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 Johannsenomyia Malloch ( Figs. 21J , 27E , 30R , 39D , 45Q , 52C , 67B , 76H–I ) DIAGNOSIS : Only pupa of Ceratopogonidae with abdominal segment 4 with all four lateral setae (L-1-IV, L-2-IV, L-3-IV, L-4-IV) arranged transversely, closely approximated and on the anterior half of segment ( Fig. 67B ). DESCRIPTION : Total length = 4.03–5.38 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 ( Fig. 21J ), 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. 27E ) with mandible well-developed, lacinia absent; palpus extending posterior to posterolateral margin of labium; labium entire (not divided medially); apex of antenna ( Fig. 39D ) posterior to posterior extent of midlength portion of midleg (portion lateral to mesosternum), narrowed posteriorly; sensilla: dorsal apotomals ( Fig. 21J )—1 elongate seta, 1 campaniform sensillum; dorsolateral cephalic sclerite sensilla—1 seta, 1 campaniform sensillum; clypeallabrals ( Fig. 27E )—2 slender setae; oculars ( Fig. 27E )—2 setae, 1 campaniform sensillum. Thorax : Prothoracic extension ( Fig. 27E ) wide, well-developed but narrow dorsolaterally, not extending to antenna; mesonotum with short tubercles, not extending posteromedially, not dividing metathorax medially ( Fig. 52C ); respiratory organ ( Fig. 45Q ) length/width = 2.41–3.09, moderately elongate, swollen apically, 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, tracheal tube straight, swollen curved apically in some, with spirals restricted to base, plates to half length; wing ( Fig. 39D ) without apical tubercle or angle, separated medially by fore-, midlegs; halter apex and hind leg (as in Fig. 33A ) broadly abutting; halter apex abutting or just anterior to anterolateral knob-like extension of tergite 2; legs ( Fig. 39D ) with lateral margin of foreleg near midlength of wing evenly curved; hind leg visible at lateral margin of wing (as in Fig. 33I ); male with apex of foreleg moderately anterior to apex of midleg, female with apex of foreleg slightly anterior to apex of midleg; apex of hind leg abutting apex of midleg laterally, small ventral lobe; sensilla: anteromedials—2 elongate setae (as in Figs. 31L–M ); anterolaterals—1 moderately long seta; dorsal setae ( Fig. 30R )—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. 52C )—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, sternites 3–7 with anterolateral spots difficult to discern, segment 2 as wide or slightly wider than segment 3, segments with undivided, thin to thick setae, with rounded to slightly pointed, short tubercles, tergites or sternites entire, each without membranous disc; segment 9 ( Figs. 76H–I ) not strongly modified, terminal processes closely approximated basally, each projecting posterodorsolaterally to somewhat laterally, tapering to pointed apex; sensilla: tergite 1 ( Fig. 52C ) 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. 67B )—D-2-IV, D-3-IV short to moderately elongate setae on short tubercles; D-5-IV, D-8-IV, D-9-IV setae; 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 short tubercle, near L-2-IV, L-3-IV, L-4-IV on anterior half of segment; L-2-IV, L-3-IV, L-4-IV short setae on short pointed tubercles, V-5-IV, V-6-IV, V-7-IV short setae on rounded tubercles; V- 5-IV, V-6-IV closely approximated; segment 8 without D-3-VIII, without L-1-VIII; segment 9 ( Figs. 76 H-I)—with D-5-IX, D-6-IX campaniform sensilla. DISTRIBUTION AND HABITAT : The genus Johannsenomyia is known from 27 species from every Region worldwide but is absent from most of the Palaearctic (known only from north Africa) ( Borkent 2014 ). Immatures have been reared from river margins, lakes and Knausenberger (1987) additionally recorded a species from a pond. Williams (1955) described how mature larvae of J. argentata burrowed into the sand on a lake beach to pupate. TAXONOMIC DISCUSSION : Only one species of Johannsenomyia has been described as a pupa ( Tables 2–3 ). Mayer (1957) also described the pupa of J. albidorsata (de Meillon) (as a Dicrohelea ) but I have examined the specimen and it is in fact a species of Bezzia . De Meillon (1937) in his description of the adult actually noted that the pupal exuviae of the holotype had been mixed with those of a Palpomyia and a Bezzia and its identification was therefore uncertain. Mayer (1957) however was clearly unaware of the uncertain association and described the purported pupal exuviae as if it was that of J. albidorsata . Debenham (1974) gave a brief synopsis of the pupa of Johannsenomyia but based it on the misidentified description by Mayer (1957) . MATERIAL EXAMINED : J. argentata : 1 pupal exuviae, Storrs, Connecticut , USA , 1-VI-1953 (USNM); 1 pupal exuviae, Patuxent Wildlife Refuge, Prince George’s County, Maryland , USA , 24-V-1977 (USNM); 6 pupal exuviae, Potomac River at Scott Run, Fairfax County, Virginia , USA , 7-VI-1955 (USNM).