Brachypogon Kieffer

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 : 55-56

Brachypogon Kieffer

( Figs. 13D , 15C , 19K , 24D , 29O , 32I , 35C , 43R , 48D , 58B , 73K )

DIAGNOSIS: Only pupa of Ceratopogonidae with setae D-1-T, D-2-T, D-4-T, and D-5-T tightly appressed and on a single, short tubercle ( Fig. 29O ). Also only pupa with prothoracic extension abutting the antenna but not extending to the palpus ( Fig. 24D ) and halter extending only to the anterior margin of tergite 2 ( Fig. 32I ).

DESCRIPTION: Total length = 1.13–2.03 mm. Without larval exuviae retained on abdomen. Exuviae with flagellum appressed against lateral margin of face ( Fig. 15C ). Ecdysial tear medial to base of antenna ( Figs. 15C , 79D ); along prothoracic extension. Head: Dorsal apotome ( Fig. 19K ), without ventral line of weakness, without dorsomedial tubercle, without central dome; dorsolateral cephalic sclerite ( Fig. 13D ) separated from scutum by thin cuticle, separate from scutum upon emergence, each side separated medially by dorsal apotome in whole pupa; mouthparts ( Fig. 24D ) with mandible well-developed, lacinia absent; palpus extending posterior to posterolateral margin of labium; labium separated medially by labrum, hypopharynx; apex of antenna ( Fig. 35C ) anterior to posterior extent of midlength portion of midleg (portion lateral to mesosternum); sensilla: dorsal apotomals ( Fig. 19K )—1 short seta present or absent, 1 campaniform sensillum; dorsolateral cephalic sclerite sensilla—1 seta, no campaniform sensillum; clypeal-labrals ( Fig. 24D )—0–1 seta or 1 seta and 1 small pit; oculars ( Fig. 24D )—2 setae, 2 campaniform sensilla. Thorax: Prothoracic extension ( Fig. 24D ) short, present only dorsolaterally abutting antenna, not extending to palpus; mesonotum with short tubercles, not extending posteromedially, not dividing metathorax medially ( Fig. 48D ); respiratory organ ( Fig. 43R ) length/width = 2.37–4.77, moderately elongate to elongate, with blunt apex, area with single subbasal spiracle slightly expanded, somewhat flattened dorsoventrally, with pores closely abutting at apex of respiratory organ, arranged in single row, with one additional subbasal pore, outer surface with annulations, without other surface modifications, with or without short, wide pedicel, base without posteromedial apodeme, membranous base of respiratory organ short, tracheal tube straight to slightly curved along length, with weak spirals restricted to base; wing ( Fig. 35C ) with apical tubercle lateral to apex of hind leg, separated medially by fore-, midlegs; halter apex and hind leg ( Fig. 32I ) just separate to just touching; halter apex abutting anterolateral knob-like extension of tergite 2; legs ( Fig. 35C ) with lateral margin of foreleg near midlength of wing evenly curved; hind leg visible at lateral margin of wing ( Fig. 32I ); with apex of foreleg well anterior to apex of midleg; apex of hind leg slightly dorsal to, partially abutting apex of midleg laterally; sensilla: anteromedials—1 seta; anterolaterals—2 setae; dorsal setae ( Fig. 29O )—D-1-T, D-2-T, D-4-T, D-5-T setae, all on one tubercle, D-3-T campaniform sensillum, D-3-T posterolateral to D-4-T; supraalar 2—campaniform sensillum; metathoracics ( Fig. 48D )—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 rounded to pointed, short tubercles, tergites or sternites entire, each without membranous disc; segment 9 ( Fig. 73K ) not strongly modified, terminal processes closely approximated basally, each projecting posterodorsolaterally to nearly laterally, tapering to pointed apex; sensilla: tergite 1 ( Fig. 48D ) with 8 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. 58B )—D-2-IV peg-like or slender seta, D-3- IV moderately elongate seta on separate tubercles; D-5-IV short seta, D-8-IV, D-9-IV moderately elongate setae; on short, separate tubercles, posterior dorsal sensilla in transverse row, arranged medially to laterally: D-5-IV, D-4- IV, D-7-IV, D-8-IV, D-9-IV; L-1-IV elongate seta on pointed tubercle, well anterior of posterior lateral setae; L-2- IV, L-3-IV, L-4-IV moderately elongate setae on pointed tubercles, V-5-IV, V-6-IV, V-7-IV short to moderately elongate setae on rounded tubercles; segment 8 without D-3-VIII, without L-1-VIII; segment 9 ( Fig. 73K )—with D-5-IX, D-6-IX campaniform sensilla.

DISTRIBUTION AND HABITAT: The genus Brachypogon is known from 200 species from every Region worldwide ( Borkent 2014 ). Immatures are generally in such small to moderately sized lentic habitats as pools, marshes, bogs and fens as well as the mud, sand or detritus at the margins of streams and creeks. At least within the Nearctic, species are diverse and often common in bogs and fens. At least two species were common as adults in the intertidial zone in western Costa Rica (pers. obs.).

TAXONOMIC DISCUSSION: There are only seven species of Brachypogon known as pupae ( Tables 2–3 ).

Some previous descriptions have included illustrations of the respiratory organ with pores only at the apex ( Mayer 1940, Kettle & Lawson 1952, Glukhova 1979 b, Harris 1981, Elson-Harris 1990, Szadziewski et al. 1994, 1997). All specimens examined here had an additional pore near the base and in some instances this was difficult to see. If present in all Brachypogon , the arrangement of pores and the squat, rectangular to somewhat funnel-shape of the respiratory organ would be unique in the Ceratopogonidae .

Debenham (1991) illustrates three species of Brachypogon from Australia and Mayer (1940) described one species from Sweden with only two posterior lateral sensilla on segment 4 (likely L-3-IV, L-4-IV) but these should be reexamined for the smaller L-2-IV possibly present.

Thienemann (1936:176) described a pupa as "genus incertum" that is almost certainly a species of Brachypogon . It has the characteristic respiratory organ shape and arrangement of pores (one basal) and the distribution of abdominal sensilla matches fairly well (there is an extra anterior lateral sensillum that would be unique to the genus but was likely an error).

MATERIAL EXAMINED: B. nitidulus : 1 pupal exuviae, Zaklin'e, Leningrad Province, Russia, 24-VI-1972 (ZIN). B. taivoi : 1 pupal exuviae, Chu river, Kochkorka, Issyk-Kul Province, Kyrgyzstan, 30-VIII-1971 (ZIN). B. ussuriensis : 2 pupal exuviae, Ussuri Nature Reserve, Primorskii Territory, Russia, 6-VI-1973 (ZIN). B. sp.: 1 pupal exuviae (in glycerin), 6 km E Salmon Arm, BC, Canada, 6-VI-1990 (CNCI); 1 pupa, 3 pupal exuviae (in glycerin), Spanish Lake, 6 km E. Falkland, 50°29.12N 119°28.07W, BC, Canada, 27-28-V-2008 (CNCI); 1 pupal exuviae, Blue Mountain Lake, New York, USA, 3-V-1960 (NYSM); 2 pupal exuviae, Flag Glade, Pocahontas County, Virginia, USA, 24-VI-1976 (VPIC); 5 pupal exuviae, 1 km downstream lower Cascade Falls, Giles County, Virginia, USA, 3-VII-1977 (5 VPIC); 1 pupal exuviae, Moung Sayaboury, Sayaboury Province, Laos, 19-II-1967 (BPBM); 1 pupal exuviae, Houay La stream, Moung Sayaboury, Sayaboury Province, Laos, 25-XI-1967 (BPBM); 1 pupal exuviae, Moung Sayaboury, Sayaboury Muong, Laos, 19-23-II-1967 (BPBM).

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 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 43. Respiratory organs, most in dorsal view, Fig. 43T in lateral view, pattern on tracheal tubes shown only in some. A. Dasyhelea pseudoincisurata (from Waugh & Wirth 1976). B. Dasyhelea grisea (from Waugh & Wirth 1976). C. Dasyhelea traverae (from Waugh & Wirth 1976). D. Dasyhelea pseudocincta (from Waugh & Wirth 1976). E. Dasyhelea leptobranchia (from Waugh & Wirth 1976). F. Dasyhelea brevicornis (from Waugh & Wirth 1976). G. Dasyhelea hitchcocki (from Wirth 1976a). H. Culicoides chiopterus (from Jamnback 1965). I. Culicoides furens (from Jamnback 1965). J. Culicoides guttipennis (from Jamnback 1965). K. Culicoides dewulfi (from Kettle & Lawson 1952, as C. pseudochiopterus). L. Culicoides punctatus (from Kettle & Lawson 1952). M. Culicoides sanguisuga (from Jamnback 1965). N. Culicoides variipennis (from Jamnback 1965). O. Paradasyhelea albipunctata (from Elson-Harris & Kettle 1985a). P. Ceratopogon inverecundus (from Borkent & Grogan 1995). Q. Ceratopogon willisi (from Borkent & Grogan 1995). R. Brachypogon sp. (from Virginia, USA). S. Alluaudomyia needhami (from Wirth & Grogan 1981). T. Alluaudomyia fumosipennis (from Debenham 1971).

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 58. Abdominal segments four, in dorsal and ventral view (shagreen not shown). A. Ceratopogon nr. inverecundus. B. Brachypogon sp. (from 6 km E. Falkland, BC, Canada). C. Alluaudomyia needhami.

FIGURE 73. Abdominal segments 9. A. Dasyhelea laeta, female, in ventral view (from Mayer 1934c). B. Dasyhelea assimilis, male, in lateral view with ventral to the right (from Mayer 1934c). C. Dasyhelea assimilis, male, in lateral view with ventral to the right (from Mayer 1934c). D. Dasyhelea grisea, male, in dorsal view (from Waugh & Wirth 1976). E. Dasyhelea chani, female, in dorsal view (from Wirth & Linley 1990). F. Dasyhelea atlantis, female, in ventral view (from Waugh & Wirth 1976). G. Dasyhelea gluchovae, female, in ventral view (from Brodskaya 1996). H. Culicoides haematopotus, female, in dorsal and ventral view (shagreen not shown). I. Paradasyhelea albipunctata, female, in dorsal view (Elson-Harris & Kettle 1985a). J. Ceratopogon boomerangus, female, in ventral view (modified from Borkent & Grogan 1995). K. Brachypogon artemis, female, in dorsal view (from Debenham 1991). L. Alluaudomyia appendiculata, female, in dorsal view (from Elson-Harris & Kettle 1985b).

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.