Clinohelea 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 : 75-76

Clinohelea Kieffer

( Figs. 13F , 16C , 20M–N , 26B , 30G , 33B , 37D , 45A–B , 50D , 63B , 75C )

DIAGNOSIS: Only pupa of Ceratopogonidae with the metathorax with only two campaniform sensilla (M-2-T, M-3-T) on the anterior margin of the metathorax, D-7-I situated anteriorly, about 1/2 way between D-5-I and D-2-I ( Fig. 50D , but see taxonomic discussion) and the dorsal apotome relatively narrow ( Figs. 20 M-N; compared to Fig. 20O ).

DESCRIPTION: Total length = 2.59–4.47 mm. Without larval exuviae retained on abdomen. Exuviae with flagellum appressed against lateral margin of midleg, wing ( Figs. 16C , 33B ). Ecdysial tear around base of antenna tear posterolaterally ( Figs. 16C , 79E ); along prothoracic extension. Head: Dorsal apotome ( Figs. 20M–N ), without ventral line of weakness, without dorsomedial tubercle, without central dome; dorsolateral cephalic sclerite ( Fig. 13F ) fused to scutum, each side separated medially by dorsal apotome in whole pupa; mouthparts ( Fig. 26B ) with mandible well-developed, lacinia absent; palpus extending posterior to posterolateral margin of labium; labium separated medially by labrum, hypopharynx; apex of antenna ( Fig. 37D ) posterior to posterior extent of midlength portion of midleg (portion lateral to mesosternum), narrowed posteriorly; sensilla: dorsal apotomals ( Figs. 20M–N )—1 elongate seta, 1 campaniform sensillum; dorsolateral cephalic sclerite sensilla—1 seta, 1 campaniform sensillum; clypeal-labrals ( Fig. 26B )—2 slender setae; oculars ( Fig. 26B )—2 elongate setae. Thorax: Prothoracic extension ( Fig. 26B ) wide, well-developed, extending from palpus to antenna; mesonotum with short tubercles, not extending posteromedially, with or without slight protuberance, not dividing metathorax medially ( Fig. 50D ); respiratory organ ( Figs. 45A–B ) length/width = 3.59–7.65, elongate, slender, gradually somewhat flattened dorsoventrally, with pores closely abutting or slightly separated at apex of respiratory organ, arranged in single row, outer surface smooth, with moderately elongate pedicel, base with elongate posteromedial apodeme, membranous base of respiratory organ moderately elongate, plain or partly annulated, tracheal tube straight to slightly curved along length, with spirals restricted to base, remainder with wrinkles; wing ( Fig. 37D ) without apical tubercle or angle, separated medially by fore-, midlegs; halter apex and hind leg ( Fig. 33B ) broadly abutting; halter apex abutting anterolateral knob-like extension of tergite 2; legs ( Fig. 37D ) with lateral margin of foreleg near midlength of wing evenly curved; hind leg visible at lateral margin of wing ( Fig. 33B ); with apex of foreleg slightly to moderately anterior to apex of midleg; apex of hind leg abutting apex of midleg laterally, small ventral lobe; sensilla: anteromedials—1 elongate or 2 short setae; anterolaterals—1 short seta; dorsal setae ( Fig. 30G )—D- 1-T, D-2-T, D-5-T peg-like, D-4-T seta, D-3-T campaniform sensillum; D-1-T, D-2-T on single tubercle, D-3-T lateral to slightly posterolateral to D-4-T; supraalar 2—campaniform sensillum; metathoracics ( Fig. 50D )—2 campaniform sensilla; M-3-T near anterior margin of metathorax. Abdomen: pigmentation very light brown or just indicated by bare patches, tergite 1 with 3 medial spots, tergites 2–7 with medial area with stripe, 2 anterolateral spots, sternites not pigmented or sternites 3–7 with bare median patch; anterolateral areas not evident, 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, each without membranous disc; segment 9 ( Fig. 75C ) not strongly modified, terminal processes separated basally, each projecting posterodorsolaterally, tapering to pointed apex; sensilla: tergite 1 ( Fig. 50D ) with 7–8 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. 63B )—D-2-IV peg-like seta, D-3-IV seta, D-2-IV on bifid tubercle, D-3-IV on short tubercle; D-5- IV peg-like seta, D-8-IV, D-9-IV short setae; D-5-IV on single tubercle, D-8-IV, D-9-IV on single bifid tubercle or separate but closely approximated, 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 seta on pointed tubercle, well anterior of posterior lateral setae; L-2-IV, L-3-IV, L-4-IV short setae on elongate tubercles, V-5-IV, V-6-IV, V-7-IV short setae on bifid tubercles, all closely approximated; segment 8 without D-3-VIII, without L-1-VIII; segment 9 ( Fig. 75C )—with D- 5-IX, D-6-IX campaniform sensilla.

DISTRIBUTION AND HABITAT: The genus Clinohelea is known from 40 species from every Region worldwide ( Borkent 2014 ). Immatures are known from the margins of streams, fens, seepage area into a lake, muddy margins of rice paddies, and rock pools ( de Meillon & Wirth 1991, Knausenberger 1987).

TAXONOMIC DISCUSSION: There are seven species of Clinohelea known as pupae ( Tables 2–3 ). Spinelli & Duret (1993) included the differences between the two Neotropical species known in their key to the adults. Elson-Harris & Kettle (1986) keyed the two known Australian species.

Wirth & Grogan (1979) illustrated the dorsal apotome of C. bimaculata with an excavated dorsal margin but reexamination of this species indicates that this was likely based on a damaged or misdrawn specimen. Elson-Harris & Kettle (1986) illustrated abdominal segment 4 of C. dryas with D-7-IV in a posterior position, which would be unique within a broader group, including the Heteromyiini + Sphaeromiini s. lat. + Palpomyiini + Stenoxenini (see character 63); the species should be reexamined. They likely overlooked both D-4-IV and D-7-IV in their illustrations of C. tasmaniensis .

Elson-Harris (1990) used the enlarged base upon which DA-1-H is situated (on the dorsal apotome) to distinguish Australian Heteromyiini . Within this region (and elsewhere), the feature does not characterize Pellucidomyia and therefore is not inclusive. The base is large and swollen in Clinohelea ( Figs. 20M–N ) but I could not consistently use this feature to distinguish this genus from some Sphaeromiini s. lat. (e.g. Fig. 21D, F ). The strongly tapering dorsal portion of the dorsal apotome of Clinohelea pointed out by Elson-Harris (1990) is distinctive within the Heteromyiini + Sphaeromiini s. lat. + Palpomyiini + Stenoxenini .

MATERIAL EXAMINED: C. bimaculata : 1 pupal exuviae, Spruce Knob Lane, Randolph County, Virginia, USA, 21-VII-1976 (VPIC). C. curriei : 2 pupal exuviae, 1 pupal exuviae (in glycerin), 6 km E of Salmon Arm, BC, Canada, 6-VI-1990 (CNCI). C. horacioi : 2 pupal exuviae, Pipeline Road, Gamboa, Canal Zone, Panama, VII-1967 (MLPA). C. nigripes : 1 pupal exuviae, Pipeline Road, Gamboa, Canal Zone, Panama, VII-1967 (MPLA). C. unimaculata : 2 pupal exuviae, Severskii Donets River, Donetsk Province, Ukraine, 2-IV-1970 (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 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 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 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 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.