Geographic Range
Microtus californicus
occurs along the Pacific Coast of North America, from central Oregon southward to
northern Baja California. It occurs in the woodlands, shrublands and grasslands of
these areas.
Habitat
California voles inhabit areas of broad-leaved chaparral, oak woodlands, and grasslands along the Pacific Coast in northern Baja California to central Oregon. This species has a restricted distribution, which is possibly due to relic populations. It seems to utilize unusual habitats in California compared to other species of voles throughout the North American continent. Marshy ground, saltwater and freshwater locations, wet meadows, coastal wetlands and dry, grassy hillsides are the preferred macrohabitats of this species.
California voles are semifossorial. Their microhabitat consists of burrows, grass
runways, and earth tunnels where piles of grass cuttings and fresh vegetation are
often found. Piles of feces are also found in the runways.
- Habitat Regions
- temperate
- terrestrial
- Terrestrial Biomes
- desert or dune
- savanna or grassland
- chaparral
- Wetlands
- marsh
- Other Habitat Features
- agricultural
- riparian
Physical Description
M. californicus present a lot of variation in size depending upon where they are found. Subspecies found in the south of the species range can be much larger than those found in the north.
In northern California, the total length veries between 139 ans 207 mm, of which only 39 to 68 mm is contributed by the tail. Males weigh between 33 and 81 g, averaging 52 g, and females can weigh between 30 and 68 g, averaging 47 g.
California voles are sexually dimorphic with the males being six percent longer and eleven percent heavier than females.
The coat of these animals is buffy brown, grayish brown or dark brown (blackish toward coast, reddish in desert) colored on top, with a reddish tinge down the middle of the back. The underside is blue-gray to white. The tail is bi-colored. The feet are pale, and the eyes are dark brown to black. M. californicus has 8 mammae.
This species can be distinguished from other voles by the following characteristics:
Microtus montanus occurs at higher elevations;
M. longicaudus is longer and its tail more bicolored;
M. townsendii has different cranial and dental features; and
M. oregoni
is smaller and has only 5 toe pads.
- Other Physical Features
- endothermic
- bilateral symmetry
- Sexual Dimorphism
- male larger
Reproduction
California voles seem mostly monogomous when populations aren't too dense. Dense populations or populations with unbalanced sex ratios will display polygynous traits. In these populations, males defend territories where grass is the staple diet, and females defend areas where fruits and forbs are the primary food source.
When polygyny is the prevailing mating system, females tend to have neighboring territories with their sisters and dispersing males may have little or no contact with close relatives.
Suppression of sexual maturation on the natal home range by the presence of the mother occurs. Kin recognition has little influence on inhibiting inbreeding, even in monogamous populations.
Adult males will cannibalize young that are not theirs. Females will also abort their
litters if exposed to the phermones produced by unfamiliar males.
- Mating System
- monogamous
- polygynous
Ovulation in M. californicus is induced by copulation. This species experiences a post-partum estrus, and breeding can occur within fifteen hours after young are born. This allows up to 4 or 5 litters a season.
The breeding season is somewhat variable throughout the range of this species. Germination of annual grasses in the fall seems to initiate reproduction in some popultions. In other populations, where the climate is very Mediterranean, reproduction begins near the time of the first rains, and ends when the hot summer dries out the vegetation. In coastal populations, where the grasses stay green all year, and temperatures are mild, breeding can occur thorughout the year.
The gestation is twenty-two days. Litters of 1 to 11 young can be roduced, but the average is 4 or 5 young. Perennial grassland populations average about two embryos less than those in annual and mixed annual-perennial grassland populations. Young are altricial and weigh approximately 2.5g at birth. The pups are quickly weaned at two weeks old.
Females reach reproductive maturity by three weeks old, and the male at five weeks. Under some conditions, males can mature more rapidly, reaching sexual maturity by about 25 days of age. Sexual maturity can be supressed by the presence of the parents on the natal range. This allows the species to rapidly colonize when population densities are low, but to limit reproduction somewhat when populations are very dense.
Microtus
species in general, appear to have a fairly plastic reproductive biology where the
emphasis seems to be on the ability to produce the correct response (phenotypic) given
the ecological conditions in which the population finds itself. Populations experience
cyclic and annual fluctuations. The population typically grows for three to four
years then declines rapidly in mild, temperate areas. In strongly seasonal habitats,
the growth phase is two to five years.
- Key Reproductive Features
- iteroparous
- seasonal breeding
- year-round breeding
- gonochoric/gonochoristic/dioecious (sexes separate)
- sexual
- induced ovulation
- fertilization
- viviparous
- post-partum estrous
As in all mammals, the female provides the young with milk for the duration of nursing. Although the young are altricial at birth, they grow quickly. Weaning occurs when the young are abut two weeks old.
In polygynous systems, females are the primary care givers, with neighboring territories being occupied by their sisters. In monogamous systems, males will participate in brood care. In these mating systems, males participate in parental care by gathering nesting materials and retreiving nestlings. Both males and females will display territorial traits, which helps to protect the young from intruders.
Nests are made with dried grasses and forbs and are located under logs or boards or
under the earth's surface a few centimeters.
- Parental Investment
- altricial
-
pre-hatching/birth
-
provisioning
- female
-
protecting
- female
-
provisioning
-
pre-weaning/fledging
-
provisioning
- female
-
protecting
- male
- female
-
provisioning
Lifespan/Longevity
A successful lifespan for a vole will be up to a year, but the average lifespan is
only a few months.
Behavior
M californicus
is most active near dawn and dusk with short bursts of activity every few hours in
between. Their crepuscular pattern is most obvious during the long, hot, dry days
of summer. These animals are actove throughout the year. They have no hibernation
and are not known to store food. California voles are a social species. Their runways
are interconnecting, but individuals are territorial, especially during the breeding
season.
- Key Behaviors
- crepuscular
- motile
- sedentary
- territorial
- social
Home Range
Adult males have larger home ranges (125 sq meters) than females (80 sq meters), but
their core area (where they spend 85% of their time) is about the same.
Communication and Perception
California voles communicate mostly through scent by depositing urine in areas of its runways as a tracking device. This tells the vole where it has been and who else inhabits a runway. Squeaks are also heard from adults during distress and young when communicating to the parent(s).
As with most other mammals, there is likely some visual communication as well.
- Other Communication Modes
- scent marks
Food Habits
M. californicus
is herbvivorous and eat mostly grasses and roots, but also relies on sedges, fruits
and forbs in certain areas. In the winter, the vole eats mostly roots and underground
plant parts. Grain will also be eaten when available.
- Primary Diet
- herbivore
- coprophage
- Plant Foods
- leaves
- seeds, grains, and nuts
- fruit
Predation
Voles spend as little time exposed to the surface as possible. Underground tunnels are commonly used as are runways throughout tall grasslands. Unfortunately for the voles, the urine they use to communicate to each other is the way their diurnial predators track them and determine vole density. Through the raptors ability to see ultraviolet light, the florescent urine shows up in the runways and directs the predator to the prey's location. This could also explain how raptors are able to locate their vole prey even after population crashes.
There are a great number of vole predators, including coyotes, kestrels, hawks, weasles,
kits, owls, snakes, herons, egrets, and ferrel cats. Because of their rapid reproduction
and periody high population densities, these voles are a keystone prey species.
Ecosystem Roles
With their potential to reproduce rapidly, these voles are prey for many carnivore
species. They form an important link in food webs. these voles also are hosts to
many species of parasites.
- Ecosystem Impact
- keystone species
- Cestodes
- Nematodes
- Dermacentor ticks
- Ixodes ticks
- Hoplopleura lice
- Polyplax lice
- Fleas of 24 different species are known to use these voles as hosts
Economic Importance for Humans: Positive
Although there is apparently no direct benefit of this species for humans, it should be noted that without this keystone species, many of the larger animals that people enjoy watching, such as hawks, kestrels, coyotes and foxes, would not be able to exist at such high densities, and therefore would be a much less visible part of local ecosystems.
Economic Importance for Humans: Negative
During peak population times, numbers of voles are said to exceed to the hundreds
per acre, and up to a thousand per hectare, causing crop problems in areas where farms
coincide with vole habitat.
- Negative Impacts
- crop pest
Conservation Status
These voles are quite common and so are not a big conservation concern. Local abundances
can vary from year to year, but overall the population cycle seems stable. However,
because thes voles are so important to their ecosystem, it is worthwile to keep an
eye on their population cycles and habitat availability, so that other species which
depend upon them for food, and whose population status may be less secure, are safeguarded.
Other Comments
Fossil Record: The size of the incisive foramina is used to help distinguish California voles from other species. In M. californicus , the foramina is wider and unconstricted, and the teeth are slightly larger than other species of vole that occur near California. It is possible M. californicus could have first appeared 1.8 million years before present in North America at the beginning of the Pleistocene, making it possibly one the first occurring in the Microtus .
Humboldtian (Northwestern California), Diablian (Central California), Californian
(Valley and Eastern California), and San Bernardinian (Southern California) provinces
exist in California. There are seventeen subspecies of
M. californicus
that occur in its region.
Additional Links
Contributors
Nancy Shefferly (editor), Animal Diversity Web.
Lisa Peronne (author), Humboldt State University, Brian Arbogast (editor), Humboldt State University.
- 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.
- 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.
- 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.
- marsh
-
marshes are wetland areas often dominated by grasses and reeds.
- agricultural
-
living in landscapes dominated by human agriculture.
- riparian
-
Referring to something living or located adjacent to a waterbody (usually, but not always, a river or stream).
- monogamous
-
Having one mate at a time.
- polygynous
-
having more than one female as a mate at one time
- 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
- year-round breeding
-
breeding takes place throughout the year
- sexual
-
reproduction that includes combining the genetic contribution of two individuals, a male and a female
- induced ovulation
-
ovulation is stimulated by the act of copulation (does not occur spontaneously)
- 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.
- crepuscular
-
active at dawn and dusk
- motile
-
having the capacity to move from one place to another.
- sedentary
-
remains in the same area
- 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.
- visual
-
uses sight to communicate
- tactile
-
uses touch to communicate
- acoustic
-
uses sound to communicate
- chemical
-
uses smells or other chemicals to communicate
- 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
- 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).
- 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
- coprophage
-
an animal that mainly eats the dung of other animals
- 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.
- 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.
References
Burt, W., R. Grossenheider. 1980. Peterson Field Guide: Mammals . New York: Houghton Mifflin Co..
Gee, H. February 2, 1995. In the eye of the kestrel. Nature , 373: 387.
Ingles, L. 1965. Mammals of the Pacific States (CA, OR, WA) . Stanford California: Stanford University Press.
Krohne, D. 1982. The Bases of Intra- and Interpopulational Reproductive Variation and their Demographic Consequences in the California Vole . Ann Arbor, Michigan: University Microfilms International.
Tamarin, R., R. Ostfeld, S. Pugh, G. Bujalska. 1990. Social Systems and Population Cycles in Voles . Boston: Birkhauser Verlag.
Tamarin, R. 1985. Biology of New World Microtus . Shippensburg, Pennsylvania: Special Publication No.8 American Society of Mammalogists.
Whitaker, Jr., J. 1998. National Audobon Society, Field Guide to Mammals . New York: Chanticleer Press.
Wilson, D., S. Ruff. 1999. The Smithsonian book of North American mammals . Washington, D.C.: Smithsonian Institution Press.
Zim, H., D. Hoffmeister. 1955. Golden Guide: Mammals . New York: Golden Press.