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Lagurus lagurussteppe lemming

By Adam Rountrey

Geographic Range

Lagurus lagurus is found on the steppes of Eurasia from the Dnieper River of Ukraine in the West to the Yenisey River northeast of Kazakhstan in the East. The range extends through Kazakhstan into northwest China and northern Mongolia while reaching its southern limit at the Tien Shan Range. The northern limit of the range occurs at about 55ºN. In the Pleistocene the range of the genus Lagurus extended west to France and Britain.

There are four subspecies of steppe lemming. Lagurus lagurus lagurus is found in Ukraine and central Kazakhstan. The eastern subspecies, L.l. altorum, is found in southeastern Kazakhstan, northwest China, and Mongolia. Lagurus lagurus agressus, the northern subspecies, is found in the northern European portion of the former U.S.S.R. and in northern Kazakhstan. The isolated subspecies L. l. abacanicus is found in the Minusinsk basin in Krasnoyarsk, Russia. (Gromov and Polyakov, 1992; Ognev, 1964)

Habitat

Steppe areas are regions with sufficient rainfall to support grasses, but too little moisture to support forest growth. Lagurus lagurus inhabits Eurasian steppes, feeding on grasses and herbs, and depending on the cover these provide to avoid predation. Steppe lemmings are found in particular abundance in areas of mixed feather grass and sheep’s fescue, but also in growths of wormwood (Artemesia). When feather grass dominates the flora, lemmings tend to be found in places lacking vegetation, such as in rocky areas. In general they prefer loose soils that allow for easier burrowing.

When living near agricultural lands, individuals often take up residence in fields and, in some cases, they have been known to live in storage buildings. However, as plowing damages their burrows, they must frequently make temporary moves away from the cultivated fields. This requirement leads to relatively low populations in agricultural areas. (Gromov and Polyakov, 1992; Ognev, 1964)

  • Range elevation
    4000 (high) m
    13123.36 (high) ft

Physical Description

Steppe lemmings are relatively small Arvicolines having masses that vary seasonally from an average of 22 g in the winter to 34.8 g in the summer. The combined length of the head and body (for adults) ranges between 80 to 120 mm. Females are slightly larger than males. Lagurus can be distinguished from Lemmiscus by possession of a tail that is shorter than the hind foot, with lengths ranging from 7 to 19 mm. The eyes are small, and the pinnae are short, rounded, and lack an antitragus. The dorsal pelage is gray to gray-brown with a central black stripe running longitudinally, while the ventral pelage is light in color. Sexual dimorphism in size or color is uncommon in Arvicolinae. The pedes are almost completely covered in fur and have four pads. Females have eight teats.

Diagnostic molar characteristics include five closed enamel loops in M3, with the elongated posterior loop angled labially, and an M1 with seven to eight closed enamel loops. The skull is flat in profile and the zygomatic arches are wide and heavily built. The sagittal crest is absent.

Geographic variation includes trends of increasing body size and narrowing of the dorsal stripe from west to east. Individuals also tend to have paler fur to the east and south. More specific differences between the subspecies include:

L. l. lagurus: Narrow dark brown spinal stripe; ventral fur gray to yellow; tail white; winter fur more yellow than summer fur.

L. l. agressus: more prominent black spinal stripe; darkest fur; tail fur darker in color than L. l. lagurus.

L. l. abacanicus: larger skull than others; larger bullae; fur more gray than L. l. lagurus; very short tail.

L. l. altorum: lightest fur color (pale yellow); spinal stripe dark. (Gromov and Polyakov, 1992; Krebs, 2001; Longrove, 2005; Nowak, 1999; Ognev, 1964; Ralls, 1976)

  • Sexual Dimorphism
  • female larger
  • Range mass
    22.0 to 34.8 g
    0.78 to 1.23 oz
  • Range length
    80 to 120 mm
    3.15 to 4.72 in

Reproduction

Within the Arvicolinae, males are usually promiscuous, though some species are monogamous. Male and female steppe lemmings are only found living together in shared burrows before the birth of young. Following this event, the male will move to another burrow and avoid involvement in raising the young. Some female arvicolines have a postpartum estrus allowing mating to occur following parturition. This, in combination with early sexual maturation and short gestation periods, leads to the high reproductive potential of arvicolines. (Krebs, 2001; Ognev, 1964)

Steppe lemmings breed primarily between the months of April and September, though winter breeding does sometimes occur. The length of the estrus cycle is seven days, and implantation takes place six or seven days following fertilization. During the normal breeding season a female will produce about six litters each consisting of three to seven young. Litters tend to be largest in the spring. The female gives birth to young weighing about 1 g each after a gestation period of approximately 20 days. Although data are unavailable for L. lagurus, weaning probably occurs after 12 to 14 days as in Lemmus lemmus. Females reach sexual maturity at 21 to 140 days. Variability in sexual maturation age is due largely to delayed maturation in those females born in late autumn. Males tend to mature slightly earlier than females, but no information is available on variability in male maturation age. (Mohi Aldeen and Finn, 1970; Nowak, 1999; Ognev, 1964; Pokrovskij, 1970; Semb-Johansson, et al., 1993)

  • Breeding interval
    Steppe lemmings breed about six times per year between April and September.
  • Breeding season
    Primarily April to September, but some mating occurs in the winter.
  • Range number of offspring
    3 to 7
  • Average number of offspring
    4.5
    AnAge
  • Average gestation period
    20 days
  • Average gestation period
    20 days
    AnAge
  • Range age at sexual or reproductive maturity (female)
    21 to 140 days
  • Average age at sexual or reproductive maturity (male)
    4 to 6 weeks
  • Average age at sexual or reproductive maturity (male)
    Sex: male
    44 days
    AnAge

Arvicoline young are altricial, and they are cared for by the female in a nest area, which, in the case of steppe lemmings, is located within the burrow. The nest is commonly located at a depth of 25 to 30 cm below the surface. It is spherical in form and lined with soft grasses. Female steppe lemmings are known for their aggressive nature in protecting young. Males take no part in caring for the young. (Krebs, 2001; Ognev, 1964)

  • Parental Investment
  • altricial
  • pre-fertilization
    • provisioning
    • protecting
      • female
  • pre-hatching/birth
    • provisioning
      • female
    • protecting
      • female
  • pre-weaning/fledging
    • provisioning
      • female
    • protecting
      • female

Lifespan/Longevity

Arvicoline lifespans are approximately 0.5 to 2.0 years. One source lists the maximum longevity for L. lagurus as 3.8 years. Most probably live no longer than one year. (de Magalhães, 2006; Krebs, 2001)

  • Range lifespan
    Status: captivity
    3.8 (high) years
  • Average lifespan
    Status: captivity
    3.8 years
    AnAge

Behavior

Steppe lemmings are burrowers that dig two types of burrows, domicile burrows and temporary burrows. The domicile burrow commonly has two to three openings, but only one distinct main entrance surrounded by a mound of raised earth. The main passage heads down at an angle of 30 to 40 degrees to a depth of up to 90 cm. A spherical nest area, located in the main passage, varies in size from about eight to 12 cm in diameter. There are also numerous blind ended passages, used when the animal is threatened, that extend up from the main burrow to within several centimeters of the surface. Separate, shallower, temporary burrows located within three meters of the main burrow are also used in times of danger.

This species is primarily crepuscular (active in the late evening and early morning). It has also been characterized as nocturnal, though it sometimes remains active through the day. Male and female Arvicolines typically maintain territories, which are marked using glandular secretions. Females engage in competition for resource-rich territories, while competition in males is focused on gaining access to females.

Lemmings are famous for their mass migrations. Steppe lemmings, though they are not true lemmings, also participate in mass migrations. Smaller scale migrations often occur locally as populations move from one food source to another due to seasonal changes in availability. However, mass migrations can occur in years when population densities rise too high due to mild autumns that allow continued breeding, or due to a lack of predators. (Krebs, 2001; Nowak, 1999; Ognev, 1964)

Home Range

A steppe lemming usually stays within six meters of its burrow. It will deplete the food supply around the burrow, then begin to maintain peripheral, temporary burrows that allow it to forage in more distant areas.

A study of the spatial distribution of L. lagurus revealed differences in distribution when population densities differed. In the spring, when population densities were low (6.1 individuals per hectare), L. lagurus lived in territorial family groups consisting of about six individuals. Each family group occupied an average territory of 0.2 hectares, and was separated from other groups by 73.3 m. In the autumn, when population densities were higher (34.0 individuals per hectare), no family groups could be distinguished. (Formozov, 1996; Mal'kova, et al., 2004)

Communication and Perception

Arvicolines communcate via scent marking and vocalization. Scent marking is used to delineate territory and establish dominance. Calls are used for various social interactions including warning others of danger and making threats. Female steppe lemmings are particularly vocal just prior to giving birth. (Krebs, 2001; Ognev, 1964)

Food Habits

Lagurus lagurus is an herbivore feeding primarily on the above ground portions of plants. Steppe lemmings have a preference for Austrian wormwood (Artemisia austriaca), but will also take herbs, grasses, lichens, roots, tubers, and bulbs. The average daily food intake is 10.7 g in the summer. During this time only the best portions of the plant are eaten. The remains are left out or packed into the burrow and will serve to provide the animal with nourishment when other food is scarce. Steppe lemmings are able to process large amounts of abrasive vegetation because they possess continuously growing molars (and incisors). (Krebs, 2001; Ognev, 1964)

  • Plant Foods
  • leaves
  • roots and tubers
  • lichens

Predation

Steppe lemmings are prey for many predatory mammals and birds including: steppe polecats, ermines, weasels, foxes, domestic cats, eagle owls, buzzards, harriers, and kestrels. They avoid predation by lying flat on the substrate, staying under the cover of vegetation, entering their burrows, or by hiding in natural crevices or temporary burrows. (Formozov, 1996; Ognev, 1964)

  • Anti-predator Adaptations
  • cryptic

Ecosystem Roles

Steppe lemmings are an important prey base for many carnivores of the steppe. In addition, their presence in an area can limit the success of other voles.

When population densities are high, the presence of steppe lemmings can be destructive to the ecosystem. In some cases they will reduce vegetation such that only hardy plants such as wormwood remain. In extreme situations where predation is limited, L. lagurus can transform steppe to semi-desert. (Ognev, 1964)

Commensal/Parasitic Species
  • fleas (Siphonaptera)
  • plague (Yersinia pestis)
  • rabbit fever (Francisella tularensis)

Economic Importance for Humans: Positive

This species is used in research related to cancer and exposure to toxic chemicals. Steppe lemmings are also kept as pets. (Bolonina, 1960; Dryden, 1979)

  • Positive Impacts
  • pet trade
  • research and education

Economic Importance for Humans: Negative

Steppe lemmings carry diseases such as tularemia (Francisella tularensis) and plague (Yersinia pestis). Incidence of these diseases is known to increase during years of high vole populations. In addition, steppe lemmings can destroy crops and pastures, leading to loss of income and cattle. (Ognev, 1964)

  • Negative Impacts
  • injures humans
    • carries human disease
  • crop pest

Conservation Status

This species is not threatened or endangered.

Contributors

Tanya Dewey (editor), Animal Diversity Web.

Adam Rountrey (author), University of Michigan-Ann Arbor, Phil Myers (editor, instructor), Museum of Zoology, University of Michigan-Ann Arbor.

Glossary

Palearctic

living in the northern part of the Old World. In otherwords, Europe and Asia and northern Africa.

World Map

acoustic

uses sound to communicate

agricultural

living in landscapes dominated by human agriculture.

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.

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.

chemical

uses smells or other chemicals to communicate

crepuscular

active at dawn and dusk

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.

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.

folivore

an animal that mainly eats leaves.

fossorial

Referring to a burrowing life-style or behavior, specialized for digging or burrowing.

herbivore

An animal that eats mainly plants or parts of plants.

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).

migratory

makes seasonal movements between breeding and wintering grounds

motile

having the capacity to move from one place to another.

native range

the area in which the animal is naturally found, the region in which it is endemic.

pet trade

the business of buying and selling animals for people to keep in their homes as pets.

polygynandrous

the kind of polygamy in which a female pairs with several males, each of which also pairs with several different females.

scent marks

communicates by producing scents from special gland(s) and placing them on a surface whether others can smell or taste them

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

social

associates with others of its species; forms social groups.

stores or caches food

places a food item in a special place to be eaten later. Also called "hoarding"

tactile

uses touch to communicate

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.

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

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.

visual

uses sight to communicate

viviparous

reproduction in which fertilization and development take place within the female body and the developing embryo derives nourishment from the female.

year-round breeding

breeding takes place throughout the year

References

Bolonina, N. 1960. Tumors induced by methylcholanthrene in the steppe lemming. A transmissible strain of lemming sarcoma (LS). Bulletin of Experimental Biology and Medicine, 49(2): 184-186.

Dryden, G. 1979. Sensitivity of European microtines to injected CdCl2. Acta Physiologica Acadamiae Scientarum Hungaricae, 53(3): 355-362.

Ferkin, M. 1999. Meadow Voles (Microtus pennsylvanicus, Arvicolidae) Over-mark and Adjacent-mark the Scent Marks of Same-sex Conspecifics. Ethology, 105: 825-837.

Formozov, A. 1996. Adaptive Modifications of Behavior in Mammals of the Eurasian Steppes. Journal of Mammalogy, 47(2): 208-223.

Gromov, I., I. Polyakov. 1992. Fauna of the USSR: Mammals Volume III, No. 8: Voles (Microtinae). Washington, D.C.: Smithsonian Institution Libraries and the National Science Foundation.

Krebs, C. 2001. Voles and Lemmings. Pp. 628-633 in D Macdonald, S Norris, eds. The New Encyclopedia of Mammals. Oxford: Oxford University Press.

Longrove, B. 2005. Seasonal thermoregulatory responses in mammals. Journal of Comparative Physiology B, 175: 231-247.

Mal'kova, M., N. Pal’chekh, V. Yakimenko, L. Kuz’min. 2004. The Spatiotemporal Structure of Rodent Populations in the Steppe Zone of Western Siberia. Russian Journal of Ecology, 35(1): 27-34.

Mohi Aldeen, K., C. Finn. 1970. The Implantation of Blastocysts in the Russian Steppe Lemming (Lagurus lagurus). Journal of Experimental Zoology, 173(1): 63-77.

Nowak, R. 1999. Walker's Mammals of the World 6th Ed. Vol II. Baltimore: Johns Hopkins University Press.

Ognev, S. 1964. Mammals of the U.S.S.R and Adjacent Countries: Mammals of Eastern Europe and Northern Asia Vol. VII Rodents. Jerusalem: Isreal Program for Scientific Translations.

Oku, Y., J. Wei, J. Chai, I. Osman, J. Wei, L. Liao, M. Asakawa, K. Hagiwara, K. Kobayashi, M. Ito. 2002. Meriones meridianus and Lagurus lagurua as alternative definitive hosts of Echinococcus multilocularis and E. granulosus. Experimental Animals / Japanese Association for Laboratory Animal Science, 51(1): 27-32.

Pokrovskij, A. 1970. Seasonal changes in biological cycles in some rodents and the problem of absolute age determination. Annales Zoologici Fennici, Vol. 8 No. 1: 94-96.

Ralls, K. 1976. Mammals in which females are larger than males. The Quarterly Review of Biology, 51(2): 245-276.

Semb-Johansson, A., C. Engh, E. Ostbye. 1993. Reproduction, litter size and survival in a laboratory strain of the Norwegian lemming (Lemmus lemmus). Pp. 329-337 in N Stenseth, R Ims, eds. The Biology of Lemmings, Vol. Linnean Society Symposium Series No. 15. London: Academic Press: Harcourt Brace & Co..

de Magalhães, J. 2006. "Human Ageing Genomic Resources" (On-line). Accessed March 19, 2006 at http://genomics.senescence.info/species/entry.php?species=Lagurus_lagurus.

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