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 Mallochohelea Wirth ( Figs. 2C, E , 12D , 13H , 17B , 21E , 27A , 30N , 33F , 38F , 45K , 51E , 66A , 76C–D ) DIAGNOSIS : Only pupa of Ceratopogonidae with the metathorax with only one campaniform sensillum (M-3-T) situated posterior from the anterior margin of the metathorax ( Fig. 51E ), abdominal segment 1 with D-2-I and D-3- I well separated ( Fig. 51E ), abdominal segment 4 with L-1-IV a short seta ( Fig. 66A ), D-5-IV on an undivided tubercle, and D-4-IV situated medial to D-8-IV and D-9-IV (not diagnosable as different from Jenkinshelea ). DESCRIPTION : Habitus as in Fig. 12D . Total length = 2.81–6.50 mm . Without larval exuviae retained on abdomen. Exuviae with flagellum appressed against lateral margin of midleg, wing ( Figs. 17B , 33F ). Ecdysial tear extending into eye sheath ( Figs. 17B , 79I ). Head : Dorsal apotome ( Fig. 21E ), without ventral line of weakness, without dorsomedial tubercle, without central dome; dorsolateral cephalic sclerite ( Fig. 13H ) fused to scutum, each side separated medially by dorsal apotome in whole pupa; mouthparts ( Fig. 27A ) with mandible well-developed, lacinia absent; palpus extending posterior to posterolateral margin of labium; labium separated medially by labrum, hypopharynx; apex of antenna ( Fig. 38F ) anterior to posterior extent of midlength portion of midleg (portion lateral to mesosternum), narrowed posteriorly; sensilla: dorsal apotomals ( Fig. 21E )—1 elongate seta, 1 campaniform sensillum; dorsolateral cephalic sclerite sensilla—1 seta, 1 campaniform sensillum; clypeal-labrals ( Fig. 27A )—2 thick setae; oculars ( Fig. 27A )—1–2 setae, 1 campaniform sensillum. Thorax : Prothoracic extension ( Fig. 27A ) wide, well-developed nearly even width to narrow dorsolaterally, extending from palpus to antenna; mesonotum with short tubercles, not extending posteromedially, not dividing metathorax medially ( Fig. 51E ); respiratory organ ( Fig. 45K ) length/width = 2.48–2.56, very to moderately short, squat, 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 without (?) or with very short posteromedial apodeme, membranous base of respiratory organ short, tracheal tube straight to slightly curved along length, with spirals restricted to base, wrinkles to half length; wing ( Fig. 38F ) without apical tubercle or angle, separated medially by fore-, midlegs; halter apex and hind leg ( Fig. 33F ) broadly abutting; halter apex abutting anterolateral knob-like extension of tergite 2; legs ( Fig. 38F ) with lateral margin of foreleg near midlength of wing evenly curved; hind leg visible at lateral margin of wing ( Fig. 33F ); with apex of foreleg moderately anterior to apex of midleg; apex of hind leg abutting apex of midleg laterally or small gap between the two; sensilla: anteromedials—1 short peg, 1 elongate seta (as in Fig. 31N ); anterolaterals—1 moderately long seta; dorsal setae ( Fig. 30N )—D-1-T, D-2-T, D-5-T peg-like setae, D-4-T elongate seta, D-3-T campaniform sensillum; D-1-T, D-2-T on single tubercle, D-3-T anterolateral to D-4-T; supraalar 2—campaniform sensillum; metathoracics ( Fig. 51E )—1 campaniform sensillum; M-3-T distant from margin of metathorax (at least 1/3 length of metathorax). Abdomen : pigmentation light brown, poorly defined in some; with tergite 1 with 3 medial spots, tergites 2–7 with medial area with stripe, 2 anterolateral spots, some with 1–2 pairs on segment tergite 8, sternites 3–7 (anterolateral spots on 5–7 poorly defined, pale) all light brown, or just very light medial strip on sternites 4, 5 and with 6–7 membranous or just very light medial strip on sternites 3, 4 and with sternites 5–7 membranous or absent, segment 2 as wide or slightly wider than segment 3, segments with undivided, peg-like or thin to thick setae, with rounded to pointed, short to moderately elongate tubercles, with sternites 5–7 or 6–7 each with membranous disc; segment 9 ( Figs. 76C–D ) not strongly modified, terminal processes closely approximated basally, each projecting posterodorsolaterally, tapering to pointed apex; sensilla: tergite 1 ( Fig. 51E ) with 8 setae, 2 campaniform sensilla, including 3 lateral sensilla, D-2-I, D-3-I well separated, D-7-I situated anteriorly near D-3-I; segment 4 ( Fig. 66A )—D-2-IV, D-3-IV moderately elongate setae, D-2-IV on bifid tubercle, D-3-IV on short tubercle; D-5-IV peg-like seta, D-8-IV short seta, D-9-IV moderately elongate seta; D-5-IV on single tubercle, D-8-IV, D-9-IV on separate but 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 short seta on pointed tubercle, just anterior of base of tubercle with L-2-IV; L-2-IV, L-3-IV, L-4-IV short setae on short tubercles, V-5-IV, V-7-IV short setae, V-6-IV elongate seta, on short or moderately elongate tubercles, all closely approximated; segment 8 without D-3-VIII, without L-1-VIII; segment 9 ( Figs. 76 C-D)—with D-5-IX, D- 6-IX campaniform sensilla. DISTRIBUTION AND HABITAT : The genus Mallochohelea is known from 48 species from every Region worldwide ( Borkent 2014 , with some moved to Anebomyia and Dibezzia ). Immatures have been reared from stream and river margins, thermal stream, reservoirs, ponds, and lakes. TAXONOMIC DISCUSSION : The pupae of 12 species of Mallochohelea have been described ( Tables 2–3 ). Wirth & Grogan (1979) provide a key to the pupae of the two species of Mallochohelea from the Potomac Valley ( Washington DC area, USA ), as well as Anebomyia atripes , then called M. atripes . Thienemann (1928) , Mayer (1934a) and Lenz (1934) provided keys to a few European species known at that time (as members of either Dicrobezzia (now a synonym of Probezzia ) or Johannsenomyia ). However, they all recognized M. dentata as distinct from M. munda (the former now considered a synonym of the latter) and both Thienemann (1928) and Mayer (1934a) additionally recognized M. breviforceps as distinct and it too is now considered a synonym of M. munda . MATERIAL EXAMINED : M. albibasis : 1 pupal exuviae, Cottonwood Lake, 5 km S of Nelson, British Columbia , Canada , 6-VII-2008 (CNCI); 2 pupal exuviae, 7 km S of Hope, British Columbia , Canada , 8-VIII-1985 (CNCI); 1 pupal exuviae, Outegamie County, Wisconsin , USA , 15-VI-1954 (USNM). M. albihalter : 1 pupal exuviae, Allegany State Park, New York , USA , 28-V-1963 (USNM). M. caudellii : 1 pupal exuviae (in glycerin), Bolean Lake, 6 km E of Falkland, BC, Canada , 12–13-VII-1989 (CNCI). M. errinae : 1 pupal exuviae (of holotype ), Klein Yukskei, Johannesburg, Transvaal, South Africa , V-1939 (SAIM); 1 pupal exuviae, Mageliesberg Agricultural School, Transvaal, South Africa , 13-XI-1973 (NMSA). M. inermis : 2 pupal exuviae, Vrevo lake, Leningrad Province , Russia , 19-VI-1968 (ZIN); 1 pupal exuviae, no locality/date (ZSMC). M. satelles : 4 pupal exuviae (of paratype ), Nattai River, Mittagong, New South Wales , Australia , 18-II-1969 (ANIC). M. setigera : 1 pupal exuviae, River Gruzskii Elanchik, Donetsk Province , Ukraine , 29-IV-1970 (ZIN). M. smithi : 4 pupal exuviae, Black Lake, Stanleyville, Ontario , Canada , 29-VI-1975 (WLGC); 1 pupal exuviae, Rideau River, Ottawa, Ontario , Canada , 29-V-1960 (USNM); 1 pupal exuviae, Snow Hill, Maryland , 19-V-1968 (USNM); 1 pupal exuviae, 10 mi SE of Middleberg, Lake Dunmore, Vermont , USA , 23-VI-1986 (CNCI). M. sybleae : 1 pupal exuviae (of paratype ), Centerville, Humboldt County, California , USA , 13-VIII-1948 (USNM). M. termophila : 2 pupal exuviae (of paratype ), Caimancito, Jujuy , Argentina , 12-VIII-1980 (MLPA). M . nr. caudellii : 1 pupal exuviae, 7 km S of Hope, British Columbia , Canada , 8-VIII-1985 (CNCI). M . nr. pullata : 3 pupal exuviae, 6 km NE of Falkland, British Columbia , Canada , 12-VII-1989 (CNCI). M . sp.: 1 pupal exuviae, Bear Creek, 0.5 mi SE of Karlsbad Springs, Ontario , Canada , 1967 (CNCI); 1 pupal exuviae, McCarrs Creek, New South Wales , Australia , 11-XI-1956 (ANIC).