Echinohelea Macfie (Figs. 20, 26A, 30F, 31K, 37C, 44T, 50C, 63A, 75B) DIAGNOSIS: Only pupa of Ceratopogonidae with abdominal segment 4 with only two lateral sensilla (L-2-IV, L- 3-IV) and with most sensilla at the bases of bifid tubercles (Fig. 63A). DESCRIPTION: Total length = 2.91 mm. Without larval exuviae retained on abdomen. Exuviae with flagellum appressed against lateral margin of face, midleg, wing (as in Figs. 16B, 33B). Ecdysial tear uncertain but not around base of antenna; at least medial to antennal base (as in Figs. 15B, 79D). Head: Dorsal apotome missing; dorsolateral cephalic sclerite (as in Fig. 13H) fused to scutum, each side uncertain in whole pupa; mouthparts (Fig. 26A) with mandible well-developed, lacinia absent; palpus extending posterior to posterolateral margin of labium; labium separated medially by labrum, hypopharynx; apex of antenna (Fig. 37C) anterior to posterior extent of midlength portion of midleg (portion lateral to mesosternum), narrowed posteriorly; sensilla: dorsal apotomals (Fig. 20)—uncertain; dorsolateral cephalic sclerite sensilla—1 elongate seta, 1 campaniform sensillum; clypeallabrals (Fig. 26A)—1 small seta; oculars (Fig. 26A)—1 campaniform sensillum. Thorax: Prothoracic extension (Fig. 26A) wide, well-developed, extending from palpus to antenna; mesonotum with short tubercles, extending posteriorly but medial division of metathorax uncertain (Fig. 50C); respiratory organ (Fig. 44T) length/width = 4.71, elongate, slender, somewhat flattened, at least apically (perhaps more), with pores closely abutting at apex of respiratory organ, arranged in single row, outer surface smooth, with short, wide pedicel, base with elongate posteromedial apodeme, membranous base of respiratory organ moderately elongate, tracheal tube straight to slightly curved along length, surface smooth; wing (Fig. 37C) with apical tubercle lateral to apex of hind leg, separated medially by fore-, midlegs; halter apex and hind leg (as in Fig. 33A) narrowly abutting; halter apex abutting anterolateral knob-like extension of tergite 2; legs (Fig. 37C) with lateral margin of foreleg near midlength of wing evenly curved; hind leg visible at lateral margin of wing (as in Fig. 32L); with apex of foreleg slightly anterior to apex of midleg; apex of hind leg abutting apex of midleg laterally; sensilla: anteromedials—2 elongate setae, 1 campaniform sensillum (Fig. 31K); anterolaterals—1 moderately long seta; dorsal setae (Fig. 30F)—D-1- T, D-2-T, D-4-T, D-5-T setae, D-3-T campaniform sensillum; D-1-T, D-2-T on single tubercle, D-3-T posteromedial to D-4-T; supraalar 2—campaniform sensillum; metathoracics (Fig. 50C)—1 seta on one side only, 2 campaniform sensilla on both sides; M-3-T near anterior margin of metathorax. Abdomen: without pigmentation pattern, segment 2 as wide or slightly wider than segment 3, segments with undivided, thin to thick setae, with rounded to bilobed, short to somewhat elongate tubercles, tergites or sternites entire, each without membranous disc; segment 9 (Fig. 75B) not strongly modified, terminal processes closely approximated basally, each projecting posterodorsolaterally, tapering to pointed apex; sensilla: tergite 1 (Fig. 50C) with 7 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. 63A)—D-2-IV short seta on bifid tubercle; D-3-IV moderately elongate seta without tubercle; D-5-IV, D-8-IV, D- 9-IV short setae; at base on bifid, separate 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 situated anterorlaterally near D-3-IV; L-1-IV absent, L-2-IV, L-3-IV short setae each at base of bifid tubercle, L-4-IV absent, V-5-IV, V-6-IV, V-7-IV moderately elongate setae, at base of bifid, slender tubercles, V-6-IV, V-7-IV closely approximated; segment 8 without D-3-VIII, without L-1- VIII; segment 9 (Fig. 75B)—with D-5-IX, D-6-IX campaniform sensilla. DISTRIBUTION AND HABITAT: The genus Echinohelea is known from 27 species from nearly every Region worldwide but is absent from the Palaearctic Region (Borkent 2014). An unpublished record from Borneo is known (pers. obs.). The only known immature, the pupa of E. lanei described here, was collected from Flanders, New York, USA. The Flanders Pond site was scrubby red maple growth flooded with a few inches of clear water (Hugo Jamnback, pers. comm.). Grogan (1975) reported a reared specimen of E. lanei collected from under bark of a rotting log in Maryland, USA and Knausenberger (1987) found this species in damp dark brown mud (his collection No. 29) in a roadside ditch in Virginia, USA. De Meillon & Wirth (1991) noted two African species reared from wet soil and from mud at a pond margin. TAXONOMIC DISCUSSION: Only one moderately well preserved pupa of Echinohelea is known, that of E. lanei (Tables 2–3). Wirth (1994b) provided an earlier description. Details of the sensilla distribution of abdominal segment 4 differs here because the specimen was distorted by compression and is reconstructed here. The dorsal apotome of the specimen was not present and cannot, therefore, be characterized. Wirth (1994b) noted that the pupa of Echinohelea was similar to some Heteromyiini (as described by Elson-Harris & Kettle 1986) and this is reflected in the phylogenetic relationship supported here (and Borkent 1995) between Echinohelea and Heteromyiini + Sphaeromiini s. lat. + Palpomyiini + Stenoxenini. MATERIAL EXAMINED: E. lanei: 1 pupal exuviae, Flanders Pond, Flanders, New York, USA, 4-VI-1963 (NYSM).

Published as part of Borkent, Art, 2014, The Pupae of the Biting Midges of the World (Diptera: Ceratopogonidae), With a Generic Key and Analysis of the Phylogenetic Relationships Between Genera, pp. 1-327 in Zootaxa 3879 (1) on pages 74-75, DOI: 10.11646/zootaxa.3879.1.1, http://zenodo.org/record/4949051