Johannsenomyia Malloch

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 : 90-91

Johannsenomyia Malloch

( Figs. 21J , 27E , 30R , 39D , 45Q , 52C , 67B , 76H–I )

DIAGNOSIS: Only pupa of Ceratopogonidae with abdominal segment 4 with all four lateral setae (L-1-IV, L-2-IV, L-3-IV, L-4-IV) arranged transversely, closely approximated and on the anterior half of segment ( Fig. 67B ).

DESCRIPTION: Total length = 4.03–5.38 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 ( Fig. 21J ), 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. 27E ) with mandible well-developed, lacinia absent; palpus extending posterior to posterolateral margin of labium; labium entire (not divided medially); apex of antenna ( Fig. 39D ) posterior to posterior extent of midlength portion of midleg (portion lateral to mesosternum), narrowed posteriorly; sensilla: dorsal apotomals ( Fig. 21J )—1 elongate seta, 1 campaniform sensillum; dorsolateral cephalic sclerite sensilla—1 seta, 1 campaniform sensillum; clypeallabrals ( Fig. 27E )—2 slender setae; oculars ( Fig. 27E )—2 setae, 1 campaniform sensillum. Thorax: Prothoracic extension ( Fig. 27E ) wide, well-developed but narrow dorsolaterally, not extending to antenna; mesonotum with short tubercles, not extending posteromedially, not dividing metathorax medially ( Fig. 52C ); respiratory organ ( Fig. 45Q ) length/width = 2.41–3.09, moderately elongate, swollen apically, somewhat flattened apically, 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, tracheal tube straight, swollen curved apically in some, with spirals restricted to base, plates to half length; wing ( Fig. 39D ) without apical tubercle or angle, separated medially by fore-, midlegs; halter apex and hind leg (as in Fig. 33A ) broadly abutting; halter apex abutting or just anterior to anterolateral knob-like extension of tergite 2; legs ( Fig. 39D ) 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 slightly anterior to apex of midleg; apex of hind leg abutting apex of midleg laterally, small ventral lobe; sensilla: anteromedials—2 elongate setae (as in Figs. 31L–M ); anterolaterals—1 moderately long seta; dorsal setae ( Fig. 30R )—D-1-T, D-2-T, D-4-T, D-5-T setae, D-3-T campaniform sensillum, D-3-T anterolateral to D-4-T; supraalar 2—campaniform sensillum; metathoracics ( Fig. 52C )—1 campaniform sensillum; M-3-T distant from margin of metathorax (at least 1/3 length of metathorax). Abdomen: with tergite 1 with 1 medial spot, tergites 2–7 with medial area with stripe, 2 anterolateral spots, sternites 3–7 with anterolateral spots difficult to discern, segment 2 as wide or slightly wider than segment 3, segments with undivided, thin to thick setae, with rounded to slightly pointed, short tubercles, tergites or sternites entire, each without membranous disc; segment 9 ( Figs. 76H–I ) not strongly modified, terminal processes closely approximated basally, each projecting posterodorsolaterally to somewhat laterally, tapering to pointed apex; sensilla: tergite 1 ( Fig. 52C ) 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. 67B )—D-2-IV, D-3-IV short to moderately elongate setae on short tubercles; D-5-IV, D-8-IV, D-9-IV setae; D-5-IV on single tubercle, D-8-IV, D-9-IV on separate 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 short seta on short tubercle, near L-2-IV, L-3-IV, L-4-IV on anterior half of segment; L-2-IV, L-3-IV, L-4-IV short setae on short pointed tubercles, V-5-IV, V-6-IV, V-7-IV short setae on rounded tubercles; V- 5-IV, V-6-IV closely approximated; segment 8 without D-3-VIII, without L-1-VIII; segment 9 ( Figs. 76 H-I)—with D-5-IX, D-6-IX campaniform sensilla.

DISTRIBUTION AND HABITAT: The genus Johannsenomyia is known from 27 species from every Region worldwide but is absent from most of the Palaearctic (known only from north Africa) ( Borkent 2014 ). Immatures have been reared from river margins, lakes and Knausenberger (1987) additionally recorded a species from a pond. Williams (1955) described how mature larvae of J. argentata burrowed into the sand on a lake beach to pupate.

TAXONOMIC DISCUSSION: Only one species of Johannsenomyia has been described as a pupa ( Tables 2–3 ). Mayer (1957) also described the pupa of J. albidorsata (de Meillon) (as a Dicrohelea ) but I have examined the specimen and it is in fact a species of Bezzia . De Meillon (1937) in his description of the adult actually noted that the pupal exuviae of the holotype had been mixed with those of a Palpomyia and a Bezzia and its identification was therefore uncertain. Mayer (1957) however was clearly unaware of the uncertain association and described the purported pupal exuviae as if it was that of J. albidorsata . Debenham (1974) gave a brief synopsis of the pupa of Johannsenomyia but based it on the misidentified description by Mayer (1957).

MATERIAL EXAMINED: J. argentata : 1 pupal exuviae, Storrs, Connecticut, USA, 1-VI-1953 (USNM); 1 pupal exuviae, Patuxent Wildlife Refuge, Prince George’s County, Maryland, USA, 24-V-1977 (USNM); 6 pupal exuviae, Potomac River at Scott Run, Fairfax County, Virginia, USA, 7-VI-1955 (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 21. Dorsal apotomes, anterior view. A. Dibezzia prominens (from Mayer 1934c). B. Hebetula tonnoiri (from Elson- Harris 1987). C. Jenkinshelea albaria (from Wirth & Grogan 1979). D. Macropeza albitarsis (from Szadziewski & Dominiak, 2007). E. Mallochohelea termophila (from Spinelli 1984, modified). F. Probezzia pallida (from Wirth & Grogan 1979). G. Neobezzia fittkaui (from Ronderos et al. 2011, modified). H. Nilobezzia basispinigera (from Elson-Harris 1987). I. Nilobezzia virago (from Elson-Harris 1987). J. Johannsenomyia argentata (from Wirth & Grogan 1979). K. Lanatomyia electra (from Elson-Harris 1987). L. Anebomyia atripes (from Wirth & Grogan 1979, modified).

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 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 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 39. Legs, wings and apices of the antennae, in ventral view. A. Probezzia sabroskyi. B. Neobezzia fittkaui. C. Nilobezzia sp. (from Australia). D. Johannsenomyia argentata. E. Lanatomyia electra. F. Anebomyia atripes.

FIGURE 45. Respiratory organs, in dorsal view, pattern on tracheal tubes shown only in some. A. Clinohelea tasmaniensis (from Elson-Harris & Kettle 1986). B. Clinohelea bimaculata (from Wirth & Grogan 1979, modified). C. Heteromyia wokei (from Wirth & Grogan 1977). D. Heteromyia clavata (from Wirth & Grogan 1977). E. Pellucidomyia leei (from Elson-Harris & Kettle 1986). F. Pellucidomyia geari (from de Meillon & Wirth 1981). G. Dibezzia prominens (from Mayer 1934c). H. Hebetula tonnoiri (from Debenham 1974). I. Jenkinshelea albaria (from Wirth & Grogan 1979). J. Macropeza albitarsis (from Szadziewski & Dominiak, 2007). K. Mallochohelea albihalter (from Wirth & Grogan 1979). L. Probezzia pallida (from Wirth & Grogan 1979). M. Probezzia xanthogaster (from Wirth & Grogan 1979). N. Neobezzia fittkaui (from Ronderos et al. 2011). O. Nilobezzia aranea (from Elson-Harris 1987). P. Nilobezzia virago (from Elson-Harris 1987). Q. Johannsenomyia argentata (from Wirth & Grogan 1979). R. Lanatomyia electra (from Debenham 1974). S. Lanatomyia miles (from Debenham 1974). T. Anebomyia atripes (from Wirth & Grogan 1979).

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 67. Abdominal segments four, in dorsal and ventral view (shagreen not shown). A. Nilobezzia curticornis. B. Johannsenomyia argentata. C. Lanatomyia miles.

FIGURE 76. Abdominal segments 9. A. Macropeza albitarsis, male, in ventral view (modified from Szadziewski & Dominiak, 2007). B. Macropeza albitarsis, female, in ventral view (from Szadziewski & Dominiak, 2007). C. Mallochohelea termophila, male, in ventral view (from Spinelli 1984). D. Mallochohelea termophila, female, in dorsal view (from Spinelli 1984). E. Probezzia pallida, male, in dorsal and ventral view (from Wirth & Grogan 1979). F. Neobezzia fittkaui, male, in dorsal view (from Ronderos, et al. 2011). G. Nilobezzia virago, female, in dorsal view (from Elson-Harris 1987). H. Johannsenomyia argentata, male, in dorsal and ventral view (from Wirth & Grogan 1979). I. Johannsenomyia argentata, female, in dorsal and ventral view (from Wirth & Grogan 1979). J. Lanatomyia miles, female, in dorsal view (from Elson- Harris 1987). K. Anebomyia atripes, male, in dorsal and ventral view (from Wirth & Grogan 1979). L. Anebomyia atripes, 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.