Phaenobezzia Haeselbarth

(Figs. 17F, 22K, 28H, 31I, 33L, 41E, 46S, 54B, 71B, 78I–K)

DIAGNOSIS: Only pupa of Ceratopogonidae with the male with the metathorax with only M-3-T present and situated at least 1/3 length of metathorax and genital lobes very short, extending no more than about 0.8 length of segment 9 (measured medially) (Figs. 78I–J); female (without genital lobes) with the metathorax with only one campaniform sensillum (M-3-T) situated at least ⅓ the length of the metathorax from its anterior margin (Fig. 54B), apex of the halter extending posteriorly to about 1/6 length of tergite 2 (Fig. 33L), abdominal segment 4 with V-5-IV, V-6-IV and V-7-IV closely approximated (Fig. 71B), abdominal segment 8 with V-5-VIII and V-6-VIII on separate tubercles or if on partially to completely fused tubercles, then V-5-VIII well-developed (not minute), without L-1-VIII (not diagnosable as different from Bezzia and Palpomyia); however, most species of Bezzia have two or more campaniform sensilla on the dorsal apotome (Figs. 22C–D), a nearly unique condition found otherwise only in P. flavipes and P. jonesi (Fig. 22J).

DESCRIPTION: Total length = 2.25–4.78 mm. Without larval exuviae retained on abdomen. Exuviae with flagellum appressed against lateral margin of midleg, wing (Figs. 17F, 33L). Ecdysial tear around base of antenna, along lateral margin of face to palpus (Figs. 17F, 79H). Head: Dorsal apotome (Fig. 22K), uncertain 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. 28H) with mandible well-developed, lacinia absent; palpus extending posterior to posterolateral margin of labium; labium separated medially by labrum, hypopharynx; apex of antenna (Fig. 41E) posterior to posterior extent of midlength portion of midleg (portion lateral to mesosternum), narrowed posteriorly; sensilla: dorsal apotomals (Fig. 22K)—1 elongate seta, 1 campaniform sensillum; dorsolateral cephalic sclerite sensilla—1 seta, 1 campaniform sensillum; clypeallabrals (Fig. 28H)—2 slender setae; oculars (Fig. 28H)—1 seta, 1 campaniform sensillum. Thorax: Prothoracic extension (Fig. 28H) wide, well-developed, extending from palpus to antenna; mesonotum with short tubercles, not extending posteromedially, not dividing metathorax medially (Fig. 54B); respiratory organ (Fig. 46S) length/width = 3.86–4.93, elongate, moderately slender, somewhat flattened apically, with pores closely abutting at apex of respiratory organ, arranged in single curved row, outer surface with some wrinkles, with short, wide pedicel, base with moderate elongate posteromedial apodeme, membranous base of respiratory organ short, annulated, tracheal tube straight to slightly curved along length, with spirals restricted to base, wrinkles to half length; wing (Fig. 41E) without apical tubercle or angle, separated medially by fore-, midlegs; halter apex and hind leg (Fig. 33L) broadly abutting; halter apex extending posteriorly to 1/6 length of tergite 2; legs (Fig. 41E) with lateral margin of foreleg near midlength of wing evenly curved; hind leg visible at lateral margin of wing (Fig. 33L); with apex of foreleg moderately anterior to apex of midleg; apex of hind leg abutting apex of midleg laterally; sensilla: anteromedials—2 elongate setae, 1 campaniform sensillum (as in Figs. 31L–M); anterolaterals—1 moderately long seta; dorsal setae (Fig. 31I)—D-1-T, D-2-T, D-4-T, D-5-T setae, D-3-T campaniform sensillum, D-3-T lateral to D- 4-T; supraalar 2—campaniform sensillum; metathoracics (Fig. 54B)—1 campaniform sensillum; M-3-T distant from margin of metathorax (at least 1/3 length of metathorax). Abdomen: pigmentation light to dark brown, with tergites 1–7 with medial area with stripe, 2 spots or 1 with single medial spot and tergites 2–7 with stripe and 2 spots, medial stripe barely discernable in some, sternites 3–7 with medial stripe, anterolateral spot, segment 2 as wide or slightly wider than segment 3, segments with undivided, thin to thick setae, with rounded to pointed, short to moderately elongate tubercles, tergites or sternites entire, each without membranous disc; segment 9 (Figs. 78I–K) not strongly modified, terminal processes closely approximated to separated basally, each projecting posterodorsolaterally, tapering to pointed apex; apex of male genital lobe anterior or to level of base of terminal process (Figs. 78I–J); sensilla: tergite 1 (Fig. 54B) with 8 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. 71B)—D-2-IV, D-3- IV moderately elongate setae on moderately developed tubercles; D-5-IV, D-8-IV short setae, D-9-IV elongate seta, D-7-IV present or absent; D-5-IV on short tubercle, D-8-IV, D-9-IV on basally fused, 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, if present, near D-3-IV or L-2-IV; L-1-IV elongate seta on rounded tubercle, just anterior of base of tubercle with L-2-IV, L-3-IV; L-2-IV, L-3-IV, L-4-IV elongate setae on pointed tubercles, L-2-IV, L-3-IV on single tubercle, V-5-IV, V-7-IV moderately elongate setae, V-6-IV elongate seta, on short tubercles, all closely approximated, with tubercles of V-5-IV, V-6-IV fused basally; segment 8 without D-3-VIII, without L-1-VIII; with V-5-VIII, V-6-VIII on single tubercle, V-5-VIII tiny, V-6-VIII elongate. Segment 9 (Figs. 78 I-K)—with D-5-IX, D- 6-IX campaniform sensilla.

DISTRIBUTION AND HABITAT: The genus Phaenobezzia is known from 34 species from every Region worldwide other than the Australasian Region ( Borkent 2014). Immatures have been collected from stream and river margins, seepage pools, rice fields, back waters of rivers, marshes (usually among vegetation), among Pistia stratiotes, lake margins and reservoirs. Knausenberger (1987) provides some further microhabitat information for P. opaca .

TAXONOMIC DISCUSSION: The pupae of nine species of Phaenobezzia are known (Tables 2–3).

Haeselbarth (1965) used the shape of the terminal processes to distinguish the pupae of the four African species he studied and indicated some further character states which may separate some species. Thomsen (1937) drew the dorsal apotome of P. opaca with two setae (and no campaniform sensillum); she likely mistook the campaniform sensillum as the broken base of a seta. Wirth & Grogan (1982) drew abdominal segment 4 of P. opaca with V-7-IV situated laterally, close to L-4-IV and this is significantly different than that observed here for this species and all other Phaenobezzia (Fig. 71B).

MATERIAL EXAMINED: P. cinnae: 2 pupal exuviae, Shaw’s Drift, Eshowe, Zululand, South Africa, 9-IX- 1936 (1 USNM, 1 SAIM); 2 pupal exuviae, no data (ZSMC) . P. fulvithorax: 3 pupal exuviae, Bainville, Roosevelt County, Montana, USA, 9-VI-1969 (USNM) ; 1 pupal exuviae, Cranberry Lake, St. Lawrence County, New York, USA, 25-VI-1963 (USNM) ; 2 pupal exuviae, Rio Frio, 5 mi NW of Leakey, Real County, Texas, USA, 23-V-1972 (USNM) ; 1 pupal exuviae, Pedernales River, Gillespie County, Texas, USA, 19-III-1972 (USNM) . P. mashonensis: 1 pupal exuviae, Tugela River, Middledrift, South Africa, 6-IX-1935 (USNM) ; 2 pupal exuviae, Burgershall, Hazyview, East Transvaal, South Africa, 3-XII-1973 (NMSA) ; 1 pupal exuviae, Sand River between White River and Hazyview, East Transvaal, South Africa, 2-XII-1973 (NMSA) ; 1 pupal exuviae, Mseleni River, North Zululand, South Africa, 7-II-1936 (1 ZSMC, 1 SAIM) ; 1 pupal exuviae, N'Kwaleni, Zululand, South Africa, 10- IX-1935 (SAIM) . P. opaca: 1 pupal exuviae, Mississippi River, Carleton County, Ontario, Canada, 24-VI-1966 (VPIC) ; 4 pupal exuviae, Turin, Lewis County, New York, USA, 21-VI-1963 (USNM); 1 pupal exuviae, Independence River, Glenfield, Lewis County, New York, USA, 22-VI-1963 (USNM) ; 1 pupal exuviae, Meachum Lake, New York, USA, 27-VI-1986 (CNCI) ; 1 pupal exuviae, Pillsburg State Park, New Hampshire, USA, 26-VI- 1986 (CNCI) ; 1 pupal exuviae, Dunmore Springs, Pocahontas County, Virginia, USA, 24-VI-1976 (VPIC); 2 pupal exuviae, Bear Creek Lake, Cumberland County, Virginia, USA, 30-IV-1977 (VPIC) ; 1 pupal exuviae, Jackson River, Allegheny County, Virginia, USA, 4-VI-1977 (VPIC) ; 1 pupal exuviae, Hamilton County, Florida, USA, 3- VI-1993 (JHEC) . P. pistiae: 1 pupal exuviae, Chipinga, Zimbabwe, VIII-1941 (SAIM) ; 1 pupal exuviae, no location/date (SAIM). P. rubiginosa: 8 pupal exuviae, Anninskoe lake, Pskov Province, Russia, 15-VII-1996 (ZIN) . P. vacunae: 4 pupal exuviae, stream nr. Umlalazi River, Eshowe, Zululand, South Africa, 19-IX-1935 (SAIM) . P. sp.: 1 pupal exuviae, Canoe, British Columbia, Canada, 7-VII-1990 (CNCI); 1 pupa, 7 pupal exuviae, Dark Canyon Creek, Eddy County, New Mexico, USA, 13-IV-1991 (CNCI) ; 1 pupal exuviae, 10 mi SE of Middleburg, Lake Dunmore, Vermont, USA, 23-VI-1986 (CNCI) ; 1 pupal exuviae, Meachum Lake, New York, USA, 21-VI-1986 (CNCI) ; 1 pupal exuviae, Dunmore Springs, Pocahontas County, West Virginia, USA, 24-VI- 1976 (VPIC); 1 pupal exuviae, Rheocrene, Rio Bento Gomes, Mato Grosso, Brazil, 29-VIII-1994 (CNCI) ; 1 pupal exuviae (in glycerin), Skulsuza, South Africa, 11-IV-1993 (ANIC); 1 pupal exuviae, Nam Houng river, Moung Sayaboury, Sayaboury Province, Laos, 17-XII-1967 (BPBM) ; 1 pupal exuviae, as previous locality, 6-I-1968 (BPBM); 2 pupal exuviae, Bamboo Creek, Kimberley, Western Australia, Australia (ANIC) ; 1 pupal exuviae, Lenard River gorge, Kimberley, Western Australia, Australia, 12-X-1995 (ANIC) ; 1 pupal exuviae, Mitchell Falls, Kimberley, Western Australia, Australia (ANIC ; 1 pupal exuviae, Lower Durack River, Kimberley, Western Australia, Australia (ANIC) ; 1 pupal exuviae, Crossing Pool, Fortescue River, Pilbara, Western Australia, Australia (ANIC) ; 2 pupal exuviae, Nattai River, Mittagong, New South Wales, Australia, 15-XI-1968 (ANIC) .