Mallochohelea Wirth

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 : 84-85

Mallochohelea Wirth

( Figs. 2C, E , 12D , 13H , 17B , 21E , 27A , 30N , 33F , 38F , 45K , 51E , 66A , 76C–D )

DIAGNOSIS: Only pupa of Ceratopogonidae with the metathorax with only one campaniform sensillum (M-3-T) situated posterior from the anterior margin of the metathorax ( Fig. 51E ), abdominal segment 1 with D-2-I and D-3- I well separated ( Fig. 51E ), abdominal segment 4 with L-1-IV a short seta ( Fig. 66A ), D-5-IV on an undivided tubercle, and D-4-IV situated medial to D-8-IV and D-9-IV (not diagnosable as different from Jenkinshelea ).

DESCRIPTION: Habitus as in Fig. 12D . Total length = 2.81–6.50 mm. Without larval exuviae retained on abdomen. Exuviae with flagellum appressed against lateral margin of midleg, wing ( Figs. 17B , 33F ). Ecdysial tear extending into eye sheath ( Figs. 17B , 79I ). Head: Dorsal apotome ( Fig. 21E ), without ventral line of weakness, without dorsomedial tubercle, without central dome; dorsolateral cephalic sclerite ( Fig. 13H ) fused to scutum, each side separated medially by dorsal apotome in whole pupa; mouthparts ( Fig. 27A ) with mandible well-developed, lacinia absent; palpus extending posterior to posterolateral margin of labium; labium separated medially by labrum, hypopharynx; apex of antenna ( Fig. 38F ) anterior to posterior extent of midlength portion of midleg (portion lateral to mesosternum), narrowed posteriorly; sensilla: dorsal apotomals ( Fig. 21E )—1 elongate seta, 1 campaniform sensillum; dorsolateral cephalic sclerite sensilla—1 seta, 1 campaniform sensillum; clypeal-labrals ( Fig. 27A )—2 thick setae; oculars ( Fig. 27A )—1–2 setae, 1 campaniform sensillum. Thorax: Prothoracic extension ( Fig. 27A ) wide, well-developed nearly even width to narrow dorsolaterally, extending from palpus to antenna; mesonotum with short tubercles, not extending posteromedially, not dividing metathorax medially ( Fig. 51E ); respiratory organ ( Fig. 45K ) length/width = 2.48–2.56, very to moderately short, squat, 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 without (?) or with very short 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 ( Fig. 38F ) without apical tubercle or angle, separated medially by fore-, midlegs; halter apex and hind leg ( Fig. 33F ) broadly abutting; halter apex abutting anterolateral knob-like extension of tergite 2; legs ( Fig. 38F ) with lateral margin of foreleg near midlength of wing evenly curved; hind leg visible at lateral margin of wing ( Fig. 33F ); with apex of foreleg moderately anterior to apex of midleg; apex of hind leg abutting apex of midleg laterally or small gap between the two; sensilla: anteromedials—1 short peg, 1 elongate seta (as in Fig. 31N ); anterolaterals—1 moderately long seta; dorsal setae ( Fig. 30N )—D-1-T, D-2-T, D-5-T peg-like setae, D-4-T elongate seta, D-3-T campaniform sensillum; D-1-T, D-2-T on single tubercle, D-3-T anterolateral to D-4-T; supraalar 2—campaniform sensillum; metathoracics ( Fig. 51E )—1 campaniform sensillum; M-3-T distant from margin of metathorax (at least 1/3 length of metathorax). Abdomen: pigmentation light brown, poorly defined in some; with tergite 1 with 3 medial spots, tergites 2–7 with medial area with stripe, 2 anterolateral spots, some with 1–2 pairs on segment tergite 8, sternites 3–7 (anterolateral spots on 5–7 poorly defined, pale) all light brown, or just very light medial strip on sternites 4, 5 and with 6–7 membranous or just very light medial strip on sternites 3, 4 and with sternites 5–7 membranous or absent, segment 2 as wide or slightly wider than segment 3, segments with undivided, peg-like or thin to thick setae, with rounded to pointed, short to moderately elongate tubercles, with sternites 5–7 or 6–7 each with membranous disc; segment 9 ( Figs. 76C–D ) not strongly modified, terminal processes closely approximated basally, each projecting posterodorsolaterally, tapering to pointed apex; sensilla: tergite 1 ( Fig. 51E ) with 8 setae, 2 campaniform sensilla, including 3 lateral sensilla, D-2-I, D-3-I well separated, D-7-I situated anteriorly near D-3-I; segment 4 ( Fig. 66A )—D-2-IV, D-3-IV moderately elongate setae, D-2-IV on bifid tubercle, D-3-IV on short tubercle; D-5-IV peg-like seta, D-8-IV short seta, D-9-IV moderately elongate seta; 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 short seta on pointed tubercle, just anterior of base of tubercle with L-2-IV; L-2-IV, L-3-IV, L-4-IV short setae on short tubercles, V-5-IV, V-7-IV short setae, V-6-IV elongate seta, on short or moderately elongate tubercles, all closely approximated; segment 8 without D-3-VIII, without L-1-VIII; segment 9 ( Figs. 76 C-D)—with D-5-IX, D- 6-IX campaniform sensilla.

DISTRIBUTION AND HABITAT: The genus Mallochohelea is known from 48 species from every Region worldwide ( Borkent 2014 , with some moved to Anebomyia and Dibezzia ). Immatures have been reared from stream and river margins, thermal stream, reservoirs, ponds, and lakes.

TAXONOMIC DISCUSSION: The pupae of 12 species of Mallochohelea have been described ( Tables 2–3 ). Wirth & Grogan (1979) provide a key to the pupae of the two species of Mallochohelea from the Potomac Valley (Washington DC area, USA), as well as Anebomyia atripes , then called M. atripes . Thienemann (1928), Mayer (1934a) and Lenz (1934) provided keys to a few European species known at that time (as members of either Dicrobezzia (now a synonym of Probezzia ) or Johannsenomyia ). However, they all recognized M. dentata as distinct from M. munda (the former now considered a synonym of the latter) and both Thienemann (1928) and Mayer (1934a) additionally recognized M. breviforceps as distinct and it too is now considered a synonym of M. munda .

MATERIAL EXAMINED: M. albibasis : 1 pupal exuviae, Cottonwood Lake, 5 km S of Nelson, British Columbia, Canada, 6-VII-2008 (CNCI); 2 pupal exuviae, 7 km S of Hope, British Columbia, Canada, 8-VIII-1985 (CNCI); 1 pupal exuviae, Outegamie County, Wisconsin, USA, 15-VI-1954 (USNM). M. albihalter : 1 pupal exuviae, Allegany State Park, New York, USA, 28-V-1963 (USNM). M. caudellii : 1 pupal exuviae (in glycerin), Bolean Lake, 6 km E of Falkland, BC, Canada, 12–13-VII-1989 (CNCI). M. errinae : 1 pupal exuviae (of holotype), Klein Yukskei, Johannesburg, Transvaal, South Africa, V-1939 (SAIM); 1 pupal exuviae, Mageliesberg Agricultural School, Transvaal, South Africa, 13-XI-1973 (NMSA). M. inermis : 2 pupal exuviae, Vrevo lake, Leningrad Province, Russia, 19-VI-1968 (ZIN); 1 pupal exuviae, no locality/date (ZSMC). M. satelles : 4 pupal exuviae (of paratype), Nattai River, Mittagong, New South Wales, Australia, 18-II-1969 (ANIC). M. setigera : 1 pupal exuviae, River Gruzskii Elanchik, Donetsk Province, Ukraine, 29-IV-1970 (ZIN). M. smithi : 4 pupal exuviae, Black Lake, Stanleyville, Ontario, Canada, 29-VI-1975 (WLGC); 1 pupal exuviae, Rideau River, Ottawa, Ontario, Canada, 29-V-1960 (USNM); 1 pupal exuviae, Snow Hill, Maryland, 19-V-1968 (USNM); 1 pupal exuviae, 10 mi SE of Middleberg, Lake Dunmore, Vermont, USA, 23-VI-1986 (CNCI). M. sybleae : 1 pupal exuviae (of paratype), Centerville, Humboldt County, California, USA, 13-VIII-1948 (USNM). M. termophila : 2 pupal exuviae (of paratype), Caimancito, Jujuy, Argentina, 12-VIII-1980 (MLPA). M. nr. caudellii : 1 pupal exuviae, 7 km S of Hope, British Columbia, Canada, 8-VIII-1985 (CNCI). M. nr. pullata: 3 pupal exuviae, 6 km NE of Falkland, British Columbia, Canada, 12-VII-1989 (CNCI). M. sp.: 1 pupal exuviae, Bear Creek, 0.5 mi SE of Karlsbad Springs, Ontario, Canada, 1967 (CNCI); 1 pupal exuviae, McCarrs Creek, New South Wales, Australia, 11-XI-1956 (ANIC).

Figures

FIGURE 2. Pupae in unusual habitats, preservation and relative sizes. A. Forcipomyia pupae, Rancho Grande, Venezuela; photo by Steve Marshall. B. Forcipomyia pupae under bark of log of Pinus sylvestris, Poland; photo by Marek W. Kozlowski. C. Pupae of Mallochohelea sp. on emergent plants at edge of Bolean Lk., 6 km NE Falkland, British Columbia. D. Palpomyia sp. showing mounting technique on pin, 8 km E. Sicamous, British Columbia. E. Permanent slide mount of dissected adult and pupal exuviae of Mallochohelea albibasis. F. Pupal exuviae, from left to right, of Baeodasymyia michaeli, Culicoides denticulatus and Sphaeromias longipennis showing size differences within the family. Total length of Baeodasymyia michaeli = 1.2 mm, of Sphaeromias = 7.3 mm.

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 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 27. Posterior portion of heads, in ventral view. A. Mallochohelea caudellii. B. Probezzia sabroskyi. C. Neobezzia fittkaui. D. Nilobezzia curticornis. E. Johannsenomyia argentata. F. Lanatomyia electra. G. Anebomyia atripes. H. Homohelea delanoe.

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 66. Abdominal segments four, in dorsal and ventral view (shagreen not shown). A. Mallochohelea nr. caudellii (from Hope, British Columbia, Canada). B. Probezzia sabroskyi. C. Neobezzia fittkaui (from Ronderos et al. 2011, modified).

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.