Hebetula Wirth & Debenham

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 : 79-80

Hebetula Wirth & Debenham

( Figs. 21B , 26F , 30K , 33D , 38B , 45H , 51B , 64C , 75J )

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. 51B ) and L-1-IV and L-3-IV slender, contrasting with the basally thick L-2-IV and L-4-IV ( Fig. 64C ); also unique with the metathorax with only one campaniform sensillum (M-3-T) situated posterior from the anterior margin of metathorax ( Fig. 51B ), halter extending just anterior of the anterolateral margin of tergite 2 ( Fig. 33D ), and abdominal segment 4 with the lateral setae scattered ( Fig. 64C ) (not forming a transverse row as in Fig. 67B ).

DESCRIPTION: Total length = 3.59 mm. Without larval exuviae retained on abdomen. Exuviae with flagellum appressed against lateral margin of midleg, wing (as in Figs. 16B , 33B ). Ecdysial tear around base of antenna, with narrow connection between face and base of antenna (as in Figs. 16E , 79G ). Head: Dorsal apotome ( Fig. 21B ), 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. 26F ) with mandible well-developed, lacinia absent; palpus extending posterior to posterolateral margin of labium; labium separated medially by labrum, hypopharynx; apex of antenna ( Fig. 38B ) posterior to posterior extent of midlength portion of midleg (portion lateral to mesosternum), narrowed posteriorly; sensilla: dorsal apotomals ( Fig. 21B )—1 elongate seta, 1 campaniform sensillum; dorsolateral cephalic sclerite sensilla—1 seta, 1 campaniform sensillum; clypeal-labrals ( Fig. 26F )—2 slender setae; oculars ( Fig. 26F )—2 setae, 1 campaniform sensillum. Thorax: Prothoracic extension ( Fig. 26F ) wide, well-developed, extending from palpus to antenna; mesonotum yes, low ones tubercles, not extending posteromedially, not dividing metathorax medially ( Fig. 51B ); respiratory organ ( Fig. 45H ) length/width = 3.67–3.81, moderately elongate, somewhat flattened apically, with pores closely abutting at apex of respiratory organ, arranged in single row, outer surface with some wrinkles, with short, wide pedicel, base with elongate posteromedial apodeme, membranous base of respiratory organ moderately elongate, annulated, tracheal tube straight to slightly curved along length, with spirals restricted to base, wrinkles to half length; wing ( Fig. 38B ) with short tubercle at apex of hind leg, separated medially by fore-, midlegs; halter apex and hind leg ( Fig. 33D ) broadly abutting; halter apex extending just anterior of anterolateral margin of tergite 2; legs ( Fig. 38B ) with lateral margin of foreleg near midlength of wing evenly curved; hind leg visible at lateral margin of wing ( Fig. 33D ); with apex of foreleg moderately anterior to apex of midleg; apex of hind leg abutting apex of midleg laterally; sensilla: anteromedials—2 elongate setae (as in Figs. 31L–M ); anterolaterals—1 moderately long seta; dorsal setae ( Fig. 30K )—D-1-T, D-2-T, D-5-T, setae, D-4-T broken seta?, D-3-T campaniform sensillum, D-3-T anterolateral to D-4-T; supraalar 2—campaniform sensillum; metathoracics ( Fig. 51B )—1 campaniform sensillum; M-3-T distant from margin of metathorax (at least 1/3 length of metathorax). Abdomen: with tergite 1 with 1 medial spot, tergites 2–7 with medial area with stripe, 2 anterolateral spots, 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, short tubercles, tergites or sternites entire, each without membranous disc; segment 9 ( Fig. 75J ) not strongly modified, terminal processes closely approximated basally, each projecting posterodorsolaterally, tapering to pointed apex; sensilla: tergite 1 ( Fig. 51B ) with 7 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. 64C )—D-2-IV, D-3-IV short to moderately elongate setae on short tubercles; D-5-IV, D-8- IV, D-9-IV moderately elongate setae; D-5-IV on single 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 moderately elongate seta on short tubercle, just anterior of base of tubercle with L-2-IV; L-2-IV, L-3-IV, L-4-IV moderately elongate setae, L-2-IV, L-4-IV on rounded tubercles, V-5- IV, V-7-IV moderately elongate, V-6-IV short seta, on rounded tubercles, all closely approximated; segment 8 without D-3-VIII, without L-1-VIII; segment 9 ( Fig. 75J )—with D-5-IX, D-6-IX campaniform sensilla.

DISTRIBUTION AND HABITAT: The genus Hebetula is known from 22 species in the Oriental, Afrotropical and Australasian Regions ( Borkent 2014). Pupae have been reared from moss in a creek and from the margin of a river.

TAXONOMIC DISCUSSION: Only one species of this genus is known as a pupa ( Tables 2–3 ). The dorsal setae on the thorax of the single pupa differed on either side, with D-3-T missing on the right side. D-4-T was either a sensillum basiconica (as shown here) or the setae were broken off on both sides. If future specimens confirm D-4- T as a sensillum basiconica, this would be a distinctive and unique feature within the Ceratopogonidae (D-4-T is absent in some taxa).

Elson-Harris (1987) illustrated the respiratory organ of H. tonnoiri with a midband of darker pigmentation but reexamination indicates the pigmentation extends to the base of the respiratory organ.

MATERIAL EXAMINED: H. tonnoiri : 1 pupal exuviae, Nattai River, Mittagong, New South Wales, Australia, 4-XI-1964 (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 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 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 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 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 38. Legs, wings and apices of the antennae, in ventral view. A. Dibezzia debenhamae (reconstructed from distorted specimen). B. Hebetula tonnoiri (reconstructed from distorted specimen). C. Jenkinshelea sp. (from Ottawa, ON, Canada). D. Macropeza natalensis. E. Macropeza sp. (from Laos). F. Mallochohelea caudellii.

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 64. Abdominal segments four, in dorsal and ventral view (shagreen not shown). A. Pellucidomyia leei. B. Dibezzia debenhamae (reconstructed from distorted and dirty specimen). C. Hebetula tonnoiri.

FIGURE 67. Abdominal segments four, in dorsal and ventral view (shagreen not shown). A. Nilobezzia curticornis. B. Johannsenomyia argentata. C. Lanatomyia miles.

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 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.