Baeodasymyia Clastrier & Raccurt

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 : 59-60

Baeodasymyia Clastrier & Raccurt

( Figs. 2F , 19N , 24F , 29Q , 32K , 35E , 44C , 48F , 59A , 74A )

DIAGNOSIS: Only pupa of Ceratopogonidae with the respiratory organ elongate and with three distinct sections: a cylindrical, smooth basal portion, a cylindrical middle portion with annulations for most of its distal length, and a compressed, rounded or concave apical portion ( Fig. 44C ); also unique with a short foreleg, with its apex well anterior of the apex of the wing ( Fig. 35E ); also unique in being very small, with total body length 0.91–1.19 mm (only Schizonyxhelea are similarly small, with a body length 1.19–2.16 mm; however, the unknown pupae of Nannohelea , Baeohelea and some others will certainly be equally tiny).

DESCRIPTION: Total length = 0.91–1.19 mm. Without larval exuviae retained on abdomen. Exuviae with flagellum appressed against lateral margin of face, in some specimens against wing (as in Figs. 15D , 32K ). Ecdysial tear medial to base of antenna (as in Figs. 15B , 79D ); along prothoracic extension. Head: Dorsal apotome ( Fig. 19N ), without ventral line of weakness, with 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. 24F ) with mandible, lacinia absent; palpus extending posterior to posterolateral margin of labium; labium entire (not divided medially); apex of antenna ( Fig. 35E ) anterior to posterior extent of midlength portion of midleg (portion lateral to mesosternum); sensilla: dorsal apotomals ( Fig. 19N )—1 elongate seta, 1 campaniform sensillum; dorsolateral cephalic sclerite sensilla—2 setae, 1 campaniform sensillum; clypeal-labrals ( Fig. 24F )—not visible; oculars ( Fig. 24F )—2 elongate setae. Thorax: Prothoracic extension ( Fig. 24F ) wide, well-developed, extending from palpus to antenna; mesonotum with short tubercles, extending posteromedially, completely dividing metathorax medially ( Fig. 48F ); respiratory organ ( Fig. 44C ) length/width = 3.75–6.67, elongate, with three distinct sections: cylindrical basal portion smooth, cylindrical middle portion with annulations for most of distal length, compressed apical portion rounded or concave apically, apical portion somewhat flattened, with pores closely abutting or slightly separated at apex of respiratory organ, arranged in single row, with one additional, more basal pore, outer surface with annulations, without other surface modifications, with moderately elongate pedicel, base without posteromedial apodeme, membranous base of respiratory organ short, tracheal tube straight to slightly curved along length, with spirals restricted to base; wing ( Fig. 35E ) with slight angle lateral to apex of hind leg, separated medially by fore-, midlegs; halter apex and hind leg ( Fig. 32K ) broadly abutting; halter apex extending posteriorly to 1/3 length of tergite 2; legs ( Fig. 35E ) with lateral margin of foreleg near midlength of wing evenly curved; hind leg visible at lateral margin of wing ( Fig. 32K ); with apex of foreleg far anterior to apex of midleg; apex of hind leg abutting apex of midleg laterally; sensilla: anteromedials—1 seta; anterolaterals—2 setae; dorsal setae ( Fig. 29Q )—D-1-T, D-2-T, D-4-T setae, D-3-T campaniform sensillum, D-5-T absent; D-1-T, D-2-T on single tubercle, D-3-T posteromedial to D-4-T; supraalar 2—campaniform sensillum; metathoracics ( Fig. 48F )—1 seta, 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, thin to thick setae, with slightly rounded to pointed, or serrate short tubercles, tergites or sternites entire, each without membranous disc; segment 9 ( Fig. 74A ) not strongly modified but with dorsolateral row of spines at near midlength, bordering shallow posteromedial concavity, terminal processes closely approximated basally, each projecting posterodorsolaterally, tapering to pointed apex; sensilla: tergite 1 ( Fig. 48F ) with 6 setae, 1 campaniform sensillum, including 3 lateral sensilla, D-2- I, D-3-I closely approximated, D-7-I situated posteriorly near D-8-I; segment 4 ( Fig. 59A )—D-2-IV, D-3-IV short setae on serrate tubercles; D-5-IV barely visible short seta, D-8-IV short seta, D-9-IV absent; D-5-IV, D-4-IV, D-7- IV, D-8-IV on short, serrate 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 absent, L-2-IV, L-3-IV short setae on pointed tubercles, L-4-IV thick seta on serrate tubercle, V-5-IV, V-6-IV, V-7-IV short setae on serrate tubercles; segment 8 without D-3-VIII, without L-1- VIII; segment 9 ( Fig. 74A )—with D-5-IX, D-6-IX campaniform sensilla.

DISTRIBUTION AND HABITAT: The genus Baeodasymyia is known from five species in the Neotropical Region ( Borkent 2014 ). The immatures of two species have been reared from small springs in Costa Rica ( Borkent & Craig 1999).

TAXONOMIC DISCUSSION: Borkent & Craig (1999) described the pupae of the only two species known in this stage ( Tables 2–3 ). Pupae of Baeodasymyia are lacking thoracic sensillum D-5-T and two sensilla on abdominal segment 4: D-9-IV and L-1-IV.

MATERIAL EXAMINED: B. christopheri : 1 pupal exuviae (of paratype), Atenas, Costa Rica, 5-X-1993 (as larva, reared) (CNCI); 1 pupa (of paratype), 2 km NE Tarcoles, Costa Rica, 17-XII-1993 (as larva, reared) (CNCI). B. michaeli : 3 pupae (of paratypes), 9 pupal exuviae (of paratypes), 1 pupal exuviae, Atenas, Costa Rica, 25-X- 1993 (as larvae, reared) (CNCI, INBC); 1 pupal exuviae (in glycerin), 2 km NE of Tarcoles, Costa Rica, 17-XII- 1993 (CNCI).

Figures

FIGURE 2. Pupae in unusual habitats, preservation and relative sizes. A. Forcipomyia pupae, Rancho Grande, Venezuela; photo by Steve Marshall. B. Forcipomyia pupae under bark of log of Pinus sylvestris, Poland; photo by Marek W. Kozlowski. C. Pupae of Mallochohelea sp. on emergent plants at edge of Bolean Lk., 6 km NE Falkland, British Columbia. D. Palpomyia sp. showing mounting technique on pin, 8 km E. Sicamous, British Columbia. E. Permanent slide mount of dissected adult and pupal exuviae of Mallochohelea albibasis. F. Pupal exuviae, from left to right, of Baeodasymyia michaeli, Culicoides denticulatus and Sphaeromias longipennis showing size differences within the family. Total length of Baeodasymyia michaeli = 1.2 mm, of Sphaeromias = 7.3 mm.

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 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 48. Metathoraces and tergites 1, in dorsal view. A. Culicoides haematopotus. B. Paradasyhelea minuta. C. Ceratopogon sp. nr. abstrusus. D. Brachypogon sp. (from 6 km E. Falkland, BC, Canada). E. Alluaudomyia needhami. F. Baeodasymyia michaeli.

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.