Genus Verrallina Theobald, 1903

Type species: 

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

Classification: 

Subfamily Culicinae, tribe Aedini. Verrallina includes 95 species divided between three subgenera: Harbachius (13 species), Neomacleaya (52 species) and Verrallina (30 species). Genus abbreviation – Ve.

Characteristics: 

The remarkable development of the male and female genitalia and the following combinations of morphological characters distinguish Verrallina from all other genera and subgenera of tribe Aedini. ADULTS - Decumbent scales of vertex all broad (infrequently with few narrow curved scales on margins of coronal suture); antenna1 pedicel with several very short fine setae and usually a few small broad scales mesally (scales not overlapping or forming a dense patch); maxillary palpus of male very short (0.09-0. 18 length of proboscis); scutum and scutellum with only narrow curved scales, paratergite bare (except Ve. indica); anterior and posterior acrostichal areas with several to numerous setae (Harbachius usually with only a few setae on posterior area); anterior and posterior dorsocentral areas with setae; mesopostnotum normally bare (2 species with 1,2 setae); postpronotum with or without narrow curved scales dorsally; prespiracular area bare; postspiracular setae present; wings with narrow dark scales (rarely with few pale scales at base of costa and/or remigium); alula with narrow to moderately broad scales on margin; tarsi are dark-scaled. FEMALE GENITALIA - Lower vaginal sclerite present; insula ill-defined and small in subgenera Neomacleaya and Verrallina, absent in subgenus Harbachius; 3 spermathecal capsules present; cerci more or less triangular, usually with numerous broad scales; tergum VIII and sternum VIII with numerous broad scales. MALE GENITLIA - Tergum IX without setae; gonocoxite with one or more well-developed lobes, fleshy projections or spiniforms dorsally and/or ventrally at apex (except in 2 species); gonostylar claw absent; paraprocts usually highly modified into short or long free projections or broad contiguous plates; cereal setae absent; phallosome complex, with opisthophallus and prosophallus; phallus with distal tergal filament. LARVAE - Seta 4-C short, multiple-branched, mesad and slightly cephalad of seta 6-C; seta 5-C caudad and slightly mesad of seta 6-C, laterad and caudad of seta 4-C; insertions of setae 5,6-C widely separated; seta 7-C usually laterad and slightly cephalad of seta 6-C, laterad and at same level as 6-C in a few species; setae 5-7-C stout, branched, approximately equal in length or seta 7-C slightly shorter than setae 5,6-C (6-C single in Ve. unca); setae 5,6-P single (double in Ve. nigrotarsis); seta 8-P long, thickened, usually single (occasionally double, rarely triple; siphon with acus, pecten with distal l-5 spines more widely spaced than others; saddle incomplete ventrally, without spines on posterior margin, acus absent; ventral brush (seta 4-X) usually with 10 setae on grid and 2 or 3 precratal setae. PUPAE - Setae 1-3-CT approximately equally developed; setae 1,3-IV branched (usually multiple-branched, rarely single); insertions of setae 2,3-I widely separated; seta 3-I branched, rarely single, short to moderately long; seta 3-II,III moderately long to long, single, seta 3-II rarely double; setae 6,9-VII both relatively short, weakly developed; seta 9-VII single, rarely double; seta 9-VIII single to 6-branched; paddle without a fringe of hair-like spicules; seta 1-Pa single, occasionally double. See Aedini.

Phylogenetic relationships: 

Verrallina was recovered as a strongly supported monophyletic group in a sister relationship with Aedes in a cladistic analysis of tribe Aedini conducted by Reinert et al. (2004). Subgenus Harbachius was recovered as the sister of subgenus Neomacleaya + subgenus Verrallina.  Different relationships were recovered in the much more comprehensive analysis of the tribe conducted by Reinert et al. (2009), where Verrallina was recovered in a clade comprised of Neomelaniconion + (Edwardsaedes + (Aedes + (Paraedes + Verrallina))) [also recovered in the phylogeny of Wilkerson et al. (2015)]; within Verrallina, subgenera Harbachius and Neomacleaya are recovered as monophyletic but subgenus Verrallina is paraphyletic with respect to Neomacleaya. Verrallina was recovered in a sister relationship with Edwardsaedes in the maximum likelihood phylogeny of Soghigian et al. (2017) based on molecular markers.

Bionomics and disease relations: 

Very little information is available for species of Verrallina. As far as is known, the immature stages are found in temporary ground pools. Some species seem to be adapted to breeding in jungle pools that are frequently flushed whereas others inhabit more open coastal pools that are not subjected to flushing. Verrallina cuccioi has only been collected in pools and tree holes at ground level in stream beds. Eggs are apparently laid in ribbon-like gelatinous strings attached to objects in locations subject to inundation by rain water. The females of several species readily attack humans, chiefly during the day and largely in shaded places or in the vicinity of forest. Verrallina carmenti and Ve. lineata are serious pests in the South Pacific. The latter seems to be a semi-domestic species that may have been dispersed by human agency.

Several species are known to attack humans, but no species of the genus appear to play a role in the transmission of pathogens that cause diseases in humans.

Distribution: 

Species of Verrallina primarily occur in the Oriental Region with extensions northward to Japan (Kyushu) and southeastward into the Caroline Islands, New Guinea, the southern Pacific islands and northeastern Australia.

Principal references: 

Belkin, 1962 (as subgenus of Aedes, South Pacific); Reinert, 1974 (as subgenus of Aedes, taxonomy); Reinert, 1984 (as subgenus of Aedes, review); Lee et al., 1987 (as subgenus of Aedes, Australasian Region, keys taxonomy, bionomics, distribution); Reinert, 1999 (taxonomy); Reinert, 2001 (female genitalia); Reinert et al, 2004, 2009 (morphology, phylogeny); Rattanarithikul et al., 2010 (Thailand, keys, bionomics); Wilkerson et al., 2015 (phylogeny); Soghigian et al., 2017 (phylogenetic relationships).

Subgenera: 
Nomina dubia

perdita Leicester, 1908

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