Leehelea Debenham

(Figs. 28A, 31B, 40C, 46C–D, 53A, 68C, 77B)

DIAGNOSIS: Only pupa of Ceratopogonidae with the abdominal tubercles all apically pointed (Fig. 68C), abdominal segment 4 with D-8-IV and D-9-IV on basally fused tubercles (Fig. 68C) and abdominal segment 8 with the two ventral sensilla (V-5-VIII, V-6-VIII) on a single tubercle and V-5-VIII tiny and V-6-VIII elongate; not diagnosable as different from Sphaeromias .

DESCRIPTION: Total length = 6.50–6.97 mm. Without larval exuviae retained on abdomen. Exuviae with flagellum appressed against lateral margin of midleg, wing (as in Figs. 16B, 33B). Ecdysial tear around base of antenna, along lateral margin of face to palpus (as in Figs. 17C, 79H). Head: Dorsal apotome (as in Fig. 22A), with ventral line of weakness, without dorsomedial tubercle, without central dome; dorsolateral cephalic sclerite (as in Fig. 13H) fused to scutum, each side separated medially by dorsal apotome in whole pupa; mouthparts (Fig. 28A) with mandible well-developed, lacinia absent; palpus extending equal to or just posterior to posterolateral margin of labium; labium entire (not divided medially); apex of antenna (Fig. 40C) anterior to posterior extent of midlength portion of midleg (portion lateral to mesosternum), narrowed posteriorly; sensilla: dorsal apotomals (as in Fig. 22A)—uncertain; dorsolateral cephalic sclerite sensilla—1 seta, 1 campaniform sensillum; clypeal-labrals (Fig. 28A)—2 slender setae; oculars (Fig. 28A)—2 setae, 1 campaniform sensillum. Thorax: Prothoracic extension (Fig. 28A) wide, well-developed but narrow dorsolaterally, not extending to antenna; mesonotum with very short tubercles, not extending posteromedially, not dividing metathorax medially (Fig. 53A); respiratory organ (Figs. 46C–D) length/width = 3.39–4.08, elongate, moderately slender, somewhat flattened apically, with pores closely abutting at apex of respiratory organ, arranged in single row, outer surface with some wrinkles, with short, wide pedicel, base with elongate posteromedial apodeme, membranous base of respiratory organ moderately elongate, annulated, tracheal tube straight to slightly curved along length, with spirals restricted to base, wrinkles to half length; wing (Fig. 40C) without apical tubercle or angle, separated medially by fore-, midlegs; halter apex and hind leg (similar to Fig. 33J) broadly abutting; halter apex extending posteriorly to 1/4 length of tergite 2; legs (Fig. 40C) with lateral margin of foreleg near midlength of wing slightly angled; hind leg visible at lateral margin of wing (as in Fig. 33I); female with apex of foreleg ventral to apex of midleg; apex of hind leg abutting apex of midleg laterally; sensilla: anteromedials—2 elongate setae (as in Figs. 31L–M); anterolaterals—1 moderately long seta; dorsal setae (Fig. 31B)—D-1-T, D-2-T, D-4-T, D-5-T setae, D-3-T campaniform sensillum, D-3-T anterolateral to D-4-T; supraalar 2—campaniform sensillum; metathoracics (Fig. 53A)—1 campaniform sensillum; M-3-T distant from margin of metathorax (at least 1/3 length of metathorax). Abdomen: with tergite 1 with 1 medial spot, tergites 2–7 with medial area with stripe, 2 anterolateral spots, 2 pairs on tergite 8, sternites 3–7 with medial stripe, anterolateral spot, 2 spots on sternite 8, segment 2 as wide or slightly wider than segment 3, segments with undivided, thin to thick setae, with pointed, short to moderately elongate tubercles, tergites or sternites entire, each without membranous disc; segment 9 (Fig. 77B) not strongly modified, terminal processes closely approximated basally, each projecting posterodorsolaterally, tapering to pointed apex; sensilla: tergite 1 (Fig. 53A) with 8 setae, 2 campaniform sensilla, including 3 lateral sensilla, D-2-I, D-3-I closely approximated, D-7-I situated anteriorly near D-3-I; segment 4 (Fig. 68C)—D-2-IV, D-3-IV short to moderately elongate setae on pointed tubercles; D-5-IV, D-8-IV, D-9-IV short to moderately elongate setae; D-5-IV on single tubercle, D-8-IV, D-9-IV on basally fused, closely approximated tubercles, posterior dorsal sensilla in transverse row, arranged medially to laterally: D-5-IV, D-4-IV, D-8-IV, D-9-IV; D-7-IV near D-3-IV; L-1-IV elongate seta from surface, moderately anterior of posterior lateral setae; L-2-IV, L-3-IV, L-4-IV short on pointed tubercles, V-5-IV, V-6-IV, V-7-IV short setae on short tubercles, all closely approximated, V-5-IV, V-6-IV with tubercles fused basally; segment 8 without D-3-VIII, without L-1-VIII; with V-5-VIII, V-6-VIII on single tubercle, V-5-VIII tiny, V-6-VIII elongate; segment 9 (Fig. 77B)—with D-5-IX, D-6-IX campaniform sensilla.

DISTRIBUTION AND HABITAT: The genus Leehelea is known from nine species in the Oriental and Australasian Regions ( Borkent 2014). Pupae have been found in sand from a river margin and from a lily pond.

TAXONOMIC DISCUSSION: The pupae of two species of Leehelea are known (Tables 2–3). Male and female pupae of Homohelea and Sphaeromias, two genera closely related to Leehelea, are sexually dimorphic in the arrangement of their forelegs. In males, the foreleg is relatively short (Figs. 40A, D) while in females the foreleg overlaps the midleg entirely (Figs. 40B, E). Only female pupae of Leehelea are known (Fig. 40C) and it is likely that the males, once discovered, will share this dimorphism (see character 44).

The dorsal apotome of one specimen was present but at too great an angle to illustrate. Seta DA-A-1 could not be seen but was likely broken off (considering that all Ceratopogonidae other than some Forcipomyia have a seta present). Overall, the shape of the dorsal apotome appeared similar to that of Sphaeromias longipennis .

Debenham (1974) noted that the pupae of Leehelea were similar to those of Sphaeromias and neither could I find any differences between the two of any significance. Considering these strong similarities, the validity of recognizing the two genera should be reconsidered. Regarding the differences given by Debenham (1974), Sphaeromias may be paraphyletic in relation to Leehelea . They are separated primarily on the basis of a single difference in the shape of the female claw (presence or absence of an inner tooth and their length) and degree of fusion of the male parameres. Further differences seem to me to be minor (size of leg setae and their bases) or invalid (the brush on the ventral surface of Ta 4 is present in at least S. longipennis). Nandi et al. (2012) lists other differences but at least most apply to both genera.

Elson-Harris (1987) keyed the pupae of the two Australian species, L. hispida and L. wasselli .

MATERIAL EXAMINED: L. hispida: 1 pupal exuviae, Nepean River, Menangle, New South Wales, Australia, 9-XII-1968 (ANIC). L. wasselli: 1 pupal exuviae, Roper River Mission, Northern Territory, Australia, 8- XI-1956 (ANIC).