Subgenus Verrallina Theobald, 1903

Type species: 

Verrallina butleri (Theobald, 1901), original combination: Aedes butleri.


Subfamily Culicinae, tribe Aedini, genus Verrallina. Subgenus Verrallina includes 30 species. Subgenus abbreviation – Ver.


The following combinations of characters distinguish species of subgenus Verrallina from those of Harbachius and Neomacleaya. ADULTS ‒ Eyes contiguous dorsally; mesepimeron with fine hair-like setae posterior and/or ventral to scale-patch; hindungues of both females and males simple (except male of Ve. prioekanensis with tooth on one unguis); FEMALE GENITALIA ‒ Tergum VIII index 0.44‒0.70, largely covered with broad scales; sternum VIII with numerous setae on caudal lobes, largely with broad scales; postgenital lobe with broadly rounded, truncate or shallowly emarginate apex; upper vaginal lip not produced into vertical shield; lower vaginal sclerite small, rather poorly developed (except Butleri Series), separated from lower vaginal lip; insula ill-defined, small, with few minute tuberculi; spermathecal eminence consisting of a pair of comma-shaped plates, without spicules (except Butleri Series) and cephalad-produced pouch. MALE GENITALIA ‒ Gonocoxite with broad basotergal apodeme; prosophallus consisting of a pair of separated sclerites each developed apically into series of overlapping leaf-like plates, more or less as long as phallus (Ve. lugubris with single leaf-like plate); phallus comprised of a pair of aedeagal sclerites joined near mid-length by narrow sternal bridge, apices of sclerites contiguous, angled outward giving phallus an A-shaped appearance; opisthophallus with caudal margin gently concave mesally; paramere noticeably shorter than phallus. LARVAE ‒ Seta 1-A with 3‒9 branches (usually 3‒6); seta 7-C with 5‒14 branches; seta 8-C single (Carmenti Series) or branched (Butleri Series); seta 13-C usually single (with 2‒4 branches in Ve. butleri, Ve. iriomotensis and Ve. lugubris of the Butleri Series); seta 7-I long, single; comb scales rounded apically, denticles usually uniformly developed (Ve. dux and Ve. iriomotensis with longer median apical denticle); seta 2-X usually double or triple (with 4‒8 branches in a few species). PUPAE – Cephalothorax with or without poorly developed ocular facets of compound eye; seta 2-IV‒VI (in most species) somewhat thicker and inserted more cephalad than seta 4-IV; seta 9-VIII usually with 3‒6 branches (except some species of the Butleri Series). See genus Verrallina.

Phylogenetic relationships: 

Subgenus Verrallina was recovered in a paraphyletic relationship to Neomacleaya in the morphology-based phylogenetic analysis of Aedini conducted by Reinert et al. (2009), and the clade comprised of species of these subgenera was the sister of Harbachius. In agreement with the earlier study of Reinert et al. (2004), Verrallina + Neomacleaya was recovered as the sister of Harbachius in the maximum likelihood phylogeny of Soghigian et al. (2017) based on molecular markers.

Bionomics and disease relations: 

Species of subgenus Verrallina typically occur in ground-water habitats in forested areas. Common larval habitats include temporary ground pools, puddles, hoof prints, ditches, marshy depressions, swamp pools, road ruts, rock holes, ponds, pot holes in streambeds and wallows. In additional to some of these habitats, larvae of Ve. butleri occur in brackish tidal pools and mangrove swamps in coastal areas, larvae of Ve. parasimilis have been found in crayfish and crab holes in forest, and larvae of Ve. lineata have once been found in an old kettle. Females of at least 13 species readily bite humans who enter their realms during the daytime. Verrallina lineata and Ve. reesi have been captured biting indoors and outdoors, Ve. carmenti sometimes enters houses in the evening and Ve. lineata and Ve. funerea are known to bite at night as well as during the day.

Dirofilaria and Japanese encephalitis, Bebaru and Getah viruses have been isolated from Ve. butleri in Malaysia, but none of the other species of subgenus Verrallina are known or suspected of being of medical importance.


Species of subgenus Verrallina occur principally in the Australasian Region, with some minor extensions into the Oriental Region. Three species, Ve. butleri, Ve. dux and Ve. lugubris, are exclusively and widely distributed in the Oriental Region. Verrallina iriomotensis is only known from localities in the Ryukyu Islands of Japan.

Principal references: 

Reinert, 1974 (several species, as species of Aedes subgenus of Verrallina, taxonomy, bionomics and distribution); Reinert, 1984 (several species, as subgenus of Aedes, species descriptions, keys, distributions, bionomics); Lee et al., 1987 (Australasian species, as species of Aedes subgenus Verrallina, keys, taxonomy, bionomics and distribution); Reinert, 1999 (subgenus and type species descriptions, keys); Reinert, 2001 (female genitalia); Reinert et al., 2004, 2009 (morphology, phylogeny); Rattanarithikul et al., 2010 (Thailand, keys, bionomics); Soghigian et al., 2017 (phylogenetic relationships).


azureosquamata (Bonne-Wepster, 1948)
bancrofti (Taylor, 1914)
bifoliata (King & Hoogstraal, 1947)
butleri (Theobald, 1901)
carmenti (Edwards, 1924)
cuccioi (Belkin, 1962)
dux (Dyar & Shannon, 1925)
embiensis (Huang, 1968)
foliformis (King & Hoogstraal, 1947)
funerea (Theobald, 1903)
iriomotensis (Tanaka & Mizusawa, 1973)
killertonis (Huang, 1968)
leilae (King & Hoogstraal, 1947)
lineata (Taylor, 1914)
lugubris (Barraud, 1928)
mccormicki (Belkin, 1962)
milnensis (King & Hoogstraal, 1947)
multifolium (King & Hoogstraal, 1947)
obsoleta (Huang, 1968)
parasimilis (King & Hoogstraal, 1947)
pipkini (Bohart, 1957)
quadrifolium (Brug, 1934)
quadrispinata (King & Hoogstraal, 1947)
reesi (King & Hoogstraal, 1947)
sentania (King & Hoogstraal, 1947)
similis (Theobald, 1910)
simpla (King & Hoogstraal, 1947)
trispinata (King & Hoogstraal, 1947)
vanapa (Huang, 1968)
variabilis (Huang, 1968)

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