Bezzia 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 : 100-102

Bezzia Kieffer

( Figs. 12F , 13I , 17D , 22C–D , 28D , 31E , 33J , 41A , 46H–K , 53D , 69C , 77E–H )

DIAGNOSIS: Only pupa of Ceratopogonidae with the metathorax with only one campaniform sensillum (M-3-T) situated at least ⅓ the length of the metathorax from its anterior margin ( Fig. 53D ), apex of the halter extending posteriorly to about 1/6 length of tergite 2 (as in Fig. 33L ), abdominal segment 4 with V-5-IV, V-6-IV and V-7-IV closely approximated (as in Fig. 70C ) or, if V-7-IV is closer to L-4-IV then L-3-IV is closer to L-2-IV than to an elongate L-1-IV ( Fig. 69C ) (not as in Fig. 70B ), abdominal segment 8 has V-5-VIII and V-6-VIII on separate tubercles or if on partially to completely fused tubercles, then V-5-VIII well-developed (not minute), and abdominal segment 8 is without L-1-VIII (not diagnosable as different from Palpomyia and Phaenobezzia ); however, most species of Bezzia have two or more campaniform sensilla on the dorsal apotome ( Figs. 22C–D ), a nearly unique condition found only in two species of Palpomyia , one of which, P. jonesi , distinctively has two setae and two campaniform sensilla ( Fig. 22J ).

DESCRIPTION: Habitus as in Fig. 12F . Total length = 2.00– 6.16 mm. Without larval exuviae retained on abdomen. Exuviae with flagellum appressed against lateral margin of midleg, wing ( Figs. 17D , 33J ). Ecdysial tear around base of antenna, along lateral margin of face to palpus (as in Figs. 17C , 79H ) or with eye appressed to antenna ( Figs. 17D ). Head: Dorsal apotome ( Figs. 22C–D ), with partial ventral line of weakness, without dorsomedial tubercle, without central dome; dorsolateral cephalic sclerite ( Fig. 13I ) fused to scutum, each side separated medially by dorsal apotome in whole pupa; mouthparts ( Fig. 28D ) with mandible well-developed, lacinia absent; palpus extending posterior to posterolateral margin of labium; labium entire or separated medially by labrum, hypopharynx; apex of antenna ( Fig. 41A ) anterior to posterior to, posterior extent of midlength portion of midleg (portion lateral to mesosternum), narrowed posteriorly; sensilla: dorsal apotomals ( Figs. 22C–D )—1 moderate to elongate seta, 1–3 campaniform sensilla; dorsolateral cephalic sclerite sensilla—1 seta, 0–1 campaniform sensillum; clypeal-labrals ( Fig. 28D )—1 slender or 2 slender or thick setae; oculars ( Fig. 28D )—1–2 seta, 1 campaniform sensillum. Thorax: Prothoracic extension ( Fig. 28D ) wide, well-developed, extending from palpus to antenna; mesonotum without tubercles, not extending posteromedially, not dividing metathorax medially ( Fig. 53D ); respiratory organ ( Figs. 46H–K ) length/width = 3.73–6.60, moderately elongate to elongate, apical portion swollen in some, somewhat flattened apically, with pores closely abutting at apex of respiratory organ, arranged in single straight to curved row, outer surface with some wrinkles, with short, wide pedicel, base with short to moderately elongate posteromedial apodeme, membranous base of respiratory organ short, annulated, tracheal tube straight to slightly curved along length, with spirals restricted to base, wrinkles to half length or more; wing ( Fig. 41A ) without apical tubercle or angle, separated medially by fore-, midlegs; halter apex and hind leg ( Fig. 33J ) broadly abutting; halter apex extending posteriorly to 1/6 length of tergite 2; legs ( Fig. 41A ) with lateral margin of foreleg near midlength of wing evenly curved; hind leg visible at lateral margin of wing ( Fig. 33J ); with apex of foreleg moderately anterior to apex of midleg; apex of hind leg abutting apex of midleg laterally or small gap between the two; sensilla: anteromedials—2 elongate setae, 1 campaniform sensillum (as in Figs. 31L–M ); anterolaterals—1 moderately long seta; dorsal setae ( Fig. 31E )—D-1-T, D-2-T, D-4-T, D-5-T setae, D-3-T campaniform sensillum, D-3-T lateral to D-4-T; supraalar 2—campaniform sensillum; metathoracics ( Fig. 53D )—1 campaniform sensillum; M-3-T distant from margin of metathorax (at least 1/3 length of metathorax). Abdomen: pigmentation light brown, with tergite 1 with 3 medial spots, tergites 2–7 with medial area with stripe, 2 anterolateral spots, sternites 3–7 with medial stripe, anterolateral spot (light brown), segment 2 as wide or slightly wider than segment 3, segments with undivided, thin to thick setae, with rounded to pointed, short to moderately elongate tubercles, tergites or sternites entire, each without membranous disc; segment 9 ( Figs. 77E–H ) not strongly modified, terminal processes closely approximated to separated basally, each projecting posterodorsolaterally, tapering to pointed apex, in some very slender; sensilla: tergite 1 ( Fig. 53D ) with 8 setae, 2 campaniform sensilla, including 3–4 lateral sensilla, D-2-I, D-3-I closely approximated, D-7-I situated anterolaterally near L-1-I to anteriorly near D-3-I; segment 4 ( Fig. 69C )—D-2-IV, D-3-IV short setae on short tubercles; D-5-IV, D-8-IV, D-9-IV short to moderately elongate setae, D-7-IV present or absent; D-5-IV on single, short tubercle, D-8-IV, D-9-IV on basally fused or 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, if present, near D-3-IV; L-1-IV short to moderately elongate seta on rounded tubercle, well anterior of posterior lateral setae; L-2- IV, L-3-IV, L-4-IV short to moderately elongate setae on rounded tubercles, L-2-IV, L-3-IV on single tubercle in some, V-5-IV, V-6-IV, V-7-IV short setate on short tubercles, all closely approximated or with V-7-IV closer to L-4- IV; segment 8 without D-3-VIII, without L-1-VIII; with V-5-VIII, V-6-VIII on single tubercle, V-5-VIII tiny, V-6- VIII elongate; segment 9 ( Figs. 77E–H )—with D-5-IX, D-6-IX campaniform sensilla.

DISTRIBUTION AND HABITAT: The genus Bezzia is known from 319 species from every Region worldwide ( Borkent 2014). Immatures have been recorded from a wide array of habitats including rice fields, springs, streams, river margins, wet moss, various phytotelmata, pools, marshes, bogs, fens, algal mats in lentic habitats, lakes, and reservoirs.

TAXONOMIC DISCUSSION: The pupae of 51 species of Bezzia are known ( Tables 2–3 ). Goeze (1780) described but did not name a species which is probably a Bezzia , providing the first morphological study describing the chaetotaxy of the larva and pupa ( Fig. 6B ) of a Ceratopogonidae . It was later named Tipula goezii by Schrank (1803). I have placed it in Bezzia as a new combination as follows but the name is so out of date that it must be considered a nomen nudum. The long slender terminal process and somewhat clavate apex of the respiratory organ ( Figs. 46H, 46J ) is a distinctive combination of many species of Bezzia .

Bezzia goezii (Schrank), 1803: 72 ( Tipula ). Quedlinburg, Germany. new combination.

Some species have been described more than once by the same author ( Table 2 ) but with different names (now synonyms) (e.g. B. circumdata , B. leucogaster , B. nobilis ), suggesting either misidentifications or the possibility of more than one species actually present.

Thomsen (1937) gave a key to three species known to her at that time but one of these, P. flavitarsis , is now recognized as a Palpomyia . Thomsen (1937) drew the dorsal apotomes of species of Bezzia with more than one seta (per side) but likely misinterpreted the multiple campaniform sensilla as the broken bases of setae.

Thienemann (1928) and Mayer (1934a) provided keys to European species groups. Lenz (1934) provided a key to European species but primarily used habitats to distinguish these and, for most, this almost certainly results in inaccurate identifications. Harris (1981) described and keyed the pupae of six unnamed species of Bezzia from Australia.

Wirth (1983a, 1983b, 1983c) diagnosed the pupae of some Nearctic and Neotropical Bezzia in the cockerelli , bicolor , and nobilis species groups but did not do so for the remaining species groups of Bezzia . At present, pupae of Bezzia cannot be diagnosed as a genus and therefore providing a key to the species in a given region is superfluous. Unless a diagnosis becomes available all Bezzia should be keyed with species of Palpomyia and Phaenobezzia . Alternately, it would be possible to key all species with more than one campaniform sensillum on the dorsal apotome, which would include nearly all Bezzia and a few Palpomyia , depending on the area being covered.

Nearly all Bezzia have more than one campaniform sensillum on the dorsal apotome (see character 6). The only exceptions seen are of B. brevicornis , B. bromeliae and B. dorsasetula , each with one seta and one campaniform sensillum.

Bezzia xanthogaster (Kieffer), 1919: 130 ( Probezzia ) is a junior homonym of Probezzia xanthogaster (Kieffer), 1917: 329 ( Bezzia ) and is here given the new name Bezzia gilvigaster (a similar specific name also meaning yellow stomach (abdomen for the adult this species).

MATERIAL EXAMINED: B. africana : 1 pupal exuviae, Burgershall , Hazyview, Transvaal, South Africa, 3- XII-1973 ( NMSA) ; 1 pupal exuviae, Magaliesberg Agricultural School , Transvaal, South Africa, 13-XI-1973 ( NMSA) . B. albicornis : 1 pupal exuviae, Kaushut , Tedjen Province, Turkmenistan, 2-IV-1972 ( ZIN) ; 1 pupal exuviae, as previous locality, 30-V-1972 ( ZIN) . B. amana : 1 pupal exuviae (of holotype), Loutpan , Transvaal, South Africa, 10-I-1974 ( NMSA) . B. annulipes : 2 pupal exuviae, Lunzer Untersee between boat house and canal, Lower Austria, Austria, 6-VI-1942 ( ZSMC) ; 1 pupal exuviae, Germany ( ZSMC) ; 3 pupal exuviae, Germany ( ZSMC) ; 12 pupal exuviae, Anninskoe lake , Pskov Province, Russia, 24-VI-1995 ( ZIN) . B. biannulata : 1 pupal exuviae (of paratype), 1 pupal exuviae (of allotype), Oceano Beach, San Luis Obispo County, California, USA, 20- VIII-1948 ( USNM) . B. bicolor : 6 pupal exuviae, Gievenbeck , Germany, 9-V-1912 ( ZSMC) ; 4 pupal exuviae, Lunz , Austria ( USNM) ; 4 pupal exuviae, Shushary , Leningrad Province, Russia, 29-V-1997 ( ZIN) ; 2 pupal exuviae, Strelna , Leningrad Province, Russia, 12-V-2002 ( ZIN) . B. bivittata : 3 pupal exuviae, Salisbury , Wicomico County, Maryland, USA, 4-V-1981 ( WLGC) ; 1 pupal exuviae, as previous locality, 13-IV-1981 ( WLGC) . B. blantoni : 2 pupal exuviae (of paratype), Escobar , Buenos Aires, Argentina, 10-I-1982 ( MPLA) . B. brevicornis : 1 pupal exuviae, Magdalena , Buenos Aires, Argentina, 24-XII-1981 ( MPLA) . B. bromeliae : 2 pupal exuviae (of paratype), Bayano Field Station , Panama Province, Panama, VI-1976 ( MPLA) . B. circumdata : 3 pupal exuviae, Black Lake , Stanleyville, Ontario, Canada, 24-VI-1975 ( USNM) ; 1 pupal exuviae, as previous locality, 25-VI-1975 ( USNM) ; 4 pupal exuviae, Rotmoos on Mittersee , Lower Austria, Austria, 10-VI-1942 ( ZSMC) ; 1 pupa, 5 pupal exuviae, Grosser Plöner See , Slesvig-Holstein, Germany, summer 1952 ( ZSMC) ; 2 pupa, 5 pupal exuviae, kerosene port on Dortmund-Ems canal, Dortmund, North Rhine-Westphalia, Germany, 28-VII-1908 ( ZSMC) ; 1 pupa, 3 pupal exuviae, Germany ( ZSMC) ; 4 pupal exuviae, Vyborg , Leningrad Province, Russia, 27-VII-1998 ( ZIN) . B. cockerelli : 1 pupal exuviae, Valdez , Alaska, USA, 1948 ( USNM) ; 1 pupal exuviae, Bainville , Roosevelt County, Montana, USA, 9-VI-1969 ( USNM) ; 1 pupal exuviae, Trails Pond , Latah County, Idaho, USA, 22-VII-1969 ( USNM) ; 5 pupal exuviae, 4 mi N of Upham , McHenry County, North Dakota, USA, 5-VI-1969 (4 USNM, 1 VPIC) . B. collessi : 1 pupal exuviae (of paratype), Singapore, 10-IX-1952 ( USNM) ; 3 pupal exuviae (of paratypes), as previous locality, 28-VIII-1952 ( USNM) . B. dorsasetula : 2 pupal exuviae, College Park , Prince George’s County, Maryland, USA, 30-V-1975 ( USNM) , 1 pupal exuviae, no locality, 6-V-1977 ( VPIC) . B. fascispinosa : 1 pupal exuviae, Lakeland Pond , College Park, Prince George’s County, Maryland, USA, 30-V-1975 ( USNM) ; 1 pupal exuviae, Beaver Lake Reservoir, Pocahontas State Park, Chesterfield County, Virginia, USA, 30-IV-1977 ( VPIC) ; 1 pupal exuviae, Iron Spring , Elliot Know, Augusta County, Virginia, USA, 22-17-1977 ( VPIC) . B. flavicornis : 1 pupal exuviae, Priluki , Belarus, 17-V-1967 ( ZIN) . B. flavicorporis : 1 pupal exuviae, Loutpan , Transvaal, South Africa, 10-I-1974 ( NMSA) . B. gibbera : 1 pupal exuviae, Dismal Swamp , Camden County, North Carolina, USA, 25-III-1976 ( VPIC) . B. glabra : 2 pupal exuviae, Morgan Arboretum , St. Anne de Bellevue, Quebec, Canada, 1964 ( USNM) ; 1 pupal exuviae, 1 mi. S of Corapeake, Dismal Swamp , Camden County, North Carolina, USA, 29-VII-1976 ( VPIC) ; 2 pupal exuviae, Keowee Reserve , Seneca , Oconee County, South Colorado, USA, 2-V-1975 ( VPIC) . B. japonica : 1 pupal exuviae, Suputinka river , Ussuri Nature Reserve, Primorskii Territory, Russia, 30-V-1973 ( ZIN) . B. kuhetiensis : 1 pupal exuviae, Issyk-Kul lake , Kyrgyzstan, 16-VI-1971 ( ZIN) . B. laciniastyla : 1 pupal exuviae, Jackson River , approx. 10 mi. upriver from Covington, Natural Well, Allegheny County, Virginia, USA, 4-VI-1977 ( VPIC) . B. leucogaster : 4 pupal exuviae, shore of Lottsee SE of Mölln, Slesvig-Holstein, Germany, 16-V-1926 ( ZSMC) , 2 pupal exuviae, shore of Kirchsee , Preetz (between Kiel and Plön), Slesvig-Holstein, Germany, 11-VII1918 ( ZSMC) ; 1 pupal exuviae, Severskii Donets River , Donetsk Province, Ukraine, 5-V-1970 ( ZIN) . B. narynica : 1 pupal exuviae, Issyk-Kul lake , Kyrgyzstan, 24-V-1971 ( ZIN) . B. nigrita : 1 pupal exuviae, Severskii Donets River , Donetsk Province, Ukraine, 5-V-1970 ( ZIN) ; 1 pupal exuviae, Malinovka , Donetsk Province, Ukraine, 8-V-1970 ( ZIN) . B. nobilis : 1 pupal exuviae, 7 km S of Hope , BC, Canada, 8–9-VII-1985 ( CNCI) ; 1 pupal exuviae, Beaver Creek , Lawrence County, South Dakota, USA, 15-VI-1969 ( USNM) ; 1 pupal exuviae, Cow Neck Salt Marsh , North Sea, New York, USA, 19-IV-1956 ( NYSM) ; 1 pupal exuviae, as previous locality, 17-IV-1956 ( NYSM) ; 4 pupal exuviae, Fishing Creek , Newcomb, New York, USA, 28-V-1958 ( NYSM) ; 1 pupal exuviae, Port Leyden , New York, USA, 4-VII-1959 ( USNM) ; 2 pupal exuviae, Letchworth State Park , New York, USA, 13-VI-1963 ( USNM) ; 1 pupal exuviae, Patuxent Refuge , Maryland, USA, 8-V-1958 ( USNM) ; 1 pupal exuviae, Vepco Property , Louisa County, Virginia, USA, 12-IX-1976 ( VPIC) ; 1 pupal exuviae, Washburn County, Wisconsin, USA, 1-IX-1951 ( VPIC) ; 2 pupal exuviae, Vepco Property , Louisa County, Virginia, USA, 12-IX-1976 ( VPIC) ; 2 pupal exuviae, Great Swamp , Isle of Wight County, Virginia, USA, 2-VIII- 1976 ( VPIC) ; 1 pupal exuviae, Cranberry Glade , Pocahontas County, Virginia, USA, 8-IX-1976 ( VPIC) ; 1 pupal exuviae, Norfolk Gardens , Norfolk County, Virginia, USA, 1-VIII-1976 ( VPIC) . 1 pupal exuviae, no locality/date ( VPIC); 1 pupal exuviae, Pecan Springs , Devils River, Juno, Texas, USA ( USNM) ; 1 pupal exuviae, Baton Rouge , Louisiana, USA, 4-III-1947 ( USNM) ; 1 pupal exuviae, Beloe lake , Alol’, Pskov Province, Russia, 27-VI-1969 ( ZIN) ; 1 pupal exuviae, Beloe lake , Alol’, Pskov Province, Russia, 5-VII-1969 ( ZIN) . B. obelisca : 2 pupal exuviae, Mer Bleue , Ottawa, Ontario, Canada, 27-V-1960 ( USNM) ; 2 pupal exuviae, McLean Reserve , Tompkins County, New York, USA, 18-VI-1963 ( USNM) ; 1 pupal exuviae, edge of Big Glade , Cranberry Glades Botanical Area, Pocahontas County, Virginia, USA, 6-IX-1976 ( VPIC) . B. perplexa : 1 pupal exuviae, Millpond Creek , North Anna Resrvoir, Louisa County, Virginia, USA, 1-III-1977 ( VPIC) . B. pulchripes : 2 pupal exuviae, Mission Rio Cururu Igarape , Brazil, 4-V-1941 ( SDEI) ; 1 pupal exuviae, Mocoreta , Corrientes, Argentina, 21-III-1985 ( MPLA) . B. roldani : 2 pupal exuviae, no locality, 20-VIII-1979 ( MPLA) . B. saileri : 1 pupal exuviae (of paratype), Fire Lake , Anchorage, Alaska, USA, 29-V-1948 ( USNM) . B. signata : 1 pupal exuviae, Luchevoi , Republic of Karelia, Russia, 14-VI-1966 ( ZIN) . B. sordida : 1 pupal exuviae, Olerna , Marin County, California, USA, 22-II-1947 ( USNM) . B. turkmenica : 1 pupal exuviae, Hodzhaz-Kala bay , Kyzyl-Arvat Province, Turkmenistan, 18-IV-1972 ( ZIN) . B. uncistyla : 2 pupal exuviae, 5 km E of Danby , Vermont, USA, 25-VI-1986 ( CNCI) ; 1 pupal exuviae, as previous locality, 25–26-VI-1986 ( CNCI) . B. varicolor : 1 pupal exuviae, Rideau River , Ottawa, Ontario, Canada, 29-V-1960 ( USNM) ; 1 pupal exuviae, Lake Temescal , Berkeley, California, USA, 1-V-1948 ( USNM) ; 1 pupal exuviae, Lake Tomahawk at Kemp Biological Station, Oneida County, Wisconsin, USA, 20-VII-1978 ( USNM) ; 2 pupal exuviae, Bar Lake , Benzie County, Michigan, USA, 8-VIII-1975 ( VPIC) ; 1 pupal exuviae (of topotype), Mill Creek Brook, Noyack, New York, USA, 6-V-1057 ( USNM) ; 1 pupal exuviae, Slaterville Spr. , New York, USA, 12- VI-1964 ( VPIC) ; 1 pupal exuviae, Lakeland Pond , College Park, Prince George’s County, Maryland, USA, 24-IV- 1977 ( VPIC) . B. nr. bicolor : 1 pupal exuviae, no locality, 30-V-1953 ( USNM) . B. nr. nobilis : 1 pupal exuviae, Back Bay National Wildlife Refuge , Virginia Beach, Virginia, USA, 30-VIII-1975 ( VPIC) . B. nr. obelisca : 1 pupal exuviae, Mer Bleue , Ottawa, Ontario, Canada, 27-V-1960 ( USNM) ; 2 pupal exuviae, no data ( USNM); 1 pupal exuviae, Mer Bleue , Ottawa, Ontario, Canada, 27-V-1960 ( VPIC) ; 1 pupal exuviae, Big Glade , Cranberry Glade Botanical Area, Pocahontas County, Virginia, USA, 8-IX-1976 ( VPIC) . B. nr. pruinosa : 1 pupal exuviae, no locality, 12-V-1953 ( USNM) . B. nr. varicolor : 1 pupal exuviae, W flank of Salt Pond Mountain , Giles County, Virginia, USA, 14-VII-1977 ( VPIC) . B. sp.: 2 pupal exuviae (in glycerin), 12 pupal exuviae, Bolean Lake , 6 km NE of Falkland, British Columbia, Canada, 12-13-VII-1989 ( CNCI) ; 1 pupal exuviae (in glycerin), Spanish Lake , 6 km E of Falkland, 50°29.12N 119°28.07W, British Columbia, Canada, 27–28-V-2008 ( CNCI) ; 4 pupal exuviae, 3 km E of Salmon Arm , British Columbia, Canada, 9-VI-1988 ( CNCI) ; 9 pupal exuviae, 6 km E of Salmon Arm , British Columbia, Canada, 6-VI-1990 ( CNCI) ; 5 pupal exuviae, 9 km S of Salmon Arm , British Columbia, Canada, 19-VII-1988 ( CNCI) ; 2 pupal exuviae, 8 km E of Sicamous , British Columbia, Canada, 1-VI-1992 ( CNCI) ; 2 pupal exuviae, 6.5 km NW of Enderby , British Columbia, Canada, 6-V-1992 ( CNCI) ; 4 pupal exuviae, 30 km N of Nakusp , British Columbia, Canada, 5-VII-1990 ( CNCI) ; 4 pupal exuviae, 20 km E of Anola , Manitoba, Canada, 16-VI-1990 ( CNCI) ; 3 pupal exuviae, Trail Bay , Manitoba, Canada ( CNCI) ; 1 pupal exuviae (in glycerin), 1 mi. E of Orleans , Ontario, Canada, 8-VII-1971 ( CNCI) ; 1 pupal exuviae, 6 km N of Eardlay , Quebec, Canada, 15-V-1986 ( CNCI) ; 2 pupal exuviae, Fire Lake , Anchorage, Alaska, USA, 29-V-1948 ( USNM) ; 5 pupal exuviae, 9 km W of Okanogan , Washington, USA, 13-VI-2008 ( CNCI) ; 9 pupal exuviae, High Creek Fen Preserve , 9 mi S of Fairplay Park, Colorado, USA, 18-VI-1995 ( CNCI) ; 1 pupal exuviae, Blue Mountain Lake , New York, USA, 30-VI-1958 ( WLGC) ; 2 pupal exuviae, Ringwood Pond , Dryden, New York, USA, 7-III-1933 ( USNM) ; 1 pupal exuviae, Keowee Reserve , Seneca, Ocouee County, South Carolina, USA, 16-V-1974 ( USNM) ; 1 pupal exuviae, as previous locality, USA, 16-VII-1974 ( USNM) ; 1 pupal exuviae, 8 km W of Atenas , Costa Rica, 19-VIII-1993 ( CNCI) ; 2 pupal exuviae, carreterra entre Santa Cecilia y Upala, 1.6 km N.O. de BVirmania, Dos Rios, Upala , Alejuela, Costa Rica, 16-X-2006 ( INBC) ; 1 pupal exuviae, Barro Colorado Island , Canal Zone, Panama, 24-VI- 1996 ( CNCI) ; 2 pupal exuviae, as previous locality, 31-X-1995 ( CNCI) ; 1 pupal exuviae, as previous locality, 30- VI-1996 ( CNCI) ; 1 pupal exuviae, as previous locality, 10-IX-1995 ( CNCI) ; 3 pupae, Lerchenauer See , Germany, 26-VII-1990, 27-VII-1990, 2-VIII-22-IX-1990 ( CNCI) ; 1 pupa, Chiemsee , Germany, 13-VIII-1990 ; 1 pupa, 1 pupal exuviae, Bottsand nr. Stein, Germany, 5-VI-1932 ( ZSMC) ; 2 pupal exuviae, Darban , Western Australia, Australia, 28-X-1986 ( ANIC) ; 9 pupal exuviae, as previous locality, 28-X-1985 ( ANIC) ; 2 pupal exuviae, 5 km N of Darban , Western Australia, Australia, 29-X-1985 ( ANIC) ; 3 pupal exuviae, Roper River Mission, Northern Territory, Australia, 8-XI-1956 ( ANIC) ; 3 pupal exuviae, Humpty Doo , Northern Territory, Australia, V-1957 ( ANIC) ; 1 pupal exuviae, Dayboro , Queensland, Australia, 10-VII-1951 ( ANIC) ; 10 pupal exuviae, Nattai River , Mittagong, New South Wales, Australia, 4-XI-1964 ( ANIC) ; 1 pupal exuviae, as previous locality, 15-XI-1968 ( ANIC) ; 1 pupal exuviae, Gap Creek , The Crags, Mittagong, New South Wales, Australia, 10-II-1966 ( ANIC) ; 5 pupal exuviae, Damin clearing, The Crags, Mittagong, New South Wales, Australia, 9-II-1966 ( ANIC) ; 2 pupal exuviae, Deep Creek , Nanasheen, New South Wales, Australia, 7-XII-1956 ( ANIC) ; 2 pupal exuviae, as previous locality, 25-XI-1956 ( ANIC) ; 21 pupal exuviae, Middle Creek , New South Wales, Australia, 8-II-1966 ( ANIC) ; 1 pupal exuviae, Colo Vale , New South Wales, Australia, 16-X-1956 ( ANIC) ; 2 pupal exuviae, Hornsby , New South Wales, Australia, 18-XI-1969 ( ANIC) ; 2 pupal exuviae, as previous locality, 25-X-1956 ( ANIC) ; 2 pupal exuviae, Nepean River , Menangle, New South Wales, Australia, 9-XII-1968 ( ANIC) ; 1 pupal exuviae, Griffith , New South Wales, Australia, 16-XI-1956 ( ANIC) ; 1 pupal exuviae, Moruya River , Merricumbene Creek, New South Wales, Australia, 2-III-1964 ( ANIC) ; 1 pupal exuviae, McCarrs Creek , New South Wales, Australia, 14-I-1969 ( ANIC) ; 1 pupal exuviae, no locality, 14-I-1937 ( SAIM) .

Figures

FIGURE 6. Previous schemes for naming of sensilla and/or tubercles on abdominal segments of pupae of Ceratopogonidae. Homologies between authors given in Table 1. A. Derham's (1713) earliest description of a ceratopogonid pupa. B. Goeze's (1780) earliest system, likely a species of Bezzia. Letters refer to various parts. y and z refer to lateral sensilla. C. Mayer's (1934a) system of sensilla on segment 3, with model and three examples shown. Black squares refer to the position of sensilla. Other abbreviations as indicated. D. Tokunaga's (1937a) system for Culicoides, for various abdominal segments. sp refers to the spiracular scar.

FIGURE 12. Habitus. A. Pellucidomyia leei, male, in dorsal view (from Debenham 1970). B. Dibezzia prominens, sex unknown, in dorsal view (form Mayer 1934c). C. Macropeza albitarsis, male, in dorsal view (from Szadziewski & Dominiak, 2007, modified). D. Mallochohelea munda, male, in ventral view (from Szadziewski et al. 1997). E. Sphaeromias fasciata, sex unknown, in dorsal view (from Mayer 1934a). F. Bezzia glabra, female, in ventral view (from Thomsen 1937). G. Palpomyia sp., male, in dorsal view (from Grogan & Wirth 1979). H. Paryphoconus oliveirai, female, in ventral view (modified from Ronderos et al. 2007).

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 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 22. Dorsal apotomes, anterior view. A. Sphaeromias longipennis (from Wirth & Grogan 1979). B. Amerohelea sordidipes (from Lane et al. 1955). C. Bezzia mollis (from Mayer 1934c). D. Bezzia roldani (from Spinelli & Wirth 1989). E. Bezzia brevicornis (from Spinelli 1983a). F. Clastrieromyia dycei (from Spinelli & Grogan 1986). G. Pachyhelea pachymera (from Spinelli 1983a). H. Palpomyia lineata (from Grogan & Wirth 1979). I. Palpomyia belkini (from Grogan & Wirth 1979, modified). J. Palpomyia flaviceps (from Grogan & Wirth 1979). K. Phaenobezzia opaca (from Wirth & Grogan 1982). L. Paryphoconus oliveirai (from Ronderos et al. 2007).

FIGURE 28. Posterior portion of heads, in ventral view. A. Leehelea hispida. B. Sphaeromias longipennis. C. Xenohelea galatea. D. Bezzia nobilis. E. Clastrieromyia dycei. F. Pachyhelea pachymera. G. Palpomyia subasper. H. Phaenobezzia opaca. I. Paryphoconus grandis.

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 41. Legs, wings and apices of the antennae, in ventral view. A. Bezzia nobilis. B. Clastrieromyia dycei (reconstructed from distorted specimen). C. Pachyhelea pachymera. D. Palpomyia occidentalis. E. Phaenobezzia opaca. F. Paryphoconus oliveirai (reconstructed from distorted specimen).

FIGURE 46. Respiratory organs, most in dorsal view, Figs. 46G. in lateral view, pattern on tracheal tubes shown only in some. A. Homohelea albitudinis (from de Meillon & Wirth 1981). B. Homohelea delanoe (from de Meillon & Wirth 1981). C. Leehelea wasselli (from Elson-Harris 1987). D. Leehelea hispida (from Elson-Harris 1987). E. Sphaeromias longipennis (from Wirth & Grogan 1979). F. Xenohelea galatea. G. Amerohelea sordidipes (from Lane et al. 1955). H. Bezzia glabra (from Wirth 1983b). I. Bezzia obelisca (from Wirth 1983a). J. Bezzia dorsasetula (from Wirth 1983a). K. Bezzia turkmenica (from Glukhova 1979b). L. Clastrieromyia dycei (from Spinelli & Grogan 1986). M. Pachyhelea pachymera (from Spinelli 1983a). N. Palpomyia lineata (from Grogan & Wirth 1979). O. Palpomyia jonesi (from Grogan & Wirth 1979). P. Palpomyia belkini (from Grogan & Wirth 1979). Q. Palpomyia rufa (from Grogan & Wirth 1979). R. Palpomyia flaviceps (from Grogan & Wirth 1979). S. Phaenobezzia opaca (from Wirth & Grogan 1982). T. Paryphoconus oliveirai (from Ronderos et al. 2007).

FIGURE 53. Metathoraces and tergites 1, in dorsal view. A. Leehelea hispida. B. Sphaeromias longipennis. C. Xenohelea galatea. D. Bezzia nobilis. E. Clastrieromyia dycei. F. Pachyhelea pachymera.

FIGURE 69. Abdominal segments four, in dorsal and ventral view (shagreen not shown). A. Sphaeromias longipennis. B. Xenohelea galatea. C. Bezzia nobilis.

FIGURE 70. Abdominal segments four, in dorsal and ventral view (shagreen not shown). A. Clastrieromyia dycei. B. Pachyhelea pachymera. C. Palpomyia jonesi.

FIGURE 77. Abdominal segments 9. A. Homohelea delanoe, female, in dorsal view (from de Meillon & Wirth 1981). B. Leehelea wasselli, female, in dorsal view (from Elson-Harris 1987). C. Sphaeromias longipennis, male, in dorsal and ventral view (from Wirth & Grogan 1979). D. Xenohelea galatea, female, in dorsal and ventral view (shagreen not shown). E. Bezzia roldani, male, in dorsal view (from Spinelli & Wirth 1989). F. Bezzia roldani, female, in dorsal view (from Spinelli & Wirth 1989). G. Bezzia brevicornis, male, in dorsal view (from Spinelli 1983a). H. Bezzia acanthodes, female, in dorsal view (from Spinelli 1983a). I. Clastrieromyia dycei, male, in ventral view (from Spinelli & Grogan 1986). J. Clastrieromyia dycei, female, in ventral view (from Spinelli & Grogan 1986). K. Pachyhelea pachymera, male, in ventral view (from Spinelli 1983a). L. Pachyhelea pachymera, female, in dorsal view (from Spinelli 1983a).

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.