Atyphohelea Borkent

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 : 71-72

Atyphohelea Borkent

( Figs. 15F , 20J , 25G , 30D , 37A , 44Q , 50A , 62A , 74K )

DIAGNOSIS: Only pupa of Ceratopogonidae with the thorax and abdomen with moderately developed rounded tubercles ( Figs. 30D , 62A , as in 33A), with three ventral setae (V-5-IV, V-6-IV, V-7-IV) on well-developed tubercles ( Fig. 62A ) (not as in Fig. 61A ) and the respiratory organ with pores restricted to its apex ( Fig. 44Q ).

DESCRIPTION: Total length = 2.20–2.94 mm. Without larval exuviae retained on abdomen. Exuviae with flagellum appressed against lateral margin of face, midleg, wing ( Fig. 15F , as in Fig. 33B ). Ecdysial tear extending into eye sheath ( Figs. 15F , 79F ). Head: Dorsal apotome ( Fig. 20J ), without 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. 25G ) with mandible well-developed, lacinia absent; palpus extending posterior to posterolateral margin of labium; labium separated medially by labrum, hypopharynx; apex of antenna ( Fig. 37A ) anterior to barely posterior to posterior extent of midlength portion of midleg (portion lateral to mesosternum), narrowed posteriorly; sensilla: dorsal apotomals ( Fig. 20J )—1 short seta, 1 campaniform sensillum; dorsolateral cephalic sclerite sensilla—1 very short seta, 1 campaniform sensillum; clypeal-labrals ( Fig. 25G )—2 short, slender setae; oculars ( Fig. 25G )—1 seta, 1 campaniform sensillum. Thorax: Prothoracic extension ( Fig. 25G ) wide, well-developed, extending from palpus to antenna; mesonotum with welldeveloped moderately sized tubercles, not extending posteromedially, with slight protuberance, not dividing metathorax medially ( Fig. 50A ); respiratory organ ( Fig. 44Q ) length/width = 2.63–3.45, moderately elongate, somewhat flattened dorsoventrally, with pores closely abutting at apex of respiratory organ, arranged in single row, outer surface with some wrinkles, with short, wide pedicel, base with elongate posteromedial apodeme, membranous base of respiratory organ moderately elongate, annulated, tracheal tube straight to slightly curved along length, with annulations to 3/4 length; wing ( Fig. 37A ) without apical tubercle or angle, 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. 37A ) 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 moderately anterior to apex of midleg; apex of hind leg slightly ventral to, partially abutting apex of midleg laterally; sensilla: anteromedials—1 very short seta, 1 campaniform sensillum; anterolaterals—1 short seta (shagreen making it difficult to interpret if more); dorsal setae ( Fig. 30D )—D-1-T, D-2-T, D-5-T peg-like setae, D-4-T seta, D-3-T campaniform sensillum; D-1-T, D-2-T, D-4-T on single tubercle, D-5-T on separate tubercle, D-3-T posterior to D- 4-T; supraalar 2—campaniform sensillum (difficult to discern among shagreen); metathoracics ( Fig. 50A )—2 campaniform sensilla; 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, peg-like or thin to thick setae, with rounded, short to moderately elongate tubercles, tergites or sternites entire, each without membranous disc; segment 9 ( Fig. 74K ) not strongly modified, terminal processes closely approximated basally, each projecting posterodorsolaterally, tapering to pointed apex; sensilla: tergite 1 ( Fig. 50A ) with 7 setae, 2 campaniform sensilla, including 4 lateral sensilla, D-2-I, D-3-I closely approximated, D-7-I situated anterolaterally near L-1-I; segment 4 ( Fig. 62A )—D-2-IV peg-like seta, D-3-IV moderately elongate seta, D-2-IV on large tubercle, D-3-IV on short tubercle; D-5-IV, D-8-IV peg-like setae, D-9-IV short seta, D-7-IV absent or not visible among shagreen; D-5-IV, D-8-IV on elongate separate tubercles, posterior dorsal sensilla in transverse row, arranged medially to laterally: D- 5-IV, D-8-IV, D-9-IV; L-1-IV elongate seta on rounded tubercle just anterior to L-3-IV; L-2-IV; L-3-IV, L-4-IV short setae on rounded tubercles, V-5-IV, V-6-IV, V-7-IV moderately elongate setae, on rounded well-developed tubercles, V-6-IV, V-7-IV closely approximated; segment 8 without D-3-VIII, without L-1-VIII; segment 9 ( Fig. 74K )—with D-5-IX, D-6-IX campaniform sensilla.

DISTRIBUTION AND HABITAT: The genus Atyphohelea is known from one Nearctic species and one unnamed species from Taiwan ( Borkent 1998 ). Adults are associated with smaller bodies of water, seeps, bogs, and small marshes. Pupae have been reared from a lake margin in North Carolina, U.S.A.

TAXONOMIC DISCUSSION: Only one species of Atyphohelea is known as a pupa ( Tables 2-3 ). The fourth abdominal segment of A. macroneura is illustrated without sensilla D-4-IV and D-7-IV but these campaniform sensilla (their form in other Ceratopogonidae ) are likely present but hidden among the strong shagreen present over much of the abdomen of this species (in addition the limited material was rather dirty, further obscuring observation of campaniform sensilla).

In nearly all Ceratopogonidae , the anteroventral margin of the wing forms a near right angle, with the margin completely or nearly perpendicular to the dorsal margin of the antenna. I found only two exceptions in the family. In the somewhat crushed specimens of A. macroneura studied here, the angle appeared less abrupt and more rounded (the wing side of the angle was greater). However, I could not convince myself that it was not a product of distortion. Fresh material would be able to test this idea. If present, the feature would be nearly unique in the family. It was also present in an exuviae of one unnamed species of Jenkinshelea (from Kruger N.P) which also had a greater angle, clearly an instance of homoplasy. Other Jenkinshelea had a nearly right angle anteroventral margin.

MATERIAL EXAMINED: A. macroneura : 3 pupal exuviae, Highlands Lake Ravenel, Macon County, North Carolina, USA, 10-VI-1986 (CNCI).

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 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 25. Posterior portion of heads, in ventral view. A. Stilobezzia antennalis. B. Stilobezzia rabelloi (from Borkent & Craig 2001). C. Schizonyxhelea bulla. D. Serromyia sp.. E. Allohelea japonica. F. Monohelea obscura (reconstructed from distorted specimen). G. Atyphohelea macroneura. H. Parabezzia huberti.

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 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 61. Abdominal segments four, in dorsal and ventral view (shagreen not shown). A. Allohelea japonica (reconstructed from distorted specimen). B. Monohelea obscura (reconstructed from distorted specimen). C. Monohelea hirsuta.

FIGURE 62. Abdominal segments four, in dorsal and ventral view (shagreen not shown). A. Atyphohelea macroneura (from Borkent 1998). B. Parabezzia huberti. C. Parabezzia alexanderi.

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.