Elpeytonius, REINERT, HARBACH & KITCHING, 2009

Reinert, John F., Harbach, Ralph E. & Kitching, Ian J., 2009, Phylogeny and classification of tribe Aedini (Diptera: Culicidae), Zoological Journal of the Linnean Society 157 (4), pp. 700-794 : 768-772

ELPEYTONIUS REINERT, HARBACH & KITCHING ,

GEN. NOV.

Type species: Ochlerotatus apicoannulatus Edwards, 1912 (nom. nov. for Aedimorphus alboannulatus Theobald, 1905 ).

Females

Head: Vertex with entire area covered with narrow, curved, decumbent scales or with few to several narrow scales on anterior, median area; occiput and vertex with numerous erect, forked scales; ocular line narrow, with narrow, pale scales; eyes above antennal pedicels touching or separated by diameter of 2 ocular facets; antennal pedicel with mesal surface bearing few short, slender setae, few small, broad, dark scales present or absent; clypeus bare; maxillary palpus dark-scaled; proboscis dark-scaled with narrow, complete or incomplete, pale-scaled band near midlength, proboscis longer than forefemur.

Thorax: Scutum covered with narrow, curved scales except bare prescutellar area; acrostichal (anterior and posterior) and dorsocentral (anterior and posterior) setae present; prescutellar area with 7–9 setae on each side; scutellum with broad, silvery scales on all lobes; paratergite with broad, pale scales; antepronota widely separated, with broad, silvery scales, several setae; postpronotum with narrow, curved, dark scales on upper area, posterior setae present; postspiracular area without scales, 3 or 4 setae present; scales absent on hypostigmal area, subspiracular area, lower proepisternum, lower and upper prealar areas and metameron; mesokatepisternum with small upper and small lower posterior patches of broad, pale scales, setae present; mesepimeron with small upper patch of pale scales, without lower setae.

Wing: Dark-scaled, with small pale-scaled patch at base of costa; upper calypter with several setae on margin; remigium with 1 or 2 short setae on dorsal surface distally; dorsal tertiary fringe scales dark.

Legs: Ante- and postprocoxal areas bare; hindfemur and hindtibia with pale scales at apex; hindtarsomeres 1–4 each dark-scaled with wide, apical, white-scaled band, tarsomere 5 entirely or nearly entirely white-scaled; fore- and midungues, equal, each with 1 tooth.

Abdomen: Tergum I with patch of broad, white scales on laterotergite; terga I– VI dark-scaled dorsally; segment VII laterally compressed.

Genitalia: Tergum VIII moderately pigmented, width greater than length, few scales on distal part; sternum VIII moderately pigmented, width greater than length, apex with moderate, median emargination separating small to moderate-sized, sublateral lobes, scales absent or few in number, seta 2-S inserted posterior to seta 1-S; tergum IX comprised of single, moderately pigmented sclerite, apex with small, median emargination separating small, rounded lobes, each with 2–4 short, slender setae; postgenital lobe moderately long, moderately wide, apex with moderate, median emargination, setae on distal part of ventral surface; upper vaginal sclerite moderately pigmented, small to moderate size; without lower vaginal sclerite; insula tongue-like, with 4–6 small tuberculi on distal area; cercus moderately long, moderately wide, apex broadly rounded, without scales; single large, spherical, spermathecal capsule.

Males

Head: Antenna with distal 2 flagellomeres disproportionally long, remainder of flagellomeres short with numerous long setae directed primarily dorsally and ventrally; maxillary palpus with 5 palpomeres, palpomeres 4 and 5 somewhat downturned, palpomeres 4 and 5 and distal part of 3 with several long setae lateroventrally.

Legs: Foreungues unequal, each with 1 tooth; midungues unequal, larger one simple, smaller one with 1 tooth; hindungues equal, both simple.

Abdomen: Terga with several moderately long setae laterally.

Genitalia: Tergum IX moderately pigmented and sclerotized, posterior margin with pair of moderately large, broadly rounded lobes each with 3–6 short, slender setae; gonocoxite moderately long, moderately wide, dorsal surface with several short and few moderately long, slender setae on mesal area, several long, stout setae on outer area and on lateral surface, broad scales on outer part of dorsal, lateral and ventral surfaces, mesal surface membranous; gonostylus attached at apex of gonocoxite, relatively long, approximately proximal 0.60 narrow, distal part broader with several minute, fine setae, terminal short, broad, leaf-like gonostylar claw attached to rounded apex, outer margin of approximately distal 0.30–0.40 with long, narrow, curved, finger-like lobe with minute seta near apex; aedeagus with 2 elongate, lateral sclerites each bearing few, elongate teeth on distal part, membrane-like dorsal flap covering lateral sclerites; proctiger moderately long, apex bluntly rounded, with 2 or 3 minute cercal setae; claspette developed as short, narrow, plaque bearing few short setae at base of gonocoxite; sternum IX moderately long, setae absent or with 3 or 4 short, slender setae on median, posterior area.

Pupae

Trumpet: Moderately long, narrow, darkly-pigmented; tracheoid area weakly developed at base.

Cephalothorax: Seta 1-CT with 3 branches, long but noticeably shorter than 3-CT; 5-CT longer than 4-CT; 7-CT longer than 6-CT; 11-CT single.

Abdomen: Seta 3-I very long, stout, single; 6-I longer than 7-I; 1- II multiple-branched, slender; 2- II, 3- II, III long, stout, single, 3- II inserted mesal or at same level anterior to 2- II; 6- II long, stout, single, longer than 3- II; 5- V longer than median, dorsal length of tergum VI; 9- VII branched, inserted anterior and lateral to and longer than 6- VII; 9- VIII with 6 or 7 stout, aciculate branches.

Paddle: Apical margin rounded; midrib extending to apex of paddle; without hair-like spicules on margins; seta 1-Pa short, with 2 or 3 branches.

Fourth-instar larvae

Head: Seta 1-C slender, single; 4-C short, with 3–9 very slender branches, inserted mesal and either slightly anterior or slightly posterior to 6-C; 5-C long, stout, with 7–10 aciculate branches, inserted posterior and mesal to 6,7-C; 6-C long, stout, with 4–6 aciculate branches, inserted close to 5-C; 7-C moderately long to long, stout, with 8–11 aciculate branches, inserted anterior and lateral to 6-C; 12-C inserted mesal to 13-C; 13-C with several relatively long branches, longer than 12-C; 14-C short, single or 2-forked; 19-C absent; antenna moderately long, narrow, with several spicules, seta 1-A with 3–6 branches.

Thorax: Setae 1–3-P not inserted on common setal support plate, 1-P> 2-P> 3-P length, 1,3-P branched, 2-P single; 5,7-P branched; 6-P single, longer than 5,7-P; 1,4-M and 1,2- T branched; 6- T single.

Abdomen: Seta 7-I long, stout, with 2 branches; 12-I absent; 1,5,8- II branched; 1- VII long; 1- VIII longer than 2- VIII; 2,4- VIII single; comb with numerous scales in patch; segment X with saddle incomplete ventrally, acus absent, seta 1-X single to 3-branched, inserted on saddle, 2-X with 3–5 moderately long branches, 3-X long, single, ventral brush with several, fan-like, multiple-branched setae attached to grid with both transverse and lateral bars, several shorter, branched, precratal setae.

Siphon: Acus present; pecten with several evenly spaced spines; seta 1-S with 2–5 branches, inserted distal to pecten.

Included species

Elpeytonius apicoannulatus and El. simulans (Newstead & Carter) .

Distribution

Central African Republic, Ghana, Liberia, Republic of Cameroon, Democratic Republic of Congo, Nigeria, Sierra Leone, Sudan and Uganda.

Bionomics

Immature stages of El. apicoannulatus have been collected from rot-holes in mango, pawpaw, cotton and various other trees, dracaenas, stumps of banana plants and cut stems of bamboos ( Evans, 1926) [we note that Hopkins (1936: 130, 1952: 170) apparently misinterpreted Evans’ (1926) comments on habitats of this species] and tree-holes ( Haddow et al., 1952, Hopkins, 1936, 1952). Elpeytonius simulans have been collected from bamboo stumps ( Kumm, 1931) and tree-holes ( Haddow et al., 1952; Hopkins, 1936, 1952). Females of El. simulans have been taken occasionally biting humans during the day in forests and plantations and rarely in forests during the night ( Haddow et al., 1952)

Discussion

Evans (1926) briefly described and partially illustrated the male genitalia and fourth-instar larva of El. apicoannulatus (description of larva was noted by Edwards, 1932: 167). Hopkins (1936, 1952) described and illustrated the fourth-instar larvae of El. apicoannulatus (utilized Evans’ illustration) and El. simulans . Apparently, the description and illustration of the fourth-instar larva of El. apicoannulatus was not included in any of the Culicidae catalogs and supplements starting with Stone, Knight & Starcke (1959) to the present. Edwards (1941) provided brief descriptions of the female, male and pupa of El. apicoannulatus and illustrated the male genitalia. He briefly described the female and male of El. simulans and illustrated the adult female and male genitalia. Pao & Knight (1970) described and illustrated the fourth-instar larval mouthparts of El. simulans . The above generic description of the pupae and fourthinstar larvae are based on specimens of El. simulans and the published partial descriptions and illustrations of both species. Additional descriptive information is provided in Appendix 1 for species included in the analysis.

Etymology

Elpeytonius is named in honour of Mr E. L. Peyton in recognition of his important contributions to the taxonomy and biology of Culicidae , for introducing the first author ( JFR) to the exciting world of mosquito biosystematics over 45 years ago, and for steering the interests of the second author ( REH) toward a career in mosquito taxonomy. The generic name is masculine, formed from his initials (E. L., which were indicated without corresponding names on his birth certificate), surname and the Latin suffix ‘ -ius ’. Recommended abbreviation = El.

OCHLEROTATUS SUBGENUS JUPPIUS REINERT, HARBACH & KITCHING , SUBGEN. NOV.

Type species: Grabhamia caballa Theobald, 1912 .

Females

Head: Vertex covered with narrow, curved, decumbent scales; occiput and vertex with numerous long, erect, forked scales; ocular line narrow, with narrow, curved, pale scales; eyes above antennal pedicels separated by diameter of approximately 4 ocular facets; antennal pedicel with numerous small, broad, non-overlapping scales and few short, slender setae on mesal surface; clypeus bare; maxillary palpus dark-scaled, with or without few pale scales intermixed; proboscis darkscaled, longer than forefemur.

Thorax: Scutum covered with narrow, curved pale and dark scales including most of median, prescutellar area; parascutellar area with 2–4 narrow, curved, pale scales; scutellum with narrow, curved scales on all lobes; acrostichal (anterior and posterior), dorsocentral (anterior and posterior) and prescutellar areas with numerous dark setae; paratergite with broad, pale scales; antepronota widely separated, with narrow, curved, pale scales, several setae; postpronotum with narrow, curved scales dorsally and few somewhat broader scales ventrally, several posterior setae; hypostigmal area with broad, pale scales; postspiracular area with numerous broad, pale scales, few setae; subspiracular area with numerous broad, pale scales; upper proepisternum with broad, pale scales, numerous setae, lower area bare; prealar area with broad, pale scales on upper and lower areas, several to numerous setae; mesokatepisternum with large upper and moderate lower posterior patches of broad, pale scales, several upper and numerous lower posterior setae; mesepimeron with large patch of broad, pale scales on upper area and extending over middle area, several upper setae, with 4 or 5 lower setae; metameron with several broad, pale scales.

Wing: Some veins with moderately broad scales, other veins with narrow scales, dark and pale scales intermixed on some or most veins ( Oc. caballus and Oc. chelli Edwards ) or dark-scaled ( Oc. juppi (McIntosh)) ; costa with pale-scaled patch at base; upper calypter with numerous setae on margin; alula with dark scales on posterior margin; dorsal tertiary fringe scales narrow, dark; remigium with 1 or 2 setae distally on dorsal surface.

Legs: Anteprocoxal membrane bare; postprocoxal membrane with broad, pale scales; femora with numerous pale scales intermixed with dark-scaled areas, apices pale-scaled; tibiae with numerous pale scales intermixed with dark scales; tarsi dark-scaled with basal, pale-scaled bands on tarsomeres 1 and 2, and tarsomere 3 of mid- and hindtarsi; fore-, mid- and hindungues equal, each with 1 tooth.

Abdomen: Tergum I with patch of broad, pale scales on laterotergite; terga II–VI with dorsobasal palescaled patch expanded as complete or incomplete median, longitudinal stripe; segment VII dorsoventrally flattened.

Genitalia: Intersegmental membrane between segments VII and VIII very long; tergum VIII moderately pigmented with small non-pigmented notches on lateral and apical margins, length greater than width, without scales (1 adventitious scale in Oc. juppi ), short setae on entire length, 2 or more long setae on lateral margins, apex flat with small, median emargination; sternum VIII moderately pigmented with narrow, median, non-pigmented strip, length greater than width, apical margin gently rounded, without scales (rarely with 1 adventitious scale), setae on nearly entire length, seta 2-S inserted slightly posterior to seta 1-S; tergum IX comprised of single moderately pigmented, moderately long sclerite, apex with shallow, median emargination separating rounded lobes, each bearing 4–8 short setae, 9–15 total setae; postgenital lobe moderately wide, dorsal length short, apex flat or with very shallow, median emargination, few setae on distal area; without upper and lower vaginal sclerites; insula lip-like, with 2 moderately long, slender setae in lateral patches, 4 total setae; cercus moderately pigmented, long, narrow, apex narrowly rounded, without scales (occasionally 1 adventitious scale in Oc. juppi ), numerous setae covering most of dorsal surface; 3 spermathecal capsules, spherical, 1 large and 2 slightly smaller.

Males

Head: Antennae with distal 2 flagellomeres disproportionally long, remainder of flagellomeres short with numerous long setae directed primarily dorsally and ventrally; maxillary palpus with 5 palpomeres, longer than proboscis, with palpomeres 4 and 5 downturned, palpomeres 4 and 5 and distal part of 3 with numerous long setae lateroventrally.

Legs: Fore- and midungues unequal, larger unguis with 2 teeth, smaller unguis with 1 tooth; hindungues equal, each with 1 tooth.

Abdomen: Terga with numerous long, curved setae laterally; tergum VIII with posterior margin flat.

Genitalia: Tergum IX moderately pigmented, posterior margin with darkly pigmented, short, narrow lobe on each side of midline bearing few short, stout, somewhat flattened setae and few short, slender setae; gonocoxite heavily pigmented, moderately long, moderately wide, dorsal surface with several short, slender setae on most of area and few long, stout setae on outer area, mesal area with small, apical lobe bearing few short, slender setae and small, basal lobe bearing few short, slender and 1 or 2 moderately long, slender setae, lateral surface with several long, stout setae, ventral surface with few moderately long, slender setae on proximal area and several long, stout setae on distal area, several broad scales on lateral and ventral surfaces, mesal surface membranous; gonostylus attached at apex of gonocoxite, long, approximately 0.70 length of gonocoxite, relatively narrow throughout length but median part slightly wider than proximal part, distal part curved mesally and narrower than proximal part, with 2 or 3 short, slender setae, gonostylar claw short, slender, apex truncate, attached at apex of gonostylus; aedeagus tube-like, lateral margins bowed outward, widest on middle 0.33, apex bluntly pointed; phallosome with basal piece short; proctiger relatively short, distal part darkly pigmented, with curved, pointed apex, 5–8 minute, cercal setae; claspette comprised of single, relatively short, slender stem bearing one, long, flattened, relatively narrow, claspette filament at apex, with distal part slightly curved; sternum IX moderately pigmented, moderately long, with several short and moderately long setae on median, posterior area.

Pupae

Trumpet: Relatively short; moderately wide distally; pinna moderately long; tracheoid area weakly developed at base.

Cephalothorax: Seta 1-CT longer than 3-CT, both branched; 4,5-CT similarly developed, branched; 6-CT much shorter than 7-CT; 11-CT single, longer than 10,12-CT.

Abdomen: Seta 6-I single or 2-branched, longer than 3,7-I; 1-II with few to several slender branches; 3-II moderately long, single (3-II,III with 3 branches in Oc. juppi , fig. 3 of McIntosh, 1973); 6-II single, longer than 3-II; 3-III single, noticeably longer than 5-III; 1-IV–VI long, moderately stout, with 2 or 3 branches; 5-IV,V with 2 very long branches, noticeably longer than median, dorsal length of following tergum; 6-VII shorter than and inserted posterior and slightly mesal to 9-VII; 9-VIII with 10–12 stout, lightly aciculate branches.

Paddle: Apical margin more or less flat, with minute spicules; without hair-like spicules on margins; midrib extends to apex of paddle; seta 1-Pa short, single (occasionally 2-branched).

Fourth-instar larvae

Head: Seta 1-C single, distal part attenuate; 4-C short, very slender, single or 2-branched, inserted mesal and in line with 6-C; 5-C long, stout, single, inserted posterior and mesal to 6,7-C; 6-C long, stout, single, inserted mesal and slightly posterior to 7-C; 7-C moderately long, moderately stout, with 4–7 aciculate branches; 12-C with 4 or 5 branches, inserted mesal to 13-C; 13-C single, longer than 12-C; 14-C single; 19-C absent; antenna relatively short, moderately pigmented, with few scattered, minute spicules, seta 1-A short, single or with 2 or 3 branches, tips not reaching apex of antenna.

Thorax: Setae 1–3-P not inserted on common setal support plate, 1-P> 2-P> 3-P length, 1,2-P single, 3-P short, with 2 or 3 branches; 4-P short, single or with 2 branches; 5-P moderately stout, long, single, longer than 6-P; 6-P long, normally 2-branched (rarely single); 7-P long, with 3 branches; 4-M branched; 2-T with 3 branches; 6-T single.

Abdomen: Seta 7-I with 2 or 3 slender branches, approximately 0.60 length of 6-I; 12-I present; 6-II with 2 branches, longer than 6-III; 8-II with 2 or 3 branches; 6-III,IV moderately long, moderately stout, with 2 branches; 1-VII short, with 2 slender branches, approximately 0.40 dorsal length of segment X; 12-VII single; 1-VIII with 5–7 branches; 2,4-VIII single; comb with several thorn-like scales in short curved row; segment X with saddle incomplete ventrally, acus present, seta 1-S short, single, inserted on saddle, 2-X moderately long, with several branches, 3-X long, single, ventral brush with numerous long, branched setae inserted on grid with well developed transverse and lateral bars, with 2 (rarely 3) shorter, branched, precratal setae.

Siphon: Relatively short; acus present; pecten with several evenly spaced spines on approximately proximal 0.45 of siphon, distal 1 or 2 spines longer; seta 1-S with 4–6 short branches, inserted distal to pecten.

Included species

Ochlerotatus caballus , Oc. chelli and Oc. juppi .

Distribution

Aden, Eritrea, Ethiopia, Iran, Kenya, Lesotho, Namibia, Republic of South Africa, Sudan and Zimbabwe.

Bionomics

Hopkins (1936, 1952) provided a summary of the immature habitats of Oc. caballus to include a rockpool, a stream, water-furrows and small or mediumsized depressions in the veldt, which were filled periodically by rain or irrigation water. The habitats usually contained vegetation. McIntosh (1973) reported the larvae of Oc. juppi occur in temporary ground pools in grassland. McIntosh (1973) indicated the females of Oc. juppi are highly anthropophilic and readily feed on larger domesticated animals and possibly also, to some extent, on birds, during the daytime but with peak feeding in the early part of the night. Ochlerotatus caballus , like the previous species, readily feed on humans and larger domesticated animals.

Discussion

Subgenus Juppius includes medium-sized mosquitoes. Confusion concerning the identity of Oc. caballus and Oc. chelli occurred until McIntosh (1973) evaluated the species included in this group. See McIntosh (1973) for descriptions, partial illustrations and a discussion of the species. Hopkins (1952) described the fourth-instar larva of Oc. caballus and Jupp (1996) provided partial illustrations of the adults of Oc. caballus and Oc. juppi and the male genitalia of the latter species. The adult habitus of the primarily African Juppius species share a number of characters with those of the Australian subgenus Lepidokeneon , however notable differences exist in adult characters, especially in the male genitalia, and fourth-instar larvae. Additional descriptive information is provided in Appendix 1 for species included in the analysis.

Etymology

Juppius is named in honour of Dr Peter G. Jupp in recognition of his important contributions to the taxonomy of mosquitoes of southern Africa and studies on mosquito-vectored pathogens. The subgeneric name is masculine, formed from his surname and the masculine Latin suffix ‘ -ius ’. Recommended subgeneric abbreviation = Jup.