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Gynaephora groenlandica

Gynaephora groenlandica, the Arctic woolly bear moth, is an erebid moth native to the High Arctic in the Canadian archipelago, Greenland and Wrangel Island in Russia.[2][4] It is known for its slow rate of development, as its full caterpillar life cycle may extend up to 7 years, with moulting occurring each spring.[5] This species remains in a larval state for the vast majority of its life.[4][6] Rare among Lepidoptera, it undergoes an annual period of diapause that lasts for much of the calendar year, as G. groenlandica is subject to some of the longest, most extreme winters on Earth.[7][8] In this dormant state, it can withstand temperatures as low as −70 °C.[7] The Arctic woolly bear moth also exhibits basking behavior, which aids in temperature regulation and digestion and affects both metabolism and oxygen consumption.[4][6] Females generally do not fly, while males usually do.[7]

Arctic woolly bear moth
Arctic woolly bear caterpillar, Greenland
Scientific classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Lepidoptera
Superfamily: Noctuoidea
Family: Erebidae
Genus: Gynaephora
Species:
G. groenlandica
Binomial name
Gynaephora groenlandica
(Wocke ex. Homeyer, 1874)
Synonyms[3]
  • Dasychira groenlandica Wocke ex. Homeyer, 1874[1]
  • Dicallomera kusnezovi Lukhtanov et Khruliova, 1989[2]

This species has an alpine subspecies which is notable for its geographic distribution south of the High Arctic.[9]

Taxonomy edit

This moth was likely first discovered on 16 June 1832 on the beach of Fury Bay, Somerset Island, northern Nunavut, Canada, by the crew of the Arctic expedition searching for the Northwest Passage (led by John Ross). John Curtis, who studied the entomological specimens obtained from the voyage, described Gynaephora rossii from the invertebrates brought-back; however, in 1897 Harrison G. Dyar showed that, when compared to the caterpillars of G. rossii he had previously collected from the heights of Mount Washington, New Hampshire, in fact caterpillars of G. groenlandica had been collected in 1832, and Curtis had based his description of the larvae on the wrong species.[10][11]

Before that, however, specimens were recovered in 1870 from northern Greenland by Gottlieb August Wilhelm Herrich-Schäffer, on board the Germania on the Second German North Polar Expedition (led by captain Karl Koldewey). These were subsequently scientifically studied and described by Alexander von Homeyer in 1874 as Dasychira groenlandica, a name which Maximilian Ferdinand Wocke (who had also examined the specimens) had given them prior, detailing this in a letter to Homeyer. Herrich was able to collect a number of specimens, and described the caterpillars as being like those of the Arctia moths, with the adults being extremely similar to D. fascelina (now Dicallomera fascelina), but having such crippled wings as to be nearly incapable of flight. Both Wocke and Homeyer noted that, while it was certainly a new species, it was closely related to the species D. rossii (now G. rossii, known at that time from the geographic vicinity in Labrador).[1]

G. groenlandica was first believed to be endemic to the High Arctic,[12] until a 2013 article reported the discovery of two populations neighbouring each other in alpine environments within southwest Yukon, 900 km south of their previously known distribution. The distinct habitats, disjunct distribution, DNA haplotype and wing patterns of these two populations were found to be distinctive enough to classify as a new subspecies, G. groenlandica beringiana.[9] In the 1980s, moth specimens were collected on Wrangel Island, Russia; these were initially described in 1989 as a new species in the Dicallomera genus, D. kusnezovi. Further study showed the moths to be G. groenlandica, and, in 2015, the taxon was subsumed as a new subspecies, G. groenlandica kusnezovi.[2]

As such, the subspecies are:

  • G. groenlandica groenlandica
  • G. groenlandica beringiana Schmidt et Cannings, 2013[2][9]
  • G. groenlandica kusnezovi (Lukhtanov et Khruliova, 1989) Lukhtanovet Khruleva 2015[2]

It has been placed in the subfamily Lymantriinae (the tussock moths), tribe Orgyiini.[13][14]

While G. groenlandica is a close relative of G. rossii, the two species are reproductively isolated, thus no hybridisation is known to occur.[9] The two species are sympatric in many areas of the northern Canadian Arctic[9][10][11] and Wrangel Island.[2]

Description edit

In general, G. groenlandica larvae are larger (~300 mg), and are densely coated in soft-looking hairs, which may actually be used as a defense mechanism to irritate the skin and soft tissues of would-be predators.[9][15][16] While they are usually a distinctive tan-brown, amber cast, colors may vary.[9] They are characterized by a distinct hair tuft on their eighth abdominal segment, which has been described as a "rudimentary hair pencil".[7][9] Later larval instars are notable for the color pattern of this dorsal hair tuft.[9] They can also be identified by the spinulose form of their hairs, which are spineless, in contrast to the finer, feather-like (plumose) hairs of their close relative, G. rossii.[9][16] They may also be distinguished from G. rossii in terms of wing pattern: G. groenlandica lack the broad, dark band along the edge of their hind wings that is characteristic of G. rossii. In general, G. rossii also have more wing patterning than G. groenlandica.[9]

The eggs are around 1.6mm.[12]

The cocoons of this species are double-layered, with a distinct pocket of air between the two layers, as opposed to the single-layered cocoons of G. rossii.[12]

Distribution edit

The nominate subspecies of Arctic woolly bear moth is native to the High Arctic of Greenland and the Canadian Arctic Archipelago including Ellesmere Island, or above approximately 70°N latitude.[4][9][12][16] It is one of the most northern members of the Lepidopteran order in the Northern hemisphere.[7] It occurs as far north as Ward Hunt Island in Canada and northernmost Greenland.[16] A new subspecies G. groenlandica beringiana was described in 2013 south of the Arctic Circle in the alpine environment of the Ruby Range in southwest Yukon, 900 km south of the previously known range.[9] A further subspecies G. groenlandica kusnezovi has as of 2015 only been found on Wrangel Island, Russia.[2]

Habitat edit

G. groenlandica is well-adapted to living in conditions of extreme cold in the High Arctic.[7]

At two distinct field sites on Ellesmere Island, it was discovered that G. groenlandica, when in a diapausal state, tend to exist in specific microhabitats rather than in a random geographic distribution. Hibernacula are frequently found secured to the base of rocks, as opposed to being attached to vegetation. At one investigation site, hibernacula were observed primarily on the leeward (on the side sheltered from the wind) side of rocks, suggesting that wind direction plays a role in the selection of hibernation sites.[8][9]

In captivity, G. groenlandica have also been observed anchoring themselves to leaf litter of Salix arctica during the diapausal period.[8]

Range edit

The G. groenlandica caterpillar moves up to several meters per day, primarily in order to acquire the necessary resources.[17] In comparing a group of caterpillars physically transferred between different Salix arctica (Arctic willow) plants and a second group in which each individual was restricted to a single willow for the duration of the larval active period, it was observed that transferred larvae demonstrated higher herbivory and growth rates compared to the stationary group. This implies that the acquisition of high quality resources is a primary reason for the movement of G. groenlandica larvae between host plants.[17]

Food resources edit

 
Salix arctica, the Arctic willow, is the larva's primary host plant.

G. groenlandica spends much of its life in a larval state, and food resources are necessary for development of the larvae. Salix arctica, the Arctic willow, is the primary host plant and food source for this species.[6][17] The larvae may also feed on plants of other families, such as the flowers of Saxifraga oppositifolia and the senescent leaves of Dryas integrifolia. In the nominate High Arctic subspecies, less than 3% of larvae, however, were found to choose these alternatives.[4] The lower latitude Canadian populations of G. g. beringiana of the alpine environments of southwest Yukon have larvae eating a broader spectrum of plants and proportionately less S. arctica.[9][18]

While larvae rarely eat the catkins (petal-less flower clusters) of S. arctica, they readily consume the plant's leaves. 97% of larvae which actively eat at the onset of their feeding season are consuming the new leaf buds of this plant. Comparing the nutrient concentrations of plant leaves to those of larval frass, has shown that larvae remove nitrogen and potassium from the plant.[17] Larvae appear to only feed in June, which is when the leaves of S. arctica reach their peak concentrations of nutrients and carbohydrates such as starches and sugars. The caterpillars decrease their food intake towards the end of the month and into the summer. At this time, the levels of carbohydrates and nutrients in S. arctica leaves tend to decrease, and the leaves become less palatable as concentrations of phenols and tannins increase.[4][6][19] The decrease in nutrients and carbohydrates, combined with an increase in secondary metabolites, may account for this decline in consumption.[19]

It appears to be adapted to a narrow thermal range. It is able to eat the most at temperatures intermediate to its range.[20]

Life history edit

The life history traits of G. groenlandica are dictated by the short, cold nature of summers in the High Arctic.[18] Due to its restricted seasonal growth period, G. groenlandica has a life cycle of approximately 7 years.[5] In contrast, its lifespan is much shorter (2–3 years) in warmer, alpine environments.[9] Arctic woolly bear moths remain larvae for the vast majority of their lives, with the exception of up to 3–4 weeks of a single summer.[6] This extended developmental period is not only attributed to low environmental temperatures, but also to the nutrition provided by its host plants .[9] While they remain in their extended larval stage, G. groenlandica experience annual winter diapauses that commence in late June or early July.[7][8] Larval mortality in an experimental caged environment on the tundra was found to be 10%.[21]

Life cycle edit

On Ellesmere Island the females typically lay their eggs in a mass on or in their cocoon, although they sometimes lay their eggs on the ground or on vegetation around the cocoon.[12]

This species spends the vast majority of its life as a late larval instar; its early larval and adult stages represent only 6% of its full life cycle. It is the later instars which experience multiple annual periods of diapause.[19] During this dominant stage of their lives (from the third to sixth instar phases), G. greenlandica moult annually.[5]

Larval activity is confined to a short period following snowmelt. The High Arctic presents a short growing season of 45–70 days, and the G. groenlandica cease foraging at the end of June, prior to mid-summer.[4] Larvae tend to spend 95% of their time either basking in the sun, feeding, or moving, and only 5% of their time fully immobile. More specifically, about 60% of their time as larvae is spent basking, 20% is spent feeding, and 15% is spent moving.[19]

In late June or early July, the larvae prepare to overwinter by weaving silken hibernacula and entering diapause until the subsequent snowmelt.[7][8][9] This typically occurs when daytime temperatures are at a maximum of 5-10 °C. In their diapausal state, G. groenlandica can withstand temperatures as low as -70 °C, and winter mortality is limited to, on average, a maximum of 13% of the population.[7][19][22]

The developmental stages of pupation, emergence, mating, egg laying, eclosion, and molting to the second instar stage are all confined to a period of 3–4 weeks during a single summer. Emergence and reproduction may occur within a single 24-hour period.[4][9]

Due to the brief lifespan of fully mature adult individuals, adult moths of this species are difficult to find in the wild.[9]

Species interactions edit

The presence of the caterpillars eating plants in a particular area appears to have a positive correlation with herbivory of the collared pika (Ochotona collaris) in southwest Yukon.[15]

Predators edit

G. groenlandica has a distinct defence reaction to bat signals.[7] The Arctic moth Psychophora sabini has some of its defensive reactions to bats, presumably due to the population being isolated from this predator. G. groenlandica and G. rossii, however, continue to possess this defensive behavior. When Arctic woolly bear moths are exposed to bat-like ultrasound (26 kHz and 110 dB sound pressure level root mean square at 1 m), males respond by reversing their flight course. Responses to the sound have been observed from up to 15–25 m away. Females, however, have a degenerated bat-sensing system. There are two presumed reasons for this. Firstly, females tend to be flightless and thus do not require this adaptation. Secondly, an auditory system would compete for space with the ovaries, and the cost of this defence mechanism may outweigh the benefit of having fully functional reproductive organs.[23]

In the High Arctic these moths are preyed upon by birds. The eggs are also eaten by small foraging birds; on Ellesmere Island such birds may tear open the cocoons when feeding on the egg mass laid on the surface of the cocoon.[12]

Parasitoids edit

Many G. groenlandica caterpillars perish during development due to parasitoids, namely the tachinid fly (Exorista thula) and the ichneumonid wasp (Hyposoter diechmanni).[17][21][24] E. thula was described from Ellesmere Island in 2012, and is a gregarious parasitoid; on the island, it killed roughly 20% of the third and fourth instars of its host. Despite co-occurring there with the closely related Gynaephora rossii, E. thula is only known to attack G. groenlandica, whereas Chetogena gelida is host specific to G. rossii.[12] In general, more than two-thirds of Gynaephora are killed by parasitoids, and parasitism in G. groenlandica causes more than 50% mortality.[6][20] The probability of parasitism increases towards the end of the species' active period, which coincides with declining rates of feeding.

The hibernaculum, in which larvae spend a dominant portion of their lives, acts as a defensive barrier to parasitism.[21]

Physiology edit

Flight edit

While females of this species have fully developed wings and may take flight for a short time, they usually do not fly. Although Arctic-inhabiting females generally remain flightless, females of southerly alpine subspecies are often more mobile.[7][9]

In contrast, males tend to fly high, fast, and erratically during the day.[9]

Thermoregulation edit

The period of maximal activity for G. groenlandica is in June, during the annual period of maximal solar radiation (24 hours of sunlight) in the High Arctic; however, temperatures at this time continue to be extremely low. Ground temperatures in June, for instance, are usually less than 10 °C.[6][19] At this time, the body temperatures of feeding larvae tend to be similar to those of molting and spinning larvae, while those of “basking” larvae tend to be higher.[6] G. groenlandica larvae spend approximately 60% of their time basking, including during periods of pupation.[19] The behavior of basking is characterized as the action of a caterpillar orienting its body so as to maximize sun exposure and avoid wind. Larvae tend to follow the direct angle of the sun's rays in order to maintain maximal absorption of sunlight. They do this by orienting perpendicularly to the sun's angle of insolation.[6] Through the act of basking, G. groenlandica larvae may raise their body temperature by up to 20 °C. Generally, maximal body temperature is approximately 30 °C.[4][7][6] This peak temperature is generally only reached when larvae lie in midday sun, surrounded by snow, on a day with minimal wind.[6]

Solar radiation promotes larval growth, and thus basking may increase developmental rates.[4][6] When comparing larval growth rates at 5, 10, and 30 °C, respectively, growth and metabolic rates were found to be lowest at 5 °C and maximized at 30 °C.[4] This trend exhibits a specific relationship: as body temperature increases due to basking, metabolic rates increase exponentially. This was found to hold true even when larvae were starved or seemingly inactive.[4][6]

In general, feeding larvae tend to have lower body temperatures than basking larvae. Therefore, larvae tend to feed when temperatures are highest, and they bask when they cannot reach the higher temperatures (more than 5 -10 °C) needed for activity.[4] It has been suggested that without the help of basking in 24-hour sunlight during High Arctic summers, larvae would rarely exceed their developmental threshold of ~5 °C.[19] This may account for the unique tendency of the Arctic woolly bear moth to have short feeding periods during times of peak insolation, followed by lengthier periods of basking and digestion.[19]

In early to mid-June, larval metabolism tends to be greatly impacted by food intake and rising temperature. Later in the active season, they become much more metabolically insensitive to temperature, and energy obtained via food consumption is conserved.[25]

Changes in metabolic state and body temperature also affect oxygen consumption.[4][6] Oxygen consumption was found to be much lower when larval body temperatures were below 10 °C.[6] Low oxygen consumption was also observed in inactive larvae. In contrast, it was found to be higher for caterpillars that were moving or starved, higher still for digesting larvae, and highest for feeding larvae.[4]

Digestion edit

Larvae frequently bask in the sunlight for roughly five hours after feeding before moving to a new site.[4][6] The consequent increase in body temperature stimulates gut enzyme activity, which enables a higher digestion rate.[6] G. groenlandica can convert ingested food at a rate of efficiency which is higher than the average value rate of efficiency of Lepidopteran species in general.[4]

Diapause edit

G. groenlandica experiences a period of winter diapause during which it remains dormant within a hibernaculum. In this state, it can withstand temperatures as low as -70 °C.[7] Encasing itself within a hibernaculum during diapause serves several functions: protection from parasitoids, avoidance of diminished nutrient concentration in their primary food source, Salix arctica, degradation of mitochondria linked to decreased metabolism (hypometabolism) and antifreeze production, and general conservation of energy reserves.[6][7][12]

These cocoons are made of silk and consist of two layers, into which larval hairs are incorporated.[9][22] In a 1995 study of experimentally caged larvae in the High Arctic of the Canadian Archipelago, 81% of larvae spun hibernacula.[21]

During the active season, larvae orient towards solar radiation, and each spins its respective hibernaculum over a 24-hour period.[7][8] They generally pupate with their head facing south, in a north–south orientation.[6] This cocoon helps the larvae to accumulate heat more effectively.[7]

G. groenlandica often anchor their hibernacula to the base of rocks. In captivity, G. groenlandica have also been observed to attach themselves to Salix arctica leaf litter during the diapausal period.[8] In the 1995 study mentioned above, in which larvae were kept in a cage-controlled environment on the High Arctic tundra, more hibernacula were actually observed on the predominant plant cover of Dryas integrifolia (mountain avens) and Cassiope tetragona (Arctic white heather) as opposed to on their principal host plant, S. arctica.[21] Almost half of the larvae which spun hibernacula did so in conjunction with other larvae, forming joint cocoons. Upwards of three caterpillars were occasionally observed sharing a common hibernaculum, but the most common case was that of two individuals sharing a joint cocoon. Higher rates of communal hibernacula occurred at lower population densities per cage.[21]

As temperatures decrease in the late Arctic summer, larvae begin synthesizing cryoprotective compounds, such as glycerol and betaine. Accumulation of these "antifreezes" (which protect cells from cold conditions) is aided by the bottlenecking of oxidative phosphorylation through mitochondrial degradation. While the larvae continue to produce energy from stored glycogen in their frozen state, this mitochondrial degradation causes their metabolism to drop so low as to almost stop entirely, inducing dormancy. Mitochondrial functioning may be fully restored in the spring after mere hours of resumed larval activity.[7][26]

Conservation edit

At warmer temperatures, arctic moth larvae generally tend to have higher respiration rates and lower growth rates. They also tend to shift their diets to more nutrient-rich foods in this type of environment. For instance, the herbivory rate of the main food source for G. groenlandica, S. arctica, is altered at elevated temperatures.[9] This implies environmentally dependent host plant plasticity in G. groenlandica. It also suggests that an increase in temperature due to global warming may have significant effects on the behavior of northern herbivore invertebrates such as G. groenlandica, as well as effects on the herbivory rates of their food sources. Thus, G. groenlandica may represent a potential indicator species for future studies on climate change.[7][18][9]

In popular culture edit

This species was highlighted in BBC's sequel to Planet Earth, called Frozen Planet.[27]

See also edit

References edit

  1. ^ a b von Homeyer, Alexander (1874). "Lepidopteren". Wissenschaftliche Ergebnisse. Die zweite Deutsche Nordpolarfahrt (in German). Vol. 2. Leipzig: F. A. Brockhaus. pp. 409–410.
  2. ^ a b c d e f g Lukhtanov, Vladimir A.; Khruleva, Olga (2015). (PDF). Folia Biologica (Kraków). 63 (4): 257–261. doi:10.3409/fb63_4.257. ISSN 1734-9168. PMID 26975140. S2CID 4837579. Archived from the original (PDF) on 2019-02-27. Retrieved 22 November 2019.
  3. ^ Savela, Markku (2 June 2019). "Gynaephora". Lepidoptera and some other life forms. Markku Savela. Retrieved 21 November 2019.
  4. ^ a b c d e f g h i j k l m n o p q Kukal, Olga; Dawson, Todd E. (1989-06-01). "Temperature and food quality influences feeding behavior, assimilation efficiency and growth rate of arctic woolly-bear caterpillars". Oecologia. 79 (4): 526–532. Bibcode:1989Oecol..79..526K. doi:10.1007/BF00378671. PMID 28313488. S2CID 22994401.
  5. ^ a b c Morewood, W. Dean; Ring, Richard A. (1998-07-01). "Revision of the life history of the High Arctic moth Gynaephora groenlandica (Wocke) (Lepidoptera: Lymantriidae)". Canadian Journal of Zoology. 76 (7): 1371–1381. doi:10.1139/z98-085. ISSN 0008-4301.
  6. ^ a b c d e f g h i j k l m n o p q r s Kukal, Olga (March 24, 1988). "Behavioral Thermoregulation in the Freeze-Tolerant Arctic Caterpillar, Gynaephora groenlandica" (PDF). The Company of Biologists Limited.
  7. ^ a b c d e f g h i j k l m n o p q r Makarova, O. L.; Sviridov, A. V.; Klepikov, M. A. (2013-04-01). "Lepidoptera (Insecta) of polar deserts". Entomological Review. 93 (2): 225–239. doi:10.1134/S0013873813020115. S2CID 18862534.
  8. ^ a b c d e f g Bennett VA, Lee RE Jr, Nauman JS, Kukal O. Selection of overwintering microhabitats used by the arctic woollybear caterpillar, Gynaephora groenlandica. Cryo Letters. 2003 May-Jun;24(3):191-200.
  9. ^ a b c d e f g h i j k l m n o p q r s t u v w x y Barrio, Isabel C.; Schmidt, B. Christian; Cannings, Sydney; Hik, David S (December 2013). "First Records of the Arctic Moth Gynaephora groenlandica (Wocke) South of the Arctic". Arctic. 66 (4): 429–434. doi:10.14430/arctic4329. hdl:10261/142753.
  10. ^ a b Curtis, John (1835). "Insects". In Ross, John (ed.). Narrative of a Second Voyage in Search of a North-west Passage, and of a Residence in the Arctic Regions during the Years 1829, 1830, 1831, 1832, 1833 - Appendix. Vol. 2. London: A.W. Webster. p. lxx, lxxi. doi:10.5962/bhl.title.11196.
  11. ^ a b Dyar, Harrison G. (December 1897). "Note on Larvae of Gynaephora groenlandica and G. rossii" (PDF). Psyche: A Journal of Entomology. 8 (260): 153. doi:10.1155/1897/67619. Retrieved 24 November 2019.
  12. ^ a b c d e f g h Morewood, Dean W.; Wood, Monty D. (August 2002). "Host utilization by Exorista thula Wood (sp. nov.) and Chetogena gelida (Coquillett) (Diptera: Tachinidae), parasitoids of arctic Gynaephora species (Lepidoptera: Lymantriidae)". Polar Biology. 25 (8): 575–582. doi:10.1007/s00300-002-0382-y. S2CID 20312863.
  13. ^ "Gynaephora rossii Species Information". BugGuide. Retrieved 2018-03-29.
  14. ^ . Pacific Northwest Moths. Archived from the original on 3 December 2019. Retrieved 21 November 2019.
  15. ^ a b Barrio, Isabel C.; Hik, David S.; Peck, Kristen; Guillermo Bueno, C. (23 June 2013). "After the frass: foraging pikas select patches previously grazed by caterpillars". Biology Letters. 9 (3): 20130090. doi:10.1098/rsbl.2013.0090. PMC 3645042. PMID 23616644.
  16. ^ a b c d Morewood, Dean W.; Lange, Petra (1997). "Immature stages of high arctic Gynaephora species (Lymantriidae) and notes on their biology at Alexandra Fiord, Ellesmere Island, Canada". The Journal of Research on the Lepidoptera. 34 (1–4): 119–141. doi:10.5962/p.266564. S2CID 248740078. Retrieved 23 November 2019.
  17. ^ a b c d e Greyson-Gaito, Christopher J.; Barbour, Matthew A.; Rodriguez-Cabal, Mariano A.; Crutsinger, Gregory M.; Henry, Gregory H. R. (April 2016). "Freedom to move: Arctic caterpillar (Lepidoptera) growth rate increases with access to new willows (Salicaceae)". The Canadian Entomologist. 148 (6): 673–682. doi:10.4039/tce.2016.22. S2CID 87098612.
  18. ^ a b c Barrio, I. C.; Hik, D. S.; Liu, J. Y. (May 2014). "Diet breadth of Gynaephora groenlandica (Lepidoptera: Erebidae): is polyphagy greater in alpine versus Arctic populations?". The Canadian Entomologist. 147 (2): 215–221. doi:10.4039/tce.2014.35. S2CID 86183593.
  19. ^ a b c d e f g h i Lee, Richard (2012-12-06). Insects at Low Temperature. Springer Science & Business Media. ISBN 9781475701906.
  20. ^ a b Birkemoe, Tone; Bergmann, Saskia; Hasle, Toril E.; Klanderud, Kari (2016-10-01). "Experimental warming increases herbivory by leaf-chewing insects in an alpine plant community". Ecology and Evolution. 6 (19): 6955–6962. doi:10.1002/ece3.2398. PMC 5513215. PMID 28725372.
  21. ^ a b c d e f Kukal, Olga (1995-04-01). "Winter mortality and the function of larval hibernacula during the 14-year life cycle of an arctic moth, Gynaephora groenlandica". Canadian Journal of Zoology. 73 (4): 657–662. doi:10.1139/z95-077.
  22. ^ a b Laity, Peter R.; Holland, Chris (2017-02-01). "Thermo-rheological behaviour of native silk feedstocks". European Polymer Journal. 87 (Supplement C): 519–534. doi:10.1016/j.eurpolymj.2016.10.054.
  23. ^ Rydell, J.; Roininen, Heikki; Philip, Kenelm W. (22 March 2000). "Persistence of Bat Defence Reactions in High Arctic Moths (Lepidoptera)". Proceedings of the Royal Society B: Biological Sciences. 267 (1443): 553–557. doi:10.1098/rspb.2000.1036. PMC 1690572. PMID 10787157.
  24. ^ Varkonyi, Gergely; Tomas Roslin (2013). "Freezing cold yet diverse: dissecting a high-Arctic parasitoid community associated with Lepidoptera hosts". Canadian Entomologist. 145 (2): 193–218. doi:10.4039/tce.2013.9. S2CID 86014193.[permanent dead link]
  25. ^ Hodkinson, Ian D. (2005). "Adaptations of invertebrates to terrestrial Arctic environments". Transactions of the Royal Norwegian Society of Sciences and Letters: 28–29 – via ResearchGate.
  26. ^ Hoffmann, Klaus H. (2014-12-19). Insect Molecular Biology and Ecology. CRC Press. ISBN 9781482231892.
  27. ^ Levin, Gary (April 8, 2008). "Another sweeping nature special when 'Planet' freezes over". USA Today. Retrieved May 25, 2010.

Further reading edit

  • Heinrich, Bernd (1993). The hot-blooded insects: strategies and mechanisms of thermoregulation. Harvard University Press. ISBN 9780674408388.
  • Chapman, R. F. (1998). The insects: structure and function. Cambridge University Press. p. 519. ISBN 978-0-521-57890-5.
  • Kukal, O.; Kevan, P.G. (1987). "The influence of parasitism on the life history of a high arctic insect, Gynaephora groenlandica (Wöcke) (Lepidoptera: Lymantriidae)". Can. J. Zool. 65: 156–163. doi:10.1139/z87-022.
  • Kukal, O (1988). "Caterpillars on ice". Natural History. 97: 36–41.
  • Kukal, O.; Duman, J.G.; Serianni, A.S. (1988). "Glycerol metabolism in a freeze-tolerant arctic insect: An in vivo 13-C NMR study". J. Comp. Physiol. B. 158 (2): 175–183. doi:10.1007/bf01075831. PMID 3170824. S2CID 19371137.
  • Kukal, O.; Heinrich, B.; Duman, J.G. (1988). "Behavioural thermoregulation in the freeze-tolerant arctic caterpillar, Gynaephora groenlandica". J. Exp. Biol. 138: 181–193. doi:10.1242/jeb.138.1.181.
  • Kukal, O.; Dawson, T.E. (1989). "Temperature and food quality influences feeding behavior, assimilation efficiency and growth rate of arctic woolly-bear caterpillars". Oecologia. 79 (4): 526–532. Bibcode:1989Oecol..79..526K. doi:10.1007/bf00378671. PMID 28313488. S2CID 22994401.
  • Kukal, O.; Duman, J.G.; Serianni, A.S. (1989). "Cold-induced mitochondrial degradation and cryoprotectant synthesis in freeze-tolerant arctic caterpillars". J. Comp. Physiol. B. 158 (6): 661–671. doi:10.1007/bf00693004. PMID 2715455. S2CID 12109546.
  • Kukal, O. 1990. Energy budget for activity of a high arctic insect, Gynaephora groenlandica (Wöcke) (Lepidoptera: Lymantriidae). In: C.R. Harington (ed) Canadian Arctic Islands: Canada's Missing Dimension. National Museum of Natural History, Ottawa, Canada.
  • Kukal, O. 1991. Behavioral and physiological adaptations to cold in a freeze-tolerant arctic insect. In: R.E. Lee and D.L. Denlinger (eds) Insects at Low Temperature. Chapman and Hall, N.Y.
  • Kukal, O. 1993. Biotic and abiotic constraints on foraging of arctic caterpillars. In: N.E. Stamp and T.M. Casey (eds) Caterpillars: Ecological and Evolutionary Constraints on Foraging. Chapman and Hall, N.Y.
  • Kevan, P.G.; Kukal, O. (1993). "A balanced life table for Gynaephora groenlandica (Lepidoptera: Lymantriidae) a long-lived high arctic insect, and implications for the stability of its populations". Can. J. Zool. 65: 156–163.
  • Danks, H.V.; Kukal, O.; Ring, R.A. (1994). "Insect cold-hardiness: Insights from the arctic". Arctic. 47 (4): 391–404. doi:10.14430/arctic1312. S2CID 55711663.
  • Kukal, O (1995). "Winter mortality and the function of larval hibernacula during the 14-year life cycle of an arctic moth, Gynaephora groenlandica". Can. J. Zool. 73 (4): 657–662. doi:10.1139/z95-077.
  • Bennett, V. A; Kukal, O.; Lee, R.E. (1999). "Metabolic opportunists: Feeding and temperature influence the rate and pattern of respiration in the high arctic woollybear caterpillar, Gynaephora groenlandica (Lymantriidae)". J. Exp. Biol. 202 (1): 47–53. doi:10.1242/jeb.202.1.47. PMID 9841894.
  • Bennett, V.A.; Lee, R. E. Jr.; Nauman, J.S.; Kukal, O. (2003). "Selection of overwintering microhabitats used by the arctic woollybear caterpillar, Gynaephora groenlandica". CryoLetters. 24 (3): 191–200. PMID 12908029.

External links edit

    gynaephora, groenlandica, arctic, woolly, bear, moth, erebid, moth, native, high, arctic, canadian, archipelago, greenland, wrangel, island, russia, known, slow, rate, development, full, caterpillar, life, cycle, extend, years, with, moulting, occurring, each,. Gynaephora groenlandica the Arctic woolly bear moth is an erebid moth native to the High Arctic in the Canadian archipelago Greenland and Wrangel Island in Russia 2 4 It is known for its slow rate of development as its full caterpillar life cycle may extend up to 7 years with moulting occurring each spring 5 This species remains in a larval state for the vast majority of its life 4 6 Rare among Lepidoptera it undergoes an annual period of diapause that lasts for much of the calendar year as G groenlandica is subject to some of the longest most extreme winters on Earth 7 8 In this dormant state it can withstand temperatures as low as 70 C 7 The Arctic woolly bear moth also exhibits basking behavior which aids in temperature regulation and digestion and affects both metabolism and oxygen consumption 4 6 Females generally do not fly while males usually do 7 Arctic woolly bear moth Arctic woolly bear caterpillar Greenland Scientific classification Domain Eukaryota Kingdom Animalia Phylum Arthropoda Class Insecta Order Lepidoptera Superfamily Noctuoidea Family Erebidae Genus Gynaephora Species G groenlandica Binomial name Gynaephora groenlandica Wocke ex Homeyer 1874 Synonyms 3 Dasychira groenlandica Wocke ex Homeyer 1874 1 Dicallomera kusnezovi Lukhtanov et Khruliova 1989 2 This species has an alpine subspecies which is notable for its geographic distribution south of the High Arctic 9 Contents 1 Taxonomy 2 Description 3 Distribution 4 Habitat 4 1 Range 5 Food resources 6 Life history 6 1 Life cycle 7 Species interactions 7 1 Predators 7 2 Parasitoids 8 Physiology 8 1 Flight 8 2 Thermoregulation 8 3 Digestion 8 4 Diapause 9 Conservation 10 In popular culture 11 See also 12 References 13 Further reading 14 External linksTaxonomy editThis moth was likely first discovered on 16 June 1832 on the beach of Fury Bay Somerset Island northern Nunavut Canada by the crew of the Arctic expedition searching for the Northwest Passage led by John Ross John Curtis who studied the entomological specimens obtained from the voyage described Gynaephora rossii from the invertebrates brought back however in 1897 Harrison G Dyar showed that when compared to the caterpillars of G rossii he had previously collected from the heights of Mount Washington New Hampshire in fact caterpillars of G groenlandica had been collected in 1832 and Curtis had based his description of the larvae on the wrong species 10 11 Before that however specimens were recovered in 1870 from northern Greenland by Gottlieb August Wilhelm Herrich Schaffer on board the Germania on the Second German North Polar Expedition led by captain Karl Koldewey These were subsequently scientifically studied and described by Alexander von Homeyer in 1874 as Dasychira groenlandica a name which Maximilian Ferdinand Wocke who had also examined the specimens had given them prior detailing this in a letter to Homeyer Herrich was able to collect a number of specimens and described the caterpillars as being like those of the Arctia moths with the adults being extremely similar to D fascelina now Dicallomera fascelina but having such crippled wings as to be nearly incapable of flight Both Wocke and Homeyer noted that while it was certainly a new species it was closely related to the species D rossii now G rossii known at that time from the geographic vicinity in Labrador 1 G groenlandica was first believed to be endemic to the High Arctic 12 until a 2013 article reported the discovery of two populations neighbouring each other in alpine environments within southwest Yukon 900 km south of their previously known distribution The distinct habitats disjunct distribution DNA haplotype and wing patterns of these two populations were found to be distinctive enough to classify as a new subspecies G groenlandica beringiana 9 In the 1980s moth specimens were collected on Wrangel Island Russia these were initially described in 1989 as a new species in the Dicallomera genus D kusnezovi Further study showed the moths to be G groenlandica and in 2015 the taxon was subsumed as a new subspecies G groenlandica kusnezovi 2 As such the subspecies are G groenlandica groenlandica G groenlandica beringiana Schmidt et Cannings 2013 2 9 G groenlandica kusnezovi Lukhtanov et Khruliova 1989 Lukhtanovet Khruleva 2015 2 It has been placed in the subfamily Lymantriinae the tussock moths tribe Orgyiini 13 14 While G groenlandica is a close relative of G rossii the two species are reproductively isolated thus no hybridisation is known to occur 9 The two species are sympatric in many areas of the northern Canadian Arctic 9 10 11 and Wrangel Island 2 Description editIn general G groenlandica larvae are larger 300 mg and are densely coated in soft looking hairs which may actually be used as a defense mechanism to irritate the skin and soft tissues of would be predators 9 15 16 While they are usually a distinctive tan brown amber cast colors may vary 9 They are characterized by a distinct hair tuft on their eighth abdominal segment which has been described as a rudimentary hair pencil 7 9 Later larval instars are notable for the color pattern of this dorsal hair tuft 9 They can also be identified by the spinulose form of their hairs which are spineless in contrast to the finer feather like plumose hairs of their close relative G rossii 9 16 They may also be distinguished from G rossii in terms of wing pattern G groenlandica lack the broad dark band along the edge of their hind wings that is characteristic of G rossii In general G rossii also have more wing patterning than G groenlandica 9 The eggs are around 1 6mm 12 The cocoons of this species are double layered with a distinct pocket of air between the two layers as opposed to the single layered cocoons of G rossii 12 Distribution editThe nominate subspecies of Arctic woolly bear moth is native to the High Arctic of Greenland and the Canadian Arctic Archipelago including Ellesmere Island or above approximately 70 N latitude 4 9 12 16 It is one of the most northern members of the Lepidopteran order in the Northern hemisphere 7 It occurs as far north as Ward Hunt Island in Canada and northernmost Greenland 16 A new subspecies G groenlandica beringiana was described in 2013 south of the Arctic Circle in the alpine environment of the Ruby Range in southwest Yukon 900 km south of the previously known range 9 A further subspecies G groenlandica kusnezovi has as of 2015 only been found on Wrangel Island Russia 2 Habitat editG groenlandica is well adapted to living in conditions of extreme cold in the High Arctic 7 At two distinct field sites on Ellesmere Island it was discovered that G groenlandica when in a diapausal state tend to exist in specific microhabitats rather than in a random geographic distribution Hibernacula are frequently found secured to the base of rocks as opposed to being attached to vegetation At one investigation site hibernacula were observed primarily on the leeward on the side sheltered from the wind side of rocks suggesting that wind direction plays a role in the selection of hibernation sites 8 9 In captivity G groenlandica have also been observed anchoring themselves to leaf litter of Salix arctica during the diapausal period 8 Range edit The G groenlandica caterpillar moves up to several meters per day primarily in order to acquire the necessary resources 17 In comparing a group of caterpillars physically transferred between different Salix arctica Arctic willow plants and a second group in which each individual was restricted to a single willow for the duration of the larval active period it was observed that transferred larvae demonstrated higher herbivory and growth rates compared to the stationary group This implies that the acquisition of high quality resources is a primary reason for the movement of G groenlandica larvae between host plants 17 Food resources edit nbsp Salix arctica the Arctic willow is the larva s primary host plant G groenlandica spends much of its life in a larval state and food resources are necessary for development of the larvae Salix arctica the Arctic willow is the primary host plant and food source for this species 6 17 The larvae may also feed on plants of other families such as the flowers of Saxifraga oppositifolia and the senescent leaves of Dryas integrifolia In the nominate High Arctic subspecies less than 3 of larvae however were found to choose these alternatives 4 The lower latitude Canadian populations of G g beringiana of the alpine environments of southwest Yukon have larvae eating a broader spectrum of plants and proportionately less S arctica 9 18 While larvae rarely eat the catkins petal less flower clusters of S arctica they readily consume the plant s leaves 97 of larvae which actively eat at the onset of their feeding season are consuming the new leaf buds of this plant Comparing the nutrient concentrations of plant leaves to those of larval frass has shown that larvae remove nitrogen and potassium from the plant 17 Larvae appear to only feed in June which is when the leaves of S arctica reach their peak concentrations of nutrients and carbohydrates such as starches and sugars The caterpillars decrease their food intake towards the end of the month and into the summer At this time the levels of carbohydrates and nutrients in S arctica leaves tend to decrease and the leaves become less palatable as concentrations of phenols and tannins increase 4 6 19 The decrease in nutrients and carbohydrates combined with an increase in secondary metabolites may account for this decline in consumption 19 It appears to be adapted to a narrow thermal range It is able to eat the most at temperatures intermediate to its range 20 Life history editThe life history traits of G groenlandica are dictated by the short cold nature of summers in the High Arctic 18 Due to its restricted seasonal growth period G groenlandica has a life cycle of approximately 7 years 5 In contrast its lifespan is much shorter 2 3 years in warmer alpine environments 9 Arctic woolly bear moths remain larvae for the vast majority of their lives with the exception of up to 3 4 weeks of a single summer 6 This extended developmental period is not only attributed to low environmental temperatures but also to the nutrition provided by its host plants 9 While they remain in their extended larval stage G groenlandica experience annual winter diapauses that commence in late June or early July 7 8 Larval mortality in an experimental caged environment on the tundra was found to be 10 21 Life cycle edit On Ellesmere Island the females typically lay their eggs in a mass on or in their cocoon although they sometimes lay their eggs on the ground or on vegetation around the cocoon 12 This species spends the vast majority of its life as a late larval instar its early larval and adult stages represent only 6 of its full life cycle It is the later instars which experience multiple annual periods of diapause 19 During this dominant stage of their lives from the third to sixth instar phases G greenlandica moult annually 5 Larval activity is confined to a short period following snowmelt The High Arctic presents a short growing season of 45 70 days and the G groenlandica cease foraging at the end of June prior to mid summer 4 Larvae tend to spend 95 of their time either basking in the sun feeding or moving and only 5 of their time fully immobile More specifically about 60 of their time as larvae is spent basking 20 is spent feeding and 15 is spent moving 19 In late June or early July the larvae prepare to overwinter by weaving silken hibernacula and entering diapause until the subsequent snowmelt 7 8 9 This typically occurs when daytime temperatures are at a maximum of 5 10 C In their diapausal state G groenlandica can withstand temperatures as low as 70 C and winter mortality is limited to on average a maximum of 13 of the population 7 19 22 The developmental stages of pupation emergence mating egg laying eclosion and molting to the second instar stage are all confined to a period of 3 4 weeks during a single summer Emergence and reproduction may occur within a single 24 hour period 4 9 Due to the brief lifespan of fully mature adult individuals adult moths of this species are difficult to find in the wild 9 Species interactions editThe presence of the caterpillars eating plants in a particular area appears to have a positive correlation with herbivory of the collared pika Ochotona collaris in southwest Yukon 15 Predators edit G groenlandica has a distinct defence reaction to bat signals 7 The Arctic moth Psychophora sabini has some of its defensive reactions to bats presumably due to the population being isolated from this predator G groenlandica and G rossii however continue to possess this defensive behavior When Arctic woolly bear moths are exposed to bat like ultrasound 26 kHz and 110 dB sound pressure level root mean square at 1 m males respond by reversing their flight course Responses to the sound have been observed from up to 15 25 m away Females however have a degenerated bat sensing system There are two presumed reasons for this Firstly females tend to be flightless and thus do not require this adaptation Secondly an auditory system would compete for space with the ovaries and the cost of this defence mechanism may outweigh the benefit of having fully functional reproductive organs 23 In the High Arctic these moths are preyed upon by birds The eggs are also eaten by small foraging birds on Ellesmere Island such birds may tear open the cocoons when feeding on the egg mass laid on the surface of the cocoon 12 Parasitoids edit Many G groenlandica caterpillars perish during development due to parasitoids namely the tachinid fly Exorista thula and the ichneumonid wasp Hyposoter diechmanni 17 21 24 E thula was described from Ellesmere Island in 2012 and is a gregarious parasitoid on the island it killed roughly 20 of the third and fourth instars of its host Despite co occurring there with the closely related Gynaephora rossii E thula is only known to attack G groenlandica whereas Chetogena gelida is host specific to G rossii 12 In general more than two thirds of Gynaephora are killed by parasitoids and parasitism in G groenlandica causes more than 50 mortality 6 20 The probability of parasitism increases towards the end of the species active period which coincides with declining rates of feeding The hibernaculum in which larvae spend a dominant portion of their lives acts as a defensive barrier to parasitism 21 Physiology editFlight edit While females of this species have fully developed wings and may take flight for a short time they usually do not fly Although Arctic inhabiting females generally remain flightless females of southerly alpine subspecies are often more mobile 7 9 In contrast males tend to fly high fast and erratically during the day 9 Thermoregulation edit The period of maximal activity for G groenlandica is in June during the annual period of maximal solar radiation 24 hours of sunlight in the High Arctic however temperatures at this time continue to be extremely low Ground temperatures in June for instance are usually less than 10 C 6 19 At this time the body temperatures of feeding larvae tend to be similar to those of molting and spinning larvae while those of basking larvae tend to be higher 6 G groenlandica larvae spend approximately 60 of their time basking including during periods of pupation 19 The behavior of basking is characterized as the action of a caterpillar orienting its body so as to maximize sun exposure and avoid wind Larvae tend to follow the direct angle of the sun s rays in order to maintain maximal absorption of sunlight They do this by orienting perpendicularly to the sun s angle of insolation 6 Through the act of basking G groenlandica larvae may raise their body temperature by up to 20 C Generally maximal body temperature is approximately 30 C 4 7 6 This peak temperature is generally only reached when larvae lie in midday sun surrounded by snow on a day with minimal wind 6 Solar radiation promotes larval growth and thus basking may increase developmental rates 4 6 When comparing larval growth rates at 5 10 and 30 C respectively growth and metabolic rates were found to be lowest at 5 C and maximized at 30 C 4 This trend exhibits a specific relationship as body temperature increases due to basking metabolic rates increase exponentially This was found to hold true even when larvae were starved or seemingly inactive 4 6 In general feeding larvae tend to have lower body temperatures than basking larvae Therefore larvae tend to feed when temperatures are highest and they bask when they cannot reach the higher temperatures more than 5 10 C needed for activity 4 It has been suggested that without the help of basking in 24 hour sunlight during High Arctic summers larvae would rarely exceed their developmental threshold of 5 C 19 This may account for the unique tendency of the Arctic woolly bear moth to have short feeding periods during times of peak insolation followed by lengthier periods of basking and digestion 19 In early to mid June larval metabolism tends to be greatly impacted by food intake and rising temperature Later in the active season they become much more metabolically insensitive to temperature and energy obtained via food consumption is conserved 25 Changes in metabolic state and body temperature also affect oxygen consumption 4 6 Oxygen consumption was found to be much lower when larval body temperatures were below 10 C 6 Low oxygen consumption was also observed in inactive larvae In contrast it was found to be higher for caterpillars that were moving or starved higher still for digesting larvae and highest for feeding larvae 4 Digestion edit Larvae frequently bask in the sunlight for roughly five hours after feeding before moving to a new site 4 6 The consequent increase in body temperature stimulates gut enzyme activity which enables a higher digestion rate 6 G groenlandica can convert ingested food at a rate of efficiency which is higher than the average value rate of efficiency of Lepidopteran species in general 4 Diapause edit G groenlandica experiences a period of winter diapause during which it remains dormant within a hibernaculum In this state it can withstand temperatures as low as 70 C 7 Encasing itself within a hibernaculum during diapause serves several functions protection from parasitoids avoidance of diminished nutrient concentration in their primary food source Salix arctica degradation of mitochondria linked to decreased metabolism hypometabolism and antifreeze production and general conservation of energy reserves 6 7 12 These cocoons are made of silk and consist of two layers into which larval hairs are incorporated 9 22 In a 1995 study of experimentally caged larvae in the High Arctic of the Canadian Archipelago 81 of larvae spun hibernacula 21 During the active season larvae orient towards solar radiation and each spins its respective hibernaculum over a 24 hour period 7 8 They generally pupate with their head facing south in a north south orientation 6 This cocoon helps the larvae to accumulate heat more effectively 7 G groenlandica often anchor their hibernacula to the base of rocks In captivity G groenlandica have also been observed to attach themselves to Salix arctica leaf litter during the diapausal period 8 In the 1995 study mentioned above in which larvae were kept in a cage controlled environment on the High Arctic tundra more hibernacula were actually observed on the predominant plant cover of Dryas integrifolia mountain avens and Cassiope tetragona Arctic white heather as opposed to on their principal host plant S arctica 21 Almost half of the larvae which spun hibernacula did so in conjunction with other larvae forming joint cocoons Upwards of three caterpillars were occasionally observed sharing a common hibernaculum but the most common case was that of two individuals sharing a joint cocoon Higher rates of communal hibernacula occurred at lower population densities per cage 21 As temperatures decrease in the late Arctic summer larvae begin synthesizing cryoprotective compounds such as glycerol and betaine Accumulation of these antifreezes which protect cells from cold conditions is aided by the bottlenecking of oxidative phosphorylation through mitochondrial degradation While the larvae continue to produce energy from stored glycogen in their frozen state this mitochondrial degradation causes their metabolism to drop so low as to almost stop entirely inducing dormancy Mitochondrial functioning may be fully restored in the spring after mere hours of resumed larval activity 7 26 Conservation editAt warmer temperatures arctic moth larvae generally tend to have higher respiration rates and lower growth rates They also tend to shift their diets to more nutrient rich foods in this type of environment For instance the herbivory rate of the main food source for G groenlandica S arctica is altered at elevated temperatures 9 This implies environmentally dependent host plant plasticity in G groenlandica It also suggests that an increase in temperature due to global warming may have significant effects on the behavior of northern herbivore invertebrates such as G groenlandica as well as effects on the herbivory rates of their food sources Thus G groenlandica may represent a potential indicator species for future studies on climate change 7 18 9 In popular culture editThis species was highlighted in BBC s sequel to Planet Earth called Frozen Planet 27 See also editBelgica antarcticaReferences edit a b von Homeyer Alexander 1874 Lepidopteren Wissenschaftliche Ergebnisse Die zweite Deutsche Nordpolarfahrt in German Vol 2 Leipzig F A Brockhaus pp 409 410 a b c d e f g Lukhtanov Vladimir A Khruleva Olga 2015 Taxonomic Position and Status of Arctic Gynaephora and Dicallomera Moths Lepidoptera Erebidae Lymantriinae PDF Folia Biologica Krakow 63 4 257 261 doi 10 3409 fb63 4 257 ISSN 1734 9168 PMID 26975140 S2CID 4837579 Archived from the original PDF on 2019 02 27 Retrieved 22 November 2019 Savela Markku 2 June 2019 Gynaephora Lepidoptera and some other life forms Markku Savela Retrieved 21 November 2019 a b c d e f g h i j k l m n o p q Kukal Olga Dawson Todd E 1989 06 01 Temperature and food quality influences feeding behavior assimilation efficiency and growth rate of arctic woolly bear caterpillars Oecologia 79 4 526 532 Bibcode 1989Oecol 79 526K doi 10 1007 BF00378671 PMID 28313488 S2CID 22994401 a b c Morewood W Dean Ring Richard A 1998 07 01 Revision of the life history of the High Arctic moth Gynaephora groenlandica Wocke Lepidoptera Lymantriidae Canadian Journal of Zoology 76 7 1371 1381 doi 10 1139 z98 085 ISSN 0008 4301 a b c d e f g h i j k l m n o p q r s Kukal Olga March 24 1988 Behavioral Thermoregulation in the Freeze Tolerant Arctic Caterpillar Gynaephora groenlandica PDF The Company of Biologists Limited a b c d e f g h i j k l m n o p q r Makarova O L Sviridov A V Klepikov M A 2013 04 01 Lepidoptera Insecta of polar deserts Entomological Review 93 2 225 239 doi 10 1134 S0013873813020115 S2CID 18862534 a b c d e f g Bennett VA Lee RE Jr Nauman JS Kukal O Selection of overwintering microhabitats used by the arctic woollybear caterpillar Gynaephora groenlandica Cryo Letters 2003 May Jun 24 3 191 200 a b c d e f g h i j k l m n o p q r s t u v w x y Barrio Isabel C Schmidt B Christian Cannings Sydney Hik David S December 2013 First Records of the Arctic Moth Gynaephora groenlandica Wocke South of the Arctic Arctic 66 4 429 434 doi 10 14430 arctic4329 hdl 10261 142753 a b Curtis John 1835 Insects In Ross John ed Narrative of a Second Voyage in Search of a North west Passage and of a Residence in the Arctic Regions during the Years 1829 1830 1831 1832 1833 Appendix Vol 2 London A W Webster p lxx lxxi doi 10 5962 bhl title 11196 a b Dyar Harrison G December 1897 Note on Larvae of Gynaephora groenlandica and G rossii PDF Psyche A Journal of Entomology 8 260 153 doi 10 1155 1897 67619 Retrieved 24 November 2019 a b c d e f g h Morewood Dean W Wood Monty D August 2002 Host utilization by Exorista thula Wood sp nov and Chetogena gelida Coquillett Diptera Tachinidae parasitoids of arctic Gynaephora species Lepidoptera Lymantriidae Polar Biology 25 8 575 582 doi 10 1007 s00300 002 0382 y S2CID 20312863 Gynaephora rossii Species Information BugGuide Retrieved 2018 03 29 Gynaephora rossi Pacific Northwest Moths Archived from the original on 3 December 2019 Retrieved 21 November 2019 a b Barrio Isabel C Hik David S Peck Kristen Guillermo Bueno C 23 June 2013 After the frass foraging pikas select patches previously grazed by caterpillars Biology Letters 9 3 20130090 doi 10 1098 rsbl 2013 0090 PMC 3645042 PMID 23616644 a b c d Morewood Dean W Lange Petra 1997 Immature stages of high arctic Gynaephora species Lymantriidae and notes on their biology at Alexandra Fiord Ellesmere Island Canada The Journal of Research on the Lepidoptera 34 1 4 119 141 doi 10 5962 p 266564 S2CID 248740078 Retrieved 23 November 2019 a b c d e Greyson Gaito Christopher J Barbour Matthew A Rodriguez Cabal Mariano A Crutsinger Gregory M Henry Gregory H R April 2016 Freedom to move Arctic caterpillar Lepidoptera growth rate increases with access to new willows Salicaceae The Canadian Entomologist 148 6 673 682 doi 10 4039 tce 2016 22 S2CID 87098612 a b c Barrio I C Hik D S Liu J Y May 2014 Diet breadth of Gynaephora groenlandica Lepidoptera Erebidae is polyphagy greater in alpine versus Arctic populations The Canadian Entomologist 147 2 215 221 doi 10 4039 tce 2014 35 S2CID 86183593 a b c d e f g h i Lee Richard 2012 12 06 Insects at Low Temperature Springer Science amp Business Media ISBN 9781475701906 a b Birkemoe Tone Bergmann Saskia Hasle Toril E Klanderud Kari 2016 10 01 Experimental warming increases herbivory by leaf chewing insects in an alpine plant community Ecology and Evolution 6 19 6955 6962 doi 10 1002 ece3 2398 PMC 5513215 PMID 28725372 a b c d e f Kukal Olga 1995 04 01 Winter mortality and the function of larval hibernacula during the 14 year life cycle of an arctic moth Gynaephora groenlandica Canadian Journal of Zoology 73 4 657 662 doi 10 1139 z95 077 a b Laity Peter R Holland Chris 2017 02 01 Thermo rheological behaviour of native silk feedstocks European Polymer Journal 87 Supplement C 519 534 doi 10 1016 j eurpolymj 2016 10 054 Rydell J Roininen Heikki Philip Kenelm W 22 March 2000 Persistence of Bat Defence Reactions in High Arctic Moths Lepidoptera Proceedings of the Royal Society B Biological Sciences 267 1443 553 557 doi 10 1098 rspb 2000 1036 PMC 1690572 PMID 10787157 Varkonyi Gergely Tomas Roslin 2013 Freezing cold yet diverse dissecting a high Arctic parasitoid community associated with Lepidoptera hosts Canadian Entomologist 145 2 193 218 doi 10 4039 tce 2013 9 S2CID 86014193 permanent dead link Hodkinson Ian D 2005 Adaptations of invertebrates to terrestrial Arctic environments Transactions of the Royal Norwegian Society of Sciences and Letters 28 29 via ResearchGate Hoffmann Klaus H 2014 12 19 Insect Molecular Biology and Ecology CRC Press ISBN 9781482231892 Levin Gary April 8 2008 Another sweeping nature special when Planet freezes over USA Today Retrieved May 25 2010 Further reading editHeinrich Bernd 1993 The hot blooded insects strategies and mechanisms of thermoregulation Harvard University Press ISBN 9780674408388 Chapman R F 1998 The insects structure and function Cambridge University Press p 519 ISBN 978 0 521 57890 5 Kukal O Kevan P G 1987 The influence of parasitism on the life history of a high arctic insect Gynaephora groenlandica Wocke Lepidoptera Lymantriidae Can J Zool 65 156 163 doi 10 1139 z87 022 Kukal O 1988 Caterpillars on ice Natural History 97 36 41 Kukal O Duman J G Serianni A S 1988 Glycerol metabolism in a freeze tolerant arctic insect An in vivo 13 C NMR study J Comp Physiol B 158 2 175 183 doi 10 1007 bf01075831 PMID 3170824 S2CID 19371137 Kukal O Heinrich B Duman J G 1988 Behavioural thermoregulation in the freeze tolerant arctic caterpillar Gynaephora groenlandica J Exp Biol 138 181 193 doi 10 1242 jeb 138 1 181 Kukal O Dawson T E 1989 Temperature and food quality influences feeding behavior assimilation efficiency and growth rate of arctic woolly bear caterpillars Oecologia 79 4 526 532 Bibcode 1989Oecol 79 526K doi 10 1007 bf00378671 PMID 28313488 S2CID 22994401 Kukal O Duman J G Serianni A S 1989 Cold induced mitochondrial degradation and cryoprotectant synthesis in freeze tolerant arctic caterpillars J Comp Physiol B 158 6 661 671 doi 10 1007 bf00693004 PMID 2715455 S2CID 12109546 Kukal O 1990 Energy budget for activity of a high arctic insect Gynaephora groenlandica Wocke Lepidoptera Lymantriidae In C R Harington ed Canadian Arctic Islands Canada s Missing Dimension National Museum of Natural History Ottawa Canada Kukal O 1991 Behavioral and physiological adaptations to cold in a freeze tolerant arctic insect In R E Lee and D L Denlinger eds Insects at Low Temperature Chapman and Hall N Y Kukal O 1993 Biotic and abiotic constraints on foraging of arctic caterpillars In N E Stamp and T M Casey eds Caterpillars Ecological and Evolutionary Constraints on Foraging Chapman and Hall N Y Kevan P G Kukal O 1993 A balanced life table for Gynaephora groenlandica Lepidoptera Lymantriidae a long lived high arctic insect and implications for the stability of its populations Can J Zool 65 156 163 Danks H V Kukal O Ring R A 1994 Insect cold hardiness Insights from the arctic Arctic 47 4 391 404 doi 10 14430 arctic1312 S2CID 55711663 Kukal O 1995 Winter mortality and the function of larval hibernacula during the 14 year life cycle of an arctic moth Gynaephora groenlandica Can J Zool 73 4 657 662 doi 10 1139 z95 077 Bennett V A Kukal O Lee R E 1999 Metabolic opportunists Feeding and temperature influence the rate and pattern of respiration in the high arctic woollybear caterpillar Gynaephora groenlandica Lymantriidae J Exp Biol 202 1 47 53 doi 10 1242 jeb 202 1 47 PMID 9841894 Bennett V A Lee R E Jr Nauman J S Kukal O 2003 Selection of overwintering microhabitats used by the arctic woollybear caterpillar Gynaephora groenlandica CryoLetters 24 3 191 200 PMID 12908029 External links edit nbsp Wikimedia Commons has media related to Gynaephora groenlandica Discovery Channel video clip Retrieved from https en wikipedia org w index php title Gynaephora groenlandica amp oldid 1210490433, wikipedia, wiki, book, books, library,

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