Alluaudomyia 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 : 57-58

Alluaudomyia Kieffer

( Figs. 13E , 19L–M , 24E , 29P , 32J , 35D , 43S–T , 44A–B , 48E , 58C , 73L )

DIAGNOSIS: Only pupa of Ceratopogonidae with the respiratory organ with two closely appressed or approximated rows of pores and with each pore oblong in shape ( Figs. 43S–T , 44A–B ). Also only pupae with the abdominal segment 4 sensilla D-4-IV and D-7-IV placed on either side of D-8-IV ( Fig. 58C ) (in some species D-7- IV is placed more laterally, closer to D-9-IV). Also only pupa with the prothoracic extension abutting the antenna but not extending to the palpus ( Fig. 24E ) and the halter extending well posterior of the anterior margin of tergite 2 ( Fig. 32J ).

DESCRIPTION: Total length = 1.81–2.75 mm. Without larval exuviae retained on abdomen. Exuviae with flagellum appressed against lateral margin of face, in some against wing (as in Figs. 15D , 32J ). Ecdysial tear medial to base of antenna or to posterior to base (as in Figs. 15B, D , 79D, E ); along prothoracic extension. Head: Dorsal apotome ( Figs. 19L–M ), with ventral line of weakness, without dorsomedial tubercle, without central dome; dorsolateral cephalic sclerite ( Fig. 13E ) separated from scutum by thin cuticle, separate from scutum upon emergence, each side separated medially by dorsal apotome in whole pupa; mouthparts ( Fig. 24E ) with mandible well-developed, lacinia absent; palpus extending posterior to posterolateral margin of labium; labium separated medially by labrum, hypopharynx; apex of antenna ( Fig. 35D ) anterior to posterior extent of midlength portion of midleg (portion lateral to mesosternum); sensilla: dorsal apotomals ( Figs. 19L–M )—1 short to moderate seta, 1 campaniform sensillum; dorsolateral cephalic sclerite sensilla—1 seta, 1 campaniform sensillum; clypeal-labrals ( Fig. 24E )—absent; oculars ( Fig. 24E )—2 setae, 2 campaniform sensilla. Thorax: Prothoracic extension ( Fig. 24E ) short, present only dorsolaterally abutting antenna, not extending to palpus; mesonotum with very short tubercles, not extending posteromedially or with slight protuberance, not dividing metathorax medially ( Fig. 48E ); respiratory organ ( Figs. 43S–T , 44A–B ) length/width = 3.56–7.20, elongate, slender to moderately thick, somewhat flattened dorsoventrally, with pores closely abutting at apex of respiratory organ, arranged in two closely appressed rows, with or without additional, more basal pores, outer surface with (nearly all species) or without spicules, spines or pointed plates, without pedicel, base without posteromedial apodeme, membranous base of respiratory organ short, tracheal tube straight to slightly curved along length, with spirals restricted to base, distally with plate; wing ( Fig. 35D ) with apical tubercle lateral to apex of hind leg, separated medially by fore-, midlegs; halter apex and hind leg ( Fig. 32J ) broadly abutting; halter apex extending posteriorly to 1/3 length of tergite 2; legs ( Fig. 35D ) with lateral margin of foreleg near midlength of wing evenly curved; hind leg visible at lateral margin of wing ( Fig. 32J ); with apex of foreleg moderately anterior to apex of midleg; apex of hind leg slightly dorsal to, partially abutting apex of midleg laterally; sensilla: anteromedials—2 setae; anterolaterals—3 setae; dorsal setae ( Fig. 29P )—D-1-T, D-2-T, D-4-T, D-5-T setae, D-3-T campaniform sensillum, D-3-T posteromedial to D-4-T; supraalar 2—campaniform sensillum; metathoracics ( Fig. 48E )—1 seta, 2 campaniform sensilla; M-3-T near anterior margin of metathorax. Abdomen: without pigmentation pattern or anteromedial margin of tergites 1–4 (fading posteriorly) or tergites 1–5 (tergite 1 with 4 medial), just medial on tergite 6, fading posteriorly, segment 2 as wide or slightly wider than segment 3, segments with undivided, thin to thick setae, with rounded to pointed, short tubercles, tergites or sternites entire, each without membranous disc; segment 9 ( Fig. 73L ) not strongly modified, terminal processes closely approximated basally, each projecting posterodorsolaterally, tapering to pointed apex; sensilla: tergite 1 ( Fig. 48E ) with 7–8 setae, 2 campaniform sensilla, including 3 lateral sensilla, D-2-I, D-3-I closely approximated, D-7-I situated posteriorly near D-8-I; segment 4 ( Fig. 58C )—D-2-IV, D-3-IV moderately elongate setae on separate tubercles; D-5-IV, D-8-IV, D-9-IV short setae, on short to moderately elongate, separate tubercles, posterior dorsal sensilla in transverse row, arranged medially to laterally: D-5-IV, D-4-IV, D-8-IV, D-7- IV, D-9-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 pointed tubercles, V-5-IV, V-6-IV, V-7-IV short to moderately elongate setae on pointed tubercles; segment 8 without D-3-VIII, without L-1-VIII; segment 9 ( Fig. 73L )—with D-5-IX, D-6-IX campaniform sensilla.

DISTRIBUTION AND HABITAT: The genus Alluaudomyia is known from 203 species from every Region worldwide ( Borkent 2014 ). Immatures have been sampled from ponds, bogs, fens, swamps, treeholes and margins of streams, creeks, rivers and lakes. Some species have been reared from wet leaves and mud associated with the aforementioned habitats.

TAXONOMIC DISCUSSION: There are 23 species of Alluaudomyia known as pupae ( Tables 2–3 ). Considering the good visibility of the larvae, which swim on the surface film, there are good prospects of rearing many additional species of this genus.

The dorsal apotomes of A. bella , A. caribbeana , A. distispinulosa , A. footei , A. megaparamera , A. paraspina , A. parv a, A. reyei and A. unguistyla have been illustrated without campaniform sensillum DA-2-H ( Elson-Harris & Kettle 1985b, Grogan & Bystrak 1976, Spinelli 1997, Spinelli & Wirth 1984, Wirth & Grogan 1981), but examination of original material of A. bella , A. distispinulosa , A. footei , A. megaparamera , A. paraspina and A. parv a indicates that it is actually present in at least these species. The campaniform sensillum has likely been overlooked in the other species considering that all species of Alluaudomyia I examined had the sensillum present among the similarly sized spicules and that the sensillum is present in most other Ceratopogonidae .

De Meillon (1939) illustrated abdominal segment 5 of A. maculithorax but misidentified the dorsal and ventral surfaces.

The distinctive placement of fourth abdominal segment sensilla D-7-IV lateral to D-8-IV has not been reported or illustrated by those previous authors describing the sensilla of the abdominal segments, likely because this sensillum coeloconica is small, difficult to discern and was overlooked ( de Meillon 1939, Debenham 1971, Elson-Harris & Kettle 1985b); this is certainly the case for the three species described by Debenham (1971) and which were reexamined here.

MATERIAL EXAMINED: A. appendiculata : 2 pupal exuviae (of paratype), Hornsby, New South Wales, Australia, 9-X-1956 (ANIC); 1 pupal exuviae, as previous locality, 18-XI-1969 (ANIC); 2 pupal exuviae, Nattai River, Mittagong, New South Wales, Australia, 25-X-1968 (ANIC). A. bella : 1 pupal exuviae, 6.5 km NW of Enderby, British Columbia, Canada, 27-VI-1990 (CNCI); 1 pupal exuviae, 20 km E of Anola, Manitoba, Canada, 16-VI-1990 (CNCI); 1 pupal exuviae (in glycerin), Charleston Provincial Park, Ontario, Canada, 18-V-1986 (CNCI); 1 pupal exuviae, Algonquin Park, Ontario, Canada, 7-VI-1960 (USNM); 6 pupal exuviae, Bat Lake, Algonquin Park, Ontario, Canada, 7-VI-1960 (USNM); 1 pupal exuviae, Notre Dame de Laus, Lac Serpent, Quebec, Canada, 22-VI-1967 (USNM); 1 pupal exuviae, as previous locality, 11-VII-1967 (USNM); 1 pupal exuviae, Patuxent Wildlife Rescue Center, Prince George’s County, Maryland, USA, 19-IV-1976 (WLGC); 2 pupal exuviae, as previous locality, 8-V-1958 (USNM); 1 pupal exuviae, as previous locality, 28-VI-1976 (WLGC); 1 pupal exuviae, as previous locality, 17-V-1976 (WLGC); 3 pupal exuviae, Lake Jimmy, Tahawus, New York, USA, 24-VI-1959 (NYSM); 1 pupal exuviae, Hamilton-Essex, Newcomb, New York, USA, 23-VII-1959 (NYSM); 1 pupal exuviae, Huntington Swamp, Newcomb, New York, USA, 4-VIII-1959 (NYSM); 1 pupal exuviae, Salmon River, Blue Mountain Lake, New York, USA, 23-VI-1959 (NYSM); 1 pupal exuviae, Allegany State Park, New York, USA, 28-V-1963 (USNM); 1 pupal exuviae, Upper Pansapoulas Pond, Montgomery County, Virginia, USA, 3-IX-1976 (VPIC); 2 pupal exuviae, Upper Pandapas Pond, Montgomery County, Virginia, USA, 2-VIII-1976 (VPIC). A. bicornis : 1 pupal exuviae, Manton River, Northern Territory, Australia, 6-VI-1958 (ANIC). A. biestroi : 1 pupal exuviae (of paratype), Artigas A “Lenguazo” y Ruta 3, Uruguay, 8-XI-1984 (MLPA). A. claudia : 1 pupal exuviae (of paratype) Salisbury, Southern Rhodesia, South Africa, 24-II-1949 (SAIM). A. distispinulosa : 1 pupal exuviae, Bulo Bulo, Cochabamba, Bolivia, 12-I-1995 (MLPA). A. fragmentum : 1 pupal exuviae (of paratype), Manton River, Northern Territory, Australia, 6-VI-1958 (ANIC); 1 pupal exuviae (of paratype), as previous locality, 1-VI-1958 (ANIC). A. latipennis : 1 pupal exuviae, Hornsby, New South Wales, Australia, 9-X-1956 (ANIC); 1 pupal exuviae, Nattai River, Mittagong, New South Wales, Australia, 4-XI-1964 (ANIC); 1 pupal exuviae, as previous locality, 15-XI-1968 (ANIC). A. megaparamera : 1 pupal exuviae, Blue Mountain Lake, New York, USA, 29-VI-1959 (NYSM); 1 pupal exuviae, Salmon River, Blue Mountain Lake, New York, USA, 8-VI- 1959 (NYSM); 1 pupal exuviae, Hamilton-Essex, Newcomb, New York, USA, 1-VI-1959 (NYSM); 1 pupal exuviae, as previous locality, 8-VI-? (NYSM). A. natalensis : 1 pupal exuviae, Empangeni, Zululand, South Africa, 16-VII-1938 (SAIM); 1 pupal exuviae, 15 mi. SE of Potgietersrus, Transvaal, South Africa, 3-I-1974 (NMSA). A. needhami : 1 pupal exuviae, Salmon Arm, British Columbia, Canada, 11-VII-1990 (CNCI); 3 pupal exuviae, 30 km N of Nakusp, British Columbia, Canada, 5-VII-1990 (CNCI); 2 pupal exuviae, Patuxent Wildlife Refuge, Prince George’s County, Maryland, USA, 1-VIII-1979 (USNM); 1 pupal exuviae, Upper Pandapas Pond, W.A., Montgomery County, Virginia, USA (VPIC); 1 pupal exuviae, 1.7 mi E of Lunter’s gate, Rt. 785, Montgomery County, Virginia, USA, 22-IV-1976 (VPIC); 1 pupal exuviae, no locality, 10-VI-1953 (USNM); 2 pupal exuviae, no locality, 20-II-1957 (USNM). A. needhami or A. megaparamera : 2 pupal exuviae, Montvale Wayside, Bedford County, Virginia, USA, 16-VI-1977 (VPIC)); 1 pupal exuviae, Beaver Lake Reservoir, Pocahontas State Park, Chesterfield County, Virginia, USA, 30-IV-1977 (VPIC). A. paraspina : 2 pupal exuviae, Lakeland Pond, College Park, Prince George’s County, Maryland, USA, 26-V-1975 (WLGC); 2 pupal exuviae, as previous locality, 26-V- 1975 (WLGC); 5 pupal exuviae, as previous locality, 30-V-1975 (WLGC); 1 pupal exuviae, Mountain Lake Biological Station, Giles County, Virginia, USA (VPIC). A. parva : 1 pupal exuviae, Blue Mountain Lake, New York, USA, 30-VI-1958 (NYSM); 1 pupal exuviae, Salmon River, Blue Mountain Lake, New York, USA, 14-V- 1959 (NYSM); 1 pupal exuviae, Allegany State Park, New York, USA, 28-V-1963 (USNM); 7 pupal exuviae, Alexandria, Virginia, USA, 6-VI-1951 (USNM). A. quadripunctata : 2 pupal exuviae, Tobolki lake, Alol’, Pskov Province, Russia, 30-VI-1969 (ZIN). A. schnacki : 2 pupal exuviae (of paratype), Los Talas (Berisso), Buenos Aires, Argentina, 29-XI-1979 (MLPA). A. splendida : 1 pupal exuviae, Ussuri Nature Reserve, Primorskii Territory, Russia, 12-VI-1973 (ZIN). A. variegata : 1 pupal exuviae, Lake Muson, Leon County, Florida, USA, 22-VIII-1987 ( JHEC). A. sp.: 1 pupal exuviae, Bolean Lake, 6 km NE of Falkland, British Columbia, Canada, 1-VII-2009 (CNCI); 1 pupal exuviae, 30 km N of Nakusp, British Columbia, Canada, 5-VII-1990 (CNCI); 1 pupal exuviae, 5 km S of Cottonwood Lake, Nelson, British Columbia, Canada, 6-VII-2008 (CNCI); 1 pupal exuviae, Meachum Lake, New York, USA, 21-VI-1986 (CNCI); 1 pupal exuviae, Ackerman Swamp, Newcomb, New York, USA, 7- VII-1958 (NYSM); 1 pupal exuviae, Moung Sayaboury, Sayaboury Province, Laos, 19-II-1967 (BPBM); 1 pupal exuviae, as previous locality, 2-XII-1967 (BPBM); 2 pupal exuviae, as previous locality, 22-II-1967 (BPBM); 1 pupal exuviae, Houay La stream, 20 kilometers NE of Moung Sayaboury, Sayaboury Province, Laos, 25-XI-1967 (BPBM); 1 pupal exuviae, Moung Sayaboury, Sayaboury Province, Laos, 9-XII-1967 (BPBM); 1 pupal exuviae, as previous locality, 7-I-1968 (BPBM); 1 pupal exuviae, Nam Houng River, Moung Sayaboury, Sayaboury Province, Laos, 17-XII-1967 (BPBM); 1 pupal exuviae, Gnieraoora Pool, Robe River, Pilbara, Western Australia, Australia (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 15. Anterior portion of heads. A. Culicoides denticulatus, in anteroventral view. B. Ceratopogon nr. abstrusus, in anterior view. C. Brachypogon sp., in anterior view (from 6 km E. Falkland, BC, Canada). D. Stilobezzia antennalis, in ventral view, dorsal apotome not present. E. Allohelea japonica, in ventral view, dorsal apotome not present (distorted slide specimen). F. Atyphohelea macroneura, in ventral view, right side of head, dorsal apotome not present.

FIGURE 19. Dorsal apotomes, anterior view. A. Culicoides circumscriptus (from Kettle & Lawson 1952). B. Culicoides cubitalis (from Kettle & Lawson 1952). C. Culicoides truncorum (from Kettle & Lawson 1952). D. Culicoides chiopterus (from Jamnback 1965). E. Paradasyhelea minuta (from Kettle & Elson 1975a). F. Paradasyhelea albipunctata (from Elson- Harris & Kettle 1985a). G. Paradasyhelea reyei (from Elson-Harris & Kettle 1985a). H. Ceratopogon boomerangus (from Borkent & Grogan 1995). I. Ceratopogon abstrusus (from Borkent & Grogan 1995). J. Ceratopogon arcanus (from Borkent & Grogan 1995). K. Brachypogon artemis (from Debenham 1991). L. Alluaudomyia appendiculata (from Elson-Harris & Kettle 1985b). M. Alluaudomyia fumosipennis (from Elson-Harris & Kettle 1985b). N. Baeodasymyia michaeli (from Borkent & Craig 1999). O. Austrohelea campbellensis. P. Stilobezzia rabelloi (from Borkent & Craig 2001).

FIGURE 24. Posterior portion of heads, in ventral view. A. Culicoides guttipennis. B. Paradasyhelea albipunctata. C. Ceratopogon arcanus. D. Brachypogon sp. (Flag Glade, Pocahontas Co., West Virginia, USA). E. Alluaudomyia needhami. F. Baeodasymyia michaeli. G. Austrohelea campbellensis.

FIGURE 29. Dorsal setae from left side of thoraces. A. Austroconops mcmillani. B. Leptoconops kerteszi. C. Leptoconops linleyi. D. Forcipomyia (Euprojoannisia) sp. (from Peru). E. Forcipomyia (Forcipomyia) bipunctata. F. Forcipomyia (Phytohelea) n. sp. nr. antiguensis. G. Atrichopogon sp. (from Lake Opinicon, Ontario, Canada). H. Dasyhelea paracincta. I. Dasyhelea atlantis. J. Dasyhelea pseudoincisurata. K. Culicoides haematopotus. L. Culicoides phlebotomus. M. Paradasyhelea albipunctata. N. Ceratopogon boomerangus. O. Brachypogon sp. (from 6 km E. Falkland, BC, Canada). P. Alluaudomyia needhami. Q. Baeodasymyia christopheri (from Borkent & Craig 1999). R. Austrohelea campbellensis. S. Stilobezzia antennalis. T. Schizonyxhelea forattinii (from Borkent 2000a).

FIGURE 32. Cephalothoraces and abdominal segments 1–3, in lateral view. A. Austroconops mcmillani. B. Leptoconops linleyi. C. Forcipomyia (Phytohelea) sp. (Florida, USA). D. Atrichopogon sp. (from 6.5 km NW Enderby, BC, Canada). E. Dasyhelea sp., in anteroventral view (Herman Lake, 4 km SE Eagle Bay, BC, Canada). F. Dasyhelea oppresa. G. Culicoides denticulatus. H. Ceratopogon nr. abstrusus. I. Brachypogon sp. (from 6 km E. Falkland, BC, Canada). J. Alluaudomyia bella. K. Baeodasymyia michaeli. L. Stilobezzia antennalis.

FIGURE 35. Legs, wings and apices of the antennae, in ventral view. A. Paradasyhelea sp. (from Borlase Stream, Lake Rotoiti, New Zealand). B. Ceratopogon sp. nr. abstrusus. C. Brachypogon sp. (from Spanish Lake, 6 km E. Falkland, BC, Canada). D. Alluaudomyia needhami. E. Baeodasymyia michaeli. F. Austrohelea campbellensis (reconstructed from distorted specimen).

FIGURE 43. Respiratory organs, most in dorsal view, Fig. 43T in lateral view, pattern on tracheal tubes shown only in some. A. Dasyhelea pseudoincisurata (from Waugh & Wirth 1976). B. Dasyhelea grisea (from Waugh & Wirth 1976). C. Dasyhelea traverae (from Waugh & Wirth 1976). D. Dasyhelea pseudocincta (from Waugh & Wirth 1976). E. Dasyhelea leptobranchia (from Waugh & Wirth 1976). F. Dasyhelea brevicornis (from Waugh & Wirth 1976). G. Dasyhelea hitchcocki (from Wirth 1976a). H. Culicoides chiopterus (from Jamnback 1965). I. Culicoides furens (from Jamnback 1965). J. Culicoides guttipennis (from Jamnback 1965). K. Culicoides dewulfi (from Kettle & Lawson 1952, as C. pseudochiopterus). L. Culicoides punctatus (from Kettle & Lawson 1952). M. Culicoides sanguisuga (from Jamnback 1965). N. Culicoides variipennis (from Jamnback 1965). O. Paradasyhelea albipunctata (from Elson-Harris & Kettle 1985a). P. Ceratopogon inverecundus (from Borkent & Grogan 1995). Q. Ceratopogon willisi (from Borkent & Grogan 1995). R. Brachypogon sp. (from Virginia, USA). S. Alluaudomyia needhami (from Wirth & Grogan 1981). T. Alluaudomyia fumosipennis (from Debenham 1971).

FIGURE 44. Respiratory organs, in dorsal view, pattern on tracheal tubes shown only in some. A. Alluaudomyia latipennis (from Debenham 1971). B. Alluaudomyia footei (from Wirth & Grogan 1981). C. Baeodasymyia christopheri (from Borkent & Craig 1999). D. Austrohelea campbellensis. E. Stilobezzia antennalis (from Wirth & Grogan 1981, modified). F. Stilobezzia glauca (from Wirth & Grogan 1981). G. Stilobezzia ochracea (from Kettle & Lawson 1952). H. Stilobezzia diversa (from Wirth & Grogan 1981). I. Stilobezzia rabelloi (from Borkent & Craig 2001). J. Stilobezzia navaiae (from Wirth & Grogan 1981). K. Stilobezzia lutea (from Wirth & Grogan 1981). L. Schizonyxhelea bulla (from Wirth & Grogan 1981, as a Stilobezzia). M. Schizonyxhelea forattinii (from Borkent 2000a). N. Serromyia femorata (from Kettle & Lawson 1952). O. Allohelea japonica. P.Monohelea hirsuta (from Wirth & Grogan 1981, modified). Q. Atyphohelea macroneura (from Borkent 1998). R. Parabezzia alexanderi (from Grogan & Wirth 1977a). S. Parabezzia huberti (from Grogan & Wirth 1977a). T. Echinohelea lanei (from Wirth 1994b).

FIGURE 48. Metathoraces and tergites 1, in dorsal view. A. Culicoides haematopotus. B. Paradasyhelea minuta. C. Ceratopogon sp. nr. abstrusus. D. Brachypogon sp. (from 6 km E. Falkland, BC, Canada). E. Alluaudomyia needhami. F. Baeodasymyia michaeli.

FIGURE 58. Abdominal segments four, in dorsal and ventral view (shagreen not shown). A. Ceratopogon nr. inverecundus. B. Brachypogon sp. (from 6 km E. Falkland, BC, Canada). C. Alluaudomyia needhami.

FIGURE 73. Abdominal segments 9. A. Dasyhelea laeta, female, in ventral view (from Mayer 1934c). B. Dasyhelea assimilis, male, in lateral view with ventral to the right (from Mayer 1934c). C. Dasyhelea assimilis, male, in lateral view with ventral to the right (from Mayer 1934c). D. Dasyhelea grisea, male, in dorsal view (from Waugh & Wirth 1976). E. Dasyhelea chani, female, in dorsal view (from Wirth & Linley 1990). F. Dasyhelea atlantis, female, in ventral view (from Waugh & Wirth 1976). G. Dasyhelea gluchovae, female, in ventral view (from Brodskaya 1996). H. Culicoides haematopotus, female, in dorsal and ventral view (shagreen not shown). I. Paradasyhelea albipunctata, female, in dorsal view (Elson-Harris & Kettle 1985a). J. Ceratopogon boomerangus, female, in ventral view (modified from Borkent & Grogan 1995). K. Brachypogon artemis, female, in dorsal view (from Debenham 1991). L. Alluaudomyia appendiculata, female, in dorsal view (from Elson-Harris & Kettle 1985b).

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.