Ptychopteridae

The Ptychopteridae, phantom crane flies, are a small family 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 has a mean of 554 eggs laid. The shape is slightly arcuated, "curiously ornamented", and roughly. 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 in P. albimana, quickly growing to nearly. They occur in moist habitats 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.

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 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”.

Evolution and relationships

The family Ptychopteridae is generally regarded as one of the most basal or primitive groups within the order Diptera, retaining several ancestral features shared with early fly lineages.
Despite their apparent antiquity, their precise relationships within the nematocerous flies remain unresolved. Morphological similarities link them to the Tanyderidae, and together these families are sometimes grouped as the Ptychopteromorpha. However, molecular and anatomical studies have yet to confirm whether this association represents a true evolutionary relationship or a convergence of structural traits.
Ptychopterids provide important insight into early dipteran evolution, preserving features that bridge the transition between primitive aquatic forms and the more derived higher flies. Their characteristic larval morphology—particularly the elongated respiratory siphon—and their simple adult structures may reflect an ecological continuity from ancestral aquatic dipterans.

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

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

Bittacomorpha
Bittacomorphella
Bittacomorphinae
*Bittacomorpha
**Bittacomorpha clavipes - Eastern NA to the edge of the Rocky Mountains
**Bittacomorpha occidentalis - Pacific Northwest
*Bittacomorphella
**Bittacomorphella jonesi - New England down to North Carolina, west to Minnesota and Michigan
**Bittacomorphella esakii - Japan
**Bittacomorphella fenderiana - Queen Charlotte Island down to Northern Oregon.
**Bittacomorphella nipponensis -
**Bittacomorphella pacifica - Northern California up through Oregon
**Bittacomorphella sackenii - Sierra Nevadas
**Bittacomorphella thaiensis -
Ptychopterinae
*Ptychoptera
**Ptychoptera byersi - California
**Ptychoptera lenis
***P. l. lenis - Pacific Northwest
***P. l. coloradensis - Colorado/Utah
**Ptychoptera metallica - Central Canada, Minnesota & Michigan
**Ptychoptera minor - California/Idaho
**Ptychoptera monoensis - Northern California
**Ptychoptera osceola - Florida
**Ptychoptera pendula - British Columbia to Utah & Colorado
**Ptychoptera quadrifasciata - Eastern North America
**Ptychoptera sculleni - Pacific Northwest
**Ptychoptera townesi - Washington & Oregon
**Ptychoptera uta - Utah
Fossil Subfamilies and genera
Subfamily Bittacomorphinae
* †Probittacomorpha
** †Probittacomorpha brisaci - Montagne d'Andance Miocene, France
** †Probittacomorpha christenseni - Fur Formation Eocene, Denmark;
* †Zhiganka
** †Zhiganka comitans - Batylykh Formation Early Cretaceous, Russia
** †Zhiganka longialata - Khasurty locality Early Cretaceous, Russia
** †Zhiganka woolgari - Weald Clay Early Cretaceous, England
Subfamily. †Eoptychopterinae
* †Architendipes
** †Architendipes tshernovskiji - Dzhil Formation Early Jurassic, Kyrgyzstan
* †Doptychoptera
** †Doptychoptera baisica - Zaza Formation Early Cretaceous, Russia
* †Eolimnobia
** †Eolimnobia geinitzi - Grimmen Formation Early Jurassic, Germany
* †Eoptychoptera
** †Eoptychoptera aequidistans - Dzhil Formation Early Jurassic, Kyrgyzstan
** †Eoptychoptera altaica - Ortsog Formation Middle Jurassic, Mongolia
** †Eoptychoptera ansorgei - Daohugou Formation Middle Jurassic, China
** †Eoptychoptera asiatica - Karabastau Formation Late Jurassic, Kazakhstan
** †Eoptychoptera aucta - Karabastau Formation Late Jurassic, Kazakhstan
** †Eoptychoptera braziliana - Crato Formation Early Cretaceous, Brazil
** †Eoptychoptera britannica - Weald Clay Early Cretaceous, England
** †Eoptychoptera cantabrica - Spanish amber, Spain
** †Eoptychoptera cretacea - Zaza Formation Early Cretaceous, Russia
** †Eoptychoptera elevata - Khaya Formation Late Jurassic, Russia
** †Eoptychoptera eximia - Ciechocinek Formation Early Jurassic, Germany
** †Eoptychoptera jurassica - Daohugou Formation Middle Jurassic, China
** †Eoptychoptera longifurcata - Lulworth Formation Early Cretaceous, England
** †Eoptychoptera magna - Karabastau Formation Late Jurassic, Kazakhstan
** †Eoptychoptera maxima - Itat Formation Middle Jurassic, Russia
** †Eoptychoptera modica - Itat Formation, Russia
** †Eoptychoptera paramaculata - Itat Formation, Russia
** †Eoptychoptera shurabica - Sagul Formation Early Jurassic, Kyrgyzstan
** †Eoptychoptera simplex - Ciechocinek Formation Early Jurassic, Germany
** †Eoptychoptera spectra - Charmouth Mudstone Formation, United Kingdom
** †Eoptychoptera tempestilla - Ulan Malgait Formation Late Jurassic, Mongolia
** †Eoptychoptera vitrea - Dzhil Formation Early Jurassic, Kyrgyzstan
*†Eoptychopterina
** †Eoptychopterina adnexa - Daohugou Formation Middle Jurassic, China
** †Eoptychopterina angularis - Karabastau Formation Late Jurassic, Kazakhstan
** †Eoptychopterina antica - Daohugou Formation Middle Jurassic, China
** †Eoptychopterina baisica - Zaza Formation Early Cretaceous, Russia
** †Eoptychopterina camura - Weald Clay Early Cretaceous, United Kingdom
** †Eoptychopterina daiensis - Glushkovo Formation Late Jurassic, Russia
** †Eoptychopterina demissa - Durlston Formation Early Cretaceous, United Kingdom
** †Eoptychopterina dimidiata - Durlston Formation Early Cretaceous, United Kingdom
** †Eoptychopterina elenae - Daohugou Formation Middle Jurassic, China
** †Eoptychopterina glabra - Itat Formation Middle Jurassic, Russia
** †Eoptychopterina kaluginae - Ulan Malgait Formation Late Jurassic, Mongolia
** †Eoptychopterina karatavica - Karabastau Formation Late Jurassic, Kazakhstan
** †Eoptychopterina mediata - Daohugou Formation Middle Jurassic, China
** †Eoptychopterina omissa - Karabastau Formation Late Jurassic, Kazakhstan
** †Eoptychopterina petri - Glushkovo Formation Late Jurassic, Russia
** †Eoptychopterina postica - Daohugou Formation Middle Jurassic, China
** †Eoptychopterina rohdendorphi - Ichetuy Formation Late Jurassic, Russia
** †Eoptychopterina transbaicalica - Uda Formation Late Jurassic, Russia
** †Eoptychopterina undensis - Glushkovo Formation Late Jurassic, Russia
* †Leptychoptera
** †Leptychoptera subgenus
*** †Leptychoptera calva - Burmese amber Mid Cretaceous, Myanmar
*** †Leptychoptera reburra - Burmese amber Mid Cretaceous, Myanmar
** †Leptychoptera subgenus
*** †Leptychoptera dimkina - Lebanese amber Early Cretaceous Lebanon
*** †Leptychoptera vovkina - Lebanese amber Early Cretaceous Lebanon
* †Nedoptychoptera
** †Nedoptychoptera karatavica - Karabastau Formation Late Jurassic, Kazakhstan
* †Neuseptychoptera
** †Neuseptychoptera carolinensis - Neuse River amber Late Cretaceous, North Carolina,
†subfamily Proptychopterininae
*†Proptychopterina
** †Proptychopterina amota - Itat Formation Middle Jurassic Russia
** †Proptychopterina evecta - Karabastau Formation Late Jurassic Kazakhstan
** †Proptychopterina gracilis - Karabastau Formation Late Jurassic Kazakhstan
** †Proptychopterina handlirschi - Itat Formation Middle Jurassic Russia
** †Proptychopterina immensa - Khaya Formation Late Jurassic, Russia
** †Proptychopterina makarova - Makarova Formation Early Jurassic, Russia
** †Proptychopterina mongolica - Ulaan-Ereg Formation Late Jurassic, Mongolia
** †Proptychopterina oleynikovi - Glushkovo Formation Late Jurassic, Russia
** †Proptychopterina opinata -Daohugou Formation Middle Jurassic, China
** †Proptychopterina sharategica - Ulan Malgait Formation Late Jurassic, Mongolia
** †Proptychopterina tenera - Khaya Formation Late Jurassic, Russia
** †Proptychopterina yeniseica - Itat Formation Middle Jurassic Russia
Subfamily Ptychopterinae
*†Brodilka
** †Brodilka mitchelli - Lulworth Formation Early Cretaceous United Kingdom
* Ptychoptera
** †Ptychoptera deleta - Cypris Formation, Czech Republic
** †Ptychoptera eocenica - Baltic amber, Russia
** †Ptychoptera mesozoica - Zaza Formation Early Cretaceous, Russia
** †Ptychoptera miocenica - Florissant Formation, Colorado
* †Sinoptychopterites
** †Sinoptychopterites paludus - Fushin amber, China
Subfamily incertae sedis
* †Bolboia
** †Bolboia mira - Byankino Formation Late Jurassic, Russia
* †Crenoptychoptera
** †Crenoptychoptera antica - Itat Formation Middle Jurassic Russia
** †Crenoptychoptera bavarica - Solnhofen Formation Late Jurassic, Germany
** †Crenoptychoptera conspecta - Cheremkhovskaya Formation, Early Jurassic, Russia
** †Crenoptychoptera decorosa - Daohugou Formation Middle Jurassic, China
** †Crenoptychoptera defossa - Itat Formation Middle Jurassic Russia
** †Crenoptychoptera dobbertinensis - Ciechocinek Formation Early Jurassic, Germany
** †Crenoptychoptera gronskayae - Khaya Formation Late Jurassic, Russia
** †Crenoptychoptera liturata - Ulan Malgait Formation Late Jurassic, Mongolia
* †Etoptychoptera
** †Etoptychoptera tertiaria - Allenby Formation, Canada