Atyphohelea Borkent

(Figs. 15F, 20J, 25G, 30D, 37A, 44Q, 50A, 62A, 74K)

DIAGNOSIS: Only pupa of Ceratopogonidae with the thorax and abdomen with moderately developed rounded tubercles (Figs. 30D, 62A, as in 33A), with three ventral setae (V-5-IV, V-6-IV, V-7-IV) on well-developed tubercles (Fig. 62A) (not as in Fig. 61A) and the respiratory organ with pores restricted to its apex (Fig. 44Q).

DESCRIPTION: Total length = 2.20–2.94 mm. Without larval exuviae retained on abdomen. Exuviae with flagellum appressed against lateral margin of face, midleg, wing (Fig. 15F, as in Fig. 33B). Ecdysial tear extending into eye sheath (Figs. 15F, 79F). Head: Dorsal apotome (Fig. 20J), without 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. 25G) with mandible well-developed, lacinia absent; palpus extending posterior to posterolateral margin of labium; labium separated medially by labrum, hypopharynx; apex of antenna (Fig. 37A) anterior to barely posterior to posterior extent of midlength portion of midleg (portion lateral to mesosternum), narrowed posteriorly; sensilla: dorsal apotomals (Fig. 20J)—1 short seta, 1 campaniform sensillum; dorsolateral cephalic sclerite sensilla—1 very short seta, 1 campaniform sensillum; clypeal-labrals (Fig. 25G)—2 short, slender setae; oculars (Fig. 25G)—1 seta, 1 campaniform sensillum. Thorax: Prothoracic extension (Fig. 25G) wide, well-developed, extending from palpus to antenna; mesonotum with welldeveloped moderately sized tubercles, not extending posteromedially, with slight protuberance, not dividing metathorax medially (Fig. 50A); respiratory organ (Fig. 44Q) length/width = 2.63–3.45, moderately elongate, somewhat flattened dorsoventrally, 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 annulations to 3/4 length; wing (Fig. 37A) without apical tubercle or angle, separated medially by fore-, midlegs; halter apex and hind leg (as in Fig. 33A) broadly abutting; halter apex abutting anterolateral knob-like extension of tergite 2; legs (Fig. 37A) 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 moderately anterior to apex of midleg; apex of hind leg slightly ventral to, partially abutting apex of midleg laterally; sensilla: anteromedials—1 very short seta, 1 campaniform sensillum; anterolaterals—1 short seta (shagreen making it difficult to interpret if more); dorsal setae (Fig. 30D)—D-1-T, D-2-T, D-5-T peg-like setae, D-4-T seta, D-3-T campaniform sensillum; D-1-T, D-2-T, D-4-T on single tubercle, D-5-T on separate tubercle, D-3-T posterior to D- 4-T; supraalar 2—campaniform sensillum (difficult to discern among shagreen); metathoracics (Fig. 50A)—2 campaniform sensilla; 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, peg-like or thin to thick setae, with rounded, short to moderately elongate tubercles, tergites or sternites entire, each without membranous disc; segment 9 (Fig. 74K) not strongly modified, terminal processes closely approximated basally, each projecting posterodorsolaterally, tapering to pointed apex; sensilla: tergite 1 (Fig. 50A) with 7 setae, 2 campaniform sensilla, including 4 lateral sensilla, D-2-I, D-3-I closely approximated, D-7-I situated anterolaterally near L-1-I; segment 4 (Fig. 62A)—D-2-IV peg-like seta, D-3-IV moderately elongate seta, D-2-IV on large tubercle, D-3-IV on short tubercle; D-5-IV, D-8-IV peg-like setae, D-9-IV short seta, D-7-IV absent or not visible among shagreen; D-5-IV, D-8-IV on elongate separate tubercles, posterior dorsal sensilla in transverse row, arranged medially to laterally: D- 5-IV, D-8-IV, D-9-IV; L-1-IV elongate seta on rounded tubercle just anterior to L-3-IV; L-2-IV; L-3-IV, L-4-IV short setae on rounded tubercles, V-5-IV, V-6-IV, V-7-IV moderately elongate setae, on rounded well-developed tubercles, V-6-IV, V-7-IV closely approximated; segment 8 without D-3-VIII, without L-1-VIII; segment 9 (Fig. 74K)—with D-5-IX, D-6-IX campaniform sensilla.

DISTRIBUTION AND HABITAT: The genus Atyphohelea is known from one Nearctic species and one unnamed species from Taiwan ( Borkent 1998). Adults are associated with smaller bodies of water, seeps, bogs, and small marshes. Pupae have been reared from a lake margin in North Carolina, U.S.A.

TAXONOMIC DISCUSSION: Only one species of Atyphohelea is known as a pupa (Tables 2-3). The fourth abdominal segment of A. macroneura is illustrated without sensilla D-4-IV and D-7-IV but these campaniform sensilla (their form in other Ceratopogonidae) are likely present but hidden among the strong shagreen present over much of the abdomen of this species (in addition the limited material was rather dirty, further obscuring observation of campaniform sensilla).

In nearly all Ceratopogonidae, the anteroventral margin of the wing forms a near right angle, with the margin completely or nearly perpendicular to the dorsal margin of the antenna. I found only two exceptions in the family. In the somewhat crushed specimens of A. macroneura studied here, the angle appeared less abrupt and more rounded (the wing side of the angle was greater). However, I could not convince myself that it was not a product of distortion. Fresh material would be able to test this idea. If present, the feature would be nearly unique in the family. It was also present in an exuviae of one unnamed species of Jenkinshelea (from Kruger N.P) which also had a greater angle, clearly an instance of homoplasy. Other Jenkinshelea had a nearly right angle anteroventral margin.

MATERIAL EXAMINED: A. macroneura: 3 pupal exuviae, Highlands Lake Ravenel, Macon County, North Carolina, USA, 10-VI-1986 (CNCI).