View source for Ptychopteridae

{{Short description|Family of flies}}
{{automatic taxobox
| taxon = Ptychopteridae
| fossil_range = {{fossilrange|Early Jurassic|Present|earliest=Late Triassic}}
| image = Ptychoptera_contaminata.jpg
| image_caption = ''[[Ptychoptera contaminata]]''
| authority = Osten-Sacken, 1862
| display_parents = 3
| subdivision_ranks = Genera
| subdivision = * ''Ptychoptera''
* ''Bittacomorpha''
* ''Bittacomorphella''
}}
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 [[Insect wing|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 {{convert|0.8|×|0.2|mm|in|abbr=on}}. Duration is reported at 7 days.

===Larvae===
The [[larva]]e are eucephalous and distinctive for the long, caudal respiratory siphon they possess. At hatching, they measure just under {{convert|4|mm|in|abbr=on}} in ''P. albimana'', quickly growing to nearly {{convert|80|mm|in|abbr=on}}. 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.

[[File:Ptychoptera 2.jpg|thumb|''Ptychoptera quadrifasciata'']]

===Adult===
The characteristic and unique feature of the family is a lobe at the base of the haltere called the prehaltere. 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; M<sub>1</sub> cell present
*''Ptychoptera'' {{small|Meigen, 1803}}
Bittacomorphinae – 20 antennomeres; M<sub>1</sub> & M<sub>2</sub> veins fused, thus without M<sub>1</sub> cell
*''Bittacomorpha'' {{small|Westwood, 1835}}
*''Bittacomorphella'' {{small|Alexander, 1916}}

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

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

==References==
{{Reflist}}

==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. [https://biostor.org/reference/55180 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: Diptera: Nematocera.'' H. DeJong, ed.

==External links==
* [http://www.catalogueoflife.org/info_about_col.php Catalog of Life]
* [http://bugguide.net/node/view/120/bgimage Bug Guide]
*[http://www.lesinsectesduquebec.com/insecta/29-diptera/Bittacomorpha_clavipes.htm Ptychopteridae at Insects of Quebec webpage]
*[http://www.troutnut.com/naturals/other/other_44_2.jpg Photograph of larva at Troutnut.com website]
*[http://www.diptera.info/photogallery.php?album_id=88 Diptera.info Gallery]
*[https://www.youtube.com/watch?v=JeeJ2sd4K4M Video]

{{Diptera|1}}
{{Taxonbar|from=Q1394972}}

[[Category:Ptychopteridae| ]]
[[Category:Nematocera families]]