Ptychopteridae

Superregnum: Eukaryota
Cladus: Unikonta
Cladus: Opisthokonta
Cladus: Holozoa
Regnum: Animalia
Subregnum: Eumetazoa
Cladus: Bilateria
Cladus: Nephrozoa
Cladus: Protostomia
Cladus: Ecdysozoa
Cladus: Panarthropoda
Phylum: Arthropoda
Subphylum: Hexapoda
Classis: Insecta
Cladus: Dicondylia
Subclassis: Pterygota
Cladus: Metapterygota
Infraclassis: Neoptera
Cladus: Eumetabola
Cladus: Endopterygota
Superordo: Panorpida
Cladus: Antliophora
Ordo: Diptera
Subordo: Nematocera
Infraordo: Ptychopteromorpha
Superfamilia: Ptychopteroidea

Familia: Ptychopteridae
Subfamiliae (5): Bittacomorphinae – Ptychopterinae – †Eoptychopterinae – †Eoptychopterininae – †Proptychopterininae
Overview of genera (3 + 12†)

†Architendipes – Bittacomorpha – Bittacomorphella – †Bolboia – †Brodilka – †Crenoptychoptera – †Doptychoptera – †Eoptychoptera – †Eoptychopterina – †Leptychoptera – †Nedoptychoptera – †Probittacomorpha – †Proptychopterina – Ptychoptera – †Zhiganka
[Source: Lukashevich (2008)]
Name

Ptychopteridae Osten Sacken, 1862
Synonyms

†Eoptychopteridae Handlirsch, 1906
†Eolimnobiidae Rohdendorf, 1962

References

Krzemiński, W., Kania, I. & Nel, A. 2012. Probittacomorpha brisaci n. sp. (Ptychopteridae, Bittacomorphinae) from the Late Miocene of Montagne d’Andance, Saint Bauzile, Ardèche (France). Zootaxa 3521: 80–88. Preview Reference page.
Nakamura, T.; Saigusa, T. 2009: Taxonomic study of the family Ptychopteridae of Japan (Diptera). Zoosymposia, 3: 273–303. Preview
Kang, Z., Yao, G. & Yang, D. 2013. Five new species of Ptychoptera Meigen with a key to species from China (Diptera: Ptychopteridae). Zootaxa 3682(4): 541–555. DOI: 10.11646/zootaxa.3682.4.5 Reference page.
Paramonov, N.M. 2013. Ptychopteridae, a family of flies (Diptera) new to the Philippine Islands with the description of a new species. Zootaxa 3682(4): 584–588. DOI: 10.11646/zootaxa.3682.4.10 Reference page.
Lukashevich, E.D. 2008. [Ptychopteridae (Insecta: Diptera): History of its study and limits of the family]. Paleontologicheskii Zhurnal 2008(1): 68–77. (in Russian). PDF. [English translation: Paleontological Journal 42(1): 66–74. DOI: 10.1007/s11492-008-1011-1 Paywall. PDF.] Reference page.
Lukashevich, E.D. 2012. [Phylogeny of Ptychopteroidea (Insecta: Diptera)]. Paleontologicheskii Zhurnal 2012(5): 32–40. (in Russian) [English translation: Paleontological Journal 46(5): 476–484. DOI: 10.1134/s003103011205005x Paywall.]

Vernacular names
English: Phantom Crane Flies
suomi: Kummitussääsket
日本語: コシボソガガンボ科

The Ptychopteridae, phantom crane flies, are a small family (three extant genera) of nematocerous Diptera. Superficially similar in appearance to other "tipuloid" families, they lack the ocelli of the Trichoceridae, the five-branched radial vein of the Tanyderidae, and the two anal veins that reach the wing margins of the Tipulidae. They are usually allied with the Tanyderidae based on similarities of the mesonotal suture, this group being called the Ptychopteromorpha.

Life history
Egg

Ptychoptera albimana (Palearctic) has a mean of 554 eggs laid. The shape is slightly arcuated, "curiously ornamented", and roughly 0.8 mm × 0.2 mm (0.0315 in × 0.0079 in). Duration is reported at 7 days.
Larvae

The larvae are eucephalous and distinctive for the long, caudal respiratory siphon they possess. At hatching, they measure just under 4 mm (0.16 in) in P. albimana, quickly growing to nearly 80 mm (3.1 in). They occur in moist habitats (described as "wet swales and meadows" for Ptychoptera; along lentic shorelines and alder swamps for Bittacomorpha) where they feed as collector-gatherers on decaying organic matter.
Pupae

The pupae possess a single, greatly elongated spiracular horn protruding from their thoraces. In Ptychoptera and Bittacomorpha, the right horn is elongated; in Bittacomorphella, the left. Reported times spent in this stage vary from 5 to 12 days.
Ptychoptera quadrifasciata
Adult

The adults are found most often from late spring through to autumn in shaded, moist environs. Presumably, adults feed little, if at all. Two generations occur per year.

The common species of Eastern North America (Bittacomorpha clavipes) is known for the odd habit of spreading out its legs while flying, using expanded, trachea-rich tarsi to waft along on air currents.

Why they are called “phantom” crane flies: Their legs are thin and black with white sheaths near the tips, and when they fly under a shady tree, everything disappears except the white spots, appearing and disappearing like a “phantom”.
Subfamilies

The general appearance of the two forms is strikingly different. The species of the Bittacomorphinae are similar in size and shape to the Tipulidae, but exhibit a striking black and white coloration — hence the common name "phantom crane flies". The two genera differ as adults in their size and the extent of white coloration on the legs. The larvae of Bittacomorphella possess unique protuberances not seen in the other two genera. Ptychoptera species resemble large mycetophilids, being generally a shiny black and often with patterned wings.

Ptychopterinae – 16 antennomeres; M1 cell present

Ptychoptera Meigen, 1803

Bittacomorphinae – 20 antennomeres; M1 & M2 veins fused, thus without M1 cell

Bittacomorpha Westwood, 1835
Bittacomorphella Alexander, 1916

Bittacomorphinae
Bittacomorpha Westwood, 1835
Bittacomorpha clavipes Fabricius, 1781 - Eastern NA to the edge of the Rocky Mountains (though not as abundant in the Plains states)
Bittacomorpha occidentalis Aldrich, 1895 - Pacific Northwest
Bittacomorphella Alexander, 1916
Bittacomorphella jonesi Johnson, 1905 - New England down to North Carolina, west to Minnesota and Michigan
Bittacomorphella esakii Tokunaga, 1938 - Japan
Bittacomorphella fenderiana Alexander, 1947 - Queen Charlotte Island down to Northern Oregon.
Bittacomorphella nipponensis Alexander, 1924 -
Bittacomorphella pacifica Alexander, 1958 - Northern California up through Oregon (possibly Washington?)
Bittacomorphella sackenii Röder, 1890 - Sierra Nevadas
Bittacomorphella thaiensis Alexander, 1953 -
Ptychopterinae
Ptychoptera Meigen 1803
Ptychoptera byersi Alexander, 1966 - California
Ptychoptera lenis Osten Sacken, 1877
P. l. lenis - Pacific Northwest
P. l. coloradensis - Colorado/Utah
Ptychoptera metallica Walker, 1848 - Central Canada, Minnesota & Michigan
Ptychoptera minor Alexander, 1920 - California/Idaho
Ptychoptera monoensis Alexander, 1947 - Northern California
Ptychoptera osceola Alexander, 1959 - Florida
Ptychoptera pendula Alexander, 1937 - British Columbia to Utah & Colorado
Ptychoptera quadrifasciata Say, 1824 - Eastern North America (Syn P. rufocinctus)
Ptychoptera sculleni Alexander, 1943 - Pacific Northwest
Ptychoptera townesi Alexander, 1943 - Washington & Oregon
Ptychoptera uta Alexander, 1947 - Utah

Fossil Subfamilies and genera

Subfamily Bittacomorphinae Alexander, 1919
†Probittacomorpha Freiwald & Willmann, 1992
†Probittacomorpha brisaci Krzemiński et al., 2012 - Montagne d'Andance (Turolian), France
†Probittacomorpha christenseni Freiwald & Willmann, 1992 - Fur Formation (Ypresian), Denmark;
†Zhiganka Lukashevich, 1995
†Zhiganka comitans Lukashevich, 1995 - Batylykh Formation (Neocomian), Russia
†Zhiganka longialata Lukashevich, 2019 - Khasurty locality (Aptian), Russia
†Zhiganka woolgari Lukashevich et al., 2001 - Weald Clay (Hauterivian), England
Subfamily. †Eoptychopterinae Handlirsch 1906
†Architendipes Rohdendorf 1962
†Architendipes tshernovskiji Rohdendorf, 1962 - Dzhil Formation (Hettangian), Kyrgyzstan
†Doptychoptera Lukashevich, 1998
†Doptychoptera baisicaLukashevich, 1998 - Zaza Formation (Aptian), Russia
†Eolimnobia Handlirsch, 1906
†Eolimnobia geinitzi Handlirsch, 1906 - Ciechocinek Formation (Toarcian), Germany
†Eoptychoptera Handlirsch 1906
†Eoptychoptera aequidistans Lukashevich, 1998 - Dzhil Formation (Hettangian), Kyrgyzstan
†Eoptychoptera altaica Kalugina, 1988 - Ortsog Formation (Bajocian), Mongolia
†Eoptychoptera ansorgei Ren & Krzemiński, 2002 - Daohugou Formation (Callovian), China
†Eoptychoptera asiatica Lukashevich, 1993 - Karabastau Formation (Callovian/Oxfordian), Kazakhstan
†Eoptychoptera aucta Lukashevich, 1993 - Karabastau Formation (Callovian/Oxfordian), Kazakhstan
†Eoptychoptera braziliana Krzemiński et al., 2015 - Crato Formation (Aptian), Brazil
†Eoptychoptera britannica Lukashevich et al., 2001 - Weald Clay (Hauterivian), England
†Eoptychoptera cantabrica Lukashevich & Arillo, 2016 - Spanish amber (Albian), Spain
†Eoptychoptera cretacea Kalugina, 1989 - Zaza Formation (Aptian), Russia
†Eoptychoptera elevata Lukashevich, 2000 - Khaya Formation (Tithonian), Russia
†Eoptychoptera eximia Bode, 1953 - Ciechocinek Formation (Toarcian), Germany
†Eoptychoptera jurassica Ren & Krzemiński, 2002 - Daohugou Formation (Callovian), China
†Eoptychoptera longifurcata Lukashevich et al., 2001 - Lulworth Formation (Berriasian), England
†Eoptychoptera magna Lukashevich, 1993 - Karabastau Formation (Callovian/Oxfordian), Kazakhstan
†Eoptychoptera maxima Kalugina, 1985 - Itat Formation (Bajocian), Russia
†Eoptychoptera modica Lukashevich, 1993 - Itat Formation (Bajocian), Russia
†Eoptychoptera paramaculata Kalugina, 1985 - Itat Formation (Bajocian), Russia
†Eoptychoptera shurabica Lukashevich, 2000 - Sagul Formation (Toarcian), Kyrgyzstan
†Eoptychoptera simplex Handlirsch, 1906 - Ciechocinek Formation (Toarcian), Germany
†Eoptychoptera spectra Whalley, 1985 - Charmouth Mudstone Formation (Sinemurian), United Kingdom
†Eoptychoptera tempestilla Lukashevich, 1998 - Ulan Malgait Formation (Tithonian), Mongolia
†Eoptychoptera vitrea Lukashevich, 1998 - Dzhil Formation (Hettangian), Kyrgyzstan
†Eoptychopterina Kalugina 1985
†Eoptychopterina adnexa Hao et al. 2009 - Daohugou Formation (Callovian), China
†Eoptychopterina angularis Lukashevich, 1993 - Karabastau Formation (Callovian/Oxfordian), Kazakhstan
†Eoptychopterina antica Hao et al. 2009 - Daohugou Formation (Callovian), China
†Eoptychopterina baisica Kalugina, 1989 - Zaza Formation (Aptian), Russia
†Eoptychopterina camura Lukashevich et al., 2001 - Weald Clay (Hauterivian), United Kingdom
†Eoptychopterina daiensis Kalugina, 1989 - Glushkovo Formation (Tithonian), Russia
†Eoptychopterina demissa Lukashevich et al., 2001 - Durlston Formation (Berriasian), United Kingdom
†Eoptychopterina dimidiata Lukashevich et al., 2001 - Durlston Formation (Berriasian), United Kingdom
†Eoptychopterina elenae Ren & Krzemiński 2002 - Daohugou Formation (Callovian), China
†Eoptychopterina glabra Lukashevich, 1993 - Itat Formation (Bajocian), Russia
†Eoptychopterina kaluginae Lukashevich, 2004 - Ulan Malgait Formation (Tithonian), Mongolia
†Eoptychopterina karatavica Lukashevich, 1993 - Karabastau Formation (Callovian/Oxfordian), Kazakhstan
†Eoptychopterina mediata Hao et al. 2009 - Daohugou Formation (Callovian), China
†Eoptychopterina omissa Lukashevich, 2004 - Karabastau Formation (Callovian/Oxfordian), Kazakhstan
†Eoptychopterina petri Lukashevich, 2004 - Glushkovo Formation (Tithonian), Russia
†Eoptychopterina postica Liu et al. 2012 - Daohugou Formation (Callovian), China
†Eoptychopterina rohdendorphi Kalugina, 1985 - Ichetuy Formation (Oxfordian), Russia
†Eoptychopterina transbaicalica Kalugina, 1985 - Uda Formation (Oxfordian/Kimmeridgian), Russia
†Eoptychopterina undensis Kalugina, 1989 - Glushkovo Formation (Tithonian), Russia
†Leptychoptera Lukashevich & Azar 2003
†Leptychoptera subgenus (Burmaptychoptera) Lukashevich, 2004
†Leptychoptera (Burmaptychoptera) calva Lukashevich, 2004 - Burmese amber (Cenomanian), Myanmar
†Leptychoptera (Burmaptychoptera) reburra Lukashevich, 2004 - Burmese amber (Cenomanian), Myanmar
†Leptychoptera subgenus (Leptychoptera) Lukashevich & Azar, 2003
†Leptychoptera (Leptychoptera) dimkina Lukashevich & Azar, 2003 - Lebanese amber (Barremian) Lebanon
†Leptychoptera (Leptychoptera) vovkina Lukashevich & Azar, 2003 - Lebanese amber (Barremian) Lebanon
†Nedoptychoptera Lukashevich 1998
†Nedoptychoptera karatavica - Karabastau Formation (Callovian/Oxfordian), Kazakhstan
†Neuseptychoptera Szadziewski et al. 2017
†Neuseptychoptera carolinensis - Neuse River amber (Campanian), North Carolina,
†subfamily Proptychopterininae Lukashevich, 1995
†Proptychopterina Kalugina, 1985
†Proptychopterina amota Lukashevich, 1993 - Itat Formation (Bajocian) Russia
†Proptychopterina evecta Lukashevich, 1993 - Karabastau Formation (Callovian/Oxfordian) Kazakhstan
†Proptychopterina gracilis Lukashevich, 1993 - Karabastau Formation (Callovian/Oxfordian) Kazakhstan
†Proptychopterina handlirschi Kalugina, 1985 - Itat Formation (Bajocian) Russia
†Proptychopterina immensa Lukashevich, 2000 - Khaya Formation (Tithonian), Russia
†Proptychopterina makarova Lukashevich, 2000 - Makarova Formation (Toarcian), Russia
†Proptychopterina mongolica Kalugina, 1988 - Ulaan-Ereg Formation (Tithonian), Mongolia
†Proptychopterina oleynikovi Kalugina, 1989 - Glushkovo Formation (Tithonian), Russia
†Proptychopterina opinata Lin & Lukashevich, 2006 -Daohugou Formation (Callovian), China
†Proptychopterina sharategica Kalugina, 1992 - Ulan Malgait Formation (Tithonian), Mongolia
†Proptychopterina tenera Lukashevich, 2000 - Khaya Formation (Tithonian), Russia
†Proptychopterina yeniseica Lukashevich, 1993 - Itat Formation (Bajocian) Russia
Subfamily Ptychopterinae Schiner 1864
†Brodilka Lukashevich et al. 2001
†Brodilka mitchelli Lukashevich et al. 2001 - Lulworth Formation (Berriasian) United Kingdom
Ptychoptera Meigen, 1803
†Ptychoptera deleta Novák, 1878 - Cypris Formation (Miocene), Czech Republic
†Ptychoptera eocenica Podenas, 2007 - Baltic amber (Priabonian), Russia
†Ptychoptera mesozoica Kalugina, 1989 - Zaza Formation (Aptian), Russia
†Ptychoptera miocenica Cockerell, 1910 - Florissant Formation (Priabonian), Colorado
†Sinoptychopterites Hong, 2002
†Sinoptychopterites paludus Hong, 2002 - Fushin amber (Ypresian), China
Subfamily incertae sedis
†Bolboia Kalugina, 1989
†Bolboia mira Kalugina, 1989 - Byankino Formation (Tithonian), Russia
†Crenoptychoptera Kalugina, 1985
†Crenoptychoptera antica Kalugina, 1985 - Itat Formation (Bajocian) Russia
†Crenoptychoptera bavarica Krzemiński & Ansorge, 1995 - Solnhofen Formation (Tithonian), Germany
†Crenoptychoptera conspecta Lukashevich, 1995 - Cheremkhovskaya Formation, (Toarcian), Russia
†Crenoptychoptera decorosa Hao et al. 2009 - Daohugou Formation (Callovian), China
†Crenoptychoptera defossa Kalugina, 1985 - Itat Formation (Bajocian) Russia
†Crenoptychoptera dobbertinensis Ansorge, 1998 - Ciechocinek Formation (Toarcian), Germany
†Crenoptychoptera gronskayae Kalugina, 1989 - Khaya Formation (Tithonian), Russia
†Crenoptychoptera liturata Lukashevich, 2011 - Ulan Malgait Formation (Tithonian), Mongolia
†Etoptychoptera Handlirsh 1910
†Etoptychoptera tertiaria Handlirsh 1910 - Allenby Formation (Ypresian), Canada

References
Further reading

Borror, D.J., C.A. Triplehorn, & N.A. Johnson. 1989. An Introduction to the Study of Insects, Sixth edition. Saunders College Publishing.
Byers, G.W. & D.A. Rossman. 2004. Preliminary survey of the crane flies of Louisiana (Diptera: Tipulidae, Ptychopteridae). Proceedings of the Entomological Society of Washington, 106: 884–890.
Stubbs, A.E. 1993. Provisional Atlas of the Ptychopteroid Craneflies (Diptera: Ptychopteridae) of Britain and Ireland. Centre for Ecology and Hydrology.
Zwick, P. 2004. Fauna Europaea: Ptychopteridae. In: Fauna Europaea: DipteraNematocera. H. DeJong, ed.