Subgenus Aedimorphus Theobald, 1903

Type species: 

Aedes domesticus (Theobald, 1901), original combination: Uranotaenia domestica.

Classification: 

Subfamily Culicinae, tribe Aedini, genus Aedes. Subgenus Aedimorphus includes 76 species. Subgenus abbreviation – Adm.

Characteristics: 

Species of subgenus Aedimorphus are distinguished from species of other generic-level taxa of Aedini by the following combinations of characters. Characters that diagnose Aedimorphus in the phylogenetic analyses of Reinert et al. (2009), based on features observed in 20 species that are currently included in the subgenus, are indicated by an asterisk (*).

ADULTS – Vertex of head with narrow decumbent scales medially, erect forked scales usually numerous, on vertex and occiput; eyes usually narrowly separated above antennal pedicels; antenna of females long, normally as long or longer than proboscis, shorter than proboscis in males with numerous flagellar setae directed mainly dorsally and ventrally; maxillary palpus of females short, about 0.15–0.30 length of proboscis, with 5 palpomeres, palpomere 5 vestigial if present; maxillary palpus of males usually longer than proboscis, with 5 palpomeres, palpomeres 4 and 5 usually thickened, setose and turned downward, rarely slender, bare and turned upward; proboscis usually distinctly longer than forefemur; scutum with narrow scales; acrostichal setae and dorsocentral setae well developed, numerous; scutellum with narrow, broad or both narrow and broad scales; paratergite with or without scales; antepronotum and postpronotum usually with narrow scales; upper proepisternum with numerous setae; pleural scaling varied, postspiracular area, subspiracular area and prealar area with or without scales (*upper prealar scales present); lower mesepimeral setae absent; hindtarsomeres entirely dark-scaled or with basal pale bands; both ungues of foreleg and midleg of females toothed, fore- and midungues of males enlarged, toothed, hindungues usually simple in both sexes but sometimes toothed; abdominal segment I usually extensively scaled, laterotergite with large scale-patch, terga of males with numerous lateral setae. FEMALE GENITALIA – Segment VIII largely or completely retracted into preceding segment; sternum VIII with deep posteromedian emargination separating broadly rounded lobes; cercus long, narrow, usually without scales; postgenital lobe with median apical indentation; insula tongue-like, with tuberculi on distal 0.25, each usually with minute seta; 3 spermathecal capsules present. MALE GENITALIA – Tergum VIII sometimes widened posteriorly; tergum IX usually narrowed medially, ninth tergal lobes more or less distinct, with or without setae; sternum IX usually with postmedian row of setae; gonocoxite without distinct apical and basal tergomesal lobes, mesal surface membranous from base to apex; gonostylus usually expanded distally, with 1 or more gonostylar claws (*most proximal gonostylar claw inserted some distance from apex); claspette represented by short process or setose plaque at base of gonocoxite; aedeagus comprised of 2 lateral plates, each with distolateral teeth; paraproct with blunt or rounded apex, *sternal arm present;  cercal setae usually absent, present in some African species. LARVAE – Extremely varied; antenna usually long, spiculate, seta 1-A strongly developed, usually multi-branched; seta 4-C small, multi-branched, inserted closer to seta 5-C than to seta 6-C; setae 5,6-C strongly developed, usually inserted close together on level or posterior to seta 7-C; thoracic and abdominal chaetotaxy not studied for most species; seta 12-I absent; seta 6-III branched; *setae 1,2-VIII inserted on common setal support plate; comb of segment VIII varied, comb scales few in single row or numerous in large patch, scales squamiform or spine-like; siphon usually long, siphon index ≥ 3.0, pecten varied, distal pecten spines usually simple, always more widely spaced than proximal spines; seta 1-S multi-branched, *length ≤ 0.40 width of siphon, inserted distal to pecten; saddle incomplete, acus present or absent; seta 1-X often single, single to 4-branched; seta 2-X usually multi-branched; ventral brush (seta 4-X) with 3 or 4 precratal setae and 8–14 setae on grid with lateral and transverse bars. PUPAE – Trumpet short, only slightly widened distally, tracheoid area present or absent, weakly developed if present; seta 8-C inserted cephalad to or at level of trumpet base; setae 10,11-C approximated, 10-C multi-branched, 11-C single; genital lobe of females usually prominent; paddle ovoid, marginal spicules usually small or inconspicuous; seta 1-Pa short, single to triple. See genus Aedes.

Phylogenetic relationships: 

The affinities of subgenus Aedimorphus, as currently defined, are indefinite and whether it is a monophyletic group or a polyphyletic assemblage of species is uncertain. Thirteen nominal taxa now in synonymy for species groups within Aedimorphus may require subgeneric recognition. Aedimorphus was recovered as the sister of a number of subgeneric taxa in the phylogenetic analysis of Aedini by Reinert et al. (2009) based on extensive morphological data recorded for 20 species of the subgenus. The sister group included subgenera divided between two clades, one comprised of Polyleptiomyia + (Bifidistylus + (Albuginosus + (Tewarius + (Christophersiomyia + (Huaedes + Leptosomatomyia))))) and the other comprised of Neomelaniconion + (Edwardsaedes + (Aedes + (Paraedes + Verrallina))). Three species of the subgenus were recovered in a paraphyletic series in a sister relationship to Edwardsaedes + (((Neomelaniconion + ((Aedes + (Paraedes + Verrallina))) in the phylogeny of Wilkerson et al. (2015). Aedimorphus was not recovered as a monophyletic group in the maximum likelihood phylogeny of Soghigian et al. (2017) based on seven molecular markers. A clade containing nine species of Aedimorphus also included two species of Catageiomyia and a species of Polyleptiomyia. A tenth species of Aedimorphus formed a dispaate lineage with a species of subgenus Neomelaniconion.

Bionomics and disease relations: 

The immature stages of species of Aedimorphus principally inhabit temporary and semi-permanent fresh-water ground pools, but have been found in a variety of habitats, including swamps, marshes, bogs, pits, wells, ground pools, puddles, flood pools, stream pools, seepage, rice fields, animal footprints, wheel tracks, rock pools, rock holes and artificial containers. Females of many species readily attack and feed on humans, and are sometimes serious pests. Females of other species prefer non-human hosts and only occasionally feed on humans. Some species feed in the daytime, chiefly outdoors towards dusk, and some feed mainly during the night.

A number of species of subgenus Aedimorphus have been implicated in the transmission of pathogens that cause diseases in humans and domestic animals. Aedes vexans has been found naturally infected with a number of arboviruses, including western equine encephalitis virus, eastern equine encephalitis virus, viruses of the California encephalitis group (see details and references in Reinert, 1973) and Japanese encephalitis virus (Burke & Leake, 1988). This species is involved in the transmission of Tahyna virus in Europe (see references in Becker et al., 2010), it is a vector of Dirofilaria immitis (dog heartworm) (Reinert, 1973) and is considered to be a potentially important bridge vector of West Nile virus (Turell et al., 2005). Other pathogens associated with various species of subgenus Aedimorphus include Batai, Bunyamwera, Middelburg, Rift Valley fever, Sindbis, Shokwe, Spondweni and Wesselsbron viruses, and microfilariae of Brugia malayi.

Distribution: 

Most species of subgenus Aedimorphus occur in the Afrotropical Region, some occur in the Oriental Region, a few occur in the Australasian Region and one, Ae. vexans, occurs in the Holarctic and Oriental Regions, Central America, South Africa and the Papuan area.

Principal references: 

Reinert, 1973 (revision, Southeast Asia); McIntosh, 1975 (southern Africa, taxonomy); Lee et al., 1982 (Australasian Region, taxonomy, literature, bionomics); Reinert, 2000 (female genitalia); Reinert et al., 2004 (morphology, phylogeny); Reinert et al., 2009 (as genus, morphology, phylogeny); Rattanarithikul et al., 2010 (as genus, Thailand, keys, bionomics); Wilkerson et al., 2015 (phylogeny, classification); Soghigian et al., 2017 (phylogenetic relationships).

Species: 
abnormalis (Theobald, 1909)
adenensis Edwards, 1941
africanus Theobald, 1909
albodorsalis Fontenille & Brunhes, 1985
alboscutellatus (Theobald, 1905)
alboventralis (Theobald, 1910)
angolae Ribeiro & Ramos, 1974
arabiensis (Patton, 1905)
argenteoscutellatus Carter & Wijesundara, 1948
bambiotai Geoffroy, 1987
bancoi Geoffroy, 1987
bevisi (Edwards, 1915)
bwamba van Someren, 1950
caecus (Theobald, 1901)
caliginosus (Graham, 1910)
centropunctatus (Theobald, 1913)
culicinus (Edwards, 1922)
cumminsii (Theobald, 1903)
dalzieli (Theobald, 1910)
davidi Basio, 1971
dentatus (Theobald, 1904)
domesticus (Theobald, 1901)
durbanensis (Theobald, 1903)
ebogoensis Rickenbach & Ferrara, 1965
eritreae (Lewis, 1942)
fowleri (de Charmoy, 1908)
gibbinsi Edwards, 1935
gouldi Reinert, 1972
hirsutus (Theobald, 1901)
holocinctus Edwards, 1941
jamesi (Edwards, 1914)
kabwachensis Edwards, 1941
karooensis Muspratt, 1961
leesoni Edwards, 1932
leptolabis Edwards, 1936
leucarthrius (Speiser, 1909)
longiseta Edwards, 1936
lowisii (Theobald, 1910)
mansouri Qutubuddin, 1959
masoalensis Fontenille & Brunhes, 1985
mathioti Fontenille & Brunhes, 1985
mattinglyi Hamon & Rickenbach, 1954
mediolineatus (Theobald, 1901)
mesostictus Harbach, 2018
natronius Edwards, 1932
nigricephalus (Theobald, 1901)
nigrostriatus (Barraud, 1927)
nipponii (Theobald, 1907)
nocturnus (Theobald, 1903)
oakleyi Stone, 1939
ochraceus (Theobald, 1901)
orbitae Edwards, 1922
ovazzai Hamon & Adam, 1959
pachyurus Edwards, 1936
pallidostriatus (Theobald, 1907)
pampangensis (Ludlow, 1905)
pipersalatus (Giles, 1902)
pubescens Edwards, 1925
punctifemoris (Ludlow, 1921)
quasiunivittatus (Theobald, 1901)
rickenbachi Hamon & Adam, 1959
semlikiensis van Someren, 1950
senyavinensis Knight & Hurlbut, 1949
stenoetrus (Theobald, 1907)
stenostylus Cornel, Kowo & Mayi, 2023 (in Kowo et al., 2023)
subdentatus Edwards, 1936
suzannae (Boussès, 2022) (in Boussès et al., 2022)
syntheticus Barraud, 1928
taeniorhynchoides (Christophers, 1911)
tauffliebi Rickenbach & Ferrara, 1965
tricholabis Edwards, 1941
trimaculatus (Theobald, 1905)
trukensis Bohart, 1957
vexans (Meigen, 1830)
wigglesworthi Edwards, 1941
Scratchpads developed and conceived by (alphabetical): Ed Baker, Katherine Bouton Alice Heaton Dimitris Koureas, Laurence Livermore, Dave Roberts, Simon Rycroft, Ben Scott, Vince Smith