Homohelea 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 : 95-96

Homohelea Kieffer

( Figs. 27H , 31A , 40A–B , 46A–B , 52F , 68B , 77A )

DIAGNOSIS: Only pupa of Ceratopogonidae with the abdominal tubercles all apically rounded ( Fig. 68B ), abdominal segment 4 with D-8-IV and D-9-IV on separate or, at most, only basally fused tubercles ( Fig. 68B ) and abdominal segment 8 with the two ventral sensilla (V-5-VIII, V-6-VIII) on a single tubercle and V-5-VIII tiny and V-6-VIII elongate.

DESCRIPTION: Total length = 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 around base of antenna, along lateral margin of face to palpus (as in Figs. 17C , 79H ). Head: Dorsal apotome as described by de Meillon & Wirth (1981), uncertain 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. 27H ) with mandible well-developed, lacinia absent; palpus extending equal to or just posterior to posterolateral margin of labium; labium separated medially by labrum, hypopharynx; apex of antenna ( Fig. 40A–B ) anterior to posterior extent of midlength portion of midleg (portion lateral to mesosternum), narrowed posteriorly; sensilla: dorsal apotomals (as described by de Meillon & Wirth 1981)—1 elongate seta, 1 campaniform sensillum; dorsolateral cephalic sclerite sensilla—1 seta, campaniform sensillum not visible (dirty specimens); clypeal-labrals ( Fig. 27H )—2 slender setae; oculars ( Fig. 27H )—2 setae, 1 campaniform sensillum. Thorax: Prothoracic extension ( Fig. 27H ) wide, well-developed but narrow dorsolaterally, not extending to antenna; mesonotum without tubercles, not extending posteromedially, not dividing metathorax medially ( Fig. 52F ); respiratory organ ( Figs. 46A–B ) length/width = 4.55–5.70, elongate, slender, somewhat flattened apically, with pores closely abutting at apex of respiratory organ, arranged in single row, outer surface smooth, with short, wide pedicel, base with elongate posteromedial apodeme, membranous base of respiratory organ short, annulated, tracheal tube straight to slightly curved along length, with spirals restricted to base, plates to half length; wing ( Fig. 40A–B ) with short tubercle at apex of hind leg, separated medially by fore-, midlegs; halter apex and hind leg (similar to Fig. 33J ) broadly abutting; halter apex extending posteriorly to 1/4 length of tergite 2; legs ( Fig. 40A–B ) with lateral margin of foreleg near midlength of wing evenly curved; hind leg visible at lateral margin of wing (as in Fig. 33I ); male with apex of foreleg moderately anterior to apex of midleg, female with apex of foreleg ventral 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 moderate seta; dorsal setae ( Fig. 31A )—D-1-T, D-2-T, D-4-T, D-5-T setae, D-3-T campaniform sensillum, D-3-T lateral to slightly posterolateral to D-4-T; supraalar 2—campaniform sensillum; metathoracics ( Fig. 52F )—1 campaniform sensillum; M-3-T distant from margin of metathorax (at least 1/3 length of metathorax). Abdomen: with tergite 1 uncertain, tergites 2-7 with medial area with stripe, 2 anterolateral spots, sternites 3–7 with anterolateral spots hard to see, 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. 77A ) not strongly modified, terminal processes closely approximated basally, each projecting posterodorsolaterally, tapering to pointed apex; sensilla: tergite 1 ( Fig. 52F ) with 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. 68B )—D-2-IV, D-3-IV moderately elongate setae, D-2-IV on short tubercle; 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 basally fused, flattened 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 arising from flat surface, moderately anterior of posterior lateral setae; L-2-IV, L-3-IV, L-4-IV short setae on rounded tubercles, those of L- 2-IV, L-4-IV flattened, V-5-IV, V-6-IV, V-7-IV short setae on rounded, flattened tubercles, all closely approximated; segment 8 without D-3-VIII, without L-1-VIII; with V-5-VIII, V-6-VIII on single tubercle, V-5-VIII tiny, V-6-VIII elongate; segment 9 ( Fig. 77A )—with D-5-IX, D-6-IX campaniform sensilla.

DISTRIBUTION AND HABITAT: The genus Homohelea is known from 19 species in the Palaearctic, Oriental, and Afrotropical Regions ( Borkent 2014 ). Pupae have been collected from a river margin, a brackish marsh, ponds, and ground pools. Ingram & Macfie (1921) noted pupae were common in "puddles of dirty water near a stand-pipe".

TAXONOMIC DISCUSSION: Three species of Homohelea are known as pupae ( Tables 2–3 ). The dorsal apotomes of the available specimens were absent and cannot, therefore, be characterized. The dorsal apotome of H. delanoe was described (but not illustrated) by de Meillon & Wirth (1981) as having "a pair of small tubercles each bearing a slender hair", which is the typical condition in most Ceratopogonidae .

In addition to the male specimens examined, photos were made available (Burgert Muller, NMSA) of the holotype female of H. albitudinis and a non-type female specimen of H. delanoe , allowing comparisons to be made of sexually dimorphic features with the male pupae at hand (see character 44).

MATERIAL EXAMINED: H. delanoe : 2 pupal exuviae, Zoutpan, Transvaal, South Africa, 10-I-1974 (USNM).

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 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 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 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 40. Legs, wings and apices of the antennae, in ventral view. A. Homohelea delanoe, male (reconstructed from distorted specimen). B. Homohelea albitudinis, female (drawn from photograph). C. Leehelea hispida, female. D. Sphaeromias longipennis, male. E. Sphaeromias longipennis, female. F. Xenohelea galatea, female (reconstructed from distorted specimen).

FIGURE 46. Respiratory organs, most in dorsal view, Figs. 46G. in lateral view, pattern on tracheal tubes shown only in some. A. Homohelea albitudinis (from de Meillon & Wirth 1981). B. Homohelea delanoe (from de Meillon & Wirth 1981). C. Leehelea wasselli (from Elson-Harris 1987). D. Leehelea hispida (from Elson-Harris 1987). E. Sphaeromias longipennis (from Wirth & Grogan 1979). F. Xenohelea galatea. G. Amerohelea sordidipes (from Lane et al. 1955). H. Bezzia glabra (from Wirth 1983b). I. Bezzia obelisca (from Wirth 1983a). J. Bezzia dorsasetula (from Wirth 1983a). K. Bezzia turkmenica (from Glukhova 1979b). L. Clastrieromyia dycei (from Spinelli & Grogan 1986). M. Pachyhelea pachymera (from Spinelli 1983a). N. Palpomyia lineata (from Grogan & Wirth 1979). O. Palpomyia jonesi (from Grogan & Wirth 1979). P. Palpomyia belkini (from Grogan & Wirth 1979). Q. Palpomyia rufa (from Grogan & Wirth 1979). R. Palpomyia flaviceps (from Grogan & Wirth 1979). S. Phaenobezzia opaca (from Wirth & Grogan 1982). T. Paryphoconus oliveirai (from Ronderos et al. 2007).

FIGURE 52. Metathoraces and tergites 1, in dorsal view. A. Neobezzia fittkaui. B. Nilobezzia curticornis. C. Johannsenomyia argentata. D. Lanatomyia miles. E. Anebomyia atripes. F. Homohelea delanoe (reconstructed from distorted and dirty specimen).

FIGURE 68. Abdominal segments four, in dorsal and ventral view (shagreen not shown). A. Anebomyia atripes. B. Homohelea delanoe. C. Leehelea hispida (reconstructed from distorted specimen).

FIGURE 77. Abdominal segments 9. A. Homohelea delanoe, female, in dorsal view (from de Meillon & Wirth 1981). B. Leehelea wasselli, female, in dorsal view (from Elson-Harris 1987). C. Sphaeromias longipennis, male, in dorsal and ventral view (from Wirth & Grogan 1979). D. Xenohelea galatea, female, in dorsal and ventral view (shagreen not shown). E. Bezzia roldani, male, in dorsal view (from Spinelli & Wirth 1989). F. Bezzia roldani, female, in dorsal view (from Spinelli & Wirth 1989). G. Bezzia brevicornis, male, in dorsal view (from Spinelli 1983a). H. Bezzia acanthodes, female, in dorsal view (from Spinelli 1983a). I. Clastrieromyia dycei, male, in ventral view (from Spinelli & Grogan 1986). J. Clastrieromyia dycei, female, in ventral view (from Spinelli & Grogan 1986). K. Pachyhelea pachymera, male, in ventral view (from Spinelli 1983a). L. Pachyhelea pachymera, female, in dorsal view (from Spinelli 1983a).

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.