Revision of Subgenus Neomacleaya Theobald, 1907 from Mon, 2022-05-23 13:54

Type species: 

Verrallina indica (Theobald, 1907), original combination: Neomacleaya indica.


Subfamily Culicinae, tribe Aedini, genus Verrallina. Subgenus Neomacleaya includes 52 species. Subgenus abbreviation – Nma.


The following combinations of characters distinguish species of Neomacleaya from those of subgenera Harbachius and Verrallina. ADULTS ‒ Eyes contiguous or nearly contiguous dorsally; mesepimeron with fine hair-like setae posterior and/or ventral to scale-patch; hindungues of both females and males simple. FEMALE GENITALIA ‒ Tergum VIII short, wider than long, index about 0.35-0.57, nearly entirely covered with broad scales; sternum VIII with numerous thin setae on apical lobes, largely covered with broad scales; postgenital lobe relatively narrow, with deep median caudal indentation; upper vaginal lip with median posterior area caudad of spermathecal eminence developed into large vertical shield; lower vaginal sclerite large, not contiguous with lower vaginal lip, usually spiculate; insula ill-defined, small, with few minute tuberculi; spermathecal eminence complex, with numerous well-developed spicules (less well-developed in Ve. incerta, Ve. panayensis and Ve. rara), without cephalad pouch. MALE GENITALIA ‒ Gonocoxite with moderately broad basotergal apodeme; prosophallus consisting of pair of sclerites separated apically, about as long as phallus; phallus comprised of a pair of aedeagal sclerites joined mesally by sternal bridge, each sclerite with narrow projection basolaterally extending at approximately right angle; opisthophallus short, caudal margin truncate or concave; paramere about as long as phallus. LARVAE ‒ Seta 1-A with 3–6 branches (rarely double); seta 7-C with 5–17 branches (usually 7‒14); seta 8-C branched; seta 13-C short, with 2–6 branches (usually 3–5); seta 7-I long, double or triple; comb scales with stout median apical spine and smaller lateral denticles; seta 2-X multi-branched (with 5 or more branches). PUPAE ‒ Cephalothorax with moderately developed ocular facets of compound eye; trumpet usually expanded apically; seta 2-IV‒VI usually thin, inserted more caudad than seta 4; seta 9-VIII usually single or double. See genus Verrallina.

Phylogenetic relationships: 

Neomacleaya was recovered as the sister of subgenus Verrallina in the morphology-based phylogenetic analysis of Aedini conducted by Reinert et al. (2004), with the two subgenera in a sister relationship to subgenus Harbachius. The latter sister relationship was recovered in the more comprehensive analysis of the tribe conducted by Reinert et al. (2009) in which Verrallina was paraphyletic relative to Neomacleaya. Interestingly, the subgeneric relationships recovered in the study of Reinert et al. (2004) were also recovered in the maximum likelihood phylogeny of Soghigian et al. (2017) based on molecular markers.

Bionomics and disease relations: 

Virtually nothing is known about the biology of at least 14 species of Neomacleaya. The immature stages of the other species are found principally in small temporary bodies of fresh water in forests and plantations, including puddles/ground pools, road ruts, flood pools, residual pools in streambeds, seepage pools and animal hoof prints. Verrallina atriisimilis, Ve. gibbosa and Ve. prioekanensis have been found in forest swamps, Ve. campylostylus in a tin can, Ve. clavata in crab holes, Ve. unca in axils of Colocasia and Ve. vallistris in a village well. Females have been collected biting cattle, horses and humans during the daylight hours, principally in forests and bamboo groves. The following species are known to bite humans: Ve. campylostylus, Ve. agrestis, Ve. atria, Ve. atriisimilis, Ve. cauta, Ve. clavata, Ve. cretata, Ve. johorensis, Ve. notabilis, Ve. panayensis, Ve. sabahensis, Ve. taeniata, Ve. unca, Ve. vallistris and Ve. varietas. It seems likely that all species of the subgenus may attack humans who enter their realms.

Species of subgenus Neomacleaya are not known to harbour or transmit pathogens of human disease.


Except for Ve. atriisimilis in Japan and Ve. neomacrodixoa and Ve. panayensis whose distributions extend south-eastward into the Australasian Region (the former reaches the Solomon Islands; both are recorded from Papua Island and Papua New Guinea), the species of subgenus Neomacleaya appear to be confined to the Oriental Region.

Principal references: 

Reinert, 1974 (several species, as species of Aedes subgenus Verrallina, taxonomy, bionomics and distribution); Reinert, 1984 (several species, as subgenus of Aedes, species descriptions, keys, distributions, bionomics); 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).


adusta (Laffoon, 1946)
agrestis (Barraud, 1931)
andamanensis (Edwards, 1922)
assamensis Bhattacharyya, Tewari, Prakash, Mohapatra & Mahanta, 2004
atria (Barraud, 1928)
atriisimilis (Tanaka & Mizusawa, 1973)
campylostylus (Laffoon, 1946)
cauta (Barraud, 1928)
ceylonica (Edwards, 1917)
clavata (Barraud, 1931)
comata (Barraud, 1931)
comosa (Reinert, 1974)
cretata (Delfinado, 1967)
cyrtolabis (Edwards, 1928)
gibbosa (Delfinado, 1967)
harrisonica (Reinert, 1974)
hispida (Delfinado, 1967)
indica (Theobald, 1907)
johnsoni (Laffoon, 1946)
johorensis (Reinert, 1974)
komponga (Klein, 1973)
latipennis (Delfinado, 1967)
leicesteri (Edwards, 1917)
macrodixoa (Dyar & Shannon, 1925)
margarsen (Dyar & Shannon, 1925)
neomacrodixoa (King & Hoogstraal, 1947)
nigrotarsis (Ludlow, 1908)
notabilis (Delfinado, 1967)
nubicola (Laffoon, 1946)
panayensis (Ludlow, 1914)
petroelephantus (Wijesundara, 1951)
philippinensis (Delfinado, 1968)
phnoma (Klein, 1973)
prioekanensis (Brug, 1931)
protuberans (Delfinado, 1967)
pseudodiurna (Theobald, 1910)
pseudomediofasciata (Theobald, 1910)
pseudovarietas (Reinert, 1974)
rami (Barraud, 1928)
rara (Delfinado, 1968)
sabahensis (Reinert, 1974)
seculata (Menon, 1950)
singularis (Leicester, 1908)
sohni (Reinert, 1974)
spermatheca (Wijesundara, 1951)
taeniata (Leicester, 1908)
torosa (Delfinado, 1967)
unca (Theobald, 190l)
vallistris (Barraud, 1928)
varietas (Leicester, 1908)
virilis Leicester, 1908
yerburyi (Edwards, 1917)

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