Echinohelea Macfie

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 : 74-75

Echinohelea Macfie

( Figs. 20 , 26A , 30F , 31K , 37C , 44T , 50C , 63A , 75B )

DIAGNOSIS: Only pupa of Ceratopogonidae with abdominal segment 4 with only two lateral sensilla (L-2-IV, L- 3-IV) and with most sensilla at the bases of bifid tubercles ( Fig. 63A ).

DESCRIPTION: Total length = 2.91 mm. Without larval exuviae retained on abdomen. Exuviae with flagellum appressed against lateral margin of face, midleg, wing (as in Figs. 16B , 33B ). Ecdysial tear uncertain but not around base of antenna; at least medial to antennal base (as in Figs. 15B , 79D ). Head: Dorsal apotome missing; dorsolateral cephalic sclerite (as in Fig. 13H ) fused to scutum, each side uncertain in whole pupa; mouthparts ( Fig. 26A ) with mandible well-developed, lacinia absent; palpus extending posterior to posterolateral margin of labium; labium separated medially by labrum, hypopharynx; apex of antenna ( Fig. 37C ) anterior to posterior extent of midlength portion of midleg (portion lateral to mesosternum), narrowed posteriorly; sensilla: dorsal apotomals ( Fig. 20 )—uncertain; dorsolateral cephalic sclerite sensilla—1 elongate seta, 1 campaniform sensillum; clypeallabrals ( Fig. 26A )—1 small seta; oculars ( Fig. 26A )—1 campaniform sensillum. Thorax: Prothoracic extension ( Fig. 26A ) wide, well-developed, extending from palpus to antenna; mesonotum with short tubercles, extending posteriorly but medial division of metathorax uncertain ( Fig. 50C ); respiratory organ ( Fig. 44T ) length/width = 4.71, elongate, slender, somewhat flattened, at least apically (perhaps more), 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 moderately elongate, tracheal tube straight to slightly curved along length, surface smooth; wing ( Fig. 37C ) with apical tubercle lateral to apex of hind leg, separated medially by fore-, midlegs; halter apex and hind leg (as in Fig. 33A ) narrowly abutting; halter apex abutting anterolateral knob-like extension of tergite 2; legs ( Fig. 37C ) with lateral margin of foreleg near midlength of wing evenly curved; hind leg visible at lateral margin of wing (as in Fig. 32L ); with apex of foreleg slightly anterior to apex of midleg; apex of hind leg abutting apex of midleg laterally; sensilla: anteromedials—2 elongate setae, 1 campaniform sensillum ( Fig. 31K ); anterolaterals—1 moderately long seta; dorsal setae ( Fig. 30F )—D-1- T, D-2-T, D-4-T, D-5-T setae, D-3-T campaniform sensillum; D-1-T, D-2-T on single tubercle, D-3-T posteromedial to D-4-T; supraalar 2—campaniform sensillum; metathoracics ( Fig. 50C )—1 seta on one side only, 2 campaniform sensilla on both sides; M-3-T near anterior margin of metathorax. Abdomen: without pigmentation pattern, segment 2 as wide or slightly wider than segment 3, segments with undivided, thin to thick setae, with rounded to bilobed, short to somewhat elongate tubercles, tergites or sternites entire, each without membranous disc; segment 9 ( Fig. 75B ) not strongly modified, terminal processes closely approximated basally, each projecting posterodorsolaterally, tapering to pointed apex; sensilla: tergite 1 ( Fig. 50C ) with 7 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. 63A )—D-2-IV short seta on bifid tubercle; D-3-IV moderately elongate seta without tubercle; D-5-IV, D-8-IV, D- 9-IV short setae; at base on bifid, separate 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 situated anterorlaterally near D-3-IV; L-1-IV absent, L-2-IV, L-3-IV short setae each at base of bifid tubercle, L-4-IV absent, V-5-IV, V-6-IV, V-7-IV moderately elongate setae, at base of bifid, slender tubercles, V-6-IV, V-7-IV closely approximated; segment 8 without D-3-VIII, without L-1- VIII; segment 9 ( Fig. 75B )—with D-5-IX, D-6-IX campaniform sensilla.

DISTRIBUTION AND HABITAT: The genus Echinohelea is known from 27 species from nearly every Region worldwide but is absent from the Palaearctic Region ( Borkent 2014). An unpublished record from Borneo is known (pers. obs.). The only known immature, the pupa of E. lanei described here, was collected from Flanders, New York, USA. The Flanders Pond site was scrubby red maple growth flooded with a few inches of clear water (Hugo Jamnback, pers. comm.). Grogan (1975) reported a reared specimen of E. lanei collected from under bark of a rotting log in Maryland, USA and Knausenberger (1987) found this species in damp dark brown mud (his collection No. 29) in a roadside ditch in Virginia, USA. De Meillon & Wirth (1991) noted two African species reared from wet soil and from mud at a pond margin.

TAXONOMIC DISCUSSION: Only one moderately well preserved pupa of Echinohelea is known, that of E. lanei ( Tables 2–3 ). Wirth (1994b) provided an earlier description. Details of the sensilla distribution of abdominal segment 4 differs here because the specimen was distorted by compression and is reconstructed here. The dorsal apotome of the specimen was not present and cannot, therefore, be characterized.

Wirth (1994b) noted that the pupa of Echinohelea was similar to some Heteromyiini (as described by Elson-Harris & Kettle 1986) and this is reflected in the phylogenetic relationship supported here (and Borkent 1995) between Echinohelea and Heteromyiini + Sphaeromiini s. lat. + Palpomyiini + Stenoxenini .

MATERIAL EXAMINED: E. lanei : 1 pupal exuviae, Flanders Pond, Flanders, New York, USA, 4-VI-1963 (NYSM).

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 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 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 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 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 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 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.