PBDB Taxon

Basic info	Taxonomic history	Classification	Included Taxa
Morphology	Ecology and taphonomy	External Literature Search	Age range and collections

Ceratopogoninae

Insecta - Diptera - Ceratopogonidae

Taxonomy

Ceratopogoninae was named by Newman (1834). It is extant. Its type is Ceratopogon.

It was assigned to Chironomidae by Kieffer (1906) and Kieffer (1911); and to Ceratopogonidae by Remm (1976), Szadziewski (1988), Szadziewski (1990), Szadziewski (1996), Borkent and Wirth (1997), Borkent (2013), Choufani et al. (2014), Stebner et al. (2016), Stebner and Szadziewski (2017), Pielowska et al. (2018), Szadziewski and Sontag (2019), Szadziewski (2019), Borkent and Dominiak (2020), Szadziewski et al. (2020), Peñalver et al. (2021) and Szadziewski et al. (2022).

Subtaxa

Ceratopogon escheri, Ceratopogonini (syn. Stilobezziini), Culicoides carri, Culicoides fossilis, Culicoides laurae, Culicoides mioceneus, Culicoidini, Heteromyiini, Johannsenomyia hotchkissae, Johannsenomyiini, Neoculicoides, Neoculicoides jeanneae, Palpomyiini, Sphaeromiini

Synonyms

Neoculicoides was named by Pierce (1966).

It was considered a nomen dubium by Borkent (2013) and Borkent and Dominiak (2020).

It was assigned to Ceratopogonidae by Carpenter (1992).
Culicoides carri was named by Pierce (1966). Its type specimen is LACMIP S9122, a nymph (pupa), and it is a 3D body fossil. Its type locality is Site 2C, Calico Hills, which is in a Hemingfordian lacustrine - large shale in the Barstow Formation of California.

It was considered a nomen dubium by Borkent and Dominiak (2020).
Culicoides fossilis was named by Pierce (1966). Its type specimen is LACMIP S9116, a nymph (pupa), and it is an inclusion in amber. Its type locality is Site 25 (LACMIP 373), Calico Hills, which is in a Hemingfordian lacustrine - large shale in the Barstow Formation of California.

It was considered a nomen dubium by Borkent and Dominiak (2020).
Culicoides laurae was named by Pierce (1966). Its type specimen is LACMIP S9114, a nymph (pupa), and it is a 3D body fossil. Its type locality is Section 19, Calico Hills, which is in a Hemingfordian lacustrine - large shale in the Barstow Formation of California.

It was considered a nomen dubium by Borkent and Dominiak (2020).
Culicoides miocenea was named by Pierce (1966). Its type specimen is LACMIP S9115, a nymph (pupa), and it is a 3D body fossil. Its type locality is Site 17F (LACMIP 365), Calico Hills, which is in a Hemingfordian lacustrine - large shale in the Barstow Formation of California.

It was corrected as Culicoides mioceneus by Borkent and Wirth (1997) and Borkent (2013); it was considered a nomen dubium by Borkent and Dominiak (2020).
Johannsenomyia hotchkissae was named by Pierce (1966). Its type specimen is LACMIP S9128, a nymph (pupa), and it is a 3D body fossil. Its type locality is Site 10 (LACMIP 357), Calico Hills, which is in a Hemingfordian lacustrine - large shale in the Barstow Formation of California.

It was considered a nomen dubium by Borkent (2013) and Borkent and Dominiak (2020).
Neoculicoides jeanneae was named by Pierce (1966). Its type specimen is LACMIP S9123, a nymph (pupa), and it is a 3D body fossil. Its type locality is Site 10 (LACMIP 357), Calico Hills, which is in a Hemingfordian lacustrine - large shale in the Barstow Formation of California.

It was considered a nomen dubium by Borkent (2013) and Borkent and Dominiak (2020).
Ceratopogon escheri was named by Giebel (1856). It is an inclusion in amber. Its type locality is Baltic amber (Leipzig University collection), which is in a Priabonian terrestrial amber in the Russian Federation.

It was considered a nomen dubium by Borkent (2013) and Borkent and Dominiak (2020).

Synonymy list

Year	Name and author
1834	Ceratopogoninae Newman
1856	Ceratopogon escheri Giebel p. 252
1906	Ceratopogoninae Kieffer p. 44
1911	Ceratopogoninae Kieffer p. 112
1966	Culicoides carri Pierce p. 91 fig. 15,16
1966	Culicoides fossilis Pierce p. 91 fig. 11
1966	Culicoides laurae Pierce p. 91 fig. 9
1966	Culicoides miocenea Pierce p. 93 fig. 10
1966	Neoculicoides Pierce p. 93
1966	Johannsenomyia hotchkissae Pierce p. 94 fig. 18
1966	Neoculicoides jeanneae Pierce p. 94 fig. 17
1976	Ceratopogoninae Remm p. 345
1988	Ceratopogoninae Szadziewski p. 25
1990	Ceratopogoninae Szadziewski p. 77
1992	Neoculicoides Carpenter
1996	Ceratopogoninae Szadziewski p. 58
1997	Ceratopogoninae Borkent and Wirth p. 59
1997	Culicoides carri Borkent and Wirth p. 64
1997	Culicoides fossilis Borkent and Wirth p. 68
1997	Culicoides laurae Borkent and Wirth p. 72
1997	Culicoides mioceneus Borkent and Wirth p. 74
2013	Ceratopogoninae Borkent p. 72
2013	Culicoides carri Borkent p. 77
2013	Culicoides fossilis Borkent p. 83
2013	Culicoides laurae Borkent p. 88
2013	Culicoides mioceneus Borkent p. 91
2014	Ceratopogoninae Choufani et al. p. 38
2016	Ceratopogoninae Stebner et al. p. 2
2017	Ceratopogoninae Stebner and Szadziewski p. 5
2018	Ceratopogoninae Pielowska et al. p. 240
2019	Ceratopogoninae Szadziewski p. 526
2019	Ceratopogoninae Szadziewski and Sontag p. 660
2020	Ceratopogoninae Borkent and Dominiak p. 102
2020	Ceratopogoninae Szadziewski et al. p. 286
2021	Ceratopogoninae Peñalver et al. p. 426
2022	Ceratopogoninae Szadziewski et al. p. 2

Is something missing? Join the Paleobiology Database and enter the data

Rank	Name	Author
kingdom	Animalia	()
—	Bilateria
—	Eubilateria	Ax 1987
—	Protostomia
—	Ecdysozoa
—	Panarthropoda
phylum	Arthropoda	Latreille 1829
superclass	Hexapoda	Latreille 1825

Rank	Name	Author
class	Insecta
order	Diptera
infraorder	Culicomorpha
superfamily	Chironomoidea	Macquart 1838
family	Ceratopogonidae
subfamily	Ceratopogoninae
subfamily	Ceratopogoninae

If no rank is listed, the taxon is considered an unranked clade in modern classifications. Ranks may be repeated or presented in the wrong order because authors working on different parts of the classification may disagree about how to rank taxa.

Subfm. Ceratopogoninae Newman 1834

show all | hide all

Tr. Ceratopogonini Newman 1834

G. Alluaudomyia Kieffer 1913

†Alluaudomyia succinea Szadziewski 1988

G. Atyphohelea Borkent 1998

†Atyphohelea cothurnata Meunier 1904

Invalid names: Ceratopogon flagellus Meunier 1904 [synonym]

G. Austrohelea Wirth and Grogan 1988

G. Baeodasymyia Clastrier and Raccurt 1979

†Baeodasymyia dominicana Szadziewski and Grogan 1994

G. Baeohelea Wirth and Blanton 1970

G. †Brachycretacea Szadziewski 1996

†Brachycretacea taimyrica Remm 1976

G. Brachypogon Kieffer 1899

Subg. Brachypogon (Brachypogon) Kieffer 1899

†Brachypogon (Brachypogon) americanus Szadziewski and Grogan 1998

†Brachypogon (Brachypogon) balticus Szadziewski 1988

†Brachypogon (Brachypogon) eocenicus Szadziewski 1988

†Brachypogon (Brachypogon) gedanicus Szadziewski 1988

†Brachypogon (Brachypogon) miocaenicus Szadziewski 1993

Subg. Brachypogon (Isohelea) Kieffer 1917

†Brachypogon (Isohelea) dominicanus Szadziewski and Grogan 1998

†Brachypogon (Isohelea) henningseni Szadziewski 1988

†Brachypogon (Isohelea) polonicus Szadziewski 1988

†Brachypogon (Isohelea) prominuloides Szadziewski and Grogan 1998

†Brachypogon (Isohelea) prominulus Meunier 1904

†Brachypogon balticus Szadziewski 1988

G. Camptopterohelea Wirth and Hubert 1960

†Camptopterohelea odora Stebner et al. 2017

G. Ceratoculicoides Wirth and Ratanaworabhan 1971

†Ceratoculicoides danicus Szadziewski 1988

G. †Ceratopalpomyia Szadziewski 1988

†Ceratopalpomyia eocenica Szadziewski 1988

G. Ceratopogon Meigen 1803

†Ceratopogon bitterfeldi Szadziewski 1993

†Ceratopogon ceranowiczi Szadziewski 1988

†Ceratopogon crypticus Szadziewski 1988

†Ceratopogon eminens Meunier 1904

†Ceratopogon forcipiformis Meunier 1904

Invalid names: Ceratopogon defectus Meunier 1904 [synonym]

†Ceratopogon gedanicus Szadziewski 1988

†Ceratopogon grogani Szadziewski 1988

†Ceratopogon hennigi Szadziewski 1988

†Ceratopogon kotejai Szadziewski 1993

†Ceratopogon margaritae Szadziewski 1988

†Ceratopogon miocaenicus Szadziewski 1993

†Ceratopogon nanalobus Borkent and Grogan 1995

†Ceratopogon paraeminens Borkent and Grogan 1995

†Ceratopogon piotrowskii Szadziewski 1988

†Ceratopogon pisinnus Borkent and Grogan 1995

†Ceratopogon remmicola Szadziewski 1988

†Ceratopogon remmicolus Szadziewski 1988

†Ceratopogon ritzkowskii Szadziewski 1988

†Ceratopogon spiniger Loew 1850

†Ceratopogon subeminens Szadziewski 1993

†Ceratopogon succinicola Szadziewski 1993

†Ceratopogon tertiaricus Szadziewski 1988

Invalid names: Ceratopogon (Ceratopogon) Meigen 1803 [invalid subgroup]

G. †Eohelea Petrunkevitch 1957

†Eohelea fossicola Szadziewski 1993

†Eohelea gedanica Szadziewski 1988

†Eohelea grogani Szadziewski 1988

†Eohelea indica Stebner and Szadziewski 2017

†Eohelea miocaenea Szadziewski 1993

†Eohelea petrunkevitchi Szadziewski 1985

†Eohelea sakhalinica Szadziewski 1990

†Eohelea sinuosa Meunier 1904

Invalid names: Eohelea stridulans Petrunkevitch 1957 [synonym]

G. Fanthamia de Meillon 1939

G. †Fossihelea Szadziewski 1988

†Fossihelea gracilitarsis Meunier 1904

†Fossihelea miocaenica Szadziewski 1993

G. †Gedanohelea Szadziewski 1988

†Gedanohelea fushunensis Stebner et al. 2016

†Gedanohelea gerdesorum Stebner and Szadziewski 2017

†Gedanohelea liaoningensis Stebner et al. 2016

†Gedanohelea loewi Szadziewski 1988

†Gedanohelea succinea Szadziewski 1988

†Gedanohelea wirthi Szadziewski 1988

G. †Mantohelea Szadziewski 1988

†Mantohelea gedanica Szadziewski 1988

†Mantohelea laca Meunier 1904

†Mantohelea sinica Stebner et al. 2016

G. Meunierohelea Szadziewski 1988

†Meunierohelea anglesensis Peñalver et al. 2021

†Meunierohelea borkenti Stebner and Szadziewski 2017

†Meunierohelea cambayana Stebner and Szadziewski 2017

†Meunierohelea fudalai Szadziewski et al. 2020

†Meunierohelea gedanicola Szadziewski 1988

†Meunierohelea miocaenica Szadziewski 1993

†Meunierohelea nielseni Szadziewski 1988

†Meunierohelea orientalis Stebner and Szadziewski 2017

†Meunierohelea wirthi Szadziewski 1988

Invalid names: Chimaerohelea Debenham 1988 [synonym]

G. †Monogedania Szadziewski et al. 2022

†Monogedania clunipes Loew 1850

G. Monohelea Kieffer 1917

G. Nannohelea Grogan and Wirth 1980

†Nannohelea eocenica Szadziewski 1988

†Nannohelea grogani Szadziewski 1988

G. †Nelohelea Szadziewski and Sontag 2019

†Nelohelea neli Szadziewski and Sontag 2019

G. †Palaeobrachypogon Borkent 1995

†Palaeobrachypogon aquilonius Boesel 1937

†Palaeobrachypogon grandiforceps Borkent 2000

†Palaeobrachypogon macronyx Remm 1976

†Palaeobrachypogon remmi Borkent 1995

†Palaeobrachypogon taimyricus Szadziewski 1996

†Palaeobrachypogon vetus Borkent 1995

G. †Peronehelea Borkent 1995

†Peronehelea chrimikalydia Borkent 1995

†Peronehelea frigida Remm 1976

†Peronehelea sibirica Szadziewski 1996

G. Schizohelea Kieffer 1917

†Schizohelea baltica Szadziewski 1988

G. Serromyia Meigen 1818

†Serromyia alphea von Heyden 1870

Invalid names: Serromyia austera Statz 1944 [synonym], Serromyia colorata Statz 1944 [synonym], Serromyia spinofemorata Statz 1944 [synonym]

†Serromyia anomalicornis Loew 1850

†Serromyia errata Szadziewski 2018

†Serromyia polonica Szadziewski 1988

†Serromyia ryszardi Borkent 1990

†Serromyia sinuosa Borkent 1990

†Serromyia spinigera Loew 1850

Invalid names: Ceratopogon elongatus Meunier 1904 [synonym]

†Serromyia succinea Szadziewski 1988

G. Stilobezzia Kieffer 1911

Subg. Stilobezzia (Acanthohelea) Kieffer 1917

†Stilobezzia (Acanthohelea) dominicana Szadziewski and Grogan 1998

†Stilobezzia (Acanthohelea) falcata Meunier 1904

Invalid names: Ceratopogon spinosus Meunier 1904 [synonym]

†Stilobezzia (Acanthohelea) kutscheri Szadziewski 1993

†Stilobezzia (Acanthohelea) saxonica Szadziewski 1993

†Stilobezzia (Acanthohelea) veterana Meunier 1920

Invalid names: Stilobezzia goetghebueri Statz 1944 [synonym]

†Stilobezzia (Acanthohelea) wirthicola Szadziewski and Grogan 1998

Invalid names: Stilobezzia (Acanthohelea) succinea Szadziewski 1993 [replaced]

Subg. Stilobezzia (Stilobezzia) Kieffer 1911

†Stilobezzia (Stilobezzia) antilleana Szadziewski and Grogan 1998

†Stilobezzia kurthi Borkent 2000

†Stilobezzia pikei Borkent 2012

†Stilobezzia roggeroi Choufani and Nel 2013

G. †Wirthohelea Szadziewski 1988

†Wirthohelea trifida Szadziewski 1988

Invalid names: Stilobezziini Wirth 1952 [synonym]

Tr. Culicoidini Kieffer 1911

G. Culicoides Latreille 1809

Subg. Culicoides (Groganomyia) Szadziewski and Dominiak 2019

†Culicoides (Groganomyia) bojarskii Szadziewski and Dominiak 2019

†Culicoides (Groganomyia) burmiticus Szadziewski and Dominiak 2019

†Culicoides (Groganomyia) ellenbergeri Szadziewski and Dominiak 2019

†Culicoides (Groganomyia) myanmaricus Szadziewski and Dominiak 2019

Subg. Culicoides (Oecacta) Poey 1853

†Culicoides (Oecacta) dasyheleiformis Szadziewski 1988

†Culicoides (Oecacta) succivarius Szadziewski 1988

†Culicoides abbreviatipennis Statz 1944

†Culicoides agamus Borkent 1995

†Culicoides ambericus Szadziewski and Grogan 1998

†Culicoides annosus Borkent 1995

†Culicoides antilleanus Szadziewski and Grogan 1998

†Culicoides atratus Statz 1944

†Culicoides austerus Statz 1944

†Culicoides balticus Szadziewski 1988

†Culicoides bicolor Statz 1944

†Culicoides bifidus Borkent 1996

†Culicoides brisaci Choufani and Nel 2013

†Culicoides brodzinskyi Szadziewski and Grogan 1998

†Culicoides bullus Borkent 1995

†Culicoides canadensis Boesel 1937

†Culicoides casei Grogan and Szadziewski 1988 [biting midge]

†Culicoides ceranowiczi Szadziewski 1988

†Culicoides dasyheleiformis Szadziewski 1988

†Culicoides doyeni Choufani et al. 2014

†Culicoides elongatulus Statz 1944

†Culicoides eoselficus Szadziewski 1988

†Culicoides filipalpis Remm 1976

†Culicoides gedanensis Szadziewski 1988

†Culicoides gracilior Statz 1944

†Culicoides grandibocus Borkent 1996

†Culicoides hispanicola Szadziewski and Grogan 1998

†Culicoides jucundus Statz 1944

†Culicoides kaluginae Remm 1976

†Culicoides liliputanus Statz 1944

†Culicoides mammalicola Szadziewski and Grogan 1998

†Culicoides megacanthus Palmer 1957

†Culicoides obesus Statz 1944

†Culicoides obscuratus Statz 1944

†Culicoides obuncus Borkent 1995

†Culicoides paleopestis Peñalver et al. 2021

†Culicoides prussicus Szadziewski 1988

†Culicoides speciosus Meunier 1904

†Culicoides sphenostylus Remm 1976

†Culicoides subgedanensis Szadziewski 1993

†Culicoides succineus Remm 1976

†Culicoides succivarius Szadziewski 1988

†Culicoides tenuipennis Statz 1944

†Culicoides truncatus Borkent 2000

†Culicoides tyrrelli Boesel 1937

†Culicoides ventralis Borkent and Wirth 1997

Invalid names: Culicoides abdominalis Statz 1944 [replaced]

†Culicoides yoosti Borkent 2000

Invalid names: Devalquia Choufani and Nel 2013 [synonym]

G. Washingtonhelea Wirth and Grogan 1988

Tr. Heteromyiini Wirth 1962

G. Heteromyia Say 1825

†Heteromyia dominicana Szadziewski and Grogan 1997

G. Metahelea Edwards 1929

†Metahelea serafini Szadziewski 1998

G. Neurohelea Kieffer 1925

G. Physohelea Grogan and Wirth 1979

†Physohelea obtusa Meunier 1904

Invalid names: Ceratopogon cothurnatulus Meunier 1904 [synonym]

Tr. Johannsenomyiini Crampton 1925

G. Johannsenomyia Malloch 1915

G. Macropeza Meigen 1818

G. Mallochohelea Wirth 1962

†Mallochohelea martae Szadziewski 2005

Tr. Palpomyiini Enderlein 1936

G. Bezzia Kieffer 1899

†Bezzia eocenica Szadziewski 1988

†Bezzia longipennis Statz 1944

G. Palpomyia Meigen 1818

†Palpomyia edwardsi Cockerell 1921

†Palpomyia erikae Szadziewski 1993

†Palpomyia freyi Pierce 1966

†Palpomyia jantari Szadziewski 1988

†Palpomyia multispinosa Pierce 1966

†Palpomyia riedeli Szadziewski 1988

†Palpomyia ryshkoffi Pierce 1966

†Palpomyia shilo Pierce 1966

†Palpomyia succinea Szadziewski 1988

†Palpomyia unca Hong 1981

†Palpomyia wirthorum Szadziewski and Grogan 1997

Invalid names: Eopalpomyitis Hong et al. 2000 [synonym], Miopalpomyia Pierce 1966 [synonym], Neopalpomyia Pierce 1966 [synonym], Parapalpomyia Pierce 1966 [synonym]

G. Phaenobezzia Haeselbarth 1965

†Phaenobezzia wirthi Szadziewski and Grogan 1997

Invalid names: Ceratopogon escheri Giebel 1856 [nomen dubium], Culicoides carri Pierce 1966 [nomen dubium], Culicoides fossilis Pierce 1966 [nomen dubium], Culicoides laurae Pierce 1966 [nomen dubium], Culicoides mioceneus Pierce 1966 [nomen dubium], Johannsenomyia hotchkissae Pierce 1966 [nomen dubium], Neoculicoides Pierce 1966 [nomen dubium], Neoculicoides jeanneae Pierce 1966 [nomen dubium], Sphaeromiini Newman 1834 [empty]

Diagnosis

No diagnoses are available

Measurements

No measurements are available

Composition:

chitinc

Environment:

terrestrialc

Locomotion:

actively mobilep

Created:

2017-04-17 09:56:18

Modified:

2017-04-17 09:56:18

Source: c = class, p = phylum

References: Kiessling 2004, Bush and Bambach 2015

Age range

Maximum range based only on fossils: base of the Turonian to the top of the Hemingfordian or 93.90000 to 15.97000 Ma
Minimum age of oldest fossil (stem group age): 93.5 Ma

Collections (84 total)

Time interval	Ma	Country or state	Original ID and collection number
Early/Lower Cenomanian	99.6 - 93.5	Myanmar	Culicoides bojarskii, Culicoides myanmaricus (161726) Culicoides burmiticus, Nelohelea neli (187929) Culicoides ellenbergeri (165560)
Late/Upper Cenomanian	99.6 - 93.5	USA (New Jersey)	Culicoides casei (110959)
Turonian	93.9 - 89.8	USA (New Jersey)	Culicoides casei, Culicoides filipalpis, Palaeobrachypogon remmi, Culicoides grandibocus, Culicoides truncatus, Culicoides yoosti, Palaeobrachypogon grandiforceps, Stilobezzia kurthi (110895) Palaeobrachypogon sp., Culicoides bifidus (123016)
Turonian	93.9 - 89.8	France	Culicoides doyeni (164386)
Early/Lower Santonian	85.8 - 83.5	France	Devalquia brisaci, Metahelea roggeroi (152462)
Late/Upper Santonian	85.8 - 83.5	Russian Federation	Culicoides sp., Culicoides filipalpis, Ceratopogon macronyx, Ceratopogon frigidus, Culicoides kaluginae, Culicoides sphenostylus, Culicoides succineus, Washingtonhelea taimyrica, Baeohelea taimyrica, Peronehelea sibirica (35056)
Late/Upper Campanian	83.5 - 70.6	Canada (Manitoba)	Culicoides filipalpis, Culicoides canadensis, Culicoides obuncus, Peronehelea chrimikalydia (122353) Culicoides filipalpis, Culicoides tyrrelli, Culicoides canadensis, Culicoides agamus, Culicoides obuncus, Culicoides annosus, Palaeobrachypogon remmi, Palaeobrachypogon vetus, Peronehelea chrimikalydia (122354) Culicoides filipalpis, Dasyhelea tyrrelli, Atrichopogon canadensis, Culicoides obuncus, Ceratopogon aquilonius, Palaeobrachypogon remmi, Palaeobrachypogon vetus (122356)
Middle Campanian	83.5 - 70.6	Canada (Alberta)	Culicoides filipalpis (122385) Culicoides filipalpis, Culicoides bullus (123143) Culicoides filipalpis, Culicoides tyrrelli, Culicoides annosus, Peronehelea chrimikalydia (104932) Culicoides filipalpis, Culicoides tyrrelli, Culicoides bullus, Palaeobrachypogon aquilonius, Peronehelea chrimikalydia (122388) Culicoides filipalpis, Culicoides tyrrelli, Culicoides canadensis, Culicoides obuncus, Palaeobrachypogon remmi, Palaeobrachypogon vetus (122384) Culicoides filipalpis, Culicoides tyrrelli, Culicoides canadensis, Palaeobrachypogon vetus, Peronehelea chrimikalydia (122383) Culicoides sp., Culicoides filipalpis, Palaeobrachypogon sp., Palaeobrachypogon vetus, Peronehelea sp., Peronehelea chrimikalydia, Stilobezzia pikei (122389) Culicoides sp., Peronehelea chrimikalydia (34896) Peronehelea chrimikalydia (122358)
Ypresian	56.0 - 47.8	China (Liaoning)	Mantohelea sinica, Gedanohelea fushunensis, Gedanohelea liaoningensis (125934) Palpomyia unca (124530)
Ypresian	56.0 - 47.8	India (Gujarat)	Mantohelea sp., Stilobezzia sp., Brachypogon sp., Serromyia sp., Camptopterohelea odora, Eohelea indica, Gedanohelea gerdesorum, Meunierohelea cambayana, Meunierohelea borkenti, Meunierohelea orientalis (124463) Meunierohelea sp., Gedanohelea gerdesorum (183742)
Lutetian	47.8 - 41.3	USA (Montana)	Alluaudomyia sp. (224195)
Lutetian	47.8 - 41.3	Russian Federation	Stilobezzia sp., Ceratopogoninae indet., Eohelea sakhalinica (107340)
Bartonian	41.3 - 38.0	Australia (Victoria)	Monohelea sp., Meunierohelea anglesensis, Culicoides paleopestis (224365)
Priabonian	38.0 - 33.9	Poland	Ceratopogon sp., Culicoides sp., Eohelea sinuosa, Meunierohelea sp., Serromyia sp., Culicoides speciosus (154301) Ceratopogon sp., Culicoides sp., Mantohelea lacus, Mantohelea gedanica, Stilobezzia sp., Stilobezzia sp. C, Stilobezzia sp. D, Stilobezzia sp. E, Ceratopogonini indet., Ceratopogon eminens, Ceratopogon forcipiformis, Ceratopogon grogani, Ceratopogon hennigi, Ceratopogon margaritae, Ceratopogon tertiaricus, Eohelea gedanica, Eohelea grogani, Eohelea petrunkevitchi, Fossihelea gracilitarsis, Gedanohelea loewi, Meunierohelea sp., Meunierohelea sp. B, Meunierohelea sp. C, Meunierohelea nielseni, Monohelea clunipes, Nannohelea sp., Serromyia sp., Serromyia sp. A, Serromyia sp. B, Serromyia spinigera, Serromyia succinea, Stilobezzia falcata, Physohelea obtusa, Neurohelea cothurnata, Bezzia eocenica, Eohelea stridulans, Culicoides sp. A, Culicoides sp. B, Culicoides sp. C, Brachypogon sp., Brachypogon sp. A, Culicoides ceranowiczi, Culicoides eoselficus, Culicoides gedanensis, Alluaudomyia succinea, Brachypogon balticus, Brachypogon eocenicus, Brachypogon gedanicus, Brachypogon henningseni, Brachypogon prominulus, Ceratopogon ceranowiczi, Ceratopogon crypticus, Ceratopogon gedanicus, Ceratopogon ritzkowskii, Gedanohelea succinea, Meunierohelea gedanicola, Monohelea baltica, Serromyia anomalicornis, Serromyia polonica, Serromyia ryszardi, Serromyia sinuosa, Wirthohelea trifida, Palpomyia jantari, Culicoides speciosus, Culicoides dasyheleiformis, Culicoides succivarius, Culicoides balticus, Ceratopogon remmicolus, Eohelea sp. A, Eohelea sp. B (120813) Ceratopogon sp., Fossihelea gracilitarsis, Meunierohelea sp. C, Ceratopogon clunipes, Serromyia sp., Neurohelea cothurnata, Ceratopogon spiniger, Culicoides prussicus, Brachypogon balticus, Brachypogon prominulus, Ceratopogon anomalicornis, Culicoides dasyheleiformis (109625) Ceratopogon sp., Fossihelea gracilitarsis, Monohelea clunipes, Brachypogon sp., Brachypogon balticus, Brachypogon polonicus, Brachypogon prominulus, Palpomyia riedeli (153481) Mantohelea gedanica, Metahelea serafini, Mallochohelea martae, Monogedania clunipes, Schizohelea baltica (110737)
Priabonian	38.0 - 33.9	Germany (Halle)	Ceratopogon sp., Culicoides sp., Culicoides speciosus, Ceratopogon bitterfeldi, Ceratopogon forcipiformis, Ceratopogon hennigi, Eohelea fossicola, Eohelea miocaenea, Eohelea sinuosa, Fossihelea sp., Fossihelea miocaenica, Monohelea clunipes, Serromyia sp., Serromyia spinigera, Stilobezzia succinea, Culicoides ceranowiczi, Culicoides subgedanensis, Brachypogon miocaenicus, Ceratopogon miocaenicus, Ceratopogon subeminens, Ceratopogon succinicolus, Chimaerohelea miocaenica, Palpomyia erikae, Serromyia alphea (129227) Ceratopogon sp., Culicoides sp., Mantohelea gedanica, Culicoides speciosus, Ceratopogon forcipiformis, Ceratopogon hennigi, Eohelea sinuosa, Fossihelea sp., Fossihelea sp. B, Monohelea clunipes, Nannohelea sp., Serromyia sp., Brachypogon sp., Brachypogon sp., Brachypogon prominulus, Ceratopogon kotejai, Stilobezzia falcata, Stilobezzia kutscheri, Stilobezzia saxonica, Chimaerohelea nielseni, Serromyia alphea (129228)
Priabonian	38.0 - 33.9	Ukraine	Ceratopogon sp. (122999) Ceratopogon sp., Stilobezzia sp., Brachypogon sp., Eohelea sinuosa, Meunierohelea sp., Monohelea clunipes (122996) Culicoides sp., Stilobezzia sp., Eohelea sinuosa, Meunierohelea sp. (122995) Palpomyia sp., Ceratopogon sp., Culicoides sp., Mantohelea laca, Stilobezzia sp., Brachypogon sp., Culicoides speciosus, Brachypogon balticus, Brachypogon prominulus, Ceratopogon bitterfeldi, Ceratopogon forcipiformis, Ceratopogon grogani, Ceratopogon hennigi, Ceratopogon margaritae, Ceratopogon tertiaricus, Eohelea fossicola, Eohelea gedanica, Eohelea miocaenea, Eohelea petrunkevitchi, Eohelea sinuosa, Fossihelea sp., Fossihelea gracilitarsis, Fossihelea miocaenica, Gedanohelea loewi, Meunierohelea sp., Meunierohelea nielseni, Monohelea sp., Monohelea clunipes, Nannohelea sp., Nannohelea grogani, Serromyia sp., Serromyia spinigera, Serromyia succinea, Stilobezzia falcata, Physohelea obtusa, Neurohelea sp., Neurohelea cothurnata, Bezzia eocenica (122997) Palpomyia sp., Ceratopogon sp., Culicoides sp., Stilobezzia sp., Brachypogon sp., Culicoides speciosus, Brachypogon balticus, Brachypogon prominulus, Ceratopogon eminens, Ceratopogon forcipiformis, Eohelea gedanica, Eohelea grogani, Eohelea sinuosa, Meunierohelea sp., Monohelea clunipes, Stilobezzia falcata, Bezzia eocenica (113621)
Priabonian	38.0 - 33.9	Denmark	Ceratopogon sp., Culicoides sp., Stilobezzia sp., Stilobezzia sp. A, Ceratopogoninae indet., Ceratopogonini indet., Culicoides speciosus, Ceratopogon eminens, Ceratopogon forcipiformis, Ceratopogon hennigi, Ceratopogon tertiaricus, Eohelea gedanica, Eohelea petrunkevitchi, Eohelea sinuosa, Fossihelea gracilitarsis, Meunierohelea sp., Meunierohelea sp. A, Meunierohelea nielseni, Monohelea clunipes, Nannohelea sp., Nannohelea grogani, Serromyia sp., Stilobezzia falcata, Neurohelea cothurnata, Bezzia eocenica, Eohelea stridulans, Culicoides sp. B, Brachypogon sp. A, Culicoides eoselficus, Alluaudomyia sp. A, Alluaudomyia succinea, Brachypogon balticus, Brachypogon eocenicus, Brachypogon gedanicus, Brachypogon henningseni, Brachypogon prominulus, Ceratoculicoides danicus, Ceratopalpomyia eocenica, Ceratopogon crypticus, Ceratopogon piotrowskii, Ceratopogon ritzkowskii, Gedanohelea wirthi, Meunierohelea wirthi, Monohelea baltica, Nannohelea eocenica, Palpomyia succinea, Culicoides balticus, Stilobezziini indet., Ceratopogon sp. B, Ceratopogon sp. A, Ceratopogon sp. C (123762)
Priabonian	38.0 - 33.9	United Kingdom (England)	Stilobezzia sp. (21976) Stilobezzia sp., Palpomyia edwardsi (123959)
Priabonian	38.0 - 33.9	Russian Federation (Kaliningrad)	Ceratopogon escheri (147344) Ceratopogon lacus, Ceratopogon speciosus, Ceratopogon prominulus, Ceratopogon eminens, Ceratopogon forcipiformis, Fossihelea sp. A, Ceratopogon gracilitarsis, Ceratopogon clunipes, Ceratopogon falcatus, Ceratopogon obtusus, Ceratopogon cothurnatus, Ceratopogon defectus, Ceratopogon flagellus, Ceratopogon elongatus, Ceratopogon cothurnatulus, Ceratopogon spiniger, Ceratopogon sinuosus, Ceratopogon spinosus (123927) Ceratopogon sp., Culicoides sp., Ceratopogoninae indet., Ceratopogonini indet., Ceratopogon tertiaricus, Monohelea sp. A, Monohelea clunipes, Brachypogon balticus, Brachypogon prominulus (123993) Ceratopogon sp., Culicoides sp., Ceratopogoninae indet., Eohelea sinuosa, Monohelea clunipes, Brachypogon sp., Culicoides ceranowiczi, Alluaudomyia succinea, Culicoides dasyheleiformis (123221) Ceratopogon sp., Culicoides sp., Stilobezzia sp. A, Stilobezzia sp. B, Ceratopogoninae indet., Ceratopogonini indet., Ceratopogon forcipiformis, Eohelea sinuosa, Fossihelea gracilitarsis, Monohelea clunipes, Nannohelea sp., Serromyia sp., Brachypogon prominulus (123962)
Priabonian	38.0 - 33.9	Russian Federation	Ceratopogon lacus (123419)
MP 30	28.4 - 23.03	Germany (Nordrhein-Westfalen)	Tetragoneura veterana (138696)
MP 30	28.4 - 23.03	Germany	Ceratopogon alpheus (123945)
Chattian	28.1 - 23.03	Germany (North Rhine-Westphalia)	Ceratopogon pupa 13, Ceratopogon pupa 12, Ceratopogon pupa 11, Ceratopogon pupa 10, Ceratopogon pupa 9, Ceratopogon pupa 8, Ceratopogon pupa 7, Ceratopogon pupa 6, Ceratopogon pupa 5, Ceratopogon pupa 4, Ceratopogon pupa 3, Ceratopogon pupa 2, Ceratopogon pupa 1, Ceratopogon sp., Serromyia sp., Culicoides abbreviatipennis, Culicoides atratus, Culicoides austerus, Culicoides bicolor, Culicoides elongatulus, Culicoides gracilior, Culicoides jucundus, Culicoides liliputanus, Culicoides obesus, Culicoides tenuipennis, Culicoides abdominalis, Serromyia austera, Serromyia colorata, Serromyia spinofemorata, Stilobezzia goetghebueri, Bezzia longipennis, Culicoides obscuratus, Stilobezzia veterana (122010)
Chattian	28.1 - 23.03	France (Provence-Alpes-Côte d'Azur)	Ceratopogon sp. (122488)
Chattian	28.1 - 23.03	New Zealand (Otago)	Austrohelea sp. (191651)
Early/Lower Miocene	23.03 - 15.97	Mexico (Chiapas)	Brachypogon sp. (124474) Nannohelea sp. (140212 140217)
Early/Lower Miocene	23.03 - 15.97	Mexico	Culicoides sp., Brachypogon sp. (21977)
Burdigalian - Langhian	20.44 - 13.82	Dominican Republic	Brachypogon dominicanus (125970) Culicoides brodzinskyi, Culicoides ambericus, Brachypogon dominicanus, Stilobezzia dominicana, Stilobezzia antilleana, Baeodasymyia dominicana, Phaenobezzia wirthi, Culicoides sp., Brachypogon sp. (122982) Heteromyia dominicana (122474) Meunierohelea fudalai (124427) Nannohelea sp. A, Culicoides antilleanus, Culicoides ambericus, Brachypogon dominicanus, Brachypogon prominuloides, Stilobezzia dominicana, Stilobezzia antilleana, Culicoides mammalicolus, Baeodasymyia dominicana, Palpomyia wirthorum, Culicoides sp., Brachypogon sp., Stilobezzia sp. A (122473) Nannohelea sp., Culicoides hispanicolus, Brachypogon americanus, Culicoides sp., Brachypogon sp., Brachypogon sp. (123857) Stilobezzia dominicana, Brachypogon sp. (140835)
Hemingfordian	20.43 - 15.97	USA (California)	Culicoides carri (152228) Culicoides fossilis (144522) Culicoides laurae, Culicoides miocenea, Neopalpomyia freyi, Parapalpomyia ryshkoffi, Johannsenomyia hotchkissae (152229) Culicoides megacanthus (123022 123030 123034) Culicoides megacanthus, Culicoides miocenea, Neopalpomyia freyi, Parapalpomyia ryshkoffi, Johannsenomyia hotchkissae (152222) Culicoides miocenea (152207 152230) Culicoides miocenea, Neopalpomyia freyi, Parapalpomyia ryshkoffi, Johannsenomyia hotchkissae (152216 152217) Johannsenomyia hotchkissae, Neoculicoides jeanneae (144521) Miopalpomyia sp., Culicoides megacanthus, Johannsenomyia hotchkissae (106431) Neopalpomyia freyi (152212) Neopalpomyia freyi, Parapalpomyia ryshkoffi (152219) Neopalpomyia freyi, Parapalpomyia ryshkoffi, Johannsenomyia hotchkissae (152220) Neopalpomyia multispinosa (152232) Parapalpomyia ryshkoffi, Johannsenomyia hotchkissae (152221) Parapalpomyia ryshkoffi, Miopalpomyia shilo (152231)