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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. The Thomas Say Foundation. Vol. VI. Entomological Society of America, College Park, Maryland, xi + 611 pp.","Lynch Arribalzaga, F. (1891 b) Dipterologia argentine. Revista del Museo de La Plata, 2, 131 - 174, 5 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","Dyar, H. G. & Knab, F. (1906 a) The species of mosquitoes in the genus Megarhinus. Smithsonian Miscellaneous Collections, 48 (3), 241 - 258.","Lutz, A. & Neiva, A. (1913) Contribuicies para a biolojia das megarininas com descricies de duas especies novas. Memorias do Instituto Oswaldo Cruz, 5 (2), 129 - 141. https: // doi. org / 10.1590 / S 0074 - 02761913000200003","Dyar, H. G. (1928) The mosquitoes of the Americas. Publication No. 387. Carnegie Institution of Washington, Washington, D. C., 616 pp.","Edwards, F. W. (1932 a) Genera Insectorum. Diptera, Fam. Culicidae. Fascicle 194. V. Verteneuil & L. Desmet, Bruxelles [Brussels], 258 pp., 5 pls.","Lane, J. (1939) Catalogo dos mosquitos neotropicos. Boletim Biologico Serie Monografica, 1, i - xi + 1 - 218. https: // doi. org / 10.5962 / bhl. title. 97955","Lane, J. (1944) A tribo Megarhinini no Brasil Meridional. Revista de Entomologia, 15 (1 - 2), 172 - 190.","Lane, J. (1953) Neotropical Culicidae. Vols. I & II. University of Sao Paulo, Sao Paulo, 1112 pp.","Vargas, L. (1953) Megarhinus de Norteamerica (Diptera: Culicidae). Memorias Congresso Ciencias Mexico), Ciencias biologicas: Zoologiea, 7, 308 - 315.","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","Giles, G. M. (1902) A handbook of the gnats or mosquitoes giving the anatomy and life history of the Culicidae together with descriptions of all species noticed up to the present date. 2 nd Edition. John Bale, Sons & Danielsson, Ltd., London, xi + (1) + 530 pp., 17 pls. https: // doi. org / 10.5962 / bhl. title. 8549","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","Coquillett, D. W. (1906) A classification of the mosquitoes of North and Middle America. U. S. Department of Agriculture, Bureau of Entomology, Technical Series, 11, 1 - 31. https: // doi. org / 10.5962 / bhl. title. 87519","Harbach, R. E. & Kitching, I. J. (1998) Phylogeny and classification of the Culicidae (Diptera). Systematic Entomology, 23 (4), 327 - 370. https: // doi. org / 10.1046 / j. 1365 - 3113.1998.00072. x","da Costa Lima, A. (1931) Sobre as especies de Megarhinus do Brasil (Diptera: Culicidae). Memorias do Instituto Oswaldo Cruz, 25 (4), 307 - 315, 2 pls. https: // doi. org / 10.1590 / S 0074 - 02761931000400002","Goeldi, E. A. (1905) Os mosquitos no Para: Reuniao de quatro trabalhos sobre os mosquitos indigenas, principalmente as especies que molestam o homem. Memorias do Museu Goeldi (Museu Paraense) de Historia Natural e Ethnographia IV. C. Wiegandt, Para, 154 pp., 21 pls. https: // doi. org / 10.5962 / bhl. title. 8538","Seguy, E. (1950) La biologie des Dipteres. Encyclopedie entomologique, Serie A, 26, 1 - 609, 10 pls.","Forattini, O. P. & Lane, J. (1952) Sobre Toxorhynchites haemorrhoidalis separatus Arribalzaga, 1891. Arquivos da Faculdade de Higiene e Saude Publica, 6 (1 - 2), 57 - 59. https: // doi. org / 10.11606 / issn. 2358 - 792 X. v 6 i 1 - 2 p 57 - 58","Lahille, F. (1904) Notes sur la classification des moustiques. Actas y Trabajos del Segundo Congreso Medico Latino-Americano, Buenos Aires, 2, 71 - 96.","Belkin, J. N. (1977) Toxorhynchites (Lyn.) gerbergi, a new species from the southern Lesser Antilles. Mosquito Systematics, 9 (3), 329 - 332.","Augier, L. M., Dantur Juri, M. J. & Molina, G. A. (2003) Redescripcion de la larva y la pupa de Toxorhynchites (Lynchiella) guadeloupensis (Diptera: Culicidae). Revista de la Sociedad Entomologica Argentina, 62 (1 - 2), 99 - 106.","Belkin, J. N., Heinemann, S. J. & Page, W. A. (1970) The Culicidae of Jamaica (Mosquito studies. XXI). Contributions of the American Entomological Institute, 6 (1), 1 - 458.","Heinemann, S. J. & Belkin, J. N. (1978 c) Collection records of the project \" Mosquitoes of Middle America \" 12. Colombia (COA, COB, COL, COM). Mosquito Systematics, 10 (4), 493 - 539.","Heinemann, S. J. & Belkin, J. N. (1977 a) Collection records of the project \" Mosquitoes of Middle America \" 7. Costa Rica (CR). Mosquito Systematics, 9 (2), 237 - 287.","Heinemann, S. J. & Belkin, J. N. (1977 b) Collection records of the project \" Mosquitoes of Middle America \" 8. Central America: Belize (BH), Guatemala (GUA), El Salvador (SAL), Honduras (HON), Nicaragua (NI, NIC). Mosquito Systematics, 9 (4), 403 - 454.","Heinemann, S. J. & Belkin, J. N. (1977 c) Collection records of the project \" Mosquitoes of Middle America \" 9. Mexico (MEX, MF, MT, MX). Mosquito Systematics, 9 (4), 483 - 534.","Heinemann, S. J. & Belkin, J. N. (1978 a) Collection records of the project \" Mosquitoes of Middle America \" 10. Panama, including Canal Zone (PA, GG). Mosquito Systematics, 10 (2), 119 - 196.","Heinemann, S. J., Aitken, T. H. G. & Belkin, J. N. (1980) Collection records of the project \" Mosquitoes of Middle America \" 14. Trinidad and Tobago (TR, TRM, TOB). Mosquito Systematics, 12 (2), 179 - 284.","Heinemann, S. J. & Belkin, J. N. (1978 b) Collection records of the project \" Mosquitoes of Middle America \" 11. Venezuela (VZ); Guianas: French Guiana (FG, FGC), Guyana (GUY), Surinam (SUR). Mosquito Systematics, 10 (3), 365 - 459.","Navarro, J. C. (1996) Toxorhynchites haemorrhoidalis superbus (Dyar & Knab 1906), nuevo registro para la fauna de Culicidae de Venezuela. Acta Biologica Venezuelica, 16 (2), 65 - 67.","Navarro, J. - C., Liria, J., Pinagmp. J. & Barrera, R. (2007) Biogeographic area relationships in Venezuela: A parsimony analysis of Culicidae - phytotelmata distribution in national parks. Zootaxa, 1547 (1), 1 - 19. https: // doi. org / 10.11646 / zootaxa. 1547.1.1"]}

Aedes (Bifidistylus) boneti Gil Collado subspecies boneti Gil Collado, 1936 —original combination: Aedes (Aedimorphus) boneti. Distribution: Known only from the type locality, Isla de Bioko (Fernando Po), Equatorial Guinea (Gil Collado 1936). subspecies kumbae Chwatt, 1948 —original combination: Aedes (Aedimorphus) boneti s.sp. kumbae. Distribution: Republic of Cameroon (Chwatt 1948). The nominotypical subspecies was described from a single male from Rebola, Bioko Island, Equatorial Guinea. The island has an area of 2,017 km 2 and is 32 km from the coast of Cameroon (https://en.wikipedia.org/wiki/ Bioko; accessed 7 April 2022). Gil Collado credited F. W. Edwards for identifying the species as new, stating: “I must tender my thanks to J. W. Edwards [sic], who... examined the material... verifying my findings and alerting me to a new species of Aedimorphus, represented by a specimen it had been impossible for me to identify due to its somewhat deficient state of preservation [translated from the Spanish].” The male holotype is in the Museum, Madrid University School of Agriculture, Madrid, Spain. We used a published translation of the description from the Spanish along with the original article for this treatment. The description is detailed and accompanied by a drawing of a dorsal view of the head and appendages (antenna, maxillary palpus, proboscis) and a hindleg. The forelegs are not described, leaving the impression that they are missing. Damage to the specimen noted by the author included: “The scutellum has very wide whitish scales in its middle lobe, those from the sides [lateral lobes] having been removed... Anterior pronotum [antepronotum] has two groups of 5 to 6 bristles [setae]; the posterior [postpronotum], some 4, though in our specimen, they are somewhat destroyed…. Supra-alar scales are dark and numerous; those from the dorso-central area of our specimen have been torn off….” Our assumption is that characteristics of the cuticle (setae, scales, pollinosity, etc.) might be modified enough to render an accurate description problematical. Re-examination of the holotype is needed to be certain. Gil Collado also noted: “This species presents traits which closest approximate lamborni Edw. [the only other species now included in the subgenus Bifidistylus], in whose group they must be included because of their tarsal rings; however, in spite of the fact that the specimen’s [as in translation] hypopygium [genitalia] was destroyed in preparation, the coxa [gonocoxite] does not seem as pronounced as in that species...”. Gil Collado then compared boneti with lamborni Edwards, 1923a. To our knowledge the holotype of boneti is the only known specimen of this species. However, given the intense study of Anopheles and malaria control on Bioko (e.g. Cook et al. 2018), it is not surprising that species of other genera might have been overlooked, with the exception of the relatively recent invasive Aedes albopictus (Skuse, 1895) (Toto et al. 2003), and pest species of Culex (e.g. Fuseini et al. 2019). Gil Collado (1936) documented about 30 mosquito species on Bioko. All but Anopheles lloreti Gil Collado, 1936 are also found on mainland Africa. The type of An. lloreti is perhaps the only specimen of this species as well. An attribute that to us stands out as unique is paired spots of erect black scales on the vertex of the head. “Head with a broad vertical zone of narrow, reclining, white scales, among which numerous dark standing scales appear; on each side there is a rounded blotch of dark scales, which in turn, has a lateral region of white, narrow, reclining scales, and in the same posterior angle of the eyes there is a small dark spot.” These spots are not noted in Ae. lamborni, but they are present in subspecies kumbae. Subspecies kumbae was described from Kumba, British Cameroons [Republic of Cameroon] by Chwatt (1948). The type localities of kumbae and the type form are 117 km apart (determined by David Pecor on 04/07/2022, https:// arcg.is/CqOTz1) and, as stated above, the island and mainland are separated by 32 km of ocean. The description and illustrations of kumbae are detailed and based on a series of specimens: 10 larvae, three pupal exuviae and two adult males with dissected genitalia mounted with the associated pupal exuviae. Of these, Townsend (1990) found two males and six larvae in the Natural History Museum, London. Chwatt (1948) apparently relied on the description of the male of boneti to compare subspecies kumbae with boneti since there is no indication that he examined the holotype of boneti. Hopkins (1952) included boneti in a key but reproduced the description and larval illustration of subspecies kumbae to represent boneti. For purposes of comparison, therefore, we only have the descriptions of the adult male of each nominal form. The following are comments and characters that Chwatt (1948) used to justify giving kumbae subspecies status: “In Edwards’s (1941) key to the Ethiopian species of Aëdes the two male adults [the two specimens used to describe subspecies kumbae] would run down to A. boneti Gil Collado, described in 1936 from a single damaged male captured on the Island of Fernando Po. The similarity between the two adults described above and the original description of A. boneti is considerable. Nevertheless, there are several notable differences―mainly the more extensive, rather differently shaped, dark scaling of the head, the pale (instead of golden) colour of the investiture of the mesonotum, the presence of prescutellar rows of scales, the scaling of the abdominal sternites [sterna], the presence of small pale apical spots on the dorsal surface of the femora, the presence of basal white spots on the costa, and the markings of the last hind tarsal segments.” We itemized the above characters and extracted text from the original descriptions of kumbae and boneti, i.e. from Gil Collado (1948) and Chwatt (1936), respectively, as follow. “...the more extensive, rather differently shaped, dark scaling of the head...”. — kumbae: “Occiput with two, dark, dorsolateral, conspicuous, oval or comma-shaped spots, formed by dark-brown upright forked scales. The remainder of the upright scales pale. Prominent dark-brown bare vertical area along the median suture [coronal suture].” [large dark spots illustrated] — boneti: “Head with a broad vertica1 zone of narrow, reclining, white scales, among which numerous dark standing scales appear; on each side there is a rounded blotch of dark scales, which in turn, has a lateral region of white, narrow, reclining scales, and in the same posterior angle of the eyes there is a small dark spot.” [dorsolateral and lateral dark spots evident in the illustration] We do not interpret these two descriptions to be substantially different, in addition the illustrations of the heads are quite similar, especially the two large dark spots of erect scales. “...the pale (instead of golden) colour of the investiture of the mesonotum [scutum]...”. — kumbae: “Mesonotum covered with a mixture of narrow, curved, dark-brown and pale scales...”. — boneti: “Mesonotum, except for its anterior edge, has sparse gold scales mixed with wider black scales.” Pale scales versus sparse gold scales could be explained by differences in lighting used for the observations. “...the presence of prescutellar rows of scales... ”. — kumbae: “...dark-brown and pale scales, the latter broader on the anterior and anterolateral borders [of the scutum] and broad and flat around the prescutellar bare area.” — boneti: “In the middle region, in front of the scutellum, there are some white scales.” Neither description mentions rows of scales in front of the scutellum or provides a clear distinction between the two. “...the scaling of the abdominal sternites [sterna]...”. — kumbae: “Sternites covered almost entirely with broad flat pale scales.” — boneti: Not noted. No applicable difference. “...the presence of small pale apical spots on the dorsal surface of the femora...”. — kumbae: “Fore femora dark, with a paler basal ventral surface and a few white scales at the distal end of the dorsal surface; middle femora similar, with a rather more extensive pale ventral area; hind femora dark on the dorsal side, except for a white distal spot, mainly pale on the ventral side...”. — boneti: Forefemur not noted. “Legs II [midlegs]: femora with their anterior and dorsal surfaces dark, and the posterior surface completely pale at the base, this coloration narrowing progressively toward the apex, where there is only a single white ventral line…”. “Legs III [hindlegs] with predominantly white femora, and a narrow black dorsal strip, while the anterior face is peppered with abundant dark scales.” The apical pale femoral spot in kumbae is described only for the hindfemur. The illustration of the hindleg of boneti does not show an apical spot. We think this remains ambiguous pending comparison of the holotypes of the two nominal forms. “...the presence of basal white spots on the Costa... ”. — kumbae: “Wings with scales dark and a small basal patch of pale scales on the lower and anterior surface of the costa.” — boneti: “The wing scales are dark, and even blackish over C and R 1.” A few pale scales at the base of the costa can be variable. “...and the markings of the last hind tarsal segments [which markings not stated].” — kumbae: “hind tarsi... first segment [tarsomere 1] dark, with an apical white band, second with one basal and one apical white band, the latter about twice the length of the former, third with one apical and one basal narrow band. The fourth tarsus [tarsomere 4] shows some variation: in one specimen it has a dark median band, while in the other the dark scales form only a narrow longitudinal line on the apical half of the lower surface. In both specimens the fifth tarsus [tarsomere 5] is white but has a similar dark line on the lower surface.” — boneti: “The metatarsi [hindtarsomeres 1] are black, with bristles of the same color, and a white spot at their apices which is a little longer than their width. The 2nd article [tarsomere 2] has a narrow pale basal zone, and another at its apex which is two and a half times its thickness; the 3rd with an apical ring the same as the 2nd and one at the base which is a little greater than its diameter. The 4th is white with a middle dark zone of one fourth its total length, and the 5th, completely white.” [roughly matches the illustration of the hindleg] No defining differences are given. Chwatt (1948) wrote: “This Aedimorphus is described here under the provisional name of A. boneti s.sp. kumbae. Should the still unknown larva of A. boneti prove to be different from the one described above, A. boneti s.sp. kumbae will have to be treated as a new species, A. (Aedimorphus) kumbae.” Our interpretation is that when kumbae was described, no significant characters were given to distinguish the two nominal taxa, and we therefore treat kumbae is a synonym of boneti until proven otherwise: kumbae Chwatt, 1948, junior subjective synonym of Aedes (Bifidistylus) boneti Gil Collado, 1936 . Should the larvae be found, the larva of kumbae possesses some potentially unique characteristics for diagnosis, such as an array of variously shaped spines on the siphon and a dense patch of comb scales. The nominal subspecies kumbae, which is listed as a species in the Encyclopedia of Life, must be removed from the list of valid species of Aedes., 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 20-22, DOI: 10.11646/zootaxa.5303.1.1, http://zenodo.org/record/8043342, {"references":["Gil Collado, J. (1936) Culicidos de la Isla de Fernando Poo recogidos por la expedicion J. Gil-F. Bonet. EOS - Revista Espanola de Entomologia, 11 (4), 311 - 329.","Chwatt, L. J. (1948) A new Aedes from the Cameroons, A. (Aedimorphus) boneti s. sp. kumbae s. sp. nov. Annals of Tropical Medicine and Parasitology, 42 (2), 184 - 189. https: // doi. org / 10.1080 / 00034983.1948.11685361","Edwards, F. W. (1923 a) Four new African mosquitos. Bulletin of Entomological Research, 13 (4), 397 - 399. [for 1922] https: // doi. org / 10.1017 / S 0007485300045466","Cook, J., Hergott, D., Phiri, W., Rivas, M. R., Bradley, J., Segura, L., Garcia, G., Schwabe, C. & Kleinschmidt, I. (2018) Trends in parasite prevalence following 13 years of malaria interventions on Bioko island, Equatorial Guinea: 2004 - 2016. Malaria Journal, 17, 62. https: // doi. org / 10.1186 / s 12936 - 018 - 2213 - 9","Skuse, F. A. A. (1895) The banded mosquito of Bengal. Indian Museum Notes, 3 (5), 20. [for 1894]","Toto, J. - C., Abaga, S., Carnevale, P. & Simard, F. (2003) First report of the oriental mosquito Aedes albopictus on the West African island of Bioko, Equatorial Guinea. Medical and Veterinary Entomology, 17 (3), 343 - 346. https: // doi. org / 10.1046 / j. 1365 - 2915.2003.00447. x","Fuseini, G., Nguema, R. N., Phiri, W. P., Donfack, O. T., Cortes, C., Von Fricken, M. E., Meyers, J. I., Kleinschmidt, I., Garcia, G. A., Maas, C., Schwabel, C. & Slotman, M. A. (2019) Increased biting rate of insecticide-resistant Culex mosquitoes and community adherence to IRS for malaria control in urban Malabo, Bioko Island, Equatorial Guinea. Journal of Medical Entomology, 56 (4), 1071 - 1077. https: // doi. org / 10.1093 / jme / tjz 025","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."]}

Trichoprosopon digitatum (Rondani) subspecies digitatum (Rondani, 1848) —original combination: Culex digitatus. Distribution: Argentina, Belize, Bolivia, Brazil, Colombia, Costa Rica, Ecuador, El Salvador, French Guiana, Guatemala, Guyana, Honduras, Lesser Antilles (includes Trinidad and Tobago), Mexico, Nicaragua, Panama, Paraguay, Peru, Suriname, Venezuela (Wilkerson et al. 2021). subspecies townsendi Stone, 1944 —original combination: Trichoprosopon (Trichoprosopon) digitatus var. townsendi (subspecific status by Harbach & Howard 2007). Distribution: Brazil, Lesser Antilles (includes Trinidad and Tobago), Panama (Wilkerson et al. 2021). Rondani (1848) described and named Tr. digitatum (as Culex digitatus) based on one or more females (number not indicated) from an unspecified locality in Brazil (Belkin 1968). The location of the type(s) remains unknown, but the type locality was restricted to the vicinity of Rio de Janeiro (Guanabara) by Belkin et al. (1971). In the absence of type material, Stone (1944) aptly noted that “It cannot be determined whether the type of digitatum possessed scales on the clypeus, but lacking proof to the contrary, it is best to retain the name for the most frequently encountered, unscaled variety [i.e. species of Trichoprosopon].” Stone then confirmed the synonymy of Tr. nivipes Theobald, 1901c (synonymy originally by Howard et al. 1915), Tr. splendens Lutz, 1904 (in Bourroul, 1904) (synonymy by Lane & Cerqueira 1942) and Joblotia splendens var. subsplendens Martini, 1931b (synonymy by Lane & Cerqueira 1942) with digitatum based on the absence of scales (presence of setae only) on the clypeus of those nominal species. Stone also noted that four females of Tr. wilsoni Ludlow, 1918 in the “ U. S. N. M.” (National Museum of Natural History, Washington, D.C.) bear type labels but neither of those specimens nor others attributable to Ludlow bear the data provided in the original description of the species; thus, it was “not considered possible to select any one as a lectotype.” In the absence of a definite type specimen, Tr. wilsoni has remained a synonym of digitatum ever since the synonymy was established by Dyar (1928). Of particular note, Stone observed that “One of the four females [‘supposed cotype’] has scales intermingled with the hairs [setae] on the clypeus and therefore should be referred to the new variety [Tr. digitatus var. townsendi] described in this paper.” The presence of only setae on the clypeus of digitatum sensu stricto has been recognized by all investigators (Theobald 1901c, as Tr. nivipes; Howard et al. 2015, as Joblotia digitatus; Dyar 1928, as Joblotia digitata; Senevet & Abonnenc 1939; Stone, 1944; Lane & Cerqueira 1942; Lane 1953; Zavortink et al. 1983). The last authors (Zavortink et al.) defined digitatum as “the only species of Trichoprosopon with 3‒5 (3‒6) pairs of large preapical teeth on the aedeagus, with these teeth becoming progressively larger distad of the smallest basal tooth.... The pupa is the stage next most easily identified, by its normal countershading and the moderately strong to strong, more-orless rigid seta 6-VI and usually also 6-V…. The adult of digitatum can be identified only by the combination of the densely setose clypeus… the presence of setae on the upper calypter of the wing… the presence of a small patch of light scales at the base of the first hindtarsal segment… and the light integument of the mesepimeron and meron. The larva of digitatum is the stage most difficult to identify. It can usually be told by the combination of a strong, long, single or double seta 0-P… a single or double seta 7-P… only a single ventral abdominal seta (12-II) arising from a sclerotized tubercle… and a long seta 6-VI”. Stone (1944) described “ var. townsendi ” from a type series comprised of the holotype and 17 paratypes (14 females and 3 males): “The holotype and 11 paratypes were collected at Boa Vista, Rio Tapajós, Pará, Brazil …. Four paratypes were reared by Dyar and Shannon at Porto Bello, Panama... along with a number of typical digitatum. One paratypes [sic] is the supposed cotype of Trichoprosopon wilsoni Ludlow, from Ancon, Panama. One paratype was reared at Montserrat, Trinidad … along with many of the typical variety.” Stone did not indicate whether he examined the genitalia of the males or if the larval and pupal exuviae of the reared specimens were available or examined. He simply stated: “This [variety] agrees with typical digitatum in all diagnostic characters save that there are violaceous scales mingled with the hairs [setae] on the sides of the clypeus.” Without mention of the male genitalia and immature stages of townsendi, Lane (1953) reiterated that “The only difference between this variety and the typical form is that there are scales as well as setae on [the] clypeus.” As noted above for Tr. compressum, Zavortink (1981) recognized Tr. digitatum as a species complex, consisting “of Tr. digitatum and at least two undescribed species.” Zavortink elaborated: “By any measure, Tr. digitatum is the most successful species in the genus Trichoprosopon. It is widespread geographically, occupying the entire range of the genus from Mexico to Ecuador and Argentina. It is diverse ecologically, with the immatures being found in water in bamboo stumps and tree holes, in fallen fruits and nuts, in Heliconia flower bracts, and in fallen leaves and palm spathes. And it is abundant, with about 95% of the specimens of Trichoprosopon that I have examined being this one species. The undescribed species of the complex occur sympatrically with Tr. digitatum, one in the Pacific versant of eastern Panama and Colombia, the other farther south in the Pacific versant of Ecuador. The immatures of only one of the undescribed species are known, and they have been found most often in leaf axils, a habitat not utilized by Tr. digitatum. The differences between Tr. digitatum and the undescribed species whose immatures are known can be used to illustrate the size of the morphological gap separating related sympatric species in this genus.” Zavortink then mentioned differences between Tr. digitatum and the undescribed species—differences in the coloration of the adults and pupae, the aedeagus and cercal setae of the male genitalia, and seta 7-P of the larvae—and emphatically stated: “The nature of the differences between Tr. digitatum and the new species are important to note, because these species are sympatric and their specific distinctness cannot be doubted.” “Even after the separation of two undescribed species, Tr. digitatum remains a variable species and may indeed still be a complex.” Zavortink did not mention townsendi, but the available evidence indicates that this form is also sympatric with the nominotypical form in Panama and Brazil. In view of the unknown immature stages of townsendi and the unique presence of scales on the clypeus in the absence of morphological information for the male genitalia, we suspect that further collections and integrated study will show that townsendi is a separate sympatric species of the Digitatum Complex; henceforth, it is afforded species status: Trichoprosopon townsendi Stone, 1944 . Trichoprosopon townsendi is currently listed as a species in the Encyclopedia of Life., Published as part of Harbach, Ralph E. & Wilkerson, Richard C., 2023, The insupportable validity of mosquito subspecies (Diptera: Culicidae) and their exclusion from culicid classification, pp. 1-184 in Zootaxa 5303 (1) on page 129, DOI: 10.11646/zootaxa.5303.1.1, http://zenodo.org/record/8043342, {"references":["Rondani, C. (1848) Esme di varie specie d'insetti ditteri brasiliani. Studi entomologici, Turino, 1, 63 - 112, 3 pls.","Wilkerson, R. C., Linton, Y. - M. & Strickman, D. (2021) Mosquitoes of the world. Vols. 1 & 2. Johns Hopkins University Press, Baltimore, Maryland, 1332 pp. https: // doi. org / 10.1186 / s 13071 - 021 - 04848 - 6","Stone, A. (1944) Notes on the genus Trichoprosopon (Diptera, Culicidae). Revista de Entomologia, 15 (3), 335 - 341.","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","Belkin, J. N., Schick, R. X. & Heinemann, S. J. (1971) Mosquito studies (Diptera, Culicidae) XXV. Mosquitoes originally described from Brazil. Contributions of the American Entomological Institute, 7 (5), 1 - 64.","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","Howard, L. O., Dyar, H. G. & Knab, F. (1915) The mosquitoes of North and Central America and the West Indies. Vol. 3. Systematic description (in two parts). Part I. Publication No. 159. Carnegie Institution of Washington, Washington, D. C., vi + 523 pp.","Bourroul, C. (1904) Mosquitos do Brasil. These, Faculdade de Medicina da Bahia, Officina Typographica, Bahia, viii + 31 pp. [Consisting of Prefacio (pp. i - viii), Novas especies (pp. l - 6, 30 - 32), Quadro das especies encontradas na Bahia (4 pp.) and Proposicies (18 pp.) by Bourroul; remainder and bulk of work by Adolpho Lutz]","Lane, J. & Cerqueira, N. L. (1942) Os sabetineos da America (Diptera, Culicidae). Arquivos de Zoologia do Estudo de Sao Paulo, 3 (9), 473 - 849.","Martini, E. (1931 b) Ueber einige s ¸ damerikanische Culiciden. Revista de Entomologia Rio de Janeiro, 1 (2), 199 - 219.","Ludlow, C. S. (1918) Trichoprosopon Theobald (Diptera; Culicidae). Psyche, 25 (3), 66 - 68. https: // doi. org / 10.1155 / 1918 / 39087","Dyar, H. G. (1928) The mosquitoes of the Americas. Publication No. 387. Carnegie Institution of Washington, Washington, D. C., 616 pp.","Senevet, G. & Abonnenc, E. (1939) Les moustiques de la Guyane Francaise. III. Les sabethines. Archives de l'Institut Pasteur d'Algerie, 17 (2), 247 - 281.","Lane, J. (1953) Neotropical Culicidae. Vols. I & II. University of Sao Paulo, Sao Paulo, 1112 pp.","Zavortink, T. J., Roberts, D. R. & Hoch, A. L. (1983) Trichoprosopon digitatum - morphology, biology, and potential medical importance. Mosquito Systematics, 15 (2), 141 - 149.","Zavortink, T. J. (1981) Species complexes in the genus Trichoprosopon. Mosquito Systematics, 13 (1), 82 - 85."]}

Uranotaenia (Uranotaenia) palmeirimi de Meillon & Rebêlo subspecies dundo da Cunha Ramos, 1993 —original combination: Uranotaenia (Uranotaenia) palmeirimi dundo. Distribution: Angola (Dundo) (da Cunha Ramos 1993). subspecies palmeirimi de Meillon & Rebêlo, 1941 —original combination: Uranotaenia palmeirimi. Distribution: Mozambique, South Africa (Jupp 1996). Uranotaenia palmeirimi was supposedly described from a male (holotype) and a female from the Portuguese East African “Colony of Moçambique ”, a northern coastal area of present-day Mozambique, but “Through an error the type locality was omitted from the text originally describing this species. It was taken at Pebane in Quelimane District” (Worth & de Meillon 1960). Today, Pebane is a town and district in Zambezia Province and Quelimane is the administrative capital of the province. All accounts of palmeirimi contain confusing contradictions. De Meillon & Rebêlo (1941) described the adult of the species without reference to either sex but indicated that two specimens were examined: “Type ♂ in the collection of the South African Institute for Medical Research [SAIMR], Johannesburg” and “One other female from the same locality.” This was taken by catalogers to denote that the species was described from a male and a female (Stone et al. 1959; Knight & Stone 1977; Wilkerson et al. 2021). As revealed by da Cunha Ramos (1993), the listing of a type male by de Meillon & Rebêlo was in error, because the holotype is actually a female, which explains the reference to “One other female…”. Despite this, da Cunha Ramos distinguished the male of the typical form from the male and female of subspecies dundo in a key (see below). Subspecies dundo was named and described based on a series of females from Dundo, Angola, including the holotype female, nine paratype females and 14 topotypic females (da Cunha Ramos 1993). In view of this, it is disturbing that the author contrasted the female and the unknown male with the unknown male of the typical form, as follows (translated from the Portuguese). 2 ― ♂: With a short but well-defined line of pale scales on the sides of the scutum, above the wing root; ppn [postpronotum] with darker, black lower half................................. palmeirimi palmeirimi ― ♂ ♀: Scutum with only a few light scales above the wing root; ppn uniformly light brown........................................................................................... palmeirimi dundo ssp. n. Prior to the introduction of subspecies dundo, Service (1990) provided a revamped version of the original description of the female of palmeirimi and indicated that the male was unknown. The irony is that Service preempted da Cunha Ramos (1993) in noting that the type specimen of palmeirimi is a female: “ Holotype female in the collection of South African Institute of Medical Research, Johannesburg, is badly rubbed and has all legs missing except for one hind leg. In addition there is another female in the collection, not marked as a paratype, but with the same date and collection site as holotype; this specimen has only one fore leg, with tarsomere 5 missing, no wings and is badly rubbed.” Nothing is known about the biology of either subspecies (Service 1990; da Cunha Ramos 1993). According to published records, palmeirimi sensu stricto has been collected at the type locality on the coast of Mozambique and Richards Bay, a town on the coast of KwaZulu-Natal Province, South Africa (Worth & de Meillon 1960; Eckard et al. 1988; Jupp 1996, see note below), whereas subspecies dundo is only known from the type locality of Dundo, Angola, where it was collected in gallery forest amid savannah (da Cunha Ramos 1993) located approximately 24 km south of the border with the Democratic Republic of the Congo. The type locality of dundo lies approximately 2,000 km northwest of the type locality of palmeirimi sensu stricto. Although the adult males, larvae and pupae of the two nominal forms are unknown, considering the distance between the type localities, the coastal versus savannah environment and the morphological features which clearly distinguish the adult females, we believe that discovery and integrated study of the unknown life stages will reveal that the two forms are distinct species. Consequently, we agree with the Encyclopedia of Life that dundo should be recognized as a separate species, at least until proven otherwise, and we hereby recognize it as such: Uranotaenia (Uranotaenia) dundo da Cunha Ramos, 1993 . Note. Wilkerson et al. (2021), in error, indicated that Jupp (1996) provided illustrations of the male and female of Ur. palmeirimi. Jupp distinguished palmeirimi from Ur. hopkinsi Edwards, 1932b in a key to the adults of the subgenus Uranotaenia in southern Africa, but he did not include illustrations for either of the two species.

Anopheles (Cellia) ludlowae (Theobald) subspecies ludlowae (Theobald, 1903a)—original combination: Myzomyia ludlowii. Distribution: India, Indonesia, People’s Republic of China, Philippines, Singapore, Sri Lanka, Taiwan, Thailand (Wilkerson et al. 2021). subspecies torakala Stoker & Waktoedi Koesoemawinangoen, 1949 —original combination: Anopheles (Myzomyia) ludlowi var. torakala (subspecific status by Harbach & Howard 2007). Distribution: Indonesia (Sulawesi) (Stoker & Waktoedi Koesoemawinangoen 1949). Theobald (1903a) described ludlowii (also spelled ludlowi; emendation to ludlowae by Stone 1957) from a series of six female syntypes from “Luzon, Philippine Islands (Miss Ludlow).” Later authors debated whether ludlowae was a fresh water, blackish water or salt water species. Ludlow (1914) further clarified the type locality: “taken by Dr. Graves in the Province of Abra, Island of Luzon, on the Benguet Road during the construction of that road. The location is definitely inland… there is no sea or blackish water within many miles.” Nominotypical ludlowae can be recognized in the adult female by the following: Maxillary palpus with three white bands, two apparently at the apices of palpomeres 2 and 3 and third at the apex of palpomere 4 continuing onto palpomere 5; wing with four main dark spots (presector, sector, preapical, apical), with an apical pale spot, an accessory sector pale spot on R 1 (sometimes continuing onto subcosta but not costa), with pale spots and pale fringe at apices of all veins, pale fringe not evident between any of the veins, vein CuA with three dark spots and 1A with two dark spots; legs with distinct speckling, speckles on the hindlegs often extending onto hindtarsomeres 2 and 3, tarsomeres usually banded apically and sometimes basally (specifically described as basal and apical by Theobald 1903a); abdomen without obvious scales; aedeagal leaflets about five or six per side with the longest leaflet somewhat serrate and sinuous (Russell & Baisas 1936). Characteristics of the larva and pupa of the nominal taxon ludlowae from the Philippines were described by King (1932) and Baisas (1936), respectively, but observations are not available for comparison to subspecies torakala. According to Urbino (1936), the egg of ludlowae does not have floats; the egg of subspecies torakala has not been documented. Subspecies ludlowae is not known as a vector of malarial parasites in the Philippines (Basio 1971). Since there are many species with the overall appearance of the adult female of ludlowae, the name was associated with other taxa as a variety or subspecies. There was initial recognition of a “salt water ludlowae ” in the Philippines. King (1932) determined this to be a separate species (currently An. (Cel.) litoralis King, 1932). Bonne-Wepster & Swellengrebel (1953) wrote: “Early writers used the name ludlowi for the common speckled-legged Anopheles species which occurs all over the Sunda Islands, Malaya and the Andamans and which in later years was also found in India, along the coast in brackish and fresh water. Rodenwaldt (1925) noted several important characters in which the species from Indonesia differs from that on the Philippine Islands and described it as ludlowi var. sundaica.” This variety was later elevated to species rank by Christophers (1933). Harrison et al. (1991) reinforced this idea in a rationale for a list of mosquitoes of Thailand and nearby countries: “ Anopheles (Cel.) ludlowae (Theobald) was listed as doubtful and needing further confirmation. Additional confirmation is not necessary. The old records of ‘ ludlowi ’ by Barnes (1923), Barraud and Christophers (1931) and Thurman (1959) were based on misidentified specimens of An. sundaicus (Rodenwaldt) as noted by Scanlon et al. (1968). Anopheles ludlowae is an insular species, not found on mainland Southeast Asia, and should not be included in the Thailand list.” Harbach & Howard (2007) noted that subspecies torakala was first published in 1938 in Dutch but neither they nor we have been able to obtain a copy of the publication. Therefore: “it is not known whether or not the name was originally introduced in 1938” or at what rank/status. Subspecies torakala is recognized as an important vector of human malarial parasites (van Hell 1952; Basio 1971). A brief description, with illustrations of the maxillary palpus of both sexes, a wing and a hindleg, was later published in English by Stoker & Waktoedi Koesoemawinangoen (1949) as a variety of ludlowi. The type locality, discussed by Kitzmiller (1982), is in South Sulawesi Province. No type specimens are known. Stoker & Waktoedi Koesoemawinangoen briefly described and illustrated, not only torakala, but also Anopheles (Myzomyia) ludlowi (sensu Walch & Soesilo 1929) and Anopheles (Myzomyia) ludlowi “ typicus ” (= sensu Theobald 1903a, i.e. the nominotypical form). The descriptions are brief and do not provide a differential diagnosis or an explanation about specimens or literature examined, but we surmise the following: torakala does not have an accessory sector pale spot on the subcosta and the accessory sector pale spot on vein R 1 is not demarcated by the additional dark mark between it and the sector pale spot (“ typicus ” has a distinct accessory sector pale spot on the subcosta and R 1, with three pale spots on R 1 below the sector dark); torakala does not have a pale fringe spot between CuA and 1A (“ typicus ” has a continuous pale fringe spot from CuA to 1A); in torakala it is difficult to interpret but it appears that vein R 3 does not end in a pale fringe spot (“ typicus ” has a continuous pale fringe spot between the ends of veins R 3 and R 4+5); torakala has CuA with two dark spots (“ typicus ” has CuA with three dark spots); torakala has three dark spots on the subcosta (“ typicus ” with two dark spots on the subcosta). In addition, judging from the illustrations, we do not see significant differences in banding on the maxillary palpi or the speckling of the legs. From the above, a likely significant character is the pale fringe between CuA and 1A, the presence of which is used in keys to distinguish Indonesian ludlowae from torakala, which does not have the spot. In contradiction, this fringe spot is not present on the wing of Philippine ludlowae (sensu stricto). Lee et al. (1987) did not distinguish between ludlowae and ludlowae var. torakala in the Australasian Region and combined them in their key, in which they state that there is usually a pale fringe spot between CuA and 1A. Since only the nominotypical form has been characterized in all life stages, no certain comparisons can be made with Indonesian forms, except for one character of the male genitalia. Bonne-Wepster & Swellengrebel (1953) compared the aedeagal leaflets of Philippine ludlowae with the aedeagal leaflets of males from Sulawesi (the Celebes). Their figure 51 shows that the leaflets of Celebes specimens are long and numerous whereas the leaflets of Philippine specimens, as discussed above, have only five or six leaflets on each side, with the longest sinuous or S-shaped. In summary, because of commonly shared characters in this group of species, we think that the concept of ludlowae from the Philippines has been mixed and confused with what probably is a species complex in Sulawesi, and elsewhere in Indonesia. How to adequately characterize torakala is not possible without further study, especially utilizing molecular methods. Because of noted differences in the transmission of malarial parasites, the presence of pale fringe between CuA and 1A in torakala and differences in the form of aedeagal leaflets, we think these are probably separate species, and therefore elevate torakala to species status: Anopheles (Cellia) torakala Stoker & Waktoedi Koesoemawinangoen, 1949 . Anopheles torakala is currently listed as a species in the Encyclopedia of Life. Three names (formosaensis, hatorii, cabrerai) are currently in synonymy with ludlowae / torakala. Since those nominal species do not occur in the geographical range of An. torakala, we believe all three are synonyms of An. ludlowae. Anopheles ludlowii var. formosaensis Koidzumi, 1917 was preoccupied and replaced by An. hatorii Koidzumi, 1920, as explained by Yamada (1925): “ Koidzumi (1917) pointed out, however, the Formosan form differs slightly from ludlowii and placed the former as a variety of the latter, giving it a name, Anopheles ludlowii var. formosensis. After that, Koidzumi (1920) leaned to separate the Formosan form specifically from ludlowii and gave it provisionally another name A. hatorii nov. sp. (?) replacing formosensis preoccupied. But the provisional name was lately made valid by the author (1924). It was, however, placed again as a synonym of ludlowii by Christophers (1924).” Anopheles (Cellia) ludlowae ssp. cabrerai Cagampang-Ramos & Darsie, 1969 (in Darsie & Cagampang-Ramos 1969) was described in comparison with ludlowae as having three dark spots on vein 1A instead of two. The authors (Darsie & Cagampang-Ramos 1977) later determined that this single character was variable and recognized cabrerai as a synonym of ludlowae., 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 67-68, DOI: 10.11646/zootaxa.5303.1.1, http://zenodo.org/record/8043342, {"references":["Theobald, F. V. (1903 a) A monograph of the Culicidae or mosquitoes. Vol. 3. British Museum (Natural History), London, xvii + 1 (errata) + 359 pp., foldout table, 17 pls. https: // doi. org / 10.5962 / bhl. title. 58067","Wilkerson, R. C., Linton, Y. - M. & Strickman, D. (2021) Mosquitoes of the world. Vols. 1 & 2. Johns Hopkins University Press, Baltimore, Maryland, 1332 pp. https: // doi. org / 10.1186 / s 13071 - 021 - 04848 - 6","Stoker, W. J. & Waktoedi Koesoemawinangoen, R. (1949) Illustrated map of the anopheline imagines of Indonesia (English translation). Ministry of Health of Indonesia, Djakarta, 69 pp.","Harbach, R. E. & Howard, T. M. (2007) Corrections in the status and rank of names used to denote varietal forms of mosquitoes (Diptera: Culicidae). Zootaxa, 1542 (1), 35 - 48. https: // doi. org / 10.11646 / zootaxa. 1542.1.3","Ludlow, C. S. (1914) Anopheles (Myzomyia) ludlowii Theobald. Psyche, 21 (1), 32 - 33. https: // doi. org / 10.1155 / 1914 / 70413","Russell, P. F. & Baisas, E. (1936) A practical illustrated key to adults of Philippine Anopheles. Philippine Journal of Science, 59 (1), 15 - 64, 34 pls.","King, W. V. (1932) The Philippine Anopheles of the rossi-ludlowi group. Philippine Journal of Science, 47 (3), 305 - 342, 4 pls.","Baisas, F. E. (1936) Notes of Philippine mosquitoes, VI. The pupal characters on anophelines of the subgenus Myzomyia. Philippine Journal of Science, 61 (2), 205 - 220, 25 pls.","Urbino, C. M. (1936) The eggs of some Philippine Anopheles. Monthly Bulletin of the Bureau of Health, 16, 261 - 275.","Basio, R. G. (1971) The mosquito fauna of the Philippines (Diptera Culicidae). Monograph No. 4. National Museum of the Philippines, Manila, v + 198 pp.","Bonne-Wepster, J. & Swellengrebel, N. H. (1953) The anopheline mosquitoes of the Indo-Australian Region. J. H. de Bussy, Amsterdam, 504 pp.","Rodenwaldt, E. (1925) Entomologische notities. III. Geneeskundig Tijdschrift voor Nederlandsch-Indie, 65, 173 - 201, 3 pls.","Christophers, S. R. (1933) The fauna of British India, including Ceylon and Burma. Diptera. Vol. IV. Family Culicidae. Tribe Anophelini. Taylor and Francis, London, vi + (iii) + 371 pp., 3 pls.","Harrison, B. A., Rattanarithikul, R., Peyton, E. L. & Mongkolpanya, K. (1991) Taxonomic changes, revised occurrence records and notes on the Culicidae of Thailand and neighboring countries. Mosquito Systematics, 22 (3), 196 - 227. [for 1990]","Christophers, S. R. & Barraud, P. J. (1931) The eggs of Indian Anopheles, with descriptions of the hitherto undescribed eggs of a number of species. Records of the Malaria Survey of India, 2 (1), 161 - 192, 5 pls.","van Hell, J. C. (1952) The anopheline fauna and malaria vectors in South Celebes. Documenta de Medicina Geographica et Tropica, 4, 45 - 56.","Kitzmiller, J. B. (1982) Anopheline names: their derivations and histories. The Thomas Say Foundation. Vol. VIII. Entomological Society of America, College Park, Maryland, vi + 639 pp. https: // doi. org / 10.4182 / UTTN 3411","Walch, E. W. & Soesilo, R. (1929) A comparative study of the pecten of the Netherlands Indies Anopheles larvae, preceded by some other morphological observations. Mededelingen van den Dienst der Volksgezondheid in Nederlandsch-Indie, 18, 453 - 468.","Koidzumi, M. (1917) On the anopheline mosquitoes from Formosa. Dobutsugaku Zassi [Zoological Magazine], 29, 131 - 142, 351 - 355. [in Japanese]","Koidzumi, M. (1920) Preventive medical study of mosquitoes in Formosa. Daiwan Kenkyujo Hokoku [Report of the Formosa Government], 8, 1 - 158. [in Japanese]","Yamada, S. (1925) A revision of the adult anopheline mosquitoes of Japan: systematic descriptions, their habits and their relations to human diseases. (Part II). Scientific Reports from the Government Institute for Infectious Diseases, Tokyo Imperial University, 4, 447 - 493.","Christophers, S. R. (1924) Provisional list and reference catalogue of the Anophelini. Indian Medical Research Memoirs, 3, 1 - 105.","Darsie, R. F., Jr. & Cagampang-Ramos, A. (1969) Manual of malaria entomology. A handbook of laboratory and field training exercises. 2 nd Edition. Malaria Eradication Training Center, Manila, vi + 142 pp.","Darsie, R. F., Jr. & Cagampang-Ramos, A. (1977) A study of Anopheles ludlowae Theobald siblings, leading to new synonymy (Diptera, Culicidae). Mosquito Systematics, 9 (2), 123 - 127."]}

Culiseta (Culicella) ochroptera (Peus) subspecies amurensis Maslov, 1964 —original combination: Culiseta silvestris amurensis (status as subspecies of ochroptera by Berlov & Kuberskaya 2023). Distribution: Central Amur region, Ussuri Basin, Southern Maritime Province, Northeast China (Maslov 1964); Primorye (Primorsky Krai, far East Region), Russia (Maslov 1964). subspecies minnesotae Barr, 1957 —original combination: Culiseta minnesotae (status as subspecies of ochroptera by Berlov & Kuberskaya 2023). Distribution: Canada, United States (Alaska, continental) (Wilkerson et al. 2021). subspecies nipponica La Casse & Yamaguti, 1950 —original combination: Culiseta (Culicella) nipponica (status as subspecies of ochroptera by Berlov & Kuberskaya 2023). Distribution: Japan, South Korea (Wilkerson et al. 2021). subspecies ochroptera (Peus, 1935) —original combination: Theobaldia (Culicella) ochroptera. Distribution: Belarus, Belgium, Czech Republic, Estonia, Finland, Germany, Hungary, Lithuania, Netherlands, People’s Republic of China, Poland, Romania, Russia, Slovakia, Sweden, Ukraine (Wilkerson et al. 2021). Soon after submission of this monograph for review, Berlov & Kuberskaya (2023) published a paper titled “Additions and corrections to the catalog of blood-sucking mosquitoes of the world (Insecta: Diptera, Culicidae) by Wilkerson et al. (2021) ” [verbatim translation from the Russian]. The authors inferred that Culiseta amurensis Maslov, 1964, Cs. minnesotae Barr, 1957, Cs. nipponica LaCasse & Yamaguti, 1950 and Cs. ochroptera (Peus, 1935) were incorrectly listed in the catalog as valid species. They pointed out that Maslov (1964, 1967, 1989) classified them as subspecies of Cs. silvestris Shingarev, 1928. However, because Cs. silvestris has been treated as a doubtful species since Dahl & White (1978) listed it, without explanation, as a nomen dubium in a footnote to a table of European mosquito species, Berlov & Kuberskaya proposed that amurensis, minnesotae and nipponica should be recognized as subspecies of ochroptera, the oldest of the four nominal taxa. To gain a better understanding of this taxonomic conundrum, we examined Maslov’s (1964) treatment of Cs. silvestris. The reasoning behind his recognition of Cs. silvestris and its purported subspecies is revealed in the following passages (translated from the Russian). … C. silvestris Sching. ‒ was described very unsuccessfully, too fragmentary and general, so much so that it was difficult to extract anything significant from the description (Shingarev, 1928). Not surprisingly, Peus (1935) later described the same species under the name C. ochroptera Peus. It is under this name that he [Peus] now appears in the literature. Unfortunately, the holotype and allotype of C. silvestris have not been preserved, but in the year of description (1928) N. I. Shingarev [the author of silvestris] gave me one male, one female, and them as paratypes. Later from the Moscow region, in the area where C. silvestris was first found, I also managed to obtain larvae of this species. Thus, at present, it seems to me necessary in the revision of the Eurasian representatives of the subgenus Culicella to establish the following taxonomic divisions: the species name C. silvestris with three subspecies 1 C. silvestris silvestris Sching. ‒ Eastern Europe, the Urals, Western Siberia; C. silvestris ochroptera Peus ‒ Central Europe and the Baltics; C. silvestris amurensis Masl. (subsp. n.) (Maslov, 1949, 1963) ‒ the extreme east of the species range ‒ Piamurye and Primorye. Below is a redescription of the species with a differential diagnosis of all subspecies and a description of C. silvestris amurensis, subsp. n. 1 It is possible that when analyzing more extensive European material, the first two forms [silvestris sensu stricto and ochroptera] will turn out to be one and the same subtype. In Japan, another species was described (La Casse а. Yamaguti, 1950), assigned by the authors to the subgenus Culicella, C. nipponica. Unfortunately, adult females and males of this species remain unknown; as for the larva of the 4th stage, then differentiating C. nipponica from C. silvestris amurensis n. is not possible (Maslov, 1963). It is probable that there are not two species here, even sub-indigenous, but one form, and then the Far Eastern C. silvestris amurensis n. would prove to be synonymous with Japanese C. nipponica, which would need to be considered as a subspecies, C. silvestris nipponica. However, at the present time, due to the lack of descriptions of the male and female already indicated, both forms have to be preserved. Recently, another species of Culicella ‒ C. minnesotae Barr, was described in the USA (Barr, 1957, 1959; Price, 1958). Among the mosquitoes I received from the USA (Wisconsin) from prof. R. Matheson, turned out to be one male matching the description (Barr, 1957), especially in the structure of the phallus. Comparison of American C. minnesotae with other representatives of the subgenus Culicella confirmed significant differences in all developmental stages from C. morsitans morsitans and C. morsitans dyari. However, an amazing similarity between C. minnesotae and C. silvestris mosquitoes was unexpectedly revealed: the most characteristic imaginal signs of one (spotted wing, the presence of not only basal but also apical bands on the abdominal tergites [terga], etc.) coincided with those of the other. Of particular interest was the similarity in the structure of the phallus [genitalia] of C. minnesotae and C. silvestris silvestris (Fig. 9, A, B). A comparison of the larval morphological structures also does not support the species independence of C. minnesotae Barr. The same is shown by the details of pupal chaetotaxy (Maslov, 1963). Thus, C. minnesotae should not be considered as an independent species, but only as a subspecies ‒ C. silvestris minnesotae Barr. Following the above, Maslov (1964) distinguished the females, males and fourth-instar larvae of subspecies amurensis, minnesotae, ochroptera and silvestris in separate keys, accompanied by descriptions of silvestris sensu lato and the new subspecies amurensis. Maslov (1967, 1989) incorporated the keys into keys for distinguishing the females, males and fourth-instar larvae of all known species of Culiseta and provided descriptions for silvestris sensu stricto and each of the four subspecies. Because Maslov (1964) had examined paratypes and topotypic specimens of silvestris, and recognized ochroptera as the same species, we were inclined to consider ochroptera as a synonym of silvestris until we noticed that Gutsevich et al. (1971, 1974) justified the retention of ochroptera: “A. V. Maslov (1964) considered C. ochroptera as a synonym of C. silvestris Shingarev, 1928, but to judge from Shingarev’s incomplete description, ‘it is certain that the two species are not identical’ (Shtakel’berg, 1937 [Stackelberg 1937]). The holotype of C. silvestris is lost and Maslov examined only paratypes. Because of the incomplete original description of C. silvestris, it seems advisable to retain the name C. ochroptera [translated from the Russian].” It is important to note here that Stackelberg (1937) treated ochroptera and silvestris as separate species (of Theobaldia Neveu-Lemaire, 1902) [the map in Maslov (1967, 1989: fig. 82) indicates they may have allopatric distributions], and distinguished them in a key for the identification of males. It seems likely that Dhal & White (1978) listed silvestris as a nomen dubium based on the explanatory note and recommendation proffered by Gutsevich et al. (1971, 1974). Until the identity of silvestris is resolved, we concur with Dahl & White that it should remain a nomen dubium. Wood et al. (1979) provided a cogent summary of the silvestris-ochroptera conundrum and decided to continue to recognize minnesotae as a separate species “Until Russian workers concur in choosing between ochroptera and silvestris ”. We agree with this rationale and reaffirm the specific rank of minnesotae established by Wood et al. (1979), and accepted by later workers (e.g. Darsie & Ward 1981, 2005; Belton 1983; Harrison et al. 2016): Culiseta (Culicella) minnesotae Barr, 1957 . Culiseta minnesotae is currently listed as a species in the Encyclopedia of Life. Berlov & Kuberskaya (2023) incorrectly asserted that Maslov (1967, 1989) had considered nipponica to be a subspecies of silvestris. All prior authors (Maslov (1964, 1967, 1989; Gutsevich et al. 1971, 1974; Tanaka et al. 1979; Lu et al. 1997) recognized nipponica as a distinct species. As there is no justification or precedence for treating nipponica as a subspecies, the specific rank of this nominal species must be retained: Culiseta (Culicella) nipponica La Casse & Yamaguti, 1950 , as listed in the Encyclopedia of Life. We have not been able to find a source for the recognition of amurensis as a species; consequently, its listing as a species by Harbach (2018) was apparently in error; it should have been listed as a subspecies of ochroptera. The error was repeated in Wilkerson et al. (2021). Although amurensis is recorded from the extreme northeast of China (Knight & Stone 1977; Wilkerson et al. 2021), it is not treated or even mentioned by Lu et al. (1997) in their monograph on the culicine mosquitoes of China (however, they do recognize and describe Cs. nipponica). As noted above, Maslov (1964) thought it “probable” that amurensis was the same as nipponica, in which case the apparent morphological similarity of the two nominal forms may explain why Lu et al. only recognized the presence of nipponica in China. In agreement with Maslov (1964), we also believe amurensis is likely to be synonymous with nipponica; hence, until proven otherwise, it is herewith consigned to synonymy: amurensis Maslov, 1964, junior subjective synonym of Culiseta (Culicella) ochroptera (Peus, 1935) . The nominal subspecies amurensis, which is listed as a species in the Encyclopedia of Life, must be removed from the list of valid species of Culiseta., 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 106-108, DOI: 10.11646/zootaxa.5303.1.1, http://zenodo.org/record/8043342, {"references":["Maslov, A. V. (1964) On the systematics of bloodsucking mosquitoes of the group Culiseta (Diptera, Culicidae). Entomologicheskoe Obozrenie, 43 (1), 193 - 217. [in Russian]","Berlov, O. E. & Kuberskaya, O. V. (2023) Additions and corrections to the Catalog of blood-sucking mosquitoes of the world (Insecta: Diptera, Culicidae) by Wilkerson et al. (2021). Amurian Zoological Journal, 15 (1), 110 - 118. [in Russian] https: // doi. org / 10.33910 / 2686 - 9519 - 2023 - 15 - 1 - 110 - 118","Barr, A. R. (1957) A new species of Culiseta (Diptera: Culicidae) from North America. Proceedings of the Entomological Society of Washington, 59 (4), 163 - 167.","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","La Casse, W. J. & Yamaguti, S. (1950) Mosquito fauna of Japan and Korea. Office of the Surgeon, Hq. 8 th Army APO 343, Kyoto, Honshu, viii + 268 + 7 (Appendix I) + 213 (Appendix II) pp.","Peus, F. (1935) Theobaldia (Subg. Culicella) ochroptera sp. n., eine bisher unbekannte Stechm ¸ cke. Markische Tierwelt: Zeitschrift f ¸ r die faunistische Erforschung der Kurmark, 1, 113 - 121.","Yamaguti, S. & LaCasse, W. J. (1950) Mosquito fauna of Guam. Office of the Surgeon, Headquarters, 8 th Army, APO 343. United States. 207 th Malaria Survey Detachment. Office of the Surgeon, Washington, D. C., 101 pp.","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.","Shingarev, N. L. (1928) Notes on Culicidae. III. Russkii Zhurnal Tropicheskoi Meditsiny [Russian Journal of Tropical Medicine], 1, 47 - 53. [in Russian]","Dahl, C. & White, G. B. (1978) Culicidae. In: Illies, J. (Ed.), Limnofauna Europaea. Gustav Fischer Verlag, New York, Stuttgart, pp. 390 - 395.","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]","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]","Neveu-Lemaire, M. (1902) Sur la classification des Culicides. Comptes Rendus Hebdomadaires des Seances et Memoires de la Societe de Biologie, 54, 1329 - 1332.","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.","Darsie Jr., R. F. & Ward, R. A. (1981) Identification and geographical distribution of the mosquitoes of North America, north of Mexico. Mosquito Systematics Supplement, 1, 1 - 313.","Darsie Jr., R. F. & Ward, R. A. (2005) Identification and geographical distribution of the mosquitoes of North America, north of Mexico. University Press of Florida, Gainesville, Florida, 383 pp.","Belton, P. (1983) The mosquitoes of British Columbia. Handbook 41. British Columbia Provincial Museum, Victoria, British Columbia, 199 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.","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.","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.","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","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."]}

Aedes (Aedimorphus) durbanensis (Theobald) subspecies angolae Ribeiro & Ramos, 1974 —original combination: Aedes durbanensis angolae. Distribution: Angola (Ribeiro & Ramos 1974). subspecies durbanensis (Theobald, 1903a)—original combination: Grabhamia durbanensis. Distribution: Angola, Congo, Democratic Republic of the Congo, Ethiopia, Madagascar, Mozambique, South Africa, Tanzania (Wilkerson et al. 2021). The nominotypical subspecies was described from a single female from Durban, South Africa [KwaZulu-Natal Province]. It was characterized as follows (after Theobald 1903a): Head with silvery gray narrow curved scales; clypeus black, with a tubercle on each side near base; proboscis pale brownish yellow overall, apex and base black; maxillary palpus densely scaled, brown basally, white at apex; antenna deep brownish yellow with first 3 flagellomeres dull brick red, first flagellomere with white scales, second and third with dark scales; thorax with rich brown ornamentation, to some extent forming ornamentation at the sides, scutellum with pale narrow curved scales, pleura brown with gray scales; femora and tibiae mottled with pale scales, tarsomeres with narrow pale basal bands; wing mottled with pale and dark scales; abdomen black, with basal white bands and apicolateral yellow spots. Theobald emphasized that “It can be at once told by the thoracic ornamentation, peculiar mottled wings, and basal abdominal banding, and apical yellow lateral spots.” The male genitalia of the type form were described and illustrated by Edwards (1941), who noted: “Style [gonostylus] with a row of long curved hairs [setae] on distal margin and many more hairs on the surface, spine [gonostylar claw] inserted well before tip at broadest part of style; no apical hair-tuft on coxite [gonocoxite], basal lobe [basal mesal lobe] peculiar in having a regular row of hairs on its margin.” These features were also reported by Ribeiro & Ramos (1974) and Jupp (1996). The pupal stage of durbanensis was very briefly described and the trumpet and paddle illustrated by de Meillon et al. (1945). The egg was described in detail by Linley & Service (1994). Subspecies angolae was described and illustrated in direct comparison with durbanensis by Ribeiro & Ramos (1974) using female, male and larval specimens of angolae from Moçâmedes, Namibe Province, coastal Angola, 15º 12′ S, 12º 09′ E [verbatim coordinates not verified]. Specimens of durbanensis used in the description were from Natal, South Africa and Lourenço Marques, Mozambique. The authors stated: “In the female sex, Ae. durbanensis angolae subsp. nov. cannot be distinguished from the nominate subspecies.” However, they pointed out diagnostic characters in the male genitalia and larva (see below). Note that Ribeiro & Ramos were motivated to sort out the identity of these taxa, along with a similar species found on the Angolan coast, Ae. natronius Edwards, 1932b: “it became clear not only that the description of the larva, given by Rebelo and Pereira (1) [1943, not seen by us] and taken by Hopkins (2) [1952], was not a satisfactory one but also that a subspecific treatment might be given to the angolan [sic] population of Ae. durbanensis.” For us, the larva of putative durbanensis described and illustrated by Hopkins (1952) is another, yet to be determined, species. It exhibits an incomplete saddle, short anal papillae (as in angolae), a straight row of comb scales and very short pecten spines. Males of the two forms differ, according to Ribeiro & Ramos (1974), as follow [figure numbers refer to the original publication]: “Males of both subspecies, however, can be readily separated on the basis of genitalic characters. While Ae. durbanensis angolae subsp. nov. (Fig. 4) has a row of l0–12 setae along the distal margin of basal lobe of coxite [gonocoxite] and has only subequal, small setae on the lateral surface of style [gonostylus], the nominate subspecies (Fig. 1) has only 5 or 6 setae along distal margin of basal lobe of coxite and the 2 or 3 most basal setae of its style are unusually long, about 1/3 or more the length of the style itself.” Other differences are illustrated but we did not undertake further study. Distinguishing differences of larvae are summarized as follow. In durbanensis: Comb scales asymmetrical, 7 or 8 (6–19) in a single row; siphon longer, index 2.2–2.5 (2.4); pecten spines moderately long and pointed, longest spine 5 or 6 times longer than width at base, mostly asymmetrical with a few basal denticles, distal 2 or 3 spines unequally spaced; anal papillae slightly longer than saddle, with round lanceolate apices. In angolae: Comb scales symmetrical, about 10 (8–20) arranged in 1 or 2 irregular rows; siphon shorter, index 1.7–2.0 (1.9); pecten spines very long, finely drawn out, longest spine about 7 to 10 time longer than width at base, also with a few basal denticles, spines equally spaced on siphon; anal papillae not or only slightly longer than saddle, ovoid or round lanceolate. Based on the above diagnostic characters, and the great distance between the type localities, without evidence of intermediate forms, we think the two nominal taxa are separate species and here elevate angolae to species status: Aedes (Aedimorphus) angolae Ribeiro & Ramos, 1974 . Aedes angolae is currently listed as a species in the Encyclopedia of Life; however, the authorship is incorrectly attributed to only the first author. Synonym Grabhamia ocellata Theobald, 1910 was described from four females and three males from Maputo Bay (‘Delagoa Bay’) [Townsend 1990]. Theobald (1910) listed the collection site as “ Habitat.―Lourenco Marques.” Maputo is the capital of Mozambique, which has the official name of Lourenço Marques. Geographically, the type localities of durbanensis and ocellata are very close. We think the description of the female of ocellata does not differ significantly from that of durbanensis, except for two prominent black spots on the thorax in ocellata, presumably on the scutal fossae. We have not examined the syntype series to verify this. However, for some reason, Theobald did not compare ocellata with durbanensis. Edwards (1911b) stated “The type of G. durbanensis is a small specimen, but evidently the same as G. ocellata.” This implied synonymy has been followed since. However, the lack of comment from Theobald, the distinctive dark eye-like thoracic spots and Theobald’s statement that “The genitalia [of ocellata] are very marked.”, referring to the large paddle-shaped gonostylus, also seen in later treatments of durbanensis, indicate that a closer look is needed to verify its status. For now, we retain it as a synonym of Ae. (Aed.) durbanensis (Theobald)., 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 12-13, DOI: 10.11646/zootaxa.5303.1.1, http://zenodo.org/record/8043342, {"references":["Ribeiro, H. & Ramos, H. C. (1974) Research on the mosquitoes of Angola. VII - Redescription of the larva of Aedes durbanensis durbanensis (Theo., 1903) and description of Aedes durbanensis angolae subsp. nov. Mosquito Systematics, 6 (2), 93 - 98.","Theobald, F. V. (1903 a) A monograph of the Culicidae or mosquitoes. Vol. 3. British Museum (Natural History), London, xvii + 1 (errata) + 359 pp., foldout table, 17 pls. https: // doi. org / 10.5962 / bhl. title. 58067","Wilkerson, R. C., Linton, Y. - M. & Strickman, D. (2021) Mosquitoes of the world. Vols. 1 & 2. Johns Hopkins University Press, Baltimore, Maryland, 1332 pp. https: // doi. org / 10.1186 / s 13071 - 021 - 04848 - 6","Edwards, F. W. (1941) Mosquitoes of the Ethiopian Region III. - Culicine adults and pupae. Printed by order of the Trustees, British Museum (Natural History), London, viii + 499 pp.","Jupp, P. G. (1996) Mosquitoes of southern Africa: Culicinae and Toxorhynchitinae. Ekogilde Publishers, Hartebeespoort, 156 pp.","de Meillon, B., Parent, M. & Black, L. O'C. (1945) Descriptions of new larvae and pupae of Ethiopian Culicini. Bulletin of Entomological Research, 36 (1), 85 - 101. https: // doi. org / 10.1017 / S 0007485300023932","Linley, J. R. & Service, M. W. (1994) Fine structure of the eggs of Aedes durbanensis, Aedes woodi, and Eretmapodites quinquevittatus (Diptera: Culicidae). Mosquito Systematics, 26 (2), 59 - 72.","Edwards, F. W. (1932 b) Mosquito notes. - XI. Bulletin of Entomological Research, 23 (4), 559 - 562. https: // doi. org / 10.1017 / S 0007485300004338","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.","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","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.","Edwards, F. W. (1911 b) The African species of Culex and allied genera. Bulletin of Entomological Research, 2 (3), 241 - 268. https: // doi. org / 10.1017 / S 0007485300001450"]}

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

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

Aedes (Ochlerotatus) fulvus (Wiedemann) subspecies fulvus (Wiedemann, 1828) —original combination: Culex fulvus. Distribution: Argentina, Belize, Bolivia, Brazil, Colombia, Costa Rica, Cuba, Ecuador, French Guiana, Guatemala, Guyana, Mexico, Panama, Peru, Suriname, Trinidad and Tobago, Venezuela (Wilkerson et al. 2021, South Korea listed in error). subspecies pallens Ross, 1943 —original combination: Aedes (Ochlerotatus) fulvus pallens. Distribution: United States (continental) (Wilkerson et al. 2021). Distributions listed in Wilkerson et al. (2021) for fulvus sensu stricto from the continental United States and for pallens from Cuba and Mexico could not be documented and should be omitted. The nominal forms are members of the Chrysoconops Group of Wilkerson et al. (2015) —genus Chrysoconops Goeldi, 1905; Group C, fulvus -group of Edwards (1932a); Ochlerotatus subgenus Chrysoconops of Reinert et al. (2008, 2009). In addition to the two nominal taxa, the group also includes bimaculatus (Coquillett, 1902b); jorgi Carpintero & Leguizamón, 2000; pennai Antunes & Lane, 1938; and stigmaticus Edwards, 1922. Wiedemann (1828) described the nominotypical subspecies from “ Brazil ”. The holotype female was seen by Belkin (1968) in the collection of the Senckenberg Naturmuseum Frankfurt, Germany. Belkin et al. (1971) later restricted the type locality to Salvador, Bahia, Brazil. Goeldi (1905) described the egg of fulvus (as Taeniorhynchus fulvus) from a female captured in Murutucú, near Belém, Brazil. He was, however, unable to rear immature stages for study. Goeldi provided a description of the adult from separate observations: “Among our mosquitoes it is one of the largest species. It is characterized, in addition to its distinct size, by the beautiful general yellow coloration, color of gold, that extends over all [body] parts, including a good part of the anterior margin of the wings, contrasting here on the wing with the distal margin having a dark spot, while the rest shines giving a beautiful iridescent effect. Also ostensibly distinctive are the dark middle articulations of all six pairs [sic] of legs, the tarsi, the tip of the proboscis and the tip of the palps [maxillary palpi]. The back of the thorax [scutum] and the posterior margins of the abdominal rings [bands] exhibit some areas of a slightly different tint, darker than the beautiful general golden yellow [translated from the Portuguese].” In his description of subspecies pallens, Ross (1943) contrasted the subspecies with fulvus, and with the very similar Ae. bimaculatus. For reference, Ross included detailed illustrations of the male genitalia and larvae of fulvus pallens and bimaculatus. He wrote:“This paper presents evidence to show that two distinct species of Aedes occurring in the United States are both at present identified as Coquillett’s bimaculatus. The true bimaculatus, described from Brownsville, Texas, and ranging from central Texas to El Salvador, is very distinct from the ‘ bimaculatus ’ collected throughout the southeastern United States which is here described as a new subspecies of the Neotropical fulvus (Wiedemann). Vargas’s rozeboomi, recently described from Campeche, Mexico, is shown to be a synonym of the true bimaculatus.” After describing the “true bimaculatus ”, Ross compared the species with fulvus as follows: “The male terminalia [genitalia] characters of bimaculatus are very distinctive and set the species well apart from fulvus [sensu lato]. The larval differences between bimaculatus and fulvus pallens, on the other hand, are relatively slight. Adults of both sexes of these species may be separated as follows: fulvus [sensu lato] has apical triangular areas of black scales on all abdominal tergites [terga], thoracic pleura with at least one black integumental spot (fulvus pallens) or two longitudinal black stripes (fulvus fulvus); bimaculatus, with scarcely any black scales on tergites except toward base of abdomen, thoracic pleura yellow—no integumental maculation”. Before describing fulvus fulvus, Ross (1943) stated: “The writer has before him the fulvus material in the U. S. National Museum from many localities in Central and South America. Representatives from Panama [not Brazil] are described...”. The following are pertinent excerpts from his description of the male: “Thorax with mesonotal [scutal] integument lemon-yellow except for the subbasal spots [posterolateral spots]; each spot is transversely divided medially by a yellow area, the spots are brownish-black with blending margins.... Pleural integument lemon-yellow with two longitudinal brownish-black bands, one extending caudad from side of anterior promontory of mesonotum to prealar sclerite [prealar area], the lower band crossing middle of sternopleural sclerite [mesokatepisternum] and covering lower half of mesepimeron.... Male terminalia [genitalia] without significant differences from that described later for fulvus pallens (fig. 1) [figure number from original].” Following his description of “ fulvus pallens, new subspecies ”, Ross noted: “Because of the lack of apparent terminalia differences between the United States series and that from Panama, and because the more superficial characters such as color and vestiture are relatively slight, though constant, the United States series is placed as a subspecies of fulvus. Pallens can be separated at once from typical fulvus by the almost complete absence of pleural maculation of the thorax and by the greater development of the mesothoracic [posterolateral scutal] spots.” We could not find any evidence of sympatry or intermediate forms of the two subspecies— pallens is found only in the southeastern United States and fulvus is distributed from southern Mexico and the Caribbean (Cuba) to Brazil, but does not occur in the United States. We disagree that lack of male genitalia differences is a reason to ignore significant diagnostic characters and quite disparate distributions. Because of these factors, we believe that fulvus and pallens are separate species. We therefore afford species status to Aedes (Ochlerotatus) pallens Ross, 1943 . Aedes pallens is currently listed as a species in the Encyclopedia of Life. Aedes pallens has one synonym, Culex flavicosta Walker, 1856. Edwards (1932a) was apparently the first to treat flavicosta as a synonym of fulvus. Alan Stone saw the type specimen, and in an unpublished taxonomic catalog research note (1955) he wrote: “British Museum: A ♀ type labeled Amaz. [Amazon] in BM, lacking head. The triangular apical areas of dark scales on the abdomen are scarcely visible, but it’s [sic] synonymy with fulvus [sic] is probably correct.” The holotype was also examined by Belkin et al. (1971) in the Natural History Museum, London, who, instead of “Amazon Region,” restricted the type locality to “Manaus (Amazonas)”. Townsend (1990) also examined the holotype and noted “head missing”, but he did not note that Belkin had restricted the type locality. While it is clear to us that pallens should be accorded species status, variation can be found in adult characters of fulvus in Central and South America and the Caribbean. This variation, and the highly unlikely occurrence of any mosquito species naturally occurring in very different zoogeographic areas, make the existence of a species complex likely. In addition, we are not aware of studies of specimens from the generalized Amazonian type localities, nor the restricted type localities, of fulvus or the synonym flavicosta. A few examples of morphological variation follow. Gutsevich & Garcia-Avila (1969) described Ae. fulvus from Cuba. Their description is not entirely clear but, in part: “The lateral surface of the thorax is also of two colors: upper half dark brown; lower half yellowish. A spot of broad silvery scales on the upper part of the sternopleuron [mesokatepisternum] and on the upper mesepimeron.... The specimens we have collected occupy an intermediate position between the southern and North American forms [between fulvus fulvus and fulvus pallens] [translated from the Spanish].” The thoracic pleura being dark above and yellowish below has not been described for any other taxon related to fulvus. The broad silvery scales usually have not been emphasized but are of possible significance in the Chrysoconops Group. These scales were also noted by Carpenter & LaCasse (1955) for fulvus pallens and by Ross (1943) for fulvus fulvus. Rodriquez-Martinez et al. (2020) documented the sympatry of bimaculatus and fulvus in southern Mexican states bordering the Gulf of Mexico and Guatemala. To identify fulvus: “I distinguish this species from the preceding [stigmaticus, which was shown by them not to occur in Mexico] by the dark bands of the pleura, by the gold-colored scales in the region of the subcosta, and by the initial dark apical part [of the wing] at the point where the subcosta ends on the costal vein [translated from the Spanish].” Martini (1935) also mentioned the apically dark wings of fulvus: “A series of example specimens from South America has, without exception, the characteristic of a dark spot at the end of the wings [translated from the Spanish].” We found wide variability in the descriptions of the extent of yellow scales and dark apical areas of the wings in this group of species. The division of the posterolateral dark scutal spots, a character used by Ross (1943) as characteristic of fulvus, is not mentioned by Rodriguez-Martinez et al. (2020), but an undivided dark scutal spot is clearly shown in a photograph (their fig. 1b). The dark pleural lines, also characteristic of fulvus according to Ross (1943), are present in a photograph of Rodriguez-Martinez et al. (2020: fig. 1d), but the authors of both publications only pointed out that there is a dark spot in the hypostigmal area, but did not discuss the two obvious dark pleural bands. In keys to the mosquitoes of Guatemala (Clark-Gil & Darsie 1983), the characters used to identify adult females of fulvus were “Scutal integument yellow with posterolateral dark marks” and “Thoracic pleuron with dark hypostigmal spot [not two dark bands]; abdominal terga yellow-scaled basally, dark-scaled apically.”, 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 34-36, DOI: 10.11646/zootaxa.5303.1.1, http://zenodo.org/record/8043342, {"references":["Wiedemann, C. R. W. (1828) Aussereuropaische zweifl ¸ gelige Insekten: als Fortsetzung des Meigenschen Werkes. Vol. 1. in der Schulzischen Buchhandlung, Hamm, xxxii + 608 pp. https: // doi. org / 10.5962 / bhl. title. 14603","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","Ross, E. S. (1943) The identity of Aedes bimaculatus (Coquillett) and a new subspecies of Aedes fulvus (Wiedemann) from the United States (Diptera, Culicidae). Proceedings of the Entomological Society of Washington, 45 (6), 143 - 151.","Wilkerson, R. C., Linton, Y. - M., Fonseca, D. M., Schultz, T. R., Price, D. C. & Strickman, D. A. (2015) Making mosquito taxonomy useful: A stable classification of tribe Aedini that balances utility with current knowledge of evolutionary relationships. PloS ONE, 10 (7), e 0133602. https: // doi. org / 10.1371 / journal. pone. 0133602","Goeldi, E. A. (1905) Os mosquitos no Para: Reuniao de quatro trabalhos sobre os mosquitos indigenas, principalmente as especies que molestam o homem. Memorias do Museu Goeldi (Museu Paraense) de Historia Natural e Ethnographia IV. C. Wiegandt, Para, 154 pp., 21 pls. https: // doi. org / 10.5962 / bhl. title. 8538","Edwards, F. W. (1932 a) Genera Insectorum. Diptera, Fam. Culicidae. Fascicle 194. V. Verteneuil & L. Desmet, Bruxelles [Brussels], 258 pp., 5 pls.","Reinert, J. F., Harbach, R. E. & Kitching, I. J. (2008) Phylogeny and classification of Ochlerotatus and allied taxa (Diptera: Culicidae: Aedini) based on morphological data from all life stages. Zoological Journal of the Linnean Society, 153 (1), 29 - 114. https: // doi. org / 10.1111 / j. 1096 - 3642.2008.00382. x","Reinert, J. F., Harbach, R. E. & Kitching, I. J. (2009) Phylogeny and classification of tribe Aedini (Diptera: Culicidae). Zoological Journal of the Linnean Society, 157 (4), 700 - 794 + 2 online appendices. https: // doi. org / 10.1111 / j. 1096 - 3642.2009.00570. x","Coquillett, D. W. (1902 b) New Diptera from North America. Proceedings of the United States National Museum, 25 (1280), 83 - 126. https: // doi. org / 10.5479 / si. 00963801.25 - 1280.83","Carpintero, D. J. & Leguizamon, M. N. (2000) Una neuva especie de mosquito en la la [sic] Provincia de Buenos Aires Aedes (Ochlerotatus) jorgi. Nueva especies. Mariposas del Mundo, 5, 20 - 23.","Antunes, P. C. A. & Lane, J. (1938) Um novo Aedes, Aedes (Ochlerotatus) pennai, encontrado em Sao Paulo. Revista do Museu Paulista, 23, 605 - 614, 3 pls. [for 1937]","Edwards, F. W. (1922) Mosquito notes. - III. Bulletin of Entomological Research, 13 (1), 75 - 102. https: // doi. org / 10.1017 / S 0007485300045259","Belkin, J. N., Schick, R. X. & Heinemann, S. J. (1971) Mosquito studies (Diptera, Culicidae) XXV. Mosquitoes originally described from Brazil. Contributions of the American Entomological Institute, 7 (5), 1 - 64.","Walker, F. (1856) Insecta Saundersiana: or characters of undescribed insects in the collection of William Wilson Saunders, Esq., F. R. S., F. L. S. & c. Vol. I. Diptera. John van Voorst, London, 474 pp., 8 pls. https: // doi. org / 10.5962 / bhl. title. 5112","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.","Gutsevich, A. V. & Garcia-Avila, I. (1969) Nuevas especies de mosquitos (Culicidae) para Cuba. Torreia, 13, 3 - 8.","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.","Martini, E. (1935) Los mosquitos de Mexico. Departamento de Salubridad Publica, Boletines Tecnicos, Serie A: Entomologia Medica y Parasitologia No. 1. Departamento de Salubridad Publica, Mexico, D. F. 65 + 1 (content) pp.","Rodriguez-Martinez, L. M., Yzquierdo-Gomez, P., Gonzalez-Acosta, C. & Correa-Morales, F. (2020) Primer Registro de Aedes (Ochlerotatus) fulvus 1 en Tabasco y notas de distribucion de otros Aedes en Mexico. Southwestern Entomologist, 45 (1), 263 - 268. https: // doi. org / 10.3958 / 059.045.0127","Clark-Gil, S. & Darsie Jr., R. F. (1983) The mosquitoes of Guatemala, their identification, distribution and bionomics, with keys to adult females and larvae. Mosquito Systematics, 15 (3), 151 - 284."]}

Aedes (Ochlerotatus) spencerii (Theobald) subspecies idahoensis (Theobald, 1903a)—original combination: Grabhamia spencerii var. idahoensis (subspecific status by Nielsen & Rees 1959). Distribution: Canada, United States (continental) (Wilkerson et al. 2021). subspecies spencerii (Theobald, 1901c) —original combination: Culex spencerii. Distribution: Canada, Mexico, United States (continental) (Wilkerson et al. 2021). Aedes spencerii was described, originally as a species of Culex, from four adult females collected in Manitoba Province of Canada, two from Stony Mountain, the type locality based on lectotype selection by Belkin (1968), and two from St. Boniface (Theobald 1901c). Theobald provided a color illustration of the adult female in dorsal aspect (Theobald 1901a [plates]): fig. 104, pl. XXVI), which shows the pattern of dark tergal scaling typical of subspecies spencerii: “Abdomen covered with creamy and white scales, with large, black-scaled patches on each side of the middle line” (Theobald 1901c). Subspecies idahoensis was originally described as a variety of spencerii based on an unspecified number of females collected at Market Lake, Idaho, United States (Theobald 1903a); however, Townson (1990) reported the presence of two syntype females in the collection of the Natural History Museum, London. Theobald indicated that idahoensis only differed from the typical form in “abdominal ornamentation”, which he described as “Abdominal basal bands almost white, broad apical bands yellowish-white to white, very narrow, almost obliterated on some segments; the abdomen not pale at the sides, so that only broad basal and narrow apical pale areas are shown, the two apical segments are mostly pale scaled, and there are scattered pale scales on the others.” In contrast, idahoensis has traditionally been characterized as having dark-scaled abdominal terga with white transverse basal bands (see references listed below). Seven COI sequences for Ae. spencerii are registered in GenBank, three from Saskatchewan Province, Canada (KR444402, 596 bp; KR443924, 551 bp; and KR440309, 596 bp); two from Manitoba Province, Canada (KF535006, 658 bp and KF535007, 658 bp); one from Utah State, USA (JX260472, 213 bp); and one from Colorado, USA (JX259672, 658 bp). BLAST comparison of sequences revealed that two of the sequences from Saskatchewan (KR444402 and KR440309) and the sequence from Colorado (JX259672) are identical, and the 551 bp of the third sequence from Saskatchewan (KR443924) is 99.82% similar to the corresponding segments of those three sequences. Additionally, a 589-bp sequence from Saskatchewan identified as “ Aedes sp. BOLD-2016” is 98.13% similar. The sequence from Utah (JX260472, 213 bp) was too short to obtain a meaningful comparison, and the two sequences from Manitoba were apparently obtained from misidentified specimens: KF535006 is identical with 11 COI sequences of Ae. dorsalis and KF535007 is 99.39% similar to a COI sequence from Alberta (KP047935) identified as “ Culicidae sp. BOLD:AAD7982”. No DNA sequences are available for specimens identified as subspecies idahoensis. Ward (1984, 1992) and Wilkerson et al. (2021) were apparently not aware that Wood et al. (1979) had synonymized idahoensis with spencerii, with the following explanation: “The status of idahoensis has not been satisfactorily resolved. A few females in the Canadian National Collection have the characteristic abdominal pattern of idahoensis, but there are no males or larvae in the collection. According to Carpenter and LaCasse (1955), larvae of idahoensis have comb scales (13–29 instead of 7–13 in spencerii), and each scale has a less pronounced apical spine. Males of the two species were not separated. Nielsen and Rees (1959) found ‘almost every degree of variation’ between the typical female abdominal pattern of idahoensis and that of spencerii in Utah, Montana, and Wyoming. Furthermore, they cited three other authors who described similar variations in spencerii. They concluded that idahoensis was a more southerly subspecies of spencerii and that a zone of integration occurred in the three states previously mentioned. Until detailed biological or genetical studies resolve the status of idahoensis, we have no choice but to consider it a synonym of spencerii.” Darsie & Ward (1981, 2005) continued to recognize the two nominal forms as subspecies, stating in 1981: “Following Nielsen and Rees (338) [number of the article in the list of references cited] we recognize two subspecies under Ae. spencerii (Theobald), the typical subspecies which inhabits the central plains of North America, and subspecies idahoensis (Theobald), a more westerly and northwesterly form, also reported from southern British Columbia (135).” Based on published records, Darsie & Ward showed that the distributions of the two subspecies overlap in Montana, Nebraska, North Dakota, South Dakota and Wyoming. They indicated that only idahoensis was known to occur in Colorado; thus, the COI sequence in GenBank (JX259672) of the specimen from Colorado identified as spencerii sensu stricto suggests that either both subspecies occur in the state or idahoensis is conspecific with the typical form that occurs in Canada. More specimens and study are needed to resolve this question. It is interesting to note that idahoensis, originally described as a variety of spencerii, has been regarded as a distinct species (Dyar & Knab 1908; Dyar 1917b; Dyar 1928; Edwards 1932a; Gjullin 1937; Rees 1943; Yamaguti & LaCasse 1951 d; Carpenter & LaCasse 1955), a synonym (Pratt 1956; Wood et al. 1979) and a subspecies (Nielsen & Rees 1959; Smith 1966; Myers 1967; Darsie & Ward 1981, 2005). In as much as idahoensis is apparently morphologically distinct in the adult and larval stages (Carpenter & LaCasse 1955; Darsie & Ward 1981, 2005) in areas southward and westward of an area of overlap or a zone of introgression with spencerii in Montana, Nebraska, North Dakota, South Dakota and Wyoming, we believe it is prudent to recognize it as a separate species, pending a comprehensive morphological and genetic analysis of specimens from throughout the distributions of the two forms: Aedes (Ochlerotatus) idahoensis (Theobald, 1903a) . Aedes idahoensis 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 42-43, DOI: 10.11646/zootaxa.5303.1.1, http://zenodo.org/record/8043342, {"references":["Theobald, F. V. (1903 a) A monograph of the Culicidae or mosquitoes. Vol. 3. British Museum (Natural History), London, xvii + 1 (errata) + 359 pp., foldout table, 17 pls. https: // doi. org / 10.5962 / bhl. title. 58067","Nielsen, L. T. & Rees, D. M. (1959) The mosquitoes of Utah - a revised list. Mosquito News, 19 (2), 45 - 47.","Wilkerson, R. C., Linton, Y. - M. & Strickman, D. (2021) Mosquitoes of the world. Vols. 1 & 2. Johns Hopkins University Press, Baltimore, Maryland, 1332 pp. https: // doi. org / 10.1186 / s 13071 - 021 - 04848 - 6","Theobald, F. V. (1901 c) A monograph of the Culicidae or mosquitoes. Vol. 2. British Museum (Natural History), London, viii + 391 pp. https: // doi. org / 10.5962 / bhl. title. 58067","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","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.","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.","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.","Darsie Jr., R. F. & Ward, R. A. (1981) Identification and geographical distribution of the mosquitoes of North America, north of Mexico. Mosquito Systematics Supplement, 1, 1 - 313.","Darsie Jr., R. F. & Ward, R. A. (2005) Identification and geographical distribution of the mosquitoes of North America, north of Mexico. University Press of Florida, Gainesville, Florida, 383 pp.","Dyar, H. G. & Knab, F. (1908) Descriptions of some new mosquitoes from tropical America. Proceedings of the United States National Museum, 35 (1632), 53 - 70. https: // doi. org / 10.5479 / si. 00963801.35 - 1632.53","Dyar, H. G. (1917 b) Notes on the Aedes of Montana (Diptera, Culicidae). Insecutor Inscitiae Menstruus, 5 (7 - 9), 104 - 121.","Dyar, H. G. (1928) The mosquitoes of the Americas. Publication No. 387. Carnegie Institution of Washington, Washington, D. C., 616 pp.","Edwards, F. W. (1932 a) Genera Insectorum. Diptera, Fam. Culicidae. Fascicle 194. V. Verteneuil & L. Desmet, Bruxelles [Brussels], 258 pp., 5 pls.","Yamaguti, S. & LaCasse, W. J. (1951) Mosquito fauna of North America. Part II - Genera Megarhinus, Wyeomyia, Uranotaenia and Culiseta. Office of the Surgeon, HQ, APO Japan Logistical Command, 343, i + 1 - 85.","Pratt, H. D. (1956) A checklist of the mosquitoes (Culicinae) of North America (Diptera: Culicidae). Mosquito News, 16 (1), 4 - 10.","Smith, M. E. (1966) Mountain mosquitoes of the Gothic, Colorado, area. American Midland Naturalist, 76 (1), 125 - 150. https: // doi. org / 10.2307 / 2423238","Myers, C. M. (1967) Identification and description of Aedes eggs from California and Nevada (Diptera: Culicidae). Canadian Entomologist, 99 (8), 795 - 806. https: // doi. org / 10.4039 / Ent 99795 - 8"]}

Anopheles (Cellia) rhodesiensis Theobald subspecies rhodesiensis Theobald, 1901a —original combination: Anopheles rhodesiensis. Distribution: Angola, Benin, Botswana, Burkina Faso, Cameroon, Central African Republic, Chad, Côte d’Ivoire, Democratic Republic of the Congo, Djibouti, Eritrea, Eswatini, Ethiopia, Gabon, Ghana, Guinea, Kenya, Malawi, Mali, Mauritania, Morocco, Mozambique, Namibia, Niger, Nigeria, Republic of the Congo, Republic of South Africa, Saudi Arabia, Senegal, Seychelles, Sierra Leone, Somalia, South Sudan, Sudan, Tanzania, Uganda, Zambia, Zimbabwe (Wilkerson et al. 2021). subspecies rupicolus Lewis, 1937 —original combination: Anopheles (Myzomyia) rupicolus (subspecific status by Mattingly & Knight 1956). Distribution: Algeria, Chad, Egypt, Eritrea, Ethiopia, Jordan, Lebanon, Morocco, Niger, Saudi Arabia, Sudan, Syria, Yemen (Wilkerson et al. 2021). The nominotypical subspecies was described from Mashonaland, British Central Africa, now in northern Zimbabwe. Theobald (1901a) noted that the species had been sent “in numbers” from “Central Africa”. It has since been documented throughout the Afrotropical Region (Kyalo et al. 2017; Irish et al. 2020, who treated all records as rhodesiensis). Theobald provided a color illustration of the habitus of the adult female (Theobald 1901a [plates]: fig. 14, pl. IV), which matches the verbal description. Of note were: Head scales erect, dark laterally, pale on occiput and frontal tuft; scutum reddish brown with a dorsocentral ash-gray stripe; maxillary palpus with three narrow pale bands, one near apex of palpomere 2, one between palpomeres 3 and 4 and one encompassing most of palpomere 5; wing with distinct pale spots on the costa and vein R 1 as follows, sector pale (on R 1 only, illustrated by others on both veins), subcostal pale, preapical pale and apical pale spots (as defined by Wilkerson & Peyton 1990); legs entirely dark brown; abdomen brown with basal and median light brownish yellow mottling. These characters have been used in most keys to identify rhodesiensis, e.g. de Meillon (1947), Gillies & de Meillon (1968) and Gillies & Coetzee (1987). Characters of the adult, larva, pupa and male genitalia are well documented and illustrated in the three publications. According to Gillies & de Meillon (1968), larvae are found in a variety of habitats, including “rock pools, stream beds, seepages with little shade, margins of streams, springs and pools, ditches and hoof-prints, artificial containers such as concrete tanks and swimming-baths, whether in exposed water or in shade.” Females are not known to feed on humans or to be a factor in malaria transmission (Gillies & de Meillon 1968). Subspecies rupicolus was described from Gebel Moya near Sennar, Blue Nile Province, southeastern Sudan. Lewis (1937) remarked that the species had almost all dark wings resembling some species of the subgenus Anopheles. The color of the head scales was described as like those of subspecies rhodesiensis, with dark scales on the back of the occiput, white scales on the vertex and a conspicuous white vertical frontal tuft. The maxillary palpus and legs were described as all dark and the wing as having variably faint to absent sector pale, subcostal pale and preapical pale spots on the costa and/or vein R 1. Specimens were collected from a metal tank but most from a deep cleft in a rock. Lewis stated: “The wing markings and characters of the pharyngeal armature [cibarial armature] and male hypopygium [genitalia] indicate that the species is allied to A. rhodesiensis Theo. It differs from the latter in having 37 teeth on the mandible which is expanded at the tip, dark palpi with a higher index [length in relation to proboscis length], very faint pale marks, less than four, on the costa, no pale area on the basal third of the first longitudinal vein [R 1], and in the larva the clypeal hairs [setae 2,3-C] usually simple [single]. The mandible of A. rhodesiensis is not expanded at the end and has 80 teeth (Christophers [& Puri] 1931).” No apical pale spot was noted. De Meillon (1947) provided full descriptions of both subspecies (as species), and included illustrations showing contrasting wing spot characters and allopatric distributions (rupicolus Saharan, rhodesiensis Afrotropical), but then noted after the description of rupicolus: “It is questionable if this species is distinct from rhodesiensis. The adult is paler, the wings and palps [maxillary palpi] with less distinct pale making but otherwise there is little to separate them. The larvae and pupae, when considering the range of variation seen in rhodesiensis, appear to be inseparable from that species.” Mattingly & Knight (1956) cited de Meillon’s (1947) note and agreed that they too doubted the two were distinct species and provisionally preferred to treat them as subspecies. In addition, they mentioned that Lewis agreed with them. The logic and utility of this decision is not clear. However, the two nominal taxa have been treated as subspecies since that time (Gillies & de Meillon 1968; Gillies & Coetzee 1987; Knight & Stone 1977; Harbach 2018; Wilkerson et al. 2021). Gillies & de Meillon (1968) included the two nominal taxa in their key to adult females but they are combined in the larval key as rhodesiensis. They stated that rupicolus “is not separable from the nominate form.” An important indicator of separate species status is evidence of sympatry. Kyalo et al. (2017) did a comprehensive review of literature records of all Anopheles in the Afrotropical Region. Their dataset (Snow 2017) was used by David Pecor (Walter Reed Biosystematics Unit) to map rhodesiensis and rupicolus. This allowed visualization of numerous instances of co-occurrence and overlapping ranges of the two nominal taxa. Also, Verrone (1962), in a paper not indexed by Kyalo et al. (2017), recorded both subspecies in close proximity in Ethiopia. The accuracy of these records is enhanced by an illustrated key to adult females clearly showing the diagnostic characters discussed above. The above nominal taxa have been closely associated since the description of rupicolus. All stages are very similar, except for diagnostic differences in the adult female (wing with an apical pale spot, banded proboscis and many fewer mandibular teeth in rhodesiensis). The assumption was made that differences in adult characters were due to clinal variation related to hotter temperatures in the Saharan taxon rupicolus. However, we can find no published evidence of clinal variation, and it is apparent the two forms are sympatric. This obviates the assumption of subspecific status. We therefore believe they are two genetically distinct species and formally reinstate rupicolus to its original specific status: Anopheles (Cellia) rupicolus Lewis, 1937 . Anopheles rupicolus is currently listed as a species in the Encyclopedia of Life. Anopheles rupicolus currently has two synonyms, which we retain: An. aegypti Salem, 1938 (type locality: Wadi Taba, Sinai, Egypt) and An. rhodesiensis var. dthalisimilis Corradetti, 1939 (type locality: Semien District, Ethiopia). Both were treated as questionable synonyms by Edwards (1941), which leaves some doubt about their status and lays open the possible existence of a species complex., 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 70-71, DOI: 10.11646/zootaxa.5303.1.1, http://zenodo.org/record/8043342, {"references":["Theobald, F. V. (1901 a) A monograph of the Culicidae or mosquitoes. Vol. 1. British Museum (Natural History), London, xviii + 424 pp., pls. viii + I - XXXVII and A - E under separate cover. https: // doi. org / 10.5962 / bhl. title. 58067","Wilkerson, R. C., Linton, Y. - M. & Strickman, D. (2021) Mosquitoes of the world. Vols. 1 & 2. Johns Hopkins University Press, Baltimore, Maryland, 1332 pp. https: // doi. org / 10.1186 / s 13071 - 021 - 04848 - 6","Lewis, D. J. (1937) A new species of Anopheles from the Anglo-Egyptian Sudan. Proceedings of the Royal Entomological Society of London, Series B: Taxonomy, 6 (9), 181 - 183, 1 pl. https: // doi. org / 10.1111 / j. 1365 - 3113.1937. tb 01685. x","Mattingly, P. F. & Knight, K. L. (1956) The mosquitoes of Arabia I. Bulletin of the British Museum (Natural History) Entomology, 4 (3), 91 - 141.","Kyalo, D., Amratia, P., Mundia, C. W., Mbogo, C. M., Coetzee, M. & Snow, R. W. (2017) A geo-coded inventory of anophelines in the Afrotropical Region south of the Sahara: 1898 - 2016. Wellcome Open Researach, 2, 57. https: // doi. org / 10.12688 / wellcomeopenres. 12187.1","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","Wilkerson, R. C. & Peyton, E. L. (1990) Standardized nomenclature for the costal wing spots of the genus Anopheles and other spotted-wing mosquitoes (Diptera: Culicidae). Journal of Medical Entomology, 27 (2), 207 - 224. https: // doi. org / 10.1093 / jmedent / 27.2.207","de Meillon, B. (1947) The Anophelini of the Ethiopian geographical region. Publications of the South African Institute for Medical Research, 10 (49), 1 - 272.","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.","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","Snow, R. W. (2017) A geo-coded inventory of anophelines in the Afrotropical Region south of the Sahara: 1898 - 2016. Harvard DataVerse, V 1. https: // doi. org / 10.7910 / DVN / NQ 6 CUN","Verrone, G. A. (1962) Outline for the determination of malarial mosquitoes in Ethiopia. Part I - Adult female anophelines. Mosquito News, 22 (1), 37 - 49.","Salem, H. H. (1938) The mosquito fauna of Sinai Peninsula (Egypt) with a description of two new species. Publications of the Egyptian University Faculty of Medicine, 16, 1 - 31, 16 pls.","Corradetti, A. (1939) Studi morfologici sulla specie anofelica precedentemente identificata come A. dthali nel semien e sua classificazione come nuova varieta di A. rhodesiensis. Rivista di Parassitologia, 3 (1), 57 - 63.","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."]}

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

Anopheles (Anopheles) pseudopunctipennis Theobald subspecies levicastilloi Levi Castillo, 1944 —original combination: Anopheles pseudopunctipennis var. levicastilloi (subspecific status by Leví Castillo 1945). Distribution: Ecuador (Levi Castillo 1944). subspecies neghmei Mann, 1950 —original combination: Anopheles pseudopunctipennis ssp. neghmei. Distribution: Chile (Mann 1950). subspecies noei Mann, 1950 —original combination: Anopheles pseudopunctipennis ssp. noei. Distribution: Chile (Mann 1950). subspecies patersoni Alvarado & Heredia, 1947 —original combination: Anopheles (Anopheles) pseudopunctipennis var. patersoni (subspecific status by Lane 1953). Distribution: Argentina (Alvarado & Heredia 1947). subspecies pseudopunctipennis Theobald, 1901c —original combination: Anopheles pseudopunctipennis. Distribution: Aruba, Curaçao, Granada, Margarita Island (Venezuela), Trinidad and Tobago (historical), Venezuela [to be verified] (revised here). subspecies rivadeneirai Leví Castillo, 1945 —original combination: Anopheles (Anopheles) pseudopunctipennis var. rivadeneirai (subspecific status by Lane 1953). Distribution: Ecuador (Leví Castillo 1945). syn. argentinus Brèthes, 1912 —original combination: Proterorhynchus argentinus. syn. peruvianus Tamayo, 1907 (in Tamayo & García 1907)—original combination: Anopheles peruvianus. syn. tucumanus Lahille, 1912 —original combination: Anopheles tucumanus. Overview. Anopheles pseudopunctipennis sensu lato (herein the Pseudopunctipennis Complex)has a wide distribution from the southern United States to northern Argentina and Chile. In some localities it is an important malaria vector, in others it is seldom attracted to humans (Rozeboom 1941). This complex belongs to the Pseudopunctipennis Group, which also includes An. chiriquiensis Komp, 1936, An. eiseni Coquillett, 1902a, An. franciscanus McCracken, 1904, An. geometricus Corrêa, 1944, An. hectoris Giaquinto Mira, 1931 and An. parapunctipennis Martini, 1932 (Reid & Knight 1961). The Pseudopunctipennis Complex (as An. pseudopunctipennis) has been diagnosed throughout its range in keys and descriptions using the following characters. Female: Maxillary palpus with narrow pale bands at articulations, palpomere 5 pale; scutum with broad silvery gray central area; legs mostly dark and unmarked; costa of wing dark except for subcostal pale and apical pale spots (according to Wilkerson & Peyton 1990), vein R 1 pale except at presector dark, sector dark and subapical dark spots, sector pale spot present at radial crossvein (r 1 -r s), vein Cu pale with postbasal and preapical dark spots, vein 1A pale basally, dark apically except for pale apex, and all posterior veins terminating at pale fringe spots. Male: Aedeagus with 1–3 pairs of small curved serrate leaflets. Larva: Seta 3-C single; 9-M,T about 2 times length of seta 10-M,T; 6-IV,V prominent, plumose; 2-IV single; median plate of spiracular apparatus with a pair of distinctive posterior-directed projections. Egg: Characters vary but an egg with the collar separate from the floats is the character state most cited and illustrated. Currently, not including the nominotypical subspecies, the complex includes five subspecies and three synonyms. We use literature-based morphological, ecological, geographic, cytological and genetic evidence to propose elevation of four subspecies and two synonyms to specific status, and synonymy of one subspecies.We transfer one elevated synonym to nomen dubium because of the lack of a primary type, or any other specimens for study, and a contradictory original description. Two junior synonyms remain associated with one of the elevated synonyms. No subspecies are retained., 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 59, 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.","Mann F., G. (1950) Dos nuevas sub-especies del Anopheles pseudopunctipennis Th. 1901. Biologica, 8 - 11, 33 - 42.","Alvarado, C. A. & Heredia, R. L. (1947) Observaciones sobre una nueva variedad del Anopheles (A.) pseudopunctipennis Theobald 1901, encontrada en la provincia de Tucuman. (Nota previa). Anales del Instituto de Medicina Regional Universidad Tucuman, 2 (1), 73 - 78.","Lane, J. (1953) Neotropical Culicidae. Vols. I & II. University of Sao Paulo, Sao Paulo, 1112 pp.","Theobald, F. V. (1901 c) A monograph of the Culicidae or mosquitoes. Vol. 2. British Museum (Natural History), London, viii + 391 pp. https: // doi. org / 10.5962 / bhl. title. 58067","Brethes, J. (1912) Los mosquitos de la Republica Argentina. Boletin de Instituto Entomologia Patagonia Vegetal, 1, 1 - 48.","Tamayo, M. O. & Garcia, C. A. (1907) Las aquas do Huacachina. Informe prosentado a la Sociedad Geografica de Lima. Memoirs Municipal de Lima, 1906, 1 - 63. [Reproduced in part 30 September 1907 as \" Las Aquas do Huacachina \" in Boletin de la Sociedad Geografica de Lima, 21, 241 - 251, including the description of Anopheles peruvianus Tamayo]","Lahille, F. (1912) Sobre un Anopheles, una Stegomyia y la notacion de las nervaduras alares de los mosquitos. Anales del Museo Nacional de Historia Natural de Buenos Aires, 23, 253 - 263, 5 pls.","Rozeboom, L. E. (1941) Distribution and ecology of the Anopheles mosquitoes of the Caribbean Region. Publication of the American Association for the Advancement of Science, 15, 98 - 107.","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.","Coquillett, D. W. (1902 a) New forms of Culicidae from North America. Journal of the New York Entomological Society, 10 (3), 191 - 194.","McCracken, I. (1904) Anopheles in California, with description of a new species. Entomological News, 15 (1), 9 - 14, 1 pl.","Correa, R. R. (1944) Desericao de uma nova subespecie do subgenero \" Anopheles \" (Diptera, Culicidae). Anals Paulistas de Medicina e Cirurgia, 47 (5), 407 - 408.","Giaquinto Mira, M. (1931) Una nueva especie de Anopheles en Guatemala: Anopheles hectoris mihi. Boletin de la Direccion General de Salubridad, Guatemala, 1 (20 / 25), 606 - 615.","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","Wilkerson, R. C. & Peyton, E. L. (1990) Standardized nomenclature for the costal wing spots of the genus Anopheles and other spotted-wing mosquitoes (Diptera: Culicidae). Journal of Medical Entomology, 27 (2), 207 - 224. https: // doi. org / 10.1093 / jmedent / 27.2.207"]}

Tripteroides (Tripteroides) powelli (Ludlow) subspecies escodae Baisas & Ubaldo-Pagayon, 1953 —original combination: Tripteroides (Tripteroides) powelli escodae. Distribution: Philippines (Baisas & Ubaldo-Pagayon 1953). subspecies laffooni Baisas & Ubaldo-Pagayon, 1953 —original combination: Tripteroides (Tripteroides) powelli laffooni. Distribution: Philippines (Baisas & Ubaldo-Pagayon 1953). subspecies mattinglyi Baisas & Ubaldo-Pagayon, 1953 —original combination: Tripteroides (Tripteroides) powelli mattinglyi. Distribution: Philippines (Baisas & Ubaldo-Pagayon 1953). subspecies powelli (Ludlow, 1909b) —original combination: Uranotaenia powelli. Distribution: Cambodia, Indonesia, Malaysia, People’s Republic of China, Philippines, Thailand, Vietnam (Wilkerson et al. 2021). The genus Tripteroides, with species distributed throughout the Australasian and Oriental Regions, has never been the subject of a comprehensive taxonomic revision; consequently, the majority of its species are poorly known. This is particularly true of species of the subgenus Tripteroides, for which the immature stages of only a “small minority” of species have been described in detail and those that are known are currently indistinguishable (Mattingly 1981). Tripteroides powelli sensu stricto and the subspecies escodae, laffooni and mattinglyi were all described from different islands of the Philippines, and only the nominotypical form has been recorded in other countries of the Oriental Region (see above). The four forms have been found on various Philippine islands: escodae —Balabac, Jolo, Mindanao and Palawan (type locality) Islands; laffooni — Leyte (type locality), Mindanao and Samar Islands; mattinglyi —Culion (type locality) and Palawan Islands; powelli sensu stricto —widely distributed from Luzon (type locality) to Mindanao (Baisas & Ubaldo-Pagayon 1953; Basio 1971). Thus, escodae, laffooni and powelli sensu stricto are found on Mindanao and escodae and mattinglyi both occur on Palawan. The immature stages are known for the four nominal forms, but they have not been studied or described in comparative detail. In fact, there is some doubt concerning the identity of the larva of powelli sensu stricto, as revealed by Baisas & Ubaldo-Pagayon (1953). The type specimen described by Ludlow, being a female with unknown larva, is not distinguishable from four kinds of powelli now known in the Philippines. Even were its larva known, it could not be separated from the others unless it was the one which had two-branched ventral siphon tufts [seta 1-S]. The other three are alike in larval characters. Brug’s description of powelli larva (1939), based on Dutch East Indian specimens, does not definitely prove that is the kind of larva Ludlow’s type powelli has. By coincidence, however, the materials we have on hand from Luzon Island (type locality) represent only the kind of larvae which has two-branched ventral siphon tufts, the same as that descried by Brug, but at the same time it differs in certain details from the Dutch East Indian form. It is not at all unlikely other forms of powelli may also be found in Luzon Island if more extensive collections are made, in which case it is only by assumption that the type specimen could be regarded as that whose larva has two-branched ventral siphon tufts. Assuming the specimens reared from larvae that were studied by Baisas & Ubaldo-Pagayon are conspecific with the type female of powelli, it seems the larvae, as well as the pupae, of the four forms recognized by the authors, based albeit on apparent cursory examination of specimens, are indistinguishable. As stated by Baisas & Ubaldo-Pagayon, “Definite differences between [the] subspecies [are] found only in [the] male terminalia [genitalia].” The anatomical terminology used by Baisas & Ubaldo-Pagayon (1953) and most other authors for structures of the phallosome of the male genitalia of Tripteroides is confusing. References to the phallosome generally apply to features of the aedeagus, which consists of two lateral plates (aedeagal sclerites) that are normally connected in tergal aspect by a narrow submedian tergal bridge, each plate with a variously developed apical sternal process (ventral arm, VA, of Baisas & Ubaldo-Pagayon) and a less distinctively developed median tergal process (dorsal arm, DA, of Baisas & Ubaldo-Pagayon). Considering this clarification, Baisas & Ubaldo-Pagayon distinguished the subspecies of powelli as follows. Phallosome of mattinglyi is quite distinct from those of others, not only in powelli complex but also from all species in Group B, because its VA is represented only by teeth which arise directly from rim of tube at point where, in other species of Group B, VA arises. Moreover, mattinglyi has a few teeth, of which three or four arise from a small prominence at base (tergal wall) of DA. Laffooni has finer and more (over twelve) teeth at this point, but they arise not from a prominence but directly from wall of DA. Mattinglyi is strikingly similar in phallosome to rozeboomi of Group A. Beyond that, however, there is no other similarity between them. VA in powelli powelli much shorter than that of escodae or mattinglyi. Viewed in whole terminalia [genitalia] mounts VA in powelli powelli may be seen beneath and at about or below middle of DA. This is because VA is only about half length of DA. VA in either escodae or laffooni measures equal or nearly equal length of DA. Because of long VA, and partly because of usual slight upward tilting of organ, the teeth are well exposed beyond apex of DA (see text fig. 2d) in whole mounts; and teeth coarser. Presence of teeth toward base of DA distinguishes laffooni from escodae; latter does not have any trace of such teeth. In many species-rich groups of animals, structures of the male genitalia provide better means for distinguishing species than any other morphological characters. Thus, evolution of genital form is thought to be involved in the origin of species. This would seem to be true of species of Tripteroides, which like many species of the genus Culex, and others, are difficult to distinguish except by features of the male genitalia. Based on the distinct differences in the structure of the aedeagal sclerites of the male genitalia and the apparent sympatry on different islands, as well as significant doubts about the identification of specimens of powelli sensu stricto in countries other than the Philippines, we are confident that the four nominal forms, escodae, laffooni, mattinglyi and powelli, are separate species, which further morphological study, coupled with molecular analyses, will corroborate. Based on this conviction, the three subspecies described by Baisas & Ubaldo-Pagayon are hereby formally raised to specific status: Tripteroides (Tripteroides) escodae Baisas & Ubaldo-Pagayon, 1953 ; Tp. (Trp.) laffooni Baisas & Ubaldo-Pagayon, 1953 ; and Tp. (Trp.) mattinglyi Baisas & Ubaldo-Pagayon, 1953 . Tripteroides escodae and Tp. mattinglyi are currently listed as species in the Encyclopedia of Life; Tp. laffooni should be added to the list., 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 130-131, DOI: 10.11646/zootaxa.5303.1.1, http://zenodo.org/record/8043342, {"references":["Baisas, F. E. & Ubaldo-Pagayon, A. (1953) Notes on Philippine mosquitoes, XVI. Genus Tripteroides. Bureau of Printing, Manila, 198 pp., 23 pls. [for 1952]","Ludlow, C. S. (1909 b) Mosquito observations. Canadian Entomologist, 41 (7), 233 - 235. https: // doi. org / 10.4039 / Ent 41233 - 7","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","Mattingly, P. F. (1981) Medical entomology studies - XIV. The subgenera Rachionotomyia, Tricholeptomyia and Tripteroides (Mabinii Group) of genus Tripteroides in the Oriental region (Diptera: Culicidae). Contributions of the American Entomological Institute, 17 (5), + 1 - 147.","Basio, R. G. (1971) The mosquito fauna of the Philippines (Diptera Culicidae). Monograph No. 4. National Museum of the Philippines, Manila, v + 198 pp."]}

Toxorhynchites (Afrorhynchus) viridibasis (Edwards) subspecies viridibasis (Edwards, 1935) —original combination: Megarhinus aeneus var. viridibasis (specific status by Edwards 1941). Distribution: Burkina Faso, Central African Republic, Côte d’Ivoire, Mali, Nigeria, South Sudan, Sudan, Uganda (Wilkerson et al. 2021); also the Democratic Republic of the Congo (see below). subspecies voltaicus Ribeiro, 2005 —original combination: Toxorhynchites (Afrorhynchus) viridibasis voltaicus. Distribution: Burkina Faso (Ribeiro 2005). Edwards (1935) described viridibasis (as a variety of Megarhinus aeneus Evans, 1926) from two adult females collected in Uganda, the type specimen from Kampala and the second female from Soroti. The description was very brief and did not include illustrations. In addition to a slightly more detailed description of the two females, Edwards (1941) provided a very brief description of a dubious adult male, stating: “Although the male is damaged I think it must belong to the same species as the females: it is clearly quite different from the male of erythrurus, but it is quite possible that both viridibasis and aeneus should be regarded as subspecies of lutescens; the available material is too scanty for any conclusion to be formed of the point.” Ribeiro (2005) based the description of subspecies voltaicus on a single adult male captured on 18.7.54 at Banankélédage (Banankeledaga), cercle (circle, area) Bobo-Dioulasso, Haute-Volta (now Burkina Faso)— Banankeledaga is a village located near the city of Bobo-Dioulasso, the capital of Houet Province. Ribeiro did not mention how he identified the specimen as being conspecific with the female of the nominate form, but he distinguished the two subspecies in a key for the identification of males of species of the subgenus Afrorhynchus Ribeiro, 1992. Oddly, the two subspecies keyed out in different couplets, with the nominate subspecies keying out in a couplet with Tx. ruwenzori (van Someren, 1948) and subspecies voltaicus keying out in a couple in which the alternate character state leads to two consecutive couplets that identify Tx. capelai Ribeiro, 1992b, Tx. lutescens (Theobald, 1901a) and Tx. zairensis Ribeiro, 2005. Scaling on the mesokatepisternum was used to distinguish subspecies viridibasis from subspecies voltaicus and the other three species: In the nominate subspecies, “Golden scales on mesokatepisternum restricted to the lower portion of the sclerite [sternum]”; in subspecies voltaicus, “A patch of golden scales present on the upper portion of the mesokatepisternum, at base of prealar knob”. According to Edwards (1941), the female of the nominate form has “much of the pre-alar area (except the knob) clothed with white scales.” It is uncertain whether the prealar area of the male also has white scaling as Edwards only mentioned that the male “resembles ♀ in colouring of body and legs (thorax much denuded).” In the diagnosis of voltaicus, Ribeiro mentioned that the male also differed from the male of the type form in having “a purple third tergum” and the coloration of the maxillary palpus, which is “extensively golden” in the nominate form and “mainly purplish golden with violet reflections” in voltaicus. He noted that “No significant differences were found between the male genitalia of Tx. v. voltaicus and those of the male Tx. v. viridibasis from Coquilhatville, Congo ” [Coquilhatville is the former name of present-day Mbandaka, a city on the Congo River in the Democratic Republic of the Congo]. For clarity, Edwards (1941) stated that the maxillary palpus of the questionable male of the nominate form that he examined was “as in lutescens ”, which he described as having the “shaft and penultimate segment golden beneath, only tip of latter dark, and with some golden scales above, terminal segment all dark”. In the original description of viridibasis, Edwards (1935) characterized the female as having the “First three abdominal segments almost entirely green-scaled, contrasting with the remainder, which are purple.” In 1941, he more precisely described the abdominal terga: “Abdomen with first three tergites [terga I–III] clothed with metallic green scales, a slight admixture of purple scales on the third, rest purple.” Although this pertains to females, it suggests that the third tergum may not always be entirely green-scaled and could prove to be predominately or entirely purple on inspection of a much larger number of specimens; perhaps more so in males. The larva and pupa of Tx. viridibasis were described by Lewis (1945, southern Sudan) and Wolfs (1947, Democratic Republic of the Congo), and minor details of the larva were provided by Hamon (1954, Burkina Faso). In each case, larvae were reared to adults, which were used to identify the species. Wolfs apparently described the larva from exuviae associated with two individually reared males. It is noteworthy that the holotype male of voltaicus designated by Ribeiro (2005) was collected and identified as Tx. viridibasis by J. Hamon, who recorded the following: “we obtained an ex larva [from a larval rearing] specimen which allows us to add the following indications to the descriptions of Lewis and Wolfs: head seta B [seta 6-C] has 5 branches; the subventral bristle of the siphon [seta 1-S] is bifid on one side and simple [single] on the other [translated from the French].” For comparison, Wolfs (1947) indicted that setae 6-C and 1-S are both 4-branched in larvae from the Democratic Republic of the Congo, whereas Lewis (1945) indicated that specimens from Sudan have seta 6-C with 2 or 3 branches “near the tip” and seta 1-S single, but “may have 1 or 2 branches near its tip.” It was not obvious until now that the holotype male of voltaicus is one of the two males which Hamon (1954) reared from larvae collected in an area that “forms approximately a circle with a radius of 50 kilometers and is entirely included in the administrative subdivision of Bobo Dioulasso [translated from the French].” Therefore, one of the larval exuviae examined by Hamon (1954) is the larval exuviae of the holotype of voltaicus. Based on the scant morphological data listed above, it would seem that subspecies voltaicus differs from the nominate form principally in features of the adults—abdominal tergum III entirely with purple scales (entirely green or with a few intermixed purple scales in the type form), upper mesokatepisternal scale-patch comprised of golden scales (these scales white in the type form), maxillary palpus of the male mainly clothed in purplish golden scales with violet reflections (mainly clothed in golden scales in the type form). Because the interpretation of color is subjective and dependent on lighting, and may be variable within a species, we consider such differences to be of questionable value in recognizing species-group taxa. With regard to the larva, the paucity of information shows that the larva of voltaicus shares the single or bifid seta 1-S with the type form in Sudan, and both differ from the form in the Democratic Republic of the Congo in which this seta is 4-branched. There seems to be a greater difference in the branching of seta 6-C, which has two or three apical branches in Sudanese larvae and is four- and five-branched in larvae from the Democratic Republic of the Congo and the type locality of subspecies voltaicus, respectively. It is impossible to know whether these differences are attributable to variation across the range of a single species or an indication of a potential species complex. In the absence of firm evidence, we feel it is prudent at this time to formally consider voltaicus as nothing more than a local morphological variant of the nominotypical form: voltaicus Ribeiro, 2005, junior subjective synonym of Toxorhynchites (Afrorhynchus) viridibasis (Edwards, 1935) . Consequently, voltaicus should be removed from the species of Toxorhynchites listed 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 116-117, DOI: 10.11646/zootaxa.5303.1.1, http://zenodo.org/record/8043342, {"references":["Edwards, F. W. (1935) Mosquito notes. - XII. Bulletin of Entomological Research, 26 (2), 127 - 136. https: // doi. org / 10.1017 / S 0007485300038116","Edwards, F. W. (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","Ribeiro, H. (2005) Three new species and one new subspecies of Toxorhynchites (Diptera: Culicidae) of the Afrotropical Region. Journal of Medical Entomology, 42 (3), 224 - 232. https: // doi. org / 10.1093 / jmedent / 42.3.224","Evans, A. M. (1926) Notes on Freetown mosquitos, with descriptions of new and little-known species. Annals of Tropical Medicine and Parasitology, 20 (1), 97 - 108, 1 pl. https: // doi. org / 10.1080 / 00034983.1926.11684481","van Someren, E. C. C. (1948) Ethiopian Culicidae: Some new mosquitoes from Uganda. Proceedings of the Royal Entomological Society of London, Series B: Taxonomy, 17 (9 - 10), 128 - 132. https: // doi. org / 10.1111 / j. 1365 - 3113.1948. tb 00902. x","Ribeiro, H. (1992 b) On the terrestrial and riverine fauna of the Democratic Republic of Sao Tome and Principe. VII - Description of Toxorhynchites capelai sp. nov. (Diptera, Culicidae). Garcia de Orta, Serie de Zoologia, 19 (1 - 2), 135 - 128.","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","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","Wolfs, J. (1947) Description des larves et des nymphes de quelques megarhines. M. phytophagus Theobald, M. viridibasis Edwards, et M. aeneus Theobald [sic, Evans]. Recueil de Travaux de Sciences Medicales au Congo Belge, 6, 63 - 72."]}

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

Subspecies nipponii Subspecies nipponii was described from a single “perfect ♀ ” from Karnizana, Japan [Karuizawa, Nagano Prefecture]. Theobald (1907) wrote: “proboscis with scattered golden scales, with black apex, palpi mottled.” “Abdomen with the basal segment testaceous, with a median patch of creamy scales and rather short pale golden hairs; the second and third segments with basal uniform creamy bands; on the fourth, fifth and sixth the bands spread out laterally, forming marked lateral areas; the sixth and seventh have yellow apical scales, and the second to the fifth have median pale scales which form a broken median line; venter pale scaled.” A distinguishing character is the median pale abdominal scales. Reinert (1973) did a comprehensive study of vexans vexans and vexans nipponii as part of a study of the subgenus Aedimorphus in Southeast Asia. He treated vexans vexans as a single variable worldwide species. He characterized subspecies nipponii in the adult female as follows: “Abdomen with terga III, IV each with an incomplete apical median longitudinal white stripe which may or may not connect with basal bands; lower subspiracular scale patch connects, or nearly so, with postspiracular scale patch.” In the pupa, the two were very similar but could be separated from other related species by having the “Ventral posterior margin of II [tergum II] with spicules mesally.” In the larval stage, he contrasted vexans vexans in Southeast Asia with subspecies nipponii by the presence of a granulose frontoclypeus in the former not present in the latter. This granularity is rarely seen in the rest of the range, sensu Reinert (1973), of the putative subspecies vexans. To us, it could also indicate a separate species in Southeast Asia. Tanaka et al. (1979) stated that only vexans nipponii occurs in Japan, Korea and the eastern Palearctic, and thus they did not provide a complete comparison of it and vexans vexans. As described by Theobald (1907) and Reinert (1973), Tanaka distinguished nipponii by noting that “Subspecies nipponii is chiefly characterized by the presence of pale median patches on the anterior abdominal terga of the adults.” The larva of subspecies nipponii is like the larva of typical vexans but setae 5–7-C are distinctly aciculate, as are setae 1,3,5-VIII. Because of its Palearctic distribution and distinct morphological characters, we believe subspecies nipponii should be afforded species status: Aedes (Aedimorphus) nipponii (Theobald, 1907) . Synonyms: None. Aedes nipponii needs to be added in the list of Aedes species recognized 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 17-18, DOI: 10.11646/zootaxa.5303.1.1, http://zenodo.org/record/8043342, {"references":["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","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.","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."]}

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

Trichoprosopon compressum Lutz subspecies compressum Lutz, 1905 —original combination: Trichoprosopon compressum. Distribution: Argentina, Bolivia, Brazil, Colombia, Ecuador, French Guiana, Guyana, Panama, Paraguay, Venezuela (Wilkerson et al. 2021). subspecies mogilasium (Dyar & Knab, 1907) —original combination: Joblotia mogilasia (varietal status by Stone 1944; subspecific status by Harbach & Howard 2007). Distribution: Brazil, Ecuador, French Guiana, Panama, Venezuela (Wilkerson et al. 2021). Trichoprosopon is a very poorly known genus.As currently defined, it only includes 14 formerly recognized species (Wilkerson et al. 2021; Rivera-García et al. 2023), but there are an unknown number of undescribed species (T. J. Zavortink and the late E. L. Peyton, pers. comm., circa 1992). Based on this and what little is known about the morphology of the immature stages of compressum sensu stricto and mogilasium, it is difficult to know for certain whether they are the same or separate species. The pupa of compressum was fully illustrated by Knight & Chamberlain (1948) and the pupa and larva were briefly described and partially illustrated by Lane (1953). The immature stages of mogilasium have not been described. The nominate subspecies was described from an “unspecified number of ♂ and ♀ from unspecified localities in the states of São Paulo and Rio de Janeiro, Brazil ” (Belkin 1968). Lane (1953) listed the type locality as “ Brasil, State of S. Paulo, Pindamonhanga”. The locality, however, has been misinterpreted to be equivalent to São Paulo (Stone et al. 1959; Belkin 1968; Knight & Stone 1977; Wilkerson et al. 2021). The type locality is Pindamonhangaba, a municipality in the state of São Paulo located in the Paraíba Valley approximately 100 km northeast of the city of Sao Paulo. Subspecies mogilasium and Joblotia trichorryes Dyar & Knab, 1907, a synonym of compressum (see below), were both described from specimens collected at Tabernilla in the Canal Zone of Panama. The three nominal forms, compressum, mogilasium and trichorryes, have an interesting history. Joblotia mogilasia and J. trichorryes were described as separate species on the same page (Dyar & Knab 1907), with the latter preceding the former. We believe the descriptions support their recognition as separate species, especially the presence of setae and scales on the clypeus of mogilasia and the presence of only setae on the clypeus of trichorryes. The two forms were listed as separate species in Panama by Dyar (1923), but five years later Dyar (1928) treated them as synonyms of J. compressa without explanation. This was in turn followed by Edwards (1932a), who listed them as synonyms of Tr. compressum. Lane (1936a), based on a single female collected on human bait at Nhandeara, a municipality in São Paulo State of Brazil, revalidated mogilasia as a species of Joblotia, based on “the fact that in our specimen the clypeus is laterally covered with dark scales and some setae that are barely perceptible” (translated from the Portuguese). Lane further noted that “If there really are no scales in the mogilasia type then our specimen is atypical or a new species.” Strangely, Lane & Cerqueira (1942) reasoned that because the presence of setae and scales on the clypeus “is the only character that separates the adults of T. mogilasium from those of T. digitatum and as the existing material is represented only by some females, we believe that it is preferable to consider T. mogilasium as a variety of T. digitatum until other phases of biology are known and the systematic position definitively established” (translated from the Portuguese). Stone (1944) noted that Lane & Cerqueira had discovered that scales may be present on the clypeus of both Tr. digitatum (Rondani, 1848) and Tr. compressum, and they had incorrectly treated mogilasium as a variety of digitatum. Stone therefore established mogilasium as a variety of compressum, stating that it “agrees with typical compressum in all diagnostic characters save that there are some scales intermingled with the hairs [setae] on the sides of the clypeus.” The taxonomic status of mogilasium was confused until it was clarified by Harbach & Howard (2007) as follows: “ Knight & Stone (1977) indicated that Joblotia mogilasia was formally recognized as a subspecies of Tr. compressum by Stone (1944) and later afforded varietal status by Stone et al. (1959). This is incorrect as Stone (1944) unambiguously treated this nominal species as a variety of Tr. compressum. Because mogilasium was adopted (originally) as the valid name of a species prior to 1985, it is deemed to be subspecific with availability from its original publication ([ICZN] Article 45.6.4.1).” In summary, setae and scales are present on the clypeus of mogilasium whereas only setae are found on the clypeus of compressum and trichorryes. In comparison with the detailed morphological descriptions of adult mosquitoes published post-Belkin (1962), the available descriptions of compressum, mogilasium and trichorryes reveal that specimens were not studied in detail and were only superficially examined. Lane (1953) provided the most recent treatment of species currently included in the genus, and for the most part his descriptions are very superficial and unsatisfactory. The three nominal forms considered here are based entirely on adult characters, principally general coloration of scaling and the length of various appendages. The adults may actually show many good characters, but these need to be studied in much greater detail than has been done by the authors cited above. The larvae and pupae of sabethine mosquitoes are known to exhibit striking anatomical differences (Zavortink 1979), but these life stages, as indicated above, are either unknown or have not been studied in sufficient detail to evaluate their usefulness in recognizing and distinguishing similar species. In the case of the nominal forms of concern here, while it appears on superficial examination that only one variable species is involved, we suspect it is likely that several distinct sympatric forms may be present. This is supported by the recognition of a Compressum Complex by Zavortink (1981) that includes two “closely related [undescribed] allopatric taxa [which] occur in bamboo internodes and are apparently indistinguishable. However, in characteristics of the adults and male genitalia, these taxa differ to a much greater extent than do the sympatric species of the Tr. digitatum complex [see below], and so I consider these allopatric taxa to be distinct species.” Zavortink did not mention subspecies mogilasium. Based on the original descriptions, recorded distributions and the distinct possibility that morphological and genetic distinctions await discovery, and because it is important to determine with certainty whether or not the nominal species of our predecessors are distinct biological species, we believe it is befitting to return mogilasium and trichorryes to their original species status: Trichoprosopon mogilasium (Dyar & Knab, 1907) and Trichoprosopon trichorryes (Dyar & Knab, 1907) , elevated from synonymy with Tr. compressum Lutz, 1905. Trichoprosopon mogilasium is currently listed as a species in the Encyclopedia of Life; Tr. trichorryes needs to be added to the list., 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 127-128, DOI: 10.11646/zootaxa.5303.1.1, http://zenodo.org/record/8043342, {"references":["Lutz, A. (1905) Novas especies de mosquitos do Brasil. Imprensa Medica, 13 (2, 3, 4, 5, 6, 7, 9, 11, 14, 15, 16, 18), 26 - 29, 48 - 52, 65 - 70, 81 - 84, 101 - 104, 125 - 128, 169 - 172, 202 - 204, 269 - 271, 287 - 290, 311 - 314, 347 - 350.","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","Dyar, H. G. & Knab, F. (1907) Descriptions of new mosquitoes from the Panama Canal Zone. Journal of the New York Entomological Society, 15 (4), 197 - 212.","Stone, A. (1944) Notes on the genus Trichoprosopon (Diptera, Culicidae). Revista de Entomologia, 15 (3), 335 - 341.","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","Rivera-Garcia, K. D., Mendez-Andrade, A. & Ibanez-Bernal, S. (2023) Trichoprosopon mixtli sp. nov., a new sabethine species (Diptera: Culicidae) from a Mexican cloud forest, with an assessment of the genus and keys for the identification of known species. Zootaxa, 5254 (1), 94 - 116. https: // doi. org / 10.11646 / zootaxa. 5254.1.5","Knight, K. L. & Chamberlain, R. W. (1948) A new nomenclature for the chaetotaxy of the mosquito pupa, based on a comparative study of the genera (Diptera: Culicidae). Proceedings of the Helminthological Society of Washington, 15 (1), 1 - 18.","Lane, J. (1953) Neotropical Culicidae. Vols. I & II. University of Sao Paulo, Sao Paulo, 1112 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.","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.","Dyar, H. G. (1923) The mosquitoes of Panama (Diptera, Culicidae). Insecutor Inscitiae Menstruus, 11 (10 - 12), 167 - 187.","Dyar, H. G. (1928) The mosquitoes of the Americas. Publication No. 387. Carnegie Institution of Washington, Washington, D. C., 616 pp.","Edwards, F. W. (1932 a) Genera Insectorum. Diptera, Fam. Culicidae. Fascicle 194. V. Verteneuil & L. Desmet, Bruxelles [Brussels], 258 pp., 5 pls.","Lane, J. (1936 a) II. Notas sobre mosquitos de Sao Paulo. Boletim de Instituto Hygiene de Sao Paulo, 60, 3 - 15, 5 pls.","Lane, J. & Cerqueira, N. L. (1942) Os sabetineos da America (Diptera, Culicidae). Arquivos de Zoologia do Estudo de Sao Paulo, 3 (9), 473 - 849.","Rondani, C. (1848) Esme di varie specie d'insetti ditteri brasiliani. Studi entomologici, Turino, 1, 63 - 112, 3 pls.","Zavortink, T. J. (1979) Mosquito studies (Diptera, Culicidae) XXXV. The new sabethine genus Johnbelkinia and a preliminary reclassification of the composite genus Trichoprosopon. Contributions of the American Entomological Institute, 17 (1), 1 - 61.","Zavortink, T. J. (1981) Species complexes in the genus Trichoprosopon. Mosquito Systematics, 13 (1), 82 - 85."]}

Toxorhynchites (Lynchiella) rutilus (Coquillett) subspecies rutilus (Coquillett, 1896) —original combination: Megarhinus rutila [sic]. Distribution: Extreme southeastern United States— Florida, Georgia (mainly coastal), North Carolina (Harrison et al. 2016), South Carolina (coastal) (Jenkins 1949; Darsie & Ward 2005). subspecies septentrionalis (Dyar & Knab, 1906a)—original combination: Megarhinus septentrionalis (subspecific status by Jenkins & Carpenter 1946). Distribution: Eastern North America, northward from northern Florida to Canada (southwestern Ontario) and westward to Kansas, Oklahoma and Texas (Jenkins 1949; Wood et al. 1979; Darsie & Ward 2005). Coquillett (1896) indicated that Tx. rutilus, as Megarhinus rutila, was described and named based on three males and five females from North Carolina and Georgiana, Florida. According to Stone & Knight (1957c), the original collection included two specimens from Georgiana, Florida and seven from undisclosed localities in Florida, but none from North Carolina. They selected a male from Georgiana bearing a type label to be the lectotype. Georgiana is located on Merritt Island of Brevard County on the eastern side of central Florida. Dyar & Knab (1906a) described and named Tx. septentrionalis, also as a species of Megarhinus Robineau-Desvoidy, 1827, from a collection of 13 males and 11 females from localities in the District of Columbia, Louisiana, Maryland, Mississippi, Missouri, North Carolina, Virginia and West Virginia. Stone & Knight (1957c) considered a male from Woodstock, Virginia bearing a type label to be the holotype. The subspecific status of septentrionalis was established by Jenkins & Carpenter (1946), and reinforced by Jenkins (1949) based on conclusions drawn from a study that included specimens form a narrow zone of overlap and apparent integration with the type form. Jenkins described the evidence for integration as follows. In the zone of overlap of ranges of the two subspecies, all types of integrations occur. In the northern part of the zone in Georgia and South Carolina, the male intergrades have segments [tarsomeres] 2 and 3 of the fore tarsi with white areas or with isolated white scales surrounded by purple. Two typical intergrades from Myrtle Beach, South Carolina, Carpenter and Jenkins (1945) [i.e. 1946], have the following markings on the fore tarsi: Specimen 1.-2nd segment [foretarsomere 2] with basal three-fourths silvery white; 3rd segment [foretarsomere 3] with three white scales. Specimen 2.- 2nd segment, (left) with six white scales, (right) with 4 white scales; 3rd segment, (left) with 3 white scales, (right) entirely purple. In the southern part of the zone of overlap in Florida, the fore tarsi of the intergrade males are usually typical T. r. rutilus with the second and basal two-thirds of the third segment silvery white, but interspersed with several dark purple scales. The intergrades were observed from Jacksonville, Tallahassee, and Bushnell, Florida. The amount of white on the fifth segment [tarsomere 5] of the hind tarsi of the males is variable. There is a decided tendency for this segment to be mostly white in Florida T. r. rutilus specimens, and mostly dark purple in the main part of the range of T. r. septentrionalis. This is quite variable and a specimen of the subspecies from Falls Church, Virginia has this segment almost entirely white. The type of intergradation described for the fore tarsi and often observed on the fifth hind tarsal segment [hindtarsomere 5]. An unusual amount of white was observed on the fore tarsi of males from Duval Co., Leon Co., and Orange Co., Florida that had the 2nd, 3rd and about half of the 4th segments [foretarsomeres 2, 3 and 4] silvery white. A few specimens exhibited white scales on the apical tip of the tibiae. Jenkins summarized his findings and previous observations (Jenkins & Carpenter 1946) in the form of conclusions: The tropical genus Toxorhynchites is represented in the United States by two closely related forms which are considered to be subspecies for the following reasons. 1. The distributions of the two forms overlap in part of their ranges. 2. A complete series of intergrades between the two forms occurs throughout the zone of overlap in ranges. 3. The only known character separating the two forms is a color difference of the males which is subject to variation. 4. No significant differences have been found to separate the two forms in the male genitalia, larvae, pupae, or females. 5. The life histories, habitats, and breeding habits of the two forms have been found to be similar. It is important to note that conclusions 3 and 4 have been repeated, apparently without further detailed morphological study, by later authors (Carpenter & LaCasse 1955; Wood et al. 1979; Harrison et al. 2016). Conclusion 5 is meaningless as it applies to species of Toxorhynchites in general. It is surprising to find that the subspecific status of septentrionalis has been accepted by taxonomists without question since the studies of Jenkins & Carpenter (1946) and Jenkins (1949), and has not since been re-examined using genetic and molecular tools. There are four records of DNA sequences in GenBank: 28S rRNA, MT-CYB and MT-ND5, Colony, Florida Medical Entomology Laboratory, recorded as Tx. rutilus (Krzywinski et al. 2001 a); mt - COI (partial), mt -COII and tRNA -Leu, York County, Pennsylvania, recorded as Tx. rutilus (Mitchell et al. 2002); mt -COI, Ontario, Canada, recorded as Tx. rutilus (Cywinska et al. 2006); and 18S rDNA, Stonington, Connecticut, recorded as Tx. r. septentrionalis (Shepard et al. 2006). Unfortunately, because the researchers did not produce the same molecular sequences, it is not possible to conduct a meaningful analysis of relationships. That aside, the identity of the records attributable to Tx. rutilus are problematic. The records from Canada and Pennsylvania must certainly be based on specimens of septentrionalis, and it is not possible to know whether the Florida colony was derived from specimens of rutilus sensu stricto or intergrades. It should be noted, however, that researchers who conducted ecological studies of treehole mosquitoes in eastern North America (Bradshaw & Holzapfel 1975, 1977; Chambers 1985) did not recognize septentrionalis. As there is no indication of intermediates in the geographically separated populations, there is little doubt that specific traits are being maintained outside the zone of introgression. This is evidence for independent species cohesion, and we believe that molecular systematic studies will reveal that the two forms are separate species that hybridize in a narrow zone where the two species overlap. This supports the taxonomic conclusion that the type form and septentrionalis are distinct species; therefore, we hereby formally return septentrionalis to its original rank as a species: Toxorhynchites (Lynchiella) septentrionalis (Dyar & Knab, 1906a) , which is currently listed as a species in the Encyclopedia of Life. Toxorhynchites septentrionalis has a single synonym, Megarhinus herrickii Theobald, 1906 (type locality: Mississippi State, USA), synonymy by Howard et al. (1917). As noted by Belkin (1968), Theobald based the description of herrickii on Herrick’s (1905) description of Megarrhinus portoricensis, apparently without having seen any specimens—probably the ultimate reason for naming the species after Herrick. Megarhinus herrickii remains a junior subjective synonym of Tx. septentrionalis., 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 120-121, DOI: 10.11646/zootaxa.5303.1.1, http://zenodo.org/record/8043342, {"references":["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.","Jenkins, D. W. (1949) Toxorhynchites mosquitoes of the United States. Proceedings of the Entomological Society of Washington, 51 (5), 225 - 229.","Darsie Jr., R. F. & Ward, R. A. (2005) Identification and geographical distribution of the mosquitoes of North America, north of Mexico. University Press of Florida, Gainesville, Florida, 383 pp.","Dyar, H. G. & Knab, F. (1906 a) The species of mosquitoes in the genus Megarhinus. Smithsonian Miscellaneous Collections, 48 (3), 241 - 258.","Jenkins, D. W. & Carpenter, S. J. (1946) Ecology of the tree hole breeding mosquitoes of nearctic North America. Ecological Monographs, 16 (1), 31 - 47. https: // doi. org / 10.2307 / 1943573","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.","Stone, A. & Knight, K. L. (1957 c) Type specimens of mosquitoes in the United States National Museum: VI, Miscellaneous genera, addenda, and summary. Journal of the Washington Academy of Sciences, 47 (6), 196 - 202.","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.","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.","Krzywinski, J., Wilkerson, R. C. & Besansky, N. (2001) Evolution of mitochondrial and ribosomal gene sequences in Anophelinae (Diptera: Culicidae): implications for phylogeny reconstruction. Molecular Phylogenetics and Evolution, 18 (3), 479 - 487. https: // doi. org / 10.1006 / mpev. 2000.0894","Mitchell, A., Sperling, F. A. H. & Hickey, D. A. (2002) Higher-level phylogeny of mosquitoes (Diptera: Culicidae): mtDNA data support a derived placement for Toxorhynchites. Insect Systematics and Evolution, 33 (2), 163 - 174. https: // doi. org / 10.1163 / 187631202 X 00118","Cywinska, A., Hunter, F. F. & Hebert. P. D. N. (2006) Identifying Canadian mosquito species through DNA barcodes. Medical and Veterinary Entomology, 20 (4), 413 - 424. https: // doi. org / 10.1111 / j. 1365 - 2915.2006.00653. x","Shepard, J. J., Andreadis, T. G. & Vossbrinck, C. R. (2006) Molecular phylogeny and evolutionary relationships among mosquitoes (Diptera: Culicidae) from the northeastern United States based on small subunit ribosomal DNA (18 S rDNA) sequences. Journal of Medical Entomology, 43 (3), 443 - 454. https: // doi. org / 10.1093 / jmedent / 43.3.443","Bradshaw, W. E. & Holzapfel, C. M. (1975) Biology of tree-hole mosquitoes: photoperiodic control of development in northern Toxorhynchites rutilus (Coq.). Canadian Journal of Zoology, 53 (7), 889 - 893. https: // doi. org / 10.1139 / z 75 - 102","Chambers, R. C. (1985) Competition and predation among larvae of three species of treehole-breeding mosquitoes. In: Lounibos, L. P., Rey, J. R. & Frank, J. H. (Eds.), Ecology of mosquitoes: proceedings of a workshop. Florida Mededical Entomology Laboratory, University of Florida, Vero Beach, Florida, pp. 25 - 53.","Theobald, F. V. (1906) A new Megarhinus. Entomologist, 39 (522), 241.","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.","Herrick, G. W. (1905) Notes on some Mississippi mosquitoes. Entomological News and Proceedings of the Entomological Section of the Academy of Natural Sciences of Philadelphia, 16 (9), 281 - 283, 1 pl."]}

Toxorhynchites (Toxorhynchites) brevipalpis Theobald subspecies abyssinicus Ribeiro, 1991 —original combination: Toxorhynchites (Toxorhynchites) brevipalpis abyssinicus. Distribution: Ethiopia (Ribeiro 1991). subspecies brevipalpis Theobald, 1901a —original combination: Toxorhynchites brevipalpis. Distribution: Angola, Benin, Burkina Faso, Cameroon, Central African Republic, Côte d’Ivoire, Democratic Republic of the Congo, Ghana, Kenya, Liberia, Malawi, Mozambique, Nigeria, Senegal, South Africa, Tanzania, Uganda, Zambia, Zimbabwe (Wilkerson et al. 2021). subspecies conradti Gr ̧nberg, 1907—original combination: Toxorhynchites conradti (subspecific status by Hopkins 1936). Distribution: Burkina Faso, Cameroon, Côte d’Ivoire, Democratic Republic of the Congo, Equatorial Guinea, Gabon, Ghana, Gambia, Liberia, Mali, Nigeria, Republic of the Congo, Sierra Leone, South Sudan, Sudan, Uganda (Wilkerson et al. 2021). Theobald (1901a) described brevipalpis from two females collected in “Natal”, now known as KwaZulu-Natal, a coastal province of South Africa. Steffan & White (1981) noted that the two specimens were originally “in Walker’s collection”, but “Since only one of the ♀♀ mentioned by Theobald is now in the BM Collection [Natural History Museum, London], it is designated as the lectotype.” This is the specimen Edwards (1941) indicated was from Durban, which is considered to be the type locality even though the name of the city does not appear on the labels that accompany the specimen (Steffan & White 1981). Edwards apparently gleaned the locality from other information associated with Walker’s collection. Subspecies conradti was originally described as a distinct species by Gŗnberg (1907) based on a female and a male captured on different dates at the Johan-Albrecht-Ĥhe station on the crater rim of Lake Barombi near Kumba in the Southwest Region of Cameroon. The species was listed as a synonym of brevipalpis, without explanation, by Edwards (1912b, 1932a) and remained so until Hopkins (1936) described the larva of “ brevipalpis ” (as a species of Megarhinus) and inadvertently referred to conradti as a variety: “The description and figures are of var. conradti Gr ̧nberg.” The figures that Hopkins ascribed to conradti were reproduced from Macfie & Ingram (1923), who described and illustrated the larva of “ Megarhinus (Toxorhynchites) brevipalpis ” from specimens collected at localities in present-day Ghana: Nsawam and Sekondi (the twin city of Sekondi-Takoradi). They prefaced their description as follows: “A figure showing the paddles and the last three abdominal segments of the pupa of this mosquito has been published, together with a few words of explanation, by Bacot (Yellow Fever Commission, West Africa, Reports, iii, p. 145); and a larva, presumed to be that of M. brevipalpis, has been briefly described by Edwards (Bull. Ent. Res. iii, p. 375). Neither of these authors, however, has given sufficient details to distinguish the species, and indeed it seems not unlikely that the characters mentioned by them are mainly generic. As we have in our possession specimens of both larvae and pupae of M. brevipalpis, we have taken the opportunity of examining them in detail.” The specimens were apparently identified to species, in the absence of associated adults, with the help of G. A. K. Marshall, Imperial Bureau of Entomology, and F. W. Edwards, British Museum. The Toxorhynchites of Africa were so very poorly known at the time that the identification of the specimens as conradti by Macfie & Ingram (1923) was probably presumptive, based on its known distribution in West Africa. Incidentally, we note that Hopkins (1936) is wrongly credited with the recognition of conradti as a subspecies in all subsequently published mosquito catalogs (Stone et al. 1959; Knight & Stone 1977; Wilkerson et al. 2021) when in fact the author clearly referred to it as a variety of brevipalpis. It appears that conradti was first formally recognized as a subspecies by Edwards (1941). Removal of the records listed for the presence of subspecies conradti in Benin and Senegal, which were not corroborated in the study of Ribeiro (1991), and discounting the separate contradictory records for the presence of conradti and the type form in the Central African Republic, Kenya and Nigeria, it would appear that the two forms are largely sympatric in the tropical rain forest region of central Africa, mainly in the Democratic Republic of the Congo. It must be borne in mind that all country records for both forms are based solely on differences in the amount of white scaling in the caudolateral tufts of abdominal tergum VI of females (nearly or completely white in the type form; all black or with a few anterior white setae in conradti) and the presence or absence of basal white scaling on foretarsomere 2 of males and females (present in the type form; absent in conradti). It must also be noted that Ribeiro (1991) found “intermediate forms” (sex not mentioned) in Angola, Burkina Faso, Nigeria and Uganda with black and white scale-tufts on abdominal tergum VI, which he considered to be brevipalpis x conradti hybrids. Because the two forms are otherwise distinct throughout their ranges, we believe the “intermediate forms” may be morphological variants of one or the other form or, perhaps more likely, a currently unrecognized species. In the same paper, Ribeiro (1991) described subspecies abyssinicus based on a single male collected in the former Keffa (or Kaffa) Province of Ethiopia, which is now a Zone in the South West Region of the country. Ribeiro distinguished abyssinicus from brevipalpis sensu stricto by the absence of white scales on foretarsomere 2 (also entirely dark-scaled in conradti), from conradti by the white-scaled tufts of abdominal tergum VI (as in the type form) and from both brevipalpis sensu stricto and conradti by the presence of fewer (8 and 10) setae on the ninth tergal lobes of the male genitalia (15–20 setae on each lobe in the other forms). Hopkins (1936), after revealing he used the illustrations of the larva of conradti published by Macfie & Ingram (1923) to represent the larva of brevipalpis, stated that “The larva of the typical form has not been distinguished, but is known to be very similar and probably indistinguishable.” He did not indicate the source of this knowledge. Hopkins (1952) repeated verbatim his 1936 treatment of brevipalpis, except for the sentences from the last paragraph quoted above, which he changed to read “The descriptions and figures are of ssp. conradti Gr ̧nberg, but the larva of the typical form is not distinguishable (E. C. C. van Someren, 1946b).” This is a delusive statement because van Someren (1946b) actually stated that the larva of the type form is “Indistinguishable from that of M. brevipalpis subsp. conradti as described by Macfie and Ingram (1922 [1923]) and Hopkins (1936).” Obviously this chain of presuppositions obscures the fact that the larva of brevipalpis sensu stricto is essentially unknown and has not been the subject of detailed comparative morphological study. If it is not obvious from the analyses of the previous three species of Toxorhynchites, it should be noted that species of Toxorhynchites are generally very poorly known taxonomically, extremely similar in all life stages and troublesome to identify. Most species have been distinguished based on pale scaling of the tarsi, which is often different in males and females of the same species, and the color of scales on areas of the thorax and abdomen, which may be variable or dependent on lighting and interpretation. Additionally, as pointed out by Steffan & Evenhuis (1985), “Species that are easily distinguishable in one stage may be entirely indistinguishable, or separable by only a single character, in another stage. …Because of the subtleties involved in distinguishing among the different species of Toxorhynchites in various stages, accurate identification necessitates well-preserved and prepared specimens of both sexes in each stage. Adults with associated pupal and larval skins [exuviae]… provide the most useful taxonomic information.” Unfortunately, individually reared adults with associated larval and pupal exuviae are lacking for Tx. brevipalpis and its currently recognized subspecies. Nevertheless, we believe that the available morphological and distributional information is sufficient to formally recognize the three nominal forms as separate species: The nominotypical species and Toxorhynchites (Toxorhynchites) conradti Grünberg, 1907 , which have largely allopatric distributions that overlap in central Africa, without demonstrable evidence of introgression, and Toxorhynchites (Toxorhynchites) abyssinicus Ribeiro, 1991 , which is diagnosed by a unique combination of characters and appears to be isolated from populations of Tx. conradti. Toxorhynchites conradti and Tx. abyssinicus are both currently listed as species in the Encyclopedia of Life. Toxorhynchites brevipalpis has a single synonym, Tx. marshallii Theobald, 1903a, described from an adult male from Salisbury, Mashonaland, a region in northern Zimbabwe which today is divided into four provinces. The type locality of marshallii is within the allopatric range of Tx. brevipalpis, and should be retained as a synonym of that species. Toxorhynchites conradti has two synonyms, Tx. schultzei Enderlein, 1931 (type locality: “Franẑsisch Äquatorial-Afrika, Mongumba am Ubangi-Fluβ”—Mongoumba is a town on the Ugangi River in the Central African Republic), and Tx. tessmanni Enderlein, 1931 (type locality: “ Spanisch-Guinea. Uam-Gebiet: Alen Benito”—Alen Benito [?Benito River; Alen is a mountain], Uam area [Rio Muni mainland], Equatorial Africa). These two nominal forms are each based on a single adult female. Their type localities are located in the allopatric range of conradti, and they should be retained as synonyms of that 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 121-123, DOI: 10.11646/zootaxa.5303.1.1, http://zenodo.org/record/8043342, {"references":["Ribeiro, H. 1991. Research on the mosquito subfamily Toxorhynchitinae (Diptera: Culicidae). I - The Afrotropical group brevipalpis (Adults) (1). Arquivos do Museu Bocage, 2 (3), 31 - 62.","Theobald, F. V. (1901 a) A monograph of the Culicidae or mosquitoes. Vol. 1. British Museum (Natural History), London, xviii + 424 pp., pls. viii + I - XXXVII and A - E under separate cover. https: // doi. org / 10.5962 / bhl. title. 58067","Wilkerson, R. C., Linton, Y. - M. & Strickman, D. (2021) Mosquitoes of the world. Vols. 1 & 2. Johns Hopkins University Press, Baltimore, Maryland, 1332 pp. https: // doi. org / 10.1186 / s 13071 - 021 - 04848 - 6","Hopkins, G. H. E. (1936) Mosquitoes of the Ethiopian Region I. - Larval bionomics of mosquitoes and taxonomy of culicine larvae. Printed by order of the Trustees, British Museum (Natural History), London, (vi) + 250 pp.","Steffan, W. A. & White, G. B. (1981) The type specimens of Toxorhynchites (Diptera: Culicidae) at the British Museum (Natural History) and Oxford University. Mosquito Systematics, 13 (1), 114 - 123.","Edwards, F. W. (1941) Mosquitoes of the Ethiopian Region III. - Culicine adults and pupae. Printed by order of the Trustees, British Museum (Natural History), London, viii + 499 pp.","Edwards, F. W. (1912 b) A synopsis of the species of African Culicidae, other than Anopheles. Bulletin of Entomological Research, 3 (1), 1 - 53. https: // doi. org / 10.1017 / S 0007485300001656","Edwards, F. W. (1932 a) Genera Insectorum. Diptera, Fam. Culicidae. Fascicle 194. V. Verteneuil & L. Desmet, Bruxelles [Brussels], 258 pp., 5 pls.","Macfie, J. W. S. & Ingram, A. (1923) The early stages of West African mosquitos. - VI. Bulletin of Entomological Research, 13 (4), 409 - 442. https: // doi. org / 10.1017 / S 000748530004551 X","Stone, A., Knight, K. L. & Starcke, H. (1959) A synoptic catalog of the mosquitoes of the world (Diptera, Culicidae). The Thomas Say Foundation. Vol. VI. Entomological Society of America, College Park, Maryland, vi + 358 pp.","Knight, K. L. & Stone, A. (1977) A catalog of the mosquitoes of the world (Diptera: Culicidae). 2 nd Edition. The Thomas Say Foundation. Vol. VI. Entomological Society of America, College Park, Maryland, xi + 611 pp.","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.","van Someren, E. C. C. (1946 b) Ethiopian Culicidae. Tribe Megarhinini - notes and descriptions. Transactions of the Royal Entomological Society of London, 97 (7), 177 - 186. https: // doi. org / 10.1111 / j. 1365 - 2311.1946. tb 00373. x","Macfie, J. W. S. & Ingram, A. (1922) On the genital armature of the female mosquito. Annals of Tropical Medicine and Parasitology, 16 (2), 157 - 188. https: // doi. org / 10.1080 / 00034983.1922.11684308","Steffan, W. A. & Evenhuis, N. L. (1985) Classification of the subgenus Toxorhynchites (Diptera: Culicidae) I. Australasian, eastern Palaearctic, and Oriental species-groups. Journal of Medical Entomology, 22 (4), 421 - 446. https: // doi. org / 10.1093 / jmedent / 22.4.421","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","Enderlein, G. (1931) Zwei neue Toxohynchites aus dem Kongogebiete. Zoologischer Anzeiger, 94 (3 / 4), 123 - 125."]}

Anopheles (Anopheles) bancroftii Giles subspecies bancroftii Giles, 1902 —original combination: Anopheles bancroftii. Distribution: Australia, Papua New Guinea, including the Admiralty Islands in the Bismarck Archipelago (Lee et al. 1987). subspecies barbiventris Brug, 1938 —original combination: Anopheles bancrofti [sic] var. barbiventris (subspecific status by Harbach & Howard 2007). Distribution: Indonesia, Sulawesi (Lee et al. 1987). Giles (1902) described Anopheles bancroftii, a widely distributed species in Australia (Lee et al. 1987), based on specimens from Burpengary, Queensland. Misidentification of An. pseudobarbirostris Ludlow, 1902 as An. bancroftii has resulted in erroneous listings of bancroftii in the Philippines, Sulawesi (Celebes) and Sri Lanka (Lee et al. 1987). Subspecies barbiventris was described from specimens collected in Kalawara, Palou, Gumbasa, Sigi Regency, Central Sulawesi, Indonesia (Gazetteer: GoeNames; Kalawara; populated place; coordinates: -1.1808, 119.9385; D. Pecor pers. comm.). In the description, it was compared to An. bancrofti [sic] var. pseudobarbirostris (now a separate species), An. tessellatus Theobald, 1901a and An. vagus D̂nitz, 1902. Foote & Cook (1959) listed barbiventris as being from “Australasia”, which includes many countries, but it’s only known occurrence is in Sulawesi (Celebes). Harbach & Howard (2007) recognized barbiventris as a subspecies of An. bancroftii per Article 45.6.4 of the International Code of Zoological Nomenclature and suggested that it was probably a distinct species. Reid (1962), commenting on the bancroftii group, noted discrepancies in existing descriptions: “ A. bancroftii var. barbiventris from Celebes seems to merit specific rank; in addition to the striking tufts of black and white scales on the abdominal sternites [sterna], the large leaflets on the phallosome [aedeagus] appear to be longer than those of bancroftii and pseudobarbirostris and have fine teeth [serration]; doubtless there are other differences.” Herein, in addition to evidence of allopatry, we agree with Reid (1962) and Harbach & Howard (2007) that barbiventris should be afforded species status: Anopheles (Anopheles) barbiventris Brug, 1938 . Anopheles barbiventris is currently listed as a species in the Encyclopedia of Life. Note. A species complex is suggested by a molecular and morphological study (Beebe et al. 2002; Beebe et al. 2013) demonstrating the probability of four bancroftii -related sibling species in northern Australia and Papua New Guinea., 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 55, DOI: 10.11646/zootaxa.5303.1.1, http://zenodo.org/record/8043342, {"references":["Giles, G. M. (1902) A handbook of the gnats or mosquitoes giving the anatomy and life history of the Culicidae together with descriptions of all species noticed up to the present date. 2 nd Edition. John Bale, Sons & Danielsson, Ltd., London, xi + (1) + 530 pp., 17 pls. https: // doi. org / 10.5962 / bhl. title. 8549","Brug, S. L. (1938) Waarnemingen bij anophelinen. Geneeskundig Tijdschrift voor Nederlandsch-Indie, 78, 520 - 523.","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","Ludlow, C. S. (1902) Two Philippine mosquitoes. Journal of the New York Entomological Society, 10 (3), 127 - 131.","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","Foote, R. H. & Cook, D. R. (1959) Mosquitoes of medical importance. Agricultural Handbook, U. S. Department of Agriculture, 152, 1 - 158.","Reid, J. A. (1962) The Anopheles barbirostris group (Diptera, Culicidae). Bulletin of Entomological Research, 53 (1), 1 - 57. https: // doi. org / 10.1017 / S 0007485300047945","Beebe, N. W. & Cooper, R. D. (2002) Distribution and evolution of the Anopheles punctulatus group (Diptera: Culicidae) in Australia and Papua New Guinea. International Journal for Parasitology, 32 (5), 563 - 574. https: // doi. org / 10.1016 / S 0020 - 7519 (01) 00359 - 9","Beebe, N. W., Russell, T. L., Burkot, T. R., Lobo, N. F. & Cooper, R. D. (2013) The systematics and bionomics of malaria vectors in the Southwest Pacific. In: Manguin, S. (Ed.), Anopheles mosquitoes: New insights into malaria vectors. InTech, Rijeka, pp. 357 - 394. https: // doi. org / 10.5772 / 55999"]}

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

Anopheles (Cellia) sergentii (Theobald) subspecies macmahoni Evans, 1936 —original combination: Anopheles (Myzomyia) macmahoni (subspecific status by Mattingly & Knight 1956). Distribution: Algeria, Burkina Faso, Côte d’Ivoire, Djibouti, Eritrea, Ethiopia, Ghana, Guinea, Kenya, Mali, Republic of South Africa, Senegal, Somalia, South Sudan, Sudan (Wilkerson et al. 2021). subspecies sergentii (Theobald, 1907) —original combination: Pyretophorus sergentii. Distribution: Albania, Algeria, Bulgaria, Burkina Faso, Cameroon, Chad, Côte d’Ivoire, Djibouti, Egypt, Eritrea, Greece, Iran, Iraq, Israel, Italy, Jordan, Libya, Kenya, Morocco, Pakistan, Portugal, Saudi Arabia, Senegal, Spain, South Sudan, Sudan, Tunisia, Turkmenistan (Wilkerson et al. 2021). The female of the nominotypical form was described by Theobald (1907) based on two adult females from an unspecified locality in Algeria (see below). He described them as a gray to brown species with three palpal bands, one of them the entire palpomere 5; with the abdomen brown without scales; legs brown, unbanded, and “Wings dark scaled; costal border with five large dark spots, four spreading evenly on to first long vein [R]; a few pale spots on wing field, notably at the cross-veins and bases of the fork-cells [R 2 and M 1].” All primary wing veins except 1A end at a pale fringe spot. There was no explicit mention of the nature of the dark and pale areas on vein R 4 +5 but “most of the veins” were described as “uniformly dark scaled”. Subsequently, Christophers (1933) described all life stages from specimens collected in presesnt-day Pakistan. Subspecies macmahoni was described as a species by Evans (1936) based on two larval cotypes and a male and a female with associated larval and pupal exuviae from Isiolo, Kenya. She noted: “The larvae of this species can be identified easily by the characters of the tergal plates…”, which are unusually broad. In the adult female, the maxillary palpus has three narrow pale bands, the tarsi lack definite pale rings and the wings are predominantly dark with the third vein (R 4+5) mainly or entirely dark-scaled. Based on additional specimens from the same locality, Evans stated that the hindtibia has a “rather distinct white spot at the base, as well as the usual apical spot.” Two years later, in her treatise on Afrotropical anophelines, Evans (1938) more precisely described variation observed for vein R 4+5: “Third vein ranging from entirely dark to about ⅔ pale, usually about ¼–½ pale; pale area when present usually situated distally as in A. funestus, but in 3 cases it was median in the vein.” Subspecies sergentii was not mentioned in Evans (1936) or Evans (1938). De Meillon (1947) followed the descriptions of Evans (1936, 1938) and additionally noted the presence of five parabasal setae on the gonocoxite of the male, the innermost borne on a prominent eminence, and vein R 4+5 all dark and the pale area, when present, at the center or with a long basal dark spot and a small apical one. Subspecies sergentii was also not mentioned in this publication. Mattingly & Knight (1956) in “Mosquitoes of Arabia” examined larvae of subspecies macmahoni from the Western Aden Protectorate (Yemen) “and consider them to be those of An. sergenti [sic] of which An. macmahoni is, in our view, a subspecies probably mainly confined to the African portion of the Somali-Arid District. The latter is therefore eliminated from our list.” Differences they noted in the cibarial armature of females, also noted by others, were deemed to be too variable to be indicative of separate species. However, two principal differentiating characters were noted. Vein R 4+ 5 in macmahoni “is normally extensively pale scaled in the middle and only exceptionally dark.” Subspecies sergentii, in contrast, “is constantly dark except at base and extreme tip”. The other feature used to separate the two nominal taxa was the large size of the main tergal plates present in larvae of macmahoni. Senevet et al. (1959) conducted a comprehensive study of the larval tergal plates of sergentii and macmahoni in Algeria. Whereas Theobald (1907) merely indicated that the type specimens of sergentii were collected in Algeria by Dr E. Sergent, Senevet et al. revealed that the specimens were found by Edmond Sergent in “El Outaya (région de Biskra, partie N. du Sahara algérien).” Based on this revelation, we hereby restrict the type locality of Anopheles sergentii Theobald, 1907 to El Outaya (35.040868, 5.56491), Biskra Province, Algeria. Senevet et al. noted that one of them had collected sergentii on the south side of the Aurès Mountains at the northern limit of the Sahara. For comparison, they obtained specimens from Tassili N’Ajjers, a national park in the Sahara located north of Djanet in southeastern Algeria. El Outaya and Djanet are about 1,000 km apart. The authors summarized their findings as follow: “we are dealing with two types: the group ́small plates» corresponds to the variety ́typicus » [sergentii]; the ́large plates» group corresponds to the subspecies macmahoni. The first of these groups does not in fact differ much from the form considered as typical by most authors [translated from the French].” The two nominal taxa belong to the demeilloni section of Gillies & de Meillon (1968) (= Demeilloni Group in Wilkerson et al. 2021). Gillies & de Meillon placed adult females of subspecies macmahoni in their key in Section XI, with the following characters: “Mosquitoes with 1 pale spot on upper branch of 5th vein [M 3+4], no pale interruption on 3rd dark area [preapical dark spot] of 1st vein [R 1]; costa with at least 1 pale spot on basal half; palps [maxillary palpi] with less than 4 bands, pale at apex; legs not speckled, hind tarsus 4 and 5 [hindtarsomeres 4 and 5] not entirely pale; abdomen without laterally projecting tufts of scales.” We point out that both nominal taxa have dark tarsomeres, maxillary palpus with three pale bands, two narrow ones, plus the entire palpomere 5, basal area of vein R 1 entirely pale-scaled, all veins except 1A ending in a pale fringe spot. Gillies & de Meillon placed fourth-instar larvae of both subspecies in their key in Section VI: “Larvae with wide abdominal plates [tergal plates], equalling on segment 5 three quarters or more the distance between bases of palmate hairs [setae]; saddle hair [seta 1-X] with less than 5 branches; no coarse spicules on sides of thorax and abdomen; outer clypeal hairs [seta 3-C] with less than 8 branches; inner clypeal hairs [seta 2-C] wide apart and not strongly branched.” Both nominal taxa have one of the long mesopleural “hairs” [seta 9- or 10-M] plumose and the inner shoulder “hair” [seta 1-P] with 15–20 branches inserted on a well-developed tubercle. In subspecies sergentii, the width of the main tergal plate on segment V is at most four-fifths the distance between the palmate setae, whereas in subspecies macmahoni the main tergal plate on segment V is equal to greater than four-fifths the distance between the palmate setae. Gillies & de Meillon (1968) summarized their view of the two taxa as follows. Anopheles sergentii sergentii: “A widespread and important vector of malaria, extending into southern Arabia. As pointed out by Mattingly and Knight (1956) it differs from subsp. macmahoni only in the larval stage. From this it is separable by the width of the main tergal plates, which are at most ⅘ the distance between the palmate hairs [setae].” “LARVAL HABITAT: According to Senevet and Andarelli (1956), sergentii occurs in oases and irrigated areas in many types of water, shaded and unshaded, with and without vegetation.” “ADULT BIOLOGY: Attacks man readily, often entering houses to do so. Often found resting indoors by day, but also makes extensive use of outdoor shelters which are sometimes remote from dwellings. It was formerly, at least, an important vector of malaria...”. Gillies & de Meillon illustrated the distribution of the two subspecies in map form (fig. 53), on which they drew a non-overlapping line of demarcation. They further commented that “We ourselves would note in passing that, despite the close similarity between sergentii and macmahoni, the behaviour of the adults of the two is very different, suggesting that there are more profound biological differences between them than their morphology would suggest.” The nature of the dark and pale spots on vein R 4+5 (“third vein”) has often been used to separate Anopheles species. In the case of sergentii and macmahoni, a great deal of variation has been noted or illustrated for R 4+ 5 in subspecies macmahoni. This could be an indicator of cryptic species or normal variation. Subspecies sergentii, according to Theobald (1907), Christophers (1933), Mattingly & Knight (1956) and Senevet & Andarelli (1956), has vein R 4+5 all dark or only with a median pale spot and/or small pale spots at the base and apex. In comparison, the scaling of vein R 4+ 5 in subspecies macmahoni has been characterized differently by several authors: “Third vein ranging from entirely dark to about ⅔ pale, usually about ¼–½ pale; pale area when present usually situated distally as in A. funestus, but in 3 cases it was median in the vein” (Evans 1938); “all dark, pale area if present may be at the center” or pale with a long dark basal spot and a small apical dark spot (de Meillon 1947); “it appears that it is normally extensively pale scaled in the middle and only exceptionally dark [all dark?]” (Mattingly & Knight 1956); and Gillies & de Meillon (1968) illustrated the wing showing R 4+5 with a long (⅓ of total length) pale area subapically and perhaps small pale spots at the base and apex, and stated that “3rd vein variable, sometimes broadly pale as figured, in others entirely dark except at base and apex as in subsp. sergenti [sic].” A problem with recognizing subspecies as legitimate ranks, at least for mosquitoes, is illustrated by Irish et al. (2020), who only recognized the nominotypical subspecies and therefore listed the occurrence of only sergentii in countries of sub-Sharan Africa, and Coetzee (2020) who did not include subspecies macmahoni in keys to the adult females of Anopheles in the Afrotropical Region. This prevents macmahoni from being known and confuses records and recognition of both putative subspecies. The two nominal taxa, sergentii and macmahoni, were not recognized as related taxonomically (Evans 1936, 1938; de Meillon 1947) until Mattingly & Knight (1956) considered macmahoni to be a subspecies of sergentii. Subspecies macmahoni is probably mainly confined to the “African portion of the Somali-Arid District” (Mattingly & Knight 1956) and zoophilic, while subspecies sergentii is a north African-Mediterranean anthropophilic malaria vector (Gillies & de Meillon 1968). There exists abundant morphological variation but the only reliable way to separate the two forms is by the relatively large size of the larval tergal plates that characterize macmahoni (Gillies & de Meillon 1968). In a recent review of the mosquitoes of Algeria, Merabti et al. (2021) stated that “Subspecies macmahoni is an Afrotropical form that occurs in many of the countries where the typical form is also found, which suggests that Evans (1936) may have correctly recognized it as a distinct species.” Mattingly & Knight (1956) eliminated subspecies macmahoni from their list for mosquitoes in Arabia while distinguishing it morphologically and bionomically from sergentii. These observations conform to our criteria for species status, i.e. allopatry (and sympatry) with morphological and bionomical distinctions. We therefore reinstate macmahoni to species status: Anopheles (Cellia) macmahoni Evans, 1936 . Anopheles macmahoni is currently listed as a species in the Encyclopedia of Life. Anopheles macmahoni has a single synonym, An. macmahoni var. barkhuusi Giaquinto-Mira, 1950 (type locality: Valley of Becilo, Ethiopia), synonymy by Stone et al. 1959. It is possible that barkhuusi may be a distinct species, but until proven otherwise, it is retained as a synonym of An. macmahoni., 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 73-75, DOI: 10.11646/zootaxa.5303.1.1, http://zenodo.org/record/8043342, {"references":["Evans, A. M. (1936) Two new anopheline mosquitoes from Kenya. Annals of Tropical Medicine and Parasitology, 30 (4), 533 - 540. https: // doi. org / 10.1080 / 00034983.1936.11684958","Mattingly, P. F. & Knight, K. L. (1956) The mosquitoes of Arabia I. Bulletin of the British Museum (Natural History) Entomology, 4 (3), 91 - 141.","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","Christophers, S. R. (1933) The fauna of British India, including Ceylon and Burma. Diptera. Vol. IV. Family Culicidae. Tribe Anophelini. Taylor and Francis, London, vi + (iii) + 371 pp., 3 pls.","Evans, A. M. (1938) Mosquitoes of the Ethiopian Region. II. - Anophelini adults and early stages. British Museum (Natural History), London, x, 404 pp.","de Meillon, B. (1947) The Anophelini of the Ethiopian geographical region. Publications of the South African Institute for Medical Research, 10 (49), 1 - 272.","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.","Senevet, G. & Andarelli, L. (1956) Les Anopheles de l'Afrique du Nord et du Bassin Mediterraneen. Encyclopedie entomologique, 33, l - 280.","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","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","Merabti, B., Boumaza, M., Ouakid, M. L., Carvajal, T. M. & Harbach, R. E. (2021) An updated checklist of the mosquitoes (Diptera: Culicidae) present in Algeria, with assessments of doubtful records and problematic species. Zootaxa, 5027 (4), 515 - 545. https: // doi. org / 10.11646 / zootaxa. 5027.4.3","Giaquinto-Mira, M. (1950) Notes on the geographical distribution and biology of \" Anophelinae \" and \" Culicinae \" in Etiopia [sic]. Revista di Malariologia, 29 (5), 281 - 313.","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."]}

Mimomyia (Mimomyia) chamberlaini Ludlow subspecies chamberlaini Ludlow, 1904b —original combination: Mimomyia chamberlaini. Distribution: Australia, Bangladesh, Cambodia, India, Indonesia, Malaysia, Myanmar, Nepal, Papua New Guinea, People’s Republic of China, Philippines, Sri Lanka, Thailand, Vietnam (Wilkerson et al. 2021). subspecies clavipalpus (Theobald, 1908) —original combination: Radioculex clavipalpus (subspecific status by Mattingly 1957a). Distribution: India, Pakistan, Sri Lanka (Wilkerson et al. 2021). subspecies metallica (Leicester, 1908)—original combination: Conopomyia metallica (subspecific status by Rattanarithikul et al. 2006a). Distribution: Australia, Cambodia, Indonesia, Laos, Malaysia, Papua New Guinea, Philippines, Singapore, Taiwan, Thailand, Vietnam (Wilkerson et al. 2021). As the most recent reviser of Mimomyia (as a subgenus of Ficalbia), Mattingly (1957a) recognized clavipalpus as a subspecies and treated metallica as a variety of chamberlaini. Curiously, Mattingly included chamberlaini sensu stricto and var. metallica in keys for the identification of adult females and males (separate keys), and included only chamberlaini sensu stricto in keys to larvae and pupae. He did not include subspecies clavipalpus in any of the keys. Mattingly & Grjebine (1958), who also recognized Mimomyia as a subgenus of Ficalbia, treated metallica as a species, but explicitly noted that “ F. metallica is probably only a color variety of F. chamberlaini [translated from the French].” It is interesting that Mattingly & Grjebine did not mention clavipalpus, perhaps because their paper was in press when Mattingly (1957a) was published and they also recognized it as a subspecies of chamberlaini. Without comment, Rattanarithikul et al. (2006a) included metallica as a subspecies of chamberlaini in keys to the adults and larvae of the mosquitoes known to occur in Thailand. Prior to these actions, clavipalpus was recognized as a synonym of chamberlaini and metallica was regarded as a separate species (Edwards 1932a). Mimomyia chamberlaini was described from a single male collected at Bayamban, Pangasinan, Luzon Island, Philippines (Ludlow 1904b). Ludlow did not provide illustrations and did not describe the genitalia of the specimen. Four years later, Theobald (1908) described Radioculex clavipalpus from a series of females and males captured at Calcutta, West Bengal, India, but the description did not include illustrations. In the same year, Leicester (1908) described the female and male of Conopomyia metallica from specimens collected at Barrack Pool, Kuala Lumpur, Malaysia. Leicester also did not provide illustrations of the nominal species. Published records indicate that Mi. chamberlaini sensu stricto occurs in most areas of the Oriental Region and tropical areas of the Australasian Region. As pointed out by Mattingly (1957a), available morphological evidence suggests that chamberlaini sensu stricto and metallica are variants of a single species. His assessment is as follows. The classical diagnostic character has been the occurrence in F. metallica of median pale markings on the abdominal tergites [terga] and the absence of such markings in F. chamberlaini. In Malaya complete inter-gradation between these conditions has been found to occur. I have specimens with a more or less continuous pale median line on the abdomen, others with this line reduced to one or two small spots on posterior tergites and one with abdomen completely dark above. As already noted… this last specimen is associated with a larval skin [exuviae] having the distal part of the antenna pale while all my other Malayan larvae have the antenna entirely dark. This is another traditional distinction between F. chamberlaini and F. metallica and were no other evidence available one would be inclined to attribute the one specimen to the first species and the remainder to the second. As against this, however, Philippines specimens with all dark abdomen have larvae with all dark antenna. Other larval differences noted by Edwards & Given… will be seen that at best they are partial differences and they seem to represent differences between the Indian and Indomalayan forms rather than between F. chamberlaini and F. metallica as such. The only significant larval variation within the Indomalayan area appears to be the occurrence of an unusually small number of branches in head seta B [seta 6-C] of the Philippines form (3–5, not less than 4 elsewhere) and a tendency for Malayan larvae to approximate to the Indian form [clavipalpus] in the reduced number of teeth [scales] in the distal row(s) of the comb (1–8, 3–10 elsewhere) and rather less strongly developed spiculation of the saddle edge. In the adult the only notable colour variation concerns the scaling of the prescutellar area. In most cases this is quite extensively pale scaled. Malayan forms, however, tend to have the pale scaling greatly reduced or even absent and my one New Guinea specimen has this area wholly or largely dark. Australian specimens appear regularly to have this area very extensively pale scaled but I have seen individual specimens with it equally pale from Sumatra, Java and Tonkin [Vietnam]. I do not think this variation is taxonomically significant. In view of the striking difference in appearance between well marked specimens of F. chamberlaini and F. metallica it has seemed to me desirable to distinguish the latter as var. metallica, using the term “var.” in the same sense as Edwards (1941: 2) as indicating “bridged variation in one area” as against the geographically representative variation associated with subspecies. In view of the observations and arguments put forward by Mattingly (1957a), until more evidence becomes available, we believe that metallica should be recognized as a synonymous name; therefore, it is here formally placed in synonymy with that species: metallica (Leicester, 1908), junior subjective synonym of Mimomyia (Mimomyia) chamberlaini Ludlow, 1904b . We should add, however, that it would seem likely that Mi. chamberlaini is a complex of species, but as Mattingly aptly stated, “Very much more material is however needed before this can be put forward as more than a most tenuous hypothesis.” For the record, Edwards & Given (1928, Singapore), Galliard & Ngu (1949, Vietnam) and Chen & Lien (1956, Taiwan) treated metallica as a separate species based principally on the characters which Mattingly (1957a) found to be attributable to variation. In agreement here, metallica is not listed as a species in the Encyclopedia of Life. Mattingly (1957a) had the following to say about clavipalpus. The Indian form of F. chamberlaini (= Radioculex clavipalpus Theobald) differs from other forms, so far as can be judged from the limited material available to me, in a number of interesting partial characters of the larva. Thus the antenna is constantly pale on the distal segments in Indian larvae… while in most Indomalayan specimens it is wholly dark. However, I have one larval skin [exuviae] with the Indian type of antenna from Malaya and it may be presumed that the larvae from Tonkin [Vietnam] provisionally assigned to F. chamberlaini by Galliard & Ngu (1949: 497) also had antennae of this type. It is interesting to find that the Malayan skin is associated with the only adult which I have from that territory with complete suppression of median pale markings on the abdominal tergites [terga]. This is, however, of doubtful significance since the U.S. National Museum has adults from the Philippines with complete suppression of these markings associated with larval skins having wholly dark antennae. Other differences between the Indian form and those occurring in the Indomalayan and Australian areas… [include (in general, using up-to-date terminology) setae 5- and 6-C with more branches, scales in distal row(s) of the comb fewer in number, saddle generally with fewer spicules on the posterior margin and pecten with more spines]. …The number of long spicules on the distal edge of the saddle is to some extent a matter of subjective estimate since some spicules are of intermediate length. However, the difference seems quite clear and constant. …It may be felt that these differences justify distinguishing the Indian form as a separate subspecies. Based on the adult and larval distinctions exhibited by clavipalpus, and because it has only been found in areas of India, Pakistan and Sri Lanka where chamberlaini and metallica have not been recorded, we feel that this nominal form should be returned to its original specific rank, at least provisionally: Mimomyia (Mimomyia) clavipalpus (Theobald, 1908 b) . Mimomyia clavipalpus 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 114-116, DOI: 10.11646/zootaxa.5303.1.1, http://zenodo.org/record/8043342, {"references":["Ludlow, C. S. (1904 b) Mosquito notes. - No. 2. Canadian Entomologist, 36 (10), 297 - 301. https: // doi. org / 10.4039 / Ent 36297 - 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","Theobald, F. V. (1908) First report on the collection of Culicidae and Corethridae in the Indian Museum, Calcutta, with descriptions of new genera and species. Records of the Indian Museum, 2 (3), 287 - 302. https: // doi. org / 10.26515 / rzsi / v 2 / i 3 / 1908 / 163338","Mattingly, P. F. (1957 a) The culicine mosquitoes of the Indomalayan Area. Part I: Genus Ficalbia Theobald. British Museum (Natural History), London, 61 pp.","Leicester, G. F. (1908) The Culicidae of Malaya. Studies from the Institute for Medical Research, Federated Malay States, 3 (3), 18 - 261.","Rattanarithikul, R., Harrison, B. A., Panthusiri, P., Peyton, E. L. & Coleman, R. E. (2006 a) Illustrated keys to the mosquitoes of Thailand. III. Genera Aedeomyia, Ficalbia, Mimomyia, Hodgesia, Coquillettidia, Mansonia, and Uranotaenia. Southeast Asian Journal of Tropical Medicine and Public Health, 37 (Supplement 1), 1 - 85.","Edwards, F. W. (1932 a) Genera Insectorum. Diptera, Fam. Culicidae. Fascicle 194. V. Verteneuil & L. Desmet, Bruxelles [Brussels], 258 pp., 5 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.","Edwards, F. W. & Given, D. H. C. (1928) The early stages of some Singapore mosquitos. Bulletin of Entomological Research, 18 (4), 337 - 357. https: // doi. org / 10.1017 / S 0007485300020265","Galliard, H. & Ngu, D. V. (1949) Culicines du Tonkin. II. Les genres Ficalbia et Mochtogenes. Annales de Parasitologie Humaine et Comparee, 24 (5 - 6), 495 - 502. https: // doi. org / 10.1051 / parasite / 1949245495","Chen, H. - H. & Lien, J. - C. (1956) Mosquitoes of genus Ficalbia in Taiwan (Formosa), China. Journal of the Formosan Medical Association, 55 (5), 199 - 215."]}

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

Aedes (Phagomyia) gubernatoris (Giles) subspecies gubernatoris (Giles, 1901a)—original combination: Culex gubernatoris. Distribution: Bangladesh, India, Nepal, People’s Republic of China, Sri Lanka (Wilkerson et al. 2021, incorrectly listed from Thailand). subspecies kotiensis Barraud, 1934 —original combination: Aedes (Finlaya) gubernatoris var. kotiensis (subspecific status by Harbach & Howard 2007). Distribution: India (Western Himalayas) (Barraud 1934). Barraud (1934) described and named kotiensis as a variety of gubernatoris based on larvae that differ in having shorter antennae and lateral palatal brush filaments with “comparatively very large teeth”; however, “adults resulting from these larvae appear to be indistinguishable from the type-form.” Information provided by Barraud indicates that the two forms may be allopatric. In the absence of features that distinguish the adults, the perceived larval distinctions may be associated with conditions that influence growth and development in the tree-hole habitats utilized by the larvae. Based on these observations, we believe that subspecies kotiensis is a morphological variant of Aedes gubernatoris, which we therefore formally recognize as a synonym: kotiensis Barraud, 1934, junior subjective synonym of Aedes (Phagomyia) gubernatoris (Giles, 1901a) . This agrees with the Encyclopedia of Life, which does not list kotiensis as a species. In addition to kotiensis, Ae. gubernatoris has another synonym, Lepidotomyia magna Theobald, 1905a, first recognized by Barraud (1934), which we retain. All three nominal forms have type localities in India., 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 46, DOI: 10.11646/zootaxa.5303.1.1, http://zenodo.org/record/8043342, {"references":["Giles, G. M. (1901 a) A plea for the collective investigation of Indian Culicidae, with suggestions as to moot points for enquiry, and a prodromus of species known to the author. Journal of the Bombay Society, 13, 592 - 610, 2 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","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.","Harbach, R. E. & Howard, T. M. (2007) Corrections in the status and rank of names used to denote varietal forms of mosquitoes (Diptera: Culicidae). Zootaxa, 1542 (1), 35 - 48. https: // doi. org / 10.11646 / zootaxa. 1542.1.3","Theobald, F. V. (1905 a) Genera Insectorum. Diptera, Fam. Culicidae. Fascicle 26. V. Verteneuil & L. Desmet, Bruxelles, 50 pp., 2 pls."]}

Anopheles pseudopunctipennis sensu stricto The nominotypical species of the Pseudopunctipennis Complex was redescribed, illustrated (except for the egg) and a neotype designated using specimens from the type locality, Granada (Rueda et al. 2004). There are two publications of note that distinguish An. pseudopunctipennis sensu stricto from other members of the complex. Manguin et al. (1995), in an isoenzyme study, sampled across the entire range of An. pseudopunctipennis sensu lato, identified three groups/clusters: 1) Granada, 2) USA, Mexico and Guatemala, and 3) Argentina to Belize. They judged that the three groups represented a single species. We agree that their method did not provide the resolution needed to separate all of the species in this complex but we think their results clearly separate the populations at the type locality as a distinct species. In addition, crossing experiments between specimens from Mexico, the USA and Granada (Coetzee et al. 1999) produced non-viable offspring and extensive asynapsis of polytene chromosomes. Coetzee et al. treated their results as evidence of a species complex. We are confident that the results of these two studies, along with adequate distances for genetic isolation, are sufficient to distinguish An. pseudopunctipennis sensu stricto from all other taxa discussed below. The distribution of An. pseudopunctipennis sensu stricto is not clear. It was reported by van der Kuyp (1953) from Curaçao, Margarita [now Venezuelan territory] and Aruba. Shannon et al. (1927) wrote that Puerto España, Trinidad was probably the eastern-most limit of this species. There are extensive records for An. pseudopunctipennis in Venezuela (Gabaldon & Cova-Garcia 1946; Sutil O. 1980). We assume these are conspecific with the Granada population, but this has not been verified. The occurrence of An. pseudopunctipennis sensu stricto in adjoining Colombia should also be considered. Taking into account the evidence/inferences of the validity and distributions of other related species in the complex summarized below, we think the distribution of An. pseudopunctipennis sensu stricto should be changed from the currently listed countries that represent all the species in the complex, found in many publications, to that shown above., 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 59, DOI: 10.11646/zootaxa.5303.1.1, http://zenodo.org/record/8043342, {"references":["Rueda, L. M., Peyton, E. L. & Manguin, S. (2004) Anopheles (Anopheles) pseudopunctipennis Theobald (Diptera: Culicidae): Neotype designation and description. Journal of Medical Entomology, 41 (1), 12 - 22. https: // doi. org / 10.1603 / 0022 - 2585 - 41.1.12","Manguin, S., Roberts, D. R., Peyton, E. L., Fernandez-Salas, I., Barreto, M., Loayza, R. F., Spinola, R. E., Granaou, R. M. & Rodriguez, M. H. (1995) Biochemical systematics and population genetic structure of Anopheles pseudopunctipennis, vector of malaria in Central and South America. American Journal of Tropical Medicine and Hygiene, 53 (4), 362 - 377. https: // doi. org / 10.4269 / ajtmh. 1995.53.362","Coetzee, M., Estrada-Franco, J. G., Wunderlich, C. A. & Hunt, R. H. (1999) Cytogenetic evidence for a species complex within Anopheles pseudopunctipennis Theobald (Diptera: Culicidae). American Journal of Tropical Medicine and Hygiene, 60 (4), 649 - 653. https: // doi. org / 10.4269 / ajtmh. 1999.60.649","van der Kuyp, E. (1953) Culicinae from the Netherlands Antilles and some other Caribbean localities. Studies on the Fauna of Curacao and other Caribbean Islands, 4 (1), 144 - 148.","Shannon, R. C., Davis, N. C. & del Ponte, E. (1927) La distribucion del Anopheles pseudopunctipennis y su relacion con el paludismo, en la Argentina. Revista del Instituto Bacteriologico, 4 (7), 679 - 705.","Gabaldon, A. & Cova-Garcia, P. (1946) Zoogeografia de los anofelinos en Venezuela: II. Los vectores secundarios y los no vectores. Tijeretazos sobre Malaria, 10 (2), 78 - 127."]}

Uranotaenia (Pseudoficalbia) unguiculata Edwards subspecies pefflyi Stone, 1961a —original combination: Uranotaenia unguiculata pefflyi. Distribution: Saudi Arabia (Stone 1961a). subspecies unguiculata Edwards, 1913c —original combination: Uranotaenia unguiculata. Distribution: Afghanistan, Albania, Algeria, Armenia, Austria, Azerbaijan, Bulgaria, Crimea, Croatia, Czech Republic, Egypt, France, FYRO Macedonia, Georgia, Germany, Greece, Hungary, India, Iran, Iraq, Israel, Italy, Jordan, Kazakhstan, Kosovo, Kyrgyzstan, Lebanon, Malta, Moldova, Montenegro, Morocco, Pakistan, Portugal, Romania, Russia, Saudi Arabia, Serbia, Slovakia, Spain, Syria, Tajikistan, Tunisia, Turkey, Turkmenistan, Ukraine, Uzbekistan (Wilkerson et al. 2021, see below regarding Saudi Arabia). Edwards (1913c) described Ur. unguiculata from a single adult male collected at Tiberias, a city in Israel located on the western shore of the Sea of Galilee. The species is widely distributed in the Mediterranean region, extending northward to Germany, eastward to southern Ukraine and the Volga delta, to middle and southwestern Asia [subspecies pefflyi], Iran and Pakistan (Becker et al. 2020; Bromley-Schnur 2021). Stone (1961a) described subspecies pefflyi from eight females and 10 males collected at “Qatif Oasis, Al Hasa Province, Saudi Arabia ”. Today, Qatif Oasis is an urban area known as Qatif or Al-Qatif Governorate on the Persian Gulf coast of Saudi Arabia’s Eastern Province (the former Al Hasa Province is now the largest governorate of the Eastern Province). Wilkerson et al. (2021) listed Saudi Arabia for the distribution of unguiculata, but this obviously came from outdated reports or records published by various researchers, e.g. Alahmed (2012) and Khater et al. (2013), who apparently were unaware of the description of subspecies pefflyi, as noted by Alahmed et al. (2019). In addition to Qatif, specimens identified merely as Ur. unguiculata have been collected at other locations on the Persian Gulf coast of the Eastern Province (Al Dammam and Al Khobar); Al Hasa (or Al Ahsa), the largest city of Al Hasa Governorate located about 65 km west of the Persian Gulf, named after Al Hasa Oasis; and Jezan (Jazan, also spelled Jizan, Gizan or Gazan), a port city and capital of Jazan Province, located on the Red Sea coast directly north of Yemen in southwestern Saudi Arabia (Wills et al. 1985; Alahmed 2012; Khater et al. 2013). Based on these records, Alahmed et al. (2019) stated that “it is unlikely that the typical form, Ur. unguiculata unguiculata, is present in the country”. The typical form is recorded as far south as Baghdad and Karbala, located about 100 km southwest of Baghdad, in central Iraq (Khattat 1955). Subspecies pefflyi has not been recorded north of Qatif in Saudi Arabia, located approximately 320 km southeast of Karbala, Iraq. Alahmed (2012) made collections of adult and larval mosquitoes at “Hafar Al Batin” (also spelled Hafr Al Batin), located about 74 km from the Iraq border in the north of Saudi Arabia’s Eastern Province, but no specimens of Ur. unguiculata were collected. Filatov (2017) mapped the available distribution records for Ur. unguiculata unquiculata in the western Palaearctic, from Portugal eastward to northern India and Kyrgyzstan, but no records were included for Saudi Arabia. Filatov then applied a species distribution modelling approach, using maximum entropy (Maxent) software, to predict the potential distribution of the subspecies in the region based on habitat suitability determined by comparing environmental data at sites where the subspecies had been recorded with sites across the region. Most areas of the region where the typical form has been recorded were ranked as highly suitable, with some intervening and flanking areas ranked as very highly suitable, including the type locality in Israel, and moderately suitable. Except for a few small areas in northern Saudi Arabia that were rated as moderate or low suitability, all areas of the country where subspecies pefflyi has been recorded were found to be unsuitable. Oddly, sites in Egypt along the Nile River some distance south of the Mediterranean coast and at Siwa Oasis near the Libyan border where Ur. unguiculata has been recorded were also classified as environmentally unsuitable. For his description of subspecies pefflyi, Stone (1961a) compared specimens from Qatif Oasis with the original description of unguiculata and 10 specimens (adults) from Algeria, France, Iran, Iraq, Israel and Macedonia. He noted that pefflyi agreed “with the typical subspecies in all particulars except as shown in the following comparative statements. ssp. pefflyi: Smaller, the wing length 1.78–2.38 mm (mean of 10 ♀♀, 10 ♂♂ 1.9 mm). Integument of thorax nearly black, paler only below the pleural scale stripe; dark scales of thorax and abdomen nearly black; usually some dark scales on venter of abdomen; lateral patches of white scales usually confined to terga 5–7. ssp. unguiculata: Larger, the wing length 2.21–3.4 mm (mean of 5 ♀♀, 5 ♂♂ 2.81 mm). Integument of thorax orange brown, slightly darker just mediad of marginal pale scale stripe of scutum; pleura mostly yellowish brown; dark scales of thorax and abdomen rich orange brown, but distinctly not black brown; scales of venter all pale; pale lateral scale patches of abdomen usually start on tergum 2. The adults of the typical subspecies have been described by a number of authors (Edwards 1913c; Barraud 1934; Harant et al. 1952; Senevet & Andarelli 1959; Gutsevich et al. 1971, 1974; Becker et al. 2020), but most of the information in the descriptions is not comparable with the information provided by Stone (1961a). None of the authors measured the length of the wings and, not surprisingly, they do not agree on the color of the thoracic and abdominal scaling. Barraud (1934) is the only author who mentioned the color of the thoracic integument: Mesonotum dark brown or black, not orange brown (Stone); pleura brownish-black, not mostly yellowish brown (Stone). It is obvious that Ur. unguiculata in Saudi Arabia is only found in isolated locations. A question which cannot be answered at this time is whether subspecies pefflyi, as described by Stone, can be identified as such in areas other than Qatif Oasis—after all, isolated populations more recently found in other parts of the country (Wills et al. 1985; Alahmed 2012; Khater et al. 2013) have been identified as merely “ unguiculata ”. Our impression is that the isolated populations in Saudi Arabia are relicts of a species that was more widespread in the past, with populations throughout the Arabian Peninsula until it began to dry up and become desert about 5,000 years ago (Brand 2018). In as much as the populations in Saudi Arabia are readily identifiable as Ur. unguiculata, and the available data indicate that it is a morphologically variable species, we believe it is in the best interests of science to regard pefflyi as nothing more than a morphological variant of a single species. Consequently, we hereby formally classify subspecies pefflyi as a synonymous name: pefflyi Stone, 1961a, junior subjective synonym of Uranotaenia (Pseudoficalbia) unguiculata Edwards, 1913c . Synonym pefflyi, which is currently listed as a species in the Encyclopedia of Life, should be removed from the list of recognized species of the genus Uranotaenia., 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 133-134, DOI: 10.11646/zootaxa.5303.1.1, http://zenodo.org/record/8043342, {"references":["Stone, A. (1961 a) A new subspecies of Uranotaenia unguiculata Edwards from Arabia (Diptera: Culicidae). Proceedings of the Entomological Society of Washington, 62 (4), 249. [for 1960]","Edwards, F. W. (1913 c) Tipulidae and Culicidae from the Lake of Tiberias and Damascus. Journal and Proceedings of the Asiatic Society of Bengal, 9, 47 - 51.","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","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","Bromley-Schnur, H. J. (2021) The mosquitoes of Israel and surrounding areas: An illustrated guide. Wageningen Academic Publishers, Wageningen, 147 pp. https: // doi. org / 10.3920 / 978 - 90 - 8686 - 913 - 8","Alahmed, A. M. (2012) Mosquito fauna (Diptera: Culicidae) of the Eastern Region of Saudi Arabia and their seasonal abundance. Journal of King Saud University - Science, 24 (1), 55 - 62. https: // doi. org / 10.1016 / j. jksus. 2010.12.001","Khater, E. I., Sowilem, M. M., Sallam, M. F. & Alahmed, A. M. (2013) Ecology and habitat characterization of mosquitoes in Saudi Arabia. Tropical Biomedicine, 30 (3), 409 - 427.","Alahmed, A. M., Munawar, K., Khalil, S. M. S. & Harbach, R. E. (2019) Assessment and an updated list of the mosquitoes of Saudi Arabia. Parasites & Vectors, 12, 356. https: // doi. org / 10.1186 / s 13071 - 019 - 3579 - 4","Wills, W. M., Jakob, W. L., Francy, D. B., Oertley, R. E., Anani, E., Calisher, C. H. & Monath, T. P. (1985) Sindbis virus isolations from Saudi Arabian mosquitoes. Transactions of the Royal Society of Tropical Medicine and Hygiene, 79 (1), 63 - 66. https: // doi. org / 10.1016 / 0035 - 9203 (85) 90238 - X","Khattat, F. H. (1955) An account of the taxonomy and biology of the larvae of culicine mosquitoes in Iraq. I. Central Iraq. Bulletin of Endemic Diseases, 1 (2), 156 - 183.","Filatov, S. (2017) Little pigeons can carry great messages: potential distribution and ecology of Uranotaenia (Pseudoficalbia) unguiculata Edwards, 1913 (Diptera: Culicidae), a lesser known mosquito species from the Western Palaearctic. Parasites & Vectors, 10, 464. https: // doi. org / 10.1186 / s 13071 - 017 - 2410 - 3","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.","Harant, H., Rioux, J. & Uzac, S. (1952) Un culicide nouveau pour la France: Uranotaenia unguiculata Edw. Etude ecologique. Annales de Parasitologie Humaine et Comparee, 27 (4), 407 - 418. https: // doi. org / 10.1051 / parasite / 1952274407","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]","Brand, M. (2018) Interview: Unraveling Arabia's green past. Nature Middle East, 9 April 2018, 44. https: // doi. org / 10.1038 / nmiddleeast. 2018.44"]}

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

Aedes (Stegomyia) aegypti (Linnaeus) subspecies aegypti (Linnaeus, 1762) —original combination: Culex aegypti. Distribution: Worldwide tropics and subtropics (Mattingly 1957b; Huang 2004; Powell et al. 2018; Soghigian et al. 2020) [see postscript note regarding current nomenclature]. subspecies formosus (Walker, 1848) —original combination: Culex formosus (subspecific status by Mattingly 1957b). Distribution: Sub-Saharan Africa (Mattingly 1957b; Huang 2004; Powell et al. 2018; Soghigian et al. 2020). It is apropos here to reiterate what we advocated in the Introduction, i.e. we follow the definitions of species and subspecies of de Queiroz (1998, 1999, 2005a, 2005b, 2007, 2020, 2021). Herewith, a brief taxonomic conspectus of Ae. aegypti aegypti and Ae. aegypti formosus is provided, followed by some examples of how formosus fits as an incompletely separated lineage. Aedes aegypti aegypti and Ae. aegypti formosus are members of the Aegypti Group (Huang 2004), which also includes Ae. mascarensis (MacGregor, 1924), type locality Mauritius, an island nation in the Indian Ocean located about 800 km east of Madagascar, and Ae. pia Le Goff & Robert, 2013 (in Le Goff et al. 2013), type locality Mayotte, officially the Department of Mayotte, an Overseas Department of France in the Comoro Archipelago in the Indian Ocean located between Madagascar and Mozambique. Using three molecular datasets, Soghigian et al. (2020) “found that: (a) the Aegypti Group diverged 16 MYA (95% HPD: 7–28 MYA) from its nearest African/Asian ancestor; (b) SWIO [Southwest Indian Ocean] populations of Ae. aegypti are basal to continental African populations; (c) after diverging 7 MYA (95% HPD: 4–15 MYA) from its nearest formally described relative (Ae. mascarensis), Ae. aegypti moved to continental Africa less than 85,000 years ago, where it recently (Ae. aegypti formosus and a human commensal, Ae. aegypti aegypti...”. The latter invaded the New World about 500 years ago via ships involved in the slave trade, and from there it invaded Asia about 150 years ago (Gloria-Soria et al. 2016). Soghigian et al. also noted that “ Ae. pia is clearly the most distantly related and thus serves as an outgroup in [their] later analyses.” They also refer to Ae. aegypti in Madagascar as ‘ aegypti ’ Madagascar due to its large genetic distance from global aegypti sensu stricto and Ae. mascarensis from Mauritius. Mattingly (1957b) considered three forms of aegypti sensu lato. 1. A. aegypti sens. str., the type form [worldwide tropics]. This form is variable in depth of colour but always either distinctly paler and browner (at least in the female) than the blackish African subspecies which is next described, or with pale scaling on the first abdominal tergite [terga], or both. In the type form extensions of pale scaling, if any, are limited either to bleaching of the two dark areas on the back of the head, or to the presence of pale scaling on the first abdominal tergite, or to both in combination. 2. A. aegypti ssp. formosus (Walker) [presumably retrieved from synonymy with aegypti in this cited publication]. This is a geographically representative form confined to Africa south of the Sahara, where it is the only form known to occur, except in coastal districts and in one or two areas of limited inland penetration. It is therefore designated as a subspecies. The name formosus was the first to be applied to any form occurring in the area (Walker, 1848). The type, a female from Sierra Leone, is in the British Museum. It is in poor condition but there is nothing about either it or Walker’s description which would suggest either of the paler forms [the type form, above, and queenslandensis, below]. This subspecies differs from the type form in the markedly blackish appearance of the dark areas of the thorax and abdomen and in entirely lacking any bleaching or extension of pale scaling on any part of the body. It never has any pale scales on the first abdominal tergite. A third form discussed by Mattingly, A. aegypti var. queenslandensis Theobald, 1901a [as Stegomyia fasciata variety] was later synonymized with aegypti sensu stricto by Huang (1979) and shown to be genetically indistinguishable from nominotypical aegypti in Australia (Rašić et al. 2016). It is hypothesized that nominotypical aegypti evolved in sub-Saharan Africa from a forest species utilizing natural containers for larval development, and then diverged into two forms, one remaining adapted to forest habitats (Aedes aegypti formosus) and the other adapted to primarily seek human hosts and to utilize artificial containers provided by humans (Aedes aegypti aegypti). Significantly, McBride et al. (2014) found a genetic change in an odorant receptor responsible for the change to a preference for human blood. A few examples of differences between aegypti and formosus follow, many demonstrating incomplete lineage separation. Note that most work comparing aegypti and formosus in Africa has been carried out in East Africa, while the type locality of formosus is in West Africa (Sierra Leone). Also note that despite the medical importance of aegypti and formosus, morphological studies of all stages have only been carried out on the invasive form in the New World and Asia. No detailed comparisons, other than documentation of adult abdominal markings, have been done for larvae and pupae of African aegypti or formosus, and therefore there are no comparisons with populations outside of Africa either. McClelland (1974) scored pale abdominal markings of Ae. aegypti sensu lato worldwide in relation to biology using an artificial “paleness” scoring scheme. He found continuous intra- and interpopulation variation and determined that identification was not reliable using abdominal markings, i.e. there were discernible but no discrete groupings. He pointed out that part of the problem of correlating markings with habitat was imprecise terminology, and gave the example of “The same population of mosquitoes regarded as ‘feral’ when breeding in a forest would become ‘peridomestic’ if it persisted after a human settlement arose in the forest and ‘urban’ if the village grew to a town.” He then provided a list of what he considered to be more usable terms. In conclusion he stated: “I would venture further that A. aegypti comprises two species or incipient species. The pale or synanthropic species competitively excludes the dark or feral species from habitats where water and man are continuously present. The reverse occurs in habitats where most breeding sites are filled naturally by rainfall.” Scott & McClelland (1975), using starch gel electrophoresis and specimens from eastern Kenya, stated: “The indoor and outdoor ecotypes differed considerably at three loci.” In the laboratory, caged indoor and outdoor ecotypes mated freely, but they stated: “We have demonstrated partial reproductive isolation between them; therefore, what we are calling two ‘ecotypes’ may actually be incipient species, partially isolated by habitat selection.” Paterson et al. (1976) concurred with Scott& McClelland (1975),stating that “the data on the alkaline phosphatase and protein loci... are sufficient to eliminate the possibility that we are dealing with a single polymorphic species. In fact, the data for these two loci provide strong evidence for the existence of positive assortative mating in the field, thus supporting the view that they are distinct genetic species.” Moore (1979), based solely on laboratory crossing experiments, determined that data for aegypti and formosus from Kenya were “consistent with the contention that Aedes aegypti is a single polytypic species.” We note that laboratory mating experiments with closely related, perhaps currently diverging species, probably do not reflect what is happening in nature. Tabachnick et al. (1979), in a larger study of aegypti and formosus, also using starch gel electrophoresis and specimens from (eastern) Kenya, were ambiguous in their judgement of the status of the two forms. They stated: “The results suggest that gene flow between the two forms is restricted. None of the loci examined in this study provided evidence for the complete absence of gene flow between forms. We feel that, at present, the two forms can still be thought of as members of the same polytypic species. However, due to the distinct behavioral differences characteristic of each, and the evidence for restricted gene flow, speciation may be a likely eventuality.” Lounibos (1981) studied the larval ecologies of mosquito species at two sites near the Kenyan coast (Rabai Location, Kilifi District and Shimba Hills National Park, Kwale District). Aedes aegypti larvae [sensu lato] were found in cultivated and ecotonal macrohabitats, predominantly in large tree holes and bamboo. Lounibos also noted that 96% of the Ae. aegypti collected for his study of tree holes and other natural containers corresponded to the morphologically dark subspecies formosus, as characterized by Mattingly (1957b). Most research on the two nominal taxa has been carried out on East African populations. In one study, in contrast to discrete local partitioning of the two forms in Kenya, Sylla et al. (2009), working in Senegal in West Africa, found a clear northwest to southeast cline in change in the proportions of individuals fitting the concept of aegypti versus formosus —in the northwest significantly more aegypti, in the southeast many more formosus. Vector competence measures also followed this cline but molecular genetic measures did not. The authors, however, pointed out many uncontrolled variables in the relatively small study. Jupp et al. (1991) concluded “that in South Africa, Ae. aegypti is a single polymorphic species showing a considerable amount of variation in the degree of white scaling on the first two abdominal tergites [terga]. The presence of some rural sylvan non-anthropophilic populations in our country... might, however, indicate incipient speciation.” Brown et al. (2011) used polymorphic microsatellite markers to test worldwide populations. They “identified two distinct genetic clusters: one included all domestic populations outside of Africa and the other included both domestic and forest populations within Africa. This suggests that human association in Africa occurred independently from that in domestic populations across the rest of the world. …Interestingly, two forms identified as subspecies aegypti and formosus were shown to coexist in a few places along the coast of East Africa, including the Rabai district of Kenya... where the former bred inside homes in villages and the latter bred in surrounding forests.” Three years later, Brown et al. (2014) “used DNA sequences of four nuclear genes and 1504 single nucleotide polymorphism (SNP) markers developed with restriction-site associated DNA (RAD) sequencing to test the hypothesis that Ae. aegypti originated in Africa, where a domestic form arose and spread throughout the tropical and subtropical world with human trade and movement.” In Sudan, Abuelmaali et al. (2021) and Elnour et al. (2022) both reported geographically separated populations of a. formosus (in the west) and a. aegypti (in the east). Elnour et al. (2022) wrote that “Analysis supports a scenario in which Ae. aegypti entered Sudan on at least two independent occasions nearly 70–80 years ago.” We think this raises the possibility of interesting scenarios as aegypti reinvades localities in Africa. For example, if a. formosus evolved into a proto- a. aegypti and then into a. aegypti (see Powell et al. 2018), which then invaded the rest of the world, where is a. aegypti in Africa and how is it evolving in relation to the invasive a. aegypti in the rest of the world? Powell & Tabachnick (2013) stated that a. aegypti, as it occurs in the New World, is not known in West Africa today, except perhaps as a reintroduction into ports (Brown et al. 2011). We think there is abundant evidence concordant with the subspecies concept of de Queiroz (2020) to hereby elevate formosus to its original specific status: Aedes (Stegomyia) formosus (Walker, 1848) . Aedes formosus is currently listed as a species in the Encyclopedia of Life. The following nominal forms are currently listed as synonyms of Aedes aegypti in Wilkerson et al. (2021). Many will remain in doubt until existing types are examined to see if any correspond to Ae. formosus. Some of these are probably from a now extinct Mediterranean (North African, European) distribution of what was thought to be reintroduced Ae. aegypti from the New World (Powell et al. 2018). There are also New World synonyms, which are probably Ae. aegypti, but verification is needed since it is possible that Ae. formosus is present in the New World. Given the unusual biogeography of the Madagascar area, and predictions of undescribed taxa by Soghigian et al. (2020), synonyms from Madagascar and Mauritius also require further scrutiny. African synonyms could potentially be Ae. agypti or Ae. formosus. Unless otherwise indicated, * = Mediterranean, † = New World, + = Madagascar and Mauritius and # = Africa in the following list of current synonyms of Aedes aegypti: * Culex argenteus Poiret, 1787; † Culex fasciatus Fabricius, 1805; * Culex calopus Meigen, 1818; † Culex mosquito Robineau-Desvoidy, 1827; † Culex frater Robineau-Desvoidy, 1827; † Culex taeniatus Wiedemann, 1828; * Culex sugens Wiedemann, 1828; * Culex kounoupi Brullé, 1833; † Culex toxorhynchus Macquart, 1838 a; * Culex annulitarsis Macquart, 1846 a; locality not known, Culex viridifrons Walker, 1848; † Culex excitans Walker, 1848; # Culex inexorabilis Walker, 1848 (could be aegypti or formosus); † Culex exagitans Walker, 1856 a; + Culex insatiabilis Bigot, 1959; Australia, Culex bancrofti Skuse, 1889; * Culex elegans Ficalbi, 1890b; India, Culex rossi Giles, 1899; Australia, Stegomyia fasciata var. queenslandensis; Theobald, 1901a; † Stegomyia fasciata var. luciensis Theobald, 1901a; # Stegomyia nigeria Theobald, 1901a (could be aegypti or formosus); + Stegomyia lamberti Ventrillon, 1904; Canary Islands, Stegomyia calopus var. canariensis Pittaluga, 1905; Philippines, Stegomyia fasciata ssp. persistans Banks, 1906b; Tenerife, Canary Islands, Culex anguste -alatus Becker, 1908; Tenerife, Canary Islands, Culex albopalposus Becker, 1908; Philippines, Duttonia alboannulis Ludlow, 1911b; Australia, Mimeteomyia pulcherrima Taylor, 1919; # Stegomyia fasciata var. atritarsis Edwards, 1920 a. Note. An explanation of the complex nomenclatural history of the nominotypical subspecies is beyond the scope of this treatment. In summary, in the interests of nomenclatural stability, Mattingly et al. (1962) petitioned the International Commission on Zoological Nomenclature (ICZN) to fix the name Aedes (Stegomyia) aegypti (Linnaeus, 1762) with designation of a neotype from Kuala Lumpur, Selangor, Malaya [Malaysia]. The petition was validated in ICZN Opinion 711 (International Commission on Zoological Nomenclature 1964). See Clements & Harbach (2018) for an alternative perspective. Since we now know that the nominotypical form is an invasive species that was restricted to sub-Saharan Africa until only about 500 years ago, we think that the choice of Malaysia for the neotype and type locality, with an even more recent history of invasion than the initial movement to the New World, is problematic., 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 48-50, DOI: 10.11646/zootaxa.5303.1.1, http://zenodo.org/record/8043342, {"references":["Linnaeus, C. (1762) Zweyter Theil, enthalt Beschreibungen verschiedener wichtiger Naturalien. In: Hasselquist, D. F., Reise nach Palastina in den Jahren von 1749 bis 1752. Auf Befehl Ihro Majestat der K ˆ niginn von Schweden. Johann Christian Koppe, Rostock, pp. 267 - 606.","Mattingly, P. F. (1957 b) Genetical aspects of the Aedes aegypti problem. I. - Taxonomy and bionomics. Annals of Tropical Medicine and Parasitology, 51 (4), 392 - 408. https: // doi. org / 10.1080 / 00034983.1957.11685829","Huang, Y. - M. (2004) The subgenus Stegomyia of Aedes in the Afrotropical Region with keys to the species (Diptera: Culicidae). Zootaxa, 700 (1), 1 - 120. https: // doi. org / 10.11646 / zootaxa. 700.1.1","Powell, J. R., Gloria-Soria, A. & Kotsakiozi, P. (2018) Recent history of Aedes aegypti: vector genomics and epidemiology records. BioScience, 68 (11), 854 - 860. https: // doi. org / 10.1093 / biosci / biy 119","Soghigian, J., Gloria-Soria, A., Robert, V., Le Goff, G., Anna-Bella Failloux, A. - B. & Powell, J. R. (2020) Genetic evidence for the origin of Aedes aegypti, the yellow fever mosquito, in the southwestern Indian Ocean. Molecular Ecology, 29 (19), 3593 - 3606. https: // doi. org / 10.1111 / mec. 15590","Walker, F. (1848) List of the specimens of dipterous insects in the collection of the British Museum. Part I. British Museum, London, 229 pp. https: // doi. org / 10.5962 / bhl. title. 57902","de Queiroz, K. (1998) The general lineage concept of species, species criteria, and the process of speciation: A conceptual unification and terminological recommendations. In: Daniel, J., Howard, D. J. & Stewart, H. & Berlocher, S. H. (Eds.), Endless forms: species and speciation, Oxford University Press, Oxford, pp. 57 - 75. https: // doi. org / 10.1046 / j. 1365 - 2540.2000.0704 b. x","de Queiroz, K. (1999) The general lineage concept of species and the defining properties of the species category. In: Wilson, R. A. (Ed.), Species: new interdisciplinary essays, MIT Press, Cambridge, Massachusetts, pp. 49 - 89. https: // doi. org / 10.7551 / mitpress / 6396.001.0001","de Queiroz, K. (2005 a) Ernst Mayr and the modern concept of species. Proceedings of the National Academy of Sciences, 102 (Supplement 1), 6600 - 6607. https: // doi. org / 10.1073 / pnas. 0502030102","de Queiroz, K. (2005 b) A unified species concept and its consequences for the future of taxonomy. Proceedings of the California Academy of Sciences, 56 (Supplement 1), 196 - 215.","de Queiroz, K. (2007) Species concepts and species delimitation. Systematic Biology, 56 (6), 879 - 886. https: // doi. org / 10.1080 / 10635150701701083","de Queiroz, K. (2020) An updated concept of subspecies resolves a dispute about the taxonomy of incompletely separated lineages. Herpetological Review, 51 (3), 459 - 461.","de Queiroz, K. (2021) Response to criticisms of an updated subspecies concept. Herpetological Review, 52 (4), 773 - 776.","MacGregor, M. E. (1924) Aedes (Stegomyia) mascarensis, MacGregor: A new mosquito from Mauritius. Bulletin of Entomological Research, 14 (4), 409 - 412. https: // doi. org / 10.1017 / S 0007485300045648","Le Goff, G., Brengues, C. & Robert, V. (2013) Stegomyia mosquitoes in Mayotte, taxonomic study and description of Stegomyia pia n. sp. Parasite, 20, 31. https: // doi. org / 10.1051 / parasite / 2013030","Gloria-Soria, A., Ayala, D., Bheecarry, A., Calderon-Arguedas, O., Chadee, D. D., Chiappero, M., Coetzee, M., Elahee, K. B., Fernandez-Salas, I., Kamal, H. A., Kamgang, B., Khater, E. K., Kramer, L. D., Kramer, V., Lopez-Solis, A., Saleh, A. A., Sanchez-Casas, R. M., Seixas, G., Sousa, CA., Tabachnick, W. J., Troyo, A. & Powell, J. R. (2016) Global genetic diversity of Aedes aegypti. Molecular Ecology, 25 (21), 5377 - 5395. https: // doi. org / 10.1111 / mec. 13866","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","Huang, Y. - M. (1979) Medical entomology studies - XI. The subgenus Stegomyia of Aedes in the Oriental region with keys to the species (Diptera: Culicidae). Contributions of the American Entomological Institute, 15 (6), 1 - 79.","Rasic, G., Filipovic, I., Callahan, A. G., Stanford, D., Chan, A., Lam-Phua, S. G., Tan, C. H. & Hoffmann, A. A. (2016) The queenslandensis and the type form of the dengue fever mosquito (Aedes aegypti L.) are genomically indistinguishable. PloS Neglected Tropical Diseases, 10 (11), e 0005096. https: // doi. org / 10.1371 / journal. pntd. 0005096","McBride, C. S., Baier, F., Omondi, A. B., Spitzer, S. A., Lutomiah, J., Sang, R., Ignell, R. & Vosshall, L. B. (2014) Evolution of mosquito preference for humans linked to an odorant receptor. Nature, 515, 222 - 227. https: // doi. org / 10.1038 / nature 13964","McClelland, G. A. H. (1974) A worldwide survey of variation in scale pattern of the abdominal tergum of Aedes aegypti (L.) (Diptera: Culicidae). Transactions of the Royal Entomological Society of London, 126 (2), 239 - 259. https: // doi. org / 10.1111 / j. 1365 - 2311.1974. tb 00853. x","Scott, J. A. & McClelland, G. A. H. (1975) Electrophoretic differences between sympatric ecotypes. Nature, 256, 405 - 406. https: // doi. org / 10.1038 / 256405 a 0","Paterson, H. E., Green, C. A. & Mahon, R. J. (1976) Aedes aegypti complex in Africa. Nature, 259, 252. https: // doi. org / 10.1038 / 259252 b 0","Moore, D. F. (1979) Hybridization and mating behavior in Aedes aegypti (Diptera: Culicidae). Journal of Medical Entomology, 16 (3), 223 - 226. https: // doi. org / 10.1093 / jmedent / 16.3.223","Tabachnick, W. J., Munstermann, L. E. & Powell, J. R. (1979) Genetic distinctness of sympatric forms of Aedes aegypti in East Africa. Evolution, 33, (1 part 2), 287 - 295. https: // doi. org / 10.1111 / j. 1558 - 5646.1979. tb 04682. x","Lounibos, L. P. (1981) Habitat segregation among African treehole mosquitoes. Ecological Entomology, 6 (2), 129 - 154. https: // doi. org / 10.1111 / j. 1365 - 2311.1981. tb 00601. x","Sylla, M., Bosio, C., Urdaneta-Marquez, L., Ndiaye, M. & Black, W. C., IV. (2009) Gene flow, subspecies composition, and dengue virus- 2 susceptibility among Aedes aegypti collections in Senegal. PLoS Neglected Tropical Diseases, 3 (4), e 408. https: // doi. org / 10.1371 / journal. pntd. 0000408","Jupp, P. G., Kemp, A. & Frangos, C. (1991) The potential for dengue in South Africa: morphology and the taxonomic status of Aedes aegypti populations. Mosquito Systematics, 23 (3), 182 - 190.","Brown, J. E., McBride, C. S., Johnson, P., Ritchie, S., Paupy, C., Bossin, H., Lutomiah, J., Fernandez-Salas, I., Ponlawat, A., Cornel, A. J., Black, W. C., Gorrochotegui-Escalante, N., Urdaneta-Marquez L., Sylla, M., Slotman, M., Murray, K. O., Walker, C. & Powell, J. R. (2011) Worldwide patterns of genetic differentiation imply multiple ' domestications' of Aedes aegypti, a major vector of human diseases. Proceedings of the Royal Society B, Biological Sciences, 278, 2446 - 2454. https: // doi. org / 10.1098 / rspb. 2010.2469","Brown, J. E., Evans, B. R., Zheng, W., Obas, V., Barrera-Martinez, L., Egizi, A., Zhao, H., Caccone, A. & Powell, J. R. (2014) Human impacts have shaped historical and recent evolution in Aedes aegypti, the dengue and yellow fever mosquito. Evolution, 68 (2), 514 - 525. https: // doi. org / 10.1111 / evo. 12281","Abuelmaali, S. A., Jamaluddin, J. A. F., Noaman, K., Allam, M., Abushama, H. M., Elnaiem, D. E., Ishak, I. H., Wajidi, M. F. F., Jaal, Z. & Abu Kassim, N. F. (2021) Distribution and genetic diversity of Aedes aegypti subspecies across the Sahelian Belt in Sudan. Pathogens, 10, 78. https: // doi. org / 10.3390 / pathogens 10010078","Elnour, M. - A. B., Gloria-Soria, A., Azrag, R. S., Alkhaibari, A. M., Powell, J. R. & Salim, B. (2022) Population genetic analysis of Aedes aegypti mosquitoes from Sudan revealed recent independent colonization events by the two subspecies. Frontiers in Genetics, 13, 825652. https: // doi. org / 10.3389 / fgene. 2022.825652","Powell, J. R. & Tabachnick, W. J. (2013) History of domestication and spread of Aedes aegypti - A review. Memorias do Instituto Oswaldo Cruz, 108 (Supplement I), 11 - 17. https: // doi. org / 10.1590 / 0074 - 0276130395","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","Meigen, J. W. (1818) Systematische Beschreibung der bekannten europaischen zweifl ¸ geligen Insekten. Vol. 1. Gedrukt bei Beaufort Sohn, Aachen, xxxvi + 332 + 1 (errata) pp., 11 pls. https: // doi. org / 10.5962 / bhl. title. 12464","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.","Wiedemann, C. R. W. (1828) Aussereuropaische zweifl ¸ gelige Insekten: als Fortsetzung des Meigenschen Werkes. Vol. 1. in der Schulzischen Buchhandlung, Hamm, xxxii + 608 pp. https: // doi. org / 10.5962 / bhl. title. 14603","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","Macquart, J. (1846) Dipteres exotiques nouveaux ou peu connus. Supplement. Memoires de la Societe Royale des Sciences, de l'Agriculture et des Arts, de Lille, Annee 1844, 133 - 364, 17 pls.","Walker, F. (1856) Insecta Saundersiana: or characters of undescribed insects in the collection of William Wilson Saunders, Esq., F. R. S., F. L. S. & c. Vol. I. Diptera. John van Voorst, London, 474 pp., 8 pls. https: // doi. org / 10.5962 / bhl. title. 5112","Skuse, F. A. A. (1889) Diptera of Australia. Part V. The Culicidae. Proceedings of the Linnean Society of New South Wales, 3 (4), 1717 - 1764, 1 pl.","Edwards, F. W. (1920) Mosquito notes. Bulletin of Entomological Research, 10 (2), 129 - 137. https: // doi. org / 10.1017 / S 0007485300043923","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"]}

Aedes (Stegomyia) gardnerii (Ludlow) subspecies gardnerii (Ludlow, 1905) —original combination: Stegomyia gardnerii. Distribution: Bangladesh, Cambodia, Indonesia, Malaysia, Philippines, Singapore, Thailand, Vietnam (Wilkerson et al. 2021). subspecies imitator (Leicester, 1908) —original combination: Stegomyia imitator (subspecific status by Mattingly 1965). Distribution: Bangladesh, Cambodia, India, Malaysia, Nepal, People’s Republic of China, Taiwan, Thailand, Vietnam (Wilkerson et al. 2021). Ludlow (1905) described the female and male of Stegomyia gardnerii from specimens captured on the Philippine Island of Mindoro (Bulacao), but she also examined specimens from Luzon Island (Angeles, Pampanga).A lectotype male was designated by Knight & Hull (1952), who described the adults in more detail, described the fourth-instar larva and illustrated the male genitalia. Leicester (1908) described Stegomyia imitator from two females captured in jungle near Kuala Lumpur, Malaysia. One of the females was selected as the lectotype by Mattingly (1965). Mattingly described and compared the adults (both sexes), pupa and fourth-instar larva of gardnerii sensu stricto and subspecies imitator, and illustrated the scutal scale-pattern and hindleg of both, the midfemur and entire pupa of imitator, the head and terminal abdominal segments of the larva of the type form, and the male genitalia of imitator. Huang (1977) provided more detailed descriptions of the male, female, pupa and larva, and illustrated the scutal and pleural scaling of both subspecies. For gardnerii sensu stricto, she also illustrated the maxillary palpus and genitalia of the female, the midfemur, male genitalia, entire pupa and entire larva. It is worth noting that in available identification keys (Mattingly 1965; Huang 1977, 1979), the two subspecies are only distinguishable as adults based on the pattern of the scutal scaling. The scutum of the nominate form bears a pair of longitudinal, midlateral stripes of white scales that extend from the anterior margin to the antealar area. The stripes are absent in subspecies imitator, which bears a large patch of white scales on the anterior margin that may be divided medially into two anterolateral patches. The two forms are included but are not distinguished in keys for the male genitalia, pupae and larvae. Huang (1977) seems to have contradicted herself, by saying on the one hand that “Although imitator (Leicester) can easily be distinguished from that of gardnerii (Ludlow) by the scutal markings”, and on the other that “this adult external morphological character (the scutal markings) seems to be geographically variable.” We believe she intended to say that due to variation in the scutal pattern of imitator, “I have here followed Mattingly (1965) in considering imitator as a sub-species of gardnerii.” Wilkerson et al. (2021) mistakenly combined the country records for imitator with those of gardnerii. In the case of gardnerii, the records for Bangladesh, Cambodia, China, Malaysia, Singapore, Thailand and Vietnam apply to subspecies imitator, based on the records of Mattingly (1965), Matsuo et al. (1974, Taiwan), Huang (1977), Darsie & Pradhan (1990, Nepal), Lu et al. (1997, China), Jeffery et al. (2010, Singapore), Rattanarithikul et al. (2010, Thailand), Irish et al. (2016, Bangladesh), Bui & Darsie (2008, Vietnam) and Maquart et al. (2021, Cambodia). As indicated by Huang (1977), the nominate subspecies “is apparently confined to the Eastern part of Indomalayan area… presently known from the Philippines, Sabah [Malaysia], Sulawesi and Alor Island.” Two partial sequences of the cytochrome c oxidase subunit I (COI) gene are registered in GenBank for Ae. gardnerii:Accession MW 321943 (667 bp), Ae. gardnerii from Singapore (Yeo et al. 2021) and accession MK654753 (661 bp), Ae. gardnerii imitator from Thailand (P. Somboon). A BLAST search revealed that the two sequences are identical. Obviously, the sequence from Singapore is derived from a specimen of subspecies imitator, which concurs with the recorded occurrence of this form in the island country (Jeffrey et al. 2010). Although the male genitalia, larva and pupa of gardnerii sensu stricto and subspecies imitator may be indistinguishable, the fact they exhibit distinct and constant differences in the pattern of white scales on the scutum and have separate distributions is a clear indication that they are probably separate species. We believe this is likely to be proven when COI sequences become available for the nominate form. Therefore, it seems prudent to formally reinstate imitator to its original specific status: Aedes (Stegomyia) imitator (Leicester, 1908) . Aedes imitator is currently listed as a species in the Encyclopedia of Life. Four nominal species originally described as species of Stegomyia Theobald, 1901 (in Howard 1901) are regarded as synonyms of Ae. imitator: argenteomaculata Theobald, 1907 (Narcondam Island; synonymy by Huang 1977); christianus Dyar, 1921b (China; synonymy by Mattingly 1965); minutissima Theobald, 1910 a (India; synonymy by Huang 1977); and indosinensis Borel, 1928 (Indochina; synonymy by Huang 1977). These nominal species should remain in synonymy with Ae. imitator; however, it is possible that argenteomaculata could be a distinct species because it is described from a remote island outside the continental range of Ae. imitator., 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 54-55, DOI: 10.11646/zootaxa.5303.1.1, http://zenodo.org/record/8043342, {"references":["Ludlow, C. S. (1905) Mosquito notes. - No. 3. Canadian Entomologist, 37 (3), 94 - 102 + 129 - 135. https: // doi. org / 10.4039 / Ent 3794 - 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","Leicester, G. F. (1908) The Culicidae of Malaya. Studies from the Institute for Medical Research, Federated Malay States, 3 (3), 18 - 261.","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.","Knight, K. L. & Hull, W. B. (1952) The Aedes mosquitoes of the Philippine Islands. II. Subgenera Skusea, Christophersiomyia, Geoskusea, Rhinoskusea, and Stegomyia (Diptera, Culicidae). Pacific Science, 6 (2), 157 - 189. [http: // hdl. handle. net / 10125 / 8813]","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.","Huang, Y. - M. (1979) Medical entomology studies - XI. The subgenus Stegomyia of Aedes in the Oriental region with keys to the species (Diptera: Culicidae). Contributions of the American Entomological Institute, 15 (6), 1 - 79.","Matsuo, K., Yoshida, Y. & Lien, J. C. (1974) Scanning electron microscopy of mosquitoes. II. The egg surface structure of 13 species of Aedes from Taiwan. Journal of Medical Entomology, 11 (2), 179 - 188. https: // doi. org / 10.1093 / jmedent / 11.2.179","Darsie Jr., R. F. & Pradhan, S. P. (1990) The mosquitoes of Nepal: their identification, distribution and biology. Mosquito Systematics, 22 (2), 69 - 130.","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.","Jeffery, J., Lee, R. M. L., Tan, S. Y., Liew, C., Ng, L. C. & Lam-Phua, S. G. (2010) New mosquito species records (Diptera: Culicidae) from Singapore. Tropical Biomedicine, 27 (1), 138 - 142.","Rattanarithikul, R., Harbach, R. E., Harrison, B. A., Panthusiri, P., Coleman, R. E. & Richardson, J. H. (2010) Illustrated keys to the mosquitoes of Thailand. VI. Tribe Aedini. Southeast Asian Journal of Tropical Medicine and Public Health, 41 (Supplement 1), 1 - 225.","Irish, S. R., Al-Amin, H. M., Alam, M. S. & Harbach, R. E. (2016) A review of the mosquito species (Diptera: Culicidae) of Bangladesh. Parasites & Vectors, 9, 559. https: // doi. org / 10.1186 / s 13071 - 016 - 1848 - z","Bui, P. & Darsie Jr., R. F. (2008) Tentative checklist of the mosquitoes of Vietnam employing new classification for tribe Aedini (Diptera, Culicidae). Journal of the American Mosquito Control Association, 24 (2), 187 - 193. https: // doi. org / 10.2987 / 5665.1","Maquart, P. - O., Fontenille, D., Rahola, N., Yean, S. & Boyer, S. (2021) Checklist of the mosquito fauna (Diptera, Culicidae) of Cambodia. Parasite, 28, 60. https: // doi. org / 10.1051 / parasite / 2021056","Yeo, H., Yeoh, T. X., Ding, H., Lee, T. T. M. & Puniamoorthy, N. (2021) Morphology and mini-barcodes: The inclusion of larval sampling and NGS-based barcoding improves robustness of ecological analyses of mosquito communities. Journal of Applied Ecology, 58 (10), 2087 - 2100. https: // doi. org / 10.1111 / 1365 - 2664.13966","Howard, L. O. (1901) Mosquitoes: how they live; how they carry disease; how they are classified; how they may be destroyed. McClure, Phillips & Co., New York, New York, xv + 241 pp. https: // doi. org / 10.5962 / bhl. title. 30512","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","Dyar, H. G. (1921 b) Two new mosquitoes from China (Diptera, Culicidae). Insecutor Inscitiae Menstruus, 9 (7 - 9), 147 - 148.","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","Borel, E. (1928) Les moustiques de la Cochinchine et du Sud-Annam (Suite). Archives des Instituts Pasteur d'Indochine, 7, 75 - 106, 8 pls."]}

Eretmapodites wansoni Edwards subspecies douceti Adam & Hamon, 1959 —original combination: Eretmapodites oedipodius [oedipodeios] ssp. douceti. Distribution: Burkina Faso, Côte d’Ivoire (Wilkerson et al. 2021). subspecies wansoni Edwards, 1941 —original combination: Eretmapodites oedipodius [oedipodeios (sic)] ssp. wansoni (specific status by Hamon 1961). Distribution: Burkina Faso, Cameroon, Central African Republic, Côte d’Ivoire, Democratic Republic of the Congo, Republic of the Congo, Togo (Wilkerson et al. 2021). Prior to Hamon (1961), the species known as Er. oedipodius Edwards, 1912b consisted of six subspecies: The nominotypical subspecies, douceti Adam & Hamon, 1959, marcelleae Adam & Hamon, 1959 (incorrectly spelled marcellei, emended by White 1975), parvipluma Edwards, 1941, stanleyi Edwards, 1941 and wansoni Edwards, 1941. Hamon (1961) synonymized stanleyi with oedipodius, raised marcelleae, parvipluma and wansoni to specific rank, and recognized douceti as a subspecies of wansoni, based mainly on characters of the genitalia and the ornamentation of the mid- and hindtarsi of males. He proffered the following taxonomic assessment (translated from the French). The subspecies oedipodius, parvipluma, wansoni, marcellae [sic] and douceti are very clearly distinguished from each other by the male genitalia and can be classified into two groups depending on whether the simple claw of the midleg is flattened (oedipodius and marcellae) or not (parvipluma, wansoni and douceti). The simultaneous presence of marcellae and oedipodius in the same region and the great differences that exist in the structure of the apical leaf of the proximal claspette make it necessary to treat them as distinct species. Although the ranges do not coincide (within the limits of our current knowledge), it also seems logical to treat parvipluma and wansoni as separate species, given the differences in the structure of the proximal claspette which, in parvipluma, bears in its apical part two long fine setae and a spiculate leaf while, in wansoni, this claspette has half a dozen long fine setae and ribbed leaves with unspiculate edges. The status of douceti is less clear and we propose to maintain it as a subspecies of wansoni, given the affinities found both in the structure of the male genitalia and the claw of the midleg and in the ornamentation of the posterior tarsus [hindtarsus]. It seems that wansoni is confined to the lower Guinean forest district and douceti exists only in the upper Guinean savannah district, but there are still very large gaps in collecting in these areas. Subspecies douceti was described from three males, two reared from larvae collected from water held in a fallen leaf in the Nasso forest of present-day Burkina Faso and one captured while flying in undergrowth of oil palms at Katiola, Ivory Coast (Adams & Hamon 1959). In addition to those specimens, Hamon (1961) also examined a male from Togo, which only differed from the type specimens of douceti in the apical leaf of the proximal claspette being bifid at the apex. Adam & Hamon (1959) described the male of douceti as follows (translated from the French). The posterior tarsus [hindtarsus] is similar to that of Eretmapodites oedipodius wansoni Edwards: the 3rd tarsomere bears neither erect scales nor long setae; the 4th and 5th tarsomeres are enlarged and pinnate, each forming a pronounced angle with the preceding tarsomere. The 5th tarsomere of the midleg does not bear an abnormally enlarged claw (fig. 3 B). The male genitalia (fig. 2) are characterized by the proximal claspette which is short, quite thick in its basal 2/3, then sharply narrows and bears half a dozen setae at the point of narrowing, but none are inserted close to the leaf as in E. oe. wansoni; the terminal leaf is quite broad, tapering at its apex and barbed laterally. The distal claspette is long, curved, with a rounded apex bearing a long lateral seta, six small joined leaves thinning into a filament at their end and 12 to 14 strong setae curved in the form of a hook. Based on these differences, and the fact that douceti is known only from localities in savannah, as opposed to wansoni being known only from forest sites, it is surprising that Haman (1961) preferred to recognize douceti as a subspecies of wansoni. We believe that when the larval and pupal stages of wansoni are known and compared with those of douceti, and molecular data become available for both forms, it will be apparent that douceti and wansoni are separate species. Until proven otherwise, we hereby formally elevate douceti to species rank: Eretmapodites douceti Adam & Hamon, 1959 . Eretmapodites douceti is currently listed as a species in the Encyclopedia of Life, however the date of authorship needs to be corrected from 1958 to 1959. The issue of the journal (no. 4) intended for publication in 1958 was not published until 1959., 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 112-113, DOI: 10.11646/zootaxa.5303.1.1, http://zenodo.org/record/8043342, {"references":["Adam, J. P. & Hamon, J. (1959) Description de deux nouvelles sous-especes d' Eretmapodites oedipodius Graham, capturees en Afrique Occidentale. Bulletin de la Societe de Pathologie exotique, 51 (4), 524 - 527. [for 1958]","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.","Hamon, J. (1961) Les Eretmapodites du groupe oedipodius Graham. 1. Descriptions complementaires et position systematique des six sous-especes actuellement cannues. Bulletin de la Societe de Pathologie exotique, 54 (4), 892 - 906.","Edwards, F. W. (1912 b) A synopsis of the species of African Culicidae, other than Anopheles. Bulletin of Entomological Research, 3 (1), 1 - 53. https: // doi. org / 10.1017 / S 0007485300001656","White, G. B. (1975) Notes on a catalogue of Culicidae of the Ethiopian Region. Mosquito Systematics, 7 (4), 303 - 344."]}

Anopheles (Cellia) wellcomei Theobald subspecies ugandae Evans, 1934 —original combination: Anopheles distinctus var. ugandae (subspecific status by Gillies & de Meillon 1968). Distribution: Angola, Botswana, Burundi, Democratic Republic of the Congo, Gabon, Gambia, Kenya, Senegal, Tanzania, Uganda (Ribeiro & Ramos 1975; Gillies & Coetzee 1987; Wilkerson et al. 2021). subspecies ungujae White, 1975 —original combination: Anopheles (Cellia) wellcomei ungujae. Distribution: Island of Zanzibar, Tanzania (White 1975). subspecies wellcomei Theobald, 1904 —original combination: Anopheles wellcomei. Distribution: Angola, Benin, Botswana, Burkina Faso, Cameroon, Central African Republic, Chad, Côte d’Ivoire, Democratic Republic of the Congo, Eritrea, Ethiopia, Gabon, Gambia, Ghana, Guinea, Kenya, Mali, Mauritania, Mozambique, Niger, Nigeria, Senegal, Seychelles, South Sudan, Sudan, Tanzania, Togo, Uganda, Zambia, Zimbabwe (Wilkerson et al. 2021). The nominal taxa treated here are members of the Wellcomei Group (Gillies & de Meillon 1968) of the Myzomyia Series (Christophers 1924), which also includes An. distinctus (Newstead & Carter, 1911), An. erepens Gillies, 1958 and An. theileri Edwards, 1912c. Species in the Wellcomei Group lack aedeagal leaflets (except theileri) and have adaptations in the immature stages for climbing out of the water (Evans 1934). The larvae have large dense hair-like spicules on the thorax and abdomen, and pupae have a row of ventrally directed hooks on the cephalothorax and the paddle is fringed externally with coarse spines. Verification of these characters for subspecies ungujae is still needed, because it is known only from a single female. The original description of An. wellcomei (adult female) includes an informative color illustration. Theobald’s (1904) diagnosis follows: “Head black with dense white, yellow and brown upright forked scales, the white ones in front and two long hair-like projecting white tufts; palpi yellow, black at the base with two white bands on the yellow area. Thorax ashy, chestnut-brown at the sides and with hair-like golden scales; abdomen brown, unbanded with brownish golden hairs. Wings mostly yellow scaled, costa jet black with two yellow spots and three or four black spots on the wing field.” Also of note are other characters from the original description and subsequent observations. The maxillary palpus is black basally and ochreous (yellow orange to orange) on approximately the distal two-thirds, with two broad white bands and a third narrow band apically. The proboscis is dark on the proximal half, and the distal half is ochreous like the palpus. The wing is distinctive and was later described by Evans (1927): “To the naked eye the wings seem yellow with a narrow, black anterior border.” Also from the original description: Costal wing spots, on costa and vein R 1, include subcostal, preapical and apical pale spots. There is also a small sector pale spot on R 1, which is almost entirely pale proximally. Wing mostly pale yellowish with small dark spots (noted by Theobald to vary) on veins R 2, R 3, R 4+5, M 1, M 2 and 1A. Fringe scales pale at ends of all veins, wing apex and entire border of the anal cell. “Legs brown with very narrow apical yellow bands.” Legs subsequently described by Gillies & de Meillon (1968) as: “Tarsus 1–3 of fore legs [foretarsomeres 1–3] with distinct but narrow apical pale bands; mid legs similar but pale bands less distinct; hind legs with tarsus 1–4 [hindtarsomeres 1–4] distinctly banded apically.” Anopheles wellcomei was described from three females from Baro and Pibor, Sudan (now in South Sudan). “Dr. Balfour states that ‘it boarded the steamer in the evening at Baro and bit freely.’” Also, “It is abundant on the Baro [River].” Townsend (1990) recorded the label data for the three syntypes as “ wellcomei Theobald, 1904 c: 64–66 (Anopheles). Syntypes (3) ‒ Sudan: 1 female, [Eastern Equatoria], Baro [5 o 45′N 31°42′E]; 2 females, [Sobat], Pibor.” David Pecor (pers. comm.) interprets the Baro and Pibor collection localities to be place names with coordinates of 5.75, 31.7 and 6.79853, 33.13045, respectively (see: https://arcg.is/0D4nD). Two other nominal species were included with the above species by Evans (1934) in the “ distinctus series” (Anopheles walravensi Edwards, 1930 and Anopheles schwetzi Evans, 1934). They “share a peculiarity in wing markings, namely, the reduction or loss of some of the pale costal spots, especially of the sector and subcostal, at least the basal 2/3 of the costa being entirely dark...”. The relative vagueness of the lack of pale spots on the costa and few dark spots on the rest of the wing (along with similarity in leg coloration) presumably led taxonomists to include similar-looking species in the concept of the “ distinctus series”. Also vague to us is the ill-defined designation of varieties and subspecies throughout the taxonomic history of this group of species. To illustrate, it is perhaps informative to look at name combinations over time. Pyretophorus distinctus Newstead & Carter, 1911: To genus Anopheles by Evans (1927). Pyretophorus distinctus var. melanocosta Newstead & Carter, 1911: To synonym of An. distinctus by Evans (1934). Anopheles theileri Edwards, 1912c: New name for Pyretophorus albipes Theobald, 1911a. Anopheles theileri var. brohieri Edwards, 1929b: To species rank, An. brohieri, by Gillies & de Meillon (1968). Anopheles distinctus var. ugandae Evans, 1934: To An. wellcomei subspecies ugandae by Gillies & de Meillon (1968) (elevated to species herein). Anopheles walravensi Edwards, 1930: No change. Anopheles schwetzi Evans, 1934: No change. Anopheles theileri var. septentrionalis Evans, 1934: To synonym of An. brohieri Edwards, 1929b by Gillies & de Meillon (1968). Anopheles walravensi var. milesi de Meillon & Evans, 1935: To synonym of An. wellcomei subspecies ugandae by Gillies & de Meillon (1968). Anopheles michaeli de Meillon & Leeson, 1940: To synonym of An. schwetzi by Gillies & de Meillon (1968). Anopheles wellcomei subspecies erepens Gillies, 1958: To species rank, An. erepens, by Gillies & Coetzee (1987). Anopheles wellcomei subspecies ungujae White, 1975 (elevated to species herein). Subspecies ugandae was described from Kampala, Uganda. The two type localities of wellcomei, Baro and Pibor, are 606 km and 716 km, respectively, north of Kampala. Larvae, pupae and adult females were available to Evans (1934), and were illustrated in part. A holotype female with associated larval and pupal exuviae is in the Natural History Museum, London. Evans (1934) did not clearly distinguish ugandae from other taxa in the “ distinctus series” but instead alluded to differences between it and “pale forms of distinctus ”, “the type of the former var. melanocosta ”, pupal paddle “Shape more nearly ovoid than usual, but apparently less so than in theileri and the type form of distinctus ”, “Larvae and pelts [exuviae] resembled those of the type form and of A. theileri ”, “Outer clypeals, as in type form and some specimens of theileri, very short and bluntly pointed distally” and, in addition, there were vague comparisons to the antennae and abdominal setae of theileri. The distinguishing character, however, provided in a key to adult females, is “Outer half of proboscis with creamy or whitish scales; palpi with dark brown scales confined to the basal one-third” in wellcomei, and “Outer half of proboscis dark-scaled; palpi with dark-scaled areas on outer two-thirds” in walravensi, distinctus var. ugandae, distinctus “type form” and schwetzi. These same characters were used by Gillies & Coetzee (1987) to distinguish wellcomei wellcomei from wellcomei ugandae. Gillies & Coetzee were not able to separate the larvae or pupae of species in the Wellcomei Group (as the wellcomei section). Gillies & de Meillon (1968) mapped the distributions of subspecies wellcomei and ugandae (their fig. 47). Subspecies wellcomei is shown across central sub-Saharan Africa and subspecies ugandae is distributed more in southeastern Africa, but with a large overlap of the two in Uganda, Republic of Congo, Tanzania, Kenya and southern South Sudan. Gillies & Coetzee (1987) stated that “ Gillies and De Meillon (1968) recognised the 3 subspecies, w. wellcomei, w. ugandae and w. erepens. The last named was a very dark form from an isolated area in Tanzania and Kenya. The pale form with flavescent palps and proboscis (wellcomei) is typical of the northern savanna while, from Uganda southwards, forms lacking this overlay of pale scales predominate (ugandae). However, material collected more recently from the western limits of the northern savanna in Senegal (Dr M Cornet) and from adjacent parts of The Gambia lacks the flavescent scales and has a rather darker wing, both characters diagnostic of subsp. ugandae. This pattern of variation, in which there is a tendency for certain peripheral as well as southern populations to become darker, means that ugandae is a rather less satisfactory taxon than appeared when Gillies and De Meillon redefined it. However, the name ugandae must be applied to Sene-gambian specimens even though this invalidates to a certain extent the picture of distribution previously presented.” This uncertainty suggests to us that the variability could indicate a species complex. Given the diagnostic character of the pale coloration of the apical half of the maxillary palpus and proboscis in subspecies wellcomei, which are dark in subspecies ugandae, the relative closeness of type localities and the extensive overlapping distributions, we believe these are genetically distinct taxa. Therefore, we hereby elevate subspecies ugandae to specific status: Anopheles (Cellia) ugandae Evans, 1934 . Anopheles ugandae is currently listed as a species in the Encyclopedia of Life. As indicated above, we agree with Gillies & de Meillon (1968) that An. walravensi var. milesi de Meillon & Evans, 1935 (type locality: Victorian Falls, Zimbabwe) is probably a synonym of An. ungandae (type locality: King’s Lake Area, Kampala, Uganda). However, the possibility that further study could reveal that milesi is a separate species cannot be ruled out. White (1975) named and designated a holotype for An. wellcomei subspecies ungujae based on a brief description by Gillies (1958) of a single female from Zanzibar, which Gillies called both Anopheles wellcomei subsp. indet. and A. wellcomei, Zanzibar form. Gillies noted that “The finding of A. wellcomei on the island of Zanzibar extends the known range of the species right across Africa. It is interesting to note that moderately intensive collecting on the adjacent East African coast has failed to reveal this species, although erepens is abundant in the arid regions 100 or so miles inland.” White (1975) stated that “Apart from its discreet distribution, this subspecies is readily separable from other forms of wellcomei by extension of the pale sector wing-spot onto the costa and by the almost completely pale first wing vein (vide Figure 4 in Gillies, 1948:10; Plate 82b in Gillies & De Meillon, 1968:177).” However, a complete description of the holotype is needed. Based on the stated discrete characters and distribution we believe, pending further morphological and molecular study, that subspecies ungujae should be afforded species status: Anopheles (Cellia) ungujae White, 1975 . Anopheles ungujae is currently listed as a species in the Encyclopedia of Life. Without a complete description of the holotype and collection and study of the adult male and immature stages, its placement in the Wellcomei Group remains uncertain., 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 80-82, DOI: 10.11646/zootaxa.5303.1.1, http://zenodo.org/record/8043342, {"references":["Evans, A. M. (1934) Further notes on African anophelines, with a description of a new group of Myzomyia. Annals of Tropical Medicine and Parasitology, 28 (4), 549 - 570. https: // doi. org / 10.1080 / 00034983.1934.11684821","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.","Ribeiro, H. & da Cunha Ramos, H. (1975) Research on the mosquitoes of Angola. VI - The genus Anopheles Meigen, 1818 (Diptera, Culicidae). Check-list with new records, keys to the females and larvae, distribution and bioecological notes. Garcia de Orta, Serie de Zoologia, 4 (1), 1 - 39, 17 pls.","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.","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","White, G. B. (1975) Notes on a catalogue of Culicidae of the Ethiopian Region. Mosquito Systematics, 7 (4), 303 - 344.","Theobald, F. V. (1904) The mosquitoes of Egypt, the Sudan and Abyssinia. In: Balfour, A. (director), First Report of the Wellcome Research Laboratories at the Gordon Memorial College Khartoum. Department of Education, Sudan Government, Khartoum, pp. 62 - 83, 2 pls.","Christophers, S. R. (1924) Provisional list and reference catalogue of the Anophelini. Indian Medical Research Memoirs, 3, 1 - 105.","Newstead, R. & Carter, H. F. (1911) On some new species of African mosquitos (Culicidae). Annals of Tropical Medicine and Parasitology, 5 (2), 233 - 244, 1 pl. https: // doi. org / 10.1080 / 00034983.1911.11686353","Gillies, M. T. (1958) Notes on the biology of a new subspecies of Anopheles wellcomei (Diptera: Culicidae) from East Africa, and on the distribution of related forms. Proceedings of the Royal Entomological Society of London, Series A, General Entomology, 33 (1 - 3), 9 - 14. https: // doi. org / 10.1111 / j. 1365 - 3032.1958. tb 00386. x","Edwards, F. W. (1912 c) A key for determining the African species of Anopheles (sensu lato). Bulletin of Entomological Research, 3 (3), 241 - 250. https: // doi. org / 10.1017 / S 0007485300001917","Evans, A. M. (1927) A short illustrated guide to the anophelines of tropical and South Africa. Memoirs of the Liverpool School of Tropical Medicine, 3, 1 - 79.","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.","Edwards, F. W. (1930) Mosquito notes. - IX. Bulletin of Entomological Research, 21 (3), 287 - 306. https: // doi. org / 10.1017 / S 0007485300021805","Theobald, F. V. (1911 a) The Culicidae or mosquitoes of the Transvaal. Report of the Division of Entomology of the Department of Agriculture, Union of South Africa, 1, 232 - 272, 1 pl.","Edwards, F. W. (1929 b) Mosquito notes. - VIII. Bulletin of Entomological Research, 20 (3), 321 - 343. https: // doi. org / 10.1017 / S 0007485300021283","de Meillon, B. & Evans, A. M. (1935) Two new anophelines from South Africa. Annals of Tropical Medicine and Parasitology, 29 (1), 91 - 94. https: // doi. org / 10.1080 / 00034983.1935.11684834","de Meillon, B. & Leeson, H. S. (1940) Notes on Ethiopian Anophelini. Bulletin of Entomological Research, 31 (1), 61 - 67. https: // doi. org / 10.1017 / S 0007485300004818"]}

Aedes (Aedimorphus) hirsutus (Theobald) subspecies adenensis Edwards, 1941 —original combination: Aedes (Aedimorphus) hirsutus var. adenensis (as ssp. in Pao & Knight 1970, explicitly elevated to ssp. by White 1975). Distribution: Yemen (Edwards 1941). subspecies hirsutus (Theobald, 1901a)—original combination: Culex hirsutum [sic]. Distribution: Benin, Burkina Faso, Côte d’Ivoire, Ethiopia, Kenya, Madagascar, Mali, Mauritania, Namibia, Nigeria, Senegal, South Africa, South Sudan, Sudan, Togo, Zambia, Zimbabwe (Wilkerson et al. 2021). The nominotypical subspecies was described from a female and a male from Salisbury, Mashonaland [Harare, Zimbabwe]. The detailed description has a separate color illustration. Theobald’s (1901a) diagnosis follows here (after Theobald 1901a [plates]): fig. 80, pl. XX): “Thorax dark chestnut-brown, with dull creamy scales forming a broad median line and four creamy spots. Abdomen dark umber-brown, with basal yellowish-white bands and with two large dull yellow apical spots on the fifth and sixth segments; venter mottled with black, yellow and white scaled patches. Legs with broad basal white bands, femora, tibiae and metatarsi [tarsomere 1] hirsute, bases of the femora pale, remainder black, with a few pale scales.” Also, “palpi short, dark brown, with numerous pale scales at the apex”. Subsequent authors (Edwards 1911b, 1912 b, 1941; Jupp 1996) noted that there is a white costal fringe on the distal part of the wing, which was not mentioned or illustrated by Theobald (1901a). Theobald (1903a) described a similar taxon, Culex transvaalensis, but he did not note a similarity with hirsutus. In the collection of the Natural History Museum, London, Townsend (1990) found three syntype females from South Africa [Transvaal], Pretoria, and a male labelled “type”, which he stated had no type status since only the female was described. Without comment Edwards (1911b) reduced transvaalensis to a synonym of hirsutum. Theobald (1913a) distinguished the two species in a key: “Abdomen basally banded” for transvaalensis and “Abdomen with two apical spots on 5th and 6th segments as well as bands” for hirsutum. He also stated: “Edwards considers this [hirsutum] the same as Culex transvaalensis, Theobald. The two marked apical spots on both fifth and sixth abdominal segments in Culex hirsutum never occur in Culex transvaalensis and the thorax is differently ornamented. The type has either been altered or this young writer has been unable to appreciate the distinctive characters.” He further added: “As pointed out under C. hirsutum, Theobald, Edwards is in complete error in vaguely assuming this [transvaalensis] is the same as that marked species.” The discussion was apparently put to rest by Edwards (1915), who stated: “Theobald... has criticised my statement that his Culex hirsutum and C. transvaalensis are one and the same species. I have re-examined the specimens, and find no reason to modify my previous opinion, except in one in particular. The male and female types of C. hirsutum belong to different species.... The female type must be taken as the type of the species, the male… being really O. nigeriensis (Theo.) [currently a synonym of Ae. (Aed.) fowleri (de Charmoy, 1908)].” Other described life stages of the nominate form hirsutus, a widespread African species, include the adult male (Edwards 1911b), pupa (Ingram & Macfie 1919), male genitalia (Edwards 1941), larva (Hopkins 1952), larval mandible and maxilla (Pao & Knight 1970) and the adult female and male genitalia (Jupp 1996). Edwards (1941) described subspecies adenensis as a variety of hirsutus from two sites in the West Aden Protectorate [southwestern Yemen], as revealed by Mattingly (1956): Jebel Jihaf, 7100 ft., ix.1937, 2 females, and Dhala, 4800 ft., 14.ix.1937, 1 male and 7 females. “Since the critical factor, if any, is likely to be altitude I have marked one of the ♀ from Jebel Jihaf as the hololectotype and the ♂ from Dhala as allolectotype.” David Pecor (pers. comm.) investigated the localities and noted that the West Aden Protectorate was renamed as part of the independent nation of Yemen, and the locality names were perhaps misspelled when translated from Arabic. Instead they are probably Jabal Jihaf [Jabal is “mountain” in Arabic] (GeoNames ID 74169, coordinates 13.7607 44.67656, altitude 2,402 m) and Dhale, (GeoNames ID 80384, coordinates 13.69572, 44.73137, altitude 1,517 m). The two sites are approximately 9 km apart. Edwards’s (1941) description of adenensis follows: “Structurally identical with the typical form, and also resembling it in the white-scaled distal part of costa, but differs in colouring of abdomen; the pale basal bands of the tergites [terga] are yellowish rather than white and all the tergites have apical median patches of yellowish scales, usually occupying an extensive area; in some extremely pale specimens the abdominal scales are almost all yellowish, with only a few black ones across the middle of each tergite.” Mattingly & Knight (1956) stated that the male genitalia [of adenensis] “are indistinguishable from those of the type form.” In their keys they noted: “Wing [of adenensis] with relatively few pale scales which are mainly confined to the basal half”. They then compared the larva of adenensis to Ae. (Aed.) natronius Edwards, 1932b: “Antenna more strongly spiculate; pecten teeth of more normal type; head seta 6 normally single, rarely double, much longer and usually stouter than 5”. The illustration in the original description showing the presence of white scaling on the costa of the wing contradicts Mattingly & Knight (1956), who reported the presence of a few pale scales, those confined to the proximal half of the costa. The head of the larva of hirsutus illustrated by Hopkins (1952), who did not mention adenensis, shows seta 5-C multi-branched and 6-C single, with the latter about twice the length of the former. Unfortunately, Mattingly & Knight (1956) did not note whether 6-C was branched or not. The inconsistent description of white scales being present or absent apically on the costa in both hirsutus and adenensis can only be solved by re-examination of the types and study of variation in these taxa. The only character that separates these two, for now, is the pattern of scaling on the abdomen of adenensis, which has median, apical patches of yellowish scales that are absent in hirsutus. No description of variation in hirsutus across its range in Africa includes median apical abdominal pale scales that would suggest adenensis represents a morphological variant of the type form. However, the type form has not been well studied morphologically. The type locality of adenensis, near the eastern border of the distribution of hirsutus, is at a fairly high altitude compared to most mosquito species and appears to us to possibly reside in an extremely dry environment, though indications of green in valleys can be seen from satellite photos. Until further study can be undertaken, we believe adenensis is sufficiently different from hirsutus to be afforded species status: Aedes (Aedimorphus) adenensis Edwards, 1941 . Aedes adenensis 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 13-14, DOI: 10.11646/zootaxa.5303.1.1, http://zenodo.org/record/8043342, {"references":["Edwards, F. W. (1941) Mosquitoes of the Ethiopian Region III. - Culicine adults and pupae. Printed by order of the Trustees, British Museum (Natural History), London, viii + 499 pp.","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.","White, G. B. (1975) Notes on a catalogue of Culicidae of the Ethiopian Region. Mosquito Systematics, 7 (4), 303 - 344.","Theobald, F. V. (1901 a) A monograph of the Culicidae or mosquitoes. Vol. 1. British Museum (Natural History), London, xviii + 424 pp., pls. viii + I - XXXVII and A - E under separate cover. https: // doi. org / 10.5962 / bhl. title. 58067","Wilkerson, R. C., Linton, Y. - M. & Strickman, D. (2021) Mosquitoes of the world. Vols. 1 & 2. Johns Hopkins University Press, Baltimore, Maryland, 1332 pp. https: // doi. org / 10.1186 / s 13071 - 021 - 04848 - 6","Edwards, F. W. (1911 b) The African species of Culex and allied genera. Bulletin of Entomological Research, 2 (3), 241 - 268. https: // doi. org / 10.1017 / S 0007485300001450","Edwards, F. W. (1912 b) A synopsis of the species of African Culicidae, other than Anopheles. Bulletin of Entomological Research, 3 (1), 1 - 53. https: // doi. org / 10.1017 / S 0007485300001656","de Meillon, B. & Rebelo, A. (1941) Culicini (Diptera, Nematocera) from the Colony of Mocambique. Mocambique Documentario trimestral, 27, 69 - 77.","Jupp, P. G. (1996) Mosquitoes of southern Africa: Culicinae and Toxorhynchitinae. Ekogilde Publishers, Hartebeespoort, 156 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","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.","Theobald, F. V. (1913 a) Second report on the mosquitoes of the Transvaal. In: Union of South Africa, Department of Agriculture, Second Report of the Director of Veterinary Research. Cape Times Limited, Government Printers, Cape Town, pp. 315 - 342, 2 pls.","Edwards, F. W. (1915) New and little-known East African Culicidae. Bulletin of Entomological Research, 5 (4), 273 - 281. https: // doi. org / 10.1017 / S 0007485300030303","de Charmoy, d'E. (1908) On three new species of Culex collected during the anti-malarial campaign in Mauritius in 1908. Annals of Tropical Medicine and Parasitology, 2 (3), 257 - 264. https: // doi. org / 10.1080 / 00034983.1908.11684887","Ingram, A. & Macfie, J. W. S. (1919) The early stages of West African mosquitos - IV. Bulletin of Entomological Research, 10 (1), 59 - 69. https: // doi. org / 10.1017 / S 0007485300043868","Hopkins, G. H. E. (1952) Mosquitoes of the Ethiopian Region I. - Larval bionomics of mosquitoes and taxonomy of culicine larvae. 2 nd Edition. British Museum (Natural History), London, viii + 355 pp.","Mattingly, P. F. & Knight, K. L. (1956) The mosquitoes of Arabia I. Bulletin of the British Museum (Natural History) Entomology, 4 (3), 91 - 141.","Edwards, F. W. (1932 b) Mosquito notes. - XI. Bulletin of Entomological Research, 23 (4), 559 - 562. https: // doi. org / 10.1017 / S 0007485300004338"]}

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