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Grylloblattidae, commonly known as the icebugs, or ice crawlers, is a
family Family (from la, familia) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its ...
of
extremophile An extremophile (from Latin ' meaning "extreme" and Greek ' () meaning "love") is an organism that is able to live (or in some cases thrive) in extreme environments, i.e. environments that make survival challenging such as due to extreme temper ...
(
psychrophile Psychrophiles or cryophiles (adj. ''psychrophilic'' or ''cryophilic'') are extremophilic organisms that are capable of growth and reproduction in low temperatures, ranging from to . They have an optimal growth temperature at . They are found in ...
) and
wingless The Wnt signaling pathways are a group of signal transduction pathways which begin with proteins that pass signals into a cell through cell surface receptors. The name Wnt is a portmanteau created from the names Wingless and Int-1. Wnt signaling p ...
insect Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body ( head, thorax and abdomen), three pairs ...
s that live in the cold on top of mountains and the edges of glaciers. They belong, along with
Mantophasmatidae Mantophasmatidae is a family (biology), family of carnivorous wingless insects within the order Notoptera, which was discovered in Africa in 2001.K.-D. Klass, O. Zompro, N.P. Kristensen, J. Adis. Mantophasmatodea: a new insect order with extant ...
(rock crawlers), to the order
Notoptera The wingless insect order Notoptera, a group first proposed in 1915, had been largely unrecognized since its original conception, until resurrected in 2004. As now defined, the order comprises five families, three of them known only from fossils ...
.Arillo, A. & M. Engel (2006) Rock Crawlers in Baltic Amber (Notoptera: Mantophasmatodea). ''American Museum Novitates'' 3539:1-10
/ref> Grylloblattids are wingless insects mostly less than 3 cm long, with a head resembling that of a
cockroach Cockroaches (or roaches) are a paraphyletic group of insects belonging to Blattodea, containing all members of the group except termites. About 30 cockroach species out of 4,600 are associated with human habitats. Some species are well-known as ...
, with long antennae and having elongated cerci arising from the tip of their abdomen. They cannot tolerate warmth (most species will die at 10 °C) and many species have small distribution ranges.


Overview

Grylloblattids, ice crawlers or icebugs puzzled the scientists who discovered them in 1914, E.M. Walker and T.B. Kurata; the first species named was ''
Grylloblatta campodeiformis ''Grylloblatta campodeiformis'', also known as the northern rock crawler, is an omnivorous species of insect in family Grylloblattidae. Like other species in the genus '' Grylloblatta'', it is endemic to North America. Taxonomy This species was ...
'', which means "
cricket Cricket is a bat-and-ball game played between two teams of eleven players on a field at the centre of which is a pitch with a wicket at each end, each comprising two bails balanced on three stumps. The batting side scores runs by striki ...
-
cockroach Cockroaches (or roaches) are a paraphyletic group of insects belonging to Blattodea, containing all members of the group except termites. About 30 cockroach species out of 4,600 are associated with human habitats. Some species are well-known as ...
shaped like a ''
Campodea ''Campodea'' is a genus of small, white, bristle-tailed arthropods in the order Diplura. The best known species, ''Campodea staphylinus'', has a wide distribution across much of Europe. It lives in damp places under stones, fallen trees, or in ro ...
''" (a kind of two-pronged bristletail). Most are nocturnal and appear to feed on
detritus In biology, detritus () is dead particulate organic material, as distinguished from dissolved organic material. Detritus typically includes the bodies or fragments of bodies of dead organisms, and fecal material. Detritus typically hosts commun ...
. They have long antennae (23–45 segments) and long cerci (5–8 segments), but no wings. Their eyes are either missing or reduced and they have no
ocelli A simple eye (sometimes called a pigment pit) refers to a form of eye or an optical arrangement composed of a single lens and without an elaborate retina such as occurs in most vertebrates. In this sense "simple eye" is distinct from a multi-l ...
(simple eyes). Their closest living relatives are the recently discovered Mantophasmatodea. Most species are less than 3 cm long, the largest being ''Namkungia magnus''. The family has its own
suborder Order ( la, ordo) is one of the eight major hierarchical taxonomic ranks in Linnaean taxonomy. It is classified between family and class. In biological classification, the order is a taxonomic rank used in the classification of organisms and ...
, Grylloblattodea (formerly its own order). It contains 5 genera and about 34 extant species. Most species have restricted distributions and small populations and with increased warming their habitats are threatened, making them endangered. In North America some species like ''
Grylloblatta barberi ''Grylloblatta barberi'' is a North American species of wingless insect in the genus '' Grylloblatta''. It is a rock crawler that lives at high altitudes in crevices under snow or glaciers. It was first described by Andrew Nelson Caudell in 192 ...
'' and '' G. oregonensis'' are known from single sites.


Habitat and distribution

Grylloblattodea are nocturnal
extremophile An extremophile (from Latin ' meaning "extreme" and Greek ' () meaning "love") is an organism that is able to live (or in some cases thrive) in extreme environments, i.e. environments that make survival challenging such as due to extreme temper ...
s typically found in leaf litter and under stones in extremely cold environments, usually at higher elevations. They are known to inhabit cold temperate forests to glaciers and the edges of ice sheets. Their optimal living temperature is between 1-4 °C (33.8-39.2 °F). They can be killed at colder temperatures due to ice formation in the body, so when the temperature drops below their optimal range they survive by living under snow pack near the soil. They have a very narrow range of temperatures that they prefer and cannot withstand high temperatures; many species are killed when the temperature rises about 5°C above their optimal temperature. They move in response to the seasons so as to maintain an optimal temperature in their foraging habitat. Grylloblattidae are patchily distributed in glaciers, caves, montane environments, and occasionally also lower-elevation forests in western North America, East Asia (Korea and Japan), Central Asia (Siberia, China, and Kazakhstan). They are predicted to occur in several other mountain chains in Asia, including parts of the
Himalayas The Himalayas, or Himalaya (; ; ), is a mountain range in Asia, separating the plains of the Indian subcontinent from the Tibetan Plateau. The range has some of the planet's highest peaks, including the very highest, Mount Everest. Over 100 ...
.


Diet

They are omnivorous, but feed primarily on dead arthropods and carrion. When arthropod carcasses are scarce, their diet relies heavily on plant material. The fossil species '' Plesioblattogryllus magnificus'' from the
Middle Jurassic The Middle Jurassic is the second epoch of the Jurassic Period. It lasted from about 174.1 to 163.5 million years ago. Fossils of land-dwelling animals, such as dinosaurs, from the Middle Jurassic are relatively rare, but geological formations co ...
had strong mandibles and is thought to have been a predator.


Evolution

Grylloblattidae is generally thought to have emerged from within the " Grylloblattida", a poorly defined group of extinct winged insects that first appeared in the Late
Carboniferous The Carboniferous ( ) is a geologic period and system of the Paleozoic that spans 60 million years from the end of the Devonian Period million years ago ( Mya), to the beginning of the Permian Period, million years ago. The name ''Carbonifero ...
.


Taxonomy

List of Grylloblattodea genera and species along with their type localities: *''
Galloisiana ''Galloisiana'' is a genus of insects in the family Grylloblattidae found in East Asia. It contains 12 species. Species These species belong to the genus ''Galloisiana'': *''Galloisiana chujoi'' Gurney 1961 – type locality: Oninoiwaya Cave, ...
'' Caudell & King 1924 – Far East Asia **''
Galloisiana chujoi ''Galloisiana chujoi'' is a species of cave-dwelling insect in the family Grylloblattidae Grylloblattidae, commonly known as the icebugs, or ice crawlers, is a family of extremophile (psychrophile) and wingless insects that live in the cold o ...
'' Gurney 1961 – type locality: Oninoiwaya Cave, Japan **'' Galloisiana kiyosawai'' Asahina 1959 – type locality: Hirayu-Onsen, Japan **'' Galloisiana kosuensis'' Namkung 1974 – type locality:
Gosu Cave Gosu Cave (고수동굴 - Kosu is also still found as an alternate spelling) is a huge limestone cave near Danyang, Chungcheongbuk-do, Danyang (North Chungcheong Province), South Korea formed over 450 million years ago. It is among the best known n ...
, South Korea **''
Galloisiana nipponensis ''Galloisiana nipponensis'' is a species of insect in the family Grylloblattidae that is endemic to Japan. Its Type locality (biology), type locality is Lake Chūzenji, Japan. Range and habitat It is found in montane habitats in central Honshu, ...
'' Caudell & King 1924 – type locality:
Lake Chūzenji , also called Sea of Happiness, is a scenic lake in Nikkō National Park in the city of Nikkō, Tochigi Prefecture, Japan. It was created 20,000 years ago when Mount Nantai (2484 m) erupted and blocked the river. The lake has a surface area of 11. ...
, Japan **''
Galloisiana notabilis ''Galloisiana notabilis'' is a species of insect in the family Grylloblattidae that is endemic to southern Japan. Its type locality is Nagasaki Prefecture is a prefecture of Japan located on the island of Kyūshū. Nagasaki Prefecture has a ...
'' Silvestri 1927 – type locality:
Nagasaki Prefecture is a Prefectures of Japan, prefecture of Japan located on the island of Kyūshū. Nagasaki Prefecture has a population of 1,314,078 (1 June 2020) and has a geographic area of 4,130 Square kilometre, km2 (1,594 sq mi). Nagasaki Prefecture borders ...
, Japan **''
Galloisiana odaesanensis ''Galloisiana odaesanensis'' is a species of insect in the family Grylloblattidae. Its Type locality (biology), type locality is Mount Odae in Gangwon Province, South Korea. Specimens have also been collected at Sangwonsa. References

Gr ...
'' Kim & Lee 2007 – type locality:
Mount Odae Odaesan, also known as Mount Odae-san or Mount Odae (), is a major cluster of mountains and national park in eastern Gangwon Province, South Korea, standing at the junction of Gangneung City, Pyeongchang County and Hongcheon County. It stands ...
, South Korea **'' Galloisiana olgae'' Vrsansky & Storozhenko 2001 – type locality: Mount Olga, Russia **'' Galloisiana sinensis'' Wang 1987 – type locality: Changbaishan,
Jilin Jilin (; alternately romanized as Kirin or Chilin) is one of the three provinces of Northeast China. Its capital and largest city is Changchun. Jilin borders North Korea (Rasŏn, North Hamgyong, Ryanggang and Chagang) and Russia (Prim ...
, PR China **''
Galloisiana sofiae ''Galloisiana sofiae'' is a species of insect in the family Grylloblattidae Grylloblattidae, commonly known as the icebugs, or ice crawlers, is a family of extremophile (psychrophile) and wingless insects that live in the cold on top of mount ...
'' Szeptycki 1987 – type locality: Mount Myoyang, South Korea **'' Galloisiana ussuriensis'' Storozhenko 1988 – type locality:
Primorsky Krai Primorsky Krai (russian: Приморский край, r=Primorsky kray, p=prʲɪˈmorskʲɪj kraj), informally known as Primorye (, ), is a federal subject (a krai) of Russia, located in the Far East region of the country and is a part of the ...
, Russia **'' Galloisiana yezoensis'' Asahina 1961 – type locality: Miyazaki-Toge, Japan **'' Galloisiana yuasai'' Asahina 1959 – type locality: Tokugo-Toge, Japan *''
Grylloblatta ''Grylloblatta'' is a genus of insects in the family Grylloblattidae. It contains 15 species, including '' Grylloblatta chirurgica'', almost exclusively from high-altitude and high-latitude regions of the United States, living in ice caves and g ...
'' Walker 1914 – western North America **''
Grylloblatta barberi ''Grylloblatta barberi'' is a North American species of wingless insect in the genus '' Grylloblatta''. It is a rock crawler that lives at high altitudes in crevices under snow or glaciers. It was first described by Andrew Nelson Caudell in 192 ...
'' Caudell 1924 – type locality: Sunny Side Mine,
Mount Lassen Mount is often used as part of the name of specific mountains, e.g. Mount Everest. Mount or Mounts may also refer to: Places * Mount, Cornwall, a village in Warleggan parish, England * Mount, Perranzabuloe, a hamlet in Perranzabuloe parish, C ...
area, California, USA **'' Grylloblatta bifratrilecta'' Gurney 1953 – type locality:
Sonora Pass Sonora Pass (el. 9,624 ft. / 2,933 m.) is a mountain pass in the Sierra Nevada in California. It is the second-highest in California and in the Sierra Nevada. lower by 321 ft. (about 98 m.) than Tioga Pass to the south. State Route 108 ...
, California, USA **''
Grylloblatta campodeiformis ''Grylloblatta campodeiformis'', also known as the northern rock crawler, is an omnivorous species of insect in family Grylloblattidae. Like other species in the genus '' Grylloblatta'', it is endemic to North America. Taxonomy This species was ...
'' Walker 1914 – type locality: Sulphur Mountain, Alberta, Canada **''
Grylloblatta chandleri ''Grylloblatta chandleri'' is a species of insect in the family Grylloblattidae. Its Type locality (biology), type locality is an ice cave in Eagle Lake (Lassen County), Eagle Lake in California, United States. It has also been recorded in Shast ...
'' Kamp 1963 – type locality:
Eagle Lake Eagle Lake may refer to: Cities, towns, townships etc. Canada * Eagle Lake, Haliburton County, Ontario * Eagle Lake, Parry Sound District, Ontario * Eagle Lake 27, Ontario (Indian reserve) * Eagle Lake, Kenora District, Ontario United States * ...
, California, USA **'' Grylloblatta chintimini'' Marshall & Lytle 2015 – type locality: Marys Peak, Oregon, USA **'' Grylloblatta chirurgica'' Gurney 1961 – type locality:
Ape Cave Mount St. Helens National Volcanic Monument is a U.S. National Monument that includes the area around Mount St. Helens in Washington. It was established on August 27, 1982, by U.S. President Ronald Reagan, following the 1980 eruption. The 110,0 ...
, Washington, USA **'' Grylloblatta gurneyi'' Kamp 1963 – type locality:
Lava Beds National Monument Lava Beds National Monument is located in northeastern California, in Siskiyou and Modoc counties. The monument lies on the northeastern flank of Medicine Lake Volcano and has the largest total area covered by a volcano in the Cascade Range. ...
, California, USA **'' Grylloblatta marmoreus'' Schoville 2012 – type locality: Marble Mountains, California, USA **'' Grylloblatta newberryensis'' Marshall & Lytle 2015 – type locality:
Newberry Volcano Newberry Volcano is a large active shield-shaped stratovolcano located about south of Bend, Oregon, United States, east of the major crest of the Cascade Range, within the Newberry National Volcanic Monument. Its highest point is Paulina ...
, Oregon, USA **''
Grylloblatta oregonensis ''Grylloblatta oregonensis'' is a species of insect in the family Grylloblattidae. Its type locality is in Oregon Caves National Monument Oregon Caves National Monument and Preserve is a protected area in the northern Siskiyou Mountains of ...
'' Schoville 2012 – type locality:
Oregon Caves National Monument Oregon Caves National Monument and Preserve is a protected area in the northern Siskiyou Mountains of southwestern Oregon in the United States. The 4,554-acre (1,843 ha) park, including the marble cave, is 20 miles (32 km) east of C ...
, USA **'' Grylloblatta rothi'' Gurney 1953 – type locality: Happy Valley, Deschutes County, Oregon, USA **'' Grylloblatta scudderi'' Kamp 1979 – type locality:
Mount Paul Mount Paul is a mountain summit towering 1134 metres above the east shore of Maligne Lake in Jasper National Park, in the Canadian Rockies of Alberta, Canada. The nearest higher peak is Maligne Mountain, to the north-northwest. Mount Paul is si ...
, Alberta, Canada **'' Grylloblatta sculleni'' Gurney 1937 – type locality: Scott Camp, Deschutes County, Oregon, USA **'' Grylloblatta siskiyouensis'' Schoville 2012 – type locality:
Oregon Caves National Monument Oregon Caves National Monument and Preserve is a protected area in the northern Siskiyou Mountains of southwestern Oregon in the United States. The 4,554-acre (1,843 ha) park, including the marble cave, is 20 miles (32 km) east of C ...
, USA **'' Grylloblatta washoa'' Gurney 1961 – type locality:
Echo Summit Echo Summit is a mountain pass over the Sierra Nevada in the western United States, located in eastern El Dorado County, California. At above sea level, it is the highest point on U.S. Route 50 in California, which traverses it at postmile 66 ...
, California, USA *''
Grylloblattella ''Grylloblattella'' is a genus of insects in the family Grylloblattidae found in the Altai-Sayan region of Central Asia. It contains 3 species restricted to montane environments in southern Siberia (Russia), China, and Kazakhstan. Species These ...
'' Storozhenko 1988 – Central Asia **'' Grylloblattella cheni'' Bai, Wang & Yang 2010 – type locality: Ake Kule Lake,
Xinjiang Xinjiang, SASM/GNC: ''Xinjang''; zh, c=, p=Xīnjiāng; formerly romanized as Sinkiang (, ), officially the Xinjiang Uygur Autonomous Region (XUAR), is an autonomous region of the People's Republic of China (PRC), located in the northwest ...
, China **'' Grylloblattella pravdini'' Storozhenko & Oliger 1984 – type locality:
Teletskoye Lake Lake Teletskoye (russian: Телецкое озеро, lit=the lake of the ; ) is the largest lake in the Altai Mountains and the Altai Republic, Russia, and has depth up to 325 meters. Situated at a height of above the sea level, the lake is ...
, Russia **'' Grylloblattella sayanensis'' Storozhenko 1996 – type locality: Sambyl Pass, Russia *''
Grylloblattina ''Grylloblattina'' is a genus of insects in the family Grylloblattidae. It is a monotypic genus consisting of the species ''Grylloblattina djakonovi''. Range and habitat ''Grylloblattina djakonovi'' is endemic to far eastern Russia, where it is ...
'' Bey-Bienko 1951 – Far East Asia **'' Grylloblattina djakonovi'' Bey-Bienko 1951 *''
Namkungia ''Namkungia'' is a genus of cave-dwelling insects in the family Grylloblattidae found in Korea. It contains 2 species, both of which are found in caves in Jeongseon County, Gangwon Province, South Korea. Species These species belong to the genu ...
'' Storozhenko & Park 2002 – Korea **'' Namkungia biryongensis'' (Namkung 1974) – type locality: Biryong Cave, South Korea **'' Namkungia magna'' (Namkung 1986) – type locality: Balgudeok Cave, South Korea In total, there are 33 extant species and 5 extant genera described as of 2015.


References


External links

* *
Video of a walking grylloblattid
in the
Three Sisters Wilderness The Three Sisters Wilderness is a wilderness area in the Cascade Range, within the Willamette and Deschutes National Forests in Oregon, United States. It comprises , making it the second largest wilderness area in Oregon, after the Eagle Cap Wilde ...
of Oregon {{Taxonbar, from=Q13582699 Insect families Psychrophiles Extant Triassic first appearances Taxa named by Edmund Murton Walker