Chionea

Life-forms

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: Tipulomorpha
Superfamilia: Tipuloidea

Familia: Limoniidae
Subfamilia: Chioneinae
Genus: Chionea
Subgenera: C. (Chionea) - C. (Sphaeconophilus)
Name

Chionea Dalman, 1816
References

De Jong, H.; Ciliberti, P. 2014: How cold-adapted flightless flies dispersed over the northern hemisphere: phylogeny and biogeography of the snow fly genus Chionea Dalman (Diptera: Limoniidae). Systematic entomology, 39(3): 563–589. DOI: 10.1111/syen.12075 Reference page.
Zhang, X.; Wang, J.; Yang, D. 2012: Chionea Dalman newly recorded from China with descriptions of two new species (Diptera, Limoniidae). Zootaxa 3316: 1–14. Preview Reference page.

Chionea is a genus of wingless limoniid crane flies. It consists of two subgenera, the holarctic Chionea and palaearctic Sphaeconophilus. About 40 species are currently recognized in the northern hemisphere, but there are probably several undescribed species. They are commonly called snow flies.

Description

Adults occur during winter, where they can be observed walking over snow. They produce glycerol in their hemolymph, preventing them from freezing.[1]

Up to 200 eggs have been found in female snow flies, which are laid singly.[1] The larvae occur under decaying leaves in wooded regions (Marchand, 1917). At least the larvae of some species seem to feed on feces in rodent burrows.[1]

Adults seem to actively seek out the coldest place they can find and drink water by pressing their proboscis against the snow (Marchand, 1917), but are not known to feed. Adult snow flies live for up to two months. They can walk at a speed of about 1.30 m per minute, and at least males have been observed to leap, without being provoked.[1]

Male C. alexandriana are about seven millimeters long.

They are easiest found on snowbanks in the afternoon, in areas that are to some extent forested. Some species have been found to be abundant on the floor of caves.[1]

The winglessness of the genus is probably attributable to the fact that at sub-freezing temperatures, it is very hard to generate enough energy for maintaining flight muscles. Also, the space normally taken by flight muscles is used to store eggs. However, the halteres have not been reduced and are innervated, thus probably serving as sensory organs.[1]
Cold tolerance

As some species of Chionea are adapted to living on snow and in subnivean habitats,[1][2] they are considered cold tolerant insects. It has been frequently reported that Chionea are observed at temperatures between 0 and -6 °C.[2] In an experiment conducted in a laboratory environment, Chionea araneoides specimens died at a mean temperature of -7.5 °C, their supercooling point;[3] another experiment on an undetermined species of European Chionea demonstrated a supercooling point of -11.2 °C.[4] The antifreeze agent found in Chionea hemolymph has been determined to be sugar trehalose.[4]
Reproduction

Mating is indiscriminate (probably because it is very hard to find a specimen of the other sex for them) and takes 30 to 70 minutes. While winged crane flies face in opposite directions, snow flies position themselves similar to many beetles, thereby requiring the male copulatory organs to be inverted 180 degrees.[1]
Predators

One of the reasons why adults emerge in winter seems to be the absence of predators. However, "rock crickets" (Grylloblattidae) have been shown to feed on them. Because the cysticercoid form of a tapeworm species has been found in two out of three C. stoneana specimens in eastern Kansas, they also have to be eaten by mice, which are the tapeworm's host. The snow fly larvae likely take up these tapeworms via mouse feces.[5]
A high magnification dorsal view of a snow fly specimen with a nematode dauer placed around the snow fly's cervix ("neck").
Parasites

Tapeworm cysticercoids, possibly of the genus Hymenolepis, have been found in the abdomens of Chionea stoneana.[1]

While most likely not a parasite of snow flies, a species of nematode, similar to Rhabditis, can sometimes be found in a ringed capsule, which is called a (dauer), around the neck of adult snow flies.[1] Immature nematodes develop inside the dauer before ultimately dispersing from it.[1][5] It has been speculated that the female nematode lays this ring around the fly's neck when it emerges from the pupa and cannot yet move properly. These nematodes are presumed to be phoretic, meaning that they use the snow fly as a mechanism for dispersal, and not snow fly parasites.[1]
Species

This list was adapted from the "Catalogue of the Craneflies of the World (Diptera, Tipuloidea: Pediciidae, Limoniidae, Cylindrotomidae, Tipulidae)". Naturalis Biodiversity Center. (January 2007).

Subgenus Chionea (Holarctic)

Chionea albertensis Alexander, 1941 (Canada, United States south to Oregon, Idaho)
Chionea alexandriana Garrett, 1922 (Canada, USA to California, Utah)
Chionea araneoides Dalman, 1816 (Europe: Sweden to Italy, Romania and Russia)
Chionea carolus Byers, 1979 (USA: California, Nevada)
Chionea crassipes Boheman, 1846
Chionea crassipes crassipes (Scandinavia, Russia to Japan)
Chionea crassipes magadanensis Narchuk, 1998 (Russia: Magadan Oblast)
Chionea durbini Byers, 1983 (USA: California)
Chionea excavata Byers, 1983 (USA: California, Nevada)
Chionea hybrida Byers, 1983 (USA: Idaho, Utah)
Chionea jellisoni Byers, 1983 (USA: Montana, Idaho, Utah)
Chionea jenniferae (Byers, 1995) (USA: California)
Chionea kanenoi Sasakawa, 1986 (Japan: Honshū)
Chionea lyrata Byers, 1983 (USA: California)
Chionea macnabeana Alexander, 1947 (Canada, USA to Washington, Oregon)
Chionea mirabilis Vanin, 2008 (Korea)
Chionea nigra Byers, 1983 (USA: Utah, Colorado)
Chionea nipponica Alexander, 1932 (Russia:Primorsky Krai, Japan)
Chionea nivicola Doane, 1900 (USA: Washington, Oregon, Idaho, Montana)
Chionea obtusa Byers, 1983 (Canada, USA to Oregon, Idaho)
Chionea pusilla Savchenko, 1983 (Russia: Primorsky Krai)
Chionea racovitzai Burghele-Balacesco, 1969 (Romania)
Chionea reclusa Byers, 1995 (USA: Illinois)
Chionea scita Walker, 1848 (Canada, USA to Kentucky, Tennessee and Georgia)
Chionea stoneana Alexander, 1940 (USA: Minnesota to Indiana, south to Kansas, Oklahoma and Missouri)
Chionea valga Harris, 1841 (Canada, USA south to Minnesota, Wisconsin, Tennessee, North Carolina and Virginia)
Chionea wilsoni Byers, 1983 (USA: Alabama, Tennessee, North Carolina)

Subgenus Sphaeconophilus (Palaearctic)

Chionea alpina Bezzi, 1908 (Europe: France to Spain, Germany to Montenegro)
Chionea ancae (Menier and Matile, 1976) (France)
Chionea arverna (Brunhes, 1986) (France: Massif Central)
Chionea austriaca (Christian, 1980) (Austria)
Chionea belgica (Becker, 1912) (Europe: Belgium, Denmark, Germany, Hungary, Netherlands, Switzerland)
Chionea besucheti (Bourne, 1979) (Switzerland)
Chionea botosaneanui (Burghele-Balacesco, 1969) (Czech Republic, Poland, Romania, Slovakia)
Chionea catalonica (Bourne, 1979) (Spain: central Pyrenees)
Chionea jurassica (Bourne, 1979) (France)
Chionea lutescens
Chionea lutescens lutescens Lundstrom, 1907 (Europe: Sweden to Portugal, Ukraine, Russia)
Chionea lutescens stelviana Suss, 1982 (Italy: Lombardia)
Chionea pyrenaea (Bourne, 1981) (France: Pyrenees)