Diversity
Twelve species are currently recognized within this genus with 32 subspecies in total.
All but
Urocitellus undulatus
are native to the Nearctic region. All are burrowing ground squirrels but only
Urocitellus brunneus
is labeled as threatened under the IUCN list. There is evidence of some interspecies
competition, but only two of these species occur in the same range. In at least one
case speciation has occurred within this genus, thus
U.richardsonii
split to form the current
Urocitellus elegans
(Durrant and Hansen, 1954). There is little fossil evidence connected to this genus
due to poor fossilization or a lack of collection, so little is known about their
evolutionary history (Black, 1972). Many of the species live in high-altitude habitats
with a preference for prairie and desert ecosystems. Research is lacking for most
of this genus because of their status as pests within agricultural regions including
reports of livestock injuries and crop destruction caused by their burrows (Howell,
1938).
Geographic Range
Urocitellus
includes ground squirrels of the Nearctic with a larger presence in northwestern
North America. Their geographic distribution is one of the identifiers for this genus
beyond morphological and genetic data. Exceptions to this are
Urocitellus undulatus
, which is found in the Oriental region, and
Urocitellus parryii
, which is found in the Arctic regions of North America. Most species have limited
geographic ranges, with some being exclusive to mountainous regions.
- Other Geographic Terms
- holarctic
Habitat
The species that makeup
Urocitellus
occupy a wide array of habitats, from alpine to prairie ecosystems and meadows. They
are primarily herbivorous, eating seeds, grasses, and other available forage (Howell,
1938). In some cases, arthropods are also part of their foraging, particularly during
the harsher seasons. The species
U. elegans
is noted along with
U. richardsonii
to be found in high altitudes and dry habitats (Durrant and Hansen, 1954). All are
colonial species that form a series of burrowing networks that are used for raising
young and hibernating. Due to their geographic locations, all are adapted to be either
true hibernators or experiencing extended periods of torpor.
- Habitat Regions
- temperate
- terrestrial
- Terrestrial Biomes
- tundra
- desert or dune
- savanna or grassland
- mountains
- Other Habitat Features
- agricultural
Systematic and Taxonomic History
Urocitellus
was first proposed in 2009 when previously part of the larger genus
Spermophilus
that included groups such as marmots. In 2009, there was a publication supporting
the splitting of
Spermophilus
into multiple genera including
Urocitells
. The grouping of
Urocitellus
is based on genetic data and morphological traits including tail length. Before the
split, there were many alterations to the identification of this and other related
groups of ground squirrels recorded as early as the 1880s. However, the only current
valid synonym is Spermophilus as this term has been used the longest. In terms of
relatedness,
Urocitellus
is closely related to other ground squirrels in the Americas and Europe such as the
golden mantel, as well as related to marmots and prairie dogs.
Physical Description
Color varies between species, with a few exhibiting a seasonal coat color despite
being hibernators. They fall in the medium-size range within
f.Spermophilus
, and all distinct tails, with one species having a notably long tail
U. undulatus
. At first glance, they appear very similar to prairie dogs, but are much smaller
and have a longer body shape in general. Most species are light to dark brown, with
a light-colored belly and a tipped tail in white or black. Coloration is used for
predation avoidance and thermoregulation with a lighter ventral side.
- Sexual Dimorphism
- sexes alike
Reproduction
Males in colonial species are responsible for defending the colony, and a harem structure
is thought to be involved. However, in some species like
U. columbianus
, both the male and female have multiple mates for an increase in fitness. In more
distantly spaced species, the males search for a female and defend her from other
males doing the same. Verbal and non-verbal attraction is common. Calling and verbal
communication are heavily used in this genus with various chirps, squeaks, and screams
used.
- Mating System
- polygynous
- polygynandrous (promiscuous)
Mating behavior heavily revolves around hibernation, with males emerging first around
early April. Females emerge soon after in mid-April to May, with mating occurring
4 days later. After, females undergo a month-long gestation period before giving birth
to a litter. All pups are born blind and naked and will spend a month in a nest burrow
in their mother's den before emerging in the late summer. Oddly, while the females
are in gestation, the previous year’s litter will emerge sexually mature despite all
mating occurring at a single time. For most species, mating takes place underground,
but a few species or even subspecies specifically have been observed above ground,
and the encounter lasts on average half an hour.
- Key Reproductive Features
- iteroparous
- seasonal breeding
- viviparous
Adults emerge first after hibernation to mate and males defend their territories or
colonies. During the month that pups stay in the burrows, the female provides food,
shelter, and protection. Near the end of the summer period, pups will start to emerge:
although fairly independent, they still live in their mother's burrow until after
their first hibernation. It is thought that the males are kicked out of the territory
after their first year while females either stay or leave on their own. There is not
much evidence for group raising of pups, but pups will be independent at emergence
from burrows.
- Parental Investment
- altricial
- female parental care
Lifespan/Longevity
While there is variation between species, overall this genus has a differing lifespan
based on sex. There is evidence that males live shorter lives than females, averaging
around 4 years. This is based on survival rates before and during hibernation, with
females surviving on average 4 to 6 years. There isn't much data on captive lifespans
as there is little to no captive care or breeding of this genus. The reason why males
don't live as long is not entirely known, but can possibly be correlated to predation,
territory disputes, and dispersion patterns.
Behavior
This genus is made almost entirely of social species, though the degree of sociality
varies. Species like the
Urocitellus parryii
are fairly solitary, but maintain territories with multiple females. Species like
the
Urocitellus elegans
and
Urocitellus richardsonii
have intermingling colonies run by one or two dominant males. All hibernate during
the cold seasons of their habitat, with females and males having separate hibernating
burrows. Territory disputes can get very aggressive, even between adults and pups,
whilst still having a high level of communication within social groups for predators.
In habitats that get very hot in the summer, a decrease in activity will occur: burrowing
species will go underground and will be most active in the morning followed by a short
period in the evening.
- Key Behaviors
- terricolous
- fossorial
- diurnal
- sedentary
- hibernation
- territorial
- colonial
- dominance hierarchies
Communication and Perception
The species of this genus are highly vocal as one can usually hear chatter from colonies
throughout the day and even at night. The vocalization varies between species but
squeaks, barks, trills, and chirps are commonly used. They are able to communicate
mating, predation avoidance, territory disputes, etc. Often there is some visual component
to communication with tail movement or body posture.
Food Habits
There is not much detail in the text on the diet or feeding behavior on
Urocitellus
, but there is a fair amount of general information. They are primarily granivores,
preferring high alpine grasses and seeds, but have also been observed eating local
arthropods including cicadas in the case of
U. canus
. When present in agricultural areas, they will also feed on crops. This causes damage
to the annual yield of the fields. Due to
Urocitellus
being hibernators, they do not stockpile food. They do, however, store a large amount
of body fat during the active season and, depending on the environmental conditions,
will either increase their foraging time closer to hibernation or go into an early
hibernation if reserves are large enough by that time. In very rare and extreme instances
cannibalism has been observed a few times in this genus. All observations involved
an adult eating a pup with no other options available for food.
Predation
Due to being a social species, there is a lot of communication used to avoid predators.
There will often be individuals acting as lookouts for threats that will signal to
others. Burrows are used to avoid predators, and are most effective against birds.
Nearly all species within this genus have contrasting coloration and possess a tan
or brown colored dorsal side with a light ventral side. This is a good form of camouflage
for their habitat.
- Anti-predator Adaptations
- aposematic
- cryptic
Ecosystem Roles
This information is based on the research of
U. canus
and
U. parryii
as there isn't information on the genus level and not all species have had in-depth
research on them.
Urocitellus
are a prey species that primarily feeds on grasses and seeds, which make up the majority
of their niche. Small and large predators including wolverines, bears, and hawks feed
on these ground squirrels with possible mutualism between the ground squirrels and
the other prey item: rabbits. When one group is doing poorly, there is more predation
on the other population and vice versa. This, however, is only suggested. Though a
mutualistic relationship has been shown between marmots and
U. parryii
, the nature of this relationship is unknown. In terms of diet, they do provide some
seed dispersal, though
Urocitellus
has not been shown to forage based on abundance but rather on palatability. They
display a preference for roots over flowers and more green growth instead of the more
abundant dry grass. Vegetation usage or coverage is possibly the reason for speciation
within the genus as there are multiple species and subspecies that are distinct despite
living right next to each other range-wise.
In terms of space usage,
Urocitellus
burrows and usually inhabits large meadows and plains over forests. This is because
they need well-drained soil to burrow in. This may initially cause a decrease in diversity
in the nearby areas of the burrows: long-term burrowing allows for higher diversity
in plant growth and even increases water retention of the soil. On top of that, the
burrows may provide second-hand homes for other species including foxes, badgers,
or other rodents. While they are not a keystone species, they can be landscape drivers
for ecosystems and allow for more niches and diversity within habitats.
- Ecosystem Impact
- disperses seeds
- creates habitat
- soil aeration
- Marmots
- siphonapterid fleas Oropsylla alaskensis
- the sucking louse Linognathoides laeviusculus
- The roundworm Ascaris laevis
- cestode Paranoplocephala wigginsi
Economic Importance for Humans: Positive
There is still work being done to determine whether or not some of the species could
be used as an ecosystem indicator for disturbances. However, these species do not
have a significant positive impact on the economy.
- Positive Impacts
- research and education
Economic Importance for Humans: Negative
Nearly all references to ground squirrel species are done in the light of them being
pests. It is incredibly common to find detailed accounts of ground squirrels being
responsible for damage to both crops and livestock with their burrowing behavior.
In all, they don't have a positive reputation in the public eye and all the preliminary
research of many of the genera has been on how to dispose of them or reduce their
presence in agricultural landscapes.
U. elegans
in particular has little research beyond how they are a pest and how to get rid of
them. There have been some cases of ground squirrels carrying plague among other infections,
but not all species within the genus act as carriers to the same degree.
- Negative Impacts
-
injures humans
- carries human disease
- crop pest
- causes or carries domestic animal disease
Conservation Status
The majority of
Urocitellus
are considered stable or of least concern. However, a couple of species are marked
as either vulnerable or endangered.
U. townsendii
subspecies
U. townsendii nancyae
are currently on a downhill trend in their population.
U. brunneus
and its subspecies are endangered but seem to be improving. On a broader scale,
Urocitellus
live in ecosystems that will see an increase in threats due to climate change, urban
development, and wildfires now and in the future. It is currently unknown if these
species will adapt well or become more vulnerable in the coming years.
Other Comments
Urocitellus
as a whole needs more research to be done. Taxonomically the relationship between
the oriental species and the Nearctic species is of interest, as well as their ability
to adapt to ecological threats. It is one of many examples of commonality resulting
in a lack of interest despite the potential knowledge gained.
Additional Links
Contributors
Kadin Samlaska (author), Colorado State University, Audrey Bowman (editor), Colorado State University, Tanya Dewey (editor), University of Michigan-Ann Arbor.
- 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.
- native range
-
the area in which the animal is naturally found, the region in which it is endemic.
- oriental
-
found in the oriental region of the world. In other words, India and southeast Asia.
- 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.
- temperate
-
that region of the Earth between 23.5 degrees North and 60 degrees North (between the Tropic of Cancer and the Arctic Circle) and between 23.5 degrees South and 60 degrees South (between the Tropic of Capricorn and the Antarctic Circle).
- 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.
- desert or dunes
-
in deserts low (less than 30 cm per year) and unpredictable rainfall results in landscapes dominated by plants and animals adapted to aridity. Vegetation is typically sparse, though spectacular blooms may occur following rain. Deserts can be cold or warm and daily temperates typically fluctuate. In dune areas vegetation is also sparse and conditions are dry. This is because sand does not hold water well so little is available to plants. In dunes near seas and oceans this is compounded by the influence of salt in the air and soil. Salt limits the ability of plants to take up water through their roots.
- tropical savanna and grassland
-
A terrestrial biome. Savannas are grasslands with scattered individual trees that do not form a closed canopy. Extensive savannas are found in parts of subtropical and tropical Africa and South America, and in Australia.
- savanna
-
A grassland with scattered trees or scattered clumps of trees, a type of community intermediate between grassland and forest. See also Tropical savanna and grassland biome.
- temperate grassland
-
A terrestrial biome found in temperate latitudes (>23.5° N or S latitude). Vegetation is made up mostly of grasses, the height and species diversity of which depend largely on the amount of moisture available. Fire and grazing are important in the long-term maintenance of grasslands.
- mountains
-
This terrestrial biome includes summits of high mountains, either without vegetation or covered by low, tundra-like vegetation.
- agricultural
-
living in landscapes dominated by human agriculture.
- 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.
- 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
- 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.
- sedentary
-
remains in the same area
- 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
- 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.
- dominance hierarchies
-
ranking system or pecking order among members of a long-term social group, where dominance status affects access to resources or mates
- visual
-
uses sight to communicate
- acoustic
-
uses sound to communicate
- visual
-
uses sight to communicate
- acoustic
-
uses sound to communicate
- aposematic
-
having coloration that serves a protective function for the animal, usually used to refer to animals with colors that warn predators of their toxicity. For example: animals with bright red or yellow coloration are often toxic or distasteful.
- 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
- causes or carries domestic animal disease
-
either directly causes, or indirectly transmits, a disease to a domestic animal
- carnivore
-
an animal that mainly eats meat
- insectivore
-
An animal that eats mainly insects or spiders.
- herbivore
-
An animal that eats mainly plants or parts of plants.
- granivore
-
an animal that mainly eats seeds
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