Heteromyia Say

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 : 76-77

Heteromyia Say

( Figs. 16D–E , 20O , 26C , 30H , 37E , 45C–D , 50E , 63C , 75D–F )

DIAGNOSIS: Only pupa of Ceratopogonidae with only one campaniform sensillum (DA-2-H) on the dorsal apotome ( Fig. 20O ), the metathorax with only two campaniform sensilla (M-2-T, M-3-T) on the anterior margin of the metathorax and with a long slender terminal process, lacking an elongate seta ( Figs. 75D–F ).

DESCRIPTION: Total length = 5.31–7.13 mm. Without larval exuviae retained on abdomen. Exuviae with flagellum appressed against lateral margin of midleg, wing ( Fig. 16E , as in Fig. 33B ). Ecdysial tear around base of antenna, with narrow connection between face and base of antenna ( Figs. 16D–E , 79G ); along prothoracic extension. Head: Dorsal apotome ( Fig. 20O ), with 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. 26C ) with mandible well-developed, lacinia absent; palpus extending posterior to posterolateral margin of labium; labium entire (not divided medially); apex of antenna ( Fig. 37E ) anterior to posterior extent of midlength portion of midleg (portion lateral to mesosternum); sensilla: dorsal apotomals ( Fig. 20O )—1 short to moderate seta, 1 campaniform sensillum; dorsolateral cephalic sclerite sensilla—1 seta, 1 campaniform sensillum; clypeal-labrals ( Fig. 26C )—2 slender setae; oculars ( Fig. 26C )—2 elongate setae, 0–1 campaniform sensillum. Thorax: Prothoracic extension ( Fig. 26C ) wide, well-developed, narrow dorsolaterally, not extending to antenna; mesonotum without or with 1 spinous tubercles, not extending posteromedially, not dividing metathorax medially ( Fig. 50E ); respiratory organ ( Figs. 45C–D ) length/width = 3.12–5.90, elongate, slender, somewhat flattened apically, with pores closely abutting at apex of respiratory organ, arranged in single curved row, outer surface with or without some wrinkles, with short to moderately elongate pedicel, base with moderate elongate posteromedial apodeme, membranous base of respiratory organ short, tracheal tube straight to slightly curved along length, with spirals restricted to base, remainder with wrinkles to near apex; wing ( Fig. 37E ) without apical tubercle or angle, separated medially by fore-, midlegs; halter apex and hind leg (as in Fig. 33A ) broadly abutting; halter apex abutting anterolateral knob-like extension of tergite 2; legs ( Fig. 37E ) with lateral margin of foreleg near midlength of wing evenly curved; hind leg visible at lateral margin of wing (as in Fig. 33B ); with apex of foreleg moderately anterior to apex of midleg; apex of hind leg abutting apex of midleg laterally in male, ventral to apex of midleg in female; sensilla: anteromedials—1 short, 1 long seta; anterolaterals—1 moderately long seta; dorsal setae ( Fig. 30H )—D-1-T, D-2-T, D-5-T peg-like or elongate setae, D-4-T seta, D-3-T campaniform sensillum or D-1-T, D-2-T, D-4-T, D-5-T all elongate, D-3-T campaniform sensillum, D-3-T lateral to slightly posterolateral to D-4-T; supraalar 2—campaniform sensillum; metathoracics ( Fig. 50E )—1-2 campaniform sensilla; M-3-T near anterior margin of metathorax. Abdomen: with tergite 1 with 3 medial spots (fused in some), tergites 2-7 with medial area with stripe, 2 spots, anterolateral spots, sternites 3-7 with medial, anterolateral spot, segment 2 as wide or slightly wider than segment 3, segments with undivided, peglike or thin to thick setae, with pointed, short to moderately elongate tubercles, tergites or sternites entire, each without membranous disc; segment 9 ( Figs. 75D–F ) not strongly modified, terminal processes separated basally, each projecting posterodorsolaterally, elongate, slender, tapering to pointed, hooked apex; sensilla: tergite 1 ( Fig. 50E ) with 8 setae, 2 campaniform sensilla, including 3 lateral sensilla, D-2-I, D-3-I closely approximated, D-7-I situated anteriorly near D-3-I; segment 4 ( Fig. 63C )—D-2-IV peg-like seta, D-3-IV seta, on short tubercles; D-5- IV peg-like seta, D-8-IV short setae, D-9-IV moderately elongate seta; D-5-IV on short tubercle, D-8-IV, D-9-IV on separate but closely approximated short 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 tooth on pointed tubercle, just anterior of L-3-IV; L-2-IV, L-4-IV short setae, L-3-IV moderately elongate seta, on pointed tubercles, V-5-IV, V-6- IV, V-7-IV on pointed tubercles; V-5-IV, V-6-IV closely approximated; segment 8 without D-3-VIII, without L-1- VIII; segment 9 ( Figs. 75 D-F)—with D-5-IX, D-6-IX campaniform sensilla.

DISTRIBUTION AND HABITAT: The genus Heteromyia is known from 13 species in the New World ( Borkent 2014 ). Immatures have been found in semi-aquatic or aquatic vegetation, with rearings from Sphagnum and Cabomba . Knausenberger (1987) provides further details of habitats, including swamps and stream and river margins. Adults have been collected from bogs, fens and, in Panama, a hotspring with abundant wet soil and vegetation on its margin.

TAXONOMIC DISCUSSION: Three species of Heteromyia are known as pupae ( Tables 2–3 ). The elongate, slender terminal process ( Figs. 75D–F ) is distinctive in the group Heteromyiini + Sphaeromiini s. lat. + Palpomyiini + Stenoxenini except for Dibezzia ( Fig. 75I ) which has an associated elongate seta (D-5-IX) and some species of Bezzia ( Fig. 77E ) which have two or more campaniform sensilla on the dorsal apotome ( Figs. 22C–D ).

Wirth & Grogan (1977) illustrated the dorsal apotome of H. wokei with two setae but reexamination shows the presence of one seta and one campaniform sensillum.

MATERIAL EXAMINED: H. clavata : 1 pupal exuviae, Juan Mina Station, Canal Zone, Panama, 3-VI-1939 (USNM). H. pratti : 1 pupal exuviae, Patuxent Rescue Center, Prince George’s County, Maryland, 13-VI-1977 (USNM). H. wokei : 4 pupal exuviae, Bluefields, Nicaragua, 25-VIII-1943 (USNM).

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 20. Dorsal apotomes, anterior view. A. Stilobezzia soror (from Mayer 1934c). B. Stilobezzia lutea (from Wirth & Grogan 1981). C. Stilobezzia antennalis (from Wirth & Grogan 1981). D. Stilobezzia pallidiventris (from Wirth & Grogan 1981). E. Schizonyxhelea forattinii (from Borkent 2000a). F. Schizonyxhelea bulla (from Wirth & Grogan 1981, modified). G. Serromyia sp.. H. Allohelea japonica. I. Monohelea obscura. J. Atyphohelea macroneura (from Borkent 1998). K. Parabezzia alexanderi male (from Grogan & Wirth 1977a, modified). L. Parabezzia alexanderi female (from Grogan & Wirth 1977a, modified). M. Clinohelea dryas (from Elson-Harris & Kettle 1986). N. Clinohelea tasmaniensis (from Elson-Harris & Kettle 1986). O. Heteromyia wokei (from Wirth & Grogan 1977, modified). P. Pellucidomyia leei (from Elson-Harris & Kettle 1986).

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 26. Posterior portion of heads, in ventral view. A. Echinohelea lanei (reconstructed from distorted specimen). B. Clinohelea curriei. C. Heteromyia wokei. D. Pellucidomyia leei. E. Dibezzia debenhamae (reconstructed from distorted specimen). F. Hebetula tonnoiri. G. Jenkinshelea magnipennis H. Macropeza bayeri.

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 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 37. Legs, wings and apices of the antennae, in ventral view. A. Atyphohelea macroneura. B. Parabezzia huberti. C. Echinohelea lanei (reconstructed from distorted specimen). D. Clinohelea currei. E. Heteromyia clavata. F. Pellucidomyia leei.

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 50. Metathoraces and tergites 1, in dorsal view. A. Atyphohelea macroneura. B. Parabezzia huberti. C. Echinohelea lanei. D. Clinohelea currei. E. Heteromyia wokei. F. Pellucidomyia leei.

FIGURE 63. Abdominal segments four, in dorsal and ventral view (shagreen not shown). A. Echinohelea lanei (reconstructed from distorted specimen). B. Clinohelea unimaculata. C. Heteromyia wokei.

FIGURE 75. Abdominal segments 9. A. Parabezzia huberti, female, in ventral view (from Grogan & Wirth 1977a). B. Echinohelea lanei (from Wirth 1994b). C. Clinohelea bimaculata, female, in ventral view (from Wirth & Grogan 1979). D. Heteromyia wokei, male, in dorsal and ventral view (from Wirth & Grogan 1977). E. Heteromyia wokei, female, in dorsal and ventral view (from Wirth & Grogan 1977). F. Heteromyia clavata, female, in dorsal and ventral view (from Wirth & Grogan 1977). G. Pellucidomyia leei, male, in dorsal view (from Elson-Harris & Kettle 1986). H. Pellucidomyia geari, male, in ventral view (from de Meillon & Wirth 1981). I. Dibezzia debenhamae, male, in dorsal and ventral view (shagreen not shown). J. Hebetula tonnoiri, female, in dorsal view (from Elson-Harris 1987). K. Jenkinshelea albaria, male, in dorsal and ventral view (from Wirth & Grogan 1979). L. Jenkinshelea albaria, female, in dorsal and ventral view (from Wirth & Grogan 1979).

FIGURE 77. Abdominal segments 9. A. Homohelea delanoe, female, in dorsal view (from de Meillon & Wirth 1981). B. Leehelea wasselli, female, in dorsal view (from Elson-Harris 1987). C. Sphaeromias longipennis, male, in dorsal and ventral view (from Wirth & Grogan 1979). D. Xenohelea galatea, female, in dorsal and ventral view (shagreen not shown). E. Bezzia roldani, male, in dorsal view (from Spinelli & Wirth 1989). F. Bezzia roldani, female, in dorsal view (from Spinelli & Wirth 1989). G. Bezzia brevicornis, male, in dorsal view (from Spinelli 1983a). H. Bezzia acanthodes, female, in dorsal view (from Spinelli 1983a). I. Clastrieromyia dycei, male, in ventral view (from Spinelli & Grogan 1986). J. Clastrieromyia dycei, female, in ventral view (from Spinelli & Grogan 1986). K. Pachyhelea pachymera, male, in ventral view (from Spinelli 1983a). L. Pachyhelea pachymera, female, in dorsal view (from Spinelli 1983a).

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.