Probezzia Kieffer

Borkent, Art, 2014, The Pupae of the Biting Midges of the World (Diptera: Ceratopogonidae), With a Generic Key and Analysis of the Phylogenetic Relationships Between Genera, Zootaxa 3879 (1), pp. 1-327 : 85-87

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).

Figures

FIGURE 13. Anterior portion of heads and thoraces, in anterior or anterodorsal view. A. Leptoconops kerteszi, ecdysial sutures of head enhanced to show position. B. Culicoides denticulatus. C. Ceratopogon nr. abstrusus. D. Brachypogon sp. (from 6 km E. Falkland, BC, Canada). E. Alluaudomyia bella, dorsal apotome absent. F. Clinohelea curriei. G. Pellucidomyia leei. H. Mallochohelea caudellii. I. Bezzia sp. (from Bolean Lake, BC, Canada).

FIGURE 16. Anterior portion of heads. A. Parabezzia huberti, in ventral view (distorted slide specimen). B. Parabezzia sp. (5 km. E. Danby, Vermont, USA), ventrolateral view. C. Clinohelea curriei, in ventral view. D. Heteromyia clavata, in ventral view, dorsal apotome not present. E. Heteromyia wokei, in ventral, slightly lateral view. F. Pellucidomyia leei, in ventral view, dorsal apotome not present.

FIGURE 17. Anterior portion of heads. A. Jenkinshelea sp., in ventral view (from Ottawa, ON, Canada). B. Mallochohelea caudellii., in ventral view. C. Sphaeromias longipennis, in ventral view. D. Bezzia sp., in ventral view (6 km E Salmon Arm, BC, Canada). E. Palpomyia sp. (Black Lake, Quebec, Canada), in ventral view. F. Phaenobezzia sp., in ventral view (from Skulsuza, South Africa).

FIGURE 21. Dorsal apotomes, anterior view. A. Dibezzia prominens (from Mayer 1934c). B. Hebetula tonnoiri (from Elson- Harris 1987). C. Jenkinshelea albaria (from Wirth & Grogan 1979). D. Macropeza albitarsis (from Szadziewski & Dominiak, 2007). E. Mallochohelea termophila (from Spinelli 1984, modified). F. Probezzia pallida (from Wirth & Grogan 1979). G. Neobezzia fittkaui (from Ronderos et al. 2011, modified). H. Nilobezzia basispinigera (from Elson-Harris 1987). I. Nilobezzia virago (from Elson-Harris 1987). J. Johannsenomyia argentata (from Wirth & Grogan 1979). K. Lanatomyia electra (from Elson-Harris 1987). L. Anebomyia atripes (from Wirth & Grogan 1979, modified).

FIGURE 27. Posterior portion of heads, in ventral view. A. Mallochohelea caudellii. B. Probezzia sabroskyi. C. Neobezzia fittkaui. D. Nilobezzia curticornis. E. Johannsenomyia argentata. F. Lanatomyia electra. G. Anebomyia atripes. H. Homohelea delanoe.

FIGURE 30. Dorsal setae from left side of thoraces. A. Serromyia sp. B. Allohelea japonica. C. Monohelea obscura. D. Atyphohelea macroneura. E. Parabezzia huberti. F. Echinohelea lanei. G. Clinohelea curriei. H. Heteromyia clavata. I. Pellucidomyia leei. J. Dibezzia debenhamae (dirty specimen, D-5-T perhaps broken setal socket). K. Hebetula tonnoiri. L. Jenkinshelea magnipennis. M. Macropeza natalensis. N. Mallochohelea caudellii. O. Probezzia sabroskyi. P. Neobezzia fittkaui (from Ronderos et al. 2011, modified). Q. Nilobezzia curticornis. R. Johannsenomyia argentata. S. Lanatomyia miles. T. Anebomyia atripes.

FIGURE 31. Dorsal setae from left side of thoraces; anteromedial setae. A. Homohelea delanoe. B. Leehelea hispida. C. Sphaeromias longipennis. D. Xenohelea galatea. E. Bezzia nobilis. F. Clastrieromyia dycei. G. Pachyhelea pachymera. H. Palpomyia jonesi. I. Phaenobezzia mashonensis. J. Paryphoconus oliveirai. K. Echinohelea lanei, partial lateral view (distorted slide specimen). L. Palpomyia (JAD685), dorsal view. M. Palpomyia (JAD685), dorsolateral view. N. Probezzia seminigra, dorsal view of left side.

FIGURE 33. Cephalothoraces and abdominal segments 1–3, in lateral view. A. Parabezzia sp. (5 km. E. Danby, Vermont, USA). B. Clinohelea curriei. C. Pellucidomyia leei. D. Hebetula tonnoiri. E. Jenkinshelea sp. (from Ottawa, ON, Canada). F. Mallochohelea caudellii. G. Probezzia seminigra. H. Nilobezzia sp. (from Kruger NP, South Africa). I. Sphaeromias longipennis. J. Bezzia sp. (6 km E. Salmon Arm, BC, Canada) K. Palpomyia sp. (Black Lake, Quebec, Canada). L. Phaenobezzia sp. (from Skulsuza, South Africa).

FIGURE 39. Legs, wings and apices of the antennae, in ventral view. A. Probezzia sabroskyi. B. Neobezzia fittkaui. C. Nilobezzia sp. (from Australia). D. Johannsenomyia argentata. E. Lanatomyia electra. F. Anebomyia atripes.

FIGURE 45. Respiratory organs, in dorsal view, pattern on tracheal tubes shown only in some. A. Clinohelea tasmaniensis (from Elson-Harris & Kettle 1986). B. Clinohelea bimaculata (from Wirth & Grogan 1979, modified). C. Heteromyia wokei (from Wirth & Grogan 1977). D. Heteromyia clavata (from Wirth & Grogan 1977). E. Pellucidomyia leei (from Elson-Harris & Kettle 1986). F. Pellucidomyia geari (from de Meillon & Wirth 1981). G. Dibezzia prominens (from Mayer 1934c). H. Hebetula tonnoiri (from Debenham 1974). I. Jenkinshelea albaria (from Wirth & Grogan 1979). J. Macropeza albitarsis (from Szadziewski & Dominiak, 2007). K. Mallochohelea albihalter (from Wirth & Grogan 1979). L. Probezzia pallida (from Wirth & Grogan 1979). M. Probezzia xanthogaster (from Wirth & Grogan 1979). N. Neobezzia fittkaui (from Ronderos et al. 2011). O. Nilobezzia aranea (from Elson-Harris 1987). P. Nilobezzia virago (from Elson-Harris 1987). Q. Johannsenomyia argentata (from Wirth & Grogan 1979). R. Lanatomyia electra (from Debenham 1974). S. Lanatomyia miles (from Debenham 1974). T. Anebomyia atripes (from Wirth & Grogan 1979).

FIGURE 51. Metathoraces and tergites 1, in dorsal view. A. Dibezzia debenhamae. B. Hebetula tonnoiri (reconstructed from distorted specimen). C. Jenkinshelea magnipennis. D. Macropeza natalensis. E. Mallochohelea caudellii. F. Probezzia sabroskyi.

FIGURE 66. Abdominal segments four, in dorsal and ventral view (shagreen not shown). A. Mallochohelea nr. caudellii (from Hope, British Columbia, Canada). B. Probezzia sabroskyi. C. Neobezzia fittkaui (from Ronderos et al. 2011, modified).

FIGURE 76. Abdominal segments 9. A. Macropeza albitarsis, male, in ventral view (modified from Szadziewski & Dominiak, 2007). B. Macropeza albitarsis, female, in ventral view (from Szadziewski & Dominiak, 2007). C. Mallochohelea termophila, male, in ventral view (from Spinelli 1984). D. Mallochohelea termophila, female, in dorsal view (from Spinelli 1984). E. Probezzia pallida, male, in dorsal and ventral view (from Wirth & Grogan 1979). F. Neobezzia fittkaui, male, in dorsal view (from Ronderos, et al. 2011). G. Nilobezzia virago, female, in dorsal view (from Elson-Harris 1987). H. Johannsenomyia argentata, male, in dorsal and ventral view (from Wirth & Grogan 1979). I. Johannsenomyia argentata, female, in dorsal and ventral view (from Wirth & Grogan 1979). J. Lanatomyia miles, female, in dorsal view (from Elson- Harris 1987). K. Anebomyia atripes, male, in dorsal and ventral view (from Wirth & Grogan 1979). L. Anebomyia atripes, female, in dorsal and ventral view (from Wirth & Grogan 1979).

FIGURE 79. Evolution of ecdysial sutures and splits in pupal exuviae of Ceratopogonidae, head and anterior portion of thorax in ventral view. Numbers refer to character states of characters 2 and 8 in text. Red indicates ecdysial splits. A. Leptoconopinae. B. Forcipomyiinae. C. Dasyheleinae. D. Most Ceratopogonini. E. Most Ceratopogonini, Clinohelea. F. Allohelea, Atyphohelea, Parabezzia. G. Heteromyia, Pellucidomyia, Hebetula. H. most Sphaeromiini s. lat., most Palpomyiini, Paryphoconus; some Bezzia are nearly completed fused (as in Fig. 79I), with only the dorsal apotome separate, along with an ecdysial split along the base of the antenna and the dorsal thoracic split. I. Jenkinshelea, Macropeza, Mallochohelea, Probezzia, Neobezzia.