Geographic Range
Shadow chipmunks, Tamias senex , occur in California, Oregon, and Nevada. The species ranges northward from the central Sierra Nevadas in California, skirting Nevada near Lake Tahoe, and into the eastern Cascade Range of Oregon. In northern California, this species is found from the Pacific Coast to the Warner mountains between the Eel and Klamath rivers.
In the Sierras, shadow chipmunks are limited to elevations above 1,500 m. They do
not move to lower elevations on the eastern or western slopes of the range, where
other species of chipmunk occur.
Habitat
Tamias senex is primarily arboreal and resides in fairly dense, moist, mature forests. It forages on low-strewn forest floors and in adjacent chaparral.
Shadow chipmunk nests are found in hollow trees and fallen logs. The most common nesting trees include Jeffrey pine ( Pinus jeffreyi ), Ponderosa pine ( Pinus ponderosa ), sugar pine ( Pinus lambertiana ), Douglas fir ( Pseudotsuga menziesii ), white fir ( Abies concolor ), red fir ( Abies magnifica ), California black oak ( Quercus kellogii ), incense cedar ( Libocedrus decurrens ), and mountain hemlock ( Tsiga mertensiana ).
The common shrubs occurring in areas inhabited by
T. senex
include manzanita (
Arctostaphylos
), California lilac and buckbrush (
Ceanothus
), chinquapin (
Castanopsis sempervirens
), and blackberry (
Rubus
).
- Habitat Regions
- temperate
- terrestrial
Physical Description
Tamias senex is a relatively large chipmunk. The total length of T. senex is between 229 and 261 mm, and the hind foot measures 34 to 38 mm. As is the case for most members of the genus Tamias , females are slightly larger than males, with an average dimorphism ratio of 1.033. Mass of females ranges between 73.0 g and 108.5 g, compared to males, which measure 66.8 g to 99.3 g. Mean head and body lengths also differ between sexes, with females measuring 144.12 mm, compared to 139.54 mm for males.
Pelage is gray. The tail, which is shorter and less bushy than seen in
Tamias merriami
and
Tamias sonomae
, has faintly buffy edging, rather than the "distintly buffy" edging seen in
Tamias speciosus
.
The pelage of inland popluations of
T. senex
varies seasonally in color, length, and quality. In both the winter and summer, pelage
is white ventrally, gray on the top of the head and the rump, and has a marked black
dorsal stripe. Two molts occur annually. The winter pelage is replaced by a summer
molt just after breeding activity in spring. The summer molt begins at the head and
continues toward the rump, resulting in short, coarse, brightly-colored pelage. The
winter molt occurs in September, October, and November (just before hibernation) beginning
at the rump and continuing toward the head.
The winter molt results in long, dense, silky, dull-colored pelage with gray upper
parts.
Tamias senex
is polymorphic. Coastal populations are larger and darker in color, whereas inland
populations are smaller and paler. The transition from the dark pelage of coastal
specimens to the paler pelage of inland specimens is quite abrupt about 32 km inland
from the Pacific coast, near the eastern margin of the coastal forest. This variation
in pelage may be an adaptive response which makes these chipmunks more cryptically
colored in different habitats.
Genital bones of each sex are diagnostic of
T. senex
, and allow these animals to be distinguished from
Tamias orchrogenys
, a species with which it is often confused. Compared to
T. orchrogenys
, the baculum is longer (3.09 mm versus 2.47 mm) and more slender.
Chipmunks are endothermic. Because they enter hibernation, their body temperature
varies, making them heterothermic. However, they may also be considered homoiothermic.
Individuals maintain their body temperature around set points; higher when they are
active, and lower when they are torpid.
- Other Physical Features
- endothermic
- heterothermic
- homoiothermic
- bilateral symmetry
- Sexual Dimorphism
- female larger
Reproduction
The mating system of this species has not been described. However, many other species
in the genus
Tamias
are polygynous. Males may travel extensively during the breeding season, seeking
females with whom to mate. It is likely that
T. senex
is similar.
Young animals start to reproduce in the breeding season following their birth. Mating season begins in mid-March, about one month after these chipmunks emerge from hibernation, and continues for about four weeks. Populations found in the Great Basin begin their mating season earlier than populations in the Sierra Nevadas and the Volcanic Plateau.
Gestation is about 28 days, and average litter size is 4.5. The young are born in
late May or early June and are nursed by the mother until early August.
As the breeding season approaches, male reproductive organs increase dramatically
in size. Testis length doubles, and the prostate and cauda epididymis triple in width.
In addition, both the diameter of the bulbo-urethral glands and the length of the
seminal vesicles increase five times. Although the swelling and post-breeding reduction
of the testes are gradual, changes in the bulbo-urethral glands and semial vesicles
occur abruptly.
In females, the vulva perforate at the onset of estrus. The genital region becomes
swollen, and the uterine horns are inflated and twisted. Nipples become enlarged during
estrus and pregnancy, and reach maximum size during lactation.
- Key Reproductive Features
- iteroparous
- seasonal breeding
- gonochoric/gonochoristic/dioecious (sexes separate)
- sexual
- fertilization
- viviparous
Details on the parental investment of these animals are not available. However, in
other species of
Tamias
, males provide no parental care.
Tamias senex
is altricial at birth. Females provide all protection and care to the young. Young
usually disperse in the late summer or autumn of their birth year.
- Parental Investment
- altricial
-
pre-fertilization
- provisioning
-
protecting
- female
-
pre-hatching/birth
-
provisioning
- female
-
provisioning
-
pre-weaning/fledging
- provisioning
-
protecting
- female
-
pre-independence
-
protecting
- female
-
protecting
Lifespan/Longevity
Members of the subgenus
Neotamias
are reported to live as long as 8 years in the wild, and may reach almost 11 years
of age in captivity. Although there are no firm data for
T. senex
, we may assume that they are similar to other members of their subgenus.
Behavior
Tamias senex
is generally social. These chipmunks are diurnal, and hibernate from about November
through March.
Tamias senex
is one of the few species in North America which puts on a layer of fat to survive
the winter months. Seasonal fat accumulation increases the weight of individuals as
much as 20% before November. They experience fewer bouts of wakefulness during hibernation
than do other members of the genus.
These chipmunks are largely arboreal, and like other members of their subgenus, build
nests in trees. Such nests may be located as high as 23 m in the trees. Other western
chipmunks may also occupy night or winter nests in rocky ground locations. Although
data for this species are not available, they are likely to resemble other arboreal
chipmunks of western North America in this regard.
Home Range
Home ranges for Tamias amoenus , a related species, are recorded at 1.6 hectares for males and 0.93 hectares for females. It is likely that T. senex occupies similar home ranges.
Communication and Perception
Tamias senex
show differences in call notes compared to other chipmunks, like
T. sonomae
. Unlike other species,
T. senex
does not produce a whistle vocalization.
Tamias senex
produces an alarm call in 3 to 5 metallic barks with maximum frequency 12.98 kHz
and minimum frequency 4.56 kHz. The average call duration is 0.81 seconds, followed
by a single chirped note.
In addition to vocal communication, these diurnal mammals use visual cues, such as
body posture, in their interactions. During mating season, there are visual changes
in female genitalia, indicating that males may use such cues in finding mates. Tactile
communication between a mother and her young undoubtedly occurs in the nest, and may
play some role in mating behaviors. The role of chemical communication has not been
studied in these animals.
Food Habits
Tamias senex
is primarily herbivorous, spending the majority of its time collecting and storing
food. Diet consists mainly of the seeds of various herbs and grasses, but also contains
fungi, fruits of shrubs and forest trees, and occasional insects. Some examples of
food choices are acorns, chinquapins, spruces, hazelnuts, cherries, hemlocks, and
berries.
- Animal Foods
- insects
- Plant Foods
- leaves
- seeds, grains, and nuts
- fruit
- flowers
- Other Foods
- fungus
- Foraging Behavior
- stores or caches food
Predation
Natural predators including coyotes, badgers, weasels, snakes, foxes, and large birds
of prey, have the most influence on the population of
T. senex
. Owl attacks are also a threat, although not very common because these chipmunks
are usually diurnal, whereas owls are usually nocturnal.
- Anti-predator Adaptations
- cryptic
Ecosystem Roles
Most small rodents, including the
T. senex
, play important roles in their ecosystem. Besides being an abundant food source for
their predators, they are also an important biotic vector in seed dispersal. They
may provide habitat for certain parisites.
- Ecosystem Impact
- disperses seeds
Economic Importance for Humans: Positive
Direct impact of this species on humans is limited.
Tamias senex
may have an indirect economic importance to humans as a result of their seed dispersal.
Economic Importance for Humans: Negative
When
T. senex
populations become too dense, this species may become a pest. Chipmunks can damage
pine seeds, preventing reforestation. This may cause negative economic consequences
to humans.
Conservation Status
Although the range of this species is not large, these chimpmunks are not thought to be at great risk.
Other Comments
The genus name
Tamias
is derived from the Greek word
tamias
, meaning steward. The species name
senex
means dim or shadowy, in Latin. Common names for
T. senex
include Allen's chipmunk, gray chipmunk, California chipmunk, Was-La of the Klamath,
large mountain chipmunk, and shadow chipmunk. Shadow chipmunk was most recently used
by Jameson and Peeters (1988), referring to the dark coloration of the pelage and
the preference of this species for densely shaded areas of the forest.
Additional Links
Contributors
Marie Winans (author), California State Polytechnic University, Pomona, Nancy Avalos (author), California State Polytechnic University, Pomona, John Demboski (editor, instructor), California State Polytechnic University, Pomona.
Nancy Shefferly (editor), Animal Diversity Web.
- 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.
- 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.
- chaparral
-
Found in coastal areas between 30 and 40 degrees latitude, in areas with a Mediterranean climate. Vegetation is dominated by stands of dense, spiny shrubs with tough (hard or waxy) evergreen leaves. May be maintained by periodic fire. In South America it includes the scrub ecotone between forest and paramo.
- forest
-
forest biomes are dominated by trees, otherwise forest biomes can vary widely in amount of precipitation and seasonality.
- 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.
- 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
- fertilization
-
union of egg and spermatozoan
- 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.
- arboreal
-
Referring to an animal that lives in trees; tree-climbing.
- diurnal
-
- active during the day, 2. lasting for one day.
- motile
-
having the capacity to move from one place to another.
- 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.
- social
-
associates with others of its species; forms social groups.
- visual
-
uses sight to communicate
- tactile
-
uses touch to communicate
- acoustic
-
uses sound to communicate
- visual
-
uses sight to communicate
- tactile
-
uses touch to communicate
- acoustic
-
uses sound to communicate
- chemical
-
uses smells or other chemicals to communicate
- 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.
- herbivore
-
An animal that eats mainly plants or parts of plants.
- granivore
-
an animal that mainly eats seeds
References
Gannon, W., R. Forbes. 1995. Tamias senex. Mammalian Species , 502: 1-6.
Hartson, T. 1999. Squirrels of the West . New York: Lone Pine Field Guide.
Harvey, T., C. Polite. 2005. "Department of Fish and Game" (On-line). Accessed November 01, 2005 at http://www.dfg.ca.gov .
Nowak, R. 1999. Walker's Mammals of the World, Sixth Edition . Baltimore and London: The Johns Hopkins University Press.
Owca, D. 2005. "www.chipmunks-chipmunks.com" (On-line). Accessed December 04, 2005 at http://www.chipmunks-chipmunks.com/chipmunks-management.htm .
Stewart, A., R. Elis, C. Krueger, A. Pertschuck. 2005. "National Wildlife Federation" (On-line). Accessed November 04, 2005 at http://enature.com/home/ .
Sutton, D., B. Patterson. 2000. Geographic variation of the western chipmunks Tamias senex and T. siskiyou with two new subspecies from California. Journal of Mammalogy , 81/2: 299-316.
Verts, B., L. Carraway. 1998. Land Mammals of Oregon . Canada: The Regents of the University of California.
Wilson, D., S. Ruff. 1999. The Smithsonian Book of North American Mammals . Canada: UBC Press.