Forcipomyia 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 : 40-41

Forcipomyia Meigen

( Figs. 2A–B , 8D–H , 9A–H , 10A–H , 11A–C , 14C–D , 18C–D , 23C–D , 29D–F , 32C , 34C , 42E–R , 47C–D , 55C , 56A , 72F–I )

DIAGNOSIS: Only pupa of Ceratopogonidae without a prothoracic extension ( Figs. 23C–D ), with the labium not divided medially by an elongate suture (as is present in Fig. 23F ) and with the terminal processes either closely abutting basally ( Figs. 8D–H , 9A–H , 10A–H , 11A–D , 72F–H ) or the terminal process spicules (if present) directed posteriorly to posterolaterally (not anteriorly); combination of retaining larval exuviae on the abdomen or with very elongate tubercles on the thorax ( Figs. 9A, C, F , 10A–E ) and/or abdomen, and the terminal process with spicules (if present) directed posteriorly to posterolaterally (not anteriorly) is unique to many species.

DESCRIPTION: Habitus as in Figs. 8D–H , 9A–H , 10A–H , 11A–C . Total length = 1.69–3.03 mm. With (as in Fig. 11F ) or without larval exuviae retained on abdomen. Exuviae with flagellum appressed against lateral margin of face ( Fig. 14D ). Ecdysial tear to anterior margin of base of antenna ( Figs. 14C–D , 79B ); along posterolateral portion of eye. Head: Dorsal apotome ( Figs. 18C–D ), without ventral line of weakness, with or without 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 ( Figs. 23C–D ) with mandible, lacinia well-developed, not overlapping apically; palpus extending posterior to posterolateral margin of labium; labium separated medially by labrum, hypopharynx; apex of antenna ( Fig. 34C ) anterior to posterior extent of midlength portion of midleg (portion lateral to mesosternum); sensilla: dorsal apotomals ( Figs. 18C–D )—absent or 1 very short seta; dorsolateral cephalic sclerite sensilla—absent or 1 short seta, 1 campaniform sensillum; clypeal-labrals ( Figs. 23C–D )—1 seta, 1 campaniform sensillum; oculars ( Figs. 23C–D )—1 campaniform sensillum. Thorax: Prothoracic extension ( Figs. 23C–D ) absent; mesonotum with or without elongate tubercles, extending posteromedially, completely dividing metathorax medially ( Figs. 47C–D ); respiratory organ ( Figs. 42E–R ) length/width = 1.05–4.75, highly variable, knob-like to elongate, slender to nearly spherical, with or without posterior basal swelling or lobe, somewhat flattened laterally or circular in cross-section, with pores closely abutting at apex of respiratory organ, arranged in single line of varying shapes or two widely spaced rows, outer surface smooth or with spicules, without or with short pedicel, base without posteromedial apodeme, membranous base of respiratory organ short, tracheal tube straight to slightly curved along length, smooth or with spirals restricted to base; wing ( Fig. 34C ) with apical tubercle or angle lateral to apex of hind leg, separated medially by fore-, midlegs; halter apex and hind leg ( Figs. 10C , 32C ) broadly to narrowly separate; halter apex abutting anterolateral edge of tergite 2; legs ( Fig. 34C ) with lateral margin of foreleg near midlength of wing slightly sinuous to evenly curved; hind leg visible at lateral margin of wing ( Fig. 32C ); with apex of foreleg moderately to well anterior to apex of midleg; apex of hind leg abutting apex of midleg laterally; sensilla: anteromedials—0–2 campaniform sensilla; anterolaterals—with or without 1 short or elongate seta or 1 campaniform sensillum; dorsal setae ( Figs. 29D–F )—D-1-T seta, D-2-T seta present or absent, D-3-T campaniform sensillum; D-2-T, D-3-T on single tubercle in some, or closely associated (if D-2-T present); supraalar 2—campaniform sensillum; metathoracics ( Figs. 47C–D )—0, 2 campaniform sensilla; M-3-T near anterior margin of metathorax. Abdomen: without pigmentation pattern, segment 2 as wide or somewhat wider than segment 3, segments with undivided, thin to thick setae, without tubercles or segments 2–8 with spines or thick tubercles or segments 1–5 with branched or thick spined elongate tubercles, tubercles rounded to pointed, tergites or sternites entire, each without membranous disc; segment 9 ( Figs. 72F–I ) highly variable in shape, terminal processes widely separated to closely approximated basally, each projecting posteriorly to laterally, tapering to pointed apex or tapering near apex to point; sensilla: tergite 1 ( Figs. 47C–D ) with 3–5 setae, 1–2 campaniform sensilla, including 1–2 lateral sensilla, D-2-I, D-3-I (if present) well separated, D-7-I absent or situated posteriorly near D-8-I; segment 4 ( Figs. 55C , 56A )—D-2-IV peg-like to slender seta, D-3-IV slender seta, without tubercles or very short separate ones; D-8-IV, when present, a moderately elongate seta; D-5-IV present or absent, D-7-IV, D-9-IV absent; D-4-IV without to on moderately elongate tubercle, posterior dorsal sensilla with D-4-IV, D-8-IV lateral to each other or D-8-IV absent, L-1-IV short to elongate seta present on short to long tubercle, close to L-2-IV, L-2-IV, L-3-IV, L-4-IV on separate rounded to elongate tubercles, in some with L-2-IV, L-3-IV closely approximated, V-5-IV, V-6-IV, V-7-IV short to moderately elongate setae, V-5-IV present or absent, on separate tubercles or V-6-IV, V-7-IV on single tubercle; segment 8 without D-3-VIII, without L-1-VIII; segment 9 ( Figs. 72F–I )—D-5-IX campaniform sensillum, D-6-IX campaniform sensillum or short or elongate seta.

DISTRIBUTION AND HABITAT: The genus Forcipomyia is known from 1150 species in 35 subgenera from every Region worldwide ( Borkent 2014). The larvae and pupae are found in many semi-terrestrial to moist terrestrial habitats, including under loose bark of logs, under decaying leaves and manure, decomposing vegetable matter and fungi, moss on soil, rotting wood or rocks, sap at tree wounds, in and above the water of various phytotelmata, mud along streams, bogs, and shallow water ponds.

TAXONOMIC DISCUSSION: There are 146 species of Forcipomyia , in 14 subgenera, known as pupae ( Tables 2–3 ). The pupae of this genus vary tremendously morphologically and are rich in numerous derived features, including overall body form, chaetotaxy, shape of respiratory organs and more. The group is ripe for a cladistic analysis and combined with their various behaviours and diverse niches, are certain to provide a rich opportunity to examine diversification within the genus. A few features were put into a phylogenetic context by Chan & LeRoux (1971c).

Although earlier work had shown that pupae of Forcipomyia have a variety of interesting modifications (e.g. Long 1902, Speiser 1910, Carter 1919, Rieth 1915), it was not until L.G. Saunders (1924, 1925, 1957, 1959, 1964), in a series of remarkable papers, studied the group that the wealth of features and their importance in their classification became apparent. He described exemplars from the 10 available subgenera (out of 14 for which immatures are known today), showing that the larvae, pupae and adults had congruent and distinctive features, indicating a solid basis for their classification (although derived features were not interpreted). He provided a consistent format for the description of the pupae and laid the groundwork for all future studies. Work by Chan & Saunders (1965), Chan & LeRoux (1965, 1970, 1971a, 1971b) and Chan & Linley (1989) added significantly with further detailed descriptions of Forcipomyia pupae.

The pupae are distinctive enough that a key to the subgenera is possible (see above).

Further differences exist at the species level and early work indicated these also could be keyed ( Malloch 1915 — Illinois; Lenz 1934 —Europe; Wirth & Howarth 1982 —Hawaiian F. ( Euprojoannisia ); Lewańczyk et al. 2010 —European species of F. ( Forcipomyia )).

Wirth & Howarth (1982), Chen et al. (1980), Spinelli et al. (2995) and Marino et al. (2010) provided SEM photomicrographs of pupae of Forcipomyia .

I initially thought that the subgenera F. ( Lepidohelea ) and F. (Schizoforcipomyia) could be separated on the basis of the relative size of the medial tubercle on the dorsal apotome. Previous descriptions of species of F. ( Lepidohelea ) show these to be long. The only species of F. (Schizoforcipomyia) to be described for this feature is F. borbonica by Chan & LeRoux (1971b, as F. petersoni ), who illustrate the pupa as having a short, rounded tubercle on the middle of the dorsal apotome, although in the text they indicate this to be "elongate". I examined the two paratype pupal exuviae of F. petersoni and found both to have an elongate tubercle, indistinguishable from those of species of F. ( Lepidohelea ).

Pupae of Forcipomyia have reduced numbers of thoracic sensilla, as well as fewer on the dorsum of the abdominal segments and the exact naming of these was uncertain. D-3-T and D-4-IV are likely correct as unique campaniform sensilla (in these areas).

MATERIAL EXAMINED: F. aeria : 2 pupal exuviae (of paratype), Mayaquez, Puerto Rico, 18-III-1953 (USNM). F. anabaenae : 1 pupal exuviae (of paratype), Singapore, 25-III-1962 (CNCI). F. antiguensis : 1 pupal exuviae (of paratype), Falmouth Harbour, Antigua, 6-V-1953 (CNCI). F. ashantii : 1 pupal exuviae, Tafo, Ghana, 17-V-1963 (ANIC); Berger's Hall, Hazyview, East Transvaal, South Africa, 3-XII-1973 (ANIC). F. attenuata : 1 pupal exuviae (of paratype), La Lola, Costa Rica, 2-VI-1956 (CNCI). F. bipunctata : 7 pupa, 23 pupal exuviae, 4 mi. W., 3 mi. S Schuyler, Nebraska, USA, various dates, 1990-1991 (CNCI). F. blantoni : 2 pupal exuviae, Turrialba, Costa Rica, 2-IX-1964 (USNM); 2 pupal exuviae, as previous locality, 26-XI-1964 (USNM); 1 pupal exuviae, as previous locality, 21-XII-1964 (USNM); 1 pupal exuviae, as previous locality, 29-XII-1964 (USNM). F. borbonica : 1 pupal exuviae (labeled as paratype of an undescribed species F. lerouxi), Wallace Way, Singapore, 1961 (LEMQ). F. brasiliensis : 1 pupal exuviae, Maracas, Trinidad, 18-VII-1957 (USNM). F. bromeliae : 1 pupal exuviae (of paratype), Rio de Janeiro, Brazil, 23-VIII-1923 (CNCI). F. bystraki : 1 pupal exuviae, Calvert, Cecil County, Maryland, USA, 29-III-1969 (USNM); 2 pupal exuviae, Headwaters, Highland County, Virginia, USA, VI-1969 (USNM). F. caerulea : 1 pupal exuviae (of paratype), Rio de Janeiro, Brazil, 8-VIII-1923 (CNCI). F. canadensis : 2 pupae (of paratypes), Pike Lake, Saskatchewan, Canada, X-1957 (USNM). F. caribbeana : 1 pupal exuviae (of paratype), Trinidad, 12-V-1953 (CNCI). F. cerifera : 1 pupal exuviae, Rio de Janeiro, Brazil, 1-VIII- 1923 (CNCI). F. cornuta : 1 pupal exuviae (of paratype), Costa Rica, 4-VI-1956 (CNCI). F. crinita : 1 pupal exuviae (of paratype), Emma Lake, Saskatoon, Saskatchewan, Canada, 19-IV-1957 (CNCI). F. edmistoni : 1 pupal exuviae (of paratype), Allegany State Park, New York, USA, 28-V-3-VI-1963 (USNM); 1 pupal exuviae (of paratype), Bittinger 4-H Campground, Garrett County, Maryland, USA, 5-V-1960 (USNM); 1 pupa (of paratype), 1 pupal exuviae (of paratype), Patuxent Wildlife Refuge, Maryland, USA, 28-VII-1979 (USNM). F. elegantula : 1 pupal exuviae, Saskatoon, Saskatchewan, Canada, 3-IX-1950 (USNM); 1 pupa, 1 pupal exuviae, Plummer's Island, Maryland, USA, 3-VI-1976 (USNM); 1 pupal exuviae, as previous locality, 10-VI-1976 (USNM); 1 pupa, 2 pupal exuviae (in glycerin), as previous locality, 10-VI-1976 (WLGC). F. flavescens : 1 pupal exuviae (of paratype), Luna, P.I., 25-X-1961 (CNCI). F. harpegonata : 1 pupa (of paratype), Puerto Rica, 23-I-1953 (USNM). F. herediae : 1 pupal exuviae (of paratype), Siquirres, Costa Rica, 1-VI-1956 (USNM). F. hutsoni : 1 pupa (of paratype), South Island, Aldabra, 1-17-II-1968 (USNM). F. hygrophila : 3 pupae, Sidney, British Columbia, Canada, 20-VIII-1947 (USNM); 2 pupal exuviae, no locality data (USNM). F. intermedia : 1 pupal exuviae (of paratype), nr. Siquirres, Costa Rica, 15-V-1956 (CNCI). F. jocosa : 1 pupal exuviae (of paratype), Mt. St. Benedict, Trinidad, 13-V-1953 (CNCI). F. lesliei : 2 pupae, Clark Hall, Dominica, 9-I-1965 (USNM); 1 pupa (of paratype of F. bicolor ), 2 pupal exuviae (of paratypes of F. bicolor ), Mayaquez, Puerto Rico, 3-II-1953 (USNM). F. luteigenua : 1 pupa (of paratype), 1 pupal exuviae (of paratype), Finca la Tigra, nr. La Virgen, Heredia Province, Costa Rica, 12-XI-1981 (USNM); 1 pupa (of paratype), as previous locality, 17-XI-1981 (USNM); 1 pupa (of paratype), as previous locality, 30-VII-1982 (USNM). F. malayae : 2 pupal exuviae (of holotype, paratype), Cameron’s Highlands, Malaysia, 29-I-1934 (CNCI). F. mortuifolii : 1 pupal exuviae, no locality data (USNM). F. musae : 2 pupal exuviae, km 123 Br 174, Manaus, Amazonas, Brazil, 29-VII-2005 (CNCI). F. nodosa : 1 pupal exuviae (of paratype), nr. Siquirres, Costa Rica, 15-VII-1956 (CNCI). F. oligarthra : 1 pupal exuviae (of paratype), Carcega Beach, Puerto Rico, 22-II-1953 (CNCI). F. pictoni : 1 pupa, 2 pupal exuviae, Siquirres, Costa Rica, 11-VI-1956 (USNM). F. pinicola : 2 pupal exuviae, no locality (USNM). F. pulchrithorax : 3 pupae, Cambridge, England, United Kingdom, 31-VIII-1922 (USNM). F. quasicornuta : 1 pupal exuviae (of paratype), nr. Siquirres, Costa Rica, 6-V-1050 (CNCI). F. seminole : 2 pupal exuviae, Hills, Puerto Rico, 6-II-1953 (USNM); 2 pupae, 2 pupal exuviae, Georgetown, Guyana, 20-V-1943 (USNM). F. sonora : 2 pupae, 2 pupal exuviae, Death Valley, Saratoga Springs, California, USA, 19-II-1955 (USNM). F.spinosa : 2 pupae (of paratypes), 1 pupal exuviae, Mayaquez, Puerto Rico, 3-II-1953 (USNM); 1 pupal exuviae (of paratype), as previous locality, 3-II-1953 (CNCI); 1 pupal exuviae, Ilheus, Bahia, Brazil, 20-X-1977 (USNM). F. terrestris : 1 pupal exuviae (of paratype), St. Augustine, Trinidad, 9-VII- 1957 (CNCI); 1 pupa (of paratype), 3 pupal exuviae (of paratype), Las Hermanas, Trinidad, 9-VII-1957 (USNM). F. trinidadensis : 1 pupal exuviae (of holotype), St. Augustine, Trinidad, 3-VII-1957 (CNCI). F. tuberculata : 1 pupa, 2 pupal exuviae, Hacienda Theobroma, Siquirres, Costa Rica, 1-VI-1956 (USNM); 1 pupa, Bayano Field Station, Panama Province, Panama, VI-1976 (USNM); 1 pupal exuviae (of paratype), Trinidad, 8-V-1953 (CNCI); 1 pupa (of paratype), 2 pupal exuviae (of paratypes), North Range, Trinidad, 8-V-1953 (USNM). F. usingeri : 3 pupae (of paratypes), Berkeley Hills, Alameda County, California, USA, X-1947 (USNM). F. varicolor : 1 pupal exuviae (of paratype), Rio de Janeiro, Brazil, 1-VIII-1923 (CNCI). F. wirthi : 1 pupal exuviae (of paratype), Alum Rock Park, Santa Clara County, California, USA, 8-IV-1948 (USNM). F. ( Phytohelea ) nr. antiguensis : 3 pupal exuviae, 1 pupal exuviae (in glycerin), Riverwoods Field Laboratory, nr. Cornwall, Highlands County, Florida, USA, 13-I-1999 (CNCI). F. ( Euprojoannisia ) sp.: 1 pupal exuviae, 52 km S of airport, Iquitos, Loreto, Peru, 1-II- 2003 (CNCI). F. ( Phytohelea ) sp.: 2 pupal exuviae, 2 pupal exuviae (in glycerin), Jardin Botanico, Rio Piedras, San Juan, Puerto Rico, 24-X-2006 (CNCI). F. sp.: 1 pupa, 1 pupal exuviae (in glycerin), Nebraska, USA (CNCI); 1 pupal exuviae, Rio Dantes, P.N. Barbilla, Cartago, Costa Rica, 10-II-2006 (CNCI); 2 pupal exuviae, Iquitos, Loreto, Peru, 1-II-2003 (CNCI); 1 pupal exuviae, Moung Sayaboury, Sayaboury Province, Laos, 19-II-1967 (BPBM); 1 pupal exuviae, as previous locality, 9-12-XII-1967 (BPBM).

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 8. Habitus. A. Austroconops mcmillani, male, in lateral view (from Borkent & Craig 2004). B. Leptoconops spinosifrons, male, in lateral view (from Laurence & Mathias 1972). C. Leptoconops gallicus, female, in dorsal and ventral view (from Clastrier 1972). D. Forcipomyia (Euprojoannisia) spatulifera, female, dorsal view (from Saunders 1957). E. Forcipomyia (Euprojoannisia) wirthi, female, in dorsal view (from Saunders 1957). F. Forcipomyia (Euprojoannisia) bromeliae, male, in dorsal view (from Saunders 1957). G. Forcipomyia (Caloforcipomyia) caerulea, female, in dorsal view (from Saunders 1957). H. Forcipomyia (Caloforcipomyia) varicolor, female, in dorsal view (from Saunders 1957).

FIGURE 9. Habitus. A. Forcipomyia (Schizoforcipomyia) petersoni, female, in dorsal view (from Chan & LeRoux 1971b). B. Forcipomyia (Metaforcipomyia) cerifera, female, in dorsal view (from Saunders 1957). C. Forcipomyia (Warmkea) lesliei, male, with female segment 9 separate, in dorsal view (from Saunders 1957). D. Forcipomyia (Warmkea) tuberculata, male, in dorsal view (from Saunders 1957). E. Forcipomyia (Microhelea) intonsa, female, with male segment 9 separate, in dorsal view (from Chan & LeRoux 1971a). F. Forcipomyia (Forcipomyia) brevipennis, female, with male segment 9 separate, in dorsal view (from Saunders 1924). G. Forcipomyia (Forcipomyia) ciliata, female, with male segment 9 separate, in dorsal view (from Saunders 1924). H. Forcipomyia (Forcipomyia) pulchrithorax, male, with female segment 9 separate, in dorsal view (from Saunders 1924).

FIGURE 10. Habitus. A. Forcipomyia (Forcipomyia) costata, female, with male segment 9 separate, in dorsal view (from Saunders 1924). B. Forcipomyia (Forcipomyia) costata, female, with male segment 9 separate, in ventral view (from Saunders 1924). C. Forcipomyia (Forcipomyia) costata, female, in lateral view (from Saunders 1924). D. Forcipomyia (Lepidohelea) bicolor, male, in dorsal view (from Saunders 1924). E. Forcipomyia (Phytohelea) grandis, male, in dorsal view (from Chan & LeRoux 1971a). F. Forcipomyia (Trichohelea) eques, female, in dorsal view (from Chan & LeRoux 1965). G. Forcipomyia (Trichohelea) crinita, male, with female segment 9 separate, in dorsal view (from Saunders 1964). H. Forcipomyia (Synthyridomyia) acidicola, male, in dorsal view (from Saunders 1957).

FIGURE 11. Habitus. A. Forcipomyia (Thyridomyia) rugosa, male, in dorsal view (from Chan & LeRoux 1970). B. Forcipomyia (Pterobosca) crinume, female, in ventral view (from Tokunaga 1932). C. Forcipomyia (Lasiohelea) rugosa, male, in dorsal view (from Saunders 1964). D. Atrichopogon hexastichus, male, in dorsal view (from Nielsen 1951). E. Atrichopogon wirthi, female, in dorsal view (from Chan & Linley 1988). F. Atrichopogon muelleri, sex unknown, in dorsal view, with larval exuviae on posterior of abdomen (from Szadziewski et al. 1997). G. Dasyhelea vericolor, sex unknown, in dorsal view (from Szadziewski et al. 1997). H. Schizonyxhelea bulla, sex unknown, in dorsal view (from Thomsen 1937).

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 14. Anterior portion of heads. A. Austroconops mcmillani, in ventral view, dorsal apotome not present. B. Leptoconops bequaerti, in anterior view (distorted slide specimen). C. Forcipomyia (Phytohelea) sp. in anterior view (Florida, USA). D. Forcipomyia (Phytohelea) sp. in anteroventral view (Florida, USA). E. Atrichopogon sp., in anterior view (from 6.5 km NW Enderby, BC, Canada). F. Dasyhelea sp., in anteroventral view (Herman Lake, 4 km SE Eagle Bay, BC, Canada).

FIGURE 18. Dorsal apotomes, anterior view. A. Austroconops mcmillani (from Borkent & Craig 2004). B. Leptoconops bequaerti. C. Forcipomyia (Phytohelea) n. sp. near F. antiguensis. D. Forcipomyia (Euprojoannisia) sp. (undescribed, from Peru). E. Atrichopogon sp. (undescribed, from Costa Rica). F. Dasyhelea insons (from Mayer 1934c). G. Dasyhelea tersa (from Mayer 1934c). H. Dasyhelea assimilis (from Mayer 1934c). I. Dasyhelea pseudocincta (from Waugh & Wirth 1976, modified). J. Dasyhelea pseudoincisurata (from Waugh & Wirth 1976, modified). K. Dasyhelea messersmithi (from Waugh & Wirth 1976, modified). L. Dasyhelea atlantis (from Waugh & Wirth 1976, modified).

FIGURE 23. Posterior portion of heads, in ventral view. A. Austroconops mcmillani. B. Leptoconops kerteszi. C. Forcipomyia (Euprojoannisia) sp. (from Peru). D. Forcipomyia (Phytohelea) n. sp. near antiguensis. E. Atrichopogon sp. (from St. Pierre de Wakefield, Quebec, Canada). F. Dasyhelea paracincta. G. Dasyhelea pseudoincisurata.

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 34. Legs, wings and apices of the antennae, in ventral view. A. Austroconops mcmillani. B. Leptoconops kerteszi. C. Forcipomyia (Euprojoannisia) sp. (from Peru). D. Atrichopogon sp. (from Lake Opinicon, Ontario, Canada). E. Dasyhelea paracincta. F. Culicoides sp. (from 3 km E. Carp, Ontario, Canada).

FIGURE 42. Respiratory organs, most in dorsal view, Fig. 42B in lateral view, pattern on tracheal tubes shown only in some. A. Austroconops mcmillani (from Borkent & Craig 2004). B. Austroconops mcmillani (from Borkent & Craig 2004). C. Leptoconops americanus (from Clastrier & Wirth 1978). D. Leptoconops asilomar (from Clastrier & Wirth 1978). E. Forcipomyia (Euprojoannisia) wirthi (from Saunders 1957). F. Forcipomyia (Euprojoannisia) longispina (from Saunders 1957). G. Forcipomyia (Warmkea) malayae (from Saunders 1957). H. Forcipomyia (Caloforcipomyia) caerulea (from Saunders 1957). I. Forcipomyia (Caloforcipomyia) varicolor (from Saunders 1957). J. Forcipomyia (Metaforcipomyia) cerifera (from Saunders 1957). K. Forcipomyia (Forcipomyia) genualis (from Saunders 1957, as raleighi). L. Forcipomyia (Forcipomyia) nigra (from Saunders 1924). M. Forcipomyia (Forcipomyia) ciliata (from Saunders 1924). N. Forcipomyia (Phytohelea) bromelicola (from Saunders 1925, (as an Apelma)). O. Forcipomyia (Synthyridomyia) acidicola (Synthyridomyia) (from Saunders 1924). P. Forcipomyia (Trichohelea) crinita (from Saunders 1964). Q. Forcipomyia (Lasiohelea) quasicornuta (from Saunders 1964). R. Forcipomyia (Lasiohelea) cornuta (from Saunders 1964). S. Atrichopogon wirthi (from Chan & Linley 1988). T. Atrichopogon thienemanni (from Lenz 1934).

FIGURE 47. Metathoraces and tergites 1 (in dorsal view). A. Austroconops mcmillani. B. Leptoconops kerteszi. C. Forcipomyia (Forcipomyia) bipunctata. D. Forcipomyia (Phytohelea) n. sp. nr. antiguensis. E. Atrichopogon sp. (from Lake Opinicon, Ontario, Canada). F. Dasyhelea pseudoincisuata.

FIGURE 55. Abdominal segments four, in dorsal and ventral view (shagreen not shown). A. Austroconops mcmillani (from Borkent & Craig 2004). B. Leptoconops kerteszi. C. Forcipomyia (Euprojoannisia) sp. (from Peru).

FIGURE 56. Abdominal segment four, in dorsal and ventral view (shagreen not shown). A. Forcipomyia dominicana. B. Atrichopogon sp. (from Lake Opinicon, Ontario, Canada). C. Dasyhelea paracincta.

FIGURE 72. Abdominal segments 9. A. Austroconops mcmillani, male, in dorsal and ventral view (from Borkent & Craig 2004). B. Leptoconops whitseli, male, in dorsal and ventral view (from Clastrier & Wirth 1978). C. Leptoconops whitseli, female, in dorsal and ventral view (from Clastrier & Wirth 1978). D. Leptoconops asilomar, male, in dorsal and ventral view (modified from Clastrier & Wirth 1978). E. Leptoconops asilomar, female, in dorsal and ventral view (modified from Clastrier & Wirth 1978). F. Forcipomyia anabaenae, female, in dorsal view (from Chan & Saunders 1965). G. Forcipomyia eques, male, in ventral view (from Chan & LeRoux 1965). H. Forcipomyia rugosa, female, in dorsal view (from Chan & LeRoux 1970). I. Forcipomyia brevis, female, in dorsal view (from Mayer 1934a). J. Atrichopogon wirthi, male, in dorsal view, arrow indicating anteriorly directed spines (from Chan & Linley 1988). K. Dasyhelea contigua, female, in ventral view (from Mayer 1934c). L. Dasyhelea grata, male, in ventral view (from Mayer 1934c).

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.