Subgenus Ochlerotatus Lynch Arribálzaga, 1891
Ochlerotatus confirmatus Lynch Arribálzaga, 1891b [subjective synonym of Aedes scapularis (Rondani, 1848)].
Subfamily Culicinae, tribe Aedini, genus Aedes. Subgenus Ochlerotatus includes 194 extant species and two extinct (fossil) species. Subgenus abbreviation – Och.
It is extremely important to point out here that the informal internal classification of Ochlerotatus previously included herein and that outlined in the catalogue of Wilkerson et al. (2021: 1078), with the exception of the Buvirilia and Culicelsa Groups, is/was entirely wrong. Those classifications were not based on a review and amalgamation of changes made to the internal classifiction in numerous independent studies conducted throughout the taxonomic history of the group. A comprehensive review of taxonomic studies has revealed that Ochlerotatus is divided into 22 species groups, three of which are subdivided into subgroups and one that has a recognised species complex (Harbach, 2024). The species groups are listed below, but pages are only provided at this time for the Buvirilia and Culicelsa Groups. Of 130 species previously listed here as Species Group uncertain, only 18 are now currently assigned to a species group (in boldface in the list below). The placement of the other 112 species in the internal classification of Ochlerotatus can be found in Part VI, Species and Their Classifications, of Harbach (2024).
Subgenus Ochlerotatus is a large highly varied group of species that generally exhibit the following characteristics. Five characters (five female genitalic and one larval) that diagnose Ochlerotatus in the phylogenetic analyses of Reinert et al. (2009), based on features observed in 48 species currently included in the subgenus, are indicated by an asterisk (*).
ADULTS – Vertex largely with narrow decumbent scales, erect forked scales numerous; interocular space with or without scales; compound eyes more or less separated above antennal pedicels; antenna of females shorter to longer than proboscis, distinctly shorter than proboscis in males, flagellar whorls of males with numerous long setae normally directed dorsally and ventrally; maxillary palpus of females 0.15–0.40 length of proboscis, with 5 palpomeres, palpomere 5 minute, maxillary palpus of males as long or longer than proboscis, palpomeres 4 and 5 subequal, downturned and setose; scutal scales usually narrow; anterior and posterior acrostichal and dorsocentral setae normally present; scutellum with narrow scales only; paratergite usually (not always) with scales; pleural scaling and setation varied; lower mesepimeral setae present or absent; remigium of wing usually with conspicuous setae; alula with extensive marginal fringe of long piliform scales; scaling of tarsi varied; ungues varied, at least fore- and midungues usually toothed, fore- and midungues of males enlarged; abdominal scaling varied, especially tergum I and laterotergite; terga with numerous lateral setae and reduced scaling in males. FEMALE GENITALIA – Segment VIII usually completely retracted within segment VII; *tergum VIII longer than wide, *with long setae on proximal 0.4 of lateral margins; *upper vaginal sclerite absent; cercus usually long and narrow, *distal part narrowly rounded; 3 spermathecal capsules. MALE GENITALIA – Segment VIII usually narrowed proximally; ninth tergal lobes usually more or less distinct, with setae; gonocoxite usually elongate, with more or less strongly developed basal dorsomesal lobe, sometimes with apical dorsomesal lobe, mesal surface membranous from base to apex; gonostylus simple, with apical usually long spiniform gonostylar claw; claspette comprised of elongate stem with single apical blade-like, sickle-like or otherwise modified claspette filament; aedeagus simple; paraproct with strong usually simple apical spine; cercal setae present. LARVAE – Head broad; antenna short to quite long, usually distinctly spiculate; seta 1-A varied, usually inserted near mid-length of antenna; *seta 1-C spiniform; seta 4-C usually small, varied in position; setae 5,6-C caudad of antennal bases; seta 13-P absent; seta 12-I present; seta 6-I,II and seta 7-I strongly developed, seta 6-III–VI at least moderately developed; comb scales varied, in large patch or single row; siphon varied in length, pecten spines evenly spaced or distal ones more widely separated; seta 1-S inserted within or beyond pecten; saddle complete or incomplete, acus present or absent; seta 1-X usually inserted on saddle; seta 2-X multi-branched; ventral brush (seta 4-X) usually with at least 7 pairs of setae on grid and 2 or more precratal setae. PUPAE – Trumpet short, broad, tracheoid area indistinct; seta 8-CT widely separated from seta 9-CT; setae 2,3-I usually approximated, seta 2-III–VII varied in position; seta 9-VI small, similar to seta 9-II–V; paddle with or without marginal spicules; seta 1-Pa single or branched. See genus Aedes.
The monophyly of subgenus Ochlerotatus is doubtful. Many species of the subgenus cannot be placed in any of the currently recognised species groups. Consequently, the affinities of Ochlerotatus are very uncertain. The subgenus was recovered as the sister-group of a clade comprised of subgenus Acartomyia + (subgenus Jarnellius + (subgenus Halaedes + genus Opifex)) in the phylogeny generated in the study of Reinert et al. (2009) based on morphological data of all life stages. Ochlerotatus was not associated with other generic-level taxa in the phylogeny of Wilkerson et al. (2015). It was recovered as a polyphyletic group in the maximum likelihood phylogeny of Soghigian et al. (2017) based on molecular markers for 85 species.
The immature stages of most species of subgenus Ochlerotatus are found in various types of temporary fresh-water ground pools, but several species inhabit rock holes, natural and artificial container habitats, and brackish water pools. Females of many species viciously attack humans and are often very troublesome pests, particularly in northern temperate regions. The majority of species bite in the daytime, particularly during crepuscular periods, but some species also feed during the night.
A number of species of subgenus Ochlerotatus are known to harbour natural infections of arboviruses and microfilariae (see Wilkerson et al., 2015: Table 1).
Subgenus Ochlerotatus is a large and abundant group with species in all temperate, subtropical and tropical areas of the world. The greatest diversity of species occur in the Australian, Nearctic and Neotropical Regions. Comparatively few species are found in the Afrotropical and Oriental Regions.
Belkin, 1962 (morphology, bionomics, distribution); Reinert, 2000 (as subgenus of Ochlerotatus, morphology); Reinert, 2002 (as subgenus of Ochlerotatus, female genitalia); Reinert et al., 2008, 2009 (as genus, morphology, phylogeny); Rattanarithikul et al., 2010 (as genus, Thailand, keys, bionomics); Huang & Rueda, 2014 (Afrotropical Region, 3 species of subgenus uncertain, key to adults); Wilkerson et al., 2015 (phylogeny, classification); Soghigian et al., 2017 (phylogenetic relationships).
aboriginis Dyar, 1917
abserratus (Felt & Young, 1904)
aculeatus (Theobald, 1903)
aenigmaticus Cerqueira & Costa, 1946
akkeshiensis Tanaka, 1998
albifasciatus (Macquart, 1838)
albineus Séguy, 1923
aloponotum Dyar, 1917
amateuri Ortega & Zavortink, 2019 (in Ortega-Morales et al., 2019)
ambreensis Rodhain & Boutonnier, 1983
andersoni Edwards, 1926
angustivittatus Dyar & Knab, 1907
annulipes (Meigen, 1830)
antipodeus (Edwards, 1920)
arundinariae Kasper, 2020
asiaticus Edwards, 1926
atactavittatus Arnell, 1976
atlanticus Dyar & Knab, 1906
auratus Grabham, 1906
aurifer (Coquillett, 1903)
behningi Martini, 1926
bejaranoi Martinez, Carcavallo & Prosen, 1960
berlandi Séguy, 1921
bimaculatus (Coquillett, 1902)
biskraensis Brunhes, 1999
bogotanus Arnell, 1976
breedensis Muspratt, 1953
burjaticus (Kuchartshuk, 1973)
burpengaryensis (Theobald, 1905)
caballus (Theobald, 1912)
cacozelus Marks, 1963
calcariae Marks, 1957
calumnior Belkin, Heinemann & Page, 1970
campestris Dyar & Knab, 1907
camptorhynchus (Thomson, 1869)
cantans (Meigen, 1818)
cantator (Coquillett, 1903)
caspius (Pallas, 1771)
cataphylla Dyar, 1916
chelli (Edwards, 1915)
churchillensis Ellis & Brust, 1973
clelandi (Taylor, 1914)
clivis Lanzaro & Eldridge, 1992
coluzzii Rioux, Guilvard & Pasteur, 1998
comitatus Arnell, 1976
communis (de Geer, 1776)
condolescens Dyar & Knab, 1907
continentalis Dobrotworsky, 1960
crinifer (Theobald, 1903)
cunabulanus Edwards, 1924
cyprioides Danilov & Stupin, 1982
cyprius Ludlow, 1920
dahlae (Nielsen, 2009)
daisetsuzanus Tanaka, Mizusawa & Saugstad, 1979
decticus Howard, Dyar & Knab, 1917
deficiens Arnell, 1976
detritus (Haliday, 1833)
diantaeus Howard, Dyar & Knab, 1913
dorsalis (Meigen, 1830)
dufouri Hamon, 1953
duplex Martini, 1926
dupreei (Coquillett, 1904)
eidsvoldensis Mackerras, 1927
eucephalaeus Dyar, 1918
euedes Howard, Dyar & Knab, 1913
euiris Dyar, 1922
euplocamus Dyar & Knab, 1906
excrucians (Walker, 1856)
explorator Marks, 1964
fitchii (Felt & Young, 1904)
flavescens (Müller, 1764)
flavifrons (Skuse, 1889)
fulvus (Wiedemann, 1828)
grossbecki Dyar & Knab, 1906
gutzevichi Dubitsky & Deshevykh, 1978
gyirongensis Ma, 1982
hakusanensis Yamaguti & Tamaboko, 1954
hargreavesi (Edwards, 1920)
harrisoni Muspratt, 1953
hastatus Dyar, 1922
hesperonotius Marks, 1959
hexodontus Dyar, 1916
hodgkini Marks, 1959
hokkaidensis Tanaka, Mizusawa & Saugstad, 1979
hungaricus Mihályi, 1955
idahoensis (Theobald, 1903)
imperfectus Dobrotworsky, 1962
impiger (Walker, 1848)
implicatus Vockeroth, 1954
incomptus Arnell, 1976
increpitus Dyar, 1916
inexpectatus Bonne-Wepster, 1948
infirmatus Dyar & Knab, 1906
intermedius Danilov & Gornostaeva, 1987
intrudens Dyar, 1919
jacobinae Serafim & Davis, 1933
jorgi Carpintero & Leguizamón, 2000
juppi McIntosh, 1973
kasachstanicus Gutsevich, 1962
lasaensis Meng, 1962
lepidus Cerqueira & Paraense, 1945
leucomelas (Meigen, 1804)
linesi Marks, 1964
longifilamentus Su & Zhang, 1988
ludlowae (Blanchard, 1905)
luteifemur Edwards, 1926
macintoshi Marks, 1959
martineti Senevet, 1937
mcdonaldi Belkin, 1962
melanimon Dyar, 1924
meprai Martínez & Prosen, 1953
mercurator Dyar, 1920
milleri Dyar, 1922
mitchellae (Dyar, 1905)
montchadskyi Dubitsky, 1968
nevadensis Chapman & Barr, 1964
nigrinus (Eckstein, 1918)
nigripes (Zetterstedt, 1838)
nigrithorax (Macquart, 1847)
nigrocanus Martini, 1927
nigromaculis (Ludlow, 1906)
niphadopsis Dyar & Knab, 1918
nivalis Edwards, 1926
normanensis (Taylor, 1915)
nubilus (Theobald, 1903)
obturbator Dyar & Knab, 1907
oligopistus Dyar, 1918
pallens Ross, 1943
patersoni Shannon & del Ponte, 1928
pectinatus Arnell, 1976
pennai Antunes & Lane, 1938
perkinsi Marks, 1949
pertinax Grabham, 1906
phaecasiatus Marks, 1964
phaeonotus Arnell, 1976
pionips Dyar, 1919
procax (Skuse, 1889)
protolepis (Cockerell, 1916) (fossil species, 23.0–33.9 Mya, Cenozoic, Oligocene)
pseudonormanensis Marks, 1949
pulcritarsis (Rondani, 1872)
pullatus (Coquillett, 1904)
punctodes Dyar, 1922
punctor (Kirby, 1837)
purpuraceus Brug, 1932
purpureifemur Marks, 1959
purpuriventris Edwards, 1926
ratcliffei Marks, 1959
raymondi del Ponte, Castro & Garcia, 1951
rempeli Vockeroth, 1954
rhyacophilus da Costa Lima, 1933
riparioides Su & Zhang, 1987
riparius Dyar & Knab, 1907
sagax (Skuse, 1889)
sallumae González & Reyes, 2017 (in González et al., 2017)
sapiens Marks, 1964
scapularis (Rondani, 1848)
schizopinax Dyar, 1929
schtakelbergi Shingarev, 1928
scutellalbum Boshell-Manrique, 1939
sedaensis Lei, 1989
serafini (Szadziewski, 1998) (fossil species, 33.9–56.0 Mya, Cenozoic, Eocene)
sergievi Danilov, Markovich & Proskuryakova, 1978
serratus (Theobald, 1901)
shannoni Vargas & Downs, 1950
silvestris Dobrotworsky, 1961
simanini Gutsevich, 1966
sinkiangensis Hsiao, 1977
sollicitans (Walker, 1856)
spencerii (Theobald, 1901)
spilotus Marks, 1963
squamiger (Coquillett, 1902)
sticticus (Meigen, 1838)
stigmaticus Edwards, 1922
stimulans (Walker, 1848)
stramineus Dubitzky, 1970
stricklandi (Edwards, 1912)
subalbirostris Klein & Marks, 1960
surcoufi (Theobald, 1912) (in Surcouf, 1912)
synchytus Arnell, 1976
taeniorhynchus (Wiedemann, 1821)
tahoensis Dyar, 1916
thelcter Dyar, 1918
theobaldi (Taylor, 1914)
thibaulti Dyar & Knab, 1910
tormentor Dyar & Knab, 1906
tortilis (Theobald, 1903)
trivittatus (Coquillett, 1902)
turneri Marks, 1963
upatensis Anduze & Hecht, 1943
vansomerenae Mattingly, 1955 (in Mattingly & Brown, 1955)
ventrovittis Dyar, 1916
vigilax (Skuse, 1889)
vittiger (Skuse, 1889)
washinoi Lanzaro & Eldridge, 1992
Albifasciatus Group
Aculeatus Group
Breedensis Group
Burpengaryensis Group
Buvirilia Group (see).
Caballus Group
Cantans Group
Caspius Group
Communis Group
Culicada Group (see).
Cunabulanus Group
Flavifrons Group
Fulvus Group
Impiger Group
Perkinsi Group
Pullatus Group
Scapularis Group
Serratus Group
Stimulans Group
Stricklandi Group
Taeniorhynchus Group
Thibaulti Group