Geographic Range
Arctic ground squirrels
are the northernmost species of ground squirrel. They occupy a Holarctic distribution
ranging from the west coast of Hudson Bay to the west cost of Alaska in North America,
and into eastern Siberia (Donker and Krebs 2011, Barker and Derocher 2010, Yensen
and Sherman 2003, Karels et al. 2000, Buck and Barnes 1999a). Recent phylogeographic
studies have identified four geographically divergent clades. The Arctic clade consists
of arctic and subarctic populations that occur north of the Yukon River in northern
Alaska and Canada. The Southeast clade includes populations south of the Yukon River,
spreading into south-central Alaska and western Canada. The Beringia clade inhabits
the Alaskan Seward Peninsula and St. Lawrence Island, stretching across the Bering
Strait into Siberia. The Southwest clade occurs south of the Yukon River in Alaska,
continuing west out onto the Alaska Peninsula and associated islands (Galbreath et
al. 2011).
Most populations of arctic ground squirrels are indigenous, though a recent study
has shown that many island populations near the Alaskan peninsula may be introduced,
resulting from the presumed introduction by early indigenous peoples who valued them
for their pelts. Likely introduced populations include several islands in the Kodiak
archipelago, Semidi islands, and Shumagin islands, as well as confirmed introduced
populations on Kavalga, Unalaska, and Amaknak islands in the Aleutians (Cook et al.
2010).
Range map at: http://mapservices.iucnredlist.org/IUCN/mapper/index.html?ID_NO=20488
- Biogeographic Regions
- nearctic
- Other Geographic Terms
- holarctic
Habitat
Arctic ground squirrels inhabit arctic and tundra habitats at northern latitudes and
higher elevations, and boreal forest and alpine meadows at lower latitudes and elevations
(Byrom et al. 2000, Donker and Krebs 2011). At their northernmost ranges, they contend
with long cold winters where temperatures can drop below -20° Celsius for seven to
eight months of the year, and where sub zero temperatures and snowfall are possible
in every month of the year. In arctic environments, the growing season is short,
lasting only six to ten weeks, and for most of the year arctic ground squirrels must
cope with frozen soils, high winds, snow accumulation of 10 to 75 cm, and extremes
of light and darkness (Buck and Barnes 1999a,b).
Arctic ground squirrels are colonial and live in shallow, subterranean burrow systems
up to 20 m in length (Nowak 1999). In many populations, burrow depth is limited to
less than 1 meter due to frozen soils (permafrost) (Buck and Barnes 1999b). Burrows
serve as maternal dens, hibernacula, and refuges from predators and climatic conditions
(Long et al. 2005, Barker and Derocher 2010). There are at least three distinct burrow
types constructed and used by arctic ground squirrels. Duck holes are short tunnels
with several exits that are used primarily as escape refuges. Permanently occupied
single and double burrow systems consist of many multilevel tunnels, nest chambers,
and exits. Double burrow systems are used by closely related breeding females as
shared places to raise young. Hibernacula burrows are used during hibernation and
can either be connected to existing burrow systems, or constructed separately elsewhere.
These often have hidden entrances that are plugged with earth upon immergence for
hibernation in the fall, and are not cleared until the animal emerges in the spring
(Iwen 1999). Within the burrows they construct nests of lichens and dry grasses,
which are 22.5 to 30 cm in diameter (Buck and Barnes 1999b). Habitats with sloped
terrains, drained soils, and sparse vegetation, are preferred by arctic ground squirrels.
Sites with wet soils are avoided due to their poor suitability for digging, and sites
with hummocks or abundant tall or shrubby vegetation are selected against since they
reduce the arctic ground squirrels ability to visually detect predators (Gillis et
al. 2005b, Barker and Derocher 2010).
Populations of arctic ground squirrels are at their highest densities in arctic tundra
and alpine environments, where they are mostly limited by food and burrow availability.
In boreal forest populations, densities are lower, and they are limited mainly by
a combination of predation and food supply. Boreal forests are considered suboptimal
habitat due to an increased number of predators, and the reduced ability of the squirrels
to detect predators. Not surprisingly, these areas have been shown to be a population
sink for the species (Karels et al. 2000, Gillis et al. 2005b, Donker and Krebs 2011).
- Habitat Regions
- polar
- terrestrial
Physical Description
All arctic ground squirrels have a cylindrical body shape with short, powerful forearms
and sharp claws adapted for digging, and short, strong hind limbs for propelling forward
movement. They have four digits on their forefeet and five on their hind feet. Their
feet have soft pads on the bottom surface that allow them to grip and manipulate food
and earth (Nowak 1999, Yensen and Sherman 2003). Their dental formula is: I 1/1,
C 0/0, P 2/1, M 3/3, and P3 is one-third to one-half the size of P4 (Hall 1981, Forsyth
1999). Arctic ground squirrels molt twice per year; in the spring after emergence
from hibernation, and in the fall prior to hibernation. They have tawny or cinnamon
dorsal pelage that is flecked with white spots. Their underparts are lighter in color,
largely buff tawny in the summer and turning an even lighter buffy color with the
fall molt. Their tail color matches their pelage at the base, fading toward darker
colors at the tip (Hall 1981, Iwen 1999). The range of length for their skull, tail,
and hind foot is 50.7 to 65.8 mm, 77 to 153 mm, and 50 to 68 mm, respectively (Yensen
and Sherman 2003). They also have large internal membranous cheek pouches that are
used to store and carry food. Females have four to six pairs of mammae (Nowak 1999).
Arctic ground squirrels are the largest of the ground squirrels with a mass of 524
to 1500 g, depending on sex and season. They range from 332 to 495 mm in length,
with an average length of 390 mm (Woods 1980, Yensen and Sherman 2003). They are sexually
dimorphic in size, and males average 740 to 1000 g and are longer in length, while
females average 600 to 1000 g and are shorter in length (Buck and Barnes 1999b, Yensen
and Sherman 2003, Hayssen 2008). They also have an enhanced ability to store fat,
and for five to seven weeks preceding hibernation they will increase their fat stores
to 30 to 41.5% of their total body weight (Frank et al. 2008).
Arctic ground squirrels are the northernmost hibernating terrestrial mammal and can
achieve wide ranges of body temperatures and metabolic rates. This plasticity is
an adaptation for surviving extended seasons of extreme temperatures and no food availability
(Sheriff et al. 2011). Their upper critical body temperature is 36° Celsius, their
lower critical body temperature is 18° Celsius, and their basal metabolic rate is
0.40 to 0.61 ml O2/g/h (Withers et al. 1979, Long et al. 2005). During hibernation
they can drop their body temperature to -2.9° Celsius. They also have the lowest
known minimum metabolic rate of any endothermic hibernator, reaching levels of 0.012
ml O2/g/h during torpor at ambient temperatures of 4° Celsius (Buck and Barnes 1999b,
2000).
- Other Physical Features
- endothermic
- heterothermic
- homoiothermic
- bilateral symmetry
- Sexual Dimorphism
- male larger
Reproduction
Arctic ground squirrels exhibit a female defense polygynous mating system (Yensen
and Sherman 2003). Mating occurs in late April shortly after emergence from hibernation
and lasts for approximately two weeks. Females are at maximal estrous three days
following emergence and they are only receptive for a short, less than 12-hour period
(Buck and Barnes 1999a, Lacy and Wieczorek 2001, Yensen and Sherman 2003, Donker and
Krebs 2011).
Males typically defend a territory where multiple females reside. During this time,
males are highly territorial and will act aggressively to all other males that enter
their territory. Severe confrontations can occur where males will severely wound
or even kill each other in defense of females (Lacy and Wieczorek 2001, Buck and Barnes
2003). Males will mate with multiple females, and extra-pair copulations between
females and multiple males also occur. Evidence of copulatory plugs in females has
been observed. The first male a female mates with is generally the one that sires
the most young in her litter (over 90%), though multiple paternity in litters is common,
and females have been observed to mate with up to four males (Lacy et al. 1997, Lacy
and Wieczorek 2001, Yensen and Sherman 2003).
During and after the breeding season, males experience a trade-off between reproduction
and survival. During breeding, males compete for territory ownership and access to
females. During this period of intense competition, the males are highly active and
often do not feed, resulting in high levels of stress, and body mass losses of up
to 21%. Many males will die after the breeding season as a result of compromised
immune systems and poor body condition (Barnes 1996, Boonstra et al. 2001, Lacy and
Wieczorek 2001).
After the breeding season, sex ratios become skewed towards females. Females are
highly philopatric and display nepotism, resulting in groups of closely-related, cooperative
females living together in close or connected burrow systems (kin clusters). They
often clump their litters together and share in the maternal responsibilities. This
is thought to serve two main functions: the first being shared duties in watching
for predators, the second being to protect the young from infanticidal males (McLean
1982,1983, Nowak 1999, Yensen and Sherman 2003). The incidence of infanticide by
males in arctic ground squirrels is high. Immigrant males that are looking to establish
new territories are most likely to engage in infanticide. Males target young that
they have most likely not sired, and litters are attacked both pre- and post-emergence
from the burrow. Females will fight to protect their young, and it is suspected that
a function of intense male territoriality is to protect the young that they have most
likely sired from other infanticidal males (McLean 1983).
- Mating System
- polygynous
- polygynandrous (promiscuous)
- cooperative breeder
Arctic ground squirrels breed in April during a two-week period shortly following
emergence from hibernation (Donker and Krebs 2011). Females give birth to one litter
per year in mid-May following a 25 to 30-day gestation period (Hayssen 2008, Donker
and Krebs 2011, Williams et al. 2011). Litters range from two to ten altricial pups
that are born hairless, toothless, blind, with unopened ears, and incapable of thermoregulation
(Yensen and Sherman 2003, Williams et al. 2011). After two days hair begins to appear,
and they are fully furred by the tenth day (Iwen 1999). Lactation lasts for 28 to
35 days, and pups come above ground around the 27th day in mid-June. Weaning mass
is approximately 199 g (Hayssen 2008, Williams et al. 2011). Within five to six weeks,
the pups undergo a six to ten fold increase in body size, reaching 80% of their adult
weight (Yensen and Sherman 2003). At eight to ten weeks, sexual dimorphism becomes
apparent, and the juveniles disperse at this time (Byrom and Krebs 1999, Yensen and
Sherman 2003). A rapid growth rate of the young is necessary to ensure that they
are able to survive the coming hibernation season (Yensen and Sherman 2003). The young
are reproductively active by the following spring (Lacy et al. 1997).
- Key Reproductive Features
- iteroparous
- seasonal breeding
- gonochoric/gonochoristic/dioecious (sexes separate)
- sexual
- viviparous
Arctic ground squirrel females spend the majority of their time underground with their
young during lactation. They will emerge several times a day for short foraging bouts
before returning to their young (Williams et al. 2011). Females will form kin clusters
with other closely-related females, where they will clump their young together and
share in the maternal responsibilities of watching for predators and defending the
young against conspecifics (McLean 1982, 1983, Nowak 1999, Yensen and Sherman 2003).
Mothers will care for their young for another one to two weeks following weaning and
emergence from the burrow (Iwen 1999). Paternal parental investments are minimal
and may be restricted to territory defense surrounding their females and sired young
(McLean 1983).
- Parental Investment
- altricial
- female parental care
-
pre-weaning/fledging
-
provisioning
- female
-
protecting
- female
-
provisioning
-
pre-independence
-
provisioning
- female
-
protecting
- female
-
provisioning
- post-independence association with parents
Lifespan/Longevity
Arctic ground squirrel mortality sources include predation, disease, starvation, freezing,
and death by conspecifics. Mortality during the first year of life is 54 to 74%.
Survival is dimorphic, with females living longer than males. Increased male mortality
is due to several factors. Males experience higher mortality rates that are associated
with increased dispersal demands and risks, and aggressive interactions between males.
Males also experience increased predation during the mating season due to their increased
visibility associated with their increased activity. They also experience a higher
overwinter mortality that is associated with competition for optimal hibernacula sites
with females (Yensen and Sherman 2003). In contrast, highly philopatric individuals
(mainly females) have a 73% survival rate, and dispersants (mainly juvenile males)
have a 20 to 40% survival rate (Byrom and Krebs 1999).
Survival also varies depending on habitat and season. In the summer, survival is
lower in the boreal forest than in the alpine habitat, due to greater predation rates
in the boreal forest habitat. In winter, survival is higher in the boreal forest
than in the alpine habitat, due to increased hibernation demands in the alpine habitat
(Gillis et al. 2005b). Burrow site is also important for survival: burrow sites with
good visibility, and deeper warmer burrows, result in increased survival (Buck and
Barnes 1999b, Yensen and Sherman 2003).
Regulation of arctic ground squirrel populations is both density-dependent and density-independent.
Annual mortality depends on: individuals obtaining variable or limited resources such
as food and burrow sites, predator abundance and success, presence of infectious diseases,
and climatic conditions such as temperature and snow depth (Yensen and Sherman 2003).
Arctic ground squirrels will survive to a maximum of eight to ten years in the wild
(Forsyth 1999).
Behavior
Arctic ground squirrels are a colonial species. Communities are based around kin
clusters of closely-related adult females and their offspring, with a single territorial
male (McLean 1982, Yensen and Sherman 2003). Adult males establish territories in
the spring that are 0.14 to 2.2 ha, and may encompass several kin clusters (Nowak
1999). Adult females make use of existing burrows, or excavate new burrows, that
they use as refuge, and to birth and wean their pups (Yensen and Sherman 2003). Home
range size varies with sex, life stage, and resource density, but is generally 0.2
to 4 ha (Hubbs and Boonstra 1998, Nowak 1999). After weaning, young are independent
and will likely disperse. The females are highly philopatric and will likely remain
in the area and kin cluster. Females will only disperse if there is already a high
population density, and competition for resources is high (Byrom and Krebs 1999).
Males are highly likely to disperse, and 57 to 100% of juveniles will disperse in
their first year, and 41 to 68% of adult males will disperse after the breeding season.
This dispersal behavior is thought to function to reduce the chances of inbreeding,
and to minimize competition for mates (Byrom and Krebs 1999, Yensen and Sherman 2003).
Dispersal comes with a cost, and disperses tend to lose body mass and experience increased
mortality as a result. One study demonstrated a positive correlation between mortality
and dispersal distance, likely as a result of predation (Buck and Barnes 1999a, Yensen
and Sherman 2003).
Population densities of arctic ground squirrels are controlled by interrelated factors
such as resource density and predation. Most of the limitations are seen through
changes in reproductive success. In the boreal forest, predation and food availability
interact to form an additive limitation on litter size, body condition, percent of
lactating females, and number of successfully weaned litters. In arctic and tundra
environments, populations are often limited by food and burrow availability (Karels
et al. 2000).
Even though in the summer months there is almost constant sunlight, arctic ground
squirrels are diurnal and are most active between 5 am and 11 pm, at temperatures
of 17 to 33 degrees Celsius. They retire to their burrows at night and during inclement
weather to minimize their thermoregulatory costs (Long et al. 2005). Arctic ground
squirrels are obligate hibernators; they are active only three to five months of the
year and spend the rest in hibernation. Hibernation is an essential physiological
adaptation they employ to survive the extreme climatic conditions and extended period
of fasting that they experience in their winter arctic habitat (Buck and Barnes 1999a,
Williams et al. 2011). Beginning in July for females, and mid-August to September
for males, individuals will start to increase body mass and fat stores in preparation
for hibernation. Females will enter hibernation first in August, and males will follow
over a one-month interval. They will remain in hibernation for 215 to 240 days, females
remaining longer than males, and adults remaining longer than juveniles. Males will
emerge before females in mid-April and adults will emerge before juveniles (Buck and
Barnes 1999a,b). Burrow site plays an important role in the costs of hibernation.
Burrows with shrubby vegetation over them will accumulate more snow and have warmer
soil temperatures than burrows located on clear, windswept sites. Competition for
favored burrow sites can be intense, and males will become particularly fierce in
late summer defending their hibernacula. Adult females appear to inhabit the favored
burrows more than males and juveniles, and juveniles are often left with the least-favored
sites (Buck and Barnes 1999b, 2003).
During hibernation, arctic ground squirrels will drastically lower their body temperature
and metabolic rate. They will supercool their body temperature to -2.9° Celsius and
remain in this state of torpor for up to three weeks at a time. Even at this low
temperature, they must still thermoregulate since the ambient temperature and soil
temperature in their burrows will drop down to -18.8° Celsius. In between bouts of
torpor they will experience arousal episodes where they rewarm their body temperature
to euthermic levels (34 to 36° Celsius) for one to two days. These rewarming episodes
are the most energetically expensive cost of hibernation. They accomplish this rewarming
through shivering and non-shivering thermogenesis. Non-shivering thermogenesis uses
brown adipose tissue and fatty acids as a fuel source to achieve rewarming. During
the endothermic arousal, high energy costs are incurred through increased metabolic
rate, and elevated activity levels of major organs, like the heart and brain. This
incurred costs depletes glucose levels in the body, which are replenished through
gluconeogenesis of protein (lean mass) tissue. It is hypothesized that they require
the endothermic arousal episodes to maintain certain homeostatic processes that are
inhibited by the colder temperatures, like brain function. It is also thought that
they arouse simply to sleep, as severely reduced body temperature during torpor prevents
this (Buck and Barnes 1999b, 2000).
Males become euthermic four to six weeks earlier than females, but will remain in
their burrow for an additional 10 to 21 days. Males must arouse from hibernation
earlier in order to achieve sexual maturation. Spermatogenesis is not possible at
sub-zero temperatures and males must become euthermic to allow gonadal development.
During this time, they consume food they have cached in their hibernacula from the
previous season. Only males cache food in their hibernacula, and their ability to
cache an adequate supply of food from the previous season will impact their reproductive
potential and survival. When males arouse, they have lost close to 30% of their body
weight. By feeding on their cache, they will regain that mass and emerge from their
burrows the same mass they were at immergence the previous fall. Females, on the
other hand, will emerge from hibernation with a greater than 30% loss in body mass
from the time of immergence (Barnes 1996, Buck and Barnes 1999a).
- Key Behaviors
- fossorial
- diurnal
- motile
- hibernation
- territorial
- social
- colonial
Home Range
Arctic ground squirrels have home ranges between 0.2 to 4 ha.
Communication and Perception
Arctic ground squirrels engage in a form of altruism by giving alarm calls when predators
are in the area. While the individual who gives the call experiences a higher predation
risk associated with drawing attention to themselves, they are thought to benefit
evolutionarily by protecting their relatives, and assuring their genes will be passed
to subsequent generations (Yensen and Sherman 2003). It has been suggested that they
have different calls for different predators. They appear to use a high-pitched whistle
for aerial predators, which is difficult to pinpoint, and a guttural chatter for terrestrial
predators (Woods 1980).
Males communicate their territories to other males by marking the boundaries with
scent glands in their cheeks and back. Territoriality, as well as other confrontations,
is resolved through violent contact that includes biting and scratching. Individuals
are often seriously injured through such confrontations, some of which result in death.
Initial social contact between individuals involves nose-to-nose contact and pressing
against each other in various poses, in which the individuals smell and get a sense
of one another. These interactions can end amicably or violently, depending on the
situation (Woods 1980).
- Other Communication Modes
- pheromones
- scent marks
Food Habits
Arctic ground squirrels are generalist foragers, feeding on everything from plants,
invertebrates, small vertebrates (including their own species), eggs, birds, and carrion
(McLean 1985, Boonstra et al. 1990, Gillis et al. 2005a, Zazula et al. 2006, Cook
et al. 2010). They have been observed to actively prey on the eggs and chicks of
birds, as well as collared
lemmings
(Boonstra et al. 1990, Cook et. 2010). Plant matter makes up the majority of their
diet however, and when caching food for winter stores, they use only plant material.
They consume all parts of plants, depending on the species, including foliage, roots,
seeds, flowers, and fruits. Commonly consumed plant families include:
Polygonaceae
,
Juncaceae
,
Cyperaceae
,
Ranunculaceaea
,
Rosaceae
,
Brassicaceae
,
Caryophllaceae
,
Poaceae
,
Saxifragaceae
,
Salicaceae
,
Gentianaceae
,
Scrophulariaceae
,
Poaceae
,
Asteraceae
,
Plantaginaceae
,
Fabaceae
, and
Santalaceae
(McLean 1985, Gillis et al. 2005a Zazula et al. 2006). They are known to selectively
choose and cache certain species, dependent on habitat location (Gillis et al. 2005a).
Arctic ground squirrels have also been shown to select for a diet with moderate levels
of polyunsaturated fatty acids (PUFAs). Polyunsaturated fatty acids are essential
fatty acids that are incorporated into cell membranes. They play a critical role for
hibernating animals that need to maintain certain levels of PUFAs in their diet, in
order to maintain flexibility in cell membranes when body temperatures drop to subnormal
levels. However, a diet too high in PUFAs results in increased lipid peroxidation,
which produces lipid peroxides that damage cells. Therefore, arctic ground squirrels
select foods that provide moderate levels of PUFAs in the fall before hibernation
(Frank et al. 2008).
- Primary Diet
-
carnivore
- eats terrestrial vertebrates
- eats eggs
- insectivore
- eats non-insect arthropods
- scavenger
- herbivore
- omnivore
- Animal Foods
- birds
- mammals
- eggs
- carrion
- insects
- terrestrial non-insect arthropods
- Plant Foods
- leaves
- roots and tubers
- seeds, grains, and nuts
- fruit
- flowers
- Foraging Behavior
- stores or caches food
Predation
Arctic ground squirrels are an important prey source for many arctic and boreal predators.
Anti-predator adaptations include extreme vigilance while above ground and away from
their protective burrows. While foraging, they will frequently sit or stand up on
their hind legs to listen and look around them. When predators are in the area, arctic
ground squirrels will give alarm calls to alert other family members. When an alarm
call is given, individuals will likely run for the protection of their burrows (Yensen
and Sherman 2003). Predation is a limiting factor in most populations and can influence
their distributions, particularly within the boreal forest habitat. In the boreal
forest, they are the third-most abundant small prey species, and during low cycles
of the snowshoe hare cycle, many predators switch from hares to arctic ground squirrels,
limiting populations even further (Karels and Boonstra 1999, Byrom et al. 2000).
The preferred habitat of arctic ground squirrels is open areas with little vegetation,
which allow them to see predators coming from greater distances (Yensen and Sherman
2003). Common predators of arctic ground squirrels include
lynxes
,
coyotes
,
wolverines
,
red foxes
,
arctic foxes
,
wolves
,
grizzly bears
,
ermines
,
northern goshawks
,
great horned owls
,
red tailed and Harlens hawks
,
common ravens
,
long tailed jaegers
,
snowy owls
,
short eared owls
,
golden eagles
,
northern harriers
,
gyrofalcons
,
rouch legged falcons
, and
peregrine falcons
(Buck and Barnes 1999a, Byrom et al. 2000, Barker and Derocher 2010).
- Anti-predator Adaptations
- cryptic
Ecosystem Roles
Arctic ground squirrels are a keystone species, having both top-down and bottom–up
effects on ecological processes (Yensen and Sherman 2003, Barker and Derocher 2010).
They are an important food source for many mammalian and avian predators. Many predator
species may be dependent on them during low cyclic phases of other prey species (Barker
and Derocher 2010).
Arctic ground squirrels are also important soil engineers. Through digging of their
burrows they help to aerate and turn over the soil, bringing nutrients to the surface
and breaking up the soil. They have been known to excavate up to 18 tons/ha/year
of soil (Barker and Derocher 2010). Their burrowing has been shown to increase a soil’s
water infiltration rates and holding capacity, cation and anion exchange capacity,
organic matter content, nutrient levels, and seed germination rates. This increases
plant productivity and species composition of microhabitats, which is a benefit to
other arctic herbivores and grazers. Through continued use of burrow sites, their
feces and urine fertilize the soil with nitrogen and phosphorus, which creates a positive
feedback loop of increased vegetation, which leads to increased snow cover over the
soil, which leads to warmer soil temperatures, which leads to increased habituation
by arctic ground squirrels (Yensen and Sherman 2003, Barker and Derocher 2010).
Arctic ground squirrels also serve as a host to several parasites including the fleas,
Oropsylla alaskensis
and
Oropsylla idahoensis
, and digestive tract dwelling protists,
Eimeria callospermophila
,
Eimeria cynomysis
,
Eimeria lateralis
, and
Eimeria morainensis
(Hass et al. 1982, Seville et al. 2005).
- Ecosystem Impact
- disperses seeds
- creates habitat
- soil aeration
- keystone species
Economic Importance for Humans: Positive
Arctic ground squirrels are a main prey species of many economically valued fur-bearing
and game species including, ermine, coyote, wolf, wolverine, lynx, red and arctic
fox, and grizzly bear. Their pelts were also used in the manufacture of parkas, and
by indigenous Alutiiq and Aleut peoples who valued them in making garments (Nowak
1999, Cook et al. 2010). In the last 30 years, arctic ground squirrels have also
been extensively studied by researchers as an uniquely-adapted endothermic arctic
mammal.
- Positive Impacts
- body parts are source of valuable material
- research and education
Economic Importance for Humans: Negative
Many species of ground squirrels serve as hosts to flea parasites that carry the sylvatic
plague. However, it is unknown whether arctic ground squirrels are host to these
species (Yensen and Sherman 2003). It is also possible that arctic ground squirrels
could be vectors for the rabies virus, however, there have been no reported cases
of rabies in arctic ground squirrels in Alaska (State of Alaska Department of Epidemiology
2011).
Conservation Status
As part of the Alaska State Department of Fish and Game’s Comprehensive Wildlife Conservation
Strategy, several subspecies of arctic ground squirrel have been identified as a G5T3
status which is, “vulnerable- at moderate risk of extinction due to a restricted range,
relatively few populations (often 80 or fewer), recent and widespread declines, or
other factors.” These subspecies include
S. p. kodiacensis
,
S. p. lyratus
,
S. p. nebulicola
, and
S. p. osgoodi
.
The International Union for the Conservation of Nature (IUCN) lists arctic ground
squirrels as a species of Least Concern. Their justification for this listing is
that “this species has a large population size and a wide distribution. It is abundant
in parts of the range. Although it is hunted for meat and skins for local trade, this
is not thought to threaten the species as a whole” (Linzey 2008).
Other Comments
The origin and meaning of arctic ground squirrel’s scientific name,
Spermophilus parryii
, is two part. Spermophilus means seed lover, and parryii was chosen by northern
explorer Sir John Richardson, after the American botanist and explorer, Dr. C. C.
Parry (1823 to 1890) (Forsyth 1999). The indigenous Inuits call the squirrel sik-sik
or sik-rik, after its chattering alarm call that they make when humans are near (Iwen
1999).
Arctic ground squirrel populations in North America have genetically diverged due
to geographic barriers, and the nature of their highly philopatric lifestyle and patchy
distribution. Currently there are eight recognized subspecies, six of which are divided
into four geographic clades. The Southeast clade includes
S. p. plesius
, which has a subarctic distribution through southeastern Alaska, extending through
the Yukon Territory and into the western Northwest Territories and northern British
Columbia in Canada. The Arctic clade includes
S. p. kennicottii
, which is the northernmost species and is distributed north of the Arctic Circle
in Alaska and in the Yukon and Northwest Territories. The Beringia clade includes
S. p. ablusus
which inhabits the Seward Peninsula of Alaska,
S. p. lyratus
, which inhabits St. Lawrence Island in Alaska, and
S. p. osgoodii
, which inhabit a small area in central Alaska. The Southwest clade includes
S. p. ablusus
, which inhabit southwestern Alaska,
S. p. kodiacensis
, which inhabit the Kodiak and Semidi Islands in Alaska, and
S. p. nebulicola
, which inhabit the Shumagin and Chernabura Islands in Alaska. The other subspecies
is
S. p. parryii
, which inhabit northern Canada through the Northwest Territories and Nunavut (Hall
1981, Galbreath et al. 2011). Recent studies have suggested that arctic ground squirrels
evolved in Beringia during the last glacial period. The earliest fossils in North
America are from the Yukon Territory in Canada, and date back to 740 thousand years
ago. In contrast, the earliest fossil evidence in Siberia only dates back to 33 thousand
years ago. This, along with recent molecular data that show three major evolutionarily
distinct lineages consisting of the Arctic, Southeast, and Beringia/Southwest geographic
clades, suggest that arctic ground squirrels evolved in North America and were separated
into isolated clades by glacial ice expansion. During the late Wisconsonian, it is
likely that populations migrated across the Bering land bridge, colonizing Siberia,
before becoming isolated again at the end of the glacial period (Galbreath et al.
2011).
Additional Links
Contributors
Nicole Torre (author), University of Alaska Fairbanks, Link Olson (editor), University of Alaska Fairbanks, Laura Podzikowski (editor), Special Projects.
- Nearctic
-
living in the Nearctic biogeographic province, the northern part of the New World. This includes Greenland, the Canadian Arctic islands, and all of the North American as far south as the highlands of central Mexico.
- introduced
-
referring to animal species that have been transported to and established populations in regions outside of their natural range, usually through human action.
- native range
-
the area in which the animal is naturally found, the region in which it is endemic.
- holarctic
-
a distribution that more or less circles the Arctic, so occurring in both the Nearctic and Palearctic biogeographic regions.
Found in northern North America and northern Europe or Asia.
- polar
-
the regions of the earth that surround the north and south poles, from the north pole to 60 degrees north and from the south pole to 60 degrees south.
- terrestrial
-
Living on the ground.
- tundra
-
A terrestrial biome with low, shrubby or mat-like vegetation found at extremely high latitudes or elevations, near the limit of plant growth. Soils usually subject to permafrost. Plant diversity is typically low and the growing season is short.
- taiga
-
Coniferous or boreal forest, located in a band across northern North America, Europe, and Asia. This terrestrial biome also occurs at high elevations. Long, cold winters and short, wet summers. Few species of trees are present; these are primarily conifers that grow in dense stands with little undergrowth. Some deciduous trees also may be present.
- mountains
-
This terrestrial biome includes summits of high mountains, either without vegetation or covered by low, tundra-like vegetation.
- endothermic
-
animals that use metabolically generated heat to regulate body temperature independently of ambient temperature. Endothermy is a synapomorphy of the Mammalia, although it may have arisen in a (now extinct) synapsid ancestor; the fossil record does not distinguish these possibilities. Convergent in birds.
- heterothermic
-
having a body temperature that fluctuates with that of the immediate environment; having no mechanism or a poorly developed mechanism for regulating internal body temperature.
- bilateral symmetry
-
having body symmetry such that the animal can be divided in one plane into two mirror-image halves. Animals with bilateral symmetry have dorsal and ventral sides, as well as anterior and posterior ends. Synapomorphy of the Bilateria.
- polygynous
-
having more than one female as a mate at one time
- polygynandrous
-
the kind of polygamy in which a female pairs with several males, each of which also pairs with several different females.
- cooperative breeder
-
helpers provide assistance in raising young that are not their own
- iteroparous
-
offspring are produced in more than one group (litters, clutches, etc.) and across multiple seasons (or other periods hospitable to reproduction). Iteroparous animals must, by definition, survive over multiple seasons (or periodic condition changes).
- seasonal breeding
-
breeding is confined to a particular season
- sexual
-
reproduction that includes combining the genetic contribution of two individuals, a male and a female
- viviparous
-
reproduction in which fertilization and development take place within the female body and the developing embryo derives nourishment from the female.
- altricial
-
young are born in a relatively underdeveloped state; they are unable to feed or care for themselves or locomote independently for a period of time after birth/hatching. In birds, naked and helpless after hatching.
- female parental care
-
parental care is carried out by females
- fossorial
-
Referring to a burrowing life-style or behavior, specialized for digging or burrowing.
- diurnal
-
- active during the day, 2. lasting for one day.
- motile
-
having the capacity to move from one place to another.
- hibernation
-
the state that some animals enter during winter in which normal physiological processes are significantly reduced, thus lowering the animal's energy requirements. The act or condition of passing winter in a torpid or resting state, typically involving the abandonment of homoiothermy in mammals.
- territorial
-
defends an area within the home range, occupied by a single animals or group of animals of the same species and held through overt defense, display, or advertisement
- social
-
associates with others of its species; forms social groups.
- colonial
-
used loosely to describe any group of organisms living together or in close proximity to each other - for example nesting shorebirds that live in large colonies. More specifically refers to a group of organisms in which members act as specialized subunits (a continuous, modular society) - as in clonal organisms.
- visual
-
uses sight to communicate
- tactile
-
uses touch to communicate
- acoustic
-
uses sound to communicate
- chemical
-
uses smells or other chemicals to communicate
- pheromones
-
chemicals released into air or water that are detected by and responded to by other animals of the same species
- scent marks
-
communicates by producing scents from special gland(s) and placing them on a surface whether others can smell or taste them
- visual
-
uses sight to communicate
- tactile
-
uses touch to communicate
- acoustic
-
uses sound to communicate
- chemical
-
uses smells or other chemicals to communicate
- carrion
-
flesh of dead animals.
- stores or caches food
-
places a food item in a special place to be eaten later. Also called "hoarding"
- cryptic
-
having markings, coloration, shapes, or other features that cause an animal to be camouflaged in its natural environment; being difficult to see or otherwise detect.
- soil aeration
-
digs and breaks up soil so air and water can get in
- keystone species
-
a species whose presence or absence strongly affects populations of other species in that area such that the extirpation of the keystone species in an area will result in the ultimate extirpation of many more species in that area (Example: sea otter).
- carnivore
-
an animal that mainly eats meat
- insectivore
-
An animal that eats mainly insects or spiders.
- scavenger
-
an animal that mainly eats dead animals
- herbivore
-
An animal that eats mainly plants or parts of plants.
- folivore
-
an animal that mainly eats leaves.
- frugivore
-
an animal that mainly eats fruit
- granivore
-
an animal that mainly eats seeds
- omnivore
-
an animal that mainly eats all kinds of things, including plants and animals
References
Barker, O., A. Derocher. 2010. Habitat selection by arctic ground squirrels (Spermophilus parryii). Journal of Mammology , 91/5: 1251-1260.
Barnes, B. 1989. Freeze avoidance in a mammal: body temperatures below O C in an arctic hibernator. Science , 244/4912: 1593-1595.
Barnes, B. 1996. Relationships between hibernation and reproduction in male ground squirrels. Adaptations to the cold: Tenth International Hibernation Symposium: 71-80.
Boonstra, R., C. Krebs, M. Kanter. 1990. Arctic ground squirrel predation on collared lemmings. Canadian Journal of Zoology , 68: 757-760.
Boonstra, R., C. McColl, T. Karels. 2001. Reproduction at all costs: the adaptive stress response of male arctic ground squirrels. Ecology , 82/7: 1930-1946.
Buck, C., B. Barnes. 2003. Androgen in free-living arctic ground squirrels: seasonal changes and influence of staged male-male aggressive encounters. Hormones and Behavior , 43: 318-326.
Buck, C., B. Barnes. 1999. Annual cycle of body composition and hibernation in free-living arctic ground squirrels. Journal of Mammalogy , 80/2: 430-442.
Buck, C., B. Barnes. 2000. Effects of ambient temperature on metabolic rate, respiratory quotient, and torpor in an arctic hibernator. The American Journal of Physiology - Regulatory, Integrative and Comparative Physiology , 279: 255-262.
Buck, C., B. Barnes. 1999. Temperatures of hibernacula and changes in body composition of arctic ground squirrels over winter. Journal of Mammology , 80/4: 1264-1276.
Byrom, A., T. Karels, C. Krebs, R. Boonstra. 2000. Experimental manipulation of predation and food supply of arctic ground squirrels in the boreal forest. Canadian Jounal of Zoology , 78: 1309-1319.
Byrom, A., C. Krebs. 1999. Natal dispersal of juvenile arctic ground squirrels in the boreal forest. Canadian Journal of Zoology , 77: 1048-1059.
Cook, J., A. Eddingsaas, J. Loxterman, S. Ebbert, S. MacDonald. 2010. Insular arctic ground squirrels (Spermophilus parryii) of the North Pacific: indigenous or exotic?. Journal of Mammology , 91/6: 1401-1412.
Donker, S., C. Krebs. 2011. Habitat-specific distribution and abundance of arctic ground squirrels (Urocitellus parryii plesius) in southwest Yukon. Canadian Journal of Zoology , 89: 570-576.
Forsyth, A. 1999. Mammals of North America: Temperate and Arctic Regions . Firefly Books Ltd..
Frank, C., S. Karpovich, B. Barnes. 2008. Dietary fatty acid compostion and the hibernation patterns in free-ranging arctic ground squirrels. Physiological and Biochemical Zoology , 81/4: 486-495.
Galbreath, K., J. Cook, A. Eddingsaas, E. Dechaine. 2011. Diversity and demography in Beringia: multilocus tests of paleodistribution models reveal the complex history of arctic ground squirrels. Evolution , 65/7: 1879-1896.
Gillis, E., D. Hik, R. Boonstra, T. Karels, C. Krebs. 2005. Being high is better: effects of elevation and habitat on arctic ground squirrel demography. OIKOS , 108: 231-240.
Gillis, E., S. Morrison, G. Zazula, D. Hik. 2005. Evidence for selective caching by arctic ground squirrels living in alpine meadows in the Yukon. Arctic , 58/4: 354-360.
Haas, G., N. Wilson, R. Zarnke, R. Barrett, T. Rumfelt. 1982. Siphonaptera from mammals in Alaska: Supplement III Western Alaska. Canadian Journal of Zoology , 60: 729-732.
Hall, E. 1981. The Mammals of North America: Sciurid rodents Volume 1: 2nd edition . John Wiley & Sons.
Hayssen, V. 2008. Reproductive efforts in squirrels: ecological, phylogenetic, allometric, and latitudinal patterns. Journal of Mammalogy , 89/3: 582-606.
Hubbs, A., R. Boonstra. 1998. Effects of food and predators on the home-range sizes of arctic ground squirrels (Spermophilus parryii). Canadian Journal of Zoology , 76: 592-596.
Iwen, F. 1999. The Smithsonian Book of North American Mammals . The Smithsonian Institution.
Karels, T., R. Boonstra. 1999. The impact of predation on burrow use by arctic ground squirrels in the boreal forest. Proceedings of the Royal Society: Biological Sciences , 266/1433: 2117-2123.
Karels, T., A. Byrom, R. Boonstra, C. Krebs. 2000. The interactive effects of food and predators on reproduction and overwinter survival of arctic ground squirrels. Journal of Animal Ecology , 69: 235-247.
Karpovich, S., O. Toien, C. Buck, B. Barnes. 2009. Energetics of arousal episodes in hibernating arctic ground squirrels. Journal of Comparative Physiology B: Biochemical, Systemic, and Environmental Physiology , 179: 691-700.
Lacy, E., J. Wieczorek, P. Tucker. 1997. Male mating behavior and patterns of sperm precedence in arctic ground squirrels. Animal Behavior , 53: 767-779.
Lacy, E., J. Wieczorek. 2001. Territoriality and male reproductive success in arctic ground squirrels. Behavioral Ecology , 12/5: 626-632.
Linzey, A. 2008. "IUCN Red List of Threatened Species Version 2011.2" (On-line). Accessed December 12, 2011 at http://www.iucnredlist.org/apps/redlist/details/20488/0 .
Long, R., T. Martin, B. Barnes. 2005. Body temperature and activity patterns in free-living arctic ground squirrels. Journal of Mammalogy , 86/2: 314-322.
McLean, I. 1983. Paternal behavior and killing of young arctic ground squirrels. Animal Behavior , 31: 32-44.
McLean, I. 1985. Seasonal patterns and sexual differences in the feeding ecology of arctic ground squirrels (Spermophilus parryii plesius). Candian Journal of Zoology , 63: 1298-1301.
McLean, I. 1982. The association of female kin in the arctic ground squirrel Spermophilus parryii. Behavioral Ecology and Sociobiology , 10/2: 91-99.
Nowak, R. 1999. Mammals of the World: Ground Squirrels or Susliks Volume 2 6th edition . The John Hopkins University.
Seville, R., C. Oliver, A. Lynch, M. Bryant, D. Duszynski. 2005. Eimeria species (Apicomplexa: Eimeriidae) from arctic ground squirrels (Spermophilus parryii) and red squirrels (Tamiasciurus hudsonicus) in Alaska and in Siberia, Russia. The Journal of Parasitology , 91/4: 857-862.
Sheriff, M., G. Kenagy, M. Richter, T. Lee, O. Toien, F. Kohl, C. Buck, B. Barnes. 2011. Phenological variation in annual timing of hibernation and breeding in nearby populations of arctic ground squirrels. Proceedings of the Royal Society: Biological Sciences , 278: 2369-2375.
Williams, C., M. Sheriff, J. Schmutz, F. Kohl, O. Toien, C. Buck, B. Barnes. 2011. Data logging of body temperatures provides precise information on phenology of reproductive events in a free-living arctic hibernator. Journal of Compartive Physiology B: Biochemical, Systemic, and Environmental Physiology , 181/8: 1101-1109.
Withers, P., T. Casey, K. Casey. 1979. Allometry of respiratory and heamatological parameter of arctic mammals. Comparative Biochemical Physiology , 64: 343-350.
Woods, S. 1980. The Squirrels of Canada . National Museums of Canada.
Yensen, E., P. Sherman. 2003. Wild Mammals of North America: Biology, Managment, and Conservation 2nd edition: Spermophilus and Ammospermophilus species . The John Hopkins University Press.
Zazula, G., R. Mathewes, A. Harestad. 2006. Cache selection by arctic ground squirrels inhabiting boreal-steppe meadows of southwest Yukon Territory, Canada. Arctic, Antarctic, and Alpine Research , 38/4: 631-638.
State of Alaska Department of Epidemiology. Alaska Rabies Prevention and Control Manual. Juneau, Alaska: State of Alaska. 2011.
Alaska Department of Fish and Game. Our wealth maintained: A strategy for conserrving Alaska's diverse wildlife and fish resources. Juneau, Alaska: Alaska Department of Fish and Game. 2006.