Genus Aedimorphus Theobald, 1903

Type species: 

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

Classification: 

Subfamily Culicinae, tribe Aedini. Aedimorphus includes 66 species, seven of which have recognised subspecies. Genus abbreviation – Am.

Tanaka (2014, 2018), in a catalogue of the mosquitoes of Japan, listed Ecculex and Lepidotomyia (synonyms of Aedimorphus) as subgenera of Aedimorphus. The former included Ad. vexans nipponii and Ad. vexans nocturnus, listed without explanation, description or formal change of status as a species rather than a subspecies, and the latter included Ad. alboscutellatus. The ‘subgenera’ were not accompanied by a description or diagnosis and were listed without an explicit indication that they were new. In the absence of this, despite being included in a dichotomous key based on features of the male genitalia, they cannot be accepted as formally (validly) established subgeneric taxa (or even species groups). It should be noted that Ad. vexans was recovered in a basal relationship to 17 other species of the genus, with low support, and Ad. alboscutellatus formed a weakly supported sister pair with Ad. culicinus in the morphology-based phylogenetic study of Reinert et al. (2009). Those results do not provide evidence for the recognition of subgeneric divisions within Aedimorphus.

Characteristics: 

Species of Aedimorphus are distinguished from species of other genera 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 genus, 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 Aedini.

Phylogenetic relationships: 

The affinities of Aedimorphus, as currently defined, are indefinite and whether the genus 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 generic-level recognition. Aedimorphus was recovered as the sister of a number of genera in the phylogenetic analysis of Aedini by Reinert et al. (2009) based on extensive morphological data recorded for 20 species of the genus. The sister group included genera 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 Aedimorphus were recovered in a paraphyletic series in a sister relationship to Edwardaedes + (((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 Neomelaniconion.

Bionomics and disease relations: 

The immature stages 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 Aedimorphus have been implicated in the transmission of pathogens that cause diseases in humans and domestic animals. Aedimorphus 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 Aedimorphus include Batai, Bunyamwera, Middelburg, Rift Valley fever, Sindbis, Shokwe, Spondweni and Wesselsbron viruses, and microfilariae of Brugia malayi.

Distribution: 

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

Principal references: 

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

Species: 
abnormalis (Theobald, 1909)
   subspecies abnormalis (Theobald, 1909)
   subspecies kabwachensis (Edwards, 1941)
aerarius (McIntosh, 1975)
africanus Theobald, 1909
albodorsalis (Fontenille & Brunhes, 1985)
alboscutellatus (Theobald, 1905)
alboventralis (Theobald, 1910)
argenteoscutellatus (Carter & Wijesundara, 1948)
bambiotai (Geoffroy, 1987)
bancoi (Geoffroy, 1987)
bevisi (Edwards, 1915)
caecus (Theobald, 1901)
caliginosus (Graham, 1910)
centropunctatus (Theobald, 1913)
culicinus (Edwards, 1922)
cumminsii (Theobald, 1903)
   subspecies cumminsii (Theobald, 1903)
   subspecies mediopunctata (Theobald, 1909)
dalzieli (Theobald, 1910)
davidi (Basio, 1971)
dentatus (Theobald, 1904)
domesticus (Theobald, 1901)
durbanensis (Theobald, 1903)
   subspecies angolae (Ribeiro & Ramos, 1974)
   subspecies durbanensis (Theobald, 1903)
ebogoensis (Rickenbach & Ferrara, 1965)
eritreae (Lewis, 1942)
fisheri Barraud, 1928c
fowleri (de Charmoy, 1908)
gibbinsi (Edwards, 1935)
gouldi (Reinert, 1972)
hirsutus (Theobald, 1901)
   subspecies adenensis (Edwards, 1941)
   subspecies hirsutus (Theobald, 1901)
holocinctus (Edwards, 1941)
jamesi (Edwards, 1914)
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)
natronius (Edwards, 1932)
nigricephalus (Theobald, 1901)
nigrostriatus Barraud, 1927
oakleyi (Stone, 1939)
ochraceus (Theobald, 1901)
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)
subdentatus (Edwards, 1936)
taeniatus (Leicester, 1908)
taeniorhynchoides (Christophers, 1911)
tauffliebi (Rickenbach & Ferrara, 1965)
tricholabis (Edwards, 1941)
   subspecies bwamba (van Someren, 1950)
   subspecies tricholabis (Edwards, 1941)
trimaculatus (Theobald, 1905)
trukensis (Bohart, 1957)
vexans (Meigen, 1830)
   subspecies arabiensis (Patton, 1905)
   subspecies nipponii (Theobald, 1907)
   subspecies 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