Probezzia Kieffer

(Figs. 21F, 27B, 30O, 31N, 33G, 39A, 45L–M, 51F, 66B, 76E)

DIAGNOSIS: Only pupa of Ceratopogonidae 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. 51F), abdominal segment 1 with D-2- I and D-3-I well separated (Fig. 51F), abdominal segment 4 with L-1-IV a short seta and D-5-IV on a bifid tubercle (Fig. 66B).

DESCRIPTION: Total length = 2.91–5.56 mm. Without larval exuviae retained on abdomen. Exuviae with flagellum appressed against lateral margin of midleg, wing (as in Fig. 16B, 33G). Ecdysial tear extending into eye sheath (as in Figs. 17A, 79I). Head: Dorsal apotome (Fig. 21F), 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. 27B) with mandible well-developed, lacinia absent; palpus extending posterior to posterolateral margin of labium; labium separated medially by labrum, hypopharynx; apex of antenna (Fig. 39A) barely anterior to equal to posterior extent of midlength portion of midleg (portion lateral to mesosternum), narrowed posteriorly; sensilla: dorsal apotomals (Fig. 21F)—1 elongate seta, 1 campaniform sensillum; dorsolateral cephalic sclerite sensilla—1 seta, 1 campaniform sensillum; clypeal-labrals (Fig. 27B)—2 moderately to very thick setae; oculars (Fig. 27B)—2 setae, 1 campaniform sensillum. Thorax: Prothoracic extension (Fig. 27B) wide, well-developed nearly even width to narrow dorsolaterally, extending from palpus to antenna; mesonotum with short to moderately sized tubercles, not extending posteromedially, not dividing metathorax medially (Fig. 51F); respiratory organ (Figs. 45L–M) length/width = 2.61–4.00, short to intermediate length, somewhat flattened apically, with pores closely abutting at apex of respiratory organ, arranged in single row, outer surface with a few to numbers of wrinkles, with short, wide pedicel, base with very short, pale 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. 39A) without apical tubercle or angle, separated medially by fore-, midlegs; halter apex and hind leg (Fig. 33G) broadly abutting; halter apex abutting anterolateral knob-like extension of tergite 2; legs (Fig. 39A) with lateral margin of foreleg near midlength of wing evenly curved; hind leg visible at lateral margin of wing (Fig. 33G); 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 or short thick seta, 1 elongate seta (Fig. 31N); anterolaterals—1 moderately long seta; dorsal setae (Fig. 30O)—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 on single tubercle, D-3-T lateral to D-4-T; supraalar 2—campaniform sensillum; metathoracics (Fig. 51F)—1 campaniform sensillum; M-3-T distant from margin of metathorax (at least 1/3 length of metathorax). Abdomen: pigmentation light brown in some; with tergite 1 with 3 medial spots, tergites 2–7 with medial area with stripe, 2 anterolateral spots, 2 pairs on tergite 8 in some, sternites 3–4 (5–7 membranous) or sternites 3–7 with medial stripe, anterolateral spot, 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, not membranous or with 5–7 or 6–7 each with membranous disc; segment 9 (Fig. 76E) not strongly modified, terminal processes closely approximated basally, each projecting posterodorsolaterally, tapering to pointed apex; sensilla: tergite 1 (Fig. 51F) with 8 setae, 2 campaniform sensilla, including 3 lateral sensilla, D-2-I, D-3-I moderately or well separated, D-7-I situated anterolaterally near L-1-I; segment 4 (Fig. 66B)—D-2-IV, D-3- IV short to moderately elongate setae on short tubercles; D-5-IV peg-like seta, D-8-IV short seta, D-9-IV moderately elongate seta; D-5-IV on single, bifid 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 to moderately elongate seta on rounded tubercle, just anterior of base of tubercle with L-2-IV; L-2-IV, L-4-IV short setae, L-3-IV moderately elongate seta on pointed 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 (Fig. 76E)—with D-5-IX, D-6-IX campaniform sensilla.

DISTRIBUTION AND HABITAT: The genus Probezzia is known from 28 species, all restricted to the Holarctic Region ( Borkent 2014). Larvae are present in the benthos of lakes, reservoirs, rivers and streams and pupae can be found on their margins. The pupae of some species are particularly common in the flotsam on beaches (Williams 1955, pers. obs.). Knausenberger (1987) provides detailed information on the behaviour, ecology and distribution of the immatures of a number of species in the eastern USA.

TAXONOMIC DISCUSSION: The pupae of 14 species of Probezzia have been described (Tables 2–3). The abdominal membranous discs are present or absent in different species of Probezzia and it is clear that the feature is homoplastic within the genus, with even otherwise very similar species with or without the discs. Knausenberger (1987) discussed the puzzling presence or absence but could not indicate a related ecological factor to explain this character state distribution. Pupae of P. seminigra have abdominal membranous discs on sternites 5–7. However, one male, certainly conspecific with others from the same locality, had discs on only the posterior half of 5 and then full ones on 6–7, indicating intraspecific variation. The feature is discussed further under Character 50.

Study of a series of Probezzia pallida indicated that at least three species are actually present under this name. Specimens from each of North Dakota, of Ottawa, Ontario and Allegeny, New York and of other localities in New York could be distinguished based on the presence or absence of pupal abdominal membranous discs, shapes and sizes of pupal CL-1-H, D-1-T and D-2-T, size and pigmentation of male adult antenna, size of female adult claws, and pigmentation of female adult postnotum.

MATERIAL EXAMINED: P. albitibia: 3 pupal exuviae (of paratypess), Algonquin Park, Ontario, Canada, 8- VI-1960 (USNM); 2 pupal exuviae, Meachum Lake, New York, USA, 21-VI-1986 (CNCI); 3 pupal exuviae (of paratypes), Blue Mountain Lake, Hamilton County, New York, USA, 10-VI-1960 (USNM); 2 pupal exuviae (of paratypes), Piercefield, St. Lawrence County, New York, USA, 26-VI-1963 (USNM); 1 pupal exuviae (of paratype), Potomac River at Scott Run, Fairfax County, Virginia, 4-VI-1955 (USNM). P. flavonigra: 2 pupal exuviae, 10 km SW of Anahim Lake, British Columbia, Canada, 3-VII-1992 (CNCI); 2 pupal exuviae, Canoe, British Columbia, Canada, 15-VII-1990 (CNCI); 5 pupal exuviae, Sicamous, British Columbia, Canada, 8-VII- 1989 (CNCI); 1 pupal exuviae, 28 km E of Enderby, British Columbia, Canada, 13-VII-1989 (CNCI). P. fuscipennis: 6 pupal exuviae, 10 mi SE of Middleburg, Lake Dunmore, Vermont, USA, 23-VI-1986 (CNCI); 1 pupal exuviae (of paratype), Blue Mountain Lake, Hamilton County, New York, USA, 10-VI-1960 (CNCI). P. pallida: 4 pupal exuviae, Rideau River, Ottawa, Ontario, Canada, 29-May-1960 (1 CNCI, 3 USNM); 6 pupal exuviae, White Earth River, Mountrail County, North Dakota, USA, 7-VI-1969 (USNM); 1 pupal exuviae, Allegany State Park, New York, USA, 28-V-1963 (USNM); 1 pupal exuviae, Piercefield, St. Lawrence County, New York, USA, 26-VI-1963 (USNM); 1 pupal exuviae, Genesee River, Portageville, New York, USA, 13-VI- 1963 (USNM); 2 pupal exuviae, no locality, VI-1953 (USNM). P. rosewalli: 1 pupal exuviae (of paratype), Kilbourne, Louisiana, USA, 10-V-1947 (USNM). P. sabroskyi: 1 pupal exuviae, 10 km S of Okanagan Falls, British Columbia, Canada, 14-V-1989 (CNCI); 1 pupal exuviae, Meachum Lake, New York, USA, 21-VI-1986 (CNCI); 1 pupal exuviae, Allegany State Park, New York, USA, 28-V-1963 (USNM). P. seminigra: 12 pupal exuviae, Cottonwood Lake, 5 km S of Nelson, British Columbia, Canada, 6-VII-1992 (CNCI); 2 pupal exuviae, Canoe, British Columbia, Canada, 15-VII-1990 (CNCI); 3 pupal exuviae (in glycerin), Sicamous, BC, Canada, 8- VII-1989 (CNCI); 1 pupal exuviae, Rutka Tartak, nr. Suwałki, Poland 2-VII-1973 (IZUG); 3 pupal exuviae, Anninskoe lake, Pskov Province, Russia, 23-26-VI-1995 (ZIN); 1 pupal exuviae, Lower Austria, Lunzer Untersee (the lowest-lying of the three major lakes in the Lunz area), Spongilla aufwuchs in 'middle zone' near S shore, Austria, 27-VI-1942 (ZSMC); 2 pupal exuviae, shore of pond at "Zillertal" inn, North Rhine-Westphalia, Riemke N of Bochum, Germany, 9-VI-1916 (ZSMC). P. smithi: 3 pupal exuviae, Wolf River, Outaganie County, Wisconsin, USA, 15-VI-1954 (USNM); 4 pupal exuviae, Potomac River at Scott Run, Fairfax County, Virginia, USA, 7-VI- 1955 (USNM). P. williamsi: 2 pupal exuviae, 10 mi SE of Middleburg, Lake Dunmore, Vermont, USA, 23-VI-1986 (CNCI). P. xanthogaster: 6 pupal exuviae, Allegany State Park, New York, USA, 28-V-1963 (USNM); 8 pupal exuviae, Taughannock Falls, Tompkins County, New York, USA, 15-VI-1963 (USNM); 1 pupal exuviae, Genesee River, Portageville, New York, USA, 13-VI-1963 (USNM). P. nr. seminigra: 1 pupal exuviae, 10 km S. Okanagan Falls, British Columbia, Canada, 14-V-1989 (CNCI). P. sp.: 2 pupal exuviae, Trail Bay, Manitoba, Canada (CNCI); 2 pupal exuviae (in glycerin), Shirley Bay, Ottawa, Ontario, Canada, 13-VI-1972, (CNCI); 4 pupal exuviae, Meachum Lake, New York, USA, 21-VI-1986 (CNCI); 1 pupal exuviae, 1 km N of Camden, Maine, USA, 1-VIII- 1985 (CNCI); 1 pupal exuviae, Potomac River at Scott Run, Fairfax County, Virginia, USA, 7-VI-1955 (USNM); 2 pupae, Lerchenauer See, Germany, 27-VI-1990, 28-VII-8-VIII-1990 (CNCI); 5 pupal exuviae, Anninskoe lake, Pskov Province, Russia, 23-26-VI-1995 (ZIN).