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Grylloblattidae, commonly known as the icebugs or ice crawlers, is a
family Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...
of
extremophile An extremophile () is an organism that is able to live (or in some cases thrive) in extreme environments, i.e., environments with conditions approaching or stretching the limits of what known life can adapt to, such as extreme temperature, press ...
(
psychrophile Psychrophiles or cryophiles (adj. ''psychrophilic'' or ''cryophilic'') are extremophile, extremophilic organisms that are capable of cell growth, growth and reproduction in low temperatures, ranging from to . They are found in places that are pe ...
) and
wingless In cellular biology, 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, pronounced "wint", is a portmanteau created from the ...
insect Insects (from Latin ') are Hexapoda, hexapod invertebrates of the class (biology), class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body (Insect morphology#Head, head, ...
s that live in the cold on top of mountains and the edges of glaciers. It is the only member of Grylloblattodea, which is generally considered an
order Order, ORDER or Orders may refer to: * A socio-political or established or existing order, e.g. World order, Ancien Regime, Pax Britannica * Categorization, the process in which ideas and objects are recognized, differentiated, and understood ...
. Alternatively, Grylloblattodea, along with
Mantophasmatodea Mantophasmatidae is a family of carnivorous wingless insects within the monotypic order Mantophasmatodea, which was discovered in Africa in 2001. Recent evidence indicates a sister group relationship with Grylloblattidae (classified in the orde ...
(rock crawlers), have been ranked as suborders of the order
Notoptera Notoptera, also known as Xenonomia, is a clade of insects belonging to Polyneoptera. It contains two living groups, Mantophasmatidae (gladiators) native to southern Africa, and Grylloblattidae (ice crawlers) native to cold montane environment ...
. Grylloblattids are wingless insects mostly less than 3 cm long, with a head resembling that of a
cockroach Cockroaches (or roaches) are insects belonging to the Order (biology), order Blattodea (Blattaria). About 30 cockroach species out of 4,600 are associated with human habitats. Some species are well-known Pest (organism), pests. Modern cockro ...
, with long antennae and having elongated cerci arising from the tip of their abdomen. They cannot tolerate warmth—most species will die at —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 games, bat-and-ball game played between two Sports team, teams of eleven players on a cricket field, field, at the centre of which is a cricket pitch, pitch with a wicket at each end, each comprising two Bail (cr ...
-
cockroach Cockroaches (or roaches) are insects belonging to the Order (biology), order Blattodea (Blattaria). About 30 cockroach species out of 4,600 are associated with human habitats. Some species are well-known Pest (organism), pests. Modern cockro ...
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 ( or ) is organic matter made up of the decomposition, decomposing remains of organisms and plants, and also of feces. Detritus usually hosts communities of microorganisms that colonize and decomposition, decompose (Reminera ...
. 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 or ocellus (sometimes called a pigment pit) is a form of eye or an optical arrangement which has a single lens without the sort of elaborate retina that occurs in most vertebrates. These eyes are called "simple" to distinguish the ...
(simple eyes). Their closest living relatives are the recently discovered
Mantophasmatodea Mantophasmatidae is a family of carnivorous wingless insects within the monotypic order Mantophasmatodea, which was discovered in Africa in 2001. Recent evidence indicates a sister group relationship with Grylloblattidae (classified in the orde ...
. Most species are less than 3 cm long, the largest being ''Namkungia magnus''. The family has its own
order Order, ORDER or Orders may refer to: * A socio-political or established or existing order, e.g. World order, Ancien Regime, Pax Britannica * Categorization, the process in which ideas and objects are recognized, differentiated, and understood ...
, Grylloblattodea (sometimes considered a suborder of Notoptera). 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 There are various disparate groups of wingless insects. Apterygota are a subclass of small, agile insects, distinguished from other insects by their lack of wings in the pr ...
'' and '' G. oregonensis'' are known from single sites.


Habitat and distribution

Grylloblattodea are nocturnal
extremophile An extremophile () is an organism that is able to live (or in some cases thrive) in extreme environments, i.e., environments with conditions approaching or stretching the limits of what known life can adapt to, such as extreme temperature, press ...
s typically found in
leaf litter Plant litter (also leaf litter, tree litter, soil litter, litterfall, or duff) is dead plant material (such as leaves, bark, needles, twigs, and cladodes) that has fallen to the ground. This detritus or dead organic material and its constituen ...
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 . 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 lower-elevation forests in western North America, East Asia (Korea and Japan), and 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 Earth's highest peaks, including the highest, Mount Everest. More than list of h ...
.


Diet

Grylloblattids are omnivorous, but feed primarily on dead arthropods and carrion. When arthropod carcasses are scarce, they subsist on plant material.


Evolution

Grylloblattidae is generally thought to have emerged from within the "
Grylloblattida Notoptera, also known as Xenonomia, is a clade of insects belonging to Polyneoptera. It contains two living groups, Mantophasmatidae (gladiators) native to southern Africa, and Grylloblattidae (ice crawlers) native to cold montane environments ...
", a poorly defined group of extinct winged insects that first appeared in the Late
Carboniferous The Carboniferous ( ) is a Geologic time scale, geologic period and System (stratigraphy), system of the Paleozoic era (geology), era that spans 60 million years, from the end of the Devonian Period Ma (million years ago) to the beginning of the ...
, over 300 million years ago. The winged '' Aristovia'' from the mid-Cretaceous
Burmese amber Burmese amber, also known as Burmite or Kachin amber, is amber from the Hukawng Valley in northern Myanmar. The amber is dated to around 100 million years ago, during the latest Albian to earliest Cenomanian ages of the mid-Cretaceous period. Th ...
of Myanmar, around 100 million years ago is thought to be closely related to modern Grylloblattidae due to its very similar mouthparts.


Taxonomy

List of Grylloblattodea genera and species along with their type localities: *'' Arctigalloisiana'' Nakahama, Yamasaki, Komazawa & Nakano, 2023 – Hokkaido, Japan **'' Arctigalloisiana poropnetopa'' Nakahama, Yamasaki, Komazawa & Nakano, 2023 – type locality: Kineusu,
Urakawa, Hokkaido is a town located in Hidaka Subprefecture, Hokkaido, Japan. As of 1 October 2020, the town has an estimated population of 12,074 and a density of 17 persons per km2. The total area is 694.24 km2. Climate Urakawa has a humid continental cl ...
, Japan **'' Arctigalloisiana yezoensis'' (Asahina 1961) – type locality: Miyazaki-Toge, Japan **'' Arctigalloisiana yubariensis'' Nakahama, Yamasaki, Komazawa & Nakano, 2023 – type locality: Niwan Pass,
Atsuma, Hokkaido is a town located in Iburi Subprefecture, Hokkaido, Japan. In March 2017, the town had an estimated population of 4,659, and a density of 12 persons per km2. The total area is 404.56 km2. The annual town festival is in mid-July, and is calle ...
, Japan *'' Galloisiana'' Caudell 1924 – Far East Asia **'' Galloisiana chujoi'' Gurney 1961 – type locality: Oninoiwaya Cave, Japan **'' Galloisiana kiyosawai'' Asahina 1959 – type locality: Hirayu-Onsen, Japan **'' Galloisiana kosuensis'' Namkung 1974 – type locality: Gosu Cave, South Korea **''
Galloisiana nipponensis ''Galloisiana nipponensis'' is a species of insect in the family Grylloblattidae that is endemic to Japan. Its type locality is Lake Chūzenji, Japan. Range and habitat It is found in montane habitats in central Honshu , historically known ...
'' (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 ar ...
, Japan **'' Galloisiana notabilis'' (Silvestri 1927) – type locality:
Nagasaki Prefecture is a Prefectures of Japan, prefecture of Japan, mainly located on the island of Kyūshū, although it also includes a number of islands off Kyūshū's northwest coast - including Tsushima and Iki. Nagasaki Prefecture has a population of 1,246,4 ...
, Japan **'' Galloisiana odaesanensis'' Kim & Lee 2007 – type locality: Mount Odae, South Korea **'' Galloisiana olgae'' Vrsansky & Storozhenko 2001 – type locality: Mount Olga, Russia **'' Galloisiana sinensis'' Wang 1987 – type locality: Changbaishan,
Jilin ) , image_skyline = Changbaishan Tianchi from western rim.jpg , image_alt = , image_caption = View of Heaven Lake , image_map = Jilin in China (+all claims hatched).svg , mapsize = 275px , map_al ...
, PR China **'' Galloisiana sofiae'' Szeptycki 1987 – type locality:
Mount Myohyang Mount Myohyang () is a mountain in North Korea. It is named for the mystic shapes and fragrances found in the area. In the foundation myth of Gojoseon, the ''Samguk sagi ''Samguk sagi'' () is a historical record of the Three Kingdoms of Korea ...
, North Korea **'' Galloisiana ussuriensis'' Storozhenko 1988 – type locality:
Primorsky Krai Primorsky Krai, informally known as Primorye, is a federal subjects of Russia, federal subject (a krais of Russia, krai) of Russia, part of the Far Eastern Federal District in the Russian Far East. The types of inhabited localities in Russia, ...
, Russia **'' Galloisiana yuasai'' Asahina 1959 – type locality: Tokugo-Toge, Japan *'' Grylloblatta'' Walker 1914 – western North America **''
Grylloblatta barberi ''Grylloblatta barberi'' is a North American species of wingless insect There are various disparate groups of wingless insects. Apterygota are a subclass of small, agile insects, distinguished from other insects by their lack of wings in the pr ...
'' Caudell 1924 – type locality: Sunny Side Mine,
Mount Lassen Lassen Peak ( ), commonly referred to as Mount Lassen, is a lava dome volcano in Lassen Volcanic National Park in Northern California. Located in the Shasta Cascade region above the northern Sacramento Valley, it is the southernmost active vo ...
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 pass with a road in California and in the Sierra Nevada. It is lower than Tioga Pass to the south. State Route 108 trav ...
, 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'' Kamp 1963 – type locality: Eagle Lake, California, USA **'' Grylloblatta chintimini'' Marshall & Lytle 2015 – type locality:
Marys Peak Marys Peak (formerly Mary's Peak and sometimes still spelled this way) is a mountain in Benton County, Oregon, Benton County, Oregon, United States, just southwest of Philomath, Oregon, Philomath. The Kalapuya people call the peak "chatímanwi" ...
, 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 Cowlitz, Lewis, and Skamania Counties, Washington.About 70% of the national monument, including Mount St. Helens itself, i ...
, Washington, USA **'' Grylloblatta gurneyi'' Kamp 1963 – type locality:
Lava Beds National Monument Lava Beds National Monument is located in northeastern California, in Siskiyou County, California, Siskiyou and Modoc County, California, Modoc counties. The monument lies on the northeastern flank of Medicine Lake Volcano, which is the largest ...
, 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 P ...
, Oregon, USA **''
Grylloblatta oregonensis ''Grylloblatta oregonensis'' 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 ...
'' 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 ...
, USA **'' Grylloblatta rothi'' Gurney 1953 – type locality: Happy Valley,
Deschutes County Deschutes County ( ) is one of the 36 counties in the U.S. state of Oregon. As of the 2020 census, the population was 198,253. The county seat is Bend. The county was created in 1916 out of part of Crook County and was named for the Deschu ...
, 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 s ...
, Alberta, Canada **'' Grylloblatta sculleni'' Gurney 1937 – type locality: Scott Camp,
Deschutes County Deschutes County ( ) is one of the 36 counties in the U.S. state of Oregon. As of the 2020 census, the population was 198,253. The county seat is Bend. The county was created in 1916 out of part of Crook County and was named for the Deschu ...
, 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 ...
, 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 6 ...
, California, USA *'' Grylloblattella'' Storozhenko 1988 – Central Asia **'' Grylloblattella cheni'' Bai, Wang & Yang 2010 – type locality:
Ake Kule Lake Ake Kule Lake () is a lake in Altay Prefecture, Xinjiang, China, close to the borders of Kazakhstan, Mongolia and Russia Russia, or the Russian Federation, is a country spanning Eastern Europe and North Asia. It is the list of countries an ...
,
Xinjiang Xinjiang,; , SASM/GNC romanization, SASM/GNC: Chinese postal romanization, previously romanized as Sinkiang, officially the Xinjiang Uygur Autonomous Region (XUAR), is an Autonomous regions of China, autonomous region of the China, People' ...
, China **'' Grylloblattella pravdini'' (Storozhenko & Oliger 1984) – type locality: Teletskoye Lake, Russia **'' Grylloblattella sayanensis'' Storozhenko 1996 – type locality: Sambyl Pass, Russia *'' Grylloblattina'' Bey-Bienko 1951 – Far East Asia **'' Grylloblattina djakonovi'' Bey-Bienko 1951 *'' Namkungia'' 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 35 extant species and 6 extant genera described as of 2023.


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 National Forest, Willamette and Deschutes National Forests in Oregon, United States. It comprises , making it the second-largest wilderness area in Oreg ...
of Oregon {{Taxonbar, from1=Q13582699, from2=Q796470 Insect families Psychrophiles Extant Triassic first appearances Taxa named by Edmund Murton Walker