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Aedes (Aedimorphus) vexans (Meigen) subspecies arabiensis (Patton, 1905) —original combination: Culex arabiensis (subspecific status by White 1975). Distribution: Gambia, Mauritania, Mauritius, Nigeria, Saudi Arabia, Somalia, South Africa, South Sudan, Sudan, Tanzania, Yemen (Wilkerson et al. 2021). subspecies nipponii (Theobald, 1907) —original combination: Culicada nipponii (subspecific status by Bohart & Ingram 1946b). Distribution: Georgia, Japan, Mongolia, People’s Republic of China, Russia, South Korea (Wilkerson et al. 2021). subspecies nocturnus (Theobald, 1903a)—original combination: Culex nocturnus (subspecific status by Bohart & Ingram 1946b). Distribution: Australia, Cook Islands (Polynesia), Fiji, Indonesia, Kiribati, Malaysia, Mariana Islands, Marshall Islands, New Caledonia, Palau, Papua New Guinea, Samoa, Solomon Islands, Timor, Tonga, Tuvalu, Vanuatu (Wilkerson et al. 2021). subspecies vexans (Meigen, 1830) —original combination: Culex vexans. Distribution [sensu lato]: Afghanistan, Albania, Algeria, Armenia, Australia, Austria, Azerbaijan, Bangladesh, Belarus, Belgium, Belize, Bosnia and Herzegovina, Bulgaria, Cambodia, Canada, Corsica, Crimean Peninsula, Croatia, Czech Republic, Denmark, Estonia, Fiji, Finland, France, Former Yugoslav Republic of Macedonia, Gambia, Georgia, Germany, Greece, Guam, Guatemala, Honduras, Hong Kong, Hungary, India, Indonesia Iran, Iraq, Italy Japan, Jordan, Kazakhstan, Kiribati, Kosovo, Laos, Latvia, Liberia, Libya, Lithuania, Macedonia, Malaysia, Malta, Mariana Islands, Mauritania, Mexico, Micronesia, Moldova, Mongolia, Montenegro, Morocco, Myanmar, Nepal, Netherlands, New Caledonia, New Zealand, Norway, Pakistan, Papua New Guinea, People’s Republic of China, Philippines, Poland, Portugal, Romania, Russia, Samoa, Saudi Arabia, Serbia, Singapore, Slovakia, Slovenia, Solomon Islands, South Africa, South Korea, Spain, Sri Lanka, Sweden, Switzerland, Taiwan, Tajikistan, Thailand, Tonga, Turkey, Turkmenistan, Tuvalu, Ukraine, United Kingdom, United States, Uzbekistan, Vanuatu, Vietnam, Yemen (Wilkerson et al. 2021). Aedes vexans sensu lato, occurs worldwide except for South America. Since Ae. vexans sensu lato is a common, medically important species, it has attracted worldwide interest. In summary, we think that two of the nominal subspecies have characteristics that adequately separate them from typical vexans: subspecies nipponii (Japan and the eastern Palaearctic) and subspecies arabiensis (northeastern Africa and Arabia [South Africa?]). However, the third subspecies, nocturnus (Central Pacific and Australasia) has generated taxonomic inconsistency since there are no clear differences to distinguish it. There are regional keys to identify the four nominal subspecies from cooccurring species of Aedes, but there are no keys to separate the subspecies from each other. A partial exception is Reinert (1973), who compared a worldwide concept of vexans vexans with the allopatric vexans nipponii. Nominotypical vexans To compare the nominal forms, we first define nominotypical vexans using a recent description from Europe, the continent of the type locality, Berlin, Germany. The nominotypical subspecies was described from a single female. The type specimen is in the Muséum National d’Histoire Naturelle, Paris, France. There are descriptions for what has been called vexans from many parts of the world, but it is sensible to use a recent description from Europe (Becker et al. 2020). The caveat is that the provenance of specimens and/or literature used for this description are not documented. It will nevertheless serve as a basis for comparison with characters of subspecies and descriptions of vexans from other parts of the world. The following is paraphrased from key characters (diagnoses) followed by more complete descriptions, a format often used by F. W. Theobald. Adult female.Tarsomeres with pale rings only; proboscis distinctly longer than forefemur; scutellum with narrow, curved, yellowish or pale scales; pale basal bands of tarsomeres very narrow, usually not exceeding 0.25 length of tarsomeres; abdominal terga with basal white bands constricted medially, giving them a bilobed appearance. Tibiae dark-scaled dorsally, pale-scaled ventrally; narrow basal bands present on foretarsomeres 2 and 3, midtarsomeres 1–4 and on all hindtarsomeres. Proboscis and maxillary palpus dark-scaled, palpus with some apical white scales; head covered with narrow curved pale and dark decumbent scales and numerous dark brown erect forked scales which extend anteriorly to interocular space. Scutal integument dark brown, covered with narrow curved dark scales and narrow pale scales forming indistinct patches on anterior submedian, prescutellar and dorsocentral areas; acrostichal and dorsocentral setae well developed; postspiracular area with large patch of narrow curved or moderately broad pale scales; upper and lower mesokatepisternal scale-patches present. Wing veins covered with moderately broad dark scales and isolated pale scales at bases of costa and subcosta. Abdominal terga with basal white bands, distally dark-scaled; basal bands on terga III–VI distinctly narrowed medially, giving a bilobed appearance; sternum VIII with distinct apical V-shaped notch. Adult male. Maxillary palpus about as long as proboscis; gonostylus attached at apex of gonocoxite, simple, not divided; gonostylus distinctly expanded apically; claspette elongate, well separated from base of gonocoxite; gonostylus gradually expanded toward apex; gonostylar claw articulated subapically, straight. Tergum IX strongly bilobed with 6–11 setae on each lobe. Gonocoxite long and moderately broad with scattered scales on lateral and ventral surfaces; basal and apical lobes absent; gonostylus widens toward apex; gonostylar claw straight, inserted on a small subapical tubercle; claspette moderately broad basally, apex slightly expanded and rounded, with crown of numerous spine-like setae, some curved apically; claspette filament absent; paraproct with pointed apex; aedeagus strongly sclerotized with lateral plates connected at base. Larva. Integument not covered with obvious spicules. Antenna less than half length of head, with numerous scattered spicules; seta 1-A 5–10 branched, inserted proximal to middle of antenna. Labral brush with median setae apically serrate (in contrast to unmodified setae in Ae. rossicus and Ae. cinereus). Setae 5–7-C arranged in a triangular pattern, 5-C posterior to 6,7-C; 5-C 1–4-branched, 6-C 1- or 2-branched, 7-C 7–9 branched. Comb with 7–13 scales arranged in 1 or 2 irregular rows; scales with long median spicules and small spicules at base. Siphon index 2.3–3.0; pecten with 13–18 spines, apical 2 or 3 spines larger and unevenly spaced; basal spines with 1–3 lateral denticles; seta 1-S inserted beyond middle of siphon, with 3–8 short branches, length about half width of siphon at setal insertion. Saddle large but incomplete; seta 1-X with 1 or 2 branches; seta 4-X with 3 or 4 precratal setae; anal papillae distinctly longer than saddle. The notion that vexans is a single species across Europe and North America was recently questioned in two studies. Krtinić et al. (2013) used allozyme data to analyze two European populations (Serbia and Germany) and one North American population (California, USA). They found unambiguous differences between European and North American vexans. “We observed that populations of Am. [as genus Aedimorphus] vexans from the Palearctic (Germany and Serbia) and from the Nearctic (USA) formed two genetically distinct populations that had no genetic exchange.” In other words, they are genetically separate species. Also, they cited Becker et al. (2020), who wrote that unlike invasive species such as Aedes albopictus, which oviposits above the water line, vexans lays eggs onto moist soil, which argues against it being an invasive species in North America. Since a single North American population was sampled, these results require verification. Lilja et al. (2018), using the mitochondrial COI gene and two nuclear genes found two sympatric genetically isolated vexans clades in Sweden and elsewhere in northern Europe. Considering COI data only, including sequences in GenBank from other parts of the world, they found four distinct groupings: “Group 1 vexans variant,” Ae. vexans nipponii, Ae. vexans North America and Ae. vexans Europe. Given a lack of morphological vouchers for study and weak branch support in some cases, Lilja et al. (2018) did not attempt to interpret the taxonomic meaning of their findings, recommending further work. These two studies generate many uncertainties, but they suggest that North American vexans could be a different species and that an unrecognized cryptic species occurs in Europe. Synonyms of Aedes (Aedimorphus) vexans (Meigen, 1830) (Wilkerson et al. 2021): Culex parvus Macquart, 1834 (type locality: Bordeaux [Gironde], France); Culex articulatus Rondani, 1872 (type locality: Italy); Culex malariae Grassi, 1898 (type locality: Italy); Culex sylvestris Theobald, 1901a (type locality: various, all in Canada) [montcalmi Blanchard, 1905 is incorrectly listed as a replacement name for sylvestris in Wilkerson et al. 2021, see Townsend (1990) for more information]; Culicada minuta Theobald, 1907 (type locality: India); Aedes eurochrus Howard, Dyar & Knab, 1917 (type locality: Popcum, British Columbia, Canada)., 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 16-17, DOI: 10.11646/zootaxa.5303.1.1, http://zenodo.org/record/8043342, {"references":["Patton, W. S. (1905) The culicid fauna of the Aden Hinterland, their haunts and habits. Journal of the Bombay Natural History Society, 16 (4), 623 - 637, 5 pls.","White, G. B. (1975) Notes on a catalogue of Culicidae of the Ethiopian Region. Mosquito Systematics, 7 (4), 303 - 344.","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. (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","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.","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","Meigen, J. W. (1830) Systematische Beschreibung der bekannten europaischen zweifl ¸ geligen Insekten. Vol. 6. Schulzische Buchhandlung, Hamm, iv + 401 pp., 12 pls. https: // doi. org / 10.5962 / bhl. title. 12464","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.","Becker, N., Petric, D., Zgomba, M., Boase, C., Madon, M., Dahl, C. & Kaiser, A. (2020) Mosquitoes: identification, ecology and control. 3 rd Edition. Springer Nature Switzerland AG, Cham, xxxi + 570 pp. https: // doi. org / 10.1007 / 978 - 3 - 030 - 11623 - 1","Krtinic, B., Francuski, L., Petric, E. & Milankov, V. (2013) Genetic diversity and differentiation between Palearctic and Nearctic populations of Aedimorphus (= Aedes) vexans (Meigen, 1830) (Diptera, Culicidae). Journal of Vector Ecology, 38 (1), 154 - 162. https: // doi. org / 10.1111 / j. 1948 - 7134.2013.12021. x","Lilja, T., Troell, K., Kirik, H. & Lindstr ˆ m, A. (2018) A distinct group of north European Aedes vexans as determined by mitochondrial and nuclear markers. Medical and Veterinary Entomology, 32 (3), 282 - 289. https: // doi. org / 10.1111 / mve. 12294","Macquart, J. (1834) Histoire naturelle des Insectes. Dipteres. Tome premier. Librairie Encyclopedique de Roret, Paris, 576 + 8 (explanation of plates) pp., 12 pls. https: // doi. org / 10.5962 / bhl. title. 14274","Rondani, C. (1872) Sulle specie Italiane del genere Culex Lin. Bollettino della Societa Entomologica Italiana, 4, 29 - 31.","Grassi, B. (1898) Rapporti tra la malaria e peculiari insetti (zanzaroni e zanzare palustri). Rendiconti delle seduti delle Reale Accademia dei Lincei. Classe de scienze fisiche, matematische e naturali, 7 (2), 163 - 177.","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","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","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.","Howard, L. O., Dyar, H. G. & Knab, F. (1917) The mosquitoes of North and Central America and the West Indies. Vol. 4. Systematic description (in two parts). Part II. Publication No. 159. Carnegie Institution of Washington, Washington, D. C., pp. 525 - 1064."]}

Aedes (Rusticoidus) rusticus (Rossi) subspecies rusticus (Rossi, 1790) —original combination: Culex rusticus. Distribution: Albania, Algeria, Austria, Belgium, Bulgaria, Crimean Peninsula, Croatia, Czech Republic, Denmark, Estonia? (see Kirik et al. 2022), France, Macedonia, Germany, Greece, Hungary, Ireland, Italy, Latvia, Lithuania, Luxembourg, Montenegro, Morocco, Netherlands, Poland, Portugal, Serbia, Slovakia, Spain, Sweden, Switzerland, Turkey, Ukraine, United Kingdom (Wilkerson et al. 2021, excluding Norway, see Mehl 1996). subspecies subtrichurus Martini, 1927 —original combination: Aedes subtrichurus (varietal status by Edwards 1932a; subspecific status by Harbach & Howard 2007). Distribution: Turkey (Martini 1927). Martini (1927) originally described subtrichurus as a distinct species of Aedes, but later (Martini 1931a) considered it to be a variety of Ae. diversus (Theobald, 1901c). It became a variety of Ae. rusticus when diversus was synonymized with that species (Edwards 1932a). Harbach & Howard (2007) recognized subtrichurus as a subspecies of Ae. rusticus because it was originally proposed as the valid name of a species, per Article 45.6.4.1 of the International Code of Zoological Nomenclature. Martini (1927) described Ae. subtrichurus based on specimens from the eastern end of the Gulf of Izmit (spelled as Ismid), located at the easternmost edge of the Sea of Marmara. Because this is the only record of subtrichurus, the locality lies within the wide distribution of Ae. rusticus and it is based merely on minor morphological differences, Aedes subtrichurus Martini, 1927 is hereby formally regarded as a synonym: subtrichurus Martini, 1927, junior subjective synonym of Aedes (Rusticoidus) rusticus (Rossi, 1790) . Consequently, “ Aedes subtrichurus ” should be removed from the list of species of Aedes in the Encyclopedia of Life. With the synonymy of subtrichurus, Ae. rusticus now includes seven junior synonyms. The previously recognized synonyms include Culex maculatus Meigen, 1804, Cx. musicus Leach, 1825, Cx. pungens Robineau-Desvoidy, 1827, Cx. quadratimaculatus Macquart, 1834, Cx. diversus Theobald, 1901c and Cx. nemorosus var. luteovittata Theobald, 1901c, all of which have type localities in Europe., 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 47-48, DOI: 10.11646/zootaxa.5303.1.1, http://zenodo.org/record/8043342, {"references":["Rossi, P. (1790) Fauna Etrusca: Sistens insecta quae in provinciis Florentina et Pisana praesertim collegit Petrus Rossius. Vol. 2. Thomae Masi & Sociorum, Liburni, 348 pp., 10 pls. https: // doi. org / 10.5962 / bhl. title. 15771","Kirik, H., Tummeleht, L. & Kurina, O. (2022) Rediscovering the mosquito fauna (Diptera: Culicidae) of Estonia: an annotated checklist with distribution maps and DNA evidence. Zootaxa, 5094 (2), 261 - 287. https: // doi. org / 10.11646 / zootaxa. 5094.2.3","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","Mehl, R. (1996) Culicidae Stikkmygg. In: Aagaard, K. & Dolmen, D. (Eds.), Limnofauna Norvegica: Katalog over norsk ferskvannsfauna. Tapir Forlag, Trondheim, pp. 205 - 203.","Martini, E. (1927) Uber zwei neue Stechm ¸ ckenarten aus Anatolien. Archiv f ¸ r Schiffs- und Tropenhygiene, 31, 386 - 390.","Edwards, F. W. (1932 a) Genera Insectorum. Diptera, Fam. Culicidae. Fascicle 194. V. Verteneuil & L. Desmet, Bruxelles [Brussels], 258 pp., 5 pls.","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","Martini, E. (1931 a) 11. u. 12. Culicidae. In: Lindner, E. (Ed.), Die Fliegen der palaearktischen Region. E. Schwizerbart'sche Verlagsbuchhandlung (Erwin Nagele) G. M. B. H., Stuttgart, pp. 1 - 398, 1 pl.","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","Meigen, J. W. (1804) Klassifikazion und Beschreibung der europaischen Zweifl ¸ geligen Insekten. (Diptera Linn.). Erster Band, ester Abtheilung, mit VIII Kupfertafeln. Karl Reichard, Braunschweig, xxviii + 152 pp., 8 pls.","Leach, W. E. (1825) Descriptions of thirteen species of Formica, and three species of Culex, found in the environs of Nice. Zoological Journal, 2 (7), 289 - 293.","Robineau-Desvoidy, J. - B. (1827) Essai sur la tribu des culicides. Memoires de la Societe d'Histoire Naturelle de Paris, 3, 390 - 413, 1 pl.","Macquart, J. (1834) Histoire naturelle des Insectes. Dipteres. Tome premier. Librairie Encyclopedique de Roret, Paris, 576 + 8 (explanation of plates) pp., 12 pls. https: // doi. org / 10.5962 / bhl. title. 14274"]}

Aedes (Neomelaniconion) lineatopennis (Ludlow) subspecies aureus Gutsevich, 1955 —original combination: Aedes (? Aedes) aureus (subspecific status by Danilov 1977). Distribution: Russia, Russian Federation (Wilkerson et al. 2021). subspecies lineatopennis (Ludlow, 1905) —original combination: Taeniorhynchus lineatopennis. Distribution: Australia, Bangladesh, Borneo (island), Cambodia, India, Indonesia, Laos, Malaysia, Nepal, Pakistan, People’s Republic of China, Philippines, Russia, South Korea, Sri Lanka, Thailand, Timor, Vietnam (Wilkerson et al. 2021). The taxonomic study of Huang(1985) provides a prelude to the treatment of these nominal taxa.Multiple identifications of lineatopennis in Africa have complicated the definition of this species. Huang, based on detailed morphological comparison of African specimens with type specimens and other material collected in the Philippines, determined that records of lineatopennis in Africa pertained to a previously unrecognized species, which she described and named Ae. mcintoshi. Since Huang excluded lineatopennis from Africa, all references to lineatopennis in Africa require re-evaluation (e.g. Ingram & de Meillon 1927; Edwards 1941; Gutsevich 1973). The nominotypical subspecies was described from two adult females captured at Camp Gregg, Bayambang, Pangasinan, Luzon, Philippines. Knight & Hull (1953) designated a lectotype, which was later validated by Stone & Knight (1956a). Its most distinctive features include the head with median golden scales, scutum dark brown with broad golden lateral stripes, legs unmarked, abdominal terga with pale basal bands and wings with pale scales on some veins. A condensed version of the original description (Ludlow 1905) follows. Taeniorhynchus lineatopennis, n. sp. — ♀. Head dark brown, with brassy yellow curved scales on median portion and extending from occiput to vertex... dark brown flat lateral scales, and a few forked scales... antennae dark brown... palpi... proboscis... clypeus dark brown.... Thorax: prothoracic lobes dark brown... no scales; mesonotum dark brown, the median portion covered with dark brown curved scales bordered by a heavy band of brassy yellow curved scales, extending cephalad from one wing joint (inverted “U”) across to the other, a very distinct and easily-recognized marking. …pleura brown and clothed only with a few brown hairs [setae]; scutellum dark brown, with brassy yellow curved scales.... Abdomen dark brown, with broad basal bands of “dirty white” scales hardly extending the full width of the terga; the first segment is dark, and the second has merely a median light spot, while on the ultimate segment the band is quite narrow; venter dark. Legs are brown throughout; coxae and trochanters and ventral side of femora somewhat lighter than the rest, a light spot near the apex of fore femora on dorsal side, i. e., the ventral colour runs up... more distal joints are darker, ranging from purplish to fawn colour.... Wings clear, clothed with brown and light typical Taeniorhynchus scales. The costa is dark throughout, the subcosta and first longitudinal [vein R 1] are mostly light scaled from the base of the wing to about the junction of the subcosta, and the stem of the fifth long vein [vein CuA] is also light, with some light scales on the lower fork. …halteres have a light stem and dark knob. Knight & Hull (1953) described the male genitalia and larva, Tanaka (2003) described the pupa and Choochote et al. (2001) described the egg of subspecies lineatopennis. Gutsevich (1955) described aureus from Kraskino, Russia, which is an urban locality in the Khasansky District of Primorsky Krai, located on the shore of Posyet Bay, 282 km southwest of Vladivostok, near the border with North Korea. If or where a type series was deposited is not known. An unpublished (1973) translation by B. F. Eldridge of the original Russian description follows. Aedes (Aedes ?) aureus Gutzevich, sp. n. Distinguishing features: Intense golden scales on lateral part of mesonotum. Large clearly outlined golden spot on occiput, presence of light scales on wings, irregularly shaped light spot on abdominal tergites [terga], entirely dark proboscis and tarsi. Female: Scales behind eye brown, abutting [eyes abutting? abutting eyes?]. Occiput with large spots of upright golden scales and hairs [setae]. Proboscis and palpi with unicolored brown scales; sometimes with middle third of proboscis having a single light scale. Proboscis as long as front femur. Length of antennae about 1/5 of length of proboscis. Antennae brown. Mesonotum with a longitudinal expanding [posteriorly?] stripe of chocolate-cinnamon scales. Lateral part of mesonotum with golden scales. Body of thorax devoid significantly of accumulated scales; the last forming a small spot on sternopleuron and mesepimeron. Bristles [Setae] (their arrangement and quantity play a part in the classification of the Oriental species of the subgenus Aedes): Proepimeral 6–8, parastigmatic [postspiracular] 5–7, upper mesepimeral 7–9, lower mesepimeral 3–5. Wing: costal vein covered with dark scales, subcosta mostly clear, cream-colored; base of radial, medial, and cubital vein with light scales, which vary among particular wings; anal vein with dark scales. Legs usually with dark scales; light longitudinal streak on posterior surface of femur, tibia, and first segment of tarsus. Tarsi without light rings. Claws equal on the anterior and middle tarsi and with teeth, the posterior tarsi without teeth. Abdomen from above usually with brown scales. Lighter yellowish-gray concentrated mainly in the middle and the anterior edge of the tergites, formed as indistinct spots. Abdominal hairs long, golden. Cerci very short. Length of body with proboscis 7–8 mm. Material (13 females) collected by K.P. Chaginem north of the Kraskino Primorskovo region in August 1947. Gutsevich (1973), after comparison of aureus with lineatopennis from South Africa [= Ae. mcintoshi Huang, 1985], determined that aureus belonged in Aedes subgenus Neomelanoconion, thus removing the initial doubt expressed in the original description. Danilov (1977) [summarized from a translation of the Russian] compared four aureus females from near the type locality with a female of lineatopennis from an undetermined [by us] locality in China (from the “IMP & TM collection”). To separate lineatopennis from aureus, Danilov determined that since there was a wide range of variation in the key character of the extent of basal pale scaling on the abdominal terga of putative aureus specimens, and also a wide range in the number of setae on the frontal stripe, that the characters were unreliable. He cited the wide distribution of lineatopennis, Afrotropical to South Korea, to conclude that aureus was simply a variant on the peripheral distribution of lineatopennis. He then decided, without explanation, that this distribution merited subspecies status for aureus. Of probable diagnostic significance, Danilov noted that aureus was much larger than lineatopennis. The larva and male genitalia of aureus were later described and illustrated by Shestakov (1980) from specimens collected near the type locality. To further our understanding of his description, we generated a rough translation of his paper using Google Translate. Most informative to us, however, are his illustrations of the head and terminal segments of the larva and the male genitalia. We compared Shestakov’s illustrations of the larva with the larva of lineatopennis illustrated by Mattingly (1961) and Knight & Hull (1953), and his illustrations of the male genitalia with the male genitalia of lineatopennis illustrated by Huang (1985). In the larval stage, the antenna of aureus has large spines on the inner surface but in lineatopennis the large spines are distributed over all surfaces. The siphon of aureus is short (index about 1.5) and widened medially whereas in lineatopennis it is longer (index about 2.5) with the widening more basal. The anal papillae of aureus are about 1.5 times the length of the saddle, gradually narrowing, while in lineatopennis they are about 2.5 times the length of the saddle, slender and tapered to a point. In the male genitalia, the stout subapical spines of the gonocoxite appear to be shorter and more numerous in aureus (the aureus illustration is a bit stylized so it is difficult to confidently compare them). Huang (1985) used differences in the nature of these spines to distinguish Ae. mcintoshi from lineatopennis. Also, tergum IX of aureus is somewhat rounded, while in lineatopennis it is slightly emarginate. The siphon of the lineatopennis larva illustrated by Lee (1999) and the siphon of aureus illustrated by Shestakov (1980) are both short and mesally expanded. Also, both nominal taxa exhibit peak activity in August and September (Shestakov 1980; Hwang et al. 2020). Since both are found in temperate climates in the same geographic region, we suspect that they are conspecific and the correct name for the form found in Korea is probably aureus, not lineatopennis. The name lineatopennis appears in all recent keys for the identification of mosquitoes that occur in the Republic of Korea (South Korea) (Lee & Egan 1985; Lee & Zorca 1987; Lee 1999; Ree 2003). The nominal taxon aureus has only been treated as a species by Gutsevich (1955, 1971, 1973, 1974) and Danilov (1977), and has only been designated and treated as a subspecies of lineatopennis by Danilov (1977). The nominal taxon lineatopennis apparently has a wide tropical distribution that extends from India to Australia. The nominal taxon aureus is only known from temperate areas ranging from the Korean peninsula to western Russia and probably China. In view of distinct ecological and morphological differences, we hereby restore subspecies aureus to species status: Aedes (Neomelanoconion) aureus Gutsevich, 1955 . Aedes aureus is currently listed as a species in the Encyclopedia of Life. Synonyms: None. Aedes (Neomelanoconion) lineatopennis (Ludlow, 1905) has one synonym: Pseudohowardina linealis Taylor, 1913. Synonymy by Taylor (1916)., 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 26-28, DOI: 10.11646/zootaxa.5303.1.1, http://zenodo.org/record/8043342, {"references":["Gutsevich, A. V. (1955) New and little known forms of mosquitoes (Diptera, Culicidae). Trudy Zoologicheskogo Instituta Akademii Nauk, 18, 320 - 324. [in Russian]","Danilov, V. N. (1977) On the synonymy of species names of Aedes mosquitoes (subgenera Finlaya and Neomelaniconion) in the Far East fauna. Prazitologiia, 11 (2), 181 - 184. [in Russian]","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","Ludlow, C. S. (1905) Mosquito notes. - No. 3. Canadian Entomologist, 37 (3), 94 - 102 + 129 - 135. https: // doi. org / 10.4039 / Ent 3794 - 3","Huang, Y. - M. (1985) A new African species of Aedes (Diptera: Culicidae). Mosquito Systematics, 17 (2), 108 - 120.","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.","Gutsevich, A. V. (1973) A subgenus of bloodsucking mosquitoes of the genus Aedes, Neomelaniconion Newst. (= Banksinella Theob.) (Diptera, Culicidae), new to the fauna of the Soviet Union. Entomological Review, 52 (1), 126 - 127.","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.","Stone, A. & Knight, K. L. (1956 a) Type specimens of mosquitoes in the United States National Museum: II, The genus Aedes (Diptera, Culicidae). Journal of the Washington Academy of Sciences, 46 (7), 213 - 228.","Tanaka, K. (2003) Studies on the pupal mosquitoes of Japan (7). Subgenera Aedimorphus, Geoskusea, Neomelaniconion and Neomacleaya of the genus Aedes (Diptera, Culicidae). Japanese Journal of Systematic Entomology, 9 (1), 11 - 28.","Choochote, W., Jitpakdi, A., Sukontason, K., Suntaravitun, T., Wongkamchal, S. & Pitsasawati, B. (2001). Scanning electron microscopy of Aedes lineatopennis (Diptera: Culicidae) eggs. Journal of Medical Entomology, 38 (5), 753 - 755. https: // doi. org / 10.1603 / 0022 - 2585 - 38.5.753","Shestakov, B. I. (1980) Description of a male and larva of mosquito Aedes aureus Guts. (Diptera, Culicidae) from the Maritime Territory. Taksonomiia Nasekomykh Dal'nego Vostoka / [red. N. V. Kurzenko, P. A. Ler, V. S. Fedikova], 1980, 115 - 117. [in Russian]","Mattingly, P. F. (1961) The culicine mosquitoes of the Indomalayan Area. Part V. Genus Aedes Meigen, subgenera Mucidus Theobald, Ochlerotatus Lynch Arribalzaga and Neomelaniconion Newstead. British Museum (Natural History), London, 62 pp.","Lee, K. W. (1999) A revision of the illustrated taxonomic keys to genera and species of mosquito larvae of Korea (Diptera, Culicidae). 5 th Medical Detachment, 168 th Medical Battalion, 18 th Medical Command, U. S. Army, Korea, APO AP 96205 - 0020, i - viii + 1 - 33.","Hwang, M. - J., Kim, H. - C., Klein, T. A., Chong, S. - T., Kisung Sim, K., Chung, Y. & Cheong, H. - K. (2020) Comparison of climatic factors on mosquito abundance at US Army Garrison Humphreys, Republic of Korea. PLoS ONE, 15 (10), e 0240363. https: // doi. org / 10.1371 / journal. pone. 0240363","Lee, K. W. & Egan, J. P. (1985) Illustrated taxonomic keys to genera and species of female mosquitoes of Korea. Part I. Department of the Army, 5 th Preventive Medicine Unit, 18 th Medical Command, APO San Francisco, 96301, i + 1 - 32.","Ree, H. - I. (2003) Taxonomic review and revised keys of the Korean mosquitoes (Diptera: Culicidae). Korean Journal of Entomology, 33 (1), 39 - 52. https: // doi. org / 10.1111 / j. 1748 - 5967.2003. tb 00047. x","Gutsevich, A. V., Monchadsky, A. S. & Stackelberg, A. A. (1971) Fauna Diptera. Vol. III. No. 4. Mosquitoes, Family Culicidae. New Series 100. Academy of Sciences of the USSR Zoological Institute, Leningrad, 384 pp. [for 1970, in Russian]","Taylor, F. H. (1913) Report of the entomologist. Report of the Australian Institute of Tropical Medicine, 1911, 49 - 74, 3 pls.","Taylor, F. H. (1916) Contributions to a knowledge of Australian Culicidae. No. iii. Proceedings of the Linnean Society of New South Wales, 41 (3), 564 - 574. https: // doi. org / 10.5962 / bhl. part. 15324"]}

Anopheles (Anopheles) parapunctipennis Martini subspecies guatemalensis de León, 1938 —original combination: Anopheles (Anopheles) chiriquiensis var. guatemalensis (subspecific status by Wilkerson 1990). Distribution: Guatemala (Wilkerson 1990). subspecies parapunctipennis Martini, 1932 —original combination: Anopheles parapunctipennis. Distribution: Guatemala, Mexico (Wilkerson 1990). The nominotypical subspecies belongs to the Pseudopunctipennis Group (Reid & Knight 1961). Most members of the group are large, high elevation species which share a number of distinctive morphological characters (see the treatment of An. pseudopunctipennis Theobald below). Wilkerson (1990) established that a synonym of An. parapunctipennis, i.e. Anopheles (Anopheles) chiriquiensis Komp, 1936, was a valid species. However, he left guatemalensis as a subspecies of parapunctipennis and stated that a single character in the female (but not the male) separated it from the nominotypical species: Vein R 1 dark at the subcostal pale spot, pale in parapunctipennis. The type locality of the nominotypical subspecies is San Cristóbal de las Casas, Chiapas Mexico (coordinates: 16.73176, -92.64126) and the type locality of subspecies guatemalensis is Cumbre del Aire, Department of Totonicapán, Guatemala (coordinates: 15.11667, -91.56667). The two localities are only 213 km apart in the same mountain range (coordinates and distance: D. Pecor pers. comm.). Given there is insufficient morphological indication that these nominal forms are genetically distinct, and that the type localities are geographically close enough to assume sympatry, we hereby recognize guatemalensis as a synonymous name: guatemalensis de León, 1938, junior subjective synonym of Anopheles (Anopheles) parapunctipennis Martini, 1932 . The nominal guatemalensis, which is listed as a species in the Encyclopedia of Life, must be removed from the list of valid species of Anopheles., 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 58, DOI: 10.11646/zootaxa.5303.1.1, http://zenodo.org/record/8043342, {"references":["de Leon, J. R. (1938) El anophelismo de altura en Guatemala. Boletin de la Direccion General de Sanidad Publica (Guatemala), 9, 411 - 424.","Martini, E. (1932) Dos nuevos mosquitos Anopheles, procedentes del Estado de Chiapas, Mexico. Revista Mexicana de Biologia, 12, 99 - 102.","Reid, J. A. & Knight, K. L. (1961) Classification within the subgenus Anopheles (Diptera, Culicidae). Annals of Tropical Medicine and Parasitology, 55 (4), 474 - 488. https: // doi. org / 10.1080 / 00034983.1961.11686077","Komp, W. H. W. (1936) Anopheles (Anopheles) chiriquiensis, a new species of Anopheles from Panama (Diptera, Culicidae). Proceedings of the Entomological Society of Washington, 38 (7), 156 - 160."]}

Anopheles pseudopunctipennis sensu lato in Ecuador and Colombia Two varieties of An. pseudopunctipennis were described from Ecuador by Leví Castillo, both currently subspecies of pseudopunctipennis. Anopheles pseudopunctipennis levicastilloi Levi Castillo, 1944, type locality: Guayas Province, and An. pseudopunctipennis rivadeneirai Leví Castillo, 1945, type locality: Cited as having been found in the provinces of Carchi, Imbabura, Pichincha, Cotopaxi, Tungurahua, Chimborazo, Cañar, Azuay and Loja. An adequately detailed map of collection records for the two varieties is in Leví Castillo (1945). The following is a brief summary [paraphrased translation of selected text from Spanish] from that publication. In Ecuador, there are two varieties in the Pseudopunctipennis Complex, completely different from each other; one coastal, An. pseudopunctipennis var. levicastilloi, and one montane, An. pseudopunctipennis var. rivadeneirai. The former is relatively small, wing length about 4 mm, the latter is the largest anopheline in Ecuador, wing length about 5 mm. The eggs of the two varieties are the best way to recognize them. Coastal levicastilloi eggs are more characteristic [of the genus Anopheles], boat-shaped with floats that give the appearance of a wasp nest [no collar is mentioned or apparent in the illustration]. The montane rivadeneirai egg is boat-shaped with the appearance of a grain of wheat, dorsocentrally concave with rounded ends, but lacks floats. Instead of floats there are many vacuoles filled with an unknown substance. Female. Wing vein R 2 in levicastilloi has a median pale spot, in rivadeneirai vein R 2 is completely dark. Coastal levicastilloi is not involved in malaria transmission while the montane rivadeneirai is the principal vector of malaria in the warm valleys of the Ecuadorian Andes. This distinction of lowland and highland Ecuadorian subspecies (as varieties) was also noted by Pinault & Hunter (2011), who collected Anopheles extensively in both ecological/altitudinal areas. They found An. pseudopunctipennis sensu lato in both lowland and montane areas but did not recognize or mention the two varieties of Leví Castillo. They described, however, finding pseudopunctipennis at very different altitudes and in different climates. In a comprehensive study of Anopheles in western Colombia using COI barcode sequence, this lowlandhighland distribution of An. pseudopunctipennis sensu lato was also reported by Ahumada et al. (2016). They found highly supported groups representing northwestern and southern Pacific coastal populations, but only referred to them as “s.l.” and did not compare their sequences with sequences from the type locality of An. pseudopunctipennis in Granada. We believe that Leví Castillo clearly described two species that differ from An. pseudopunctipennis sensu stricto, and each other, both geographically and morphologically. Furthermore, we think that Pinault & Hunter (2011) and Ahumada et al. (2016) studied these same species in Ecuador and Colombia. Accordingly, we elevate both to species status: Anopheles (Anopheles) levicastilloi Levi-Castillo, 1944 and Anopheles (Anopheles) rivadeneirai Levi-Castillo, 1945 . Further studies are needed to clarify their overall distributions. Anopheles levicastilloi and An. rivadeneirai 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 pages 60-61, DOI: 10.11646/zootaxa.5303.1.1, http://zenodo.org/record/8043342, {"references":["Levi Castillo, R. (1944) El complejo \" Pseudopunctipennis \" en el Ecuador (Diptera - Culicidae). Imprenta de la Universidad, Guayaquil, 7 pp.","Levi Castillo, R. (1945) Los anofelinos de la Republica del Ecuador. Tomo Primero. Artes Graficas Senefelder, Guayaquil, Ecuador, vi + 72 pp.","Pinault, L. L. & Hunter, F. F. (2011) New highland distribution records of multiple Anopheles species in the Ecuadorian Andes. Malaria Journal, 10, 236. https: // doi. org / 10.1186 / 1475 - 2875 - 10 - 236","Ahumada, M. L., Orjuela, L. I., Pareja, P. X., Conde, M., Cabarcas, D. M., Cubillos, E. F. G., Lopez, J. A., Beier, J. C., Herrera, S. & Quinones, M. L. (2016) Spatial distributions of Anopheles species in relation to malaria incidence at 70 localities in the highly endemic Northwest and South Pacific coast regions of Colombia. Malaria Journal, 15, 407. https: // doi. org / 10.1186 / s 12936 - 016 - 1421 - 4"]}

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."]}

Aedes (Stegomyia) annandalei (Theobald) subspecies annandalei (Theobald, 1910 a)—original combination: Stegomyia annandalei. Distribution: Bangladesh, Cambodia, India, Indonesia, Myanmar, Nepal, Papua New Guinea, People’s Republic of China, Taiwan, Thailand, Vietnam (Wilkerson et al. 2021). subspecies quadricinctus (Barraud, 1923b) —original combination: Stegomyia annandalei var. quadricincta (subspecific status by Harbach & Howard 2007). Distribution: India (Barraud 1923b). The taxonomic history of Ae. annandalei involves three nominal forms: Stegomyia annandalei Theobald, 1910 a, Aedes horishensis Yamada, 1921 and Stegomyia annandalei var. quadricincta Barraud, 1923b. Aedes horishensis, described from a single female collected at or near Horisha in present-day Taiwan, was recognized as a subspecies of annandalei by Lien (1962). Its taxonomic status thereafter is a matter of confusion. Mattingly (1965) listed it as a synonym of annandalei, but also indicated that he considered it to be a variety: “I agree with Lien that the evidence no longer warrants this being treated as a distinct species. However, the apparently low incidence of this type of marking [slight posterior shifting of pale bands on abdominal terga IV–VI] in Taiwan and its simultaneous occurrence in Indonesia, in both cases in areas where normal markings are the rule, seems to me to render his suggestion that it should be treated as a subspecies unacceptable. Since this character reappears in the Ae. scutellaris complex and is, therefore of particular interest, I consider that specimens showing it should be distinguished as var. horishensis Yamada. ” Likewise, Huang (1977) also made conflicting statements about the status of horishensis. In the abstract of her publication, she stated that “ Aedes horishensis Yamada is regarded as a variety of annandalei Theobald ”, whereas in her discussion of annandalei, she sided with Mattingly in noting that the posterior shifting of the abdominal bands is variable, and stated “Therefore, I consider horishensis Yamada to be a synonym of annandalei.” Despite these contradictions, horishensis was listed as a synonym of annandalei by Knight & Stone (1977), attributed to Mattingly (1965), Harbach (2018) and Wilkerson et al. (2021), attributed to both Mattingly (1965) and Huang (1977). Lu et al. (1997) agreed with the synonymy. Based on the findings and actions taken by Mattingly (1965), Huang (1977) and Lu et al. (1997), we agree that horishensis is probably conspecific with annandalei and should continue to be recognized as a synonymous nominal taxon. Barraud (1923b) described Stegomyia annandalei var. quadricincta based on “One female from Nongpoh, Assam, July, 1922 (Barraud).” He noted there was “another female specimen from the same place which agrees with the above [description of quadricincta] in the markings of the hind tarsi, but the fore and mid legs have only two rings, as in the type form.” This is obviously an indication that the two forms were sympatric. Barraud (1934) more explicitly described the distinctive tarsal banding of quadricinctus as follows: “Differs from the type-form in having basal white markings to first four tarsal segments on all legs; markings at base of segments 3 and 4 on fore and mid-legs small and not forming complete rings; complete white rings on tarsal segments 1–4 on hind legs, that on 4 occupying nearly whole segment.” Concomitant with the recognition of Stegomyia as a subgenus of Aedes by Edwards (1932a), the variety described by Barraud (1923b) became known as Aedes (Stegomyia) annandalei var. quadricinctus, and was subsequently listed as such in the catalogs of Stone et al. (1959) and Knight & Stone (1977). Mattingly (1965) and Huang (1977) both listed quadricinctus as a synonym of annandalei without explicit mention or discussion of its taxonomic status. Mattingly, however, noted that “Var. quadricinctus Barraud … has white rings on the first 4 tarsi [tarsomeres] of all legs. An otherwise similar ♀ from the type locality had only the first 2 segments [tarsomeres] banded on the fore and mid legs. I have not seen this type of variation in any Indomalayan material.” Perplexingly, Huang (1977) vaguely (doubtfully?) indicated that the tarsi of annandalei may sometimes exhibit the condition described for quadricinctus, i.e. that tarsomeres 1–4 of all legs have white bands. She described the tarsi of males as having the “fore- and midtarsi with basal white band on tarsomere 1, sometimes midtarsus with a few white scales on basal area of tarsomere 2 as well; hindtarsus with basal white bands on tarsomeres 1,2; tarsomere 3 dark [emphasis ours]; tarsomere 4 with basal 0.67 white band to all dark…”. She noted females are essentially the same except that “Foretarsomere 2 sometimes with a few white scales on basal area; midtarsomere 2 with basal white band; hindtarsomere 4 with basal 0.83 white band; sometimes hindtarsomere 3 [emphasis ours] with a few white scales on basal area as well…”. The tacit synonymy of quadricinctus with annandalei published by Huang (1977) was not published in the world catalog of the Culicidae of Knight & Stone (1977) because the cut-off date for additions to the catalog was the end of 1973, but neither was it recorded in the three subsequent catalog supplements (Knight 1978; Ward 1984, Ward 1992). Unaware of the synonymy by Huang (1977), Harbach & Howard (2007) followed Knight & Stone in recognizing quadricinctus as a variety, which they deemed to be subspecific under provisions of Article 45.6.4 of the International Code of Zoological Nomenclature. Despite knowing that the leg markings of annandalei are subject to variation (Barraud 1923b; Mattingly 1965; Huang 1977) and recognizing that “Although it is likely that the name quadricincta applies to an infrasubspecific form…”, Harbach & Howard were obligated to reveal that quadricinctus “officially has subspecific rank” under provisions of the Code. In view of the earlier synonymy by Huang (1977), the change of rank from variety to subspecies was unnecessary and unsupportable. A number of different DNA sequences are available in GenBank for specimens of Ae. annandalei from China and Singapore, but until molecular data become available for specimens of quadricinctus from its type locality in India, we must agree with Mattingly (1965) and Huang (1977) that quadricinctus is merely a morphological variant and synonym of Ae. annandalei. To avoid any doubt, quadricincta is hereby formally regarded as a synonymous name: quadricincta Barraud, 1923b, junior subjective synonym of Aedes (Stegomyia) annandalei (Theobald, 1910 a) . Consequently, “ Aedes quadricinctus ” should be removed from the list of species of Aedes included 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 51-52, DOI: 10.11646/zootaxa.5303.1.1, http://zenodo.org/record/8043342, {"references":["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","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","Barraud, P. J. (1923 b) A revision of the culicine mosquitoes of India. Part V. Further notes on the genera Stegomyia, Theo. and Finlaya, Theo. with descriptions of new species. Indian Journal of Medical Research, 11 (1), 224 - 228, 3 pls.","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","Yamada, S. (1921) Description of ten new species of Aedes found in Japan, with notes on the relation between some of these mosquitoes and the larva of Filaria bancrofti Cobbold. Annotationes Zoologicae Japonenses, 10 (6), 45 - 81.","Lien, J. C. (1962) Non-anopheline mosquitoes of Taiwan: Annotated catalog and bibliography. Pacific Insects, 4 (3), 615 - 649.","Mattingly, P. F. (1965) The culicine mosquitoes of the Indomalayan area. Part VI. Genus Aedes Meigen, subgenus Stegomyia Theobald (Groups A, B and D). British Museum (Natural History), London, 67 pp.","Huang, Y. - M. (1977) Medical entomology studies - VII. The subgenus Stegomyia of Aedes in Southeast Asia. II - The edwardsi group of species. III - The w-albus group of species. (Diptera: Culicidae). Contributions of the American Entomological Institute, 14 (1), 1 - 111.","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.","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","Lu, B., Li, B., Ji, S., Chen, H., Meng, Q., Su, L., Qu, I. Gong, Z. & Zhang, Z. (1997) Fauna Sinica, Insecta. Vol. 8. Diptera: Culicidae 1. Science Press, Beijing, xiii + 884 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.","Edwards, F. W. (1932 a) Genera Insectorum. Diptera, Fam. Culicidae. Fascicle 194. V. Verteneuil & L. Desmet, Bruxelles [Brussels], 258 pp., 5 pls.","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. (1978) Supplement to a catalog of the mosquitoes of the world (Diptera: Culicidae). Thomas Say Foundation, Supplement to Vol. VI. Entomological Society of America, College Park, Maryland, (iii) + 107 pp.","Ward, R. A. (1984) Second supplement to \" A catalog of the mosquitoes of the world \" (Diptera: Culicidae). Mosquito Systematics, 16 (3), 227 - 270.","Ward, R. A. (1992) Third supplement to \" A catalog of the mosquitoes of the world \" (Diptera: Culicidae). Mosquito Systematics, 24 (3), 177 - 230."]}

Anopheles (Cellia) moucheti Evans subspecies moucheti Evans, 1925b —original combination: Anopheles marshalli var. moucheti (specific status by Evans 1931). Distribution: Burundi, Central African Republic, Côte d’Ivoire, Democratic Republic of the Congo, Equatorial Guinea, Gabon, Ghana, Kenya, Republic of the Congo, Rwanda, Sierra Leone, South Sudan, Tanzania, Uganda (Wilkerson et al. 2021, Cameroon, Guinea and Nigeria are excluded). subspecies nigeriensis Evans, 1931 —original combination: Anopheles moucheti var. nigeriensis. Distribution: Cameroon, Guinea, Niger, Nigeria (Wilkerson et al. 2021). Anopheles moucheti was originally described as a variety of An. marshallii (Theobald, 1903a) and elevated to species status by Evans (1931). The taxon was described from a series of adult males and females from the Belgium Congo, present-day Democratic Republic of the Congo. The holotype male was collected at Buta, which today is a city and capital of Bas-Uiele Province located in the northern area of the country. When Evans recognized moucheti as a distinct species, she also described nigeriensis as a variety of the species based on a series of syntypes (“co-types”), including one male and two females with associated larval and pupal exuviae, collected at Yaba, a suburb of Lagos, Lagos State in southern Nigeria. Townson (1990) verified the presence of nine syntypes in the Natural History Museum, London— two females from Yaba, and two males, two females and three larvae from near Lagos. Gillies & de Meillon (1968) and Gillies & Coetzee (1987) did not distinguish the adults of moucheti sensu stricto and nigeriensis in their keys to the Anopheles of the Afrotropical Region; however, they did distinguish the larvae based on the development of head seta 3-C—with three or more branches in moucheti sensu stricto and single, occasionally bifid in nigeriensis. The complete chaetotaxy of the larvae, and also the pupae, has not been 8studied comparatively. Gillies & de Meillon (1968) specified that nigeriensis “Differs from the nominate subspecies only in the following characters: Pharynx [cibarium]: Spines on pediment of cone apparently rather longer. Wing: In a series of specimens from Lagos, reared from eggs and with associated larvae from the same eggbatch (C. D. Ramsdale), a 6th pale fringe spot is present in 4 out of 5 females. Outer clypeal hairs [seta 3C]: Simple or bifid apically. We have not been able to confirm Evans’s description of the mesonotal hairs as being narrower than in the type form.” The taxonomic history of An. moucheti (Moucheti Complex, Brunhes et al. 1998 b) involves the closely related An. bervoetsi D’Haenens, 1961, which was originally described as a subspecies of moucheti. It was afforded specific status by Gillies & de Meillon (1968) based on distinctive features of the larva, and returned to subspecific status by Brunhes et al. (1998 b), who also considered nigeriensis to be a synonym of moucheti: “All of these [published] observations lead us, at least while awaiting new information, to consider An. moucheti as a polymorphic species and An. moucheti nigeriensis as a synonym of An. moucheti Evans, 1925 [translated from the French].” In agreement with Gillies & de Meillon, bervoetsi was reinstated as a separate species by Antonio-Nkondjio et al. (2008) based on high levels of differentiation between 10 microsatellite loci of non-coding DNA. Kengne et al. (2007) investigated the use of the mitochondrial cytochrome oxidase b (CytB) gene and the ribosomal internal transcribed spacers ITS1 and ITS2 to distinguish bervoetsi, moucheti sensu stricto and nigeriensis, which were recognized as subspecies of moucheti at the time. The results of their study suggested that moucheti sensu stricto occurs in the forested areas of central Africa while bervoetsi and nigeriensis are only found in areas of the Democratic Republic of the Congo and Nigeria, which encompass their type localities, respectively. For ITS1, the mean genetic distance was found to be greater between nigeriensis and bervoetsi (0.166) than between moucheti sensu stricto and bervoetsi (0.141), and lower between moucheti sensu stricto and nigeriensis (0.084). The authors stated that “This degree of differentiation between morphological forms is relatively high, indicating significant genetic divergence among the three taxa, which is comparable with that among species.” In comparison, the ITS2 sequences exhibited a lower degree of divergence, ranging from 0.03 between moucheti sensu stricto and nigeriensis to 0.063 between moucheti sensu stricto and bervoetsi. Sequence differences included 30 fixed indel or substitutions between moucheti sensu stricto and bervoetsi, 29 between nigeriensis and bervoetsi, and 16 between moucheti sensu stricto and nigeriensis. For the CytB gene, 26 fixed differences in the first codon position were identified among the three forms. The CytB gene sequences, in agreement with the ITS2 sequences, showed consistent but low divergence of 0.035 –0.047 between the three forms. The authors concluded that “The concordance of genetic variability of the three markers and the lack of intraspecific geographical variation suggests that the three members of the group are true species.” In summary they stated that “The three genomic regions revealed sequence differences between the three morphological forms similar in degree to the differences shown previously for members of other anopheline species groups or complexes (genetic distance d = 0.047–0.05 for CytB, 0.084 –0.166 for ITS1 and 0.03–0.05 for ITS2).” In view of the genetic evidence, we are suprised that Kengne et al. did not formally elevate nigeriensis to specific rank. But having said that, the taxonomic status of nigeriensis was not relevant to the objective of their study, which was to elucidate molecular means for distinguishing and identifying the three nominal forms. Despite the available morphological, molecular and distribution data, nigeriensis continued to be recognized as a subspecies (Harbach 2018; Coetzee 2020; Irish et al. 2020; Wilkerson et al. 2021). In fact, nigeriensis as a separate entity has largely been ignored, e.g., in the recent list of countries in which Afrotropical species of Anopheles have been recorded (Irish et al. 2020), the authors stated “As An. nigeriensis refers to a subspecies of An. moucheti, it is not included here.” In view of the morphological, distributional and in particular the molecular distinctions detailed above, we are compelled to formally recognize nigeriensis as a separate species of the Moucheti Complex: Anopheles (Cellia) nigeriensis Evans, 1931 . Anopheles nigeriensis is not currently included as a species in the Encyclopedia of Life and should be added to the list of species of the genus. As currently understood, the Moucheti Complex includes three species, bervoetsi D’Haenens, moucheti Evans and nigeriensis Evans., 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 69-70, DOI: 10.11646/zootaxa.5303.1.1, http://zenodo.org/record/8043342, {"references":["Evans, A. M. (1925 b) A new variety of Anopheles marshalli Theobald from the Belgian Congo. Annals of Tropical Medicine and Parasitology, 19 (2), 211 - 213. https: // doi. org / 10.1080 / 00034983.1925.11684451","Evans, A. M. (1931) Notes on African anophelines. Annals of Tropical Medicine and Parasitology, 25 (1), 129 - 143. https: // doi. org / 10.1080 / 00034983.1931.11684674","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. (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","Gillies, M. T. & de Meillon, B. (1968) The Anophelinae of Africa south of the Sahara (Ethiopian Zoogeographical Region). Publications of the South African Institute for Medical Research, 54, 1 - 343.","Gillies, M. T. & Coetzee, M. (1987) A supplement to the Anophelinae of Africa south of the Sahara (Afrotropical Region). Publications of the South African Institute for Medical Research, 55, 1 - 143.","Brunhes, J., Le Goff, G., Manga, L. & Geoffroy, B. (1998) Anopheles afrotropicaux. - IV. Mises au point sur le complexe Anopheles moucheti, rehabilitation d' An. garnhami basilewskyi (Diptera: Culicidae). Annales de la Societe Entomologique de France, 34 (4), 397 - 405.","Antonio-Nkondjio, C., Ndo, C., Kengne, P., Mukwaya, L., Awono-Ambene, P., Fontenille, D. & Simard, F. (2008) Population structure of the malaria vector Anopheles moucheti in the equatorial forest region of Africa. Malaria Journal, 7, 120. https: // doi. org / 10.1186 / 1475 - 2875 - 7 - 120","Kengne, P., Antonio-Nkondjio, C., Awono-Ambene, H. P., Simard, F., Awolola, T. S. & Fontenille, D. (2007) Molecular differentiation of three closely related members of the mosquito species complex, Anopheles moucheti, by mitochondrial and ribosomal DNA polymorphism. Medical and Veterinary Entomology, 21 (2), 177 - 182. https: // doi. org / 10.1111 / j. 1365 - 2915.2007.00681. x","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","Coetzee, M. (2020) Key to the females of Afrotropical Anopheles mosquitoes (Diptera: Culicidae). Malaria Journal, 19, 70. https: // doi. org / 10.1186 / s 12936 - 020 - 3144 - 9","Irish, S. R., Kyalo, D., Snow, R. W. & Coetzee, M. (2020) Updated list of Anopheles species (Diptera: Culicidae) by country in the Afrotropical Region and associated islands. Zootaxa, 4747 (3), 401 - 449. https: // doi. org / 10.11646 / zootaxa. 4747.3.1"]}

Aedes (Ochlerotatus) canadensis (Theobald) subspecies canadensis (Theobald, 1901c) —original combination: Culex canadensis. Distribution: Canada, Dominican Republic, Mexico, United States (Alaska and continental) (Wilkerson et al. 2021). subspecies mathesoni Middlekauff, 1944 —original combination: Aedes (Ochlerotatus) mathesoni (subspecific status by Rings & Hill 1948). Distribution: Southeast United States (Alabama, Florida, Georgia, South Carolina) (Rings & Hill 1948). The nominotypical subspecies was described from specimens collected near Toronto, Canada. Theobald (1901c) stated: “A very distinct species, in which the legs are very characteristically marked, the last tarsal joint [tarsomere 5] of the hind legs being entirely dull white and the banding of the legs involving both sides of the joints. The dusky scaled abdomen, with the creamy-white basal lateral patches, is also very characteristic.” Additionally we note: Maxillary palpus with apical half of palpomere 2 pale; scutum dull purplish brown to deep chestnut brown, covered with varying patterns of curved golden-brown scales; wing all dark; and venter of abdomen covered with pale yellowish scales. In the Natural History Museum, London, Townsend (1990) found: “ Lectotype male, designated by Belkin, 1968b: 4 ‒ Canada: Ontario, near Toronto, Trout Creek. Paralectotypes (14) ‒ Canada: 1 male, 13 female.” “Belkin misread the type locality as ‘Front Creek’, and overlooked some of the paralectotypes.” Other pertinent descriptions include: Adult female, larva and male genitalia (Carpenter & LaCasse 1955); larva and male genitalia (Ross 1947); pupa (Darsie 1951); and egg (Craig 1956; Craig & Horsfall 1958; Ross & Horsfall 1965; Kalpage & Brust 1968; Horsfall et al. 1970). Notably, canadensis is cold adapted, emerging very early in the spring (reviewed by Horsfall 1972). Notable characters of the larva include: Setae 1-A and 5–7-C multi-branched, 5–7-C not in a straight line, 6-C inserted far forward of 5-C; seta 1-M small and short, not reaching bases of prothoracic setae; comb with many scales in a patch, individual scales pointed and fringed with rather slender subequal spinules; siphon index 3.0–4.0, pecten with 13–24 evenly spaced spines on basal two-fifths of siphon, siphonal tuft (seta 1-S) usually 3–8-branched, inserted beyond pecten; saddle incomplete, extending about 0.67 down side of segment, ventral brush (seta 4-X) large, usually with 2 precratal setae; anal papillae tapered, about 1.5 times as long as saddle (Carpenter & LaCasse 1955; Harrison et al. 2016). Subspecies mathesoni was described, as a species, by Middlekauff (1944) from specimens collected in south-central Florida (Kissimmee) in late summer and autumn [early season occurrence not noted]. Middlekauff recognized mathesoni as quite similar to canadensis: “This species [mathesoni] is most closely related to Aedes canadensis (Theob.), but that species [canadensis] lacks the nearly black scales of the mesonotum and the white areas on the legs are more extensive. The male genitalia of canadensis differ only slightly, the spines of the ninth tergites [terga] being less regularly placed and more abundant, and the tenth sternites [paraprocts] being somewhat curved and acutely [sic] apically.” “ Holotype, allotype and one paratype [are] in the collection of the United States National Museum.” Rings & Hill (1946) described the larva of mathesoni using reared-associated specimens from Georgia (Camp Gordon) and from northeastern Florida (Camp Blanding). They found the larvae of canadensis and mathesoni “to be remarkably similar, the only apparent difference being in the degree of branching in the head hairs [setae] and antennal tufts [seta 1-A].” In general, mean branch counts for setae 5- and 6-C were lower in specimens from further north (canadensis) than southern specimens. They counted setal branches of 89 canadensis in places ranging from British Columbia to Florida and 33 mathesoni from Georgia and Florida: Seta 5-C, mean number of branches 6.77 (4–10) (canadensis), 9.8 (6–14) (mathesoni); seta 6-C 5.06 (range not given) (canadensis), 7.1 (4–9) (mathesoni). Rings & Hill compared specimens of mathesoni from Georgia and Florida and found the same tendency: “... individuals collected in the more southern portions of the geographical range showed relatively more branching than those from the north.” Seta 5-C, mean 8.9 (Georgia), 10.7 (Florida); seta 6-C, 7.0 (Georgia), 7.2 (Florida). We think these numbers are suggestive of clinal variation. Rings & Hill concluded: “...there is a very close phylogenetic relationship between the two forms and that these forms do not possess differences of specific rank.” Rings & Hill (1948) documented and illustrated the scutal patterns of canadensis and mathesoni and noted that there were many “intergrades”. They stated: “The data here presented are interpreted by the authors as evidence that mathesoni is a melanistic, geographical variation of A. canadensis.” They illustrated intergrades showing the extent of pale scaling on the hindtarsomeres and mapped the geographical extent of intergradation and showed a broad area of intergrades. Those north and south of this zone were easily identified as either canadensis or mathesoni, respectively. Darker coloration seen on the legs was explained as directly related to higher temperatures at decreasing latitudes. They found little or no differences (intergrades) in the larvae or male genitalia of the two forms. The notion that mathesoni was a subspecies of canadensis was introduced in this paper; however, their unattributed definition of subspecies allows for sympatry (which for us precludes subspecific status). They concluded: “The phylogenetic rank of subspecies is generally applied to a part of a species showing geographical variations which intergrade with a subspecies occupying different though usually adjacent and overlapping parts of the general range of the species.” They, however, continued to use the subspecies rank as a convention for pointing out observed variability. Bickley (1981) furthered the observations of Rings & Hill (1946, 1948) in an analysis of adults and larvae of more than 1,400 specimens in the United States National Museum (Smithsonian Institution, National Museum of Natural History (USNM), Washington, D.C.). He concluded: “From the survey described here, involving specimens from 43 North American states and provinces, it may be concluded that the geographical range of Ae. canadensis mathesoni does not extend northward from southern Georgia and South Carolina. Only 1 character, the narrow pale rings on the hindtarsomeres, is available as a means of recognizing this rare subspecies, the validity of which is certainly complicated by the existence of intergrades.” Harrison et al. (2016) also used the pale bands of the hindtarsomeres to separate canadensis canadensis from canadensis mathesoni, but they could not separate them in the larval stage. They stated (Note 14): “The two subspecies of Ae. canadensis are separated in the adult female key, but are not separated in the larval key. Those subspecies, Ae. canadensis canadensis and Ae. canadensis mathesoni represent a real taxonomic enigma that begs to be resolved with more study and the use of modern techniques.” An explanation for this enigma may lie in the abundant examples of mosquito morphological and physiological variation influenced by the environment, for example: Factors affecting larval setal branching (Colless 1956; Mattingly 1975); altitude and latitude influencing diapause and the size of anal papillae in Wyeomyia smithii (Coquillett, 1901) (Bradshaw & Lounibos 1977); season and temperature affecting melanism in species of the genus Anopheles (Harrison 1980); and the effect of temperature on morphological attributes of Anopheles merus (D̂nitz, 1902) (Le Sueur & Sharp 1991). Ongoing sampling of mosquitoes at several locations in North America is being carried out by the National Ecological Observatory Network (NEON). Their sampling is based on light trapping of adult females, which are then identified morphologically. A subsample is kept as morphological and DNA vouchers, with single legs used for sequencing of the barcoding region of the mtDNA COI gene. The NEON database (https://biorepo.neonscience. org/portal/collections/list.php?usethes=1&taxa=80835) has 571 records for canadensis canadensis (also written simply as canadensis) and canadensis mathesoni. The database includes a limited number of sympatric records of nominotypical canadensis with canadensis mathesoni in the area of historical “intergrades,” i.e. Florida Disney Wilderness Preserve near Kissimmee and near Gainesville, Florida, which is relatively close to Camp Blanding. The Barcode of Life Data System (BOLD: boldsystems.org) includes a single entry for canadensis mathesoni (http://www.boldsystems.org/index.php/Taxbrowser_Taxonpage?taxon= Aedes %20canadensis) that corresponds to a specimen collected by NEON (specimen number MOSN4427-20), which allowed us to visualize, using tools available in BOLD, its genetic distance from more than 300 other nominal canadensis in the database. This specimen clustered very closely with all others at a distance of COI barcode criteria for an individual species. This is a single data point, from a single gene, from unpublished data, but it suggests that further focused collecting and sequencing will probably reveal that canadensis and mathesoni are conspecific, at least using the COI gene. We do not think there is any indication of genetic divergence, only easily explicable clinal variation. Plus, the two nominal taxa are apparently sympatric and, although weak, the single molecular indicator shows they are genetically very similar. We therefore think that subspecies mathesoni is a synonym of Aedes (Ochlerotatus) canadensis, which we formally assert here: mathesoni Middlekauff, 1944, junior subjective synonym of Aedes (Ochlerotatus) canadensis (Theobald, 1901c) . Future work may prove we have not interpreted the literature correctly, but until then mathesoni belongs as a synonym. An alternate hypothesis, that there is a northern species and a southern species with a zone of hybridization, is valid. We do not see signs of that, but it should not be totally discounted. The nominal subspecies mathesoni, which is listed as a species in the Encyclopedia of Life, must be removed from the list of valid species of Aedes. There is one other synonym of Ae. canadensis, Culex nivitarsis Coquillett, 1904, synonymized by Howard et al. (1917), who wrote: “Coquillett described Culex nivitarsis from two specimens and none have been since collected. Dr. C. S. Ludlow suggested to us that these specimens were only aberrations of A. canadensis, and we have adopted this view.”, 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 28-30, DOI: 10.11646/zootaxa.5303.1.1, http://zenodo.org/record/8043342, {"references":["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","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","Middlekauff, W. W. (1944) A new species of Aedes from Florida (Diptera: Culicidae). Proceedings of the Entomological Society of Washington, 46 (2), 42 - 44.","Rings, R. W. & Hill, S. O. (1948) The taxonomic status of Aedes mathesoni. Proceedings of the Entomological Society of Washington, 50 (2), 41 - 49.","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.","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.","Ross, H. H. (1947) The mosquitoes of Illinois (Diptera, Culicidae). Bulletin of the Illinois Natural History Survey, 24 (1), 1 - 96. https: // doi. org / 10.21900 / j. inhs. v 24.195","Darsie, R. F., Jr. (1951) Pupae of the culicine mosquitoes of the northeastern United States (Diptera, Culicidae, Culicini). Memoir 304. Cornell University Agricultural Experiment Station, Ithaca, New York, 67 pp.","Craig, G. B. Jr. (1956) Classification of eggs of nearctic aedine mosquitoes (Diptera: Culicidae). PhD Thesis, University of Illinois, Urbana-Champaign, Illionis, vii + 135 pp.","Craig, G. B., Jr. & Horsfall, W. R. (1958) Taxonomic and ecological significance of eggs of aedine mosquitoes. Proceedings of the Tenth International Congress of Entomology, 3, 853 - 857.","Ross, H. H. & Horsfall, W. R. (1965) A synopsis of the mosquitoes of Illinois (Diptera, Culicidae). Biological Notes No. 52. Illinois Natural History Survey, Urbana, Illinois, 50 pp. https: // doi. org / 10.5962 / bhl. title. 15153","Kalpage, K. S. & Brust, R. A. (1968) Mosquitoes of Manitoba. I. Descriptions and a key to Aedes eggs (Diptera: Culicidae). Canadian Journal of Zoology, 46 (4), 699 - 718. https: // doi. org / 10.1139 / z 68 - 098","Horsfall, W. R., Voorhees, F. R. & Cupp, E. W. (1970) Eggs of floodwater mosquitoes. XIII. Chorionic sculpturing. Annals of the Entomological Society of America, 63 (6), 1709 - 1716. https: // doi. org / 10.1093 / aesa / 63.6.1709","Horsfall, W. R. (1972) Mosquitoes, their bionomics and relation to disease. Hafner Publishing Company, New York, x + 723 pp.","Harrison, B. A., Byrd, B. D., Sither, C. B. & Whitt, P. B. (2016) The Mosquitoes of the Mid-Atlantic Region: An Identification Guide. Mosquito and Vector-borne Infectious Diseases Laboratory Publication 2016 - 1. Western Carolina University, Cullowhee, North Carolina, v + (1) + 201 pp.","Rings, R. W. & Hill, S. O. (1946) The larva of Aedes (Ochlerotatus) mathesoni Middlekauff (Diptera, Culicidae). Proceedings of the Entomological Society of Washington, 48 (9), 237 - 240.","Bickley, W. E. (1981) Notes on the geographical distribution of Aedes canadensis mathesoni. Mosquito Systematics, 13 (2), 150 - 152.","Colless, D. H. (1956) Environmental factors affecting hairiness in mosquito larvae. Nature, 177, 229 - 230. https: // doi. org / 10.1038 / 177229 a 0","Mattingly, P. F. (1975) Mosquito larvae. III. The hairiness factor again? Mosquito Systematics, 7 (3), 179 - 184.","Coquillett, D. W. (1901) Three new species of Culicidae. Canadian Entomologist, 33 (9), 258 - 260. https: // doi. org / 10.4039 / Ent 33258 - 9","Bradshaw, W. E. & Lounibos, L. P. (1977) Evolution of dormancy and its photoperiodic control in pitcher-plant mosquitoes. Evolution, 31 (3), 546 - 567. https: // doi. org / 10.2307 / 2407521","Harrison, B. A. (1980) Medical entomology studies - XIII. The Myzomyia Series of Anopheles (Cellia) in Thailand, with emphasis on intra-interspecific variations (Diptera: Culicidae). Contributions of the American Entomological Institute, 17 (4), iv + 1 - 195.","Le Sueur, D. & Sharp, B. L. (1991) Temperature-dependent variation in Anopheles merus larval head capsule width and adult wing length: implications for anopheline taxonomy. Medical and Veterinary Entomology, 5 (1), 55 - 62. https: // doi. org / 10.1111 / j. 1365 - 2915.1991. tb 00521. x","Coquillett, D. W. (1904) New North American Diptera. Proceedings of the Entomological Society of Washington, 6 (3), 166 - 192.","Howard, L. O., Dyar, H. G. & Knab, F. (1917) The mosquitoes of North and Central America and the West Indies. Vol. 4. Systematic description (in two parts). Part II. Publication No. 159. Carnegie Institution of Washington, Washington, D. C., pp. 525 - 1064."]}

Aedes (Aedimorphus) tricholabis Edwards subspecies bwamba van Someren, 1950 —original combination: Aedes (Aedimorphus) tricholabis ssp. bwamba. Distribution: Benin, Kenya, Nigeria, Uganda (Wilkerson et al. 2021). subspecies tricholabis Edwards, 1941 —original combination: Aedes (Aedimorphus) tricholabis. Distribution: Benin, Burkina Faso, Central African Republic, Côte d’Ivoire, Kenya, Mali, Nigeria (Wilkerson et al. 2021). This taxon is an example of the scant taxonomic work carried out on mosquitoes in Africa on all but the most important vectors of human and animal pathogens. Aedes tricholabis was described from the adult female, male and male genitalia, but not the larval or pupal stages. Subspecies bwamba was described, in addition to the larva and pupa, from the adult male and female, which were distinguished from tricholabis based on a single, but we think significant, character. Because the larval and pupal stages of tricholabis are not known, descriptions of the immature stages of bwamba have served to represent tricholabis (see below). The dearth of specimens needed to carry out direct comparisons of all life stages is surprising given that tricholabis sensu lato has been found to harbor Dengue, Sindbis and other viruses (e.g. Ochieng et al. 2013; Munyao et al. 2020; Musa et al. 2020). In a sample of published studies, we found that most authors cited Edwards (1941) as the authority for identifications, and therefore subspecies bwamba, not yet described in 1941, was subsequently ignored in post-1950 studies. Aedes tricholabis was described from two males and five females collected in Gede (or Gedi), located north of Mombasa, Kenya. Townsend (1990) found two females and the lectotype male (designated by Mattingly 1956) in the collection of the Natural History Museum, London. Edwards distinguished tricholabis in a key using the following characters: Scutellum with narrow scales only; decumbent scales of vertex all narrow; femora and tibiae not striped; hindtarsus entirely dark; head in female with a distinct patch of dark decumbent scales on vertex (this patch absent in males); abdominal terga usually with complete basal pale bands; postspiracular scales all broad and flat; hindfemur pale beneath almost to apex; hindungues (as hindtarsal claws) simple. The description also states that all scutellar scales are narrow and yellowish, and, in addition to pale basal bands on all terga, there are also dark apical bands on the sterna. The adult male, as noted above, does not have a patch of dark decumbent scales on the vertex, which led Edwards to question if the males and females were the same species, because there were no associated larval and pupal exuviae to prove otherwise. To our knowledge, this query has not been answered. The most remarkable distinguishing feature of tricholabis (and subspecies bwamba), however, is a gonostylus, which on the ventral surface (“outer edge”) is “provided with a row of long soft hairs [setae], most of which are branched, the one nearest the base having most branches.” The egg, larva and pupa have not been described and only the male genitalia have been illustrated. Subspecies bwamba was described from the male holotype, six male paratypes, and five larval and nine pupal exuviae from Bwamba Province, Toro District, in western Uganda. Townsend (1990) found the holotype and four paratypes in the collection of the Natural History Museum, London. Van Someren (1950) provided detailed descriptions of the adult male, male genitalia, and larval and pupal stages. Only the male genitalia and pupa were illustrated. Van Someren stated: “The male terminalia of this species do not differ from the figure of A. (A.) tricholabis...”. The description of the male of bwamba does not appear to us to differ from tricholabis, including the lack of a patch of dark decumbent scales on the vertex. However, the scutellum is “clothed with broad creamy white scales on all lobes.” This is in contrast to the scutellar scales being all narrow and yellowish in tricholabis. The larval stage of subspecies bwamba is apparently quite distinctive, as relayed in a personal communication to van Someren: “Dr. Haddow (personal communication) states that the larvae are very pale and transparent in life, with the eye spots very large and black. Usually the swimming is done with the mouth brushes, but when they are disturbed, body swimming movements occur. They are markedly restless, often looping the loop under water and have a habit of browsing. …The larval skins are very fragile, with all the setae pale and transparent and easily detached.” The larval stage of tricholabis is not known, precluding a comparison with subspecies bwamba. Hopkins (1952) used van Someren’s description of the larva of subspecies bwamba to represent the type form and Pao & Knight (1970) described the larval mandible and maxilla of subspecies bwamba to represent tricholabis sensu lato. No explanation was given of how the taxonomic determination was made. Similarly, Cordellier & Geoffroy (1976) represented tricholabis sensu lato as both a species and as a species with a subspecies, without explanation. The distributions of these nominal taxa, therefore, have not been documented, other than for the type localities and for surveys that used Edwards (1941) as an identification resource. Service (1960), however, added to the description of the larva of subspecies bwamba from Lokoja, Nigeria (7° 47′ 48″ N, 6° 44′ 25″ E). This site is far from the type locality of bwamba in Uganda and gives possible insight into the broad distribution of at least subspecies bwamba. Service (1960) also “fully endorsed” Haddow’s statement that “the larvae in life are pale and transparent with large dark contrasting eyes.” We do not note consensus regarding the identity and distributions of the nominotypical form or subspecies bwamba. The only clear and significant distinction is that tricholabis adults have all scutellar scales narrow and yellowish whereas adults of bwamba have all scutellar scales broad and creamy white. We consider this a significant difference that stands out in a confused understanding of these taxa, which we think justifies elevation of bwamba to species status: Aedes (Aedimorphus) bwamba van Someren, 1950 . Aedes bwamba 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 14-15, DOI: 10.11646/zootaxa.5303.1.1, http://zenodo.org/record/8043342, {"references":["van Someren, E. C. C. (1950) Ethiopian Culicidae: A description of a new sub-species of Aedes (Aedimorphus) tricholabis Edwards. Proceedings of the Royal Entomological Society of London, Series B: Taxonomy, 19 (5 - 6), 67 - 69. https: // doi. org / 10.1111 / j. 1365 - 3113.1950. tb 00933. x","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.","Ochieng, C., Lutomiah, J., Makio, A., Koka, H., Chepkorir, E., Yalwala, S., Mutesya, J., Musila, L., Kharmadi, S., Richardson, J., Bast, J., Schnabel, D., Wurapa, E. & Sang, R. (2013) Mosquito-borne arbovirus surveillance at selected sites in diverse ecological zones of Kenya; 2007 - 2012. Virology Journal, 10, 140. https: // doi. org / 10.1186 / 1743 - 422 X- 10 - 140","Munyao, V., Karisa, J., Munyao, C. M., Ngari, M., Menza, N., Peshu, N., Rono, M., Mbogo, C. & Mwangangi, J. (2020) Surveillance of culicine mosquitoes in six villages of Taita-Taveta County, Kenya, with host determinations from blood-fed females. Journal of Medical Entomology, 57 (6), 1972 - 1982. https: // doi. org / 10.1093 / jme / tjaa 109","Musa, A. A., Muturi, M. W., Musyoki, A. M., Ouso, D. O., Oundo, J. W., Makhulu, E. E., Wambua, L, Villinger, J. & Jeneby, M. W. (2020) Arboviruses and blood meal sources in zoophilic mosquitoes at human-wildlife interfaces in Kenya. Vector-Borne and Zoonotic Diseases, 20 (6), 444 - 453. https: // doi. org / 10.1089 / vbz. 2019.2563","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.","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.","Pao, B. & Knight, K. L. (1970) The fourth instar larval mandible and maxilla of selected Aedes (Aedimorphus) species (Diptera, Culicidae). Mosquito Systematics Newsletter, 2 (3), 98 - 131.","Cordellier, R. & Geoffroy, B. (1976) Les moustiques de Republique Centrafricaine. Distribution, abundance et frequence des culicines dans l'ouest du pays. Les arbovirus isoles. Travaux et Documents de L'O. R. S. T. O. M. No. 49. O. R. S. T. O. M., Paris, 105 pp.","Service, M. W. (1960) Taxonomic notes on some larvae of the genus Aedes (Diptera: Culicidae) from Lokoja in northern Nigeria. Entomologist's Monthly Magazine, 96, 229 - 230."]}

Culiseta (Culiseta) alaskaensis (Ludlow) subspecies alaskaensis (Ludlow, 1906) —original combination: Theobaldia alaskaensis. Distribution: Armenia, Austria, Belarus, Canada, Crimean Peninsula, Czech Republic, Denmark, Estonia, Finland, France, Georgia, Germany, Hungary, Iran, Ireland, Latvia, Lithuania, Moldova, Mongolia, Netherlands, Norway, Poland, Romania, Russia, Slovakia, Slovenia, Sweden, Switzerland, Turkey, Ukraine, United Kingdom, United States (Wilkerson et al. 2021). subspecies indica (Edwards, 1920) —original combination: Theobaldia indica (subspecific status by Maslov 1964). Distribution: Armenia, Azerbaijan, Georgia, India, Kazakhstan, Kyrgyzstan, Pakistan, Russia, Tajikistan, Turkmenistan, Ukraine, Uzbekistan (Wilkerson et al. 2021). Culiseta alaskaensis was described from five females and one male collected at Fort Egbert, Alaska (Ludlow 1906). According to published works, Cs. alaskaensis sensu stricto is a Holarctic species. In the Nearctic Region, it is found in western North America from Colorado to Alaska and across Canada to Newfoundland. In the Palaearctic Region, it occurs in northern Europe southward to the northern slopes of the Alps and eastward to the far east of Siberia. Subspecies indica was originally described as a species based on specimens collected in the hilly and mountainous areas of Haryana State (Ambala) and Himachal Pradesh State (Bakloh and Dalhousie) in the far north of India (Edwards 1920) and was reclassified as a subspecies of alaskaensis by Maslov (1964). Both subspecies are recorded from localities in Armenia, Georgia, Russia and Ukraine. If the identifications are correct, it would appear that the distributions of the two forms overlap, indicating that they may occur in sympatry. However, those countries lie far north of the type locality of indica, which is located on the south side of the Himalaya Mountains, and they are thus likely to be based on misidentified specimens of the typical form. This agrees with the allopatric distributions of the two forms depicted by Maslov (1967, 1989: map, fig. 60). According to Maslov, individuals of the typical form are “Mostly forest mosquitoes found as frequently in the hills as in the plains” whereas individuals of indica “are mosquitoes of the plains in forest-free regions”. Wood et al. (1979) confirmed that the type form “is widely distributed in the boreal forest across northern Europe, the USSR, and northern Canada.” Maslov (1967, 1989) and Gutsevich et al. (1971, 1974) distinguished the two subspecies as follows. Culiseta alaskaensis sensu stricto is a dark mosquito, integument and scales dark brown or black; dark and pale scaling of tarsi and abdominal terga in strong contrast; wing entirely dark-scaled or with few pale scales on anterior veins, clusters of dark scales distinct; posterior half of abdominal terga entirely dark-scaled. In general, indica is a paler mosquito, integument light brown or ochreous brown, scutum covered with golden-yellowish scales; dark and pale scales of tarsi and abdominal terga not well contrasted; most veins of wing with scattered pale scales, clusters of dark scales indistinct; posterior half of abdominal terga with few scattered pale scales. Maslov (1967, 1989) is the only researcher to distinguish the male genitalia and larvae of the two forms. In the type form, 2 large setae on basal mesal lobe of gonocoxite bent in distal one-third; larva with seta 4-C (postclypeal) 3-branched, very rarely with 4 branches; head and siphon very dark, often almost black. In indica, 2 large setae of basal mesal lobe of gonocoxite bent just beyond mid-length; larval seta 4-C with 3–7 (usually 5 or 6) branches; head and siphon light brown, sometimes yellowish brown. Qutubuddin (1952) purportedly described and illustrated the larva of indica from Pakistan. Although he received comments on his manuscript from Peter Mattingly in London, there is no doubt the larva he described is that of an undescribed species of the subgenus Allotheobaldia Brolemann, 1919. The larva is very similar to the larva of Cs. longiareolata (Macquart, 1838), the only currently recognized species of the subgenus, but it bears some distinct differences. The following characteristics place the larva in Allotheobaldia: Antenna short, seta 1-A weakly developed; siphon short and stout, not sclerotized at base, siphon index 1.5, pecten comprised of simple spines (one bifid spine is illustrated), setae 1a-S and 2a-S absent; saddle incomplete ventrally, covering dorsal half of segment X; seta 2-X multi-branched, seta 3-X double; ventral brush (seta 4-X) extended anteriorly on ventral midline of segment, with about 9 pairs of setae. The larva was described from exuviae associated with three reared females. Unlike larvae of alaskaensis, the exuviae examined by Qutubuddin were those of larvae collected “from foul-smelling water in an unused masonry well”. “Several adults were, later on, taken from the same place.” It seems that the adults were misidentified and Peter Mattingly, if he saw the illustrations, did not notice that the larva could not be the larva of indica, which does “not differ [substantially] from those of the nominate subspecies” (Gutsevich et al. 1971, 1974). The larva illustrated by Qutubuddin (1952) differs distinctly from the larva of Cs. longiareolata (based on the description of Hopkins 1952) in having setae 5- and 6-C with multiple branches (single in longiareolata), 5-C more or less pectinate with branches arising from a short stout stem; dorsomentum shorter, less acute distally (a straight-sided triangle in longiareolata); some comb scales distinctly asymmetrical (spicules on one side) (evenly fringed in longiareolata); siphon shorter (index about 2 in longiareolata); pecten comprised of 5–7 spines born entirely on the siphon (pecten with 3 or 4 small spines proximal to the base of the siphon and about 9 on the siphon in longiareolata). In contrast, as described and illustrated by Carpenter & LaCasse (1955), Maslov (1967, 1989), Gutsevich et al. (1971, 1974), Wood et al. (1979) and Becker et al. (2020), the larva of alaskaensis bears the following comparable traits: Setae 5- and 6-C fan-like with multiple aciculate branches; dorsomentum short, edges not exactly straight; comb scales elongate and evenly fringed; siphon longer, index 2.5–3.5; pecten comprised of numerous short spines on proximal 0.2 followed by a row of 16–18 filamentous spines extending to about distal 0.25 of the siphon. Two nominal species, Theobaldia arctica Edwards, 1920 and Culiseta siberiensis Ludlow, 1920, are currently recognized as synonyms of the nominotypical subspecies, and the nominal Theobaldia wassilievi Shingarev, 1927 is a synonym of subspecies indica. Theobaldia arctica was described from a single male collected at Arkhangel (English for Arkhangelsk), located far north of Moscow on the Northern Dvina River where it empties into the White Sea, and Cs. siberiensis was described from 24 females collected at three places, including Verkhne-Udinsk (former name of present-day Ulan-Ude), located southeast of Lake Baikal in south-central Siberia. The type localities of both nominal forms reside within the distribution of alaskaensis sensu stricto (Maslov 1967, 1989), and both have been listed as synonyms of the type form since Edwards (1921d). In fact, when Edwards (1920) described arctica, he stated that “In coloration and genital structure this insect agrees almost entirely with T. alaskaensis, Ludlow, and may in fact be the same.” Theobaldia wassilievi was recognized as a distinct species until Martini (1930) treated it as a variety of alaskaensis and Stackelberg (1937) later treated it as a subspecies. Contrary to Knight & Stone (1977), Harbach (2018) and Wilkerson et al. (2021), wassilievi was placed in synonymy with indica (when it was still recognized as a species) by Edwards (1932a), not by Maslov (1967, 1989). Theobaldia wassilievi was originally described as a species from Turkestan, a city in the Kazakh Desert in the southernmost region of Kazakhstan. Turkestan lies at the northern limit of the distribution of indica mapped by Maslov (1967, 1989). Theobaldia wassilievi should therefore remain a synonym of indica. Based on the available morphological, ecological and distributional information, it seems prudent to recognize indica as a separate species: Culiseta (Culiseta) indica (Edwards, 1920) . Culiseta indica is currently listed as a species in the Encyclopedia of Life. Based on the wide distribution of Cx. alaskaensis, we think it is likely that molecular data will show it is a complex of 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 108-110, DOI: 10.11646/zootaxa.5303.1.1, http://zenodo.org/record/8043342, {"references":["Ludlow, C. S. (1906) An Alaskan mosquito. Canadian Entomologist, 38 (10), 326 - 328. https: // doi. org / 10.4039 / ent 38326 - 10","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. (1920) Mosquito notes. Bulletin of Entomological Research, 10 (2), 129 - 137. https: // doi. org / 10.1017 / S 0007485300043923","Maslov, A. V. (1964) On the systematics of bloodsucking mosquitoes of the group Culiseta (Diptera, Culicidae). Entomologicheskoe Obozrenie, 43 (1), 193 - 217. [in Russian]","Maslov, A. V. (1967) Krovososushchie komarȳ podtribȳ Culisetina (Diptera, Culicidae) mirovoi faunȳ [Bloodsucking mosquitoes of the subtribe Culisetina (Diptera, Culicidae) of the world]. Academie Imperiale des Sciences, Musee Zoologique, St Petersburg. Opredeliteli po faune SSSR, Vȳp. 93, 1 - 182, 1 pl. https: // doi. org / 10.5962 / bhl. title. 46351","Maslov, A. V. (1989) Blood-sucking mosquitoes of the subtribe Culisetina (Diptera, Culicidae) in world fauna. Smithsonian Institution Libraries and the National Science Foundation, Washington, D. C., xv + 248 pp., 1 pl.","Wood, D. M., Dang, P. T. & Ellis, R. A. (1979) The insects and arachnids of Canada. Part 6. The mosquitoes of Canada. (Diptera: Culicidae). Canadian Government Publishing Centre, Supply and Services Canada, Hull, Quebec, 390 pp.","Gutsevich, A. V., Monchadsky, A. S. & Stackelberg, A. A. (1971) Fauna Diptera. Vol. III. No. 4. Mosquitoes, Family Culicidae. New Series 100. Academy of Sciences of the USSR Zoological Institute, Leningrad, 384 pp. [for 1970, in Russian]","Qutubuddin, M. (1952) A description of the thitherto unknown larva of Theobaldia indica Edwards (Diptera, Culicidae), with a key to larvae of the Pakistan species of Theobaldia Neveu-Lemaire, 1902. Proceedings of the Royal Entomological Society of London, Series B: Taxonomy, 21 (3 - 4), 37 - 40. https: // doi. org / 10.1111 / j. 1365 - 3113.1952. tb 01045. x","Brolemann, H. - W. (1919) Sur quelques Culex des Pyrenees II. Campagne 1918. Annales de la Societe Entomologique de France, 88 (1 - 2), 65 - 103.","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","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.","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.","Becker, N., Petric, D., Zgomba, M., Boase, C., Madon, M., Dahl, C. & Kaiser, A. (2020) Mosquitoes: identification, ecology and control. 3 rd Edition. Springer Nature Switzerland AG, Cham, xxxi + 570 pp. https: // doi. org / 10.1007 / 978 - 3 - 030 - 11623 - 1","Ludlow, C. S. (1920) New Siberian Culicidae (Diptera). Insecutor Inscitiae Menstruus, 7 (10 - 12), 151 - 161. [for 1919]","Shingarev, N. L. (1927) Notes on Culicidae. II. Russkii Zhurnal Tropicheskoi Meditsiny [Russian Journal of Tropical Medicine], 5, 545 - 550. [in Russian]","Edwards, F. W. (1921 d) A revision of the mosquitos of the Palaearctic Region. Bulletin of Entomological Research, 12 (3), 263 - 351. https: // doi. org / 10.1017 / S 0007485300040207","Martini, E. (1930) 11. u. 12. Culicidae. In: Lindner, E. (Ed.), Die Fliegen der palaerktischen Region. Band III. E. Schweizerbart'sche Verlagsbuchhandlung (Erwin Nagele), Stuttgart, pp. 145 - 320.","Stackelberg, A. A. (1937) Fauna de l'URSS. Insectes, Dipteres. Vol. III. n 0 4. Fam. Culicidae (Subfam. Culicinae). Nouvelle Serie no 11. Institut Zoologique de l'Academie des Sciences de l'URSS, Moscow, x + 258 pp. [in Russian]","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.","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","Edwards, F. W. (1932 a) Genera Insectorum. Diptera, Fam. Culicidae. Fascicle 194. V. Verteneuil & L. Desmet, Bruxelles [Brussels], 258 pp., 5 pls."]}

Uranotaenia (Pseudoficalbia) novobscura Barraud subspecies novobscura Barraud, 1934 —original combination: Uranotaenia novobscura. Distribution: Bangladesh, Cambodia, India, [Palaearctic] Japan, Laos, Malaysia, People’s Republic of China, Taiwan, Thailand (Wilkerson et al. 2021). subspecies ryukyuana Tanaka, Mizusawa & Saugstad, 1979 —original combination: Uranotaenia (Pseudoficalbia) novobscura ryukyuana. Distribution: Japan, Ryukyu Archipelago (Amani, Okinawa, Yaeyama) (Tanaka et al. 1979). Uranotaenia novobscura ryukyuana is endemic to the Ryukyu Archipelago of Japan. It has an interesting taxonomic history. Roth (1946) recognized two larval forms of this taxon on Okinawa Island, which he identified as Forms A and B of Ur. bimaculata Leicester, 1908. Since its original description, Ur. novobscura was mistakenly identified as Ur. bimaculata until its identity was resolved by Peyton (1977). Although not explicitly stated, it is obvious that Roth considered the two larval forms to belong to a single variable species. Peyton, in his revision of the subgenus Pseudoficalbia in Southeast Asia, likewise concluded that the various populations of Ur. novobscura that he studied “represent a single plastic species and that the adult population from Okinawa is a recognizable variant.” Tanaka et al. (1979) disagreed that specimens from Okinawa were merely a variant of novobscura sensu stricto and distinguished and described the form as a distinct subspecies, ryukyuana, as follows: “The population of the Ryukyu Archipelago is characterized by the entirely pale pleura; the scutum is also rather paler, and the supraalar dark patches are smaller than in the populations of Palaearctic Japan. The latter is identical with specimens from Taiwan and Malaya *. [The asterisk refers to specimens from the two countries that were examined.] The female antenna appears slightly shorter, 1.30–1.40 (8) length of proboscis and male antennal flagellomere 12 is shorter relative to Flm 13, viz., Flm 12 1.11–1.25 (7) length of Flm 13. Remigium scales are all dark in the Amami and Okinawa populations, but pale ochreous in the basal half in the Yaeyama population. Bristles [setae] on each side of tergum IX of the male genitalia are fewer, 1–4 (x = 2.7, mode = 2) in Amami and Okinawa, 1–3 (x = 1.6, mode = 1) in Yaeyama, while they are 2–6 (x = 3.4, mode = 3) in Palaearctic Japan. Differences in the larvae are rather obscure…. Six larvae from Yaeyama do not appear distinctly different from specimens from Okinawa Is.” Despite the morphological similarity of other subgeneric forms described from the Ryukyu Archipelago, molecular studies have revealed that these forms are genetically distinct, reproductively isolated allopatric populations which required recognition as independent species (Toma et al. 2019; Somboon et al. 2020a; Wilkerson et al. 2022). In view of those studies, and in accordance with the above treatments of the former subspecies of Culex hayashii Yamada, 1917 and Toxorhynchites manicatus (Edwards, 1921a), we believe it is likely that molecular and genetic studies will show that ryukyuana is a separate species. Accordingly, ryukyuana is hereby formally accorded species status: Uranotaenia (Pseudoficalbia) ryukyuana Tanaka, Mizusawa & Saugstad, 1979 . Uranotaenia ryukyuana is currently listed as a species in the Encyclopedia of Life; however, the date of authorship of the species is incorrectly listed as 1975, and should be corrected to 1979., 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 132-133, DOI: 10.11646/zootaxa.5303.1.1, http://zenodo.org/record/8043342, {"references":["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.","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.","Roth, L. (1946) The larva and pupa of Uranotaenia bimaculata Leicester on Okinawa Shima, Ryukyu Retto (Diptera: Culicidae). Proceedings of the Entomological Society of Washington, 48 (4), 67 - 75.","Leicester, G. F. (1908) The Culicidae of Malaya. Studies from the Institute for Medical Research, Federated Malay States, 3 (3), 18 - 261.","Peyton, E. L. (1977) Medical entomology studies - X. A revision of the subgenus Pseudoficalbia of the genus Uranotaenia in Southeast Asia (Diptera: Culicidae). Contributions of the American Entomological Institute, 14 (3), iv + 1 - 273.","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","Yamada, S. (1917) On two new species of mosquitoes from Japan. Dobutsugaku zasshi [Zoological Magazine], 29 (341), 61 - 72. [in Japanese]","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"]}

Subspecies arabiensis Patton (1905) described subspecies arabiensis from Ulub Camp and Crater, West Aden Protectorate [Yemen] from an unspecified number of adult males, adult females and larvae. No type specimen was designated and type material is presumed to be lost or never existed. White (1975), when he removed arabiensis from synonymy (synonymy by Hamon et al. 1966) with vexans and elevated it to a subspecies of vexans, reported finding a male and two females of arabiensis from Aden, “topotypic arabiensis material”, in the collection of the Natural History Museum, London, but did not designate a neotype. Patton’s (1905) original description of arabiensis is vague and roughly follows the description of vexans by Becker et al. (2020) paraphrased above. Confusingly, Patton stated: Thorax “with a dark line down the center and two at the sides. Abdomen brown with apical black bands. … Male genitalia, basal lobes narrow... apical segment thin and somewhat club shaped.” The apical black abdominal bands probably refer to contrast with the basal pale bands in all other taxa related to vexans. Edwards (1921d) apparently listed it as a synonym, perhaps “in part”, in a treatment of vexans: “(?) Culex arabiensis, Patton (adult, not larva) [alternatively, this could mean he thought the larva was a different species].” Edwards (1925) listed it with the “ vexans series” as “ A. vexans, Mg. (=? arabiensis, Patton.).” A year later, Edwards (1926b) explicitly listed arabiensis as a synonym of vexans, but then in 1941 he placed it as a species of the vexans group. Edwards (1941) stated that arabiensis was “Distinguished from other Ethiopian species of the subgenus [Aedimorphus] by the combination of the following features: basally-ringed tarsal segments, dark-scaled wings, broadly banded abdomen, and pale posterior surface of middle tibiae. A. arabiensis is very closely related to the Palearctic A. vexans Mg., from which it differs in having the male palpi rather shorter; abdominal bands in both sexes broader and not emarginate in the middle; middle tibia dark above (in all European and Central Asian females of A. vexans examined the middle tibia is conspicuously pale above as well as posteriorly).” Further, abdominal “tergites [terga] 2–6 with broad creamy-white basal bands, 2–7 with lateral whitish patches extending most of their length, 6 and 7 with narrow pale apical bands; sternites almost entirely pale scaled.” Lewis (1945) added that the larva has seta A [7-C] with 7 or 8 branches, seta B [6-C] single and seta C [5-C] single or double. “Comb a patch or irregular row of 8–12 sharp-pointed spines with small basal denticles. Siphon with index about 2.5... pecten reaching slightly beyond middle… of the last 1–3 teeth [spines] usually 2 larger and widely spaced.... Gills [anal papillae] subequal, lanceolate, much longer than saddle.” The relatively long pointed anal papillae are a particularly obvious characteristic. Hopkins (1952) expanded the description of Lewis (1945) and provided an illustration. “The only other Aëdes larvae with the comb composed of a small number of spines... head-setae B and C [6- and 5-C] single and the last 2–3 pecten spines wider-spaced are cumminsi, fowleri and durbanensis. From the first two of these the larva of arabiensis is easily separated... by the shape of the ‘gills’ [anal papillae].” Carpenter & LaCasse (1955), without comment, listed nocturnus, arabiensis and nipponii as synonyms of vexans. Perhaps those authors were unaware of or disagreed with the recognition of nocturnus and nipponii as subspecies of vexans by Bohart & Ingram (1946b). The story became less clear to us with Muspratt’s (1955) description of vexans from Transvaal, South Africa, and its comparison with vexans from England and the USA (Washington State). Muspratt pointed out some differences, such as “the tibiae of the former [South Africa] are practically all dark except for the narrow basal pale bands and apical spots, whereas the tibiae of the latter [England; USA] are extensively pale posteriorly...” [as described for arabiensis from Yemen]. Also, “on the South African form the basal pale bands of the abdominal tergites [terga] are not narrowed in the middle, thus being as described for arabiensis...”. Jupp (1996) illustrated and keyed, from South Africa, as vexans, a taxon with narrow basal emarginate abdominal tergal bands and with the sterna not all pale but with dark markings. This suggests two species in South Africa, and we think neither is arabiensis, which is distributed in northeastern Africa and Arabia. Mattingly & Knight (1956) treated arabiensis from Arabia as a species but wrote: “This is very closely related to the holarctic [sic], Oriental and Australasia Aë. vexans Meigen, of which it is possibly no more than a subspecies…”. Hamon et al. (1966) considered all observed differences as only variation. Regarding this, White (1975) wrote: “Having not seen the types of either arabiensis or sudanensis [a synonym of arabiensis], the former having been lost, Hamon et al [sic] (1966:373) formally synonymised both with vexans s. str. ” As noted above, White (1975) removed arabiensis from synonymy with vexans and elevated it to subspecific rank, and affirmed that sudanensis Theobald, 1911b was a junior synonym of arabiensis. He also listed the known distribution of vexans arabiensis as Aden, Gambia, Mauritania, Nigeria, Saudi Arabia, Somalia, South Africa (Transvaal) [we think doubtful] and Sudan. Since that time, vexans arabiensis has been identified in a number of studies, and has been implicated as a possible vector of the Rift Valley Fever virus in northeastern Africa and Arabia (Miller et al. 2002; Mondet et al. 2005; Fall et al. 2011; Clements 2012; Krtinić et al. 2013; Francuski et al. 2016; Mohamed et al. 2017; Azari-Hamidian et al. 2019). Mohamed et al. (2017), using the keys in Edwards (1941) and Hopkins (1952), found vexans arabiensis to be very common in collections made in the Republic of Sudan. Of special note was the presence of two larval types (“X” and “Z”), both of which keyed to vexans arabiensis in Hopkins (1952). Since no larvae were reared to the adult stage, it is not known which might be true arabiensis. Nor is it known if the adult females collected during the study belonged to more than one species. Given the distinctive morphological characters of the adult female and larval stage of subspecies arabiensis, we think it should be formally afforded species status: Aedes (Aedimorphus) arabiensis (Patton, 1905) . Synonym: Culex sudanensis Theobald, 1911b. Aedes arabiensis 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 18-19, DOI: 10.11646/zootaxa.5303.1.1, http://zenodo.org/record/8043342, {"references":["Patton, W. S. (1905) The culicid fauna of the Aden Hinterland, their haunts and habits. Journal of the Bombay Natural History Society, 16 (4), 623 - 637, 5 pls.","White, G. B. (1975) Notes on a catalogue of Culicidae of the Ethiopian Region. Mosquito Systematics, 7 (4), 303 - 344.","Hamon, J., Maffi, M., Grenier, P., Ouedraogo, C. S. & Djime, D. (1966) Notes sur les moustiques de la Republique islamique de Mauritanie [Dipt. Culicidae] (IIe partie). Annales de la Societe Entomologique de France, 2 (2), 371 - 383.","Becker, N., Petric, D., Zgomba, M., Boase, C., Madon, M., Dahl, C. & Kaiser, A. (2020) Mosquitoes: identification, ecology and control. 3 rd Edition. Springer Nature Switzerland AG, Cham, xxxi + 570 pp. https: // doi. org / 10.1007 / 978 - 3 - 030 - 11623 - 1","Edwards, F. W. (1921 d) A revision of the mosquitos of the Palaearctic Region. Bulletin of Entomological Research, 12 (3), 263 - 351. https: // doi. org / 10.1017 / S 0007485300040207","Edwards, F. W. (1925) Mosquito notes. - V. Bulletin of Entomological Research, 15 (3), 257 - 270. https: // doi. org / 10.1017 / S 0007485300046204","Edwards, F. W. (1926 b) Una revisione delle zanzare delle regioni paleartiche. Rivista di Malariologia, 5 (3 - 6), 1 - 152.","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. (1945) Observations on the distribution and taxonomy of Culicidae (Diptera) in the Sudan. Transactions of the Royal Entomological Society of London, 95 (1), 1 - 24, 2 maps. https: // doi. org / 10.1111 / j. 1365 - 2311.1945. tb 00259. 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.","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.","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.","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.","Jupp, P. G. (1996) Mosquitoes of southern Africa: Culicinae and Toxorhynchitinae. Ekogilde Publishers, Hartebeespoort, 156 pp.","Mattingly, P. F. & Knight, K. L. (1956) The mosquitoes of Arabia I. Bulletin of the British Museum (Natural History) Entomology, 4 (3), 91 - 141.","Theobald, F. V. (1911 b) A new genus and two new species of Culicidae from the Sudan. In: Balfour, A. (Dir.), Fourth Report of the Wellcome Tropical Research Laboratories at the Gordon Memorial College, Khartoum. Vol. B. - General Science. Bailliere, Tindall & Cox, London, pp. 151 - 156.","Miller, B. R., Godsey, M. S., Crabtree, M. B., Savage, H. M., Al-Mazrao, Y., Al-Jeffri, M. H., Abdoon, A. - M. M., Al-Seghayer, S., M., Al-Shahrani, A. M. & Ksiazek, T. G. (2002) Isolation and genetic characterization of Rift Valley fever virus from Aedes vexans arabiensis, Kingdom of Saudi Arabia. Emerging Infectious Diseases, 8 (12), 1492 - 1494. https: // doi. org / 10.3201 / eid 0812.020194","Mondet, B., Diaite, A., Fall, A. G. & Chevalier, V. (2005) Relations entre la pluviometrie et le risque de transmission virale par les moustiques: cas du virus de la Rift Valley Fever (RVF) dans le Ferlo (Senegal). Environnement, Risques & Sante, 4 (2), 125 - 129.","Fall, A. G., Diaite, A., Etter, E., Bouyer, J., Ndiaye, T. D. & Konate, T. D. (2011) The mosquito Aedes (Aedimorphus) vexans arabiensis as a probable vector briging the West Nile virus between birds and horses in Barkedji (Ferlo, Senegal). Medical and Veterinary Entomology, 26 (1), 106 - 111. https: // doi. org / 10.1111 / j. 1365 - 2915.2011.00974. x","Clements, A. N. (2012) The biology of mosquitoes. Vol. 3. Transmission of viruses and interactions with bacteria. CABI, Wallingford, xvii + 571 pp. https: // doi. org / 10.1079 / 9781845932428.0000","Krtinic, B., Francuski, L., Petric, E. & Milankov, V. (2013) Genetic diversity and differentiation between Palearctic and Nearctic populations of Aedimorphus (= Aedes) vexans (Meigen, 1830) (Diptera, Culicidae). Journal of Vector Ecology, 38 (1), 154 - 162. https: // doi. org / 10.1111 / j. 1948 - 7134.2013.12021. x","Francuski, L., Milankov, V., Ludoski, J., Krtinic, B., Lundstrom, J. O., Kemenesi, G. & Ferenc, J. (2016) Genetic and phenotypic variation in central and northern European populations of Aedes (Aedimorphus) vexans (Meigen, 1830) (Diptera, Culicidae). Journal of Vector Ecology, 41 (1), 160 - 171. https: // doi. org / 10.1111 / jvec. 12208","Mohamed, A. H., Ali, A. M., Harbach, R. E., Reeves, R. G., Ibrahim, K. M., AhmedMohmed, M. A. & Azrag, R. S. (2017) Aedes mosquitoes in the Republic of the Sudan, with dichotomous keys for the adult and larval stages. Journal of Natural History, 51 (9 - 10), 513 - 529. https: // doi. org / 10.1080 / 00222933.2017.1285069","Azari-Hamidian, S., Norouzi, B. & Harbach, R. E. (2019) A detailed review of the mosquitoes (Diptera: Culicidae) of Iran and their medical and veterinary importance. Acta Tropica, 194, 106 - 122. https: // doi. org / 10.1016 / j. actatropica. 2019.03.019"]}

Toxorhynchites (Lynchiella) haemorrhoidalis (Fabricius) subspecies haemorrhoidalis (Fabricius, 1787) —original combination: Culex haemorrhoidalis. Distribution: Argentina, Bolivia, Brazil, Colombia, Ecuador, French Guiana, Guyana, Paraguay, Peru, Suriname, Trinidad and Tobago, Venezuela (updated from Knight & Stone 1977). subspecies separatus (Lynch Arribálzaga, 1891b) —original combination: Megarhina [sic] separata (subspecific status by Lane 1951). Distribution: Argentina, Bolivia, Brazil, Paraguay (Wilkerson et al. 2021, Nicaragua deleted). subspecies superbus (Dyar & Knab, 1906a)—original combination: Megarhinus superbus (subspecific status by Lane 1951). Distribution: Belize, Colombia, Costa Rica, Cuba, Ecuador, French Guiana, Guatemala, Honduras, Mexico, Nicaragua, Panama, Suriname, Trinidad and Tobago, Venezuela (Wilkerson et al. 2021). The taxonomic history of Tx. haemorrhoidalis involves four nominal species: Culex haemorrhoidalis Fabricius, 1787, Megarhina separata Lynch Arribálzaga, 1891b, Megarhinus lynchi Dyar & Knab, 1906a and Megarhinus superbus Dyar & Knab, 1906a. Megarhina separata was considered a synonym of haemorrhoidalis as early as Lutz & Neiva (1913). Dyar (1928) followed earlier workers in recognizing Megarhinus lynchi as a valid species; it was regarded as a questionable synonym of haemorrhoidalis by Edwards (1932a) and formally synonymized with separatus by Lane (1939). Five years later, Lane (1944) treated haemorrhoidalis, separatus and superbus as separate species; however, he distinguished the last two based only on distribution: superbus in Central America and separatus in Argentina. It seems Lane (1951), without explanation, was unable to further support the specific rank of separatus and superbus and reduced them to subspecies of haemorrhoidalis. Lane (1953), in accordance with his earlier interpretation, distinguished the two subspecies based only on distribution, and stated in a note that “We have placed T. superbus and T. separatus as subspecies of T. haemorrhoidalis. Such a course taken by us is strengthened by the fact that the zoogeographical distribution of the three forms is quite distinct.” This is not clear, but we believe Lane intended to say that the distributions of each of the three forms are distinct from one another. Except for Vargas (1953), who either disagreed with or was unaware of Lane (1951, 1953) and recognized superbus as a species in Central America, separatus and superbus have continued to be recognized as subspecies to this day. Dyar & Knab (1906a), in a discussion explaining why they gave the new name lynchi to the species in Argentina previously identified as haemorrhoidalis by Lynch Arribálzaga (1891b), Theobald (1901a), Giles (1902) and Blanchard (1905), stated that “Great confusion has been caused by basing the diagnosis on the tarsal markings without reference to sex. We find that when the sexual differences are considered the tarsal markings are a useful guide in the diagnosis of the species and are a much more constant character than has been supposed.” It is noteworthy that Dyar & Knab treated separatus, also originally described from Argentina, as a synonym of haemorrhoidalis, and described superbus as a new species based on specimens from Trinidad and Mexico. Their concept of superbus also included the identification of haemorrhoidalis by Williston (1900) based on specimens from Cuba, French Guiana and Mexico, as well as the identification of Megarhinus violaceus by Dyar & Knab (1906a) and Coquillett (1906) based on specimens collected in Central America. Dyar (1928) subsequently separated lynchi and superbus from haemorrhoidalis based on the presence or absence of a basal pale band on hindtarsomere 2, present in haemorrhoidalis and absent in the other two. He distinguished lynchi and superbus based on environmental location: superbus “From the northern edge of the tropics” and lynchi “From the southern edge of the tropics”. These distinctions were restated, using slightly different terminology, by Lane (1944, 1953), an indication that he either accepted the observations of Dyar without further study or he was unable to find additional characters to distinguish the three nominal forms. In his treatment of the Toxorhynchites (as Megarhinus) of “ Brasil Meridional” (southern Brazil), Lane (1944) stated the following (translated from the Portuguese). It is very interesting to note that, while this species [haemorrhoidalis] occurs in the Guianas and the Amazon Valley, the two related species are found, one in the North (superbus) and the other in the South (separatus). The distinguishing characteristics of these three species reside in the development of the abdominal tufts of the males and in the marking of the tarsi of the females. We think it very likely that they represent a single species and that both superbus and separatus are just geographic forms. A definitive solution of this case is impossible for us due to lack of material. The available descriptions of the leg markings of both sexes are confusing and lack explicit detail, but it appears that they are the same in all four nominal forms except for hindtarsomere 2 of females, which is pale basally in haemorrhoidalis and completely dark-scaled in lynchi, separatus and superbus. The lateral tufts on the posterior abdominal segments of males are said to be more strongly developed in haemorrhoidalis than they are in the other three nominal forms, but the degree of development has not been made explicit. Dyar & Knab (1906a) indicated that strictly red tufts are only present on abdominal segment VII in superbus but are present on segments VI and VII in haemorrhoidalis and lynchi. In contrast, Dyar (1928) stated that the male of haemorrhoidalis has “Abdominal red tufts on the last four segments”, and this is quoted verbatim by Lane (1953). In their identification keys, Dyar (1928) and Lane (1953) merely indicated that the “abdominal red tufts [are] well developed” in haemorrhoidalis and are “less developed” in lynchi (separatus of Lane) and superbus. It is interesting to note that Dyar & Knab (1906a) distinguished the males of haemorrhoidalis and lynchi based on the length of “segments 3 and 4” [palpomeres 3 and 4] of the maxillary palpus—equal in length in haemorrhoidalis and 3 longer than 4 in lynchi. Considering what is now known about the development of the maxillary palpus of mosquitoes (Harbach & Kitching 1998), “segments 3 and 4” are actually palpomeres 4 and 5. Oddly, this character was not mentioned in later works, and the descriptions of the maxillary palpi provided by Dyar (1928) and Lane (1953) are ambiguous. Dyar stated that haemorrhoidalis has “Palpi with the third joint long and pointed” (quoted verbatim by Lane), which surely must refer to the terminal palpomere, and Lane added that superbus has the “Last palpal segment long and acuminate” and separatus is “Similar to T. haemorrhoidalis superbus.” As revealed by da Costa Lima (1931), Dyar (1928) failed to notice that Goeldi (1905) had described the egg, larva and pupa of separatus, and provided a color illustration of the adult male. It is interesting to note that hindtarsomere 2 is pale basally in the male illustrated by Goeldi, indicating that the species he described is not separatus, nor any of the other three nominal forms under discussion here, all of which have hindtarsomere 2 completely dark-scaled. Séguy (1950) provided a similar color illustration, which seems to correctly depict the male of haemorrhoidalis. The larva and pupa of separatus were briefly described by Forattini & Lane (1952) based on a single larva captured in the Serra do Diabo region in the western area of São Paulo State in southern Brazil that was reared to an adult. The sex of the adult was not mentioned, but it is presumed to have been a female otherwise the specimen could not have been identified as separatus. The descriptions (in Portuguese) were repeated (in English) by Lane (1953). Dyar (1928) and Lane (1953) provided brief descriptions of the larva of separatus. Their descriptions lack comparable information except the former author reported that the siphon is “over four times as long as wide” whereas the latter author described the siphon as being “three and a half times basal width.” Lane (1953) also provided brief descriptions of the larva and pupa of the nominotypical form. He unintelligibly characterized the larval siphon as “slightly more than one time as broad as wide.” However, judging from his illustration of the terminal abdominal segments, he obviously meant to say the siphon is slightly longer than broad. Vargas (1953), in a key for the identification of larvae of species of Toxorhynchites (as Megarhinus) known to occur in Venezuela, characterized the larval siphon of superbus as being twice as long as the saddle of segment X (transliterated from the Spanish). In comparison, Lane (1953) described the length of the siphon of superbus as being “two and a half times greatest width.” Although these authors expressed siphon length in different ways, the degree of actual difference seems to be greater than expected for individuals of the same species. The subgenus Lynchiella Lahille, 1904, to which haemorrhoidalis and 16 other species belong, is predominantly Neotropical, with an extension into eastern areas of the United States and southeastern Canada represented by Tx. rutilus (Coquillett, 1896) (see below). For the most part, the current taxonomy of Lynchiella dates back to Lane (1953) and Vargas (1953). Their studies were based almost entirely on adult mosquitoes, and the immature stages, as noted above, were described very superficially. With the exception of Tx. gerbergi Belkin, 1977, Tx. guadeloupensis (Dyar & Knab, 1906a) and Tx. portoricensis (von R̂der, 1885), the larva and pupa of which were fully described and illustrated by Belkin (1977), Augier et al. (2003) and Belkin et al. (1970), respectively, the complete larval and pupal chaetotaxy has not been studied for any other species of the subgenus. As noted by Belkin et al., Toxorhynchites “is amazingly similar in all [life] stages throughout its nearly worldwide distribution and great difficulty is experienced in identifying species. New World species have been diagnosed largely on the basis of light markings of the tarsi, which frequently differ in the 2 sexes and are not always reliable. The metallic coloration of the thoracic scales shows considerable differences among species but is subject to some variation and is difficult to describe accurately owing to marked changes in color depending on the angle of observation. Few specific differences have been noted in the male genitalia. To date no reliable characters have been found to separate any of the species of a group in the larval and pupal stages….”, which is obviously due to the fact that they have not been studied in comparative detail. It is worth noting that Belkin et al. provisionally applied the name superbus to the species in Cuba. Likewise, Belkin and his colleagues applied the name superbus to specimens reared from collections made in Colombia (Heinemann & Belkin 1978c); Costa Rica (Heinemann & Belkin 1977a); Guatemala, Honduras and Nicaragua (Heinemann & Belkin 1977b); Mexico (Heinemann & Belkin 1977c); Panama (Heinemann & Belkin 1978a); Trinidad (the type locality of superbus) (Heinemann et al. 1980); and Venezuela (Heinemann & Belkin 1978b; Navarro 1996). Those researchers identified haemorrhoidalis sensu stricto in collections made in northern Brazil and Ecuador (Heinemann & Belkin 1979); Colombia (Heinemann & Belkin 1978c); and also French Guiana, Guyana and Surinam (Heinemann & Belkin 1978b). It is certain that haemorrhoidalis sensu stricto and superbus are sympatric in at least Venezuela (Navarro et al. 2007). It is interesting to note that the collections made in Colombia, French Guiana, Guyana and Surinam also included a species identified as sp. D, near superbus. From the foregoing, it should be evident that the three subspecific forms may prove to be morphologically distinct, particularly in the larval stage; there is evidence that the distributions of haemorrhoidalis sensu stricto and superbus overlap in northern countries of South America; and available information indicates that separatus may be geographically separated from haemorrhoidalis sensu stricto and is restricted to areas southward of approximately latitude 20° south. In view of these indicators, we believe it is likely that haemorrhoidalis, separatus and superbus are separate species; thus, we here formally return separatus and superbus to their original specific status: Toxorhynchites (Lynchiella) separatus (Lynch Arribálzaga, 1891b) and Toxorhynchites (Lynchiella) superbus (Dyar & Knab, 1906a) . We anticipate that molecular data will confirm these two forms and haemorrhoidalis are three separate species. Toxorhynchites separatus and Tx. superbus are currently listed as species in the Encyclopedia of Life. Based on available morphological data and having type localities in Argentina, Megarhinus lynchi Dyar & Knab, 1906a is retained as a synonym of Tx. separatus (Lynch Arribálzaga, 1891b)., 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 118-120, DOI: 10.11646/zootaxa.5303.1.1, http://zenodo.org/record/8043342, {"references":["Fabricius, J. C. (1787) Mantissa Insectorum, sistens eorum species nuper detectas: adjectis synonymis, observationibus, descriptionibus, emendationibus. Vol. 2. Christ. Gottl. Proft, Hafniae [Copenhagen], 382 pp. https: // doi. org / 10.5962 / bhl. title. 36471","Knight, K. L. & Stone, A. (1977) A catalog of the mosquitoes of the world (Diptera: Culicidae). 2 nd Edition. 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