Phaenobezzia Haeselbarth

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 : 110-111

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) .

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 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 22. Dorsal apotomes, anterior view. A. Sphaeromias longipennis (from Wirth & Grogan 1979). B. Amerohelea sordidipes (from Lane et al. 1955). C. Bezzia mollis (from Mayer 1934c). D. Bezzia roldani (from Spinelli & Wirth 1989). E. Bezzia brevicornis (from Spinelli 1983a). F. Clastrieromyia dycei (from Spinelli & Grogan 1986). G. Pachyhelea pachymera (from Spinelli 1983a). H. Palpomyia lineata (from Grogan & Wirth 1979). I. Palpomyia belkini (from Grogan & Wirth 1979, modified). J. Palpomyia flaviceps (from Grogan & Wirth 1979). K. Phaenobezzia opaca (from Wirth & Grogan 1982). L. Paryphoconus oliveirai (from Ronderos et al. 2007).

FIGURE 28. Posterior portion of heads, in ventral view. A. Leehelea hispida. B. Sphaeromias longipennis. C. Xenohelea galatea. D. Bezzia nobilis. E. Clastrieromyia dycei. F. Pachyhelea pachymera. G. Palpomyia subasper. H. Phaenobezzia opaca. I. Paryphoconus grandis.

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 41. Legs, wings and apices of the antennae, in ventral view. A. Bezzia nobilis. B. Clastrieromyia dycei (reconstructed from distorted specimen). C. Pachyhelea pachymera. D. Palpomyia occidentalis. E. Phaenobezzia opaca. F. Paryphoconus oliveirai (reconstructed from distorted specimen).

FIGURE 46. Respiratory organs, most in dorsal view, Figs. 46G. in lateral view, pattern on tracheal tubes shown only in some. A. Homohelea albitudinis (from de Meillon & Wirth 1981). B. Homohelea delanoe (from de Meillon & Wirth 1981). C. Leehelea wasselli (from Elson-Harris 1987). D. Leehelea hispida (from Elson-Harris 1987). E. Sphaeromias longipennis (from Wirth & Grogan 1979). F. Xenohelea galatea. G. Amerohelea sordidipes (from Lane et al. 1955). H. Bezzia glabra (from Wirth 1983b). I. Bezzia obelisca (from Wirth 1983a). J. Bezzia dorsasetula (from Wirth 1983a). K. Bezzia turkmenica (from Glukhova 1979b). L. Clastrieromyia dycei (from Spinelli & Grogan 1986). M. Pachyhelea pachymera (from Spinelli 1983a). N. Palpomyia lineata (from Grogan & Wirth 1979). O. Palpomyia jonesi (from Grogan & Wirth 1979). P. Palpomyia belkini (from Grogan & Wirth 1979). Q. Palpomyia rufa (from Grogan & Wirth 1979). R. Palpomyia flaviceps (from Grogan & Wirth 1979). S. Phaenobezzia opaca (from Wirth & Grogan 1982). T. Paryphoconus oliveirai (from Ronderos et al. 2007).

FIGURE 54. Metathoraces and tergites 1, in dorsal view. A. Palpomyia jonesi. B. Phaenobezzia opaca. C. Paryphoconus oliveirai.

FIGURE 71. Abdominal segments four, in dorsal and ventral view (shagreen not shown). A. Palpomyia occidentalis. B. Phaenobezzia opaca. C. Paryphoconus oliveirai.

FIGURE 78. Abdominal segments 9. A. Palpomyia lineata, male, in dorsal and ventral view (from Grogan & Wirth 1979). B. Palpomyia lineata, female, in dorsal and ventral view (from Grogan & Wirth 1979). C. Palpomyia jonesi, male, in dorsal and ventral view (from Grogan & Wirth 1979. D. Palpomyia jonesi, female, in dorsal and ventral view (from Grogan & Wirth 1979). E. Palpomyia belkini, male, in dorsal and ventral view (from Grogan & Wirth 1979). F. Palpomyia belkini, female, in dorsal and ventral view (from Grogan & Wirth 1979). G. Palpomyia rufa, male, in dorsal and ventral view (from Grogan & Wirth 1979). H. Palpomyia flaviceps, female, in dorsal and ventral view (from Grogan & Wirth 1979). I. Phaenobezzia vacunae, male, in ventral view (from Haeselbarth 1965). J. Phaenobezzia mashonensis, male, in ventral view (from Haeselbarth 1965). K. Phaenobezzia opaca, female, in dorsal and ventral view (from Wirth & Grogan 1982). L. Paryphoconus oliveirai, female, in dorsal view (from Ronderos et al. 2007).

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.