Ceratpogonidae -- fossil Diptera cat.

Family CERATOPOGONIDAE

The Ceratopogonidae, members of which are commonly called punkies, no-see-ums, or biting midges, are a cosmopolitan family of flies with over 5,000 fossil and extant species (A. Borkent, pers. comm.). Adults of extant forms feed on a variety of food sources including nec tar and vertebrate and invertebrate blood. Some species are predaceous, others are known as vectors of arboviruses and various parasitic diseases. Larval habitats are diverse and include moss, rotting wood, decaying plant matter, animal excrement, ponds, lakes (some in benthos), streams and rivers, wet soil, the water of leaf axils, and partially submerged substrates where they feed on detritus, fungi, algae, or small organisms.

Over 200 species of fossil Ceratopogonidae have been described in over 40 genera. Many undescribed species exist in collections. Zherichin & Sukacheva (1973) recorded 305 undetermined specimens of Ceratopogonidae from the Cretaceous amber deposits of Siberia. Only a portion of these have been described (Remm, 1976; Szadziewski, 1996). Langenheim et al. (1960) recorded an undetermined specimen from the Cretaceous Alaskan amber. Other Cretaceous amber fossils of Ceratopogonidae have been described by Szadziewski & Schlüter (1992), Grogan & Szadziewski (1988), Boesel (1937), and Borkent (1995, 1996). The oldest records of this family are specimens from the Lower Cretaceous amber of Lebanon and Austria (Schlee, 1984). Some Lebonese amber specimens have been described recently by Szadziewski (1996).

Metz (1987) discussed the possibility that trace fossils from the Eocene Green River shales mentioned by Moussa (1970) were ceratopogonid larval trails. Additional records of the family in the Tertiary are by Poinar (1992), who recorded undetermined material from the Oligocene/Miocene amber of Chiapas, Mexico, and Soom (1984), who recorded material of this family from the Paleocene amber of Switzerland.

Crosskey (1988) removed two species of Simulium from Simuliidae and transferred them to "?Ceratopogonidae". They are listed here under questionably placed species at the end of the family chapter.
Ref.: Szadziewski (1988, world checklist of amber taxa; 1993, Saxonian amber taxa), Borkent (1995, Cretaceous Canadian spp., world cat. fossil spp.).

Genus ADELOHELEA Borkent

ADELOHELEA Borkent, 1995: 45. Type species: Adelohelea glabra Borkent, 1995, by original designation.
glabra Borkent, 1995: 46. NE: Canada (Cretaceous) [A].

Genus ALAUTUNMYIA Borkent

ALAUTUNMYIA Borkent, 1996: 11. Type species: Alautunmyia elongata Borkent, 1996, by original designation.
elongatus Borkent, 1996: 13. USA (New Jersey) (Cretaceous) [A].

Genus ALLUAUDOMYIA Kieffer

*ALLUAUDOMYIA Kieffer, 1913b: 12. Type species: Alluaudomyia imparunguis Kieffer, 1913, by original designation.
succinea Szadziewski, 1988: 112. PA: Baltic Region (Eocene/Oligocene) [A].

Genus ARCHAUSTROCONOPS Szadziewski

ARCHIAUSTROCONOPS Szadziewski, 1996: 34. Type species: Archiaustroconops ceratoformis Szadziewski, 1996, by original designation.
ceratoformis Szadziewski, 1996: 34. PA: Lebanon (Lower Creatceous) [A].

Genus ARCHICULICOIDES Szadziewski

ARCHICULICOIDES Szadziewski, 1996: 29. Type species: Archiculicoides schleei Szadziewski, 1996, by original designation.
schleei Szadziewski, 1996: 29. PA: Lebanon (Lower Cretaceous) [A].

Genus ATRICHOPOGON Kieffer

*ATRICHOPOGON Kieffer, 1906: 53 (as Ceratopogon subgenus). Type species: Ceratopogon exilis Coquillett, 1902 [= Ceratopogon levis Coquillett, 1901], by subsequent designation of Coquillett (1910: 512).
brunnescens Statz, 1944c: 141. PA: Germany (Oligocene) [C].
eocenicus Szadziewski, 1988: 223. PA: Baltic Region (Eocene/Oligocene) [A].
Unidentified sp.-PA: Japan (Pleistocene) [K] (Saigusa, 1974: 425).
Unidentified sp.-PA: Germany (Miocene) [A] (Schumann & Wendt, 1989: 41).
Unidentified sp. NT: Dominican Republic (Oligocene/Miocene) [A] (Szadziewski & Grogan, 1994: 221).

Genus ATRICULICOIDES Remm

ATRICULICOIDES Remm, 1976: 345 (108). Type species: Atriculicoides macrophthalmus Remm, 1976, by original designation.
cenomanensis Szadziewski & Schlüter, 1992: 74. PA: France (Upper Cretaceous) [A].
dasyheleis Szadziewski, 1996: 55. PA: Russia (Siberia) (Upper Cretaceous) [A].
globosa Boesel, 1937: 47 (Lasiohelea). NE: Canada (Upper Cretaceous) [A].

cretea Boesel, 1937: 46 (Lasiohelea). NE: Canada (Upper Cretaceous) [A].

;
incompletus Szadziewski & Schlüter, 1992: 76. PA: France (Upper Cretaceous) [A].
macrophthalmus Remm, 1976: 346 (108). PA: Russia (Siberia) (Upper Cretaceous) [A].

squamiciliatus Remm, 1976: 346 (110). PA: Russia (Siberia) (Upper Cretaceous) [A].

sibiricus Szadziewski, 1996: 36. PA: Russia (Siberia) (Upper Cretaceous) [A].
taimyricus Szadziewski, 1996: 56. PA: Russia (Siberia) (Upper Cretaceous) [A].

Genus AUSTROCONOPS Wirth & Lee

*AUSTROCONOPS Wirth & Lee, 1959: 337. Type species: Austroconops mcmillani Wirth & Lee, 1959, by original designation.
borkenti Szadziewski & Schlüter, 1992: 78. PA: France (Upper Cretaceous) [A].
cretaceous Szadziewski, 1996: 36. PA: Lebanon (Lower Cretaceous) [A].
fossilis Szadziewski, 1996: 38. PA: Lebanon (Lower Cretaceous) [A].
gondwanicus Szadziewski, 1996: 38. PA: Lebanon (Lower Cretaceous) [A].
sibiricus Szadziewski, 1996: 36. PA: Russia (Siberia) (Upper Cretaceous) [A].
?Unidentified sp.-AU: Australia (Upper Cretaceous) [C] (Szadziewski, 1990a: 231).

Genus BAEODASYMYIA Clastrier & Raccurt

*BAEODASYMYIA Clastrier & Raccurt, 1979: 100. Type species: Baeodasymyia modesta Clastrier & Raccurt, 1979, by original designation.
dominicana Szadziewski & Grogan, 1994: 220. NT: Dominican Republic (Oligocene/Miocene) [A].

Genus BEZZIA Kieffer

*BEZZIA Kieffer, 1899: 69. Type species: Ceratopogon ornatus Meigen, 1830, by original designation.
eocenica Szadziewski, 1988: 187. PA: Baltic Region (Eocene/Oligocene) [A].
longipennis Statz, 1944c: 151. PA: Germany (Oligocene) [C].

Genus BRACHYCRETACEA Szadziewski

BRACHYCRETACEA Szadziewski, 1996: 63. Type species: Baeohelea taimyricxa Remm, 1976, by original designation.
taimyrica Remm, 1976: 115 (351] (Baeohelea). PA: Russia (Siberia) (Upper Cretaceous) [A].

Genus BRACHYPOGON Kieffer

*BRACHYPOGON Kieffer, 1899: 69. Type species: Ceratopogon vitiosus Winnertz, 1852, by original designation.

Subgenus BRACHYPOGON Kieffer

balticus Szadziewski, 1988: 93. PA: Baltic Region (Eocene/Oligocene) [A].
eocenicus Szadziewski, 1988: 91. PA: Baltic Region (Eocene/Oligocene) [A].
gedanicus Szadziewski, 1988: 95. PA: Baltic Region (Eocene/Oligocene) [A].
miocaenicus Szadziewski, 1993: 610. PA: Germany (Miocene) [A].
Unidentified sp. -NT: Dominican Republic (Oligocene/Miocene) [A] (Szadziewski & Grogan, 1994: 221).

Subgenus ISOHELEA Kieffer

*ISOHELEA Kieffer, 1917b: 295 (as genus). Type species: Ceratopogon lacteipennis Winnertz, 1852 [preoccupied, = Ceratopogon sociabilis Goetghebuer, 1920], by original designation.
henningseni Szadziewski, 1988: 89. PA: Baltic Region (Eocene/Oligocene) [A].
polonicus Szadziewski, 1988: 88. PA: Baltic Region (Eocene/Oligocene) [A].
prominulus Meunier, 1904c: 228 [1904d: 239] (Ceratopogon). PA: Baltic Region (Eocene/Oligocene) [A]; PA: Germany (Miocene) [A].

Unplaced to Subgenus of Brachypogon Kieffer

?Unidentified sp.-PA: Sicily (Miocene) [A] (Szadziewski, 1988: 12).

Genus CERATOCULICOIDES Wirth & Ratanaworabhan

*CERATOCULICOIDES Wirth & Ratanaworabhan, 1971: 170. Type species: Helea longipennis Wirth, 1952, by original designation.
danicus Szadziewski, 1988: 102. PA: Baltic Region (Eocene/Oligocene) [A].

Genus CERATOPALPOMYIA Szadziewski

CERATOPALPOMYIA Szadziewski, 1988: 171. Type species: Ceratopalpomyia eocenica Szadziewski, 1988, by original designation.
eocenica Szadziewski, 1988: 173. PA: Baltic Region (Eocene/Oligocene) [A].

Genus CERATOPOGON Meigen

*CERATOPOGON Meigen, 1803: 261. Type species: Tipula barbicornis Linnaeus, 1767 (as "barbicornis Fabr.") [misidentification, = Ceratopogon communis Meigen, 1804], by monotypy.
*CERAPOGON Rafinesque, 1815: 130 (unnecessary new replacement name for Ceratopogon Meigen). Type species: Tipula barbicornis Meigen, 1803 [misidentification, = Ceratopogon communis Meigen, 1804], automatic.
APEDELIA Loew, 1850: 30 (as Ceratopogon subgenus). Nomen nudum.
EUPEDELIA Loew, 1850: 30 (as Ceratopogon subgenus). Nomen nudum.
bitterfeldi Szadziewski, 1993: 613. PA: Germany (Miocene) [A].
ceranowiczi Szadziewski, 1988: 77. PA: Baltic Region (Eocene/Oligocene) [A].
crypticus Szadziewski, 1988: 70. PA: Baltic Region (Eocene/Oligocene) [A].
eminens Meunier, 1904c: 229 [1904d: 240]. PA: Baltic Region (Eocene/Oligocene) [A].

eucerus Keilbach, 1982: 348. Nomen nudum.

forcipiformis Meunier, 1904c: 235 [1904d: 246]. PA: Baltic Region (Eocene/ Oligocene) [A]; PA: Germany (Miocene) [A].

defectus Meunier, 1904c: 229 [1904d: 240]. PA: Baltic Region (Eocene/ Oligocene) [A].

gedanicus Szadziewski, 1988: 74. PA: Baltic Region (Eocene/Oligocene) [A].
grogani Szadziewski, 1988: 64. PA: Baltic Region (Eocene/Oligocene) [A].
heini Keilbach, 1982: 348. Nomen nudum.
hennigi Szadziewski, 1988: 60. PA: Baltic Region (Eocene/Oligocene) [A]; PA: Germany (Miocene) [A].
kotejai Szadziewski, 1993: 616. PA: Germany (Miocene) [A].
longicornis Keilbach, 1982: 348. Nomen nudum.
macronyx Remm, 1976: 113 (350). PA: Russia (Siberia) (Upper Cretaceous) [A].
margaritae Szadziewski, 1988: 80. PA: Baltic Region (Eocene/Oligocene) [A].
minimus Keilbach, 1982: 348. Nomen nudum.
miocaenicus Szadziewski, 1993: 617. PA: Germany (Miocene) [A].
nanalobus Borkent & Grogan, 1995: 97. PA: Baltic Region (Eocene/Oligocene) [A].
paraeminens Borkent & Grogan, 1995: 99. PA: Baltic Region (Eocene/Oligocene) [A].
pectinatus Keilbach, 1982: 348. Nomen nudum.
pilosus Keilbach, 1982: 348. Nomen nudum.
piotrowskii Szadziewski, 1988: 75. PA: Baltic Region (Eocene/Oligocene) [A].
pisinnus Borkent & Grogan, 1995: 97. PA: Baltic Region (Eocene/Oligocene) [A].
remmicola Szadziewski, 1988: 71. PA: Baltic Region (Eocene/Oligocene) [A].
ritzkowskii Szadziewski, 1988: 78. PA: Baltic Region (Eocene/Oligocene) [A].
subeminens Szadziewski, 1993: 617. PA: Germany (Miocene) [A].
succinicola Szadziewski, 1993: 621. PA: Germany (Miocene) [A].
terminalis Keilbach, 1982: 349. Nomen nudum.
tertiaricus Szadziewski, 1988: 62. PA: Baltic Region (Eocene/Oligocene) [A].
ungulatus Keilbach, 1982: 349. Nomen nudum.
ungulinus Keilbach, 1982: 349. Nomen nudum.
Unidentified sp.-PA: Sicily (Miocene) [A] (Guérin-Méneville, 1838a: 170).

Genus CULICOIDES Latreille

*CULICOIDES Latreille, 1809: 251. Type species: Culicoides punctata Latreille, 1809 [preoccupied, =Ceratopogon punctatus Meigen, 1804], by monotypy.
*PADROSIA Rafinesque, 1815: 130 (unnecessary replacement name for Culicoides Meigen). Type species: Ceratopogon punctatus Meigen, 1804, automatic.

Subgenus OECACTA Poey

*OECACTA Poey, 1853: 238 (as genus). Type species: Oecacta furens Poey, 1853, by monotypy.
balticus Szadziewski, 1988: 41. PA: Baltic Region (Eocene/Oligocene) [A].
dasyheleiformis Szadziewski, 1988: 36. PA: Baltic Region (Eocene/Oligocene) [A].
speciosus Meunier, 1904c: 229 [1904d: 240] (Ceratopogon). PA: Baltic Region (Eocene/Oligocene) [A]; PA: Germany (Miocene) [A].
succineus Remm, 1976: 113 (349). PA: Russia (Siberia) (Upper Cretaceous) [A].
succivarius Szadziewski, 1988: 40. PA: Baltic Region (Eocene/Oligocene) [A].

Unplaced to Subgenus of Culicoides Latreille

abbreviatipennis Statz, 1944c: 142. PA: Germany (Oligocene) [C].
abdominalis Statz, 1944c: 145. PA: Germany (Oligocene) [C]. [Preoccupied by abdominalis Santos Abreu, 1918; no new replacement name proposed here.]
agamus Borkent, 1995: 62. NE: Canada (Cretaceous) [A].
annosus Borkent, 1995: 65. NE: Canada (Cretaceous) [A].
atratus Statz, 1944c: 147. PA: Germany (Oligocene) [C].
austerus Statz, 1944c: 144. PA: Germany (Oligocene) [C].
bicolor Statz, 1944c: 146. PA: Germany (Oligocene) [C].
bifidus Borkent, 1996: 19. NE: USA (Cretaceous) [A].
bullus Borkent, 1995: 61. NE: Canada (Cretaceous) [A].
canadensis Boesel, 1937: 47 (Atrichopogon). NE: Canada (Cretaceous) [A].
carri Pierce, 1966: 91. NE: USA (Miocene) [P].
casei Grogan & Szadziewski, 1988: 809. NE: USA (Cretaceous) [A].
ceranowiczi Szadziewski, 1988: 45. PA: Baltic Region (Eocene/Oligocene) [A]; PA: Germany (Miocene) [A].
elongatulus Statz, 1944c: 143. PA: Germany (Oligocene) [C].
eoselficus Szadziewski, 1988: 43. PA: Baltic Region (Eocene/Oligocene) [A].
filipalpis Remm, 1976: 111 (347). PA: Russia (Siberia) (Upper Cretaceous) [A]; NE: Canada (Upper Cretaceous) [A].
fossilis Pierce, 1966: 91. NE: USA (Miocene) [P].
gedanensis Szadziewski, 1988: 46. PA: Baltic Region (Eocene/Oligocene) [A].
gracilior Statz, 1944c: 143. PA: Germany (Oligocene) [C].
grandibocus Borkent, 1996: 19. NE: USA (Cretaceous) [A].
jucundus Statz, 1944c: 147. PA: Germany (Oligocene) [C].

jucundus Statz, 1941: fig. 8. Nomen nudum.

kaluginae Remm, 1976: 110 (347). PA: Russia (Siberia) (Upper Cretaceous) [A].
laurae Pierce, 1966: 91. NE: USA (Miocene) [P].
liliputanus Statz, 1944c: 142. PA: Germany (Oligocene) [C].
megacanthus Palmer, 1957: 272. NE: USA (Miocene) [P].
miocenicus Pierce, 1966: 93. NE: USA (Miocene) [P].
obesus Statz, 1944c: 145. PA: Germany (Oligocene) [C].
obscuratus Statz, 1944c: 144. PA: Germany (Oligocene) [C].
obuncus Borkent, 1995: 64. NE: Canada (Cretaceous) [A].
prussicus Szadziewski, 1988: 48. PA: Baltic Region (Eocene/Oligocene) [A].
sphenostylus Remm, 1976: 112 (347). PA: Russia (Siberia) (Upper Cretaceous) [A].
subgedanicus Szadziewski, 1993: 608. PA: Germany (Miocene) [A].
tenuipennis Statz, 1944c: 147. PA: Germany (Oligocene) [C].
tyrrelli Boesel, 1937: 48 (Dasyhelea). NE: Canada (Upper Cretaceous) [A].
Unidentified sp.-NE: Canada (Upper Cretaceous) [A] (Downes, 1978: 58).
Unidentified sp. -NT: Dominican Republic (Oligocene/Miocene) [A] (Szadziewski & Grogan, 1994: 221).

Genus DASYHELEA Kieffer

*DASYHELEA Kieffer, 1911b: 5. Type species: Dasyhelea halophila Kieffer, 1911, by monotypy.
antiqua Palmer, 1957: 266 (as australis Wirth subspecies). NE: USA (Miocene) [P].
browneae Pierce, 1966: 87. NE: USA (Miocene) [P].
dara Pierce, 1966: 88. NE: USA (Miocene) [P].
eodicryptoscenica Szadziewski, 1988: 228. PA: Baltic Region (Eocene/Oligocene) [A].
gedanica Szadziewski, 1988: 226. PA: Baltic Region (Eocene/Oligocene) [A].
judithae Pierce, 1966: 88. NE: USA (Miocene) [P].
kanakoffi Pierce, 1966: 89. NE: USA (Miocene) [P].
miocaenica Szadziewski, 1993: 651. PA: Germany (Miocene) [A].
stanislavi Szadziewski, 1988: 230. PA: Baltic Region (Eocene/Oligocene) [A].
stenoceras Palmer, 1957: 271. NE: USA (Miocene) [P].
Unidentified sp. -NT: Dominican Republic (Oligocene/Miocene) [A] (Szadziewski & Grogan, 1994: 221).

Genus EOHELEA Petrunkevitch

EOHELEA Petrunkevitch, 1956: 675 [1957: 208]. Type species: Eohelea stridulans Petrunkevitch, 1956, by monotypy.
fossicola Szadziewski, 1993: 624. PA: Germany (Miocene) [A].
gedanica Szadziewski, 1988: 165. PA: Baltic Region (Eocene/Oligocene) [A].
grogani Szadziewski, 1988: 164. PA: Baltic Region (Eocene/Oligocene) [A].
miocaenica Szadziewski, 1993: 625. PA: Germany (Miocene) [A].
petrunkevitchi Szadziewski, 1984: 39 [1985: 127]. PA: Baltic Region (Eocene/ Oligocene) [A].
sakhalinica Szadziewski, 1990b: 78. PA: Russia (Sakhalin I) (Paleocene/Eocene) [A].
sinuosa Meunier, 1904c: 234 [1904d: 245] (Ceratopogon). PA: Baltic Region (Eocene/Oligocene) [A]; PA: Germany (Miocene) [A].

stridulans Petrunkevitch, 1956: 675 [1957: 209]. PA: Baltic Region (Eocene/Oligocene) [A]

Genus FORCIPOMYIA Meigen

*FORCIPOMYIA Meigen, 1818: 73, 75. Type species: Tipula bipunctata Linnaeus, 1767, automatic (by designation of type species for Labidomyia).
[Generic name originally proposed in synonymy with Ceratopogon, but available under I.C.Z.N. Code Article 11(d).]
*LABIDOMYIA Stephens, 1829a: 52 [1829b: 239] (unnecessary new replacement name for Forcipomyia Meigen). Type species: Tipula bipunctata Linnaeus, 1767, by subsequent designation of Westwood (1840: 126).

Subgenus EUPROJOANNISIA Brèthes

*EUPROJOANNISIA Brèthes, 1914: 155 (as genus). Type species: Euprojoannisia platensis Brèthes, 1914, by original designation.
berendti Szadziewski, 1988: 209. PA: Baltic Region (Eocene/Oligocene) [A].
henningseni Szadziewski, 1988: 211. PA: Baltic Region (Eocene/Oligocene) [A].
miocaenica Szadziewski, 1993: 647. PA: Germany (Miocene) [A].
piriformis Meunier, 1904c: 228 [1904d: 239] (Ceratopogon). PA: Baltic Region (Eocene/Oligocene) [A].
tuberculosa Szadziewski, 1993: 648. PA: Germany (Miocene) [A].

Subgenus FORCIPOMYIA Meigen

eocostata Szadziewski, 1988: 202. PA: Baltic Region (Eocene/Oligocene) [A].
gedanicola Szadziewski, 1988: 200. PA: Baltic Region (Eocene/Oligocene) [A]; PA: Germany (Miocene) [A].
lyneborgi Szadziewski, 1988: 196. PA: Baltic Region (Eocene/Oligocene) [A].
pseudomicrohelea Szadziewski, 1988: 205. PA: Baltic Region (Eocene/Oligocene) [A].
subgedanicola Szadziewski, 1993: 645. PA: Germany (Miocene) [A].
turbinata Meunier, 1904c: 227 [1904d: 238] (Ceratopogon). PA: Baltic Region (Eocene/Oligocene) [A].
uncula Meunier, 1904c: 227 [1904d: 238] (Ceratopogon). PA: Baltic Region (Eocene/Oligocene) [A].
unculiformis Szadziewski, 1993: 646. PA: Germany (Miocene) [A].
Unidentified sp.-PA: Russia (Sakhalin I) (Paleocene/Eocene) [A] (Szadziewski, 1990: 79).

Subgenus LASIOHELEA Kieffer

*LASIOHELEA Kieffer, 1921: 115 (as genus). Type species: Atrichopogon pilosipennis Kieffer, 1919 [= Ceratopogon velox Winnertz, 1852], by original designation.
succinea Szadziewski, 1988: 194. PA: Baltic Region (Eocene/Oligocene) [A].
Unidentified sp.-PA: Russia (Sakhalin I) (Paleocene/Eocene) [A] (Szadziewski, 1990: 79).

Subgenus PHYTOHELEA Remm

*PHYTOHELEA Remm, 1971: 189. Type species: Ceratopogon bromelicola Lutz, 1914, by original designation.
eophytoheleana Szadziewski, 1988: 215. PA: Baltic Region (Eocene/Oligocene) [A].

Subgenus TRICHOHELEA Goetghebuer

*TRICHOHELEA Goetghebuer, 1920: 39 (as genus). Type species: Trichohelea tonnoiri Goetghebuer, 1920 [= Apelma auronitens Kieffer, 1919], by original designation.
bifidicola Szadziewski, 1993: 649. PA: Germany (Miocene) [A].
eotrichoheleana Szadziewski, 1988: 213. PA: Baltic Region (Eocene/Oligocene) [A].
succinicola Szadziewski, 1993: 650. PA: Germany (Miocene) [A].

Unplaced to Subgenus of Forcipomyia Meigen

eobreviflagellata Szadziewski, 1988: 220. PA: Baltic Region (Eocene/Oligocene) [A].
krzeminskii Szadziewski, 1988: 217. PA: Baltic Region (Eocene/Oligocene) [A].
kulickae Szadziewski, 1988: 219. PA: Baltic Region (Eocene/Oligocene) [A].
Unidentified sp.-NE: Canada (Upper Cretaceous) [A] (Downes & Wirth, 1981: 396).
Unidentified sp.-PA: Russia (Sakhalin I) (Paleocene/Eocene) [A] (Szadziewski, 1990: 78).
Unidentified sp. -NT: Dominican Republic (Oligocene/Miocene) [A] (Szadziewski & Grogan, 1994: 221).

Genus FOSSIHELEA Szadziewski

FOSSIHELEA Szadziewski, 1988: 104. Type species: Ceratopogon gracilitarsis Meunier, 1904, by original designation.
gracilitarsis Meunier, 1904c: 235 [1904d: 246] (Ceratopogon). PA: Baltic Region (Eocene/Oligocene) [A].
miocaenica Szadziewski, 1993: 627. PA: Germany (Miocene) [A].

Genus FOSSILEPTOCONOPS Szadziewski

FOSSILEPTOCONOPS Szadziewski, 1996: 46. Type species: Fossileptoconops lebanicus Szadziewski, 1996, by original designation.
lebanicus Szadziewski, 1996: 47. PA: Lebanon (Lower Cretaceous) [A].

Genus GEDANOHELEA Szadziewski

GEDANOHELEA Szadziewski, 1988: 166. Type species: Gedanohelea loewi Szadziewski, 1988, by original designation.
loewi Szadziewski, 1988: 168. PA: Baltic Region (Eocene/Oligocene) [A].
succinea Szadziewski, 1988: 169. PA: Baltic Region (Eocene/Oligocene) [A].
wirthi Szadziewski, 1988: 170. PA: Baltic Region (Eocene/Oligocene) [A].

Genus HELEAGERON Borkent

HELEAGERON Borkent, 1995: 47. Type species: Heleageron arenatus Borkent, 1995, by original designation.
arenatus Borkent, 1995: 48. NE: Canada (Cretaceous) [A].
grimaldii Borkent, 1996: 15. NE: USA (New Jersey) (Cretaceous) [A].

Genus HETEROMYIA Say

*HETEROMYIA Say, 1825: pl. 35. Type species: Heteromyia fasciata Say, 1825, by monotypy.
Unidentified sp.-NT: Dominican Republic (Oligocene/Miocene) [A] (Schlee, 1980: 50).

Genus LEBANOCONOPS Szadziewski

LEBANOCONOPS Szadziewski, 1996: 40. Type species: Lebanoconops lebanicus Szadziewski, 1996, by original designation.
lebanicus Szadziewski, 1996: 42. PA: Lebanon (Lower Cretaceous) [A].
minutus Szadziewski, 1996: 43. PA: Lebanon (Lower Cretaceous) [A].
wirthi Szadziewski, 1996: 44. PA: Lebanon (Lower Cretaceous) [A].

Genus LEBANOCULICOIDES Szadziewski

LEBANOCULICOIDES Szadziewski, 1996: 30. Type species: Lebanoculicoides mesozoicus Szadziewski, 1996, by original designation.
mesozoicus Szadziewski, 1996: 30. PA: Lebanon (Lower Cretaceous) [A].

Genus LEPTOCONOPS Skuse

*LEPTOCONOPS Skuse, 1889: 288. Type species: Leptoconops stygius Skuse, 1889, by monotypy.
boreus Kalugina, 1991: 78 (1992: 76). PA: Russia (Siberia) (Upper Cretaceous) [A].
copiosus Borkent, 1996: 3. USA (Upper Cretaceous) [A].
curvachelus Borkent, 1996: 7. USA (Upper Cretaceous) [A].
primaevus Borkent, 1995: 30. NE: Canada (Cretaceous) [A].
sibiricus Szadziewski, 1996: 49. PA: Russia (Siberia) (Upper Cretaceous) [A].
succineus Szadziewski, 1988: 233. PA: Baltic Region (Eocene/Oligocene) [A].
Unidentified sp.-PA: Russia (Sakhalin I) (Paleocene/Eocene) [A] (Szadziewski, 1990: 79).
Unidentified sp.-NE: Canada (Upper Cretaceous) [A] (Downes & Wirth, 1981: 396).
Unidentified sp.-PA: France (Upper Cretaceous) [A] (Szadziewski & Schlüter, 1992: 78).

Genus MANTOHELEA Szadziewski

MANTOHELEA Szadziewski, 1988: 144. Type species: Ceratopogon lacus Meunier, 1904, by original designation.
gedanica Szadziewski, 1988: 146. PA: Baltic Region (Eocene/Oligocene) [A]; PA: Germany (Miocene) [A].
laca Meunier, 1904c: 232 [1904d: 243] (Ceratopogon). PA: Baltic Region (Eocene/Oligocene) [A].

Genus MEUNIEROHELEA Szadziewski

*MEUNIEROHELEA Szadziewski, 1988: 148. Type species: Meunierohelea nielseni Szadziewski, 1988, by original designation.
*CHIMAEROHELEA Debenham, 1988: 801. Type species: Chimaerohelea caligula Debenham, 1988, by original designation.
gedanicola Szadziewski, 1988: 153. PA: Baltic Region (Eocene/Oligocene) [A].
miocaenica Szadziewski, 1993: 622 (Chimaerohelea). PA: Germany (Miocene) [A].
nielseni Szadziewski, 1988: 150. PA: Baltic Region (Eocene/Oligocene) [A].
wirthi Szadziewski, 1988: 157. PA: Baltic Region (Eocene/Oligocene) [A].

Genus MINYOHELEA Borkent

MINYOHELEA Borkent, 1995: 68. Type species: Minyohelea pumilus Borkent, 1995, by original designation.
pumilus Borkent, 1995: 69. NE: Canada (Cretaceous) [A].
schleei Szadziewski, 1996: 45. PA: Lebanon (Lower Cretaceous) [A].

Genus MONOHELEA Kieffer

*MONOHELEA Kieffer, 1917b: 294. Type species: Monohelea hieroglyphica Kieffer, 1917, by original designation.
baltica Szadziewski, 1988: 130. PA: Baltic Region (Eocene/Oligocene) [A].
clunipes Loew, 1850b: 30 (Ceratopogon). PA: Baltic Region (Eocene/Oligocene) [A]; PA: Germany (Miocene) [A].

Genus NANNOHELEA Grogan & Wirth

*NANNOHELEA Grogan & Wirth, 1980: 374. Type species: Ceratopogon bourioni Clastrier, 1961, by original designation.
eocenica Szadziewski, 1988: 101. PA: Baltic Region (Eocene/Oligocene) [A].
grogani Szadziewski, 1988: 99. PA: Baltic Region (Eocene/Oligocene) [A].
Unidentified sp. -NT: Dominican Republic (Oligocene/Miocene) [A] (Szadziewski & Grogan, 1994: 221).

Genus NEOCULICOIDES Pierce

NEOCULICOIDES Pierce, 1966: 93. Type species: Neoculicoides jeanneae Pierce, 1966, by original designation.
jeanneae Pierce, 1966: 94. NE: USA (Miocene) [P].

Genus NEUROHELEA Kieffer

*NEUROHELEA Kieffer, 1925: 112. Type species: Ceratopogon luteitarsis Meigen, 1838, by monotypy.
cothurnata Meunier, 1904c: 231 [1904d: 242] (Ceratopogon). PA: Baltic Region (Eocene/Oligocene) [A].

flagellus Meunier, 1904c: 230 [1904d: 241] (Ceratopogon). PA: Baltic Region (Eocene/Oligocene) [A].

Genus PALAEOBRACHYPOGON Borkent

PALAEOBRACHYPOGON Borkent, 1995: 70. Type species: Palaeobrachypogon vetus Borkent, 1995, by original designation.
remmi Borkent, 1995: 76. NE: USA (Cretaceous) [A].
vetus Borkent, 1995: 78. NE: Canada (Cretaceous) [A].

Genus PALPOMYIA Meigen

*PALPOMYIA Meigen, 1818: 82. Type species: Ceratopogon flavipes Meigen, 1804 (as flavipes with Palpomyia geniculata Meigen in synonymy), by monotypy.
[Generic name originally published in synonymy with Ceratopogon but available under I.C.Z.N. Code Art. 11(d).]
MIOPALPOMYIA Pierce, 1966: 95. Type species: Miopalpomyia shilo Pierce, 1966, by original designation.
NEOPALPOMYIA Pierce, 1966: 95. Type species: Neopalpomyia freyi Pierce, 1966, by original designation.
PARAPALPOMYIA Pierce, 1966: 97. Type species: Parapalpomyia ryshkoffi Pierce, 1966, by original designation.
edwardsi Cockerell, 1921c: 469. PA: UK (England) (Eocene/Oligocene) [C].
erikae Szadziewski, 1993: 642. PA: Germany (Miocene) [A].
freyi Pierce, 1966: 95 (Neopalpomyia). NE: USA (Miocene) [P].
jantari Szadziewski, 1988: 182. PA: Baltic Region (Eocene/Oligocene) [A].
multispinosa Pierce, 1966: 97 (Neopalpomyia). NE: USA (Miocene) [P].
riedeli Szadziewski, 1988: 183. PA: Baltic Region (Eocene/Oligocene) [A].
ryshkoffi Pierce, 1966: 98 (Parapalpomyia). NE: USA (Miocene) [P].
shilo Pierce, 1966: 95 (Miopalpomyia). NE: USA (Miocene) [P].
succinea Szadziewski, 1988: 185. PA: Baltic Region (Eocene/Oligocene) [A].
unca Hong, 1981: 58. PA: China (Eocene) [A].
Unidentified sp. -NT: Dominican Republic (Oligocene/Miocene) [A] (Szadziewski & Grogan, 1994: 221).

Genus PARACULICOIDES Pierce

PARACULICOIDES Pierce, 1966: 94. Type species: Paraculicoides rouseae Pierce, 1966, by original designation.
rouseae Pierce, 1966: 94. NE: USA (Miocene) [P].

Genus PERONEHELEA Borkent

PERONEHELEA Borkent, 1995: 80. Type species: Peronehelea chrimikalydia Borkent, 1995, by original designation.
chrimikalydia Borkent, 1995: 82. NE: Canada (Cretaceous) [A].
frigidus Remm, 1976: 115 (350) (Ceratopogon). PA: Russia (Siberia) (Upper Cretaceous) [A].
sibiricus Szadziewski, 1996: 69. PA: Russia (Siberia) (Upper Cretaceous) [A].

Genus PHAENOBEZZIA Haeselbarth

PHAENOBEZZIA Haeselbarth, 1965: 297. Type species: Probezzia pistiae Ingram & Macfie, 1921, by original designartion.
Unidentified sp. -NT: Dominican Republic (Oligocene/Miocene) [A] (Szadziewski & Grogan, 1994: 221).

Genus PHYSOHELEA Grogan & Wirth

*PHYSOHELEA Grogan & Wirth, 1979: 53. Type species: Neurohelea oedidactyla Ingram & Macfie, 1931, by original designation.
obtusa Meunier, 1904c: 235 [1904d: 246] (Ceratopogon). PA: Baltic Region (Eocene/Oligocene) [A].

cothurnatulus Meunier, 1904c: 231 [1904d: 242] (Ceratopogon). PA: Baltic Region (Eocene/Oligocene) [A].

Genus PROTOCULICOIDES Boesel

PROTOCULICOIDES Boesel, 1937: 50. Type species: Protoculicoides depressus Boesel, 1937, by original designation.
depressus Boesel, 1937: 51. NE: Canada (Upper Cretaceous) [A].
succineus Szadziewski, 1996: 31. PA: Lebanon (Lower Cretaceous) [A].

Genus SERROMYIA Meigen

*SERROMYIA Meigen, 1818: 83. Type species: Ceratopogon femoratus Meigen, 1804, by monotypy.
alphea Heyden, 1870: 251 (Ceratopogon). PA: Germany (Oligocene) [C]; PA: Germany (Miocene) [A].

austera Statz, 1941: fig. 10. Nomen nudum.

austera Statz, 1944c: 150. PA: Germany (Oligocene) [C].

colorata Statz, 1944c: 150. PA: Germany (Oligocene) [C].

spinofemorata Statz, 1941: fig. 9. Nomen nudum.

spinofemorata Statz, 1944c: 151. PA: Germany (Oligocene) [C].

anomalicornis Loew, 1850b: 30 (Ceratopogon). PA: Baltic Region (Eocene/ Oligocene) [A].
polonica Szadziewski, 1988: 135. PA: Baltic Region (Eocene/Oligocene) [A].
ryszardi Borkent in Borkent & Bisset, 1990: 207. PA: Baltic Region (Eocene/ Oligocene) [A].
sinuosa Borkent in Borkent & Bisset, 1990: 207. PA: Baltic Region (Eocene/ Oligocene) [A].
spinigera Loew, 1850b: 30 (Ceratopogon). PA: Baltic Region (Eocene/Oligocene) [A]; PA: Germany (Miocene) [A].

elongatus Meunier, 1904c: 242 [1904d: 253] (Ceratopogon). PA: Baltic Region (Eocene/Oligocene) [A].

succinea Szadziewski, 1988: 136. PA: Baltic Region (Eocene/Oligocene) [A].

Genus STILOBEZZIA Kieffer

*STILOBEZZIA Kieffer, 1911a: 118. Type species: Stilobezzia festiva Kieffer, 1911, by original designation.

Subgenus ACANTHOHELEA Kieffer

*ACANTHOHELEA Kieffer, 1917a: 198 (as genus). Type species: Acanthohelea pruinosa Kieffer, 1917, by monotypy.
PALEOTETRAGONEURA Meunier, 1920b: 1221 (897). Type species: Tetragoneura veterana Meunier, 1920, by monotypy.
falcata Meunier, 1904c: 233 [1904d: 244] (Ceratopogon). PA: Baltic Region (Eocene/Oligocene) [A]; PA: Germany (Miocene) [A].

spinosa Meunier, 1904c: 234 [1904d: 245] (Ceratopogon). PA: Baltic Region (Eocene/Oligocene) [A].

kutscheri Szadziewski, 1993: 637. PA: Germany (Miocene) [A].
saxonica Szadziewski, 1993: 638. PA: Germany (Miocene) [A].
succinea Szadziewski, 1993: 639. PA: Germany (Miocene) [A].
veterana Meunier, 1920b: 897 (1221) (Tetragoneura). PA: Germany (Oligocene) [C].

goetghebueri Statz, 1944c: 148. PA: Germany (Oligocene) [C].

Unidentified sp.-Unidentified sp.-PA: Russia (Sakhalin I) (Paleocene/Eocene) [A] (Szadziewski, 1990: 78).

Subgenus STILOBEZZIA Kieffer

Unidentified sp.-PA: Baltic Region (Eocene/Oligocene) [A] (Szadziewski, 1988: 124).

Unplaced to Subgenus of Stilobezzia Kieffer

Unidentified sp. -NT: Dominican Republic (Oligocene/Miocene) [A] (Szadziewski & Grogan, 1994: 221).

Genus WASHINGTONHELEA Wirth & Grogan

WASHINGTONHELEA Wirth & Grogan, 1988: 97. Type species: Washingtonhelea frommeri Wirth & Grogan, 1988, by original designation.
aquilonius Boesel, 1937: 449 (Ceratopogon). NE: Canada (Cretaceous) [A].
taimyrica Szadziewski, 1996: 71. PA: Russia (Siberia) (Upper Cretaceous) [A].

Genus WIRTHOHELEA Szadziewski

WIRTHOHELEA Szadziewski, 1988: 158. Type species: Wirthohelea trifida Szadziewski, 1988, by original designation.
trifida Szadziewski, 1988: 159. PA: Baltic Region (Eocene/Oligocene) [A].

Unplaced Species of Ceratopogonidae

crassipalpis Meunier, 1912c: 191 [1912d: 365] (Ceratopogon). AF: Madagascar (Holocene) [K].
escheri Giebel, 1856: 252 (Ceratopogon). PA: Baltic Region (Eocene/Oligocene) [A].
hotchkissae Pierce, 1966: 94 (Johannsenomyia). NE: USA (Miocene) [P].
madagascariensis Meunier, 1912c: 191 [1912d: 365] (Ceratopogon). AF: Madagascar (Holocene) [K].
swinhoei Cockerell, 1919b: 243 (Johannsenomyia). OR: Myanmar (Oligocene) [A].
tenuipes Meunier, 1912d: 366 (Ceratopogon). AF: Madagascar (Holocene) [K].
viscatus Meunier, 1912c: 191 [1912d: 364] (Ceratopogon). AF: Madagascar (Holocene) [K].

Questionably Placed in Ceratopogonidae

pasitheum Heyden, 1870: 238 (Simulium). PA: Germany (Oligocene) [C].
terribile Förster, 1891: 467 (Simulium). PA: Germany (Oligocene) [C].

This page last revised 17 November 2001