Hibernaculum (zoology): Difference between revisions

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A hibernaculum plural form: hibernacula (Latin, "tent for winter quarters") is a place in which a creature seeks refuge, such as a bear using a cave to overwinter. The word can be used to describe a variety of shelters used by many kinds of animals, including insects, toads, lizards, snakes, bats, rodents, and primates of various classes.

An artificial hibernaculum or 'Bug Hotel'
Toad hibernaculum
Gray bats hibernating in a cave

Insects

Insects range in their size, structure, and general appearance but are all primarily exothermic^[1]. For this reason, extremely cold temperatures, such as those experiences in the winter season cause their metabolic systems to shut down and may lead to death. Insects survive winter through the process of overwintering, which occurs at all stages of development and may include migration or hibernation for different insects. Monarch butterflies are notorious for migrating to Mexico and California to wait out the winter^[2]. For other insects that do not migrate, they must halt their growth to avoid freezing to death, in a process called diapause ^[3]. Insects prepare to overwinter through a variety of mechanisms, such as using anti-freeze proteins or cryoprotectants in freeze-avoidant insects, such as the soybean aphids and inoculative freezing in freeze-tolerant insects that can survive being frozen, like second-generation corn borers ^[4]. Overwintering occurs in hibernacula to protect them from extreme weather and temperature changes and range in size and structure depending on the insects using them ^[5].

Lady Beetles

Some insects, like convergent lady beetles, reuse the same ones in groups, converging and migrating to those same hibernacula, used by prior generations, likely due to hydrocarbons previously left by their feet, causing them to retrace their footsteps to return to these locations ^[6]. Their tendency to aggregate and overwinter in groups is likely due to their attraction to similar environments and conspecifics. Beetles use rock crevices as hibernacula, often clumping in them in groups. These rock crevices are found in rock fields the beetle are attracted to for high levels of vegetation and greenery ^[7].

Other Insects

Other types of insect hibernacula include self-spun silk hibernacula, such as those made and used by Spruce budworms as they moult and overwinter in their second instars^[8]. Woolly bear caterpillars overwinter as caterpillars and grow to be Isabella tiger moths. They use plant debris as make-shift hibernacula, to protect themselves from extreme elements ^[9]. For freeze-avoidant insects, ideal hibernacula are dry, as freeze-avoidant insects are less likely to dampen and freeze in them, however moist hibernacula promote inoculative freezing for freeze-tolerant insects, which overwinter by freezing ^[10].

Amphibians

Amphibians that hibernate include several species of frogs and salamanders from the northern continental climates of North America and Eurasia and also from extreme southern hemisphere climates^[11]. These amphibians slow their metabolism during winter to avoid unsuitable conditions, such as freezing. Most freeze avoidance strategies include overwintering in aquatic situations or burrowing in the soil to depths below the frostline.^[12] A hibernaculum for amphibians should provide the following^[11]:

Optimum temperatures;
Maintenance of oxygen and humidity levels;
Low-intensity, short-photoperiod lighting;
Minimum disturbance
Frogs wintering at their hibernaculum at Gunnersbury Triangle in London

Species from cool continental climates hibernate at temperatures from 0 to 4°C. Some species will not survive hibernation at temperatures that exceed 4°C.^[11]

Generally, for amphibians that hibernate under ice, it is necessary for the animal to be submerged in water that is 10 to 15 cm deep and to maintain the temperature between 2 and 3°C and not above 4°C. Water should be well aerated, with maintained low-intensity light levels and minimal disturbance of the amphibians.

Frogs

Like other amphibians, frogs show minimal capacities for freezing tolerance and survive winter by using terrestrial hibernacula where they avoid freezing. However, frogs may exhibit greater freeze-tolerance capacity at high latitude range limits, where winter climate is more severe. For example, data suggests that while cricket frogs in South Dakota survive winter by locating hibernacula that prevent freezing, their toleration of short freezing bouts may expand the range of suitable hibernacula^[12]. However, overwinter mortality may be high at the northern range boundary due to colder temperatures and might limit cricket frogs from expanding their range northward^[12].

The microclimate refers to the the climate of a very small or restricted area, like the hibernaculum, especially when this differs from the climate of the surrounding area. Overwinter survival in these cricket frogs among other frogs is dependent on using hibernacula with appropriate physical microclimate characteristics, such as moist soil, that buffer frogs from temperatures that drop below the freezing point of the body fluids for extended periods.^[12] Although, determining if frogs can identify sites with appropriate microclimates to support overwinter survival and what factors might inform such choices are still unknown and will require further study. Therefore, it is still not known to what extent various types of prospective hibernacula for frogs might be suitable in the years to come, especially factoring in climate change.

Newts

As part the Highways Agency Biodiversity Action Plan (HABAP) in the UK, the Species Action Plan (SAP) for great crested newts aims to maintain and enhance existing newt populations through appropriate management of suitable habitat. As part of steps to implement the HABAP, newt hibernacula (e.g. log piles) have been constructed to improve the quality of the terrestrial habitat through increasing the number of potential overwintering sites^[13]. It was also determined that habitat surrounding breeding ponds with plenty of cover and suitable overwintering sites may have less need for provision of artificial hibernacula than landscapes with less woodland, hedgerows, scrub etc.^[13] Because great crested newts show high loyalty to over-wintering locations, returning to such established areas year after year, artificial hibernacula could be important in future years to conserve newts and other amphibians^[13]. Although, monitoring in the vicinity of these hibernacula in autumn using felt roofing tiles did not reveal the presence of any great crested newts even though they are know to breed in nearby ponds.^[13] Common toads and frogs did surround the area however. Therefore, further studies need to be conducted in order to create species-appropriate artifical hibernacula.

Reptiles

Many reptiles undergo hibernation or a process called brumation, which is similar to hibernation; both processes require usage of a hibernaculum. Staying inside an insulated hibernaculum is a strategy to avoid the harsh winter months when the frigid outside temperatures may kill an ectothermic reptile. They depress their metabolism and heart rates to reduce energy consumption so they don't need to exit their hibernacula. Hibernating reptiles are also safer from predation inside of their concealed and protected hibernacula. Various species of turtles, snakes, and lizards all use hibernacula, the forms of which can vary greatly.^[14]^[15]

Turtles

Common snapping turtles generally hibernate for about 6 months from early October to mid April. They live in lakes during their active months, then travel to small offshoot streams to hibernate. Hibernacula are about 100-150 meters away from the main body of the home lake. Most snapping turtles hibernate by burrowing into the banks of alder streams or vegetated streams, but some use other structures such as abandoned beaver dens. These streams are typically less than 0.3 m deep and 0.7 m wide, covered by sunken alder roots or fallen trees, and not covered by ice in the winter. Many animals return to the same stream to hibernate in subsequent years.^[14]

Snakes

Unlike more solitary snapping turtles, snakes may either hibernate alone or in large aggregations of up to several thousand individuals of the same or different species. They use a wide variety of hibernacula, including: rock piles, debris-filled wells, caves, crevices, unused burrows made by other animals, and ant mounds. The common European viper has actually been observed using all of the hibernacula listed above. Most species seem to prefer finding an already-present suitable site rather than constructing one of their own, but they do expand upon present structures and may make their own burrows if there aren't any quality sites available^[15]. Pine snakes and the closely related Louisiana pine snakes are two of the most well-studied hibernating snake species, and share similar hibernacula characteristics. These species sometimes constructs their own burrows, or use tunnels formed from the decay of tree roots or by gophers. The tunnels form complex networks, and have side chambers which each house one snake.^[15]^[16]

Lizards

Most initial discoveries of hibernating lizards have been accidental, but many species do in fact hibernate. Mesquite lizards in Mexico and the southern United States have been found hibernating in groups of 2-8 in cracks or under slabs of bark in mesquite trees.^[17] Common collared lizards spend about 6 months hibernating, almost always solitarily, though pairs of juvenile females have been observed within the same hibernacula. They use the undersides of rock slabs as hibernacula, digging a small chamber in the dirt just large enough for their bodies with a small tunnel for outside access. Adults use larger rock slabs, dig deeper chambers, and have longer tunnels than juveniles.^[18] Perhaps the most extreme example is seen in the viviparous lizard, the most northern of all lizards. They can burrow into the soil, go under leaf litter, or use shelters like rocks as hibernacula. Although the air temperature in West Siberia can drop to −10 °C, the soil temperature at the depths where these lizards hibernate remains higher than −10 °C. This enables them to survive the harshest temperatures of any lizard.^[19]

Mammals

Black bear mother and cubs hibernating, utilizing a hibernaculum as a maternity den

Like other animals, mammals hibernate during seasons of harsh environmental conditions and resource scarcity. As it requires less energy to maintain homeostasis and survive when an individual is hibernating, this is a cost-effective strategy to increase survival rates.^[20]^[21] Hibernation is usually perceived as taking place during winter, as in the most well-known hibernators bears and bats,^[22]^[21] but can also occur during the dry season when there is little food or water, as in the mouse lemurs of Madagascar.^[20] Given that mammals can spend anywhere from 1–9 months hibernating, their choice in hibernaculum in essential in determining their survival.^[21]

Bears

Many bears occupy similar hibernacula to smaller mammals, but theirs are, of course, much larger and can vary greatly across and within species. Most black bears excavate dens into a hillside or at the base of a tree, stump, or shrub, but some make dens at the bases of hollow trees, in hollow logs, or in rock caves or cavities. Den resuse is observed in this species, but very rarely. There were no significant den size differences between age or sex classes, except adult males creating larger entrances.^[23] Grizzly bears likewise don't show age or sex class differences in den dimensions. Grizzlies prefer hibernacula sites with abundant ground and canopy cover, and abundant sweet-vetch.^[21] Polar bears differ from black bears, grizzlies, and other bear species where both sexes hibernate in that only females use hibernacula. Like other female bears, polar bears use hibernacula as maternity dens. Similarly to other species, though, they tend to dig dens into the earth, although their arctic hibernacula are usually covered with snow by the time they emerge.^[24]

Bats

Bats favor larger hibernacula where large groups may roost together, including natural caves, mines, cellars, and other kinds of underground sites and man-made structures, like ice-houses.^[25] Within these hibernacula, the bats are still highly-tuned to environmental factors. Little brown bats in northern latitudes hibernate for up to 8 months during the winter, and leave their roosts in the warm spring weather when insect prey is plentiful again. Bats gauge the outside temperature by being attuned to the airflow at the hibernacula entrance, which is driven by temperature differences between inside and outside the hibernacula, allowing bats to leave when the temperature begins to warm.^[22] Some hibernacula are shared between multiple species, such as common pipistrelles roosting with soprano pipistrelles. Behavior other than hibernating can also occur at hibernacula; common pipistrelles produce most of their mating calls and mate with each other in and near their hibernacula.^[26]

Other Small Mammals

Many mammals that use hibernacula are small-bodied creatures, and include marsupials, rodents, and primates, among others. Most of these small-bodied mammals hibernate in simple holes in the ground, though some use complex systems of tunnels and burrows. Mountain pygmy possums in New South Wales, Australia awaken occasionally and leave their hibernacula for up to five days at a time. This occurs most often during the first few months of hibernation, with the possums leaving one hibernaculum in favor of another, seemingly in an effort to find the hibernaculum with the most suitable microclimate.^[27] The reddish-grey mouse lemur also wakes and leaves the hibernacula spontaenously and for brief periods of time.^[20] There can be hibernacula differences even within a species. In Columbian ground squirrels, hibernacula size is proportional to the weight of the individual occupying it, with adults having deeper hibernacula than juveniles, unlike black bears. Most juveniles choose to hibernate within 20 meters of their mother's burrow; those that don't have higher mortality rates.^[28]

References

^ Jung, M.-P., Kim, K.-H., Lee, S.-G. and Park, H.-H. (2013), Effect of climate change on the occurrence of overwintered moths of orchards in South Korea. Entomological Research, 43: 177–182. doi:10.1111/1748-5967.12016
^ Alerstam, T., et al. (2011). Convergent patterns of long-distance nocturnal migration in noctuid moths and passerine birds. Proc. Biol. Sci., 278: 3074–3080.
^ McMullen, David C. (March 2008). "Mitochondria of cold hardy insects: Responses to cold and hypoxia assessed at enzymatic, mRNA and DNA levels". Insect Biochemistry and Molecular Biology. 38 (3).
^ How insects survive cold weather. (2012, March 29). Western Farm Press. Retrieved from http://go.galegroup.com/ps/i.do?p=ITOF&sw=w&u=duke_perkins&v=2.1&it=r&id=GALE%7CA284721999&sid=summon&asid=b00124cf877e3cecb509af79899c4cd8
^ Layne, J. R., Jr., Edgar, C. L., & Medwith, R. E. (1999). Cold hardiness of the woolly bear caterpillar (Pyrrharctia isabella Lepidoptera: arctiidae). The American Midland Naturalist, 141(2), 293. Retrieved from http://go.galegroup.com/ps/i.do?p=ITOF&sw=w&u=duke_perkins&v=2.1&it=r&id=GALE%7CA54455057&sid=summon&asid=af754cb4b1f0f5c46bfae4c6c6de0514
^ Wheeler, C. A., & Cardé, R.,T. (2014). Following in their footprints: Cuticular hydrocarbons as overwintering aggregation site markers in hippodamia convergens. Journal of Chemical Ecology, 40(5): 418-28. doi:http://dx.doi.org/10.1007/s10886-014-0409-1
^ Honek, A., Martinková, Z., & Stano Pekár. (2007). Aggregation characteristics of three species of coccinellidae (coleoptera) at hibernation sites. European Journal of Entomology, 104(1): 51-56. Retrieved from http://search.proquest.com/docview/213976009
^ Volney, W. Jan A (17 November 2000). "Climate change and impacts of boreal forest insects". Agriculture, Ecosystems & Environment. 82 (1–3).
^ Layne, J. R., Jr., Edgar, C. L., & Medwith, R. E. (1999). Cold hardiness of the woolly bear caterpillar (Pyrrharctia isabella Lepidoptera: arctiidae). The American Midland Naturalist, 141(2): 293. Retrieved from http://go.galegroup.com/ps/i.do?p=ITOF&sw=w&u=duke_perkins&v=2.1&it=r&id=GALE%7CA54455057&sid=summon&asid=af754cb4b1f0f5c46bfae4c6c6de0514
^ Kawarasaki, Y., Teets, N. M., Denlinger, D. L., & Lee Jr, R.,E. (2014). Wet hibernacula promote inoculative freezing and limit the potential for cryoprotective dehydration in the antarctic midge, belgica antarctica. Polar Biology, 37(6): 753-761. doi:http://dx.doi.org/10.1007/s00300-014-1475-0
^ ^a ^b ^c Browne, Robert (2007). "Facility Design and Associated Services for the Study of Amphibians". ILAR. 48: 188–202.
^ ^a ^b ^c ^d Burdick, Seth L.; Swanson, David L. (2010). "Overwintering Physiology and Hibernacula Microclimates of Blanchard's Cricket Frogs at Their Northwestern Range Boundary". Copeia. 2: 247–253.
^ ^a ^b ^c ^d Knowles, Mark; Latham, Dorian (2008). "Assessing the use of artificial hibernacula by great crested newts Triturus cristatus and other amphibians for habitat enhancement, Northumberland, England". Conservation Evidence. 5: 74–79.
^ ^a ^b Strain, Gabriel F.; Anderson, James T.; Michael, Edwin D.; Turk, Philip J. (2012-01-01). "Hibernacula Use and Hibernation Phenology in the Common Snapping Turtle (Chelydra serpentina) in Canaan Valley, West Virginia". Journal of Herpetology. 46 (2): 269–274.
^ ^a ^b ^c Burger, Joanna; Zappalorti, Robert T.; Gochfeld, Michael; Boarman, William I.; Caffrey, Michael; Doig, Victor; Garber, Steven D.; Lauro, Brook; Mikovsky, Maria (1988-01-01). "Hibernacula and Summer Den Sites of Pine Snakes (Pituophis melanoleucus) in the New Jersey Pine Barrens". Journal of Herpetology. 22 (4): 425–433. doi:10.2307/1564337.
^ D.C., Rudolph,; R.R., Schaefer,; S.J., Burgdorf,; M., Duran,; R.N., Conner, (2007-01-01). "Pine snake (Pituophis ruthveni and Pituophis mellanoleucus lodingi) hibernacula". Journal of Herpetology. 41(4): 560-565.{{cite journal}}: CS1 maint: extra punctuation (link) CS1 maint: multiple names: authors list (link)
^ Axtell, Ralph W.; Axtell, Carol A. (1970-01-01). "Hibernacula, Birth and Young of Sceloporus grammicus disparilis (Iguanidae)". The Southwestern Naturalist. 14 (3): 363–366. doi:10.2307/3668971.
^ Legler, John M.; Fitch, Henry S. (1957-01-01). "Observations on Hibernation and Nests of the Collared Lizard, Crotaphytus collaris". Copeia. 1957 (4): 305–307. doi:10.2307/1439168.
^ Berman, Daniil I.; Bulakhova, Nina A.; Alfimov, Arcady V.; Meshcheryakova, Ekaterina N. (2016-12-01). "How the most northern lizard, Zootoca vivipara, overwinters in Siberia". Polar Biology. 39 (12): 2411–2425. doi:10.1007/s00300-016-1916-z. ISSN 0722-4060.
^ ^a ^b ^c Kobbe S. & Dausmann K. H. (2009). Hibernation in Malagasy mouse lemurs as a strategy to counter environmental challenge. Naturwissenschaften, 96(10):1221-1227. Jump up^
^ ^a ^b ^c ^d Pigeon, Karine E.; Côté, Steeve D.; Stenhouse, Gordon B. (2016-07-01). "Assessing den selection and den characteristics of grizzly bears". The Journal of Wildlife Management. 80 (5): 884–893. doi:10.1002/jwmg.1069. ISSN 1937-2817.
^ ^a ^b Meyer, Gretchen A.; Senulis, Joseph A.; Reinartz, James A. (2016-12-05). "Effects of temperature and availability of insect prey on bat emergence from hibernation in spring". Journal of Mammalogy. 97 (6): 1623–1633. doi:10.1093/jmammal/gyw126. ISSN 0022-2372.
^ Beecham, John J.; Reynolds, Doyle G.; Hornocker, Maurice G. (1983-01-01). "Black Bear Denning Activities and Den Characteristics in West-Central Idaho". Bears: Their Biology and Management. 5: 79–86. doi:10.2307/3872522.
^ Ramsay, Malcolm A.; Stirling, Ian (1990-05-21). "Fidelity of Female Polar Bears to Winter-Den Sites". Journal of Mammalogy. 71 (2): 233–236. doi:10.2307/1382172. ISSN 0022-2372.
^ Van der Meij, Thomas; Van Strien, A. J.; Haysom, K. A.; Dekker, J.; Russ, J.; Biala, K.; Bihari, Z.; Jansen, E.; Langton, S. (2015-05-01). "Return of the bats? A prototype indicator of trends in European bat populations in underground hibernacula". Mammalian Biology - Zeitschrift für Säugetierkunde. Special Issue: Bats as Bioindicators. 80 (3): 170–177. doi:10.1016/j.mambio.2014.09.004.
^ Bartoničková, Lenka; Reiter, Antonín; Bartonička, Tomáš (2016-11-22). "Mating and Courtship Behaviour of Two Sibling Bat Species (Pipistrellus pipistrellus, P. pygmaeus) in the Vicinity of a Hibernaculum". Acta Chiropterologica. 18 (2): 467–475. doi:10.3161/15081109ACC2016.18.2.013. ISSN 1508-1109.
^ Körtner, Gerhard; Geiser, Fritz (1998-01-01). "Ecology of natural hibernation in the marsupial mountain pygmy-possum (Burramys parvus)". Oecologia. 113 (2): 170–178. doi:10.1007/s004420050365. ISSN 0029-8549.
^ Young, Paul J. (1990-01-01). "Structure, Location and Availability of Hibernacula of Columbian Ground Squirrels (Spermophilus columbianus)". The American Midland Naturalist. 123 (2): 357–364. doi:10.2307/2426563.

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[1] Jung, M.-P., Kim, K.-H., Lee, S.-G. and Park, H.-H. (2013), Effect of climate change on the occurrence of overwintered moths of orchards in South Korea. Entomological Research, 43: 177–182. doi:10.1111/1748-5967.12016

[2] Alerstam, T., et al. (2011). Convergent patterns of long-distance nocturnal migration in noctuid moths and passerine birds. Proc. Biol. Sci., 278: 3074–3080.

[3] McMullen, David C. (March 2008). "Mitochondria of cold hardy insects: Responses to cold and hypoxia assessed at enzymatic, mRNA and DNA levels". Insect Biochemistry and Molecular Biology. 38 (3).

[4] How insects survive cold weather. (2012, March 29). Western Farm Press. Retrieved from http://go.galegroup.com/ps/i.do?p=ITOF&sw=w&u=duke_perkins&v=2.1&it=r&id=GALE%7CA284721999&sid=summon&asid=b00124cf877e3cecb509af79899c4cd8

[5] Layne, J. R., Jr., Edgar, C. L., & Medwith, R. E. (1999). Cold hardiness of the woolly bear caterpillar (Pyrrharctia isabella Lepidoptera: arctiidae). The American Midland Naturalist, 141(2), 293. Retrieved from http://go.galegroup.com/ps/i.do?p=ITOF&sw=w&u=duke_perkins&v=2.1&it=r&id=GALE%7CA54455057&sid=summon&asid=af754cb4b1f0f5c46bfae4c6c6de0514

[6] Wheeler, C. A., & Cardé, R.,T. (2014). Following in their footprints: Cuticular hydrocarbons as overwintering aggregation site markers in hippodamia convergens. Journal of Chemical Ecology, 40(5): 418-28. doi:http://dx.doi.org/10.1007/s10886-014-0409-1

[7] Honek, A., Martinková, Z., & Stano Pekár. (2007). Aggregation characteristics of three species of coccinellidae (coleoptera) at hibernation sites. European Journal of Entomology, 104(1): 51-56. Retrieved from http://search.proquest.com/docview/213976009

[8] Volney, W. Jan A (17 November 2000). "Climate change and impacts of boreal forest insects". Agriculture, Ecosystems & Environment. 82 (1–3).

[9] Layne, J. R., Jr., Edgar, C. L., & Medwith, R. E. (1999). Cold hardiness of the woolly bear caterpillar (Pyrrharctia isabella Lepidoptera: arctiidae). The American Midland Naturalist, 141(2): 293. Retrieved from http://go.galegroup.com/ps/i.do?p=ITOF&sw=w&u=duke_perkins&v=2.1&it=r&id=GALE%7CA54455057&sid=summon&asid=af754cb4b1f0f5c46bfae4c6c6de0514

[10] Kawarasaki, Y., Teets, N. M., Denlinger, D. L., & Lee Jr, R.,E. (2014). Wet hibernacula promote inoculative freezing and limit the potential for cryoprotective dehydration in the antarctic midge, belgica antarctica. Polar Biology, 37(6): 753-761. doi:http://dx.doi.org/10.1007/s00300-014-1475-0

[:52-11] Browne, Robert (2007). "Facility Design and Associated Services for the Study of Amphibians". ILAR. 48: 188–202.

[:62-12] Burdick, Seth L.; Swanson, David L. (2010). "Overwintering Physiology and Hibernacula Microclimates of Blanchard's Cricket Frogs at Their Northwestern Range Boundary". Copeia. 2: 247–253.

[:72-13] Knowles, Mark; Latham, Dorian (2008). "Assessing the use of artificial hibernacula by great crested newts Triturus cristatus and other amphibians for habitat enhancement, Northumberland, England". Conservation Evidence. 5: 74–79.

[:3-14] Strain, Gabriel F.; Anderson, James T.; Michael, Edwin D.; Turk, Philip J. (2012-01-01). "Hibernacula Use and Hibernation Phenology in the Common Snapping Turtle (Chelydra serpentina) in Canaan Valley, West Virginia". Journal of Herpetology. 46 (2): 269–274.

[:4-15] Burger, Joanna; Zappalorti, Robert T.; Gochfeld, Michael; Boarman, William I.; Caffrey, Michael; Doig, Victor; Garber, Steven D.; Lauro, Brook; Mikovsky, Maria (1988-01-01). "Hibernacula and Summer Den Sites of Pine Snakes (Pituophis melanoleucus) in the New Jersey Pine Barrens". Journal of Herpetology. 22 (4): 425–433. doi:10.2307/1564337.

[16] D.C., Rudolph,; R.R., Schaefer,; S.J., Burgdorf,; M., Duran,; R.N., Conner, (2007-01-01). "Pine snake (Pituophis ruthveni and Pituophis mellanoleucus lodingi) hibernacula". Journal of Herpetology. 41(4): 560-565.{{cite journal}}: CS1 maint: extra punctuation (link) CS1 maint: multiple names: authors list (link)

[17] Axtell, Ralph W.; Axtell, Carol A. (1970-01-01). "Hibernacula, Birth and Young of Sceloporus grammicus disparilis (Iguanidae)". The Southwestern Naturalist. 14 (3): 363–366. doi:10.2307/3668971.

[18] Legler, John M.; Fitch, Henry S. (1957-01-01). "Observations on Hibernation and Nests of the Collared Lizard, Crotaphytus collaris". Copeia. 1957 (4): 305–307. doi:10.2307/1439168.

[19] Berman, Daniil I.; Bulakhova, Nina A.; Alfimov, Arcady V.; Meshcheryakova, Ekaterina N. (2016-12-01). "How the most northern lizard, Zootoca vivipara, overwinters in Siberia". Polar Biology. 39 (12): 2411–2425. doi:10.1007/s00300-016-1916-z. ISSN 0722-4060.

[:0-20] Kobbe S. & Dausmann K. H. (2009). Hibernation in Malagasy mouse lemurs as a strategy to counter environmental challenge. Naturwissenschaften, 96(10):1221-1227. Jump up^

[:1-21] Pigeon, Karine E.; Côté, Steeve D.; Stenhouse, Gordon B. (2016-07-01). "Assessing den selection and den characteristics of grizzly bears". The Journal of Wildlife Management. 80 (5): 884–893. doi:10.1002/jwmg.1069. ISSN 1937-2817.

[:2-22] Meyer, Gretchen A.; Senulis, Joseph A.; Reinartz, James A. (2016-12-05). "Effects of temperature and availability of insect prey on bat emergence from hibernation in spring". Journal of Mammalogy. 97 (6): 1623–1633. doi:10.1093/jmammal/gyw126. ISSN 0022-2372.

[23] Beecham, John J.; Reynolds, Doyle G.; Hornocker, Maurice G. (1983-01-01). "Black Bear Denning Activities and Den Characteristics in West-Central Idaho". Bears: Their Biology and Management. 5: 79–86. doi:10.2307/3872522.

[24] Ramsay, Malcolm A.; Stirling, Ian (1990-05-21). "Fidelity of Female Polar Bears to Winter-Den Sites". Journal of Mammalogy. 71 (2): 233–236. doi:10.2307/1382172. ISSN 0022-2372.

[25] Van der Meij, Thomas; Van Strien, A. J.; Haysom, K. A.; Dekker, J.; Russ, J.; Biala, K.; Bihari, Z.; Jansen, E.; Langton, S. (2015-05-01). "Return of the bats? A prototype indicator of trends in European bat populations in underground hibernacula". Mammalian Biology - Zeitschrift für Säugetierkunde. Special Issue: Bats as Bioindicators. 80 (3): 170–177. doi:10.1016/j.mambio.2014.09.004.

[26] Bartoničková, Lenka; Reiter, Antonín; Bartonička, Tomáš (2016-11-22). "Mating and Courtship Behaviour of Two Sibling Bat Species (Pipistrellus pipistrellus, P. pygmaeus) in the Vicinity of a Hibernaculum". Acta Chiropterologica. 18 (2): 467–475. doi:10.3161/15081109ACC2016.18.2.013. ISSN 1508-1109.

[27] Körtner, Gerhard; Geiser, Fritz (1998-01-01). "Ecology of natural hibernation in the marsupial mountain pygmy-possum (Burramys parvus)". Oecologia. 113 (2): 170–178. doi:10.1007/s004420050365. ISSN 0029-8549.

[28] Young, Paul J. (1990-01-01). "Structure, Location and Availability of Hibernacula of Columbian Ground Squirrels (Spermophilus columbianus)". The American Midland Naturalist. 123 (2): 357–364. doi:10.2307/2426563.

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 == Reptiles ==
-Many reptiles undergo hibernation or a process called [[Dormancy|brumation]], which is similar to hibernation; both processes require usage of a hibernacula. Staying inside an insulated hibernacula is a strategy to avoid the harsh winter months when the frigid outside temperatures may kill an [[Ectotherm|ectothermic]] reptile. They depress their metabolism and heart rates to reduce energy consumption so they don't need to exit their hibernacula. Hibernating reptiles are also safer from predation inside of their concealed and protected hibernacula. Various species of turtles, snakes, and lizards all use hibernacula, the forms of which can vary greatly.<ref name=":3">{{Cite journal|last=Strain|first=Gabriel F.|last2=Anderson|first2=James T.|last3=Michael|first3=Edwin D.|last4=Turk|first4=Philip J.|date=2012-01-01|title=Hibernacula Use and Hibernation Phenology in the Common Snapping Turtle (Chelydra serpentina) in Canaan Valley, West Virginia|url=http://www.jstor.org/stable/41515049|journal=Journal of Herpetology|volume=46|issue=2|pages=269–274}}</ref><ref name=":4">{{Cite journal|last=Burger|first=Joanna|last2=Zappalorti|first2=Robert T.|last3=Gochfeld|first3=Michael|last4=Boarman|first4=William I.|last5=Caffrey|first5=Michael|last6=Doig|first6=Victor|last7=Garber|first7=Steven D.|last8=Lauro|first8=Brook|last9=Mikovsky|first9=Maria|date=1988-01-01|title=Hibernacula and Summer Den Sites of Pine Snakes (Pituophis melanoleucus) in the New Jersey Pine Barrens|url=http://www.jstor.org/stable/1564337|journal=Journal of Herpetology|volume=22|issue=4|pages=425–433|doi=10.2307/1564337}}</ref>[[File:Snapping turtle in Mud.JPG|thumb|A snapping turtle burrowed into a stream bank|223x223px]]
+Many reptiles undergo hibernation or a process called [[Dormancy|brumation]], which is similar to hibernation; both processes require usage of a hibernaculum. Staying inside an insulated hibernaculum is a strategy to avoid the harsh winter months when the frigid outside temperatures may kill an [[Ectotherm|ectothermic]] reptile. They depress their metabolism and heart rates to reduce energy consumption so they don't need to exit their hibernacula. Hibernating reptiles are also safer from predation inside of their concealed and protected hibernacula. Various species of turtles, snakes, and lizards all use hibernacula, the forms of which can vary greatly.<ref name=":3">{{Cite journal|last=Strain|first=Gabriel F.|last2=Anderson|first2=James T.|last3=Michael|first3=Edwin D.|last4=Turk|first4=Philip J.|date=2012-01-01|title=Hibernacula Use and Hibernation Phenology in the Common Snapping Turtle (Chelydra serpentina) in Canaan Valley, West Virginia|url=http://www.jstor.org/stable/41515049|journal=Journal of Herpetology|volume=46|issue=2|pages=269–274}}</ref><ref name=":4">{{Cite journal|last=Burger|first=Joanna|last2=Zappalorti|first2=Robert T.|last3=Gochfeld|first3=Michael|last4=Boarman|first4=William I.|last5=Caffrey|first5=Michael|last6=Doig|first6=Victor|last7=Garber|first7=Steven D.|last8=Lauro|first8=Brook|last9=Mikovsky|first9=Maria|date=1988-01-01|title=Hibernacula and Summer Den Sites of Pine Snakes (Pituophis melanoleucus) in the New Jersey Pine Barrens|url=http://www.jstor.org/stable/1564337|journal=Journal of Herpetology|volume=22|issue=4|pages=425–433|doi=10.2307/1564337}}</ref>[[File:Snapping turtle in Mud.JPG|thumb|A snapping turtle burrowed into a stream bank|223x223px]]
 === Turtles ===
 [[Common snapping turtle|Common snapping turtles]] generally hibernate for about 6 months from early October to mid April. They live in lakes during their active months, then travel to small offshoot streams to hibernate. Hibernacula are about 100-150 meters away from the main body of the home lake. Most snapping turtles hibernate by burrowing into the banks of [[alder]] streams or vegetated streams, but some use other structures such as abandoned beaver dens. These streams are typically less than 0.3 m deep and 0.7 m wide, covered by sunken alder roots or fallen trees, and not covered by ice in the winter. Many animals return to the same stream to hibernate in subsequent years.<ref name=":3" />
 === Snakes ===
 [[File:Snake Hole.jpg|thumb|An outside view of a snake hibernacula]]
-Unlike more solitary snapping turtles, snakes may either hibernate alone or in large aggregations of up to several thousand individuals of the same or different species. They use a wide variety of hibernacula, including: rock piles, debris-filled wells, caves, crevices, unused burrows made by other animals, and ant mounds. The [[Vipera berus|common European viper]] has actually been observed using all of the hibernacula listed above. Most species seem to prefer finding an already-present suitable site rather than constructing one of their own, but they do expand upon present structures and may make their own burrows if there aren't any quality sites available. [[Pituophis melanoleucus|Pine snakes]] and the closely related [[Louisiana pinesnake|Louisiana pine snakes]] are two of the most well-studied hibernating snake species, and share similar hibernacula characteristics. These species sometimes constructs their own burrows, or use tunnels formed from the decay of tree roots or by gophers. The tunnels form complex networks, and have side chambers which each house one snake.<ref name=":4" /><ref>{{Cite journal|last=D.C.|first=Rudolph,|last2=R.R.|first2=Schaefer,|last3=S.J.|first3=Burgdorf,|last4=M.|first4=Duran,|last5=R.N.|first5=Conner,|date=2007-01-01|title=Pine snake (Pituophis ruthveni and Pituophis mellanoleucus lodingi) hibernacula|url=https://www.srs.fs.usda.gov/pubs/31517|journal=Journal of Herpetology. 41(4): 560-565|language=en}}</ref>
+Unlike more solitary snapping turtles, snakes may either hibernate alone or in large aggregations of up to several thousand individuals of the same or different species. They use a wide variety of hibernacula, including: rock piles, debris-filled wells, caves, crevices, unused burrows made by other animals, and ant mounds. The [[Vipera berus|common European viper]] has actually been observed using all of the hibernacula listed above. Most species seem to prefer finding an already-present suitable site rather than constructing one of their own, but they do expand upon present structures and may make their own burrows if there aren't any quality sites available<ref name=":4" />. [[Pituophis melanoleucus|Pine snakes]] and the closely related [[Louisiana pinesnake|Louisiana pine snakes]] are two of the most well-studied hibernating snake species, and share similar hibernacula characteristics. These species sometimes constructs their own burrows, or use tunnels formed from the decay of tree roots or by gophers. The tunnels form complex networks, and have side chambers which each house one snake.<ref name=":4" /><ref>{{Cite journal|last=D.C.|first=Rudolph,|last2=R.R.|first2=Schaefer,|last3=S.J.|first3=Burgdorf,|last4=M.|first4=Duran,|last5=R.N.|first5=Conner,|date=2007-01-01|title=Pine snake (Pituophis ruthveni and Pituophis mellanoleucus lodingi) hibernacula|url=https://www.srs.fs.usda.gov/pubs/31517|journal=Journal of Herpetology. 41(4): 560-565|language=en}}</ref>
 === Lizards ===
 Most initial discoveries of hibernating lizards have been accidental, but many species do in fact hibernate. [[Sceloporus grammicus|Mesquite lizards]] in Mexico and the southern United States have been found hibernating in groups of 2-8 in cracks or under slabs of bark in [[Mesquite|mesquite trees]].<ref>{{Cite journal|last=Axtell|first=Ralph W.|last2=Axtell|first2=Carol A.|date=1970-01-01|title=Hibernacula, Birth and Young of Sceloporus grammicus disparilis (Iguanidae)|url=http://www.jstor.org/stable/3668971|journal=The Southwestern Naturalist|volume=14|issue=3|pages=363–366|doi=10.2307/3668971}}</ref> [[Common collared lizard|Common collared lizards]] spend about 6 months hibernating, almost always solitarily, though pairs of juvenile females have been observed within the same hibernacula. They use the undersides of rock slabs as hibernacula, digging a small chamber in the dirt just large enough for their bodies with a small tunnel for outside access. Adults use larger rock slabs, dig deeper chambers, and have longer tunnels than juveniles.<ref>{{Cite journal|last=Legler|first=John M.|last2=Fitch|first2=Henry S.|date=1957-01-01|title=Observations on Hibernation and Nests of the Collared Lizard, Crotaphytus collaris|url=http://www.jstor.org/stable/1439168|journal=Copeia|volume=1957|issue=4|pages=305–307|doi=10.2307/1439168}}</ref> Perhaps the most extreme example is seen in the [[viviparous lizard]], the most northern of all lizards. They can burrow into the soil, go under leaf litter, or use shelters like rocks as hibernacula. Although the air temperature in West Siberia can drop to −10 °C, the soil temperature at the depths where these lizards hibernate remains higher than −10 °C. This enables them to survive the harshest temperatures of any lizard.<ref>{{Cite journal|last=Berman|first=Daniil I.|last2=Bulakhova|first2=Nina A.|last3=Alfimov|first3=Arcady V.|last4=Meshcheryakova|first4=Ekaterina N.|date=2016-12-01|title=How the most northern lizard, Zootoca vivipara, overwinters in Siberia|url=https://link.springer.com/article/10.1007/s00300-016-1916-z|journal=Polar Biology|language=en|volume=39|issue=12|pages=2411–2425|doi=10.1007/s00300-016-1916-z|issn=0722-4060}}</ref>
 == Mammals ==
+[[File:Bear hibernating.jpg|thumb|Black bear mother and cubs hibernating, utilizing a hibernaculum as a maternity den|200x200px]]Like other animals, mammals hibernate during seasons of harsh environmental conditions and resource scarcity. As it requires less energy to maintain [[homeostasis]] and survive when an individual is hibernating, this is a cost-effective strategy to increase survival rates.<ref name=":0">Kobbe S. & Dausmann K. H. (2009). Hibernation in Malagasy mouse lemurs as a strategy to counter environmental challenge. Naturwissenschaften, 96(10):1221-1227.
-[[File:Columbian Ground Squirrel Roger's Pass.jpg|thumb|Columbian ground squirrel outside its hibernaculum|201x201px]]
-Like other animals, mammals hibernate during seasons of harsh environmental conditions and resource scarcity. As it requires less energy to maintain [[homeostasis]] and survive when an individual is hibernating, this is a cost-effective strategy to increase survival rates.<ref name=":0">Kobbe S. & Dausmann K. H. (2009). Hibernation in Malagasy mouse lemurs as a strategy to counter environmental challenge. Naturwissenschaften, 96(10):1221-1227.
 '''[[Talk:Hibernaculum (zoology)#cite ref-12|Jump up^]]'''
 </ref><ref name=":1">{{Cite journal|last=Pigeon|first=Karine E.|last2=Côté|first2=Steeve D.|last3=Stenhouse|first3=Gordon B.|date=2016-07-01|title=Assessing den selection and den characteristics of grizzly bears|url=http://onlinelibrary.wiley.com/doi/10.1002/jwmg.1069/abstract|journal=The Journal of Wildlife Management|language=en|volume=80|issue=5|pages=884–893|doi=10.1002/jwmg.1069|issn=1937-2817}}</ref> Hibernation is usually perceived as taking place during winter, as in the most well-known hibernators bears and bats,<ref name=":2">{{Cite journal|last=Meyer|first=Gretchen A.|last2=Senulis|first2=Joseph A.|last3=Reinartz|first3=James A.|date=2016-12-05|title=Effects of temperature and availability of insect prey on bat emergence from hibernation in spring|url=https://academic.oup.com/jmammal/article-lookup/doi/10.1093/jmammal/gyw126|journal=Journal of Mammalogy|language=en|volume=97|issue=6|pages=1623–1633|doi=10.1093/jmammal/gyw126|issn=0022-2372}}</ref><ref name=":1" /> but can also occur during the dry season when there is little food or water, as in the mouse lemurs of Madagascar.<ref name=":0" /> Given that mammals can spend anywhere from 1–9 months hibernating, their choice in hibernaculum in essential in determining their survival.<ref name=":1" />
-=== Small Mammals ===
-[[File:Bear hibernating.jpg|thumb|Black bear mother and cubs hibernating, utilizing a hibernaculum as a maternity den|200x200px]]Many mammals that use hibernacula are small-bodied creatures, and include marsupials, rodents, and primates, among others. Most of these small-bodied mammals hibernate in simple holes in the ground, though some use complex systems of tunnels and burrows. [[Mountain pygmy possum|Mountain pygmy possums]] in New South Wales, Australia awaken occasionally and leave their hibernacula for up to five days at a time. This occurs most often during the first few months of hibernation, with the possums leaving one hibernacula in favor of another, seemingly in an effort to find the hibernacula with the most suitable microclimate.<ref>{{Cite journal|last=Körtner|first=Gerhard|last2=Geiser|first2=Fritz|date=1998-01-01|title=Ecology of natural hibernation in the marsupial mountain pygmy-possum (Burramys parvus)|url=https://link.springer.com/article/10.1007/s004420050365|journal=Oecologia|language=en|volume=113|issue=2|pages=170–178|doi=10.1007/s004420050365|issn=0029-8549}}</ref> The [[Reddish-gray mouse lemur|reddish-grey mouse lemur]] also wakes and leaves the hibernacula spontaenously and for brief periods of time.<ref name=":0" /> There can be hibernacula differences even within a species. In [[Columbian ground squirrel|Columbian ground squirrels]], hibernacula size is proportional to the weight of the individual occupying it, with adults having deeper hibernacula than juveniles. Most juveniles choose to hibernate within 20 meters of their mother's burrow; those that don't have higher mortality rates.<ref>{{Cite journal|last=Young|first=Paul J.|date=1990-01-01|title=Structure, Location and Availability of Hibernacula of Columbian Ground Squirrels (Spermophilus columbianus)|url=http://www.jstor.org/stable/2426563|journal=The American Midland Naturalist|volume=123|issue=2|pages=357–364|doi=10.2307/2426563}}</ref>
 === Bears ===
-Many bears occupy similar hibernacula to smaller mammals, but theirs are, of course, much larger and can vary greatly across and within species. Most [[American black bear|black bears]] excavate dens into a hillside or at the base of a tree, stump, or shrub, but some make dens at the bases of hollow trees, in hollow logs, or in rock caves or cavities. Den resuse is observed in this species, but very rarely. Unlike Columbian ground squirrels, there were no significant den size differences between age or sex classes, except adult males creating larger entrances.<ref>{{Cite journal|last=Beecham|first=John J.|last2=Reynolds|first2=Doyle G.|last3=Hornocker|first3=Maurice G.|date=1983-01-01|title=Black Bear Denning Activities and Den Characteristics in West-Central Idaho|url=http://www.jstor.org/stable/3872522|journal=Bears: Their Biology and Management|volume=5|pages=79–86|doi=10.2307/3872522}}</ref> [[Grizzly bear|Grizzly bears]] likewise don't show age or sex class differences in den dimensions. Grizzlies prefer hibernacula sites with abundant ground and canopy cover, and abundant [[Hedysarum|sweet-vetch]].<ref name=":1" /> [[Polar bear|Polar bears]] differ from black bears, grizzlies, and other bear species in that only females use hibernacula, which function as [[Maternity den|maternity dens]]. Similarly to other species, though, they tend to dig dens into the earth, although their arctic hibernacula are usually covered with snow by the time they emerge.<ref>{{Cite journal|last=Ramsay|first=Malcolm A.|last2=Stirling|first2=Ian|date=1990-05-21|title=Fidelity of Female Polar Bears to Winter-Den Sites|url=https://academic.oup.com/jmammal/article-abstract/71/2/233/835345/Fidelity-of-Female-Polar-Bears-to-Winter-Den-Sites|journal=Journal of Mammalogy|volume=71|issue=2|pages=233–236|doi=10.2307/1382172|issn=0022-2372}}</ref>
+[[File:Indiana bats (5600274705).jpg|thumb|203x203px|[[Indiana bat|Indiana bats]] hibernating in a cave]]Many bears occupy similar hibernacula to smaller mammals, but theirs are, of course, much larger and can vary greatly across and within species. Most [[American black bear|black bears]] excavate dens into a hillside or at the base of a tree, stump, or shrub, but some make dens at the bases of hollow trees, in hollow logs, or in rock caves or cavities. Den resuse is observed in this species, but very rarely. There were no significant den size differences between age or sex classes, except adult males creating larger entrances.<ref>{{Cite journal|last=Beecham|first=John J.|last2=Reynolds|first2=Doyle G.|last3=Hornocker|first3=Maurice G.|date=1983-01-01|title=Black Bear Denning Activities and Den Characteristics in West-Central Idaho|url=http://www.jstor.org/stable/3872522|journal=Bears: Their Biology and Management|volume=5|pages=79–86|doi=10.2307/3872522}}</ref> [[Grizzly bear|Grizzly bears]] likewise don't show age or sex class differences in den dimensions. Grizzlies prefer hibernacula sites with abundant ground and canopy cover, and abundant [[Hedysarum|sweet-vetch]].<ref name=":1" /> [[Polar bear|Polar bears]] differ from black bears, grizzlies, and other bear species where both sexes hibernate in that only females use hibernacula. Like other female bears, polar bears use hibernacula as [[Maternity den|maternity dens]]. Similarly to other species, though, they tend to dig dens into the earth, although their arctic hibernacula are usually covered with snow by the time they emerge.<ref>{{Cite journal|last=Ramsay|first=Malcolm A.|last2=Stirling|first2=Ian|date=1990-05-21|title=Fidelity of Female Polar Bears to Winter-Den Sites|url=https://academic.oup.com/jmammal/article-abstract/71/2/233/835345/Fidelity-of-Female-Polar-Bears-to-Winter-Den-Sites|journal=Journal of Mammalogy|volume=71|issue=2|pages=233–236|doi=10.2307/1382172|issn=0022-2372}}</ref>
-[[File:Indiana bats (5600274705).jpg|thumb|203x203px|[[Indiana bat|Indiana bats]] hibernating in a cave]]
 === Bats ===
+[[File:Columbian Ground Squirrel Roger's Pass.jpg|thumb|Columbian ground squirrel outside its hibernaculum|201x201px]]
 Bats favor larger hibernacula where large groups may roost together, including natural caves, mines, cellars, and other kinds of underground sites and man-made structures, like ice-houses.<ref>{{Cite journal|last=Van der Meij|first=Thomas|last2=Van Strien|first2=A. J.|last3=Haysom|first3=K. A.|last4=Dekker|first4=J.|last5=Russ|first5=J.|last6=Biala|first6=K.|last7=Bihari|first7=Z.|last8=Jansen|first8=E.|last9=Langton|first9=S.|date=2015-05-01|title=Return of the bats? A prototype indicator of trends in European bat populations in underground hibernacula|url=http://www.sciencedirect.com/science/article/pii/S1616504714001098|journal=Mammalian Biology - Zeitschrift für Säugetierkunde|series=Special Issue: Bats as Bioindicators|volume=80|issue=3|pages=170–177|doi=10.1016/j.mambio.2014.09.004}}</ref> Within these hibernacula, the bats are still highly-tuned to environmental factors. [[Little brown bat|Little brown bats]] in northern latitudes hibernate for up to 8 months during the winter, and leave their roosts in the warm spring weather when insect prey is plentiful again. Bats gauge the outside temperature by being attuned to the airflow at the hibernacula entrance, which is driven by temperature differences between inside and outside the hibernacula, allowing bats to leave when the temperature begins to warm.<ref name=":2" /> Some hibernacula are shared between multiple species, such as [[Common pipistrelle|common pipistrelles]] roosting with [[Soprano pipistrelle|soprano pipistrelles]]. Behavior other than hibernating can also occur at hibernacula; common pipistrelles produce most of their mating calls and mate with each other in and near their hibernacula.<ref>{{Cite journal|last=Bartoničková|first=Lenka|last2=Reiter|first2=Antonín|last3=Bartonička|first3=Tomáš|date=2016-11-22|title=Mating and Courtship Behaviour of Two Sibling Bat Species (Pipistrellus pipistrellus, P. pygmaeus) in the Vicinity of a Hibernaculum|url=http://www.bioone.org/doi/10.3161/15081109ACC2016.18.2.013|journal=Acta Chiropterologica|volume=18|issue=2|pages=467–475|doi=10.3161/15081109ACC2016.18.2.013|issn=1508-1109}}</ref>
+=== Other Small Mammals ===
+Many mammals that use hibernacula are small-bodied creatures, and include marsupials, rodents, and primates, among others. Most of these small-bodied mammals hibernate in simple holes in the ground, though some use complex systems of tunnels and burrows. [[Mountain pygmy possum|Mountain pygmy possums]] in New South Wales, Australia awaken occasionally and leave their hibernacula for up to five days at a time. This occurs most often during the first few months of hibernation, with the possums leaving one hibernaculum in favor of another, seemingly in an effort to find the hibernaculum with the most suitable microclimate.<ref>{{Cite journal|last=Körtner|first=Gerhard|last2=Geiser|first2=Fritz|date=1998-01-01|title=Ecology of natural hibernation in the marsupial mountain pygmy-possum (Burramys parvus)|url=https://link.springer.com/article/10.1007/s004420050365|journal=Oecologia|language=en|volume=113|issue=2|pages=170–178|doi=10.1007/s004420050365|issn=0029-8549}}</ref> The [[Reddish-gray mouse lemur|reddish-grey mouse lemur]] also wakes and leaves the hibernacula spontaenously and for brief periods of time.<ref name=":0" /> There can be hibernacula differences even within a species. In [[Columbian ground squirrel|Columbian ground squirrels]], hibernacula size is proportional to the weight of the individual occupying it, with adults having deeper hibernacula than juveniles, unlike black bears. Most juveniles choose to hibernate within 20 meters of their mother's burrow; those that don't have higher mortality rates.<ref>{{Cite journal|last=Young|first=Paul J.|date=1990-01-01|title=Structure, Location and Availability of Hibernacula of Columbian Ground Squirrels (Spermophilus columbianus)|url=http://www.jstor.org/stable/2426563|journal=The American Midland Naturalist|volume=123|issue=2|pages=357–364|doi=10.2307/2426563}}</ref>
 == See also ==