Macropeza Meigen

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 : 82-84

Macropeza Meigen

( Figs. 12C , 21D , 26H , 30M , 38D–E , 45J , 51D , 65B–C , 76A–B )

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. 51D ), abdominal segment 4 with L-1- IV a short seta ( Fig. 65B ) and L-3-IV situtated well anterior of L-2-IV; also unique with D-2-I and D-3-I well separated ( Fig. 51D ) and L-1-VIII present (for all species other than M. natalensis ); also unique 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. 51D ) and D-4-IV situated between D-8-IV and D-9-IV (except for M. natalensis but in that species D-4-V and D-4-VI are between D-8 and D-9 of their respective segments).

DESCRIPTION: Habitus as in Fig. 12C . Total length = 2.59–4.88 mm. Without larval exuviae retained on abdomen. Exuviae with flagellum appressed against lateral margin of midleg, wing (as in Figs. 16B , 33B ). Ecdysial tear extending into eye sheath (as in Figs. 17A , 79I ). Head: Dorsal apotome ( Fig. 21D ), 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. 26H ) with mandible well-developed, lacinia absent; palpus extending posterior to posterolateral margin of labium; labium separated medially by labrum, hypopharynx; apex of antenna ( Figs. 38D–E ) anterior to posterior to, posterior extent of midlength portion of midleg (portion lateral to mesosternum), narrowed posteriorly; sensilla: dorsal apotomals ( Fig. 21D )—1 elongate seta, 1 campaniform sensillum; dorsolateral cephalic sclerite sensilla—1 seta, 1 campaniform sensillum; clypeal-labrals ( Fig. 26H )—2 slender or thick setae; oculars ( Fig. 26H )—2 setae, 1 campaniform sensillum. Thorax: Prothoracic extension ( Fig. 26H ) wide, well-developed, extending from palpus to antenna; mesonotum without or with well-developed moderately sized tubercles, not extending posteromedially, not dividing metathorax medially ( Fig. 51D ); respiratory organ ( Fig. 45J ) length/width = 2.15–4.31, very short, squat to 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 to moderately elongate pedicel, base with short to elongate 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 ( Figs. 38D–E ) 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 ( Figs. 38D–E ) with lateral margin of foreleg near midlength of wing evenly curved; hind leg visible at lateral margin of wing (as in Fig. 33E ); with apex of foreleg moderately anterior to apex of midleg; apex of hind leg abutting apex of midleg laterally, small ventral lobe; sensilla: anteromedials—1 short, 1 elongate seta, 0–1 campaniform sensillum (as in Fig. 31N ); anterolaterals—1 moderately long seta; dorsal setae ( Fig. 30M )—D-1-T, D-2-T, D-5-T peg-like setae or setae, D-4- T elongate seta or peg-like 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. 51D )—1 campaniform sensillum; M-3-T distant from margin of metathorax (at least 1/3 length of metathorax). Abdomen: with tergite 1 with 3 medial spots (in female only), 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, 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 sternites 5–7 each with membranous disc; segment 9 ( Figs. 76A–B ) not strongly modified, terminal processes closely approximated basally, each projecting posterolaterally to nearly laterally, tapering to pointed apex; sensilla: tergite 1 ( Fig. 51D ) with 8 setae, 2 campaniform sensilla, including 4 lateral sensilla, D-2-I, D-3-I well separated, D-7-I situated anterolaterally near L-1-I; segment 4 ( Figs. 65 B-C)—D-2-IV, D-3-IV elongate setae; D-2-IV on elongate or bifid tubercle, D-3-IV on bump or bifid tubercle; D-5-IV peg-like seta, D-8-IV, D-9- IV short to moderately elongate setae, D-7-IV present or absent; 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 or D-5-IV, D-8-IV, D-4-IV, D-9-IV; D-7-IV, if present, near D-3-IV; L-1- IV short to moderately elongate seta on short tubercle, near L-3-IV; L-2-IV, L-4-IV short setae, L-3-IV moderately elongate seta, L-2-IV, on pointed or bifid tubercles, V-5-IV, V-6-IV, V-7-IV short to moderately elongate setae, on short bilobed tubercles, all closely approximated or with only V-5-IV, V-6-IV close; segment 8 without D-3-VIII, with L-1-VIII; segment 9 ( Figs. 76 A-B)—with D-5-IX, D-6-IX campaniform sensilla.

DISTRIBUTION AND HABITAT: The genus Macropeza is known from 22 species in the Nearctic, Palaearctic, Oriental, and Afrotropical Regions ( Borkent 2014 ). Macropeza geari is transferred to Pellucidomyia as a new combination above. Immatures have been collected from mud and among plants at stream and river margins, reservoirs, and from aquatic plants ( Pistia stratiotes ).

TAXONOMIC DISCUSSION: The pupae of four named species of Macropeza have been described ( Tables 2–3 ).

All species of Macropeza , other than M. natalensis , are nearly unique in having L-1-VIII on abdominal segment 8 (see character 71). In addition, nearly all species have another synapomorphy, with D-4-IV between D- 8-IV and D-9-IV. In M. natalensis the sensilla is medial to D-8-IV and D-9-IV but is between the two comparable sensilla on abdominal segments 5 and 6. These observations may suggest that M. natalensis is the sister group to the remaining Macropeza examined or alternately, that the single specimen available had aberrantly distributed sensilla on segment 4.

MATERIAL EXAMINED: M. albitarsis : 2 pupal exuviae, Piława river, Szwecja nr Wałcz, Poland, 5-VII- 2007 ( IZUG). M. bayeri : 1 pupal exuviae (of paratype), M’posa River, Empangeni, Zululand, South Africa, 14-I- 1937 (SAIM). M. natalensis : 1 pupal exuviae, Manypuza River, INYONI, Zululand, South Africa, 19-VIII-1936 (SAIM). M. sp.: 1 pupal exuviae, Nam Houng River Margin, Moung Sayaboury, Sayaboury Province, Laos, 17- XII-1967 (BPBM); 2 pupal exuviae (different species), as previous locality, 17-20-XII-1967 (BPBM); 1 pupal exuviae, as previous locality, 6-9-I-1968 (BPBM).

Figures

FIGURE 12. Habitus. A. Pellucidomyia leei, male, in dorsal view (from Debenham 1970). B. Dibezzia prominens, sex unknown, in dorsal view (form Mayer 1934c). C. Macropeza albitarsis, male, in dorsal view (from Szadziewski & Dominiak, 2007, modified). D. Mallochohelea munda, male, in ventral view (from Szadziewski et al. 1997). E. Sphaeromias fasciata, sex unknown, in dorsal view (from Mayer 1934a). F. Bezzia glabra, female, in ventral view (from Thomsen 1937). G. Palpomyia sp., male, in dorsal view (from Grogan & Wirth 1979). H. Paryphoconus oliveirai, female, in ventral view (modified from Ronderos et al. 2007).

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 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 65. Abdominal segments four, in dorsal and ventral view (shagreen not shown). A. Jenkinshelea magnipennis. B. Macropeza natalensis. C. Macropeza sp. (from Laos).

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.