Ceratopogon Meigen

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 : 54-55

Ceratopogon Meigen

( Figs. 13C , 15B , 19H–J , 24C , 29N , 32H , 35B , 43P–Q , 48C , 57C , 58A , 73J )

DIAGNOSIS: Only pupa of Ceratopogonidae with the dorsal apotome with a pronounced central dome from which both dorsal apotome setae arise ( Figs. 19H–J ); also unique in having an inner tracheal tube of the respiratory organ strongly hooked apically, appearing as an inverted "J" ( Figs. 43P–Q ).

DESCRIPTION: Total length = 1.88–3.38 mm. Without larval exuviae retained on abdomen. Exuviae with flagellum appressed against lateral margin of face ( Fig. 15B ). Ecdysial tear medial or posteromedial to base of antenna ( Figs. 15B , 79D ); along prothoracic extension. Head: Dorsal apotome ( Figs. 19H–J ), without ventral line of weakness, without dorsomedial tubercle, with central dome; dorsolateral cephalic sclerite ( Fig. 13C ) separated from scutum by thin cuticle, separate from scutum upon emergence, each side separated medially by dorsal apotome in whole pupa; mouthparts ( Fig. 24C ) with only mandible or mandible ventral to lacinia; palpus extending equal to or just posterior to posterolateral margin of labium; labium separated medially by labrum, hypopharynx; apex of antenna ( Fig. 35B ) anterior to posterior extent of midlength portion of midleg (portion lateral to mesosternum); sensilla: dorsal apotomals ( Figs. 19H–J )—1 elongate seta, 1 campaniform sensillum; dorsolateral cephalic sclerite sensilla—1 seta, 1 campaniform sensillum; clypeal-labrals ( Fig. 24C )—2 short, slender setae; oculars ( Fig. 24C )—2 setae, 1 campaniform sensillum. Thorax: Prothoracic extension ( Fig. 24C ) wide, welldeveloped, extending from palpus to antenna; mesonotum with short tubercles, not extending posteromedially or with slight protuberance, not dividing metathorax medially ( Fig. 48C ); respiratory organ ( Figs. 43P–Q ) length/ width = 2.74–3.68, moderately elongate, apical half somewhat flattened dorsoventrally, with pores closely abutting or slightly separated at apex of respiratory organ, arranged in single straight or curve row, outer surface smooth or with some wrinkles, with or without short, wide pedicel, base with slender, strung out posteromedial apodeme, membranous base of respiratory organ short, tracheal tube apically J-shaped, surface smooth or distally with plates; wing ( Fig. 35B ) without apical tubercle or angle, separated medially by fore-, midlegs; halter apex and hind leg ( Fig. 32H ) just separate; halter apex abutting anterolateral knob-like extension of tergite 2; legs ( Fig. 35B ) with lateral margin of foreleg near midlength of wing evenly curved; hind leg visible at lateral margin of wing ( Fig. 32H ); with apex of foreleg ventral to or moderately 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. 29N )—D-1-T, D-2-T, D-4-T, D-5-T setae, D-3-T campaniform sensillum, D-3-T posterior to D-4-T; supraalar 2—campaniform sensillum; metathoracics ( Fig. 48C )—1 seta, 2 campaniform sensilla; M-3-T near anterior margin of metathorax. Abdomen: without pigmentation pattern or pigmentation patterns only evident in alcohol/ glycerin material, with tergite 1 with 3 medial spots, tergites 2–8 with 3 medial spots, anterolateral spots, sternites 3–7 with medial spot, anterolateral spot, segment 2 as wide or slightly wider than segment 3, segments with undivided, thin to thick setae, with bilobed, short and triangular to elongate and slender tubercles, tergites or sternites entire, each without membranous disc; segment 9 ( Fig. 73J ) not strongly modified, terminal processes closely approximated basally, each projecting posterodorsolaterally to nearly laterally, tapering to pointed apex; sensilla: tergite 1 ( Fig. 48C ) 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 ( Figs. 57C , 58A )—D-2-IV, D-3-IV setae on either short tubercle, bilobed or bifid elongate tubercles; D-5-IV present or absent, D-8-IV, D-9-IV short to elongate setae; each at base of short bilobed or elongate bifid, 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 or D-5-IV absent and D-7-IV anterior to D-9-IV; L-1-IV short seta at base of short bilobed or elongate bifid tubercle, well anterior of posterior lateral setae; L-2-IV, L-3-IV, L-4- IV short setae at base of bilobed elongate bifid tubercles, V-5-IV, V-6-IV, V-7-IV on bilobed or at base of bifid, elongate tubercles; segment 8 without D-3-VIII, without L-1-VIII; segment 9 ( Fig. 73J )—with D-5-IX, D-6-IX campaniform sensilla.

DISTRIBUTION AND HABITAT: The genus Ceratopogon is known from 43 species in the Holarctic Region with adults emerging in cold environments (Arctic, mountainous, or in early spring) ( Borkent & Grogan 1995). Immatures have been found in moss in cold forest pools, at the margins of shallow water bodies, bogs, fens, in moist soil, on flood plains and on lake shores.

TAXONOMIC DISCUSSION: Six species of Ceratopogon are known as pupae ( Tables 2–3 ) and the five Nearctic ones are keyed by Borkent & Grogan (1995) as part of a comprehensive revision of the genus.

Ceratopogon boomerangus is the only member of the genus missing the full complement of abdominal sensilla present in most Ceratopogonidae (and all other Ceratopogon ), with fourth abdominal segment sensillum D-5-IV absent. The figure of the fourth abdominal segment of C. boomerangus shown in Fig. 57C (from Borkent & Grogan 1995) is distorted by compression and the distribution of the sensilla are broader than is natural (i.e. the dorsal and ventral sensilla are actually more lateral, L-1-IV, L-2-IV and L-3-IV are shown as ventral). This is true for all the species illustrated by Borkent & Grogan (1995:183). The fourth abdominal segment of C. nr. inverecundus in Fig. 58A was drawn from an uncompressed specimen in glycerin.

Mayer (1934a:238) described a pupa referred to as "Genus incertum" from Germany which was reexamined for this study. It is a species of Ceratopogon but cannot be identified to species at present.

MATERIAL EXAMINED: C. abstrusus : 1 pupal exuviae (of paratype), Sand Pond Br., Blue Ridge, New York, USA, 26-V-1959 (NYSM); 1 pupal exuviae (of paratype), The Gulf Brook, Blue Ridge, New York, USA, 25- V-1958 (NYSM); 1 pupal exuviae (of paratype), New York (?) (NYSM); 1 pupal exuviae (of paratype), Hamilton-Essex, Newcomb, New York, USA, 28-V-1959 (NYSM). C. arcanus : 2 pupal exuviae (of paratypes), Fishing Creek, Newcomb, New York, USA, 5-V-1958 (NYSM); 2 pupal exuviae (of paratype), Huntington Rd. Marsh, Newcomb, New York, USA, 14-V-1958 (NYSM); 1 pupal exuviae (of paratype), Blue Mountain Lake, New York, USA, 3-V-1960 (NYSM); 1 pupal exuviae (of paratype), High Rock Pond outlet, Inlet, New York, USA, 8-V-1958 (NYSM); 2 pupal exuviae (of paratypes), Hamilton-Essex, Newcomb, New York, USA, 11-V-1959 (NYSM). C. boomerangus : 1 pupal exuviae (of allotype), College Park, Prince George County, Maryland, USA, 14-IV-1976 (USNM). C. inverecundus : 2 pupal exuviae (of paratypes), Edge 6 Mile Creek, Long Lake, New York, USA, 18-V- 1959 (NYSM). C. naccinervis : 1 pupal exuviae, Severskii Donets River, Donetsk Province, Ukraine, 16-IV-1970 (ZIN). C. willisi : 1 pupa (paratype), College Park, Prince Georges County, Maryland, 17-III-1976 (USNM); 1 pupa (paratype), as previous locality, 26-III-1976 (USNM). C. nr. abstrusus : 1 pupal exuviae (in glycerin), 2 pupal exuviae, Bolean Lake, 6 km NE Falkland, BC, Canada, 12-13-VII-1989 (CNCI). C. nr. inverecundus : 1 pupal exuviae (in glycerin), 1 pupal exuviae, Bolean Lake, 6 km NE Falkland, BC, Canada, 12-13-VII-1989 (CNCI). C. sp.: 1 pupa, Haspertalsperre Sauerland, Westfalen, Germany 1-VI-? (ZSMC).

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 57. Abdominal segments four, in dorsal and ventral view (shagreen not shown). A. Culicoides haematopotus (from Borkent 2012). B. Paradasyhelea albipunctata. C. Ceratopogon boomerangus (from Borkent & Grogan 1995, compressed dorsoventrally).

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.