal bands are variable, even in the same locality. The other differences are just errors in description. As confirmation of the errors in Edwards’s description, we note that seta g of the subapical lobe of the gonocoxite illustrated by Knight as having “a distinctive angular and serrate margin and with a distinctive median pigmented area” bears no resemblance to the simple leaf illustrated by Edwards (fig. 83a). Although White (1975) synonymized naudeanus with capensis, this was apparently unknown to or ignored by Jupp (1996), who treated naudeanus as a subspecies of salisburiensis. He distinguished the larva of naudeanus from the larva of the typical form in a key and an illustration of the terminal abdominal segments, and illustrated the gonocoxite and phallosome of the male genitalia. His study was based on specimens collected in southern Africa that were housed in the former National Institute for Virology and the South African Institute for Medical Research, which were united in the year 2000 to form the South Africa National Institute for Communicable Diseases. The two forms in southern Africa are clearly easily distinguished by the form of the comb scales and pecten spines, as originally noted by Muspratt (1961). Subspecies naudeanus differs from the typical form in having mostly spine-like comb scales with saw tooth-like lateral margins and simple curved darkly pigmented pecten spines. In the typical form, the comb scales are evenly fringed and the pecten spines have a row of proximal denticles. It should also be noted that head seta 5-C is about as long and thick as seta 6-C in larvae of naudeanus, as opposed to being about half as long and thinner in the type form (Muspratt 1961). A comparison of the drawings of the gonocoxite provided by Knight (1953) and Jupp (1996) indicates that setal group d–f of naudeanus (= capensis) consists of fewer setae and seta g has a more globose head and a narrower stem. One can only imagine how many other differences would be found to distinguish capensis from the typical form if the adults, larvae and pupae were subjected to a comprehensive comparative anatomical study. Based on the distinctive differences in structures of the larvae and male genitalia, and also because capensis and the type form occur in the same geographical areas, we believe that further study will reveal that the former is a genetically separate species; thus, we believe that capensis should be formally afforded species status: Culex (Maillotia) capensis de Meillon, 1935 . Culex capensis needs to be added to the list of Culex species recognized in the Encyclopedia of Life. Subspecies coursi was described as a full species based on larvae collected at the edge of a rice field on the road between Bejofo and Amparafaravola in the Alaotra-Mangoro Region of Madagascar (Doucet 1949). The adults are unknown. White (1975) dismissed the specific status of coursi based on his perception that “the published description differs insufficiently from that of [the] salisburiensis type-form to warrant maintaining coursi above subspecific rank.” We have carefully examined the description (in French, translated into English by Hopkins 1952) and illustrations of coursi provided by Doucet and it seems obvious that White completely misjudged the degree of morphological distinction from salisburiensis. The larva of coursi bears the following exceptional differences from the larva of salisburiensis: Dorsomentum differently shaped, resembling an equilateral triangle with six acute lateral teeth (more convex with four or five larger blunter lateral teeth in salisburiensis); comb comprised of about 35 rather broad scales with a fringe of rather coarse spicules (about 40–50 narrower scales with an even fringe of slender spicules in salisburiensis); siphon relatively short, broad at the base and tapered to the apex, index about 5 (siphon longer, more or less cylindrical, index about 7 in salisburiensis); three pairs of seta 1-S, each seta with 2–4 branches and distinctly shorter than the diameter of the siphon (four to six pairs of seta 1-S, each seta with 5–8 branches and distinctly longer that the diameter of the siphon in salisburiensis); siphon with two anterolateral setae, each with 2 or 3 branches (siphon with three anterolateral setae, each with 1 or 2 branches in salisburiensis); anal papillae about as long as the saddle, dorsal pair about the same length as the ventral pair (anal papillae shorter than the saddle, dorsal pair distinctly shorter than the ventral pair in salisburiensis). Based on these striking differences, we agree with Doucet (1949) and Hopkins (1952) that coursi is unlikely to be conspecific with salisburiensis, which it does not resemble as closely as was perceived by White (1975). Additionally, coursi is apparently sympatric with salisburiensis in the Lake Alaotra region on the eastern slopes of the central highlands of Madagascar (Doucet 1949; Tantely et al. 2016). Obviously, it is necessary for coursi to be reinstated to its original status as a distinct species: Culex (Maillotia) coursi Doucet (1949) . Culex coursi is currently listed as a species in the Encyclopedia of Life. Two nominal species are recognized as junior synonyms of Cx. salisburiensis: Cx. bostocki Theobald, 1905c (type locality: Transvaal Province, South Africa) and Cx. salisburiensis var. amboannulatus Theobald, 1913a (type locality: Hout Bay, Pinelands and Palmiet River, Cape Province, South Africa). These nominal forms are based on descriptions of adult specimens—the type specimen of bostocki is a female and the type of amboannulatus is a male (genitalia not described or illustrated). In the absence of information about the larvae and male genitalia, it is not possible to know whether the two nominal forms might be conspecific with Cx. capensis rather than Cx. salisburiensis; consequently, for the time being they should remain as synonyms of the latter species., Published as part of Harbach, Ralph E. & Wilkerson, Richard C., 2023, The insupportable validity of mosquito subspecies (Diptera: Culicidae) and their exclusion from culicid classification, pp. 1-184 in Zootaxa 5303 (1) on pages 100-101, DOI: 10.11646/zootaxa.5303.1.1, http://zenodo.org/record/8043342, {"references":["White, G. B. (1975) Notes on a catalogue of Culicidae of the Ethiopian Region. Mosquito Systematics, 7 (4), 303 - 344.","Doucet, J. (1949) Etude des Culicidae (Diptera) du lac Alaotra. Memoires de l'Institut Scientifique de Madagascar, Serie A, 3 (2), 121 - 145.","Theobald, F. V. (1901 b) The classification of mosquitoes. Journal of Tropical Medicine, 4, 229 - 235.","Theobald, F. V. (1901 c) A monograph of the Culicidae or mosquitoes. Vol. 2. British Museum (Natural History), London, viii + 391 pp. https: // doi. org / 10.5962 / bhl. title. 58067","Simsaa, M. A. A., Harbach, R. E., Almalik, A. M. A., Ahmed, E. M., Esia, A. A., Mohamed, A. H. & Azrag, R. S. (2021) Culex mosquitoes (Diptera: Culicidae) recorded along the Nile River in northern Sudan, with a key for the identification of all species of the genus known to occur in the country. Zootaxa, 4963 (3), 401 - 411. https: // doi. org / 10.11646 / zootaxa. 4963.3.1","Wilkerson, R. C., Linton, Y. - M. & Strickman, D. (2021) Mosquitoes of the world. Vols. 1 & 2. Johns Hopkins University Press, Baltimore, Maryland, 1332 pp. https: // doi. org / 10.1186 / s 13071 - 021 - 04848 - 6","Edwards, F. W. (1941) Mosquitoes of the Ethiopian Region III. - Culicine adults and pupae. Printed by order of the Trustees, British Museum (Natural History), London, viii + 499 pp.","Jupp, P. G. (1996) Mosquitoes of southern Africa: Culicinae and Toxorhynchitinae. Ekogilde Publishers, Hartebeespoort, 156 pp.","Hopkins, G. H. E. (1952) Mosquitoes of the Ethiopian Region I. - Larval bionomics of mosquitoes and taxonomy of culicine larvae. 2 nd Edition. British Museum (Natural History), London, viii + 355 pp.","Tantely, M. L., Le Goff, G., Boyer, S. & Fontenille, D. (2016) An updated checklist of mosquito species (Diptera: Culicidae) from Madagascar. Parasite, 23, 20. https: // doi. org / 10.1051 / parasite / 2016018","Theobald, F. V. (1905 c) New Culicidae from India, Africa, British Guiana, and Australia. Journal of Economic Biology, 1, 17 - 36, 2 pls.","Theobald, F. V. (1913 a) Second report on the mosquitoes of the Transvaal. In: Union of South Africa, Department of Agriculture, Second Report of the Director of Veterinary Research. Cape Times Limited, Government Printers, Cape Town, pp. 315 - 342, 2 pls."]}

Culex (Culex) annulirostris Skuse subspecies annulirostris Skuse, 1889 —original combination: Culex annulirostris. Distribution: Australia, Bismarck Archipelago, Cook Island, Easter Island, Fiji, French Polynesia, Guam, Indonesia, Kiribati, Nauru, New Caledonia, Palau, Papua New Guinea, Philippines, Samoa, Solomon Islands, Tahiti, Timor, Tonga, Tuvalu, Vanuatu, Wake Island (Wilkerson et al. 2021, except Mariana Islands). subspecies marianae Bohart & Ingram, 1946b —original combination: Culex (Culex) annulirostris marianae. Distribution: Mariana Islands (Bohart & Ingram 1946b). The nominotypical subspecies is widely distributed in the Australasian Region and occurs in eastern islands of Indonesia (Maluku Islands and Timor) and northward into the Philippines. As indicated by Bohart (1957), “The typical subspecies is replaced in the Mariana Islands by subspecies marianae ”, which he recorded from Aguiguan, Anatahan, Pagan, Saipan and Tinian Islands in the contemporary Northern Mariana Islands and from Guam, the southernmost of the Mariana Islands (Bohart & Ingram 1946b; Yamaguti & LaCasse 1950; Bohart 1957). The adults of subspecies marianae differ from those of the typical subspecies mainly in having a narrow line of pale scales on the caudal margins of abdominal terga II–IV and sometimes also on terga V–VII or VIII. Larvae usually have blunter dorsomental teeth and shorter anal papillae, normally shorter than the saddle (Yamaguti & LaCasse 1950; Bohart 1957). Based on these seemingly consistent differences and the great distances between the Mariana Islands and islands that harbour populations of the typical form, and because evidence suggests that Cx. annulirostris is a complex of species (Lee et al. 1989a), it seems likely that marianae is genetically distinct and should therefore be afforded specific status: Culex (Culex) marianae Bohart & Ingram, 1946b . Culex marianae is currently listed as a species in the Encyclopedia of Life. Five nominal species are recognized as junior synonyms of Cx. annulirostris: Cx. jepsoni Bahr, 1912 (type locality: Fiji Islands), Cx. somerseti Taylor, 1912 (type locality: Somerset, Queensland, Australia), Culicelsa consimilis Taylor, 1913 (type locality: Ayr, Queensland, Australia), Culicelsa simplex Taylor, 1914 (type locality: Townsville, Queensland, Australia) and Cx. palmi Baisas, 1938 (type locality: Parañaque, now officially Pasay, Rizal Province, Luzon, Philippines). The type locality of Cx. annulirostris is located in the Blue Mountains of New South Wales, Australia. As it is probable that Cx. annulirostris is a complex of species (Lee et al. 1989a), it is possible that the five nominal forms may represent at least three separate species: One in the Fiji Islands (Cx. jepsoni), one in the Philippines (Cx. palmi) and a third species in Queensland (Cx. somerseti, Culicelsa consimilis and Culicelsa simplex). For the time being, however, these nominal forms are retained as synonyms of Cx. annulirostris pending further consideration of their taxonomic status., Published as part of Harbach, Ralph E. & Wilkerson, Richard C., 2023, The insupportable validity of mosquito subspecies (Diptera: Culicidae) and their exclusion from culicid classification, pp. 1-184 in Zootaxa 5303 (1) on page 83, DOI: 10.11646/zootaxa.5303.1.1, http://zenodo.org/record/8043342, {"references":["Skuse, F. A. A. (1889) Diptera of Australia. Part V. The Culicidae. Proceedings of the Linnean Society of New South Wales, 3 (4), 1717 - 1764, 1 pl.","Wilkerson, R. C., Linton, Y. - M. & Strickman, D. (2021) Mosquitoes of the world. Vols. 1 & 2. Johns Hopkins University Press, Baltimore, Maryland, 1332 pp. https: // doi. org / 10.1186 / s 13071 - 021 - 04848 - 6","Bohart, R. M. & Ingram, R. L. (1946 b) Mosquitoes of Okinawa and islands in the central Pacific. Navmed 1055. Bureau of Medicine and Surgery, Navy Department, Washington, D. C., 110 pp.","Bohart, R. M. (1957) Insects of Micronesia, Diptera: Culicidae. Insects of Micronesia, 12 (1), 1 - 85. [for 1956]","Yamaguti, S. & LaCasse, W. J. (1950) Mosquito fauna of Guam. Office of the Surgeon, Headquarters, 8 th Army, APO 343. United States. 207 th Malaria Survey Detachment. Office of the Surgeon, Washington, D. C., 101 pp.","Lee, D. J., Hicks, M. M., Debenham, M. L., Griffiths, M., Marks, E. N., Bryan, J. H. & Russell, R. C. (1989 a) The Culicidae of the Australasian Region. Vol. 7. Nomenclature, synonymy, literature, distribution, biology and relation to disease. Genus Culex. Subgenera Acallyntrum, Culex. Monograph Series, Entomology Monograph No. 2. Australian Government Publishing Service, Canberra, ix + 281 pp.","Bahr, P. H. (1912) Filariasis and elephantiasis in Fiji: being a report to the London School of Tropical Medicine. Dysentry [sic] in Fiji during the year 1910. [Series]. Journal of the London School of Tropical Medicine, Supplement 1, viii + 1 - 192, 5 pls., map.","Taylor, F. H. (1912) Contribution to a knowledge of the Culicidae and Ixodidea of North Queensland and the Torres Strait. Annual Report of Community Public Health Queensland, 1912, 27 - 29, 1 pl.","Taylor, F. H. (1913) Report of the entomologist. Report of the Australian Institute of Tropical Medicine, 1911, 49 - 74, 3 pls.","Taylor, F. H. (1914) The Culicidae of Australia. - I. Transactions of the Entomological Society of London, 1913, 683 - 708, 4 pls. https: // doi. org / 10.1111 / j. 1365 - 2311.1914. tb 01794. x","Baisas, F. E. (1938) Notes on Philippine mosquitoes, VII. A - Culex (Culex) with banded proboscis and tarsi. B - Anopheles: the pupae of three rare species; the leucosphyrus - subgroup. Monthly Bulletin of the Bureau of Health, Manila, 18 (5), 175 - 232, 18 pls."]}

Culex (Maillotia) hortensis Ficalbi subspecies hortensis Ficalbi, 1889 —original combination: Culex hortensis. Distribution: Afghanistan, Albania, Algeria, Andorra, Armenia, Austria, Azerbaijan, Belgium, Bosnia and Herzegovina, Bulgaria, Crimean Peninsula, Cyprus, Czech Republic, France, FYRO Macedonia, Georgia, Germany, Greece, Hungary, India, Iran, Iraq, Israel, Italy, Kosovo, Lebanon, Liechtenstein, Malta, Morocco, Poland, Portugal (includes Madeira), Romania, Russia, Serbia, Slovakia, Slovenia, Spain, Switzerland, Tajikistan, Tunisia, Turkey, Ukraine (Wilkerson et al. 2021). subspecies maderensis Mattingly, 1955—original combination: Culex (Neoculex) hortensis ssp. maderensis. Distribution: Madeira Islands (Portugal) (Mattingly 1955a; Ribeiro et al. 1988). Culex hortensis, originally described and named from specimens collected at locations in Tuscany, Italy (Ficalbi 1889), is widely distributed in western areas of the Palaearctic Region. Mattingly (1955) described and named subspecies maderensis for a population on Madeira Island of Portugal for which the adults have “scarcely any apical banding of the abdominal segments” (Christophers 1929). Mattingly stated “In my opinion, its color very clearly different from that of all the other forms of C. hortensis that I have been able to examine, as well as the clear presumption that it is characteristic for a determined geographical area, give me the right to re-describe it as a separate subspecies” (translated from the French). Christophers and Mattingly both noted that the genitalia of the two forms are identical. Schaffner et al. (2001) suspected that maderensis also occurs in the Canary Islands, but the presence of hortensis in the Canaries was confirmed by F. M. Edwards, as indicated in a footnote in Christophers (1929): “Specimens from the Canaries have been very kindly examined by Mr. Edwards who very kindly informs me that the hypopygeal [genitalic] characters are, as I suspected, identical.” More recently, Rogozi et al. (2012) identified maderensis in Albania, presumably based on reduced apical pale banding of the abdominal terga; thus, indicating sympatric variation with regard to this anatomical feature. It seems likely that Cx. hortensis was transported to Madeira Island in historical times and has not evolved into a separate species, and is, as suggested by Christophers (1929), nothing more than a “locally less banded variety”. Henceforth, unless molecular data show otherwise, which seems unlikely, the nominal subspecies maderensis must be formally recognized as a synonymous name: maderensis Mattingly, 1955, junior subjective synonym of Culex (Maillotia) hortensis Ficalbi, 1889 . The nominal maderensis, which is listed as a species in the Encyclopedia of Life, must be removed from the list of valid species of Culex. With the synonymy of maderensis, Cx. hortensis now includes three junior synonyms.The previously recognized synonyms include Maillotia pilifera Theobald, 1907 (type locality: Bornand, Haute Savoie, France) and Cx. lavieri Larrousse, 1925 (type locality: Algeria). These are undoubtedly conspecific with Cx. hortensis., Published as part of Harbach, Ralph E. & Wilkerson, Richard C., 2023, The insupportable validity of mosquito subspecies (Diptera: Culicidae) and their exclusion from culicid classification, pp. 1-184 in Zootaxa 5303 (1) on page 99, DOI: 10.11646/zootaxa.5303.1.1, http://zenodo.org/record/8043342, {"references":["Ficalbi, E. (1889) Notizie preventive sulle zanzare italiane. Ia Nota preventiva. Alcune generalita. - Descrizione di una specie nuova (Culex hortensis.). Bullettino della Societa Entomologica Italiana, 21, 20 - 30.","Wilkerson, R. C., Linton, Y. - M. & Strickman, D. (2021) Mosquitoes of the world. Vols. 1 & 2. Johns Hopkins University Press, Baltimore, Maryland, 1332 pp. https: // doi. org / 10.1186 / s 13071 - 021 - 04848 - 6","Christophers, S. R. (1929) Note on a collection of anopheline and culicine mosquitoes from Madeira and the Canary Islands. Indian Journal of Medical Research, 17 (2), 518 - 530, 2 pls.","Schaffner, F., Angel, G., Geoffroy, B., Hervy, J. P., Rhaim, A., Brunhes, J. (2001) The mosquitoes of Europe. An identification and training programme. IRD-EID Mediterranee, IRD Editions, Paris. [CD-ROM]","Rogozi, E., Velo, E., Bino, S., Severini, F., Schaffner, F. (2012) An update of the mosquito fauna of Albania, based on a country-wide field survey 2011 - 12. Poster ESOVE, Montpellier. Available from: https: // www. researchgate. net / publication / 271485432 _ An _ Update _ of _ the _ Mosquito _ Fauna _ of _ Albania _ based _ on _ a _ Country-Wide _ Field _ Survey _ 2011 - 12 j (accessed 15 May 2023)","Theobald, F. V. (1907) A monograph of the Culicidae or mosquitoes. Vol. 4. British Museum (Natural History), London, xix + 639 pp., 16 pls. https: // doi. org / 10.5962 / bhl. title. 58067","Larrousse, F. (1925) Larve de Culex a branchies tres developpees nouvelle pour la faune Francaise (Culex lavieri n. sp.). Annales de Parasitologie Humaine et Comparee, 3 (1), 68 - 70. https: // doi. org / 10.1051 / parasite / 1925031068"]}

Culex (Culex) striatipes Edwards subspecies joanae Muspratt, 1955 —original combination: Culex (Culex) striatipes ssp. joanae. Distribution: Republic of South Africa (Muspratt 1955). subspecies striatipes Edwards, 1941 —original combination: Culex (Culex) striatipes. Distribution: Burkina Faso, Democratic Republic of the Congo, Ethiopia, Ivory Coast, Kenya, Madagascar, Zambia, Zimbabwe (Wilkerson et al. 2021). Edwards (1941) described and named Cx. striatipes based on three adults collected in Kenya, a male and a female from Nairobi and a male from Mt Elgon. The larva and pupa of the species were described by de Meillon et al. (1945) from exuviae, derived from two larvae collected at Livingstone, Zambia (as Northern Rhodesia), which were apparently reared to adults that were presumably used to identify the species. Hopkins (1952) repeated verbatim the description of the larva published by de Meillon et al. and included illustrations of the head and terminal abdominal segments which were reconstructed from the two larval exuviae used for the original description. The illustrations included details that were not shown in the drawings provided by de Meillon et al. (1945). Muspratt (1955) described subspecies joanae from six males and six females, one with an associated larval exuviae. The adults were apparently reared from larvae collected “near the Lydenburg to Sabie road over Mount Anderson” in the former Transvaal Province of South Africa. Muspratt stated that the subspecies differed from the typical form “in having no anterior white stripe on the middle femur and the middle tibia either all dark or with at most a faint anterior pale stripe. The hind femur is pale anteriorly only to about half instead of the usual 4/5. The latter character is probably the most reliable distinction as on some otherwise typical specimens from southern Rhodesia [now Zimbabwe] one or more of the white lines on the anterior side of the front and middle femora, and on the middle tibia, are faint or absent. The type series of ssp joanae are rather larger and darker than striatipes from southern Rhodesia.” He stated that the larva “appears to be practically as the typical form”. No further morphotaxonomic work has been published on striatipes, but interestingly Jupp (1996), who included the species (without joanae) in a key to the adult mosquitoes of South Africa, observed that subspecies joanae “is indistinguishable from typical striatipes in the larva, and probably also in the ♂ genitalia. Adult ♀ ’s differ in having no anterior white stripe on the midfemur and midtibia, although midtibia sometimes has very faint stripe. Hindfemur is pale anteriorly only to about 0.5 instead of the usual 0.8.” As so few specimens have been historically collected and studied, it is difficult to know the degree of morphological variation that exists in populations of the species between Zimbabwe and South Africa, indeed throughout the range of the species, which is also imperfectly known. For the time being, we consider that there is insufficient evidence to retain joanae as a subspecies and hereby formally consign it to synonymy: joanae Muspratt, 1955, junior subjective synonym of Culex (Culex) striatipes Edwards, 1941 . Subspecies joanae Muspratt, 1955, which is currently listed as a species in the Encyclopedia of Life, should be removed from the list of recognized species of the genus Culex., Published as part of Harbach, Ralph E. & Wilkerson, Richard C., 2023, The insupportable validity of mosquito subspecies (Diptera: Culicidae) and their exclusion from culicid classification, pp. 1-184 in Zootaxa 5303 (1) on page 90, DOI: 10.11646/zootaxa.5303.1.1, http://zenodo.org/record/8043342, {"references":["Muspratt, J. (1955) Research on South African Culicini (Diptera, Culicidae). III. - A check-list of the species and their distribution, with notes on taxonomy, bionomics and identification. Journal of the Entomological Society of Southern Africa, 18 (2), 149 - 207.","Edwards, F. W. (1941) Mosquitoes of the Ethiopian Region III. - Culicine adults and pupae. Printed by order of the Trustees, British Museum (Natural History), London, viii + 499 pp.","Wilkerson, R. C., Linton, Y. - M. & Strickman, D. (2021) Mosquitoes of the world. Vols. 1 & 2. Johns Hopkins University Press, Baltimore, Maryland, 1332 pp. https: // doi. org / 10.1186 / s 13071 - 021 - 04848 - 6","de Meillon, B., Parent, M. & Black, L. O'C. (1945) Descriptions of new larvae and pupae of Ethiopian Culicini. Bulletin of Entomological Research, 36 (1), 85 - 101. https: // doi. org / 10.1017 / S 0007485300023932","Hopkins, G. H. E. (1952) Mosquitoes of the Ethiopian Region I. - Larval bionomics of mosquitoes and taxonomy of culicine larvae. 2 nd Edition. British Museum (Natural History), London, viii + 355 pp.","Jupp, P. G. (1996) Mosquitoes of southern Africa: Culicinae and Toxorhynchitinae. Ekogilde Publishers, Hartebeespoort, 156 pp."]}

Culex (Culiciomyia) nebulosus Theobald subspecies nebulosus Theobald, 1901d —original combination: Culex nebulosus. Distribution: Benin, Burkina Faso, Cameroon, Central African Republic, Comoros, Côte d’Ivoire, Democratic Republic of the Congo, Ethiopia, Gabon, Ghana, Kenya, Liberia, Madagascar, Mali, Niger, Nigeria, Republic of the Congo, Republic of South Africa, Saudi Arabia, Senegal, Sierra Leone, Tanzania, Togo, Uganda, Yemen, Zambia (Wilkerson et al. 2021, listed in error from Malaysia). subspecies pseudocinereus Theobald, 1901c —original combination: Culex pseudocinereus (Stone et al. 1959 and Knight & Stone 1977 erroneously attributed subspecific status to Edwards 1941, who actually treated the taxon as a variety; however, as explained by Harbach 2018, pseudocinereus is deemed to have subspecific rank). Distribution: Botswana, Democratic Republic of the Congo, Lesotho, Malawi, Namibia, Nigeria, Republic of South Africa, Tanzania, Zimbabwe (Wilkerson et al. 2021). Culiciomyia is a very difficult group taxonomically. Females are generally difficult to distinguish, some are indistinguishable, and the identification of species depends mainly on anatomical features of males and fourth-instar larvae. Nineteen species of the subgenus are currently recognized in the Afrotropical Region (Cornel et al. 2020). The adults of both sexes and larvae are known for 12 of the species, albeit inadequately or incompletely described, only males are known for six species and one species is only known from the larval stage. As for most species of the subgenus, the adults and larva of Cx. nebulosus sensu stricto and subspecies pseudocinereus are known but have not been studied and described in detail. Their identification relies on subtle differences in the extent of scaling on areas of the thorax of adults, the development of the maxillary palpal comb scales of males and the form of the pecten spines and spiracular apodeme of larvae. Note. The palpal comb (Cornel et al. 2020) is a ventral row of long outstanding hair-like scales (Belkin 1962) on the distal half of palpomere 3 (Edwards 1941). Cornel et al. (2020) quoted Edwards (1941) for the development of the palpal comb scales of the typical form, but their photographic illustration differs slightly from Edwards’s line drawing. The palpal comb depicted by Edwards consists of a proximal set of 6 long spear-like scales separated by a small gap from a distal set comprised of 7 smaller, distally broader and apically hooked scales. In contrast, the proximal comb set illustrated by Cornel et al. consists of 7 scales of two sizes, a proximal group of 4 distinctly longer scales similar to those illustrated by Edwards and a distal group of 3 shorter, broader, apically pointed scales; a gap separates the proximal set from the distal set, which consists of 8 or 9 scales about the same length as the distal scales of the proximal set but developed as illustrated by Edwards. Variation in the composition of the palpal comb of the type form is further illustrated by Jupp (1996): The proximal comb set consists of 7 scales separated by a gap from the distal set comprised of 8 scales, the scales of both sets are as illustrated by Edwards. In summary, the palpal comb of the type form consists of a proximal set of 6 or 7 lanceolate scales separated by a gap from a distal set of 7–9 distally broadened, apically hooked scales. The palpal comb of subspecies pseudocinereus has never been illustrated; however, Edwards (1941) noted that “Scales in distal part of palpal comb slightly different in shape from typical nebulosus, being bluntly pointed but without strongly hooked tips.” During the course of this study, we had an opportunity to examine the holotype male of pseudocinereus and noted that the comb scales gradually grade from large scales proximally to smaller scales distally with little change in shape and no apparent gap or clear delimitation of proximal and distal sets. In addition to differences in the palpal comb scales, Edwards also noted that pseudocinereus “Closely resembles typical nebulosus but differs in both sexes in having much more numerous flat white scales on the mesepimeron; propleura [proepisternum] with a large patch of scales; lower mesepimeral bristle [seta] often accompanied by from one to three short hairs [small setae] (between it and the scales).” Edwards did not find any differences in the male genitalia of the two forms, but this is likely to be due to inattention to delicate structural detail. Bram (1967) stated that species of Culiciomyia are most reliably distinguished in the larval stage; for this reason, it is surprising that Hopkins (1952) did not treat the larva of pseudocinereus, the omission of which may have lead Mattingly & Lips (1953) to consider that larvae of nebulosus and pseudocinereus were indistinguishable. As is the case with most species of Afrotropical Culex, the larva of nebulosus is superficially and inadequately described. For this reason, it was only possible for Peters (1955) to compare larvae of pseudocinereus with the description of the type form provided by Hopkins (1952). In doing so, Peters was only able to determine that the larva of pseudocinereus differs from the type form in the development of the pecten spines and the spiracular apodeme (“stirrup-shaped piece”). The pecten spines of the former have a broad base and a single denticle as opposed to a narrower base and usually at least two denticles in the type form. More striking is the development of the spiracular apodeme, which is much broader with the arms of the U-shaped distal end more widely separated and the narrower arm longer and differently shaped than in pseudocinereus. As shown for two species of the subgenus Culex in Central America (Strickman 1990), the form of the spiracular apodeme is indicative of different species. Collection records indicate that nebulosus sensu stricto probably occurs throughout sub-Saharan Africa. Discounting records of subspecies pseudocinereus in Nigeria and the Democratic Republic of the Congo, it appears that this form is prevalent in southeastern and southern countries of the continent where it is probably sympatric with the type form. This is supported by the occurrence of the adults of both forms in Elizabethville, Belgian Congo (Peters 1955), now known as Lubumbashi in the Democratic Republic of the Congo, which is located in the southeast of the country adjacent to the border with Zambia. Based on the forgoing analysis of morphological and distributional information, it seems highly likely that nebulosus and pseudocinereus are separate species; hence, the latter form is herewith formally returned to its original specific status: Culex (Culiciomyia) pseudocinereus Theobald, 1901c . Culex pseudocinereus is currently listed as a species in the Encyclopedia of Life. Culex nebulosus has a single synonym, Cx. nigrochaetae Theobald, 1901c (♀ only of type series; type locality: Lagos, Nigeria) and Cx. pseudocinereus has three synonyms, Cx. freetownensis Theobald, 1901c (type locality: Freetown, Sierra Leone), Cx. invenustus Theobald, 1901d (type locality: Degama, Rivers State, Nigeria) and Pectinopalpus fuscus Theobald, 1909 (type locality: Obuasi, Ashanti Region, Ghana). These four nominal forms are based on superficial descriptions of adults: The type specimens of the first three are females and the type of fuscus is a male. Until reared specimens from the type localities of these nominal species are available for critical comparative study, they must remain synonyms of their senior synonyms., Published as part of Harbach, Ralph E. & Wilkerson, Richard C., 2023, The insupportable validity of mosquito subspecies (Diptera: Culicidae) and their exclusion from culicid classification, pp. 1-184 in Zootaxa 5303 (1) on pages 94-95, DOI: 10.11646/zootaxa.5303.1.1, http://zenodo.org/record/8043342, {"references":["Theobald, F. V. (1901 d) Notes on a collection of mosquitoes from West Africa, and descriptions of new species. Memoirs of the Liverpool School of Tropical Medicine, 4 (Appendix), i - xiv, 3 pls.","Wilkerson, R. C., Linton, Y. - M. & Strickman, D. (2021) Mosquitoes of the world. Vols. 1 & 2. Johns Hopkins University Press, Baltimore, Maryland, 1332 pp. https: // doi. org / 10.1186 / s 13071 - 021 - 04848 - 6","Theobald, F. V. (1901 c) A monograph of the Culicidae or mosquitoes. Vol. 2. British Museum (Natural History), London, viii + 391 pp. https: // doi. org / 10.5962 / bhl. title. 58067","Stone, A., Knight, K. L. & Starcke, H. (1959) A synoptic catalog of the mosquitoes of the world (Diptera, Culicidae). The Thomas Say Foundation. Vol. VI. Entomological Society of America, College Park, Maryland, vi + 358 pp.","Knight, K. L. & Stone, A. (1977) A catalog of the mosquitoes of the world (Diptera: Culicidae). 2 nd Edition. The Thomas Say Foundation. Vol. VI. Entomological Society of America, College Park, Maryland, xi + 611 pp.","Edwards, F. W. (1941) Mosquitoes of the Ethiopian Region III. - Culicine adults and pupae. Printed by order of the Trustees, British Museum (Natural History), London, viii + 499 pp.","Harbach, R. E. (2018) Culicipedia: Species-group, genus-group and family-group names in Culicidae (Diptera). CABI, Wallingford, Oxfordshire, xviii + 378 pp. https: // doi. org / 10.1079 / 9781786399052.0000","Cornel, A. J., Mayi, M. P. A., Kowo, C., Foncha, D., Andongma, E., Anong, D. N., Elad, M., Djomo, C., Tchuinkam, T., Brisco, K. K. & Sehgal, R. (2020) New species of Culex (Culiciomyia) (Diptera: Culicidae) from Talangaye Forest in Cameroon and descriptions and identification keys for males of the Afrotropiccal species of the subgenus. Zootaxa, 4858 (4), 451 - 506. https: // doi. org / 10.11646 / zootaxa. 4858.4.1","Belkin, J. N. (1962) The mosquitoes of the South Pacific (Diptera, Culicidae). Vols 1 & 2. University of California Press, Berkeley and Los Angeles, California, xii + 608 pp., 412 figs.","Jupp, P. G. (1996) Mosquitoes of southern Africa: Culicinae and Toxorhynchitinae. Ekogilde Publishers, Hartebeespoort, 156 pp.","Bram, R. A. (1967) Contributions to the mosquito fauna of Southeast Asia. II. The genus Culex in Thailand (Diptera: Culicidae). Contributions of the American Entomological Institute, 2 (1), 1 - 296.","Hopkins, G. H. E. (1952) Mosquitoes of the Ethiopian Region I. - Larval bionomics of mosquitoes and taxonomy of culicine larvae. 2 nd Edition. British Museum (Natural History), London, viii + 355 pp.","Mattingly, P. F. & Lips, M. (1953) Notes on the Culicini of the Katanga (Diptera, Culicidae). Revue de Zoologie et de Botanique Africaines, 47 (3 - 4), 311 - 343.","Peters, W. (1955). The mosquitoes of Liberia (Diptera: Culicidae). Proceedings of the Royal Entomological Society of London Series B Taxonomy, 24 (5 - 6), 81 - 90. https: // doi. org / 10.1111 / j. 1365 - 3113.1955. tb 01487. x","Strickman, D. (1990) Culex pseudostigmatosoma, Cx. yojoae and Cx. aquarius: New Central American species in the subgenus Culex (Diptera: Culicidae). Mosquito Systematics, 21 (3), 143 - 177. [for 1989]","Theobald, F. V. (1909) Descriptions of the new mosquitoes collected by Dr. Graham in Ashanti. Gold Coast Miscellaneous Publication No. 237. Great Britain Colonial Office. London, 31 pp., 2 pls."]}

Culex (Culex) weschei Edwards subspecies gediensis Edwards, 1941 —original combination: Culex (Culex) weschei ssp. gediensis. Distribution: Coastal Kenya (Edwards 1941; van Someren et al. 1955). subspecies weschei Edwards, 1935 —original combination: Culex weschei. Distribution: Burkina Faso, Cameroon, Democratic Republic of the Congo, Ghana, Guinea, Madagascar, Mali, Mozambique, Nigeria, Senegal, Seychelles, South Sudan, Sudan, Togo (Wilkerson et al. 2021), Zimbabwe (Jupp 1996). As is the case of most species of Culex in the Afrotropical Region, knowledge of the morphology and bionomics of Cx. weschei is very scanty. The adult female and male and the fourth-instar larva of the type form are known but they have not been studied in detail. The original description was based on the scaling of the legs and abdomen of adults that distinguished them from the adults of Cx. guiarti Blanchard, 1905. Subspecies gediensis was described as a form that resembles the type form “closely in colouring and in structure of the ♂ terminalia [genitalia], but differs as follows: antenna of ♀ with only 10 instead of 15 hairs [setae] in each whorl; propleura [antepronota] in both sexes with fewer setae (about 10 in ♂ and about 6 in ♀ instead of about 20 and 15 respectively); sternopleura [mesokatepisterna] with fewer scales” (Edwards 1941). The mesokatepisternal scales are numerous in the type form, with the upper and lower patches nearly joined (Edwards 1941). The larva of gediensis is unknown. It is interesting to note that the larva of the type form illustrated as “ Culex sp.?” by Macfie & Ingram (1916) was recognized as the larva of Cx. weschei by Edwards (1941), but it was not described until Hopkins (1952) included it, along with the illustrations of Macfie & Ingram, in his treatise on the larvae of the Mosquitoes of the Ethiopian Region. The type form was originally described from Ghana (as Gold Coast) and has been recorded from a number of countries in sub-Saharan Africa. Subspecies gediensis was originally described (Edwards 1935) and subsequently recorded (van Someren et al. 1955) from the coastal fringe of Kenya. It has not been found elsewhere and the type form has not been recorded from inland areas of Kenya, and except for South Sudan to the northwest, it has not been recorded from the adjoining countries of Ethiopia (north), Somalia (northeast), Tanzania (south) and Uganda (west). In view of its apparent allopatric coastal distribution and the exceptional differences in the vestiture of the antenna, antepronotum and mesokatepisternum compared to the type form, we conclude that gediensis is probably a distinct species and therefore formally afford it specific status: Culex (Culex) gediensis Edwards, 1941 . Culex gediensis is currently listed as a species in the Encyclopedia of Life., Published as part of Harbach, Ralph E. & Wilkerson, Richard C., 2023, The insupportable validity of mosquito subspecies (Diptera: Culicidae) and their exclusion from culicid classification, pp. 1-184 in Zootaxa 5303 (1) on page 94, DOI: 10.11646/zootaxa.5303.1.1, http://zenodo.org/record/8043342, {"references":["Edwards, F. W. (1941) Mosquitoes of the Ethiopian Region III. - Culicine adults and pupae. Printed by order of the Trustees, British Museum (Natural History), London, viii + 499 pp.","van Someren, E. C. C., Teesdale, C. & Furlong, M. (1955) The mosquitoes of the Kenya coast; records of occurrence, behaviour and habitat. Bulletin of Entomological Research, 46 (3), 463 - 493. https: // doi. org / 10.1017 / S 0007485300039481","Edwards, F. W. (1935) Mosquito notes. - XII. Bulletin of Entomological Research, 26 (2), 127 - 136. https: // doi. org / 10.1017 / S 0007485300038116","Wilkerson, R. C., Linton, Y. - M. & Strickman, D. (2021) Mosquitoes of the world. Vols. 1 & 2. Johns Hopkins University Press, Baltimore, Maryland, 1332 pp. https: // doi. org / 10.1186 / s 13071 - 021 - 04848 - 6","Jupp, P. G. (1996) Mosquitoes of southern Africa: Culicinae and Toxorhynchitinae. Ekogilde Publishers, Hartebeespoort, 156 pp.","Blanchard, R. (1905) Les moustiques, histoire naturelle et medicale. F. R. de Rudeval, Imprimeur-Editeur, Paris, xiii + 673 pp. https: // doi. org / 10.5962 / bhl. title. 100899","Macfie, J. W. S. & Ingram, A. (1916) New culicine larvae from the Gold Coast. Bulletin of Entomological Research, 7 (1), 1 - 18. https: // doi. org / 10.1017 / S 000748530001703 X","Hopkins, G. H. E. (1952) Mosquitoes of the Ethiopian Region I. - Larval bionomics of mosquitoes and taxonomy of culicine larvae. 2 nd Edition. British Museum (Natural History), London, viii + 355 pp."]}

Culex (Eumelanomyia) hayashii Yamada subspecies hayashii Yamada, 1917 —original combination: Culex hayashii. Distribution: Japan, People’s Republic of China, Russia, South Korea, Taiwan (Wilkerson et al. 2021). subspecies ryukyuanus Tanaka, Mizusawa & Saugstad, 1979 —original combination: Culex (Eumelanomyia) hayashii ryukyuanus. Distribution: Japan (Ryukyu Archipelago) (Tanaka et al. 1979). Sirivanakarn (1972), in his revisionary study of the subgenus Eumelanomyia in Southeast Asia and adjacent areas, stated that “The adults of C. hayashii show a great deal of variation in size, color, texture of scutal scales and in the length of male palpus. The specimens from the Ryukyu Islands differ from the specimens from Japan and Korea in smaller size, darker coloration, finer scutal scales and shorter male pulpus [sic]. However, these differences are not correlated with any differentiation in the male terminalia [genitalia], indicating that there is in all probability only one species involved.” Tanaka et al. (1979) interpreted ryukyuanus to be a subspecies distinct from the type form based on the following characteristics: Generally smaller body size, wing with a smaller ratio of the length of cell R 2 to the length of vein R 2+3, a slight difference in the ratio of the length of hindtarsomere 1 to the length of the hindtibia (0.98–1.11 as opposed to 0.85–1.03 in the type form), male with shorter maxillary palpus (palpomere 2 shorter than it is in the type form). They stated that the larva of ryukyuanus did not appear to be significantly different from hayashii sensu stricto on Japan’s main island (Honshu). The Ryukyu Archipelago comprises an arc of 55 islands and islets that extend about 1,000 km southwestward from 40 km south of Kyushu, the southernmost of Japan’s main islands, to 100 km east of northern Taiwan. The chain of islands is divided by two large gaps, a northern gap of about 270 km between Yakushima Island and the Amami island group, and a southern gap of about 280 km between the islands of Okinawa and Miyako. The flora and fauna tend to be very different on either side of these gaps, and forms that occur on islands of the Archipelago that have been regarded as subspecies are proving to be genetically distinct (Toma et al. 2019; Somboon et al. 2020a; Wilkerson et al. 2022). For example, Toma et al. (2019) clearly showed that Tripteroides bambusa yaeyamensis Tanaka, Mizusawa & Saugstad, 1979 in the central and southern regions of the archipelago is molecularly and genetically distinct from Tp. bambusa (Yamada, 1917) in the northern Palaearctic region of Japan. Based on the results of their study and similar findings regarding the specific status of Aedes (Hulecoeteomyia) yaeyamensis Tanaka, Mizusawa & Saugstad, 1979 (Somboon et al. 2020a; Wilkerson et al. 2022), despite the morphological similarity observed by Sirivanakarn (1972) and Tanaka et al. (1979), we believe it is likely molecular study will reveal that ryukyuanus and hayashii sensu stricto are separate species. Therefore, unless genetic evidence proves otherwise, we hereby formally elevate ryukyuanus to specific rank: Culex (Eumelanomyia) ryukyuanus Tanaka, Mizusawa & Saugstad, 1979 . Culex ryukyuanus is currently listed as a species in the Encyclopedia of Life. For comparison, see the analogous treatments of the subspecies of Toxorhynchites manicatus (Edwards, 1921a) and Uranotaenia novobscura Barraud, 1934 presented below., Published as part of Harbach, Ralph E. & Wilkerson, Richard C., 2023, The insupportable validity of mosquito subspecies (Diptera: Culicidae) and their exclusion from culicid classification, pp. 1-184 in Zootaxa 5303 (1) on page 96, DOI: 10.11646/zootaxa.5303.1.1, http://zenodo.org/record/8043342, {"references":["Yamada, S. (1917) On two new species of mosquitoes from Japan. Dobutsugaku zasshi [Zoological Magazine], 29 (341), 61 - 72. [in Japanese]","Wilkerson, R. C., Linton, Y. - M. & Strickman, D. (2021) Mosquitoes of the world. Vols. 1 & 2. Johns Hopkins University Press, Baltimore, Maryland, 1332 pp. https: // doi. org / 10.1186 / s 13071 - 021 - 04848 - 6","Tanaka, K., Mizusawa, K. & Saugstad, E. S. (1979) A revision of the adult and larval mosquitoes of Japan (including the Ryukyu Archipelago and the Ogasawara Islands) and Korea (Diptera: Culicidae). Contributions of the American Entomological Institute, 16, vii + 1 - 987.","Sirivanakarn, S. (1972) Contributions to the mosquito fauna of Southeast Asia. XIII. The genus Culex, subgenus Eumelanomyia Theobald in Southeast Asia and adjacent areas. Contributions of the American Entomological Institute, 8 (6), i, 1 - 86.","Toma, T., Taira, K., Oosiro, A., Azama, Y., Fukuchi, Y. & Miyagi, I. (2019) New taxonomic status of Tripteroides bambusa (Yamada), 1917 [sic] from Japan, based on experimental crossing and COI sequence divergence. Medical Entomology and Zoology, 70 (1), 15 - 23. https: // doi. org / 10.7601 / mez. 70.15","Somboon, P., Phanitchakun, T., Namgay, R. & Harbach, R. E. (2020 a) Description of Aedes (Hulecoeteomyia) bhutanensis n. sp. (Diptera: Culicidae) from Bhutan. Acta Tropica, 203, 105280. https: // doi. org / 10.1016 / j. actatropica. 2019.105280","Wilkerson, R. C., Somboon, P. & Harbach, R. E. (2022) Reconsideration of the status of subspecies in the Japonicus Group of the subgenus Hulecoeteomyia Theobald of Aedes Meigen (Diptera: Culicidae). Zootaxa, 5162 (2), 198 - 200. https: // doi. org / 10.11646 / zootaxa. 5162.2.8","Edwards, F. W. (1921 a) H. Sauter's Formosan collections: Culicidae. Annals and Magazine of Natural History, Series 9, 8 (48), 629 - 632. https: // doi. org / 10.1080 / 00222932108632632","Barraud, P. J. (1934) The fauna of British India, including Ceylon and Burma. Diptera. Vol. V. Family Culicidae. Tribes Megarhinini and Culicini. Taylor and Francis, London, xxviii + 463 pp., 8 pls."]}

Culex (Culex) invidiosus Theobald subspecies invidiosus Theobald, 1901d —original combination: Culex invidiosus. Distribution: Sub-Saharan Africa, countries north of approximately 10° S latitude—including Benin, Burkina Faso, Cameroon, Côte d’Ivoire, Democratic Republic of the Congo, Gabon, Gambia, Ghana, Kenya, Liberia, Mali, Nigeria, Republic of the Congo, São Tomé and Príncipe, Senegal, Sierra Leone, South Sudan, Sudan, Tanzania, Uganda (Wilkerson et al. 2021). subspecies vexillatus Edwards, 1941 —original combination: Culex (Culex) invidiosus var. vexillatus (subspecific status by Harbach & Howard 2007). Distribution: Cameroon, Democratic Republic of the Congo, Nigeria, Republic of the Congo, Uganda (Wilkerson et al. 2021). subspecies vicinalis de Meillon & Lavoipierre, 1944 —original combination: Culex (Culex) invidiosus ssp. vicinalis. Distribution: Democratic Republic of the Congo (de Meillon & Lavoipierre 1944). Like many species of Afrotropical Culex, Cx. invidiosus is very poorly known. It was last treated by Edwards (1941), who distinguished vexillatus as a variety based on five males and a female from Kampala, Uganda (Mattingly 1956) that closely resembles the typical form except for the shape of seta f of the subapical lobe of the male gonocoxite. Three years later, de Meillon & Lavoipierre (1944) described vicinalis as a subspecies of Cx. invidiosus based on a single male collected at Yangambi, a town located on the Congo River in the central region of present-day Democratic Republic of the Congo. A review of the taxonomic history of Cx. invidiosus following its original description by Theobald (1901c) begins with Edwards (1911b), who indicated that it belonged to a group of species that are “very difficult to classify” because they lack “clearly marked distinctions.” Ironically, Edwards formally recognized Cx. euclastus Theobald, 1903b, Cx. chloroventer Theobald, 1909 and Cx. aquilus Graham, 1910 as synonyms of Cx. invidiosus, and those names have remained as synonyms of Cx. invidiosus to the present. Wesché (1910) described the larva of Cx. invidiosus (as Cx. aquilus) and included it in a key to the larvae of African Culicidae. Edwards (1912d) updated the key and illustrated the head and terminal abdominal segments of Cx. invidiosus. Although Edwards was “unable to separate the larvae of C. decens [now considered a valid species]… and C. invidiosus ”, and found that the characters given by Wesché were unreliable, he believed the two species were distinct because Cx. decens could be distinguished by the reddish thorax (brown in Cx. invidiosus) and the banded abdominal segments of males. Despite treating them as separate species, he concluded that “It is therefore quite possible that the two are really only forms of one species; they generally occur together, but specimens bred from one batch of larvae exhibit little variation.” Edwards (1914) acknowledged that species of Culex are most readily separated by features of the male genitalia, but indicted “As previously stated, I can detect no difference whatever between the hypopygia [genitalia] of this species [Cx. invidiosus] and of C. decens.” Despite this comment, Edwards treated Cx. invidiosus as a distinct species and surmized that it “is probably a purely West African species.” He illustrated the phallosomes and gonocoxites of Cx. invidiosus, Cx. antennatus (Becker, 1903) (as Cx. laurenti Newstead, 1907, in Newstead et al. 1907) and Cx. perfuscus sp. nov. to show the close similarity of the genitalia of seemingly unrelated species. Macfie & Ingram (1920) conducted a detailed comparison of the pupae of Cx. decens and Cx. invidiosus, but were unable to find differences to distinguish them. In summary, they stated: “The question then arises, are C. decens and C. invidiosus separate species or are they varieties of a single mosquito. There are indeed certain differences in the adults, but the genitalia of the males are identical according to Edwards, the larvae cannot be separated [reiterated by Hopkins 1936, 1952], and the same remark applies to the pupae. Under these circumstances we think there can be little doubt that they should be regarded as varieties and not as distinct species; we propose to retain for the species the name C. decens.” Based on these findings, Edwards (1932a) listed invidiosus as a variety of Cx. decens, with the three synonyms noted above. Edwards (1941) is the last reviser of the subgenus Culex in the Afrotropical Region. He reiterated his earlier observation (Edwards 1912d) that the adults of Cx. invidiosus differ from those of Cx. decens in having brown instead of reddish mesonotal (scutal) scales and abdominal terga without pale bands, and added that the male genitalia of Cx. invidiosus differ from those of Cx. decens in having seta f longer and distally expanded on one side and seta h with a kink at mid-length. He concluded that “Since it has now been found that small differences exist in the male terminalia [genitalia], supporting the more obvious differences in colouring, it may be more correct to treat C. invidiosus as a distinct species rather than as a variety of C. decens.” He then described vexillatus as a variety with male genitalia that “Closely resemble those of C. invidiosus in all respects except as regards the shape of appendage f of the coxite lobe [gonocoxite subapical lobe]; this is greatly expanded at the tip, like a small flag; seta h sinuous as in typical invidiosus.” In addition to the character of seta f, comparison of Edwards’s illustrations of the partial phallosomes of invidiosus and vexillatus reveals a difference not noticed by Edwards, i.e. the dorsal arms of the lateral plates are larger, distally tapered and project beyond the largest tooth of the lateral arm in the type form whereas in vexillatus the dorsal arms are smaller, slightly enlarged distally and do not reach beyond the largest tooth of the lateral arm. Culex invidiosus vicinalis was described by de Meillon & Lavoipierre (1944) as “agreeing with invidiosus Theo. in all respects the only differences to be found in the male terminalia [genitalia]”, i.e. seta f of the subapical lobe is not enlarged apically, seta h is not sinuous and a unique double row of rather sharply bent setae is borne adjacent to the subapical lobe on the lateral surface of the gonocoxite. The authors pointed out that a similar double row of setae is present in an unnamed variety of Cx. ornatothoracis Theobald, 1909 (see Edwards 1941: fig. 118g), but because vicinalis bears an overall closer resemblance to Cx. invidiosus, de Meillon & Lavoipierre regarded it “as a subspecies of that species rather than of ornatothoracis.” Based on many years of taxonomic work (REH) on species of the genus Culex, especially species of the subgenus Culex, it is apparent that seemingly minor differences in features of the male genitalia are indicative of separate species. In the case of vexillatus and the typical form, differences now apparent in the development of the dorsal arms of the phallosome, supporting the previously noted differences in the form of setae f and h of the subapical lobe, it is more appropriate to treat vexillatus as a distinct species: Culex (Culex) vexillatus Edwards, 1941 . The situation with vicinalis is very different. Based on the diagnostic presence of a unique double row of setae on the gonocoxite, coupled with the distinctive development of setae f and h, it is surprising that vicinalis was not originally recognized as a distinct species; thus, it is hereby afforded specific status: Culex (Culex) vicinalis de Meillon & Lavoipierre, 1944 . Both Cx. vexillatus and Cx. vicinalis are listed as species in the Encyclopedia of Life. The larva of Cx. invidiosus is partially known (Wesché 1910; Edwards 1912d); the larvae of Cx. vexillatus and Cx. vicinalis are unknown. Once the larvae of all three species are known and have been studied and compared in detail, it is likely that morphological differences will be found that support their recognition as separate species. Molecular data are also expected to support their specific status. Three nominal species are recognized as junior synonyms of Cx. invidiosus: Cx. euclastus Theobald, 1903b (type locality: Gambia), Cx. chloroventer Theobald, 1909 (type locality: Accra, Ashanti Region, Ghana) and Cx. aquilus Graham, 1910 (type locality: Lagos, Nigeria). The type locality of Cx. invidiosus is the island town of Bonny in Rivers State in southern Nigeria.As there is no evidence that one or more of the three nominal forms might be conspecific with either Cx. vexillatus (type locality: Kampala, Uganda) or Cx. vicinalis (type locality: Yangambi, Tshopo Province, Democratic Republic of the Congo), they must remain as junior synonyms of Cx. invidiosus., Published as part of Harbach, Ralph E. & Wilkerson, Richard C., 2023, The insupportable validity of mosquito subspecies (Diptera: Culicidae) and their exclusion from culicid classification, pp. 1-184 in Zootaxa 5303 (1) on pages 86-87, DOI: 10.11646/zootaxa.5303.1.1, http://zenodo.org/record/8043342, {"references":["Theobald, F. V. (1901 d) Notes on a collection of mosquitoes from West Africa, and descriptions of new species. Memoirs of the Liverpool School of Tropical Medicine, 4 (Appendix), i - xiv, 3 pls.","Wilkerson, R. C., Linton, Y. - M. & Strickman, D. (2021) Mosquitoes of the world. Vols. 1 & 2. Johns Hopkins University Press, Baltimore, Maryland, 1332 pp. https: // doi. org / 10.1186 / s 13071 - 021 - 04848 - 6","Edwards, F. W. (1941) Mosquitoes of the Ethiopian Region III. - Culicine adults and pupae. Printed by order of the Trustees, British Museum (Natural History), London, viii + 499 pp.","Harbach, R. E. & Howard, T. M. (2007) Corrections in the status and rank of names used to denote varietal forms of mosquitoes (Diptera: Culicidae). Zootaxa, 1542 (1), 35 - 48. https: // doi. org / 10.11646 / zootaxa. 1542.1.3","de Meillon, B. & Lavoipierre, M. (1944) New records and species of biting insects from the Ethiopian region. Journal of the Entomological Society of Southern Africa, 7 (1), 38 - 67.","Theobald, F. V. (1901 c) A monograph of the Culicidae or mosquitoes. Vol. 2. British Museum (Natural History), London, viii + 391 pp. https: // doi. org / 10.5962 / bhl. title. 58067","Edwards, F. W. (1911 b) The African species of Culex and allied genera. Bulletin of Entomological Research, 2 (3), 241 - 268. https: // doi. org / 10.1017 / S 0007485300001450","Theobald, F. V. (1903 b) Report on a collection of mosquitoes or Culicidae, etc, from Gambia, and descriptions of new species. Memoirs of the Liverpool School of Tropical Medicine, 10 (Appendix), i - xi, 2 pls.","Theobald, F. V. (1909) Descriptions of the new mosquitoes collected by Dr. Graham in Ashanti. Gold Coast Miscellaneous Publication No. 237. Great Britain Colonial Office. London, 31 pp., 2 pls.","Graham, W. M. (1910) On new species of West-African Culicidae. Annals and Magazine of Natural History, including Zoology, Botany, and Geology, Series 8, 5 (27), 264 - 273. https: // doi. org / 10.1080 / 00222931008692764","Wesche, W. (1910) On the larval and pupal stages of West African Culicidae. Bulletin of Entomological Research, 1 (1), 7 - 50, 7 pls. https: // doi. org / 10.1017 / S 0007485300000043","Edwards, F. W. (1912 d) Revised keys to the known larvae of African Culicinae. Bulletin of Entomological Research, 3 (4), 373 - 385. https: // doi. org / 10.1017 / S 0007485300002042","Edwards, F. W. (1914) New species of Culicidae in the British Museum, with notes on the genitalia of some African Culex. Bulletin of Entomological Research, 5 (1), 63 - 81. https: // doi. org / 10.1017 / S 0007485300034568","Newstead, R., Dutton, J. E. & Todd, J. L. L (1907) Insects and other Arthropoda collected in the Congo Free State. Annals of Tropical Medicine and Parasitology, 1 (1 - 5), 3 - 113. https: // doi. org / 10.1080 / 00034983.1907.11719252","Macfie, J. W. S. & Ingram, A. (1920) The early stages of West African mosquitos. V. - Culex decens, Theo. and Culex invidiosus, Theo. Bulletin of Entomological Research, 11 (2), 104 - 112. https: // doi. org / 10.1017 / S 0007485300044515","Hopkins, G. H. E. (1936) Mosquitoes of the Ethiopian Region I. - Larval bionomics of mosquitoes and taxonomy of culicine larvae. Printed by order of the Trustees, British Museum (Natural History), London, (vi) + 250 pp.","Hopkins, G. H. E. (1952) Mosquitoes of the Ethiopian Region I. - Larval bionomics of mosquitoes and taxonomy of culicine larvae. 2 nd Edition. British Museum (Natural History), London, viii + 355 pp.","Edwards, F. W. (1932 a) Genera Insectorum. Diptera, Fam. Culicidae. Fascicle 194. V. Verteneuil & L. Desmet, Bruxelles [Brussels], 258 pp., 5 pls."]}

Culex (Culex) toroensis Edwards & Gibbins subspecies macrophyllus Edwards & Gibbins, 1939 —original combination: Culex vansomereni spp. macrophyllus (subspecific status by Edwards, 1941 concomitant with the elevation of toroensis to specific status). Distribution: Cameroon, Uganda (Knight & Stone 1977). subspecies toroensis Edwards & Gibbins, 1939 —original combination: Culex vansomereni toroensis (specific status by Edwards 1941). Distribution: Burundi, Cameroon, Democratic Republic of the Congo, Ethiopia, Kenya, Malawi, Republic of South Africa, Rwanda, South Sudan, Sudan, Uganda (Wilkerson et al. 2021). The typical form and subspecies macrophyllus were originally described as subspecies of Cx. vansomereni Edwards, 1926a. Edwards (1941) elevated toroensis to specific status based on “well-marked larval distinctions from C. vansomereni, together with the constant difference in pleural scaling”; consequently, macrophyllus became a subspecies of Cx. toroensis. Edwards & Gibbins (1939) and Edwards (1941) distinguished the subspecies based solely on features of the male genitalia. In the typical form, foliform seta g of the subapical lobe is about as long as seta f and more or less pointed, the lateral plate of the phallosome has about five denticles and the gonostylus is long and markedly narrowed distally. In comparison, seta g is much larger than seta f in subspecies macrophyllus, the lateral plate has a slightly different shape with about three denticles and the gonostylus is distinctly shorter and not narrowed distally. The larva of the typical form is known from specimens reared to adults (Edwards 1941; Service 1959). The larva of subspecies macrophyllus is not definitely known, but larvae associated with adults collected at the type locality (Edwards & Gibbins 1939) “bore a close resemblance” to the larva of Cx. andersoni Edwards, 1914, which is very distinct from the larva of subspecies toroensis. The larva of the typical form has not been studied in detail—only features of the head and terminal abdominal structures have been described, with special emphasis on the anterior (dorsal) and posterior (ventral) clusters of spines near the apex of the siphon. As noted by Service (1959), specimens of toroensis from Kenya and Sudan examined in the “British Museum” did not exhibit the arrangement of 18 anterior siphonal spines illustrated by Hopkins (1952), and specimens from elsewhere had 0–9 anterior spines. It should be mentioned, however, that a male from Kisomoro, Uganda, the type locality of toroensis, was reared from a “larva similar to that figured by Hopkins [1936] … as sp. indet.” (Edwards & Gibbins 1939), and was illustrated by Hopkins (1952) as the larva of Cx. toroensis. Both Hopkins and Service noted the variable presence of 0–3 posterior spines. In agreement with Service (1959) and Jupp (1996), the variable number of apical siphonal spines makes it impossible to reliably identify the species, e.g. to distinguish larvae from those of Cx. vansomereni. On the other hand, we feel that such variation may be an indication of a complex of closely related species. Oddly, Edwards & Gibbins (1939) described macrophyllus (p. 31) prior to describing toroensis (p. 32); hence, macrophyllus could have been taken as the nominotypical species with toroensis as a subspecies of Cx. macrophyllus. Mattingly (1956) revealed that macrophyllus was described from six males, five with dissected genitalia, and four females collected at “Lugezi Camp between Mt. Mgahinga [Mt Gahinga] and Mt. Sabinio [Mt Sabyinyo]” located at the extreme southwestern fringe of Uganda along the border with Rwanda [actually, Mt Sabyinyo, an extinct volcano, is situated at the intersection of present-day Democratic Republic of the Congo, Rwanda and Uganda, and Mt Gahinga, a dormant/extinct volcano, is on the border between Rwanda and Uganda]. Lugezi Camp was apparently located on the Uganda side of the current border with Rwanda. The two mountains were located in the former Kigezi District, which included flanking areas of the two currently neighboring countries. The former Kigezi District now includes the Kabale, Kanungu, Kisoro and Rukungiri Districts and the two mountains are situated at the margins of the Kisoro District. Edwards (1941) indicated that macrophyllus was found at the “Saddle between Mts. Mgadinga and Sabinio, 8000 ft. [2,438 m]”, but according to Edwards & Gibbins (1939) Lugezi Camp was located at the side of “Mt. Sabinio” at an elevation of “ 7–8000 ft. [2,135 –2,438 m]”. Mattingly (1956) observed that toroensis was described from five males, two incomplete and three with dissected genitalia, and one female from Kisomoro, Uganda and nine males and nine females from Kararama Camp in the Namwamba Valley, and designated a male with dissected genitalia from Kisomoro as the name-bearing lectotype. Kisomoro (elevation 1,492 m), situated in the Bunyangabu District in the Western Region of Uganda, is located approximately 240 km north of the type locality of macrophyllus. It is interesting that the following statement made by Edwards & Gibbins (1939) regarding toroensis was apparently overlooked by later workers: “These specimens [from Kararama Camp, Namwamba Valley] have the male hypopygium [genitalia] almost exactly as in the typical form [vansomereni], and do not show the enlarged leaf and other slight peculiarities seen in specimens from Kigezi [i.e. Lugezi Camp].” The Namwamba Valley is located approximately 37 km southwest of Kisomoro, indicating that toroensis is probably widely distributed in the mountainous regions of western Uganda. In view of the differences in structures of the male genitalia of the two forms, especially the very different gonostylus, the extensive variation observed in the anterior siphonal spines of larvae identified as toroensis suggesting that it may be a species complex, the likelihood that the larva of macrophyllus is similar to the larva of Cx. andersoni, and the probability that toroensis and macrophyllus are likely to occur in sympatry, we feel confident that the latter form is a separate species and hereby formally accord it specific status: Culex (Culex) macrophyllus Edwards & Gibbins, 1939 . Culex macrophyllus is currently listed as a species in the Encyclopedia of Life.

Culex (Culex) comorensis Brunhes subspecies comorensis Brunhes, 1977 —original combination: Culex (Culex) comorensis. Distribution: Anjouan and Mohéli Islands, Union of the Comoros and Ankaratra Massif, Madagascar (Brunhes 1977). subspecies kartalae Brunhes, 1977 —original combination: Culex (Culex) comorensis ssp. kartalae. Distribution: Grande Comore [Union of the Comoros], Mayotte [overseas department of France] (Brunhes 1977). Culex comorensis was described and named from specimens discovered on Anjouan Island, an autonomous high island that forms part of the Union of the Comoros. Brunhes (1977) indicated that it was also found on Mohéli Island and the Ankaratra Massif of Madagascar. Subspecies kartalae was briefly described and named in the same publication from specimens collected on Grande Comore Island. Brunhes pointed out morphological “peculiarities” of the male genitalia and larvae that distinguish the two forms. In the typical form, the subapical lobe of the gonocoxite bears a few minute setae at the bases of setae a-f; in the larva, head seta 13-C is long (0.6 mm), seta 1-S of the siphon usually has 4 pairs of setae on the posterior margin, the pecten consists of 10–14 spines (mean 11.2), and the dorsal and ventral anal papillae are equal in length. In subspecies kartalae, the subapical lobe of the gonocoxite bears a dense cluster of minute setae at the bases of setae a-f; the larva differs in having a short seta 13-C (0.2–0.3 mm), the siphon usually has 5 pairs of seta 1-S on the posterior margin, the pecten consists of 4–10 spines (mean 7.2) and the dorsal pair of anal papillae are longer than the ventral pair. The author, however, overlooked some additional important differences of the male genitalia. Subspecies kartalae differs from the typical form as follows: The ventral arms of the phallosome are longer and more slender, the dorsal arms are more distinctly minutely toothed and more sharply pointed, setae a, b, c and h of the subapical lobe are distinctly longer than seta g, setae a and b are stouter and more distinctly hooked apically and the insertion of seta c is borne mesad of setae a and b whereas it is inserted distal to these two setae in the typical form. The complexity of morphological distinctions and the geographical isolation are a clear indication that kartalae is a distinct species; thus, this form is hereby afforded specific status: Culex (Culex) kartalae Brunhes, 1977 . Culex kartalae is currently listed as a species in the Encyclopedia of Life., Published as part of Harbach, Ralph E. & Wilkerson, Richard C., 2023, The insupportable validity of mosquito subspecies (Diptera: Culicidae) and their exclusion from culicid classification, pp. 1-184 in Zootaxa 5303 (1) on page 84, DOI: 10.11646/zootaxa.5303.1.1, http://zenodo.org/record/8043342, {"references":["Brunhes, J. (1977) Les moustiques de l'archipel des Comores I. - Inventaire, repartition et description de quatre especes ou sous-especes nouvelles. Cahiers ORSTOM Serie Entomologie medicale et Parasitologie, 15 (2), 131 - 152."]}

Culex (Culex) trifilatus Edwards subspecies aenescens Edwards, 1941 —original combination: Culex (Culex) trifilatus ssp. aenescens. Distribution: Eritrea (Mara 1945), Uganda (Edwards 1941). subspecies trifilatus Edwards, 1914 —original combination: Culex trifilatus. Distribution: Angola, Cameroon, Democratic Republic of the Congo, Ethiopia, Gabon, Guinea, Kenya, Madagascar, Malawi, Mozambique, Republic of the Congo, Republic of South Africa, South Sudan, Sudan, Tanzania, Uganda, Zimbabwe, Zambia (Wilkerson et al. 2021). Culex trifilatus was descried by Edwards (1914) based on adult males and females collected in Kabete, Kenya. In 1941, he described subspecies aenescens from adults captured in the Toro District of Uganda. Subspecies aenescens was distinguished principally from the type form in having seta c of the subapical lobe of the male gonocoxite as long as seta b (seta c is shorter than b in the type form), setae d and e absent (both present in the type form) and seta f very slender (broad and flattened in the type form). However, the illustrations of the genitalia provided by Edwards (1941) clearly indicate that the two laterally bent projections of the lateral plate are more slender, the dorsal one is tapered and pointed, and the dorsal arm is narrower and longer in subspecies aenescens (the dorsal projection is broadened apically in the nominate subspecies). The larva of subspecies aenescens was described by Hopkins (1952) from many specimens collected in the Toro District. The larva of the nominate subspecies was not known until it was described by Ribeiro et al. (1982) based on specimens from Angola (Ribeiro & da Cunha Ramos 1980) and Tanzania (V. N. Danilov). As described by these researchers, larvae of subspecies aenescens differ from those of the type form as follows: Seta 5-C with 3 branches (4 or more in the type form); siphonal seta 1-S comprised of 3 pairs of alternating setae with 2–5 branches (with 3–9 branches in the type form); seta 1-X single or double (always single in the type form); dorsal pair of anal papillae 1.5 times as long as the ventral pair, which are about as long as the saddle (dorsal pair 3 times as long as the ventral pair in the type form, but the ventral pair are only about half as long as the saddle). Ribeiro et al. (1982) stated that the two subspecies are allopatric based on then present knowledge of their geographical distributions. This is apparently incorrect as both forms are recorded from localities in Uganda and a footnote in Hopkins (1952) provided by P. F. Mattingly indicates that Mara (1945) recorded the presence of subspecies aenescens in Eritrea. Stone et al. (1959), and later catalogs, indicate that the type form has been found in Ethiopia and Sudan, but without identifying the sources of those records and whether they are based on adult or larval mosquitoes. Assuming the identifications are correct, it seems likely that the two subspecies occur in sympatry in northeastern areas of the Afrotropical Region. It is an undeniable fact that most species of Afrotropical Culex are incompletely (inadequately) described and illustrated or otherwise only superficially known. As a result of critical revisionary studies, it is known that many currently recognized species of the genus are distinguished by a combination of seemingly minor differences in individual life stages, e.g. species of the Oriental Vishnui Group (Sirivanakarn 1976), and some nominal species thought to be conspecific have been found to be distinct species with very similar male genitalia that exhibit distinctive minor differences, e.g. Cx. bidens Dyar, 1922 and Cx. interfor Dyar, 1928 (Harbach et al. 1986). It is well known that male genitalia generally provide a better means for distinguishing species than any other morphological characters, especially members of the genus Culex. Evolution of genital form is thought to be involved in the origin of species and the reproductive isolation between species. Thus, the differences in the male genitalia noted above would alone provide prima facie evidence of separate species; however, those differences coupled with the larval differences and the probable sympatry of the two forms further strengthens the likelihood that aenescens is a valid species. For these reasons, we feel justified in formally recognizing this nominal subspecific taxon as a distinct species: Culex (Culex) aenescens Edwards, 1941 . Culex aenescens is currently listed as a species in the Encyclopedia of Life., Published as part of Harbach, Ralph E. & Wilkerson, Richard C., 2023, The insupportable validity of mosquito subspecies (Diptera: Culicidae) and their exclusion from culicid classification, pp. 1-184 in Zootaxa 5303 (1) on page 92, DOI: 10.11646/zootaxa.5303.1.1, http://zenodo.org/record/8043342, {"references":["Edwards, F. W. (1941) Mosquitoes of the Ethiopian Region III. - Culicine adults and pupae. Printed by order of the Trustees, British Museum (Natural History), London, viii + 499 pp.","Mara, L. (1945) Considerazioni sul rinvenimento dell' Aedes aegypti (Dip. Aedinae) ad altitudini d'eccezione' e brevi note sulla fauna culicidica de 1 M. Bizen (Eritrea, A. O.). Bollettino. Societa italiana di medicina e igiene tropicale. Sezione Eritrea, 5, 189 - 198.","Edwards, F. W. (1914) New species of Culicidae in the British Museum, with notes on the genitalia of some African Culex. Bulletin of Entomological Research, 5 (1), 63 - 81. https: // doi. org / 10.1017 / S 0007485300034568","Wilkerson, R. C., Linton, Y. - M. & Strickman, D. (2021) Mosquitoes of the world. Vols. 1 & 2. Johns Hopkins University Press, Baltimore, Maryland, 1332 pp. https: // doi. org / 10.1186 / s 13071 - 021 - 04848 - 6","Hopkins, G. H. E. (1952) Mosquitoes of the Ethiopian Region I. - Larval bionomics of mosquitoes and taxonomy of culicine larvae. 2 nd Edition. British Museum (Natural History), London, viii + 355 pp.","Ribeiro, H., Danilov, V. N. & da Cunha Ramos, H. (1982) Description of the fourth instar larva of Culex (Culex) trifilatus trifilatus Edwards, 1914 (Diptera, Culicidae). Mosquito Systematics, 14 (2), 94 - 100.","Stone, A., Knight, K. L. & Starcke, H. (1959) A synoptic catalog of the mosquitoes of the world (Diptera, Culicidae). The Thomas Say Foundation. Vol. VI. Entomological Society of America, College Park, Maryland, vi + 358 pp.","Sirivanakarn, S. (1976) Medical entomology studies - III. A revision of the subgenus Culex in the Oriental Region (Diptera: Culicidae). Contributions of the American Entomological Institute, 12 (2), iii, 1 - 272.","Dyar, H. G. (1922) Notes on tropical American mosquitoes (Diptera, Culicidae). Insecutor Inscitiae Menstruus, 10 (10 - 12), 188 - 196, 1 pl.","Dyar, H. G. (1928) The mosquitoes of the Americas. Publication No. 387. Carnegie Institution of Washington, Washington, D. C., 616 pp.","Harbach, R. E., Jakob, W. L. & Peyton, E. L. (1986) Recognition of Culex bidens Dyar and Culex interfor Dyar (Diptera: Culicidae) as separate species. Mosquito Systematics, 18 (2), 139 - 144."]}

Culex (Eumelanomyia) horridus Edwards subspecies horridus Edwards, 1922 —original combination: Culex horridus. Distribution: Angola, Benin, Burkina Faso, Cameroon, Central African Republic, Comoros, Côte d’Ivoire, Democratic Republic of the Congo, Ghana, Kenya, Liberia, Madagascar, Mali, Mozambique, Nigeria, Republic of the Congo, Republic of South Africa, South Sudan [but not Sudan (Simsaa et al. 2021)], Tanzania, Togo, Uganda, Zambia, Zimbabwe (Wilkerson et al. 2021). subspecies rageaui (Hamon & Rickenbach, 1955) —original combination: Neoculex horridus var. rageaui (subspecific status by Harbach & Howard 2007). Distribution: Cameroon (Hamon & Rickenbach 1955). These authors considered specimens from Benin and Burkina Faso that they compared with subspecies rageaui to be specimens of the nominotypical form (see below); thus, those two countries were erroneously included in the distribution of rageaui in the catalogs of Knight & Stone (1977) and Wilkerson et al. (2021). Edwards (1922) proposed horridus as a replacement name for Protomelanoconion fusca Theobald, 1909, type locality Accra in present-day Ghana, which was preoccupied by Taeniorhynchus fuscus Theobald, 1905d, a synonym of Culex (Culiciomyia) fragilis Ludlow, 1903. Unfortunately, Edwards (1922, 1941) only briefly described (with lack of detail) and did not illustrate the male genitalia of horridus. Hamon & Rickenbach (1955) examined two series of specimens, one consisting of three males, one from Benin and two from Burkina Faso, and the other consisting of six males from Cameroon. They provided two illustrations of the subapical lobe of the gonocoxite, one drawn from a male from Burkina Faso labelled as horridus and the other drawn from the holotype of rageaui from Cameroon. The two illustrations are similar except in the former there is only one seemingly flexible simple seta (d or e) in group d–f and the three rod-like setae a–c are closely aligned parallel to each other with c inserted slightly distal to a and b. Subspecies rageaui has two straight (stiff?) setae (d and e) in group d–f and seta c projects at a 90-degree angle from setae a and b, which is very unusual and could be due to distortion. Except for setae d and e, the characteristics (shapes) of the other setae (a–c and f–h) are otherwise the same. Jupp (1996) also illustrated the subapical lobe of horridus, but setae d–g are very different than those illustrated by Hamon & Rickenbach. Whereas group d–f of the males from Burkina Faso and Cameroon consists of one or two simple setae (d and e) and four blades (f) with rounded tips and three or four apical barbs, group d–f of the specimen illustrated by Jupp consists of setae d and e and a single seta f with a bifid tip (which Jupp indicated may be an artefact). Whereas seta g is distinctly asymmetrical and as long as or slightly longer than seta f in the specimens illustrated by Hamon & Rickenbach, it is symmetrical and shorter than seta f in the specimen illustrated by Jupp. Obviously the species illustrated by Jupp is not conspecific with the species examined by Hamon & Rickenbach. Hamon & Rickenbach stated that the genitalia of their two series of specimens are identical, and noted that they differed from Edwards’s description of horridus as follows: “the three rods (rods or blades) [setae a–c] are pointed at their apex, and at least two are curved into a hook; the four strong setae (blunt tipped setae) [seta f] are in fact narrow blades, rounded at their apex, and each bearing 3 to 4 subapical barbs; they are accompanied by one or two setae [d and e] thinner and shorter than themselves; the leaf [seta g], apparently unstriated, is very asymmetrical in shape and is accompanied by a strong seta [h] as long or longer than it…” (translated from the French). However, whereas the scutal scales are dark brown in the specimens of horridus from Benin and Burkina Faso, as described by Edwards for the typical form, the scutum of rageaui is “uniformly covered with yellowish-white scales” (translated from the French), whence the characteristic that distinguishes subspecies rageaui from the description of the type form provided by Edwards. The nature of the differences, especially the major difference in the color of the scutal scaling, prompted one of us (REH) to examine the syntype males of Protomelanoconion fusca Theobald, 1909 (for which Edwards replaced with the name horridus) in the Natural History Museum (NHM), London. This resulted in an unexpected discovery. The subapical lobe shown in the two drawings of Hamon & Rickenbach (1955) is very different than the subapical lobe of the syntypes, and also different than the drawing of Jupp (1996). Out of curiosity, drawings of subapical lobes in Edwards (1941) were scanned to see if any of the other species he treated was similar to the syntypes of fusca (= horridus). Edwards classified most species currently in the subgenera Eumelanomyia and Maillotia as species of the subgenus Neoculex. Astonishingly, the illustration of the subapical lobe he provided for Cx. salisburiensis Theobald, 1901c, a species of the subgenus Maillotia, agrees with that of fusca, particularly the shape of setae d–f and the distinctive foliform seta g with an elongate stem. Consequently, it seemed necessary to look at other treatments of salisburiensis, which resulted in another unexpected discovery. Knight (1953), in his paper on the mosquitoes of Yemen, consulted Peter Mattingly at the British Museum (Natural History) (now the NHM), concerning differences between the male genitalia of specimens they identified as salisburiensis and the genitalia of the species illustrated by Edwards (1941). Mattingly responded with the following. I am afraid almost all of the differences you noted are due to errors in Edwards’ description. The only difference of any significance is that, while the Nairobi males have terminalia [genitalia] identical with the Yemen form, those from further south lack the longest of three accessory bristles [seta f] on the subapical lobe and one of the two small setae on the dististyle [gonostylus]. It is clear therefore that we have a northern and a southern form but it is impossible to say which is the type form since I have no males from Salisbury [Harare, Zimbabwe]…. It may be possible eventually to distinguish two subspecies. The southern forms are certainly much darker than yours, especially with respect to the scutal scaling but the Nairobi specimens are intermediate and, I should think, would probably intergrade in Eritrea. Also the two specimens from Chilanga [Zambia] suggest that there is considerable seasonal variation since one, collected in January, is very dark, while the other collected in November, is as pale as the Nairobi form. Edwards’ figure for the length of the female palps [maxillary palpi] seems to have been based on a single aberrant or shrunken specimen; and the tergal bands are variable, even in the same locality. The other differences are just errors in description. It appears that the figure of the subapical lobe that Edwards’s attributed to Cx. salisburiensis is that of Cx. horridus (i.e. P. fusca). His descriptions of the genitalia of both species are brief, but the description given for Cx. horridus and the illustration attributed to Cx. salisburiensis clearly correspond with the syntypes of P. fusca. Therefore, there is little doubt that the species illustrated by Hamon & Rickenbach, and also Jupp, are not Cx. horridus and are separate species. The only problem that remains is the question about the color of the scutal scaling. Hamon & Rickenbach cite Edwards’s description of Cx. horridus, which says the scutal scales are dark. The scutal scales of the syntypes of P. fusca definitely are not dark, but are golden and those on the anterior margin are white. Examination of specimens in the NHM from several East African countries all have golden scutal scales with white scales on the anterior margin. This would suggest an error in the description, or perhaps Edwards’s interpretation of dark scutal scaling due the optics and light sources that were available at the time. However, as the genitalia of the males with dark and yellowish white scutal scales are the same, it would seem possible that this is due to seasonal or geographical variation within a single species, or they may be two different closely related species. In any case, the subspecific form described by Hamon & Rickenbach must be considered to be a distinct species, Culex (Eumelanomyia) rageaui Hamon & Rickenbach, 1955 , and the question of whether or not the form with dark scutal scaling is the same or a different species must await further study. Culex rageaui is currently listed as a species in the Encyclopedia of Life., Published as part of Harbach, Ralph E. & Wilkerson, Richard C., 2023, The insupportable validity of mosquito subspecies (Diptera: Culicidae) and their exclusion from culicid classification, pp. 1-184 in Zootaxa 5303 (1) on pages 96-97, DOI: 10.11646/zootaxa.5303.1.1, http://zenodo.org/record/8043342, {"references":["Edwards, F. W. (1922) Mosquito notes. - III. Bulletin of Entomological Research, 13 (1), 75 - 102. https: // doi. org / 10.1017 / S 0007485300045259","Simsaa, M. A. A., Harbach, R. E., Almalik, A. M. A., Ahmed, E. M., Esia, A. A., Mohamed, A. H. & Azrag, R. S. (2021) Culex mosquitoes (Diptera: Culicidae) recorded along the Nile River in northern Sudan, with a key for the identification of all species of the genus known to occur in the country. Zootaxa, 4963 (3), 401 - 411. https: // doi. org / 10.11646 / zootaxa. 4963.3.1","Wilkerson, R. C., Linton, Y. - M. & Strickman, D. (2021) Mosquitoes of the world. Vols. 1 & 2. Johns Hopkins University Press, Baltimore, Maryland, 1332 pp. https: // doi. org / 10.1186 / s 13071 - 021 - 04848 - 6","Hamon, J. & Rickenbach, A. (1955) Contribution a l'etude des Neoculex (Dipteres: Culicides) de la Region Ethiopienne. 1. - Corrections de quelques descriptions de terminalia males donnees par Edwards, avec etude d'une nouvelle variete. Bulletin de la Societe de Pathologie exotique, 48 (6), 848 - 859.","Harbach, R. E. & Howard, T. M. (2007) Corrections in the status and rank of names used to denote varietal forms of mosquitoes (Diptera: Culicidae). Zootaxa, 1542 (1), 35 - 48. https: // doi. org / 10.11646 / zootaxa. 1542.1.3","Knight, K. L. & Stone, A. (1977) A catalog of the mosquitoes of the world (Diptera: Culicidae). 2 nd Edition. The Thomas Say Foundation. Vol. VI. Entomological Society of America, College Park, Maryland, xi + 611 pp.","Theobald, F. V. (1909) Descriptions of the new mosquitoes collected by Dr. Graham in Ashanti. Gold Coast Miscellaneous Publication No. 237. Great Britain Colonial Office. London, 31 pp., 2 pls.","Theobald, F. V. (1905 d) Some new mosquitoes from Ceylon. Journal of the Bombay Natural History Society, 16 (2), 237 - 250, 2 pls.","Ludlow, C. S. (1903) Some Philippine mosquitoes. Journal of the New York Entomological Society, 11 (3), 137 - 144.","Edwards, F. W. (1941) Mosquitoes of the Ethiopian Region III. - Culicine adults and pupae. Printed by order of the Trustees, British Museum (Natural History), London, viii + 499 pp.","Jupp, P. G. (1996) Mosquitoes of southern Africa: Culicinae and Toxorhynchitinae. Ekogilde Publishers, Hartebeespoort, 156 pp.","Theobald, F. V. (1901 c) A monograph of the Culicidae or mosquitoes. Vol. 2. British Museum (Natural History), London, viii + 391 pp. https: // doi. org / 10.5962 / bhl. title. 58067"]}

Culex (Lophoceraomyia) curtipalpis (Edwards) subspecies curtipalpis (Edwards, 1914 c) —original combination: Lophoceratomyia curtipalpis. Distribution: Cambodia, Indonesia, Malaysia, Singapore, Thailand, Vietnam (Wilkerson et al. 2021). subspecies sumatranus Brug, 1931 —original combination: Culex (Culex) sumatranus (subspecific status by Meng & Chen 1980). Distribution: Cambodia, Indonesia, Macau, People’s Republic of China, Vietnam (Wilkerson et al. 2021). Edwards (1914 c) described curtipalpis from five males collected at Kuching Reservoir in Sarawak, Malaysia. His description did not mention or include an illustration of the male genitalia. Edwards (1928) added some details to his previous description, including a brief description of the genitalia and an illustration of the gonostylus. The larva and pupa were described by Edwards & Given (1928) based on association with reared adults. The male and larva were later described and illustrated in more detail by Colless (1965), Bram (1967) and Sirivanakarn (1977), and the pupa was described by the last author. Subspecies sumatranus was originally described by Brug (1931) as a species of the subgenus Culex, described from a single male reared from a larva taken from a pitcher plant in the vicinity of Dermajoe, Benkoelen, Sumatra. Brug illustrated the male genitalia and the head and terminal abdominal segments of the larval exuviae. The species was transferred to the subgenus Neoculex by Brug & Edwards (1931), and this was accepted (see Stone et al. 1959) until Sirivanakarn (1971) recognized it as a species of the subgenus Lophoceraomyia. It was later regarded as a subspecies of Cx. curtipalpis by Meng & Chen (1980). As revealed by Barraud (1934), in a footnote on page 351, and discussed by Sirivanakarn (1977), the larva described and illustrated by Brug (1931) was wrongly associated with the adult male, and adults reared from larvae collected in Hong Kong and elsewhere are identical with the larva of curtipalpis described by Edwards & Given (1928). Sirivanakarn treated sumatranus as a distinct species, but noted that “All stages of sumatranus are exceedingly similar to and indistinguishable from curtipalpis except for the male which differs from the latter rather strikingly in the absence of the modified tufts of the male antenna.” For clarity, unlike curtipalpis, the flagellar whorls of sumatranus are weakly verticillate or comprised of relatively fewer long setae and tufts of modified setae/scales are absent on flagellomeres 5–9 or a rudimentary or inconspicuous tuft of four very short setae may be present on the mesal surface of flagellomere 7. Contrary to Sirivanakarn (1977), Meng & Chen (1980) considered this striking difference to be nothing more than interspecific variation. In view of the poor illustration of the male genitalia of sumatranus provided by Brug (1931), we decided it was important to examine the holotype in the Natural History Museum, London. The dissected genitalia of the holotype are poorly mounted on a microscope slide. The left gonocoxite seems to be positioned differently than indicated in Brug’s illustration, and the gonostylus, which is tapered to the apex in the illustration, is expanded and strongly modified as illustrated by Colless (1965: fig. 29d), Bram (1967: fig. 22) and Sirivanakarn (1977: fig. 69). The setae of the subapical lobe are inaccurately drawn, but the two setae proximal to setae a–c are clearly present. Setae a–c are more tapered and pointed than illustrated, and also tapered more distally than illustrated by Colless, Bram and Sirivanakarn for curtipalpis. Foliform seta g is not shown in the drawing, but it probably was not seen by Brug because it is very faint and difficult to see even under differential interference contrast microscopy, which we used to examine the genitalia. Surprisingly, it is much longer and narrower than seta g shown in the illustrations of the genitalia of curtipalpis provided by Colless, Bram and Sirivanakarn, which is short and very broad, about as broad as long. It is likely that Sirivanakarn (1977) also did not see seta g when he examined the holotype of sumatranus (he undoubtedly used traditional bright-field microscopy), otherwise he would have noted this obvious difference in his description of the species. These differences, along with the absence of modified tufts of setae or scales on the antennal flagellum, are a clear indication that sumatranus is a species distinct from curtipalpis. Based solely on the antennal characteristics, both forms have been found in areas of Cambodia, Indonesia (Sumatra) and Vietnam. Considering this apparent sympatry and the morphological distinctions exhibited by the adult male, we believe curtipalpis and sumatranus are genetically distinct species and hereby reinstate the latter to its original specific status: Culex (Lophoceraomyia) sumatranus Brug, 1931 . Culex sumatranus is currently listed as a species in the Encyclopedia of Life. Culex sumatranus has a single synonym, Cx. gudouensis Chang, Zhao, Hang & Chen, 1975 (type locality: Xin-hui Shien, Kwangtung Province, People’s Republic of China), which was synonymized with sumatranus by Meng & Chen (1980). The illustration of the subapical lobe of Cx. gudouensis provided by Chang et al. (1975) is very poor. There is no indication of seta g, but Meng & Chen examined the type specimens and noted the presence of a “broad leaf” that was “ignored in the original description” (translated from the Chinese). This does not clearly indicate, however, whether seta g is like that of Cx. curtipalpis or Cx. sumatranus, but Meng & Chen also noted that a tuft of specialized short setae was present only on the seventh flagellomere of the antenna, and used this as the primary character to support the synonymy of Cx. gudouensis. Unless other information becomes available to indicate otherwise, Cx. gudouensis should continue to be recognized as junior synonym of Cx. sumatranus., Published as part of Harbach, Ralph E. & Wilkerson, Richard C., 2023, The insupportable validity of mosquito subspecies (Diptera: Culicidae) and their exclusion from culicid classification, pp. 1-184 in Zootaxa 5303 (1) on pages 98-99, DOI: 10.11646/zootaxa.5303.1.1, http://zenodo.org/record/8043342, {"references":["Edwards, F. W. (1914) New species of Culicidae in the British Museum, with notes on the genitalia of some African Culex. Bulletin of Entomological Research, 5 (1), 63 - 81. https: // doi. org / 10.1017 / S 0007485300034568","Wilkerson, R. C., Linton, Y. - M. & Strickman, D. (2021) Mosquitoes of the world. Vols. 1 & 2. Johns Hopkins University Press, Baltimore, Maryland, 1332 pp. https: // doi. org / 10.1186 / s 13071 - 021 - 04848 - 6","Meng, Q. & Chen, H. (1980) Arguments on the taxonomic positions of six species of Culex from China. Entomotaxonomia, 2 (2), 161 - 164. [in Chinese]","Edwards, F. W. & Given, D. H. C. (1928) The early stages of some Singapore mosquitos. Bulletin of Entomological Research, 18 (4), 337 - 357. https: // doi. org / 10.1017 / S 0007485300020265","Colless, D. H. (1965) The genus Culex, subgenus Lophoceraomyia in Malaya (Diptera: Culicidae). Journal of Medical Entomology, 2 (3), 261 - 307. https: // doi. org / 10.1093 / jmedent / 2.3.261","Bram, R. A. (1967) Contributions to the mosquito fauna of Southeast Asia. II. The genus Culex in Thailand (Diptera: Culicidae). Contributions of the American Entomological Institute, 2 (1), 1 - 296.","Sirivanakarn, S. (1977) Medical entomology studies - VI. A revision of the subgenus Lophoceraomyia of the genus Culex in the Oriental Region (Diptera: Culicidae). Contributions of the American Entomological Institute, 13 (4), iii + 1 - 245.","Brug, S. L. & Edwards, F. W. (1931) Fauna Sumatrensis. (Bijdrage Nr. 68), Culicidae (Diptera). Tijdschrift voor Entomologie, 74, 251 - 261.","Stone, A., Knight, K. L. & Starcke, H. (1959) A synoptic catalog of the mosquitoes of the world (Diptera, Culicidae). The Thomas Say Foundation. Vol. VI. Entomological Society of America, College Park, Maryland, vi + 358 pp.","Barraud, P. J. (1934) The fauna of British India, including Ceylon and Burma. Diptera. Vol. V. Family Culicidae. Tribes Megarhinini and Culicini. Taylor and Francis, London, xxviii + 463 pp., 8 pls.","Chang, H. - T., Zhao, S. X., Hang, B. - X. & Chen, S. - F. (1975) A new species of Culex. Acta Entomologica Sinica, 18 (3), 338 - 340. [in Chinese]"]}

Culex (Oculeomyia) annulioris Theobald subspecies annulioris Theobald, 1901a —original combination: Culex annulioris. Distribution: Angola, Benin, Burkina Faso, Cameroon, Central African Republic, Côte d’Ivoire, Democratic Republic of the Congo, Ethiopia, Ghana, Guinea, Kenya, Liberia, Madagascar, Malawi, Mali, Nigeria, Republic of the Congo, São Tomé and Príncipe, Senegal, Seychelles, South Africa, South Sudan, Sudan, Tanzania, Uganda, Zimbabwe (Wilkerson et al. 2021). subspecies consimilis Newstead, 1907 (in Newstead et al. 1907)—original combination: Culex tigripes var. consimilis (subspecific status by Edwards 1941). Distribution: Cameroon, Democratic Republic of the Congo, Gabon, Ghana, Kenya, Liberia, Mozambique, Nigeria, Sierra Leone, South Sudan (Lewis 1956, but not Sudan, see Simsaa et al. 2021), Tanzania, Uganda (Wilkerson et al. 2021). Culex annulioris was described from a single female collected in Salisbury, Southern Rhodesia (present-day Harare, Zimbabwe), “distinguished by the abdominal ornamentation, which is very marked and peculiar”, “each segment with a triangular basal patch of creamy yellow scales, and with apical lateral patches of yellow scales” (Theobald 1901a). Subspecies consimilis, originally proposed as a variety of Lutzia tigripes (de Grandpré & de Charmoy, 1901) and subsequently recognized as a subspecies of annulioris by Edwards (1941), was described from females collected at several localities in the Belgium Congo (present-day Democratic Republic of the Congo). The abdomen of the female was described by Newstead (in Newstead et al. 1907) as having “narrow basal bands; basal segment with two median black apical spots; sixth and seventh segments with two lateral apical pale spots.” As pointed out in a footnote, Edwards (1941) observed that “in the lectotype the slight appearance of banding is merely due to the pale marginal hairs [setae] and not to pale scales.” He also noted that “only small apical lateral spots, if any…” were present on segments VI and VII. Oddly, except for Edwards, there is no mention of a lectotype of consimilis in the catalogs of Stone et al. (1959), Knight & Stone (1977) and Wilkerson et al. (2021); however, Townson (1990) indicated the presence of two syntype females in the British Museum (Natural History) (now the Natural History Museum, London), one from Kasongo, Zaire (present day Democratic Republic of the Congo) and the other one with no locality. As confirmation of the lectotype, Hamon & Ovazza (1956) stated that consimilis was “described from a female from Kasongo, Belgium Congo,” which “differs from the type form by: the absence of basal pale markings on the terga 1–7, the apicolateral pale triangular spots being reduced and not appearing clearly as on segments 6 and 7” (translated from the French). Edwards (1911b) treated consimilis as a species of Culex, distinct from Cx. annulioris, and recognized Cx. pseudoannulioris Theobald, 1909, described from three females captured at Obuasi, Gold Coast (present-day Ghana), as its junior synonym. Theobald recognized pseudoannulioris as “A very pronounced, banded proboscis species, coming near C. annulioris, Theobald ”, differing in having the “Abdomen unbanded, deep blackish brown with lateral creamy scales, which to some extent form apical spots above”. In addition to pseudoannulioris, Edwards (1941) recognized Cx. annulioris var. congolensis Evans, 1923 and Cx. bitaeniorhynchus var. mayumbae Galliard, 1931 as synonyms of subspecies consimilis. The former was described from two males collected at Leopoldville, Belgian Congo (present-day Kinshasa, Democratic Republic of the Congo), characterized as having the “ abdomen entirely dark scaled above [sic], the median basal and lateral apical white markings characteristic of annulioris being entirely absent.” The variety mayumbae was based on features of the phallosome of males reared from larvae collected in Tchibanga and Mouila, Gabon. Two other nominal forms, Cx. annulioris var. gambiensis Theobald, 1903b and Cx. annulioris var. major Edwards, 1935, are presently considered to be junior synonyms. The former, listed as a synonym of the type form by Edwards (1932a), was described from a single female reared from a larva collected at [Bathurst, renamed Banjul in 1973], Gambia as having the abdomen “like the type, but the triangular basal white spots are very indistinct, but can be detected on each segment by a few white scales”. The variety major, subsequently raised to subspecific rank by Edwards (1941) and synonymized with subspecies consimilis by Hamon & Ovazza (1956), was described from a series of specimens, including the type male, from Nairobi, Kenya, which differed from the type form in being larger and having “abdominal tergites [terga] with distinct basal pale bands which are somewhat widened in the middle; apical lateral pale spots very small or even absent.” Although Edwards (1935) mentioned that the male genitalia of major were not obviously different from those of the typical form, he later (Edwards 1941) noted that the phallosome differed “slightly from both the typical form and from ssp. consimilis.” Actually, the lateral plate of major is more than slightly different than those of the type form and subspecies consimilis (see below). In addition to the specimens from Nairobi, Edwards also identified specimens of major from localities in the Democratic Republic of the Congo (as the Belgian Congo) and Uganda. Based on the examination of type material and specimens from Benin (as Dahomey), Cameroon, Côte d’Ivoire, Ethiopia, Kenya, Republic of Upper Volta (as Haute-Volta) and Senegal, Hamon & Ovazza (1956) concluded “that [in] Culex annulioris there are very many variations in colouring which do not correspond at all to variations in the structure of the phallosome, the same structure of the phallosome being able according to the regions to correspond to very dark forms or to very pale forms. The systematic value of the types of C. annulioris and C. annulioris consimilis is therefore almost zero since they are females. For the sake of simplicity, we propose to continue to call C. annulioris sensu stricto the specimens with smooth phallosome [i.e. lateral plate, see below] and C. annulioris var. consimilis those with phallosome bearing numerous and long spicules. Although an almost pure population is generally observed at a given point, there are some forms of passage between the two types of phallosome; given the variations recorded here both in the exterior colouring and in the structure of the phallosome, it does not seem appropriate to continue to regard C. annulioris major as a different variety from C. annulioris consimilis and we propose to treat it as a synonym [translated from the French].” Cutting to the chase, we find that the conclusions reached by Hamon & Ovazza (1956) are not convincing because they are based on presumed probability that features of the lateral plates of the male genitalia are not correlated with observed differences in the ornamentation of the adult females, principally the absence or presence and development of pale scaling on the abdominal terga. For the most part, neither the quantity nor quality of material available for study was adequate for resolving the taxonomy of the nominal forms. The study of Hamon & Ovazza was based entirely on adult mosquitoes, and characteristics of the immature stages were not considered. Indeed, the larval and pupal stages for all of the nominal forms are unknown, except the type form, which has not been definitely associated with topotypic material. As communicated by Macfie & Ingram (1923), “The larva of this mosquito [Cx. annulioris] has been figured by Edwards [1912d] … and included in his key to the larvae of African CULICIDAE; it is therefore unnecessary for us to do more than mention a few additional characters.” As usual, Edwards only illustrated the head (dorsal view) and abdominal segments VIII and X (lateral view). Macfie & Ingram also provided a description of the pupa, based on a single exuviae, and an illustration of the trumpet. The descriptions were based on specimens found in pools at Accra, Ghana. The authors did not indicate how the specimens were identified to species, but they were presumably associated with reared adults that were identifiable as the type form of Cx. annulioris. Hopkins (1936, 1952) also described and illustrated the head and terminal abdominal segments of the annulioris larva, with notable differences from the illustrations of Edwards (1912d): Antenna slightly curved and slightly more slender distal to seta 1A, with more numerous branches; comb with fewer large spine-like scales; siphon shorter (index about 6.0 as opposed to about 8.4); pecten short, on approximately the basal 0.06 of the siphon (on approximately the basal 0.15 of the siphon illustrated by Edwards). With regard to the male genitalia, it is disappointing that attention has only been given to the structure of the lateral plate of the phallosome, which, in the case of the nominal taxa considered here, has been referred to inaccurately as the “hypopygium” (Galliard 1931; Edwards 1935), “mesosome” (Galliard 1931) and “phallosome” (Edwards 1941; Hamon & Ovazza 1956). Jupp (1996) correctly referred to it, for the most part, as the “lateral plate of phallosome”. More specifically, distinctions between the nominal forms for which the male genitalia are known (i.e. annulioris, consimilis and major) have focused on the curved ventrocaudal surface of the inner division of the lateral plate (the “posterior margin of the inner division” of Edwards 1941). In the type form, that surface of the inner division bears numerous very minute spicules (characterized as “smooth” by Hamon & Ovazza 1956) whereas in consimilis and major it bears many semi-recumbent thorn-like projections. However, the lateral plate is a complicated structure that exhibits differences in the shape and development of the dorsocaudal angle and its dorsally projecting recurved teeth, differences which seem to have been overlooked but are readily apparent in the illustrations provided by Galliard (1931), Edwards (1941), Hamon & Ovazza (1956) and Jupp (1996). Although the lateral plates of consimilis and major both have thorn-like projections on the ventrocaudal surface, the contour and development of the surface between the projections and the dorsally projecting teeth is very different, as shown in figures 101f (major) and 101g (consimilis) of Edwards (1941). As illustrated by Edwards, and also Hamon & Ovazza (1956), in consimilis the region of the plate bearing the thorn-like projects is more or less evenly rounded and a distinct group of small dorsally directed denticles is present at the base of the recurved teeth whereas in major the region bearing the thorn-like processes is more angular and there are no denticles at the base of the recurved teeth. Surprisingly, none of the illustrations of the lateral plates which Hamon & Ovazza illustrated as those of consimilis are similar to the lateral plate of major illustrated by Edwards. We anticipate that other differences in structures of the male genitalia, e.g. the development of the subapical lobe of the gonocoxite and its specialized setae, will be found that further indicate the existence of a number of closely related species. In addition to overlooking information about larvae and pupae, Hamon & Ovazza (1956) also failed to consider the descriptions and illustrations of structures of the female genitalia of annulioris and consimilis (as separate species) provided by Macfie & Ingram (1922). Differences were noted as follows: In annulioris, the postgenital lobe (their tenth segment) bears four or five small setae on each side of the ventral surface and the posterior margin is emarginate; the spermathecal capsules are “sub-equal, oval; length 99 μ, breadth 68 μ, the chitinised portion of the ducts very short, about 2 μ.” In contrast, the postgenital lobe of consimilis “bears about nine small setae on each side on its ventral aspect” and the posterior margin is not emarginate; the spermathecal capsules are larger, “the middle one measured 137 μ in length by 84 μ in breadth, and the chitinised portion of its duct was about 7 μ long, and in the other two spermathecae the corresponding measurements were 129 μ, 80 μ, and 4μ respectively.” The authors did not mention how the specimens were identified to species or where they were collected, but the observed differences clearly indicate potential specific distinctions. A look at the collection records for the type form and subspecies consimilis and major reveals that the three forms occur in sympatry throughout central (sub-Saharan) and eastern Africa. It is interesting to note that the type localities of all nominal forms currently regarded as junior synonyms are in countries of central and western Africa, far north of the type locality of annulioris sensu stricto in Zimbabwe. It is particularly interesting to note that the type locality of gambiensis, the sole synonym of the type form, is located more than 6,500 km (by land) from Zimbabwe. Considering the ecological and topographical differences over the very wide distribution of Cx. annulioris sensu lato, it is surprising that Hamon & Ovazza found no correlation of the marked variation in the coloration of the adults with geographical distribution. This is very puzzling because such variation is often suggestive of a species complex. Whereas Hamon & Ovazza (1956) interpreted Cx. annulioris as an extremely variable species, we believe that the available morphological and distributional data provide evidence of a species complex. In the absence of a comprehensive study of topotypic material, the nominal forms remain poorly and inadequately known, particularly because the immature stages have never been used to help define them—the complete larval and pupal chaetotaxy has not been studied even for the two currently recognized subspecies. As in the case of several species of the Vishnui Subgroup of the subgenus Culex, whose lateral plates are very similar to those of Cx. annulioris and which exhibit marked similarities in the adult stage (see Colless 1957 and Sirivanakarn 1976), we anticipate that larvae of reared specimens collected in the type localities are likely to show that annulioris, consimilis and major are clearly differentiated species. In addition to morphology, bionomical and molecular data are needed to resolve the composition of the Annulioris Complex. For the time being, we believe it is prudent to recognize the following nominal taxa as separate species of the complex: Culex (Oculeomyia) annulioris Theobald, 1901a , Culex (Oculeomyia) consimilis Newstead, 1907 and Culex (Oculeomyia) major Edwards, 1935 . Culex consimilis and Cx. major need to be added to the list of Culex species recognized in the Encyclopedia of Life. The following nominal forms are provisionally retained as junior synonyms: Cx. annulioris var. gambiensis Theobald, 1903b (synonym of Cx. annulioris Theobald, 1901a); Cx. pseudoannulioris Theobald, 1909, Cx. annulioris var. congolensis Evans, 1923 and Cx. bitaeniorhynchus var. mayumbae Galliard, 1931 (synonyms of Cx. consimilis Newstead, 1907)., Published as part of Harbach, Ralph E. & Wilkerson, Richard C., 2023, The insupportable validity of mosquito subspecies (Diptera: Culicidae) and their exclusion from culicid classification, pp. 1-184 in Zootaxa 5303 (1) on pages 103-105, DOI: 10.11646/zootaxa.5303.1.1, http://zenodo.org/record/8043342, {"references":["Theobald, F. V. (1901 a) A monograph of the Culicidae or mosquitoes. Vol. 1. British Museum (Natural History), London, xviii + 424 pp., pls. viii + I - XXXVII and A - E under separate cover. https: // doi. org / 10.5962 / bhl. title. 58067","Wilkerson, R. C., Linton, Y. - M. & Strickman, D. (2021) Mosquitoes of the world. Vols. 1 & 2. Johns Hopkins University Press, Baltimore, Maryland, 1332 pp. https: // doi. org / 10.1186 / s 13071 - 021 - 04848 - 6","Newstead, R., Dutton, J. E. & Todd, J. L. L (1907) Insects and other Arthropoda collected in the Congo Free State. Annals of Tropical Medicine and Parasitology, 1 (1 - 5), 3 - 113. https: // doi. org / 10.1080 / 00034983.1907.11719252","Edwards, F. W. (1941) Mosquitoes of the Ethiopian Region III. - Culicine adults and pupae. Printed by order of the Trustees, British Museum (Natural History), London, viii + 499 pp.","Lewis, D. J. (1956) The Culex mosquitoes of the Sudan. Bulletin of Entomological Research, 47 (4), 703 - 721. https: // doi. org / 10.1017 / S 0007485300046940","Simsaa, M. A. A., Harbach, R. E., Almalik, A. M. A., Ahmed, E. M., Esia, A. A., Mohamed, A. H. & Azrag, R. S. (2021) Culex mosquitoes (Diptera: Culicidae) recorded along the Nile River in northern Sudan, with a key for the identification of all species of the genus known to occur in the country. Zootaxa, 4963 (3), 401 - 411. https: // doi. org / 10.11646 / zootaxa. 4963.3.1","Stone, A., Knight, K. L. & Starcke, H. (1959) A synoptic catalog of the mosquitoes of the world (Diptera, Culicidae). The Thomas Say Foundation. Vol. VI. Entomological Society of America, College Park, Maryland, vi + 358 pp.","Knight, K. L. & Stone, A. (1977) A catalog of the mosquitoes of the world (Diptera: Culicidae). 2 nd Edition. The Thomas Say Foundation. Vol. VI. Entomological Society of America, College Park, Maryland, xi + 611 pp.","Ovazza, M., Hamon, J. & Neri, P. (1956) Contribution a l'etude des dipteres vulnerants de l'empire d'Ethiopie. I. Culicidae. Bulletin de la Societe de Pathologie exotique, 49 (1), 151 - 182.","Edwards, F. W. (1911 b) The African species of Culex and allied genera. Bulletin of Entomological Research, 2 (3), 241 - 268. https: // doi. org / 10.1017 / S 0007485300001450","Theobald, F. V. (1909) Descriptions of the new mosquitoes collected by Dr. Graham in Ashanti. Gold Coast Miscellaneous Publication No. 237. Great Britain Colonial Office. London, 31 pp., 2 pls.","Evans, A. M. (1923) A new species and a new variety of Culex from the Belgian Congo. Annals of Tropical Medicine and Parasitology, 17 (1), 89 - 91. https: // doi. org / 10.1080 / 00034983.1923.11684352","Galliard, H. (1931) Culicides du Gabon. I. - Culicines, avec la description d'une espece et de deux varietes nouvelles. Annales de Parasitologie Humaine et Comparee, 9 (3), 225 - 232. https: // doi. org / 10.1051 / parasite / 1931093225","Theobald, F. V. (1903 b) Report on a collection of mosquitoes or Culicidae, etc, from Gambia, and descriptions of new species. Memoirs of the Liverpool School of Tropical Medicine, 10 (Appendix), i - xi, 2 pls.","Edwards, F. W. (1935) Mosquito notes. - XII. Bulletin of Entomological Research, 26 (2), 127 - 136. https: // doi. org / 10.1017 / S 0007485300038116","Edwards, F. W. (1932 a) Genera Insectorum. Diptera, Fam. Culicidae. Fascicle 194. V. Verteneuil & L. Desmet, Bruxelles [Brussels], 258 pp., 5 pls.","Macfie, J. W. S. & Ingram, A. (1923) The early stages of West African mosquitos. - VI. Bulletin of Entomological Research, 13 (4), 409 - 442. https: // doi. org / 10.1017 / S 000748530004551 X","Edwards, F. W. (1912 d) Revised keys to the known larvae of African Culicinae. Bulletin of Entomological Research, 3 (4), 373 - 385. https: // doi. org / 10.1017 / S 0007485300002042","Hopkins, G. H. E. (1936) Mosquitoes of the Ethiopian Region I. - Larval bionomics of mosquitoes and taxonomy of culicine larvae. Printed by order of the Trustees, British Museum (Natural History), London, (vi) + 250 pp.","Hopkins, G. H. E. (1952) Mosquitoes of the Ethiopian Region I. - Larval bionomics of mosquitoes and taxonomy of culicine larvae. 2 nd Edition. British Museum (Natural History), London, viii + 355 pp.","Jupp, P. G. (1996) Mosquitoes of southern Africa: Culicinae and Toxorhynchitinae. Ekogilde Publishers, Hartebeespoort, 156 pp.","Macfie, J. W. S. & Ingram, A. (1922) On the genital armature of the female mosquito. Annals of Tropical Medicine and Parasitology, 16 (2), 157 - 188. https: // doi. org / 10.1080 / 00034983.1922.11684308","Colless, D. H. (1957) Notes on the culicine mosquitoes of Singapore II. - The Culex vishnui group (Diptera, Culicidae), with descriptions of two new species. Annals of Tropical Medicine and Parasitology, 51 (1), 87 - 101. https: // doi. org / 10.1080 / 00034983.1957.11685798","Sirivanakarn, S. (1976) Medical entomology studies - III. A revision of the subgenus Culex in the Oriental Region (Diptera: Culicidae). Contributions of the American Entomological Institute, 12 (2), iii, 1 - 272."]}

Subspecies nocturnus Descriptions of subspecies nocturnus are very similar to descriptions of vexans, which has led to many different opinions concerning its status. It was described by Theobald (1903a) from “several ♀ ’s” from “Ba in Fiji ”. Townsend (1990) found two syntype females in the collection of the Natural History Museum, London. Theobald (1903a) described the species as having the “Thorax deep brown, covered densely with narrow-curved, bright brown and golden scales, scarcely showing any definite ornamentation; proboscis with a minute black tip and base, remainder ochraceous.” The legs are as in the description of vexans above. “Abdomen deep brown with basal white bands curved in the middle…. ♀. Head brown, with narrow curved pale golden scales on the crown, with black upright forked scales, flat black scales at the sides with a median white patch…. Abdomen …sixth and seventh segments with narrow apical yellow bands, no basal pale band or mark to the last; laterally are median white spots…. Legs …tibiae deep brown, paler ventrally…. Observations.... The abdominal ornamentation is very characteristic; the species cannot well be mistaken for any other. The proboscis is really very broadly pale banded.” Theobald included descriptions of all life stages except the egg, none of which provided distinguishing characters. The following publications concerning the nominal taxon nocturnus are arranged by taxonomic status. A few authors gave expanded rationale for their judgements (Bohart & Ingram 1946b; Belkin 1962; Reinert 1973; Lee et al. 1982), which were summarized by Lee et al. (1982), as shown below. This is not intended to be an exhaustive listing (* = with some sort of illustration). As a synonym of vexans. Dyar (1920, 1921a); Senior-White (1923); Freeborn (1926); Edwards (1924); Edwards (1932a); Natvig (1948); Senevet & Andarelli (1954) [who followed Edwards 1932a]; Carpenter & LaCasse (1955) [subspecies arabiensis ? and nipponii also listed as synonyms of vexans]; Reinert (1973); Knight & Stone (1977) [who we assume followed Reinert 1973]; Reinert et al. (2004). As a synonym of Aedes (Ochlerotatus) vigilax (Skuse, 1889). Cooling (1924). As a subspecies of vexans. * Bohart & Ingram (1946b); * Yamaguti & LaCasse (1950); Knight & Hull (1951); * Knight & Hull (1953); Laird (1955); Bohart (1957); Stone et al. (1959); Iyengar (1960); Stone (1961b); Lien (1962); * Joyce & Nakagawa (1963); * Holway & Bridges (1970); Basio (1971); * Baisas (1974); Tanaka et al. (1979); Harbach (2018); Wilkerson et al. (2021). As species nocturnus. * Theobald (1903a); Edwards (1912a), in genus Ochlerotatus but “Doubtfully distinct from the preceding [vigilax Skuse, 1889]”; * Penn (1949), as vexans but possibly represents subspecies nocturnus; * Belkin (1962); Peters & Christian (1963), included as both a species and apparently inadvertently as a subspecies of vexans; Stone (1963); Belkin (1965); Stephan (1966); Standfast (1967); Maffi (1977). Lee et al. (1982), as species nocturnus, stated the following in a footnote. Although F.W. Edwards (1924) synonymised nocturnus with vexans it was later considered a variety or subspecies of vexans by regional authors, commencing with Bohart & Ingram (1946). Belkin (1962) provisionally raised nocturnus to specific rank, mainly on the basis of branching of larval head hairs [setae]. He recognised that the taxonomic status of nocturnus was uncertain, clarification awaiting study of the vexans complex. Reinert (1973) sank nocturnus as a synonym of vexans vexans “in the absence of sufficient biological, behavioural and genetical data on the Pacific Island populations of vexans, and since specimens of these populations fall within the variable range of the morphological characters of other populations within the distribution of the species”. For the time being we have chosen to retain nocturnus as a species partly because the synonymising of nocturnus under vexans gives vexans an extraordinarily wide distribution far greater than that achieved by any other non-domestic species. We agree with the summary and conclusions of Lee et al. (1982) and thus hereby return nocturnus to its original species status: Aedes (Aedimorphus) nocturnus (Theobald, 1903a) . Synonyms: Culicada eruthrosops Theobald, 1910; Culex nocturnus var. niger Theobald, 1913b. Aedes nocturnus is currently listed as a species in the Encyclopedia of Life., Published as part of Harbach, Ralph E. & Wilkerson, Richard C., 2023, The insupportable validity of mosquito subspecies (Diptera: Culicidae) and their exclusion from culicid classification, pp. 1-184 in Zootaxa 5303 (1) on pages 19-20, DOI: 10.11646/zootaxa.5303.1.1, http://zenodo.org/record/8043342, {"references":["Theobald, F. V. (1903 a) A monograph of the Culicidae or mosquitoes. Vol. 3. British Museum (Natural History), London, xvii + 1 (errata) + 359 pp., foldout table, 17 pls. https: // doi. org / 10.5962 / bhl. title. 58067","Townsend, B. C. (1990) Culicidae. In: Townsend, B. C., Chainey, J. E., Crosskey, R. W., Pont, A. C., Lane, R. P., Boorman, J. P. T. & Crouch, C. A. (Eds.), A catalogue of the types of bloodsucking flies in the British Museum (Natural History), Natural History Museum, London, pp. 35 - 152.","Bohart, R. M. & Ingram, R. L. (1946 b) Mosquitoes of Okinawa and islands in the central Pacific. Navmed 1055. Bureau of Medicine and Surgery, Navy Department, Washington, D. C., 110 pp.","Belkin, J. N. (1962) The mosquitoes of the South Pacific (Diptera, Culicidae). Vols 1 & 2. University of California Press, Berkeley and Los Angeles, California, xii + 608 pp., 412 figs.","Reinert, J. F. (1973) Contributions to the mosquito fauna of Southeast Asia. - XVI. Genus Aedes Meigen, subgenus Aedimorphus Theobald in Southeast Asia. Contributions of the American Entomological Institute, 9 (5), 1 - 218.","Lee, D. J., Hicks, M. M., Griffiths, M., Russell, R. C. & Marks, E. N. (1982) The Culicidae of the Australasian Region. Vol. 2. Nomenclature, synonymy, literature, distribution, biology and relation to disease. Genus Aedeomyia. Genus Aedes (subgenera [Aedes], Aedimorphus, Chaetocruiomyia, Christophersiomyia, Edwardsaedes and Finlaya). Monograph Series, Entomology Monograph No. 2. Australian Government Publishing Service, Canberra, v + 286 pp.","Dyar, H. G. (1920) The Aedes of the mountains of California and Oregon (Diptera, Culicidae). Insecutor Inscitiae Menstruus, 8 (10 - 12), 165 - 173.","Dyar, H. G. (1921 a) The mosquitoes of Canada. Transactions of the Royal Canadian Institute, 13 (1), 71 - 120.","Senior-White, R. (1923) Catalogue of Indian insects. Part 2 - Culicidae. Superintendent Government Printing, Calcutta, ii + 124 pp.","Freeborn, S. B. (1926) The mosquitoes of California. University of California Publications: Technical Bulletins. Entomology, 3 (5), 333 - 460.","Edwards, F. W. (1924) A synopsis of the adult mosquitos of the Australasian Region. Bulletin of Entomological Research, 14 (4), 351 - 401. https: // doi. org / 10.1017 / S 0007485300045624","Edwards, F. W. (1932 a) Genera Insectorum. Diptera, Fam. Culicidae. Fascicle 194. V. Verteneuil & L. Desmet, Bruxelles [Brussels], 258 pp., 5 pls.","Natvig, L. R. (1948) Contributions to the knowledge of the Danish and Fennoscandian mosquitoes: Culicini. Norsk Entomologisk Tidsskrift Supplement, 1, i - xxi + 567 pp., 12 pls., map.","Senevet, G. & Andarelli, L. (1954) Le genre Aedes en Afrique du Nord. I. - Les larves. Archives de l'Institut Pasteur d'Algerie, 32 (4), 310 - 351.","Carpenter, S. J. & LaCasse, W. J. (1955) Mosquitoes of North America (north of Mexico). University of California Press, Berkeley, Los Angeles, London, vii + 360 pp., 127 pls.","Knight, K. L. & Stone, A. (1977) A catalog of the mosquitoes of the world (Diptera: Culicidae). 2 nd Edition. The Thomas Say Foundation. Vol. VI. Entomological Society of America, College Park, Maryland, xi + 611 pp.","Reinert, J. F., Harbach, R. E. & Kitching, I. J. (2004) Phylogeny and classification of Aedini (Diptera: Culicidae) based on morphological characters of all life stages. Zoological Journal of the Linnean Society, 142 (3), 289 - 368. https: // doi. org / 10.1111 / j. 1096 - 3642.2004.00144. x","Skuse, F. A. A. (1889) Diptera of Australia. Part V. The Culicidae. Proceedings of the Linnean Society of New South Wales, 3 (4), 1717 - 1764, 1 pl.","Cooling, L. E. (1924) A synonymic list of the more important species of Culicidae of the Australian Region. Commonwealth of Australia, Department of Health. Service Publication (Tropical Division), 2, 1 - 61.","Yamaguti, S. & LaCasse, W. J. (1950) Mosquito fauna of Guam. Office of the Surgeon, Headquarters, 8 th Army, APO 343. United States. 207 th Malaria Survey Detachment. Office of the Surgeon, Washington, D. C., 101 pp.","Knight, K. L. & Hull, W. B. (1951) The Aedes mosquitoes of the Philippine Islands. I. Keys to species. Subgenera Mucidus, Ochlerotatus, and Finlaya (Diptera, Culicidae). Pacific Science, 5 (3), 211 - 251.","Knight, K. L. & Hull, W. B. (1953) The Aedes mosquitoes of the Philippine Islands. III. Subgenera Aedimorphus, Banksinella, Aedes, and Cancraedes (Diptera, Culicidae). Pacific Science, 7 (4), 453 - 481.","Laird, M. (1955) Notes on the mosquitos of the Gilbert, Ellice and Tokelau Islands, and on filariasis in the latter group. Bulletin of Entomological Research, 46 (2), 291 - 300. https: // doi. org / 10.1017 / S 0007485300030923","Bohart, R. M. (1957) Insects of Micronesia, Diptera: Culicidae. Insects of Micronesia, 12 (1), 1 - 85. [for 1956]","Stone, A., Knight, K. L. & Starcke, H. (1959) A synoptic catalog of the mosquitoes of the world (Diptera, Culicidae). The Thomas Say Foundation. Vol. VI. Entomological Society of America, College Park, Maryland, vi + 358 pp.","Iyengar, M. O. T. (1960) A review of the mosquito fauna of the South Pacific (Diptera: Culicidae). South Pacific Commission Technical Paper, 130, (ii) + 1 - 102.","Stone, A. (1961 b) A synoptic catalog of the mosquitoes of the world, supplement I (Diptera: Culicidae). Proceedings of the Entomological Society of Washington, 63 (1), 29 - 52.","Lien, J. C. (1962) Non-anopheline mosquitoes of Taiwan: Annotated catalog and bibliography. Pacific Insects, 4 (3), 615 - 649.","Joyce, C. R. & Nakagawa, P. Y. (1963) Aedes vexans nocturnus (Theobald) in Hawaii. Proceedings, Hawaiian Entomological Society, 18 (2), 273 - 280.","Holway, R. T. & Bridges, J. R. (1970) Illustrated key to the adult mosquitoes of the Marianas. U. S. Navy, Preventative Medicine, Unit- 6, 1 - 6.","Basio, R. G. (1971) The mosquito fauna of the Philippines (Diptera Culicidae). Monograph No. 4. National Museum of the Philippines, Manila, v + 198 pp.","Baisas, F. E. (1974) The mosquito fauna of Subic Bay Naval Reservation. Republic of the Philippines. Headquarters, First Medical Service Wing (PACAF), APO San Francisco 96274, Technical Report No. 72 - 2, 1 - 179, 79 figs.","Tanaka, K., Mizusawa, K. & Saugstad, E. S. (1979) A revision of the adult and larval mosquitoes of Japan (including the Ryukyu Archipelago and the Ogasawara Islands) and Korea (Diptera: Culicidae). Contributions of the American Entomological Institute, 16, vii + 1 - 987.","Harbach, R. E. (2018) Culicipedia: Species-group, genus-group and family-group names in Culicidae (Diptera). CABI, Wallingford, Oxfordshire, xviii + 378 pp. https: // doi. org / 10.1079 / 9781786399052.0000","Wilkerson, R. C., Linton, Y. - M. & Strickman, D. (2021) Mosquitoes of the world. Vols. 1 & 2. Johns Hopkins University Press, Baltimore, Maryland, 1332 pp. https: // doi. org / 10.1186 / s 13071 - 021 - 04848 - 6","Edwards, F. W. (1912 a) A key to the Australasian species of Ochlerotatus (Culicidae). Annals and Magazine of Natural History, Series 8, 9 (53), 521 - 527. https: // doi. org / 10.1080 / 00222931208693166","Penn, G. H. (1949) The pupae of the mosquitoes of New Guinea. Pacific Science, 3 (1), 3 - 85.","Peters, W. & Christian, S. H. (1963) The bionomics, ecology and distribution of some mosquitoes (Diptera: Culicidae) in the territory of Papua and New Guinea. Acta Tropica, 20 (1), 35 - 79. https: // doi. org / 10.5169 / seals- 311101","Stone, A. (1963) A synoptic catalog of the mosquitoes of the world, supplement II (Diptera: Culicidae). Proceedings of the Entomological Society of Washington, 65 (2), 117 - 140.","Standfast, H. A. (1967) Biting times of nine species of New Guinea Culicidae (Diptera). Journal of Medical Entomology, 4 (2), 192 - 196. https: // doi. org / 10.1093 / jmedent / 4.2.192","Maffi, M. (1977) Contribution to the knowledge of the mosquito fauna of the New Hebrides island group proper (Diptera: Culicidae). Rivista di Parassitologia, 38 (2 / 3), 194 - 214, table 2.","Theobald, F. V. (1910) A monograph of the Culicidae or mosquitoes. Vol. 5. British Museum (Natural History), London, xiv + (1) + 646 pp., 6 pls. https: // doi. org / 10.5962 / bhl. title. 58067","Theobald, F. V. (1913 b) Culicidae from New Caldeonia and the Loyalty Islands. In: Sarasin, F. & Roux, J., Nova Caledonia. Forschungen in Neu-Caledonien und auf den Loyalty-Inseln. Recherches scientifiques en Nouvelle-Caledonie et aux iles Loyalty. A. Zoologie. Vol. I. L. III. C. W. Kreidels Verlag, Wiesbaden, pp. 163 - 164."]}

Culex (Culex) pipiens Linnaeus subspecies pallens Coquillett, 1898 —original combination: Culex pallens (subspecific status by Tanaka 2004). Distribution: China, Japan, Mexico, South Korea, United States (continental) (Wilkerson et al. 2021). subspecies pipiens Linnaeus, 1758 —original combination: Culex pipiens. Distribution: Temperate regions of Africa, Asia, Australia, Europe and North and South America (for specific country records, see Wilkerson et al. 2021). Without question, the taxonomic status of Cx. pipiens and its allied forms has received more attention than any other culicine taxon. As pointed out in a review of the taxonomic history of the species (Harbach 2012), the essence of decades of work conducted on Cx. pipiens was to determine whether it is a single polytypic species or a species complex. As a starting point, Edwards (1932a) listed Cx. pipiens and its sister species Cx. quinquefasciatus Say, 1823 (as Cx. fatigans Wiedemann, 1828) as separate species, and Cx. pallens as a variety of the former. In a review of the systematics of these nominal species, and taking into consideration evidence of hybridization in areas where the distributions of Cx. pipiens and Cx. quinquefasciatus overlap, Mattingly et al. (1951) and Mattingly (1967) concluded that they should be considered members of a single polymorphic species comprised of subspecies pipiens and quinquefasciatus and several varieties, with pallens being one of them. Beginning with Belkin (1962), the recognition of Cx. quinquefasciatus as a separate species gradually gained acceptance, resulting in the current recognition of Cx. pipiens and Cx. quinquefasciatus as closely related sister species and pallens as a subspecies of the former species (Smith & Fonseca 2004; Harbach 2012; Fonseca et al. 2009; Aardema et al. 2020; Wilkerson et al. 2021). Culex pipiens pallens has been regarded as a subspecies since the studies of Japanese mosquitoes by Tanaka et al. (1979) and Tanaka (2004). Since then, genetic and molecular studies have shed light on the status of pallens in eastern Asia. A multilocus genotype analysis conducted by Fonseca et al. (2009) revealed the occurrence of hybridization between Cx. p. pallens and Cx. quinquefasciatus in China, South Korea and southern Japan, but not in northern Japan. However, Ohashi et al. (2014) found that Cx. p. pallens and Cx. pipiens hybridize in northern Japan, but hybrids have lower fitness than the offspring of either parental taxon. Finally, recent detailed analyses of DNA sequence data conducted by Aardema et al. (2020), which included sequence for specimens of pallens from China, cogently support the hypothesis that pallens is a genetically distinct entity derived from ancestral hybridization between Cx. pipiens and Cx. quinquefasciatus. The authors note, however, that the “hypothesis warrants further examination.” The presence of pallens in Mexico and the United States must also be assessed. As indicated by Mattingly et al. (1951), some authors have conjectured that the pallens form was introduced into California, presumably from Japan. Despite the concerns and implications of hybridization, we feel it is prudent at this time to accept the evidence for the genetic distinction of pallens provided by Aardema et al. (2020) and hereby re-establish its original species status pending further investigation: Culex (Culex) pallens Coquillett, 1898 . It is interesting to note that Cx. pipiens and Cx. quinquefasciatus are listed as species in the Encyclopedia of Life, but Cx. pallens is not included; thus, it needs to be included with the Culex species listed therein. Oddly, the nomen dubium Culex molestus Kollar, 1832 (in Pohl & Kollar 1832) is listed as a species, probably mistakenly included instead of Cx. molestus Forskål, 1775, a recognized synonym and physiological variant (molestus form) of Cx. pipiens. The list of mosquitoes in the Encyclopedia of Life was not compiled by mosquito taxonomic experts and does not, in many cases, reflect the current taxonomy of various taxa. Thirty-six nominal taxa (disregarding the name melanorhinus Giles, 1900, which was proposed as a replacement name for Cx. pallipes Macquart, 1838) are currently recognized as synonyms of Cx. pipiens. We consider all synonymous taxa named from localities in Europe (England, France, Germany and Portugal), northern Africa (Algeria, Egypt and Tunisia), Mexico and the United States to be synonyms of Cx. pipiens. These include all of the synonyms listed by Harbach (2018) and Wilkerson et al. (2021), as well as Cx. comitatus Dyar & Knab, 1909a (California) and Cx. quinquefasciatus race dipseticus Dyar & Knab, 1909a (western Mexico and California), which were listed as synonyms of Cx. pipiens pallens Coquillett, 1898. Only a single nominal species, Cx. osakaensis Theobald, 1907 (type locality: Osaka, Honshu Island, Japan), is retained as a synonym of Cx. pallens., Published as part of Harbach, Ralph E. & Wilkerson, Richard C., 2023, The insupportable validity of mosquito subspecies (Diptera: Culicidae) and their exclusion from culicid classification, pp. 1-184 in Zootaxa 5303 (1) on page 88, DOI: 10.11646/zootaxa.5303.1.1, http://zenodo.org/record/8043342, {"references":["Coquillett, D. W. (1898) Report on a collection of Japanese Diptera, presented to the U. S. National Museum by the Imperial University of Tokyo. Proceedings of the United Stated National Museum, 21 (1146), 301 - 340. https: // doi. org / 10.5479 / si. 00963801.21 - 1146.301","Tanaka, K. (2004) Studies on the pupal mosquitoes of Japan (10) Culex (Culex) (Diptera, Culicidae). Japanese Journal of Systematics Entomology, 10 (1), 9 - 42.","Wilkerson, R. C., Linton, Y. - M. & Strickman, D. (2021) Mosquitoes of the world. Vols. 1 & 2. Johns Hopkins University Press, Baltimore, Maryland, 1332 pp. https: // doi. org / 10.1186 / s 13071 - 021 - 04848 - 6","Linnaeus, C. (1758) Systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus 1. Editio Decima. Impensis Direct. Laurentii Salvii, Holmiae, 824 pp. https: // doi. org / 10.5962 / bhl. title. 542","Harbach, R. E. (2012) Culex pipiens Linnaeus: Species versus species complex - taxonomic history and perspective. Journal of the American Mosquito Control Association, 28 (4 s), 10 - 23. https: // doi. org / 10.2987 / 8756 - 971 X- 28.4.10","Edwards, F. W. (1932 a) Genera Insectorum. Diptera, Fam. Culicidae. Fascicle 194. V. Verteneuil & L. Desmet, Bruxelles [Brussels], 258 pp., 5 pls.","Say, T. (1823) Descriptions of dipterous insects of the United States. Journal of the Academy of Natural Sciences of Philadelphia, 3 (1), 9 - 54.","Wiedemann, C. R. W. (1828) Aussereuropaische zweifl ¸ gelige Insekten: als Fortsetzung des Meigenschen Werkes. Vol. 1. in der Schulzischen Buchhandlung, Hamm, xxxii + 608 pp. https: // doi. org / 10.5962 / bhl. title. 14603","Mattingly, P. F., Rozeboom, L. E., Knight, K. L., Laven, H., Drummond, F. H., Christophers, S. R. & Shute, P. G. (1951) The Culex pipiens complex. Transaction of the Royal Entomological Society of London, 102 (7), 331 - 382. https: // doi. org / 10.1111 / j. 1365 - 2311.1951. tb 00752. x","Mattingly, P. F. (1967) The systematics of the Culex pipiens complex. Bulletin of the World Health Organization, 37, 257 - 261.","Belkin, J. N. (1962) The mosquitoes of the South Pacific (Diptera, Culicidae). Vols 1 & 2. University of California Press, Berkeley and Los Angeles, California, xii + 608 pp., 412 figs.","Smith, J. L., Fonseca, D. M. (2004) Rapid assays for identification of members of the Culex (Culex) pipiens complex, their hybrids, and other sibling species (Diptera: Culicidae). American Journal of Tropical Medicine and Hygiene, 70 (4), 339 - 345. https: // doi. org / 10.4269 / ajtmh. 2004.70.339","Fonseca, D. M., Smith, J. L., Kim, H. - C. & Mogi, M. (2009) Population genetics of the mosquito Culex pipiens pallens reveals sex-linked asymmetric introgression by Culex quinquefasciatus. Infection, Genetics and Evolution, 9 (6), 1197 - 1203. https: // doi. org / 10.1016 / j. meegid. 2009.06.023","Aardema, M. L., vonHoldt, B. M., Fritz, M. L. & Davis, S. R. (2020) Global evaluation of taxonomic relationships and admixture within the Culex pipiens complex of mosquitoes. Parasites & Vectors, 13, 8. https: // doi. org / 10.1186 / s 13071 - 020 - 3879 - 8","Tanaka, K., Mizusawa, K. & Saugstad, E. S. (1979) A revision of the adult and larval mosquitoes of Japan (including the Ryukyu Archipelago and the Ogasawara Islands) and Korea (Diptera: Culicidae). Contributions of the American Entomological Institute, 16, vii + 1 - 987.","Ohashi, K., Tsuda, Y., Kasai, S., Kawada, H. & Takagi, M. (2014) Hybridization between sympatric populations of Culex pipiens pallens and Culex pipiens f. molestus (Diptera: Culicidae) in Nagasaki, Japan. Medical Entomology and Zoology, 65 (2), 67 - 72. https: // doi. org / 10.7601 / mez. 65.67","Pohl, J. E. & Kollar, V. (1832) Brasiliens vorz ¸ glich lastige Insecten. Besonderer Abdruck aus der Reise im Innern von Brasilien von Dr. Pohl [extracted from Pohl's \" Reise im Innern von Brasilien \"], Wien, 1832, 1 - 20, 1 pl.","Forskal, P. (1775) Descriptiones animalium, avium, amphibiorum, piscium, insectorum, vermium; quae in itinere orientali observavit Petrus Forskal. Post mortem auctoris edidit Carsten Niebuhr. Adjuncta est materia medica kahirina atque tabula maris Rubri geographica. ex officina M ˆ lleri, Hauniae, 164 pp. https: // doi. org / 10.5962 / bhl. title. 2154","Giles, G. M. (1900) A handbook of the gnats or mosquitoes, giving the anatomy and life history of the Culicidae. John Bale, Sons & Danielsson, Ltd, London, viii + 374 pp., 8 pls. https: // doi. org / 10.5962 / bhl. title. 101828","Macquart, J. (1838) Dipteres exotiques nouveaux ou peu connus. Memoires de la Societe Royale des Sciences, de l'Agriculture et des Arts, de Lille, 2, 9 - 225, 25 pls. https: // doi. org / 10.5962 / bhl. title. 15792","Harbach, R. E. (2018) Culicipedia: Species-group, genus-group and family-group names in Culicidae (Diptera). CABI, Wallingford, Oxfordshire, xviii + 378 pp. https: // doi. org / 10.1079 / 9781786399052.0000","Dyar, H. G. & Knab, F. (1909 a) On the identity of Culex pipiens Linnaeus. Proceedings of the Entomological Society of Washington, 11 (1), 30 - 39, 3 pls.","Theobald, F. V. (1907) A monograph of the Culicidae or mosquitoes. Vol. 4. British Museum (Natural History), London, xix + 639 pp., 16 pls. https: // doi. org / 10.5962 / bhl. title. 58067"]}

Culex (Culex) andersoni Edwards subspecies abyssinicus Edwards, 1941 —original combination: Culex (Culex) andersoni ssp. abyssinicus. Distribution: Ethiopia (Edwards 1941). subspecies andersoni Edwards, 1914 —original combination: Culex andersoni. Distribution: Comoros, Democratic Republic of the Congo, Eritrea, Kenya, Malawi, Republic of South Africa, Rwanda, Tanzania, Uganda (Wilkerson et al. 2021). subspecies bwambanus Edwards, 1941 —original combination: Culex (Culex) andersoni ssp. bwambanus. Distribution: Lesotho, Republic of South Africa, Uganda (Wilkerson et al. 2021). Edwards (1914) described Culex andersoni from specimens collected at Kabete, located outside the borders of Nairobi in the Central Province of Kenya, based principally on features of the male habitus and male genitalia. Subspecies abyssinicus is only known from Ethiopia, and according to Edwards (1941) the adult is ornamented differently than the typical form but the male genitalia are essentially the same. Because the treatment of the genitalia is rather superficial, it is likely that some differences might be evident upon closer examination. Based on differences in ornamentation and allopatry, abyssinicus is hereby considered a separate species: Culex (Culex) abyssinicus Edwards, 1941 . This species should be added to the species of Culex listed in the Encyclopedia of Life. Subspecies bwambanus and the nominotypical form are both reported from Uganda, and Edward (1941) indicated that they both occur in the Ruwenzori Mountains. However, Edwards described bwambanus from a Ruwenzori location while recognizing the presence of the typical form elsewhere in the same mountains. Although the Ruwenzori Mountains appear to be the only place where the two forms may be in sympatry, Edwards described bwambanus as a new subspecies because of differences in the pale scaling of the hindfemur of the adults, and nearly “hairless” maxillary palpi (“may be rubbed in the type ”), gonocoxite with a more prominent subapical lobe and a narrower gonostylus in the male. Since the available evidence indicates that bwambanus is morphological distinct in the absence of clinal variation, it is hereby afforded species status: Culex (Culex) bwambanus Edwards, 1941 . Culex bwambanus is currently listed as a species in the Encyclopedia of Life., Published as part of Harbach, Ralph E. & Wilkerson, Richard C., 2023, The insupportable validity of mosquito subspecies (Diptera: Culicidae) and their exclusion from culicid classification, pp. 1-184 in Zootaxa 5303 (1) on page 82, DOI: 10.11646/zootaxa.5303.1.1, http://zenodo.org/record/8043342, {"references":["Edwards, F. W. (1941) Mosquitoes of the Ethiopian Region III. - Culicine adults and pupae. Printed by order of the Trustees, British Museum (Natural History), London, viii + 499 pp.","Edwards, F. W. (1914) New species of Culicidae in the British Museum, with notes on the genitalia of some African Culex. Bulletin of Entomological Research, 5 (1), 63 - 81. https: // doi. org / 10.1017 / S 0007485300034568","Wilkerson, R. C., Linton, Y. - M. & Strickman, D. (2021) Mosquitoes of the world. Vols. 1 & 2. Johns Hopkins University Press, Baltimore, Maryland, 1332 pp. https: // doi. org / 10.1186 / s 13071 - 021 - 04848 - 6"]}

Culex (Culex) guiarti Blanchard subspecies guiarti Blanchard, 1905 —original combination: Culex guiarti [nomen novum for Culex viridis Theobald, 1903a]. Distribution: Benin, Burkina Faso, Cameroon, Central African Republic, Côte d’Ivoire, Democratic Republic of the Congo, Gabon, Gambia, Ghana, Kenya, Liberia, Madagascar, Mali, Mozambique, Nigeria, Republic of the Congo, Republic of South Africa, Senegal, South Sudan, Tanzania, Uganda (Wilkerson et al. 2021, but not Sudan, see Simsaa et al. 2021). subspecies sudanicus Edwards, 1941 —original combination: Culex guiarti var. sudanicus (subspecific status by Harbach & Howard 2007). Distribution: Ghana, Nigeria (Wilkerson et al. 2021). Culex guiarti sudanicus was originally described and named as a variety that was considered to be morphologically “Intermediate between C. guiarti and C. weschei ” (Edwards 1941). It was recognized as a subspecies of Cx. guiarti by Harbach & Howard (2007). Edwards (1941) noted that sudanicus resembled the nominotypical form in having the proboscis entirely dark-scaled, the abdominal sterna with dark apical bands and the distal half of the midfemur extensively dark-scaled dorsally in posterior view, but exhibited the following differences: Scutal integument uniformly brownish, not paler on the fossae (shoulders), scales nearly entirely pale and coarser; mesokatepisternal scales more numerous, the upper and lower patches almost contiguous; proepisternal scales and setae also more numerous; anterior scales of forecoxa all pale; male genitalia as in the typical form, gonostylus not unusually broad (moderately broad and rather suddenly narrowed apically in the typical form). The larva is known for the typical form (Hopkins 1952), but it is not known for sudanicus. The typical form has been recorded from many countries in sub-Sharan Africa whereas sudanicus is only recorded from Ghana and Nigeria (Wilkerson et al. 2021). Based on the paucity of morphological data and the fact that the typical form has also been recorded from Ghana and Nigeria, we believe that Edwards (1941) was correct in describing sudanicus as a morphological variant of Cx. guiarti. Consequently, it seems prudent to formally recognize sudanicus as a synonymous name: sudanicus Edwards, 1941, junior subjective synonym of Culex (Culex) guiarti Blanchard, 1905 . “ Culex sudanicus ” should be removed from the list of Culex species included in the Encyclopedia of Life. We retain a second synonym of Culex guiarti, Culex viridis Theobald, 1903a, described from Buse, Uganda., Published as part of Harbach, Ralph E. & Wilkerson, Richard C., 2023, The insupportable validity of mosquito subspecies (Diptera: Culicidae) and their exclusion from culicid classification, pp. 1-184 in Zootaxa 5303 (1) on pages 85-86, DOI: 10.11646/zootaxa.5303.1.1, http://zenodo.org/record/8043342, {"references":["Blanchard, R. (1905) Les moustiques, histoire naturelle et medicale. F. R. de Rudeval, Imprimeur-Editeur, Paris, xiii + 673 pp. https: // doi. org / 10.5962 / bhl. title. 100899","Theobald, F. V. (1903 a) A monograph of the Culicidae or mosquitoes. Vol. 3. British Museum (Natural History), London, xvii + 1 (errata) + 359 pp., foldout table, 17 pls. https: // doi. org / 10.5962 / bhl. title. 58067","Wilkerson, R. C., Linton, Y. - M. & Strickman, D. (2021) Mosquitoes of the world. Vols. 1 & 2. Johns Hopkins University Press, Baltimore, Maryland, 1332 pp. https: // doi. org / 10.1186 / s 13071 - 021 - 04848 - 6","Simsaa, M. A. A., Harbach, R. E., Almalik, A. M. A., Ahmed, E. M., Esia, A. A., Mohamed, A. H. & Azrag, R. S. (2021) Culex mosquitoes (Diptera: Culicidae) recorded along the Nile River in northern Sudan, with a key for the identification of all species of the genus known to occur in the country. Zootaxa, 4963 (3), 401 - 411. https: // doi. org / 10.11646 / zootaxa. 4963.3.1","Edwards, F. W. (1941) Mosquitoes of the Ethiopian Region III. - Culicine adults and pupae. Printed by order of the Trustees, British Museum (Natural History), London, viii + 499 pp.","Harbach, R. E. & Howard, T. M. (2007) Corrections in the status and rank of names used to denote varietal forms of mosquitoes (Diptera: Culicidae). Zootaxa, 1542 (1), 35 - 48. https: // doi. org / 10.11646 / zootaxa. 1542.1.3","Hopkins, G. H. E. (1952) Mosquitoes of the Ethiopian Region I. - Larval bionomics of mosquitoes and taxonomy of culicine larvae. 2 nd Edition. British Museum (Natural History), London, viii + 355 pp."]}

Culex (Culex) grahamii Theobald subspecies farakoensis Hamon, 1955 —original combination: Culex grahami [sic] var. farakoensis (subspecific status by Harbach & Howard 2007). Distribution: Southern Mali (Hamon 1955). subspecies grahamii Theobald, 1910 —original combination: Culex grahamii [nomen novum for Culex pullatus Graham, 1910]. Distribution: Angola, Benin, Burkina Faso, Cameroon, Central African Republic, Côte d’Ivoire, Democratic Republic of the Congo, Ghana, Guinea, Kenya, Liberia, Madagascar, Mali, Nigeria, Republic of the Congo, Senegal, Sudan and South Sudan, The Gambia, Togo, Uganda (Knight & Stone 1977; Wilkerson et al. 2021). Graham (1910) described the male and female of this species (as Cx. pullatus) from specimens collected in the vicinity of Lagos, Nigeria (Stone et al. 1959; Townson 1990). The larva was described by Wesché (1910), as Cx. pullatus, from specimens collected around Lagos, described by Macfie & Ingram (1923) from specimens collected in the vicinity of Accra, Ghana (as Gold Coast) and by Hopkins (1936, 1952) from undisclosed localities, except for reference to specimens described by Macfie & Ingram. Hamon (1955) described farakoensis from specimens collected “aux cascades de Farako, Cercle et Subdivision de Sikasso, Soudan Français ” (at Farako waterfalls, Sikasso Cercle and Subdivision, French Sudan = Mali). Farako Waterfalls are located about 30 km east of Sikasso in Sikasso Cercle, which is one of seven administrative subdivisions of the Sikasso Region of southern Mali. Based on this, the type locality of farakoensis that is listed as “Farako, Sikasso, French Sudan ” in the catalogs of Stone et al. (1959), Knight & Stone (1977) and Wilkerson et al. (2021) must be corrected to Farako Waterfalls, Sikasso Cercle, Mali (formerly French Sudan). The listing of Nigeria in the distribution given by Wilkerson et al. (2021) appears to be in error. Hamon (1955) provided a brief description of farakoensis, stating that the adults are “almost identical to the typical form and probably not separable with certainty. Male terminalia [genitalia] do not show any difference…” and the “general morphology [of the larva] is that of Cx. grahami [sic] except for the eighth abdominal segment…” (translated from the French). Hamon recorded the following distinctions: Comb with 13–19 spine-like scales, siphon index 7.9–10.4, pecten with 6–9 simple spines, seta 1-S with about 16 setae (this is misleading as the illustration provided includes a ventral row of 12 setae beyond the pecten and a lateral row of 5 setae on the distal half of the siphon) with 2–4 branches and the 8 proximal setae at least as long as the diameter of the siphon; anal papillae 2.5–4.0 times as long as the saddle. Hamon stated that the dorsal pair of anal papillae were usually longer than the ventral pair, but the illustration shows the ventral pair are longer than the dorsal pair. In the typical form, these morphological traits, based on the descriptions of Wesché (1910), Macfie & Ingram (1923) and Hopkins (1936, 1952), are characterized as follows: Comb with 14–25 spine-like scales, siphon index 12–15, pecten also with 6–9 simple spines, seta 1-S with about 4 setae (6 are illustrated by Hopkins), minute, with 1–3 branches, one seta inserted within the pecten; anal papillae slightly longer than the saddle, usually equally long or dorsal pair slightly longer. Thus, only three of these characters are distinctive for farakoensis, i.e. the significantly shorter siphon, the more numerous and longer siphonal setae all borne distal to the pecten and the greater length of the anal papillae. However, the comb scales illustrated for farakoensis appear to differ from those illustrated for the typical form. The lateral spicules progressively increase in length to the sides of the median apical spine in farakoensis whereas in the typical form they comprise lateral fringes of equal length borne proximal to a stronger apical spine. Hamon (1955) concluded the description of farakoensis with the following observation: “This species [farakoensis] was already known from various places of the A. O. F. [Afrique-Occidentale française (French West Africa)], but the only locality having given us larvae identical to the description of Hopkins is Danané (Forest Zone of lower Côte d’Ivoire). All the larvae collected in the savannah zone: Agba-titoé (Togo), Thiès (Senegal), Bobo Dioulasso (Upper Volta) [Burkina Faso] and in various sites of Casamance [Senegal] have in fact the color and chaetotaxy of the typical form but their siphon indexes rarely exceed 11 [translated from the French].” This indicates that the siphon index of farakoensis may be slightly greater than noted in the description, but it is still smaller than that of the typical form. This observation, the morphological distinctions noted above and the apparently close association with savannah strongly suggest that farakoensis is likely to be genetically distinct from Cx. grahamii sensu stricto; accordingly, this nominal form is formally elevated to species status: Culex (Culex) farakoensis Hamon, 1955 . Culex farakoensis is currently listed as a species in the Encyclopedia of Life, however the date of authorship for the former species needs to be corrected from 1954 to 1955., Published as part of Harbach, Ralph E. & Wilkerson, Richard C., 2023, The insupportable validity of mosquito subspecies (Diptera: Culicidae) and their exclusion from culicid classification, pp. 1-184 in Zootaxa 5303 (1) on pages 84-85, DOI: 10.11646/zootaxa.5303.1.1, http://zenodo.org/record/8043342, {"references":["Harbach, R. E. & Howard, T. M. (2007) Corrections in the status and rank of names used to denote varietal forms of mosquitoes (Diptera: Culicidae). Zootaxa, 1542 (1), 35 - 48. https: // doi. org / 10.11646 / zootaxa. 1542.1.3","Theobald, F. V. (1910) A monograph of the Culicidae or mosquitoes. Vol. 5. British Museum (Natural History), London, xiv + (1) + 646 pp., 6 pls. https: // doi. org / 10.5962 / bhl. title. 58067","Graham, W. M. (1910) On new species of West-African Culicidae. Annals and Magazine of Natural History, including Zoology, Botany, and Geology, Series 8, 5 (27), 264 - 273. https: // doi. org / 10.1080 / 00222931008692764","Knight, K. L. & Stone, A. (1977) A catalog of the mosquitoes of the world (Diptera: Culicidae). 2 nd Edition. The Thomas Say Foundation. Vol. VI. Entomological Society of America, College Park, Maryland, xi + 611 pp.","Wilkerson, R. C., Linton, Y. - M. & Strickman, D. (2021) Mosquitoes of the world. Vols. 1 & 2. Johns Hopkins University Press, Baltimore, Maryland, 1332 pp. https: // doi. org / 10.1186 / s 13071 - 021 - 04848 - 6","Stone, A., Knight, K. L. & Starcke, H. (1959) A synoptic catalog of the mosquitoes of the world (Diptera, Culicidae). The Thomas Say Foundation. Vol. VI. Entomological Society of America, College Park, Maryland, vi + 358 pp.","Wesche, W. (1910) On the larval and pupal stages of West African Culicidae. Bulletin of Entomological Research, 1 (1), 7 - 50, 7 pls. https: // doi. org / 10.1017 / S 0007485300000043","Macfie, J. W. S. & Ingram, A. (1923) The early stages of West African mosquitos. - VI. Bulletin of Entomological Research, 13 (4), 409 - 442. https: // doi. org / 10.1017 / S 000748530004551 X","Hopkins, G. H. E. (1936) Mosquitoes of the Ethiopian Region I. - Larval bionomics of mosquitoes and taxonomy of culicine larvae. Printed by order of the Trustees, British Museum (Natural History), London, (vi) + 250 pp.","Hopkins, G. H. E. (1952) Mosquitoes of the Ethiopian Region I. - Larval bionomics of mosquitoes and taxonomy of culicine larvae. 2 nd Edition. British Museum (Natural History), London, viii + 355 pp."]}

Culex (Culex) pruina Theobald subspecies eschirasi Galliard, 1931 —original combination: Culex pruina var. eschirasi (subspecific status by Harbach & Howard 2007). Distribution: Burkina Faso, Cameroon, Central African Republic, Gabon, Côte d’Ivoire, Nigeria, Republic of the Congo, Uganda (Wilkerson et al. 2021). subspecies pruina Theobald, 1901d —original combination: Culex pruina. Distribution: Cameroon, Central African Republic, Côte d’Ivoire, Democratic Republic of the Congo, Gabon, Ghana, Liberia, Nigeria, Republic of the Congo, Sierra Leone, South Sudan, Uganda (Wilkerson et al. 2021, but not Sudan, see Simsaa et al. 2021). With the exception of Burkina Faso, subspecies eschirasi has been recorded from eight of the 12 countries in which the type form has been recorded. The larvae of the two forms, which are apparently sympatric, exhibit significant differences. The larva of the type form was described by Macfie & Ingram (1916), and their description and illustrations of the head and terminal abdominal segments were utilized by Hopkins (1936, 1952), with little additional information. Galliard (1931) illustrated the terminal abdominal segments of eschirasi and noted two major distinctions from the type form. Hopkins (1936, 1952) illustrated a longer siphon for eschirasi and reiterated the differences between the two forms noted by Galliard (1931). In the type form, the posterior surface of the siphon between the pecten of either side is covered with microtrichium-like spicules (aculeae) whereas in subspecies eschirasi the entire surface of the siphon, except narrowly at the base, is covered with aculeae. Both authors indicate that the integument of the thorax and abdominal segments I–VII and part of VIII bears a dense covering of aculeae. Harbach et al. (2017) recorded the presence of thoracic and abdominal aculeae in Cx. pruina, but they did not indicate which subspecific form was examined. Other differences noted in the descriptions and illustrations of the two forms include the following: Comb scales evenly fringed in eschirasi whereas some scales are only fringed at the apex and on one side in the type form; the siphonal setae (seta 1-S) are about 0.6 the diameter of the siphon in the former and about 0.5 the diameter in the latter; seta 1-X is short and single in eschirasi and longer with three branches in the type form; and the anal papillae are equally long in eschirasi, slightly longer than the length of the saddle, whereas the dorsal and ventral papillae are unequal and the dorsal pair is slightly shorter than the length of the saddle in the type form. The male genitalia of the typical form were partially described and illustrated by Edwards (1914, 1929 b, 1941). The genitalia of eschirasi have not been illustrated, but Galliard (1931) noted that they look a little different than those of the type form: “appendices chitineux du tube génital (mésosome) sont profondément divisés en deux” [lateral plates of the genital tube (aedeagus + lateral plates) are deeply split in two]. However, Edwards (1941) stated that he could not find any differences between the genitalia of the two forms. That aside, in view of the morphological distinctions of the larvae and the sympatry of the two forms, it is likely that eschirasi is genetically distinct and should therefore be afforded specific status: Culex (Culex) eschirasi Galliard, 1931 . Culex eschirasi is currently listed as a species in the Encyclopedia of Life. Culex pruina has a single synonym, Cx. pallidothoracis Theobald, 1909 (type locality: Obuasi, Ashanti Region, Ghana). Until topotypic larvae of this nominal species are available for examination, Cx. pallidothoracis should remain a synonym of Cx. pruina., Published as part of Harbach, Ralph E. & Wilkerson, Richard C., 2023, The insupportable validity of mosquito subspecies (Diptera: Culicidae) and their exclusion from culicid classification, pp. 1-184 in Zootaxa 5303 (1) on pages 88-89, DOI: 10.11646/zootaxa.5303.1.1, http://zenodo.org/record/8043342, {"references":["Galliard, H. (1931) Culicides du Gabon. I. - Culicines, avec la description d'une espece et de deux varietes nouvelles. Annales de Parasitologie Humaine et Comparee, 9 (3), 225 - 232. https: // doi. org / 10.1051 / parasite / 1931093225","Harbach, R. E. & Howard, T. M. (2007) Corrections in the status and rank of names used to denote varietal forms of mosquitoes (Diptera: Culicidae). Zootaxa, 1542 (1), 35 - 48. https: // doi. org / 10.11646 / zootaxa. 1542.1.3","Wilkerson, R. C., Linton, Y. - M. & Strickman, D. (2021) Mosquitoes of the world. Vols. 1 & 2. Johns Hopkins University Press, Baltimore, Maryland, 1332 pp. https: // doi. org / 10.1186 / s 13071 - 021 - 04848 - 6","Theobald, F. V. (1901 d) Notes on a collection of mosquitoes from West Africa, and descriptions of new species. Memoirs of the Liverpool School of Tropical Medicine, 4 (Appendix), i - xiv, 3 pls.","Simsaa, M. A. A., Harbach, R. E., Almalik, A. M. A., Ahmed, E. M., Esia, A. A., Mohamed, A. H. & Azrag, R. S. (2021) Culex mosquitoes (Diptera: Culicidae) recorded along the Nile River in northern Sudan, with a key for the identification of all species of the genus known to occur in the country. Zootaxa, 4963 (3), 401 - 411. https: // doi. org / 10.11646 / zootaxa. 4963.3.1","Macfie, J. W. S. & Ingram, A. (1916) New culicine larvae from the Gold Coast. Bulletin of Entomological Research, 7 (1), 1 - 18. https: // doi. org / 10.1017 / S 000748530001703 X","Hopkins, G. H. E. (1936) Mosquitoes of the Ethiopian Region I. - Larval bionomics of mosquitoes and taxonomy of culicine larvae. Printed by order of the Trustees, British Museum (Natural History), London, (vi) + 250 pp.","Hopkins, G. H. E. (1952) Mosquitoes of the Ethiopian Region I. - Larval bionomics of mosquitoes and taxonomy of culicine larvae. 2 nd Edition. British Museum (Natural History), London, viii + 355 pp.","Edwards, F. W. (1914) New species of Culicidae in the British Museum, with notes on the genitalia of some African Culex. Bulletin of Entomological Research, 5 (1), 63 - 81. https: // doi. org / 10.1017 / S 0007485300034568","Edwards, F. W. (1929 b) Mosquito notes. - VIII. Bulletin of Entomological Research, 20 (3), 321 - 343. https: // doi. org / 10.1017 / S 0007485300021283","de Meillon, B. & Rebelo, A. (1941) Culicini (Diptera, Nematocera) from the Colony of Mocambique. Mocambique Documentario trimestral, 27, 69 - 77.","Edwards, F. W. (1941) Mosquitoes of the Ethiopian Region III. - Culicine adults and pupae. Printed by order of the Trustees, British Museum (Natural History), London, viii + 499 pp.","Theobald, F. V. (1909) Descriptions of the new mosquitoes collected by Dr. Graham in Ashanti. Gold Coast Miscellaneous Publication No. 237. Great Britain Colonial Office. London, 31 pp., 2 pls."]}

Culex (Microculex) imitator Theobald subspecies imitator Theobald, 1903a —original combination: Culex imitator. Distribution: Argentina, Brazil, Colombia, Ecuador, French Guiana, Guyana, Honduras, Lesser Antilles (includes Trinidad and Tobago), Mexico, Panama, Suriname, Uruguay, Venezuela (Wilkerson et al. 2021). subspecies retrosus Lane & Whitman, 1951 —original combination: Culex (Microculex) imitator retrosus. Distribution: Brazil (Lane & Whitman 1951). Despite being included in many taxonomic studies, Cx. imitator has only been superficially described, especially the larval and pupal stages. This also applies to subspecies retrosus, which has only been described once (Lane & Whitman 1951; summarized by Lane 1953). However, Lane & Whitman provided distinctions, some unmentioned but illustrated, which seem to indicate that retrosus is a separate species. The differences include the following. In the male genitalia, ninth tergal lobe (as illustrated) short with relatively weak apical setae only slightly longer than the lobe (lobe elongate with strong apical setae about as long as the lobe in imitator); subapical lobe of the gonocoxite (as illustrated) relatively long, setae a and b equally broad with narrow hooked apices, proximal side of lobe with a row of relatively long setae (lobe shorter, setae a and b constricted before short somewhat leaf-like apical segment, seta b distinctly stouter than seta a, proximal side of lobe with a row of relatively short setae in imitator); basal hook of the lateral plate of the phallosome strongly curved/sharply bent (slightly curved, not bent in imitator). In the larva, based on the “prothoracic hair formula”, seta 4-P 3-branched and setae 6,8-P 2-branched (seta 4-P 2- branched and setae 6,8-P single in imitator). This, coupled with molecular evidence that Cx. imitator is a species complex in Brazil (Demari-Silva et al. 2011), based on high intraspecific divergence (3.5%) in the mitochondrial COI gene and associated morphological differences in the pupal stage, we believe it is likely that further study will reveal retrosus is a separate species. Thus, retrosus is hereby elevated to specific status: Culex (Microculex) retrosus Lane & Whitman, 1961 . Culex retrosus is currently listed as a species in the Encyclopedia of Life. Three nominal species are currently considered to be synonyms of Cx. imitator: Cx. daumasturus Dyar & Knab, 1906b (type locality: Trinidad and Tobago), Cx. vector Dyar & Knab, 1906b (type locality: Trinidad and Tobago) and Microculex argenteoumbrosus Theobald, 1907 (type locality: Rio de Janeiro, Brazil). It is possible the first two nominal forms represent a distinct species and the last one may be conspecific with Cx. retrosus, but we believe it is prudent to retain them as synonyms of Cx. imitator for the time being., Published as part of Harbach, Ralph E. & Wilkerson, Richard C., 2023, The insupportable validity of mosquito subspecies (Diptera: Culicidae) and their exclusion from culicid classification, pp. 1-184 in Zootaxa 5303 (1) on page 102, DOI: 10.11646/zootaxa.5303.1.1, http://zenodo.org/record/8043342, {"references":["Theobald, F. V. (1903 a) A monograph of the Culicidae or mosquitoes. Vol. 3. British Museum (Natural History), London, xvii + 1 (errata) + 359 pp., foldout table, 17 pls. https: // doi. org / 10.5962 / bhl. title. 58067","Wilkerson, R. C., Linton, Y. - M. & Strickman, D. (2021) Mosquitoes of the world. Vols. 1 & 2. Johns Hopkins University Press, Baltimore, Maryland, 1332 pp. https: // doi. org / 10.1186 / s 13071 - 021 - 04848 - 6","Lane, J. & Whitman, L. (1951) The subgenus \" Microculex \" in Brazil (Diptera, Culicidae). Revista Brasileira de Biologia, 11 (3), 341 - 366.","Lane, J. (1953) Neotropical Culicidae. Vols. I & II. University of Sao Paulo, Sao Paulo, 1112 pp.","Demari-Silva, B., Vesgueiro, F. T., Sallum, M. A. M. & Marrelli, M. T. (2011) Taxonomic and phylogenetic relationships between species of the genus Culex (Diptera: Culicidae) from Brazil inferred from the cytochrome c oxidase I mitochondrial gene. Journal of Medical Entomology, 48 (2), 272 - 279. https: // doi. org / 10.1603 / me 09293","Dyar, H. G. & Knab, F. (1906 b) The larvae of Culicidae classified as independent organisms. Journal of the New York Entomological Society, 14 (4), 169 - 230, 13 pls.","Theobald, F. V. (1907) A monograph of the Culicidae or mosquitoes. Vol. 4. British Museum (Natural History), London, xix + 639 pp., 16 pls. https: // doi. org / 10.5962 / bhl. title. 58067"]}

quinquefasciatus Say Culex quinquefasciatus Say, 1823: 10. Gerecke et al. 1995: 133 [aquatic invertebrates]; Whiteman et al. 2005: 844 [records]; Fonseca et al. 2006: 285 [pathways of expansion]; Causton et al. 2006: 125 [introduced species]; Causton & Sevilla 2007: 144 [introduced species]; Sinclair 2009: 112 [introduced species]; Levin et al. 2009 [avian malaria]; Asigau et al. 2017: 247 [altitudinal range]. Culex pipiens quinquefasciatus: Peck 1996: 122 [mode of introduction]; Peck et al. 1998: 228 [introduced species]. Distribution. Introduced. Cosmotropical; Galápagos: Baltra, Floreana, Isabela, San Cristóbal, Santa Cruz (see Eastwood et al. 2019). Remarks. This is a night-biting, domestic mosquito. Larvae are found in foul water, water containers and rain barrels. Adult females are generally believed to be predominantly bird feeders (Carpenter & LaCasse 1955). This species is a vector of the filarial nematode Wuchereria bancrofti, avian pox Aviposvirus spp. and avian malaria Plasmodium relictum (Carpenter & LaCasse 1955; Whiteman et al. 2005; Levin et al. 2009). Culex quinquefasciatus is an introduced species, first recorded in 1985 (Causton et al. 2006). This species was ranked by Causton et al. (2006) as “highly invasive” because of the high potential impact on the endemic bird fauna as a vector of avian malaria and West Nile virus (Eastwood et al. 2011, 2019)., Published as part of Sinclair, Bradley J., 2023, An annotated checklist of the Diptera of the Galápagos Archipelago (Ecuador), pp. 1-102 in Zootaxa 5283 (1) on page 22, DOI: 10.11646/zootaxa.5283.1.1, http://zenodo.org/record/7912667, {"references":["Say, T. (1823) Descriptions of dipterous insects of the United States. Journal of the Academy of Natural Sciences of Philadelphia, 3, 9 - 54, 73 - 104.","Gerecke, R., Peck, S. B. & Pehofer, H. E. (1995) The invertebrate fauna of the inland waters of the Galapagos Archipelago (Ecuador) - a limnological and zoogeographical summary. Archiv fur Hydrobiologie, Supplement, 107 (2), 113 - 147.","Whiteman, N. K., Goodman, S. J., Sinclair, B. J., Walsh, T., Cunningham, A. A., Kramer, L. D. & Parker, P. G. (2005) Establishment of the avian disease vector Culex quinquefasciatus Say, 1823 (Diptera: Culicidae) on the Galapagos Islands, Ecuador. Ibis, 147 (4), 844 - 847. https: // doi. org / 10.1111 / j. 1474 - 919 X. 2005.00468. x","Fonseca, D. M., Smith, J. L., Wilkerson, R. C. & Fleischer, R. C. (2006) Pathways of expansion and multiple introductions illustrated by large genetic differentiation among worldwide populations of the southern house mosquito. The American Journal of Tropical Medicine and Hygiene, 74 (2), 284 - 289. https: // doi. org / 10.4269 / ajtmh. 2006.74.284","Causton, C. E., Peck, S. B., Sinclair, B. J., Roque-Albelo, L., Hodgson, C. J. & Landry, B. (2006) Alien insects: threats and implications for conservation of Galapagos Islands. Annals of Entomological Society of America, 99 (1), 121 - 143. https: // doi. org / 10.1603 / 0013 - 8746 (2006) 099 [0121: AITAIF] 2.0. CO; 2","Causton, C. & Sevilla, C. (2007) Latest records of introduced invertebrates in Galapagos and measures to control them. Galapagos Report, 2006 - 2007, 142 - 145.","Sinclair, B. J. (2009) Dipteran biodiversity of the Galapagos. In: Pape, T., Bickel, D. & Meier, R. (Eds.), Diptera Diversity: Status, Challenges and Tools. Brill, Leiden, pp. 98 - 120. https: // doi. org / 10.1163 / ej. 9789004148970. I- 459.21","Levin, I. I, Outlaw, D. C., Vargas, F. H. & Parker, P. G. (2009) Plasmodium blood parasite found in endangered Galapagos penguins (Spheniscus mendiculus). Biological Conservation, 142, 3191 - 3195. https: // doi. org / 10.1016 / j. biocon. 2009.06.017","Asigau, S., Hartman, D. A., Higashiguchi, J. M. & Parker, P. G. (2017) The distribution of mosquitoes across an altitudinal gradient in the Galapagos Islands. Journal of Vector Ecology, 42 (2), 243 - 253. https: // doi. org / 10.1111 / jvec. 12264","Peck, S. B. (1996) Origin and development of an insect fauna on a remote archipelago: The Galapagos Islands, Ecuador. In: Keast, A. & Miller, S. E. (Eds.), The origin and evolution of Pacific Island biotas, New Guinea to Eastern Polynesia: patterns and processes. Academic Publishing, Amsterdam, pp. 91 - 122.","Peck, S. B., Heraty, J., Landry, B. & Sinclair, B. J. (1998) Introduced insect fauna of an oceanic archipelago: the Galapagos Islands, Ecuador. American Entomologist, 44 (4), 218 - 237. https: // doi. org / 10.1093 / ae / 44.4.218","Eastwood, G., Cunningham, A. A., Kramer, L. D. & Goodman, S. J. (2019) The vector ecology of introduced Culex quinquefasciatus populations, and implications for future risk of West Nile virus emergence in the Galapagos archipelago. Medical and Veterinary Entomology, 33 (1), 44 - 55. https: // doi. org / 10.1111 / mve. 12329","Carpenter, S. J. & LaCasse, W. J. (1955) Mosquitoes of North America (North of Mexico). University of California Press, Berkley, California, 360 pp. https: // doi. org / 10.1525 / 9780520325098","Eastwood, G., Kramer, L. D., Goodman, S. J. & Cunningham, A. A. (2011) West Nile virus vector competency of Culex quinquefasciatus mosquitoes in the Galapagos Islands. American Journal of Tropical Medicine and Hygiene, 85 (3), 426 - 433. https: // doi. org / 10.4269 / ajtmh. 2011.10 - 0739"]}

Glyphosate is one of the most common herbicides used to control weeds in the USA. This herbicide can impact the mosquito life cycle through being carried to mosquito habitats by runoff. This study investigates the effects of glyphosate on the larval development and oviposition of a laboratory colony of Culex quinquefasciatus. Our experiments indicated that a concentration of 1 g/liter glyphosate was lethal to larval instars or the larvae impacted were either unable to molt to pupae or never emerged as adults. Larval instars exposed to 0.5 g/liter glyphosate experienced a similar impact; however, the larval stage was extended and pupation was considerably delayed. Mosquitoes oviposited in 0.5 g/liter glyphosate laid the same number of egg rafts as in water, but a considerably lower number of eggs exposed to glyphosate hatched as first instars. If gravid females laid their eggs in 1 g/liter glyphosate versus water, the difference between the 2 groups would be statistically significant and a very low number of eggs exposed to glyphosate could ever hatch into larvae. In an oviposition assay carried out in a cage with 3 choices of water, 0.5 g/liter, and 1 g/liter glyphosate, a higher number of egg rafts were laid in water, followed by 0.5 g/liter and 1 g/liter glyphosate concentrations, respectively. Our study indicated such a severe effect of glyphosate on all immature stages of Cx. quinquefasciatus that no adult could ever emerge.

Two Culex pipiens form Pipiens colony strains and a field population of Cx. restuans from Michigan were susceptible and a Cx. pipiens form Molestus colony strain was comparatively less susceptible to a dose of 43 μg/ml of permethrin in Centers for Disease Control and Prevention (CDC) bottle bioassays. Using this diagnostic dose and these populations as controls, adult female Cx. pipiens and Cx. restuans were reared from egg rafts from 28 sites in Illinois, Ohio, Michigan, Iowa, Indiana, Wisconsin, and Minnesota. Tested mosquitoes showed high mortality in populations from 12 sites, less mortality (90–96%) at 9 sites, and less than 90% mortality from 7 sites during 30-min exposures. However, all tested populations showed 97–100% mortality at 60 min, indicating low phenotypic penetrance of resistance factors. These results indicate variation in susceptibility to permethrin in populations of West Nile virus vectors in the Great Lakes region of the United States, with evidence of modest resistance at 7 of 28 (25%) of the sampled populations.

West Nile virus (WNV) is a zoonotic disease that is endemic in North America and is known to cause a range of symptoms from mild to life threatening in humans. Culex quinquefasciatus is one of the most prominent vectors of WNV in Southern California. The goal of this study was to identify which animal species are most fed upon by these mosquitoes in various habitats in the West Valley area of San Bernardino County, California, and determine the relationship between blood-feeding patterns and WNV activity in the region. Culex quinquefasciatus specimens were collected by West Valley Mosquito and Vector Control District during 2011 from 32 different sites. The bloodmeals of 683 individuals (92.4% of those tested) were identified using the mitochondrial gene cytochrome c oxidase 1 (COI). These bloodmeals comprised 29 vertebrate species across four different habitats. Species richness (ranging from 10 to 17) was not significantly different between habitats when rarified to account for sample size. Across habitats, the highest percentage of avian bloodmeals were taken from house sparrows (18.8–39.1%) and house finches (2.6–31.5%). Bloodmeals were identified from five mammalian species, accounting for 5.1–59.2% of bloodmeals by habitat, including humans (0–4.1%). A seasonal shift towards increased mammalian bloodmeal prevalence, specifically for domestic dog and human bloodmeals, was observed in urban habitats. The WNV activity during 2011 in San Bernardino County occurred mostly in urban and suburban areas as indicated by minimum infection rate (MIR) in Culex quinquefasciatus, notable as all human bloodmeals were identified from these two habitats.

As climate change proceeds, the management of the population of mosquitoes becomes more and more challenging. Insect adulticides and larvicides constitute significant control techniques, with the latter being considered the leading mosquito control method. However, the development of mosquito resistance development and the adverse side effects caused by the extensive use of synthetic insecticides have turned research towards the discovery of environmentally-friendly solutions. Plants and bacteria have historically proven to be a good source of insecticidally active compounds, which may possess novel modes of action to overcome current resistance mechanisms and could also possess favorable human and environmental safety profiles. A previous study demonstrated that the naturally occurring prenylated acyl phloroglucinol deoxycohumulone is a potent larvicidal agent against Culex pipiens. Herein the structural characteristics that improve it are explored by evaluating colupulone and novel geranylated analogues.Colupulone, a prenylated acyl phloroglucinol isolated from Humulus lupulus, colupone, and novel geranylated acyl phloroglucinol congeners, were synthesized and evaluated against Cx. pipiens larva. Results indicated that selected derivatives exhibited superior potency than deoxycohumulone (LCThe results determined the relationship between the target activity and the chemical structure of the tested compounds, and they revealed significantly improved larvicidal candidates. These results highlight the potential of the acyl phloroglucinol chemistry for further development of mosquito larvicides. © 2022 Society of Chemical Industry.

Mosquitoes cause life threatening diseases such as yellow fever, malaria, filariasis, encephalitis infection etc. The focus of this research is to evaluate the larvicidal and fumigant properties of Moringa oleifera and Momordica charantia extracts on the larvae and adult mosquitoes. The leaves of both plants were dried and pulverized into fine powder. Rotary evaporator was used to extract the plant oils. The results showed that Momordica charantia was more effective as larvicide than Moringa oleifera as its evoked 100% larval mortality at 20% concentration for 3 hours with LC50 and LC90 of 0.5% and 8.5% respectively. Moringa oleifera produced 100% larval mortality and LC50 (0.75%) and LC90 (10%) at 25% concentration for 4 hours. Similarly, the leaf extract of Momordica charantia is a better fumigant than Moringa oleifera, the former produced LC50 of 0.5% and LC90 of 3.75%, while the latter produced LC50 and LC90 of 1.05% and 4.25% respectively. The significantly higher larvicidal and fumigant activities observed in Momordica charantia is due to the presence of cardiac glycosides only in the plant in addition to saponins, tannins, flavonoids and alkaloids which are common to both plants. Therefore botanicals are advocated to be included in vector control programs. This is because botanicals are relatively safe, cheap and easy to obtain in many parts of the world.

The Educator Group of the Melanoconion Section of Culex (Melanoconion) (Diptera, Culicidae) is reviewed. Currently, the group includes the following valid species: Culex (Melanoconion) aphyllusTalaga, 2020, Culex (Melanoconion) cristovaoi Duret, 1968, Culex (Melanoconion) educatorDyar & Knab, 1906, Culex (Melanoconion) eknomiosForattini & Sallum, 1992, Culex (Melanoconion) inadmirabilisDyar, 1928, Culex (Melanoconion) rachoui Duret, 1968, Culex (Melanoconion) theobaldi (Lutz), 1904, Culex (Melanoconion) vaxusDyar, 1920, Culex (Melanoconion) angularis Sá & Sallum n. sp., Culex (Melanoconion) spiniformis Sá & Hutchings n. sp., Culex (Melanoconion) longistriatus Sá & Hutchings n. sp., Culex (Melanoconion) anelesDyar & Ludlow, 1922, Culex (Melanoconion) apeteticus Howard, Dyar & Ludlow, 1913, and Culex (Melanoconion) bibulusDyar, 1920. The last three species were resurrected from synonymy of either Cx. educator or Cx. vaxus. Descriptions, differential diagnoses, bionomics, and geographical data are provided for each species when available.

Plastic – derived Bisphenol A (BPA) contaminated sewages in Trivandrum, Kerala, India are mosquito breeding sites. After summer rain, BPA in the stagnant water samples ranged between 0.86 and 1.14ppm. 1.0 ppm BPA is considered as environmentally relevant concentration. Present study revealed that BPA is a developmental agonist of Culex quinquefasciatus. Embryonic and larval development are shortened by BPA but pupal development is unaffected. Under the atmospheric temperature of 26 to 310C lifecycle was completed within 312 hours but during summer it was completed within 278 hours, meanwhile atmospheric temperature ranged between 30 and 370C. Reduction in the duration of development due to BPA was 50 to 70 hours during summer and 60 to 80 hours in other seasons. Larval – pupal stadium of the mosquito has three surges of 20-hydroxy ecdysone(20-HE), at 24,32 and 48th hour of the 4th instar. BPA resulted dose-dependent advancement of 20-HE peaks, Phospholipase A2 induction and expression of Ecdysone receptor EcRA and Ecdysone inducible gene E75A, which culminated in early pupation. Adults emerging from 1.0 ppm BPA treatment did not show significant difference in sanguivory and fecundity compared to control but pupae developed in 2 and 4 ppm BPA were significantly small.

Blood-engorged Culex quinquefasciatus and Cx. nigripalpus were collected from 140 locations throughout Sarasota County, FL, from 2017 to 2020 to determine local, habitat-specific, and seasonal variations in the host usage patterns of these 2 important arbovirus vectors. Mosquitoes were collected using light traps, gravid traps, and via aspiration of resting shelters. Host was determined from 920 samples using multiple polymerase chain reaction protocols that target mitochondrial sequences specific to mammals, birds, and reptiles. The data were analyzed to test for statistical associations between host class and season or with habitat categories (urban, suburban, and rural). Culex quinquefasciatus took significantly more blood meals from birds compared to mammals, though a seasonal shift to a higher ratio of mammalian host usage was observed in fall. There was a habitat-dependent pattern of host usage by Cx. nigripalpus, with significantly more mammalian hosts identified from mosquitoes captured in rural habitats and a similar ratio of mammalian and avian hosts in urban habitats. In general humans were used as hosts by Cx. nigripalpus less often compared to Cx. quinquefasciatus. In contrast to previous studies, Cx. nigripalpus utilized ectothermic hosts (mostly reptiles) at a much higher ratio and exhibited no apparent seasonal shift in host selection.

Mosquito blood feeding plays a key role in epidemiology. Despite its importance and large number of studies worldwide, less attention has been paid in South America. We summarized some general concepts and methodological issues related to the study of mosquito blood feeding habits, and compiled and analyzed all published information regarding the subject in the continent until 2020. Available literature comprised 152 scientific studies, that pursued different approaches: human landing catches (102 studies), baited trap (19), and blood meal analyses of collected specimens (38). Among the latter, 23 used serological and 15 molecular techniques. Species most frequently studied were those incriminated in malaria transmission, whereas relevant vectors such as Aedes aegypti, Ae. albopictus, and Haemagogus janthinomys were surprisingly neglected. Brazil was the leading country both in number of works and species studied. For over 70% of the species and three out of 13 South American countries there is no single information on mosquito blood feeding habits. Data from baited traps included 143 mosquito species, 83.9% of which were attracted to humans, either exclusively (10.5%) or in combination with other vertebrates (73.4%). Host blood identification of field collected specimens provided data on 102 mosquito species, and 60.8% of these fed on humans (55.9% combined with other vertebrates). Only 17 of the 73 species assessed by both methods yielded similar feeding patterns. Finally, supplementary tables are provided in a comprehensive summary of all information available and information gaps are highlighted for future research in the continent.

Culex vishnui Theobald, 1901, a main vector of Japanese encephalitis virus (JEV), is widely distributed in the Oriental region where it often accounts for a great part of the culicid fauna. This species also has been found naturally infected with at least 13 other arboviruses of medical and veterinary importance. Females blood feed predominantly upon pigs and birds, but may readily bite cattle and humans. Because of its abundance, medical importance, and presence throughout ecological gradients among urban, peri-urban, and rural areas, Cx. vishnui potentially may serve as a bridge vector transmitting viruses from natural and wild hosts to humans. Being zoo- and anthropophagic, omnipresent in the Oriental region, and presenting strong resistance to many insecticide families, this overlooked mosquito species may pose a serious health risk in one of the most densely populated regions of the world.

Background Mosquitoes are vectors of many pathogens, such as malaria, dengue virus, yellow fever virus, filaria and Japanese encephalitis virus. Wolbachia are capable of inducing a wide range of reproductive abnormalities in their hosts, such as cytoplasmic incompatibility. Wolbachia has been proposed as a tool to modify mosquitoes that are resistant to pathogen infection as an alternative vector control strategy. This study aimed to determine natural Wolbachia infections in different mosquito species across Hainan Province, China. Methods Adult mosquitoes were collected using light traps, human landing catches and aspirators in five areas in Hainan Province from May 2020 to November 2021. Species were identified based on morphological characteristics, species-specific PCR and DNA barcoding of cox1 assays. Molecular classification of species and phylogenetic analyses of Wolbachia infections were conducted based on the sequences from PCR products of cox1, wsp, 16S rRNA and FtsZ gene segments. Results A total of 413 female adult mosquitoes representing 15 species were identified molecularly and analyzed. Four mosquito species (Aedes albopictus, Culex quinquefasciatus, Armigeres subalbatus and Culex gelidus) were positive for Wolbachia infection. The overall Wolbachia infection rate for all mosquitoes tested in this study was 36.1% but varied among species. Wolbachia types A, B and mixed infections of A × B were detected in Ae. albopictus mosquitoes. A total of five wsp haplotypes, six FtsZ haplotypes and six 16S rRNA haplotypes were detected from Wolbachia infections. Phylogenetic tree analysis of wsp sequences classified them into three groups (type A, B and C) of Wolbachia strains compared to two groups each for FtsZ and 16S rRNA sequences. A novel type C Wolbachia strain was detected in Cx. gelidus by both single locus wsp gene and the combination of three genes. Conclusion Our study revealed the prevalence and distribution of Wolbachia in mosquitoes from Hainan Province, China. Knowledge of the prevalence and diversity of Wolbachia strains in local mosquito populations will provide part of the baseline information required for current and future Wolbachia-based vector control approaches to be conducted in Hainan Province. Graphical Abstract

Culex (Phalangomyia) apicinus Philippi, 1865 (Figs 1–3) Culex apicinus Philippi, 1865: 596 (A). Type locality: Santiago, Chile (NE). Refs: Dyar 1928: 360 (M*, F, L*, as Culex (Phalangomyia)); Edwards 1932: 206 (as Culex (Culex)); Lane 1953: 341 (M*, F, L*); Forattini 1965: 143 (M*, F); Bram 1967: 26 (M*, F, L; syn.); Belkin, 1968 (type info.). Phalangomyia debilis Dyar & Knab, 1914: 58 (M*, F). Type locality: Matucana, (Lima), Peru (USNM). Refs: Dyar 1926: 42 (as Culex (Phalangomyia)); Dyar 1928: 359 (M*, F, L*); Edwards 1930: 107, 1932: 206 (syn. with Cx. apicinus Philippi, as Culex (Culex)). Culex escomeli Brèthes, 1920: 41 (M*, F). Type locality: Arequipa, Peru (NE). Refs: Dyar 1926: 42 (as Culex (Phalangomyia)); Dyar 1928: 359 (M*, F, L*); Edwards 1930: 107, 1932: 206 (syn. with Cx. apicinus Philippi, as Culex (Culex)). Culex (Phalangomyia) alticola Martini, 1931: 216 (A). Type locality: Serrata, 2,300 m, Bolivia (NHM). Refs: Lane 1953: 341 (M*, F, L; syn. with Cx. apicinus Philippi); Stone 1957: 340 (M*); Belkin 1968: 12 (tax.); Bram 1967: 24, 27 (syn.). Diagnosis. Diagnostic morphological characteristics of Cx. apicinus, contrasted with other species of the subgenus Culex, are as follows (Table 2). Adult: Flagellomere 1 of antenna with scales on median border (Fig. 1A). Proboscis longer than forefemur. Maxillary palpus of female dark brown, with white-scales (Fig. 1A, B). Scutum with 2 narrow denuded stripes between acrostichal and dorsocentral setae; acrostichal, dorsocentral, and supra-alar setae dark brown (Fig. 1C). Wing with white scale-spot near middle of veins M 2 and M 3 (Fig. 1D). Forefemur (2.23–3.01 mm) longer than foretarsomere 1 (1.00– 1.84 mm). Foretarsomere 1 as long as sum of length of foretarsomeres 2–5. Terga II‒VII dark brown-scaled with a complete basal band of pale scales (Fig. 1E). Female genitalia: Upper vaginal lip and sclerite strongly sclerotized, cerci longer than postgenital lobe. Male genitalia: Tergum VIII deeply emarginate. Gonocoxite with 8–20 strong apicolateral setae (Fig. 2A). Subapical lobe with 2 setae: A large leaflet with serration on distal margin and ending in a long filament (seta h), and the other tubular, ending in 2 open arms (seta g) (Fig. 2B). Median lobe partially divided: Larger proximal part bearing 9–28 strong rod-like setae and 2–3 normal setae and the small distal part bearing a strong apically hooked seta, a longer stout seta with a bent fimbriate apex and a shorter stout seta with tip bifurcate (Fig. 2C). Gonostylus with a spiny apical crest (Fig. 2A), with 5–8 subapical setae (Fig. 2D). Dorsal arm of phallosome short (Fig. 2E). Paraproct with a small basolateral arm (Fig. 2E), with 7–20 cercal setae not visible in Fig. 2E), IXth tergal lobe with 2–13 setae in 1 or 2 rows. Pupa: Trumpet conical (Fig. 3A) opening wide, large and oval. At least alveolus of seta 13-III–VI present and paddle with minute spicules on the surface (Fig. 3B). Larva (fourth instar): Head, segment X and siphon heavily tanned (Fig. 3C). Siphon unique (Fig. 3C), as follows: siphon index 2.50–3.33 (mean 2.94), pecten on basal 0.60, with 5–11 spines, seta 1-S usually in 5 pairs, 1a, 1b-S inserted between pecten spines, 1e/f-S out of line. Distribution. Argentina, Bolivia, Chile and Peru., Published as part of González, Christian R., Reyes-Valenzuela, Carolina, Rossi, Gustavo C. & Laurito, Magdalena, 2023, Revalidation of Phalangomyia Dyar & Knab as a subgenus of Culex L. (Diptera: Culicidae) based on morphological and molecular evidence, pp. 544-564 in Zootaxa 5256 (6) on page 552, DOI: 10.11646/zootaxa.5256.6.2, http://zenodo.org/record/7762560, {"references":["Philippi, R. A. (1865) Aufzahlung der chilenischen Dipteren. Verhandlungen der Kaiserlich-Ko ¨ niglichen Zoologisch-Botanischen Gesellschaft in Wien, 15, 595 - 782, 7 pls.","Dyar, H. G. (1928) The mosquitoes of the Americas. Carnegie Institute Washington Publishing, Washington, D. C., 616 pp.","Edwards, F. W. (1932) Genera Insectorum. Diptera, Fam. Culicidae. Fascicle 194. V. Verteneuil & L. Desmet, Bruxelles [Brussels], 258 pp., 5 pls.","Lane, J. (1953) Neotropical Culicidae. Vol. 1. Editora da Universidade de Sao Paulo, Sao Paulo, 548 pp.","Forattini, O. P. (1965) Entomologia Medica. Editora da Universidad de SaTo Paulo, SaTo Paulo, 506 pp.","Bram, R. A. (1967) Classification of Culex subgenus Culex in the New World (Diptera: Culicidae). Proceedings of the United States National Museum, 120 (3557), 1 - 122.","Belkin, J. N. (1968) Mosquito studies (Diptera: Culicidae). IX. The type specimens of New World mosquitoes in European museums. Contributions of the American Entomological Institute, 3 (4), 1 - 69.","Dyar, H. G. & Knab, F. (1914) New mosquitoes from Peru. Insecutor Inscitiae Menstruus, 2 (4), 58 - 62.","Dyar, H. G. (1926) A note on Phalangomyia. Insecutor Inscitiae Menstruus, 14, 41 - 43.","Edwards, F. W. (1930) Bibionidae, Scatopsidae, Cecidomyiidae, Culicidae, Thaumaleidae (Orphnephilidae), Anisopodidae (Rhyphidae). Fascicle 3. In: Diptera of Patagonia and South Chile based mainly on material in the British Museum (Natural History). Pt. 2. British Museum of Natural History, London, pp. 77 - 120.","Brethes, J. (1920) Description d´un nouveau moustique du Perou. Revista Chilena de Historia Natural, 24, 41 - 43.","Martini, E. (1931) Ueber einige s ¸ damerikanische Culiciden. Revista de Entomologia Rio de Janeiro, 1 (2), 199 - 219.","Stone, A. (1957) Corrections in the taxonomy and nomenclature of mosquitoes (Diptera: Culicidae). Proceedings of the Entomological Society of Washington, 58 (6), 333 - 344. [for 1956]"]}

Introduction During evolution, blood-feeding arthropods developed a complex salivary mixture that can interfere with host haemostatic and immune response, favoring blood acquisition and pathogen transmission. Therefore, a survey of the salivary gland contents can lead to the identification of molecules with potent pharmacological activity in addition to increase our understanding of the molecular mechanisms underlying the hematophagic behaviour of arthropods. The southern house mosquito, Culex quinquefasciatus, is a vector of several pathogenic agents, including viruses and filarial parasites that can affect humans and wild animals. Results Previously, a Sanger-based transcriptome of the salivary glands (sialome) of adult C. quinquefasciatus females was published based on the sequencing of 503 clones organized into 281 clusters. Here, we revisited the southern mosquito sialome using an Illumina-based RNA-sequencing approach of both male and female salivary glands. Our analysis resulted in the identification of 7,539 coding DNA sequences (CDS) that were functionally annotated into 25 classes, in addition to 159 long non-coding RNA (LncRNA). Additionally, comparison of male and female libraries allowed the identification of female-enriched transcripts that are potentially related to blood acquisition and/or pathogen transmission. Conclusion Together, these findings represent an extended reference for the identification and characterization of the proteins containing relevant pharmacological activity in the salivary glands of C. quinquefasciatus mosquitoes.

Pale-banded variants of Culex territans Walker larvae were found in southern New Jersey in 2016 and in subsequent years. The larvae lacked pigmentation on the prothoracic and mesothoracic segments as well as on abdominal segment (IV). The amount of pigmentation varied between populations. Specimens were found in six locations within a 5-yr period; of which, two locations recurred within the same season and in subsequent seasons. This is the first report in the State of New Jersey despite extensive larval surveillance over many years among the study area.

Culex pipiens (Linnaeus), one of the most abundant mosquito species in Europe, plays a crucial role in the endemic transmission of West Nile virus and caused the large outbreak with1600 human cases in 2018. Although evidence of resistance to pyrethroids has been reported for Cx. pipiens populations from Spain and Greece, resistance monitoring has been largely neglected in Italy. Herein, we investigate susceptibility of Italian Cx. pipiens populations to the pyrethroids permethrin and deltamethrin. Results from WHO-tube-bioassays revealed mortalities ranging from 14-54%, indicating high levels of resistance, in four out of 10 populations exposed to permethrin (0.75%) and of 63% in one of three populations exposed to deltamethrin (0.05%). Reduced susceptibility (mortality98%) was detected in almost all other populations. A clear association is shown between the resistant phenotype and the presence of kdr-alleles in position 1014 of the VSSC, strongly suggesting its role in reducing susceptibility. The study provides the first evidence of pyrethroid-resistance in Italian Cx. pipiens populations and reports levels of resistance paralleled in the European region only in Turkey. This highlights the urgent need to implement insecticide-resistance management plans to restore the efficacy of the nowadays only chemical weapon available to control arbovirus transmission in Europe.

The extensive use of insecticides in agriculture and public health has resulted in the rapid development of insecticide resistance in mosquito populations. Therefore, in this study, we aimed to evaluate insecticide resistance costs on the fitness of Culex pipiens. Two Cx. pipiens field populations (Beheira malathion-resistant and Gharbia malathion-susceptible) were compared to the reference (sensitive) population. The biochemical composition and expression of four genes relevant to insecticide resistance were estimated in third instar larvae. Adult survival, female fecundity and egg hatchability were also determined. As per our findings, it was found that the total protein and carbohydrate contents in Beheira malathion-resistant larvae were significantly lower than that in the reference larvae. Beheira malathion-resistant larvae had higher phenoloxidase (PO) specific activity than the reference population. In terms of the relevant genes, only cytochrome P450 (CYP6F1) expression showed elevated levels in the Gharbia malathion-susceptible population compared to the Beheira malathion-resistant population. In esterases (Estα and Estβ) and glutathione S-transferase, the tested populations did not show any significant differences. Compared to the reference mosquito population, Gharbia malathion-susceptible Cx. pipiens males exhibited significantly longer median survival. Female fecundity and hatchability showed nonsignificant differences among the populations tested. In conclusion, malathion resistance can induce lower protein and carbohydrate contents, but higher PO activity in larvae.

Information on the mosquito species that transmit canine filariosis is scanty. Hence, an experimental study was conducted to identify the potential vectors responsible for the transmission of D. immitis Leidy and B. pahangi Buckley & Edeson. A total of 367 mosquitoes belonging to six species containing both laboratory and field strains (i.e. Aedes togoi Theobald, Aedes aegypti Linnaeus, Aedes albopictus Skuse, Culex quinquefasciatus Say, Culex vishnui Theobald and Anopheles dirus Peyton & Harrison) were used in this study. All mosquitoes were artificially fed on either D. immitis or B. pahangi microfilariae (mfs) infected blood by using the Hemotek™ membrane feeding system. Out of 367 mosquitoes, 228 (64.9%) were fully engorged. After feeding on D. immitis (20%) and B. pahangi (33%) mfs positive blood, the mortality rates for Cx. quinquefasciatus were found to be slightly lower than that of other species of mosquitoes. On the other hand, majority of An. dirus were found to be incapable to withstand the infection of mfs as the mortality rates were relatively high (D. immitis = 71.4%; B. pahangi = 100.0%). Brugia pahangi was detected in Ae. togoi and Cx. quinquefasciatus with infection rates of 50% and 25%, respectively. Aedes togoi was the only species infected with D. immitis with an infection rate of 69%. Our results showed that Ae. togoi was an excellent experimental vector for both D. immitis and B. pahangi. This study also documented the observation of B. pahangi, for the first time in the head region of Cx. quinquefasciatus under a laboratory setting.

The northern house mosquito Culex pipiens sensu stricto is one of the most important disease vector mosquitoes in temperate zones across the northern hemisphere, responsible for the emergence of West Nile Virus over the last two decades. It comprises two ecologically distinct forms - an aboveground form, pipiens, diapauses in winter and primarily bites birds, while a belowground form, molestus, thrives year-round in subways, basements and other human-made, belowground habitats, bites mammals, and can even lay eggs without a blood meal. The two forms hybridize in some but not all places, leading to a complex ecological mosaic that complicates predictions of vectorial capacity. Moreover, the origin of the belowground molestus is contentious, with iconic populations from the London Underground subway system being held up by evolutionary biologists as a preeminent example of rapid, in situ, urban adaptation and speciation. We review the recent and historical literature on the origin and ecology of this important mosquito and its enigmatic forms. A synthesis of genetic and ecological studies spanning 100+ years clarifies a striking latitudinal gradient - behaviorally divergent and reproductively isolated forms in northern Europe gradually break down into what appear to be well-mixed, intermediate populations in North Africa. Moreover, a continuous narrative thread dating back to the original description of form molestus in Egypt in 1775 refutes the popular idea that belowground mosquitoes in London evolved in situ from their aboveground counterparts. These enigmatic mosquitoes are more likely derived from populations in the Middle East, where human-biting and other adaptations to human environments may have evolved on the timescale of millennia rather than centuries. We outline several areas for future work and discuss the implications of these patterns for public health and for our understanding of urban adaptation in the Anthropocene.

Mosquito surveillance is critical to reduce the risk of West Nile virus (WNV) transmission to humans. In response to surveillance indicators such as elevated mosquito abundance or increased WNV levels, many mosquito control programs will perform truck-mounted ultra-low volume (ULV) adulticide application to reduce the number of mosquitoes and associated virus transmission. Despite the common use of truck-based ULV adulticiding as a public health measure to reduce WNV prevalence, limited evidence exists to support a role in reducing viral transmission to humans. We use a generalized additive and fused ridge regression model to quantify the location-specific impact of truck-mounted ULV adulticide spray efforts from 2010 to 2018 in the North Shore Mosquito Abatement District (NSMAD) in metropolitan Chicago, IL, on commonly assessed risk factors from NSMAD surveillance gravid traps: Culex abundance, infection rate, and vector index. Our model also takes into account environmental variables commonly associated with WNV, including temperature, precipitation, wind speed, location, and week of year. Since it is unlikely ULV adulticide spraying will have the same impact at each trap location, we use a spatially varying spray effect with a fused ridge penalty to determine how the effect varies by trap location. We found that ULV adulticide spraying has an immediate temporary reduction in abundance followed by an increase after 5 days. It is estimated that mosquito abundance increased more in sprayed areas than if left unsprayed in all but 3 trap locations. The impact on infection rate and vector index were inconclusive due to the large error associated with estimating trap-specific infection rates.