Austrohelea Wirth & Grogan

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 : 60-61

Austrohelea Wirth & Grogan

( Figs. 19O , 24G , 29R , 35F , 44D , 49A , 59B , 74B )

DIAGNOSIS: Only pupa of Ceratopogonidae with the apex of the hind leg ventral to the apex of the midleg ( Fig. 35F ) (only the female is known; perhaps the male, without the elongate hind leg claw of the developing adult will not have this feature).

DESCRIPTION: Total length = 2.50 mm. Without larval exuviae retained on abdomen. Exuviae with flagellum appressed against lateral margin of face (as in Fig. 15D ). Ecdysial tear around base of antenna (as in Figs. 15D , 79E ); along prothoracic extension. Head: Dorsal apotome ( Fig. 19O ), without ventral line of weakness, without dorsomedial tubercle, without central dome; dorsolateral cephalic sclerite (as in Fig. 13B ) separated from scutum by thin cuticle, separate from scutum upon emergence, each side separated medially by dorsal apotome in whole pupa; mouthparts ( Fig. 24G ) 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. 35F ) anterior to posterior extent of midlength portion of midleg (portion lateral to mesosternum); sensilla: dorsal apotomals ( Fig. 19O )—1 moderately elongate seta, 1 campaniform sensillum; dorsolateral cephalic sclerite sensilla—1 seta, no campaniform sensillum; clypeal-labrals ( Fig. 24G )—2 elongate setae; oculars ( Fig. 24G )—2 setae, 1 campaniform sensillum. Thorax: Prothoracic extension ( Fig. 24G ) wide, well-developed, extending from palpus to antenna; mesonotum with short tubercles, extending posteromedially, completely dividing metathorax medially ( Fig. 49A ); respiratory organ ( Fig. 44D ) length/width = 6.25–6.25, elongate, slender, somewhat flattened dorsoventrally, with pores closely abutting at apex of respiratory organ, arranged in single row, with additional more basal pores, outer surface smooth, with moderately elongate pedicel, base with short 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. 35F ) with slight angle lateral to apex of hind leg, 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 ( Fig. 35F ) 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 ventral to apex of midleg, apex of each hind leg nearly abutting apically and abutting apices of foreleg; sensilla: anteromedials—2 setae; anterolaterals—3 setae; dorsal setae ( Fig. 29R )—D-1- T, D-2-T, D-4-T setae, D-3-T campaniform sensillum, D-5-T absent, D-3-T posteromedial to D-4-T; supraalar 2—campaniform sensillum; metathoracics ( Fig. 49A )—1 seta, 2 campaniform sensilla; M-3-T near anterior margin of metathorax. Abdomen: with anterior margins of tergites 3–7, anterior margins of sternites 3–7 pigmented, segment 2 as wide or slightly wider than segment 3, segments with undivided, thin to thick setae, with bilobed or rounded, short tubercles, tergites or sternites entire, each without membranous disc; segment 9 ( Fig. 74B ) not strongly modified, terminal processes widely separated basally, each projecting posterodorsolaterally, tapering to pointed apex; sensilla: tergite 1 ( Fig. 49A ) with 6 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. 59B )—D-2-IV, D-3-IV short or moderately elongate setae on separate tubercles; D-5-IV, D-8-IV short seta, D-9-IV absent; on short bilobed tubercles, posterior dorsal sensilla in transverse row, arranged medially to laterally: D-5-IV, D-4-IV, D-7-IV, D-8- IV; L-1-IV 1 seta on bilobed tubercle, well anterior of posterior lateral setae; L-2-IV, L-3-IV, L-4-IV short setae on bilobed tubercles, V-5-IV, V-6-IV, V-7-IV short setae on bilobed tubercles; segment 8 without D-3-VIII, without L- 1-VIII; segment 9 ( Fig. 74B )—with D-5-IX, D-6-IX campaniform sensilla.

DISTRIBUTION AND HABITAT: The genus Austrohelea is known from seven species from Australia, New Zealand and Argentina ( Borkent 2014 ). The only pupa known was reared from the subantarctic Campbell Island but the specific habitat was not recorded.

TAXONOMIC DISCUSSION: The unique pupal exuviae was previously described by Sublette & Wirth (1980) and reexamined here ( Tables 2–3 ).

MATERIAL EXAMINED: A. campbellensis : 1 pupal exuviae, Tucser Cove, Campbell Island, New Zealand, 22-I-1969 (NZAC).

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 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 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 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 49. Metathoraces and tergites 1, in dorsal view. A. Austrohelea campbellensis. B. Stilobezzia antennalis. C. Schizonyxhelea forattinii. D. Serromyia sp.. E. Allohelea japonica. F. Monohelea obscura.

FIGURE 59. Abdominal segments four, in dorsal and ventral view (shagreen not shown). A. Baeodasymyia christopheri (from Borkent & Craig 1999). B. Austrohelea campbellensis. C. Stilobezzia antennalis.

FIGURE 74. Abdominal segments 9. A. Baeodasymyia michaeli, female, in dorsal view (modified from Borkent & Craig 1999). B. Austrohelea campbellensis, female, in dorsal and ventral view (reconstructed from distorted specimen, shagreen not shown). C. Stilobezzia lutea, male, in dorsal and ventral view (from Wirth & Grogan 1981). D. Stilobezzia lutea, female, in dorsal and ventral view (from Wirth & Grogan 1981). E. Stilobezzia rabelloi, male, in ventral view (from Borkent & Craig 2001). F. Schizonyxhelea bulla, female, in dorsal and ventral view (from Wirth & Grogan 1981). G. Schizonyxhelea forattinii, female, in ventral view (from Borkent 2000a). H. Serromyia sp., female (in dorsal and ventral view (reconstructed from distorted specimen, shagreen not shown). I. Allohelea japonica, female, in dorsal and ventral view (shagreen not shown). J. Monohelea obscura, male, in dorsal and ventral view (reconstructed from distorted specimen, shagreen not shown). K. Atyphohelea macroneura, female, in dorsal view (from Borkent 1998). L. Parabezzia huberti, male, in ventral view (from Grogan & Wirth 1977a).

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.