Subgenus Ochlerotatus Lynch Arribálzaga, 1891

Type species: 

Ochlerotatus confirmatus Lynch Arribálzaga, 1891b [subjective synonym of Aedes scapularis (Rondani, 1848)].

Classification: 

Subfamily Culicinae, tribe Aedini, genus Aedes. Subgenus Ochlerotatus includes 207 species. Thirteen species groups are recognised for 76 species. Subgenus abbreviation – Och.

Characteristics: 

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 Aedes.

Phylogenetic relationships: 

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.

Bionomics and disease relations: 

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).

Distribution: 

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.

Principal references: 

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).

Species - Group uncertain: 

aboriginis Dyar, 1917
abserratus (Felt & Young, 1904)
akkeshiensis Tanaka, 1998
albifasciatus (Macquart, 1838)
albineus Séguy, 1923
aloponotum Dyar, 1917
ambreensis Rodhain & Boutonnier, 1983
andersoni Edwards, 1926
annulipes (Meigen, 1830)
antipodeus (Edwards, 1920
arundinariae Kasper, 2020
asiaticus Edwards, 1926
aurifer (Coquillett, 1903)
behningi Martini, 1926
bejaranoi Martinez, Carcavallo & Prosen, 1960
berlandi Séguy, 1921
biskraensis Brunhes, 1999
breedensis Muspratt, 1953
burjaticus (Kuchartshuk, 1973)
burpengaryensis (Theobald, 1905)
cacozelus Marks, 1963
calumnior Belkin, Heinemann & Page, 1970
campestris Dyar & Knab, 1907
camptorhynchus (Thomson, 1869)
cantans (Meigen, 1818)
cantator (Coquillett, 1903)
caspius (Pallas, 1771)
cataphylla Dyar, 1916
churchillensis Ellis & Brust, 1973
clivis Lanzaro & Eldridge, 1992
coluzzii Rioux, Guilvard & Pasteur, 1998
communis (de Geer, 1776)
continentalis Dobrotworsky, 1960
cunabulanus Edwards, 1924
cyprioides Danilov & Stupin, 1982
cyprius Ludlow, 1920
dahlae (Nielsen, 2009
daisetsuzanus Tanaka, Mizusawa & Saugstad, 1979
decticus Howard, Dyar & Knab, 1917
detritus (Haliday, 1833)
dorsalis (Meigen, 1830)
dufouri Hamon, 1953
duplex Martini, 1926
eidsvoldensis Mackerras, 1927
euedes Howard, Dyar & Knab, 1913
euiris Dyar, 1922
excrucians (Walker, 1856)
explorator Marks, 1964
fitchii (Felt & Young, 1904)
flavescens (Müller, 1764)
grossbecki Dyar & Knab, 1906
gutzevichi Dubitsky & Deshevykh, 1978
gyirongensis Ma, 1982
hakusanensis Yamaguti & Tamaboko, 1954
hargreavesi (Edwards, 1920)
harrisoni Muspratt, 1953
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, 1954a
increpitus Dyar, 1916
inexpectatus Bonne-Wepster, 1948
intermedius Danilov & Gornostaeva, 1987
jacobinae Serafim & Davis, 1933
kasachstanicus Gutsevich, 1962
lasaensis Meng, 1962
lepidus Cerqueira & Paraense, 1945
leucomelas (Meigen, 1804)
linesi Marks, 1964
longifilamentus Su & Zhang, 1988
luteifemur Edwards, 1926
macintoshi Marks, 1959
martineti Senevet, 1937
melanimon Dyar, 1924
mercurator Dyar, 1920
milleri Dyar, 1922
montchadskyi Dubitsky, 1968
nevadensis Chapman & Barr, 1964
nigrinus (Eckstein, 1918)
nigripes (Zetterstedt, 1838)
nigrithorax (Macquart, 1847)
nigrocanus Martini, 1927
niphadopsis Dyar & Knab, 1918
nivalis Edwards, 1926
normanensis (Taylor, 1915)
perkinsi Marks, 1949
phaecasiatus Marks, 1964
pionips Dyar, 1919
procax (Skuse, 1889)
pseudonormanensis Marks, 1949
pulcritarsis (Rondani, 1872)
pullatus (Coquillett, 1904)
punctodes Dyar, 1922
punctor (Kirby, 1837)
purpureifemur Marks, 1959
ratcliffei Marks, 1959
rempeli Vockeroth, 1954
riparioides Su & Zhang, 1987
riparius Dyar & Knab, 1907
sagax (Skuse, 1889)
sallumae González & Reyes, 2017 (in González et al., 2017)
sapiens Marks, 1964
schizopinax Dyar, 1929
schtakelbergi Shingarev, 1928
scutellalbum Boshell-Manrique, 1939
sedaensis Lei, 1989
sergievi Danilov, Markovich & Proskuryakova, 1978
shannoni Vargas & Downs, 1950
silvestris Dobrotworsky, 1961
simanini Gutsevich, 1966
sinkiangensis Hsiao, 1977
spencerii (Theobald, 1901)
squamiger (Coquillett, 1902)
sticticus (Meigen, 1838)
stimulans (Walker, 1848)
stramineus Dubitzky, 1970
subalbirostris Klein & Marks, 1960
surcoufi (Theobald, 1912) (in Surcouf, 1912)
tahoensis Dyar, 1916
theobaldi (Taylor, 1914)
thibaulti Dyar & Knab, 1910
upatensis Anduze & Hecht, 1943
ventrovittis Dyar, 1916
vittiger (Skuse, 1889)
washinoi Lanzaro & Eldridge, 1992

Scratchpads developed and conceived by (alphabetical): Ed Baker, Katherine Bouton Alice Heaton Dimitris Koureas, Laurence Livermore, Dave Roberts, Simon Rycroft, Ben Scott, Vince Smith