The Pupae of the Biting Midges of the World (Diptera: Ceratopogonidae), With a Generic Key and Analysis of the Phylogenetic Relationships Between Genera Author Borkent, Art text Zootaxa 2014 2014-10-31 3879 1 1 327 journal article 3916 10.11646/zootaxa.3879.1.1 4b976ba3-1a7b-4dd5-8afb-630b7cc8a695 1175-5326 4949051 6423894B-97D9-4286-ABB9-D4AF072B57FD Clinohelea Kieffer ( Figs. 13F , 16C , 20M–N , 26B , 30G , 33B , 37D , 45A–B , 50D , 63B , 75C ) DIAGNOSIS : Only pupa of Ceratopogonidae with the metathorax with only two campaniform sensilla (M-2-T, M-3-T) on the anterior margin of the metathorax, D-7-I situated anteriorly, about 1/2 way between D-5-I and D-2-I ( Fig. 50D , but see taxonomic discussion) and the dorsal apotome relatively narrow ( Figs. 20 M-N; compared to Fig. 20O ). DESCRIPTION : Total length = 2.59–4.47 mm . Without larval exuviae retained on abdomen. Exuviae with flagellum appressed against lateral margin of midleg, wing ( Figs. 16C , 33B ). Ecdysial tear around base of antenna tear posterolaterally ( Figs. 16C , 79E ); along prothoracic extension. Head : Dorsal apotome ( Figs. 20M–N ), without ventral line of weakness, without dorsomedial tubercle, without central dome; dorsolateral cephalic sclerite ( Fig. 13F ) fused to scutum, each side separated medially by dorsal apotome in whole pupa; mouthparts ( Fig. 26B ) with mandible well-developed, lacinia absent; palpus extending posterior to posterolateral margin of labium; labium separated medially by labrum, hypopharynx; apex of antenna ( Fig. 37D ) posterior to posterior extent of midlength portion of midleg (portion lateral to mesosternum), narrowed posteriorly; sensilla: dorsal apotomals ( Figs. 20M–N )—1 elongate seta, 1 campaniform sensillum; dorsolateral cephalic sclerite sensilla—1 seta, 1 campaniform sensillum; clypeal-labrals ( Fig. 26B )—2 slender setae; oculars ( Fig. 26B )—2 elongate setae. Thorax : Prothoracic extension ( Fig. 26B ) wide, well-developed, extending from palpus to antenna; mesonotum with short tubercles, not extending posteromedially, with or without slight protuberance, not dividing metathorax medially ( Fig. 50D ); respiratory organ ( Figs. 45A–B ) length/width = 3.59–7.65, elongate, slender, gradually somewhat flattened dorsoventrally, with pores closely abutting or slightly separated at apex of respiratory organ, arranged in single row, outer surface smooth, with moderately elongate pedicel, base with elongate posteromedial apodeme, membranous base of respiratory organ moderately elongate, plain or partly annulated, tracheal tube straight to slightly curved along length, with spirals restricted to base, remainder with wrinkles; wing ( Fig. 37D ) without apical tubercle or angle, separated medially by fore-, midlegs; halter apex and hind leg ( Fig. 33B ) broadly abutting; halter apex abutting anterolateral knob-like extension of tergite 2; legs ( Fig. 37D ) with lateral margin of foreleg near midlength of wing evenly curved; hind leg visible at lateral margin of wing ( Fig. 33B ); with apex of foreleg slightly to moderately anterior to apex of midleg; apex of hind leg abutting apex of midleg laterally, small ventral lobe; sensilla: anteromedials—1 elongate or 2 short setae; anterolaterals—1 short seta; dorsal setae ( Fig. 30G )—D- 1-T, D-2-T, D-5-T peg-like, D-4-T seta, D-3-T campaniform sensillum; D-1-T, D-2-T on single tubercle, D-3-T lateral to slightly posterolateral to D-4-T; supraalar 2—campaniform sensillum; metathoracics ( Fig. 50D )—2 campaniform sensilla; M-3-T near anterior margin of metathorax. Abdomen : pigmentation very light brown or just indicated by bare patches, tergite 1 with 3 medial spots, tergites 2–7 with medial area with stripe, 2 anterolateral spots, sternites not pigmented or sternites 3–7 with bare median patch; anterolateral areas not evident, segment 2 as wide or slightly wider than segment 3, segments with undivided, peg-like or thin to thick setae, with rounded to pointed or bifid, short to moderately elongate tubercles, tergites or sternites entire, each without membranous disc; segment 9 ( Fig. 75C ) not strongly modified, terminal processes separated basally, each projecting posterodorsolaterally, tapering to pointed apex; sensilla: tergite 1 ( Fig. 50D ) with 7–8 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. 63B )—D-2-IV peg-like seta, D-3-IV seta, D-2-IV on bifid tubercle, D-3-IV on short tubercle; D-5- IV peg-like seta, D-8-IV, D-9-IV short setae; D-5-IV on single tubercle, D-8-IV, D-9-IV on single bifid tubercle or separate but closely approximated, 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 short seta on pointed tubercle, well anterior of posterior lateral setae; L-2-IV, L-3-IV, L-4-IV short setae on elongate tubercles, V-5-IV, V-6-IV, V-7-IV short setae on bifid tubercles, all closely approximated; segment 8 without D-3-VIII, without L-1-VIII; segment 9 ( Fig. 75C )—with D- 5-IX, D-6-IX campaniform sensilla. DISTRIBUTION AND HABITAT : The genus Clinohelea is known from 40 species from every Region worldwide ( Borkent 2014 ). Immatures are known from the margins of streams, fens, seepage area into a lake, muddy margins of rice paddies, and rock pools ( de Meillon & Wirth 1991 , Knausenberger 1987 ). TAXONOMIC DISCUSSION : There are seven species of Clinohelea known as pupae ( Tables 2–3 ). Spinelli & Duret (1993) included the differences between the two Neotropical species known in their key to the adults. Elson-Harris & Kettle (1986) keyed the two known Australian species. Wirth & Grogan (1979) illustrated the dorsal apotome of C. bimaculata with an excavated dorsal margin but reexamination of this species indicates that this was likely based on a damaged or misdrawn specimen. Elson-Harris & Kettle (1986) illustrated abdominal segment 4 of C. dryas with D-7-IV in a posterior position, which would be unique within a broader group, including the Heteromyiini + Sphaeromiini s. lat. + Palpomyiini + Stenoxenini (see character 63); the species should be reexamined. They likely overlooked both D-4-IV and D-7-IV in their illustrations of C. tasmaniensis . Elson-Harris (1990) used the enlarged base upon which DA-1-H is situated (on the dorsal apotome) to distinguish Australian Heteromyiini . Within this region (and elsewhere), the feature does not characterize Pellucidomyia and therefore is not inclusive. The base is large and swollen in Clinohelea ( Figs. 20M–N ) but I could not consistently use this feature to distinguish this genus from some Sphaeromiini s. lat. (e.g. Fig. 21D, F ). The strongly tapering dorsal portion of the dorsal apotome of Clinohelea pointed out by Elson-Harris (1990) is distinctive within the Heteromyiini + Sphaeromiini s. lat. + Palpomyiini + Stenoxenini . MATERIAL EXAMINED : C. bimaculata : 1 pupal exuviae, Spruce Knob Lane, Randolph County, Virginia , USA , 21-VII-1976 (VPIC). C. curriei : 2 pupal exuviae, 1 pupal exuviae (in glycerin), 6 km E of Salmon Arm, BC, Canada , 6-VI-1990 (CNCI). C. horacioi : 2 pupal exuviae, Pipeline Road, Gamboa, Canal Zone, Panama , VII-1967 (MLPA). C. nigripes : 1 pupal exuviae, Pipeline Road, Gamboa, Canal Zone, Panama , VII-1967 (MPLA). C. unimaculata : 2 pupal exuviae, Severskii Donets River, Donetsk Province , Ukraine , 2-IV-1970 (ZIN).