Birch mice, jumping mice, and jerboas

(Dipodidae)

Class Mammalia

Order Rodentia

Suborder Sciurognathi

Family Dipodidae

Thumbnail description
Pygmy to medium-sized quadrupedal or bipedal rodents with hind limbs slightly or much longer than forelimbs; long tails and eyes and external ears ranging in size; coarse or soft brown, brownish-yellow or jonquil fur; a dichromatic, black-and-white banner tail tip characteristic of many jerboa species specialized for the fast bipedal gallop

Size
1.8–9 in (45–230 mm); weight 0.2–15 oz (6–415 g)

Number of genera, species:
15 genera, 49–51 species

Habitat
Inhabitants of forests, meadows, steppe and desert plains and mountain regions

Conservation status
Endangered: 4 species; Vulnerable: 4 species; Lower Risk/Near Threatened: 6 species; Data Deficient: 4 species

Distribution
North America, Northern Africa, Arabian Peninsula, Europe, and Asia

Evolution and systematics

Traditional classifications based mostly on morphological similarities consider two related families: Dipodidae, including all jerboas, and Zapodidae, including jumping mice and birch mice. The family of jerboas include three subfamilies (Euchoreutinae, Cardiocraniinae, and Dipodinae), and the family of jumping and birch mice include two subfamilies (Zapodinae and Smintinae = Sicistinae). Vorontsov et al. in 1971 divided jerboas into four subfamilies (Euchoreutinae, Cardiocraniinae, Allactaginae, and Dipodinae), and Gambarianin in 1983 divided them into three families (Euchoreutidae, Cardiocraniidae, and Dipodidae with two subfamilies, Allactaginae and Dipodinae). The recent revision of plylogenetic relationships of all groups of jerboas, jumping mice, and birch mice, based on palaeontological data and complex morphological comparisons, suggests one superfamily Dipodoidea with four families: Zapodidae, Smintidae (with subfamilies Euchoreutinae and Smintinae), Dipodidae (with subfamilies Cardiocraniinae, Paradipodinae, and Dipodinae), and Allactagidae. The position of Euchoreutes is not clearly defined, and there are reasonable arguments to consider long-eared jerboa to represent a separate family. Considering Dipodidae as an extended interpretation, one must be aware of long-term independent evolution of jumping mice, birch mice, and at least three lineages of jerboas. Formal resemblance of long-eared jerboa (subfamily Euchoreutinae), five-toed jerboas (subfamily Allactaginae), and three-toed jerboas (subfamilies Cardiocraniinae, Paradipodinae, Didodinae) is considered now to be a result of parallel evolution of bipedality in arid open and flat landscapes. Among three-toed jerboas very deep differences, signifying early divergence, exist between pygmy jerboas (Cardiocraniinae), genus Paradipus, and true three-toed jerboas (Dipodinae). Fossil remains of Dipodidae have been discovered from the Eocene in North America and from the Oligocene in Eurasia. Three-toed jerboa fossils from the middle Miocene were discovered in Asia, and five-toed jerboa fossils from the middle Miocene were unearthed in North Africa as well as upper Miocene specimens from Europe and Asia.

Physical characteristics

Birch mice and jumping mice are small, mouse-like quadrupedal rodents with fairly long semi-prehensile tails. Jumping mice hind limbs are moderately elongated, whereas birch mice have no elongation. Jerboas are small to medium-sized bipedal, nocturnal rodents adapted to run fast in sparse vegetation. Both birch and jumping mice have relatively small and narrow heads, and jerboas have relatively large heads with wide muzzles and flat snouts. Jerboas also have big eyes, long vibrissa, and auricles that vary in size from relatively short among three-toed jerboas to extremely long in long-eared jerboa

(Euchoreutes naso). In pygmy jerboas (Cardiocraniinae), the auricle is tubular. Among jerboas, the neck is weakly pronounced or not pronounced at all; the neck vertebrae are fused in the Cardiocraniinae, Paradipodinae, and Dipodinae, but unfused in the other subfamilies.

All jerboas have short and compact bodies with short forelimbs and long strong hind limbs. The tail is usually long, ending as a rule, but not always, with the dichromatic black-and-white banner. The brush of long hairs may resemble a bird feather. Among pygmy jerboas (Cardiocraniinae) and in Pygeretmus platyurus, a specialized representative of five-toed jerboas, the tail is relatively short, bannerless, and fat. Forelimbs have five fingers. Claws are short and blunt in birch mice, jumping mice, and five-toed jerboas, or long and narrow in three-toed jerboas. Hind feet have five, four, or three toes. If toes are more than three, the first and the fifth toes are much shorter than three middle ones. In Allactaga tetradactyla the first toe is absent. The three central bones of the hind foot are fused to form a single cannon bone among all fast running jerboas (except in Cardiocraniinae). The pelage is coarse in the Smintinae and Zapodinae with light or dark brown or brownish yellow back hair and paler, lighter brown hair on the ventrum. Jerboas have dense, soft earthen-colored fur with lighter, sand-colored fur on the back and white on the belly. The dental formula of the Dipodidae is (I1/1 C0/0 P0–1/0 M3/3) × 2 = 16 or 18. The upper incisors are weakly grooved in the Zapodinae, but smooth in the other subfamilies. Premolar is intent for reduction. Molars are either low-(Smintinae) or high-crowned (other subfamilies) and have adult roots.

Distribution

Thirteen birch mice species are endemic to Eurasia (from Austria in Europe to Sakhalin island in the Far East). Five species inhabit mostly plains. All others are present in mountain terrains (Caucasus, Altai, Tien Shan, mountains of Central China, Pakistan, Kashmir). Jumping mice (genus Eozapus) occur in Central China (Quinghai, southern Gansu, Ningxia, Shaanxi, western Sichuan, northwestern Yunan provinces). Another two genera of jumping mice, Zapus (Z. hudsonius, Z. princeps, Z. trinotatus) inhabit North America from southern Alaska to southwestern United States, and Napaeozapus (N. insignis) from southeastern Manitoba to Labrador and Pennsylvania and south along Appalachian Mountains to northern Georgia. Jerboas are confined to open semi-arid and arid lands of the Palaeoarctic region. The range covers North Africa, Eastern Europe, Central, middle, and southwestern Asia, including Arabia.

Their origin is considered connected with Asia where they demonstrate the greatest species and genera diversity. By the middle Miocene, jerboas penetrated Africa but did not diverge. Ranges of nine genera are restricted to Eurasia and ranges of two genera are distributed in arid lands of both Eurasian and North African continents. Euchoreutinae and Cardiocraniinae are strictly Asian groups. Asia accounts for 30–31 species; Europe five species; Africa three species. Two centers of species diversity of jerboas in Asia, one connected with deserts of Middle Asia (Uzbekistan, Turkmenistan, and southwestern and southern portion of Kazakhstan), and the other with deserts of Mongolia and China. Euchoreutes naso, Salpingotus kozlovi, Stylodipus andrewsi, Allactaga bullata, and A. balicunica are endemic to Mongolian-Chinese deserts. Salpingotus heptneri, Paradipus ctenodactylus, Eremodipus lichtensteini, Allactodipus bobrinskii, and Allactaga severtzovi are endemic to desert middle Asia. Pygeretmus shitkovi and P. platyurus are endemic to cold deserts of Kazakhtan. Species diversity in Asia decreases from subzone of cold deserts to the south (subtropical deserts) and to the north (steppe). Jerboas did not cross the Indus River. Accordingly, there are no jerboas in the Thar Desert of India. Three species of jerboas (Allactaga tetradactyla, Jaculus jaculus, and J. orientalis)

are present in North Africa. Genus Jaculus are also common in deserts located in the Arabian Peninsula, Middle East, Iran, Iraq, and Afganistan.

Habitat

Birch mice inhabit forests, thickets, moors, meadows, and steppe. In the mountain regions, they occur in the forest and subalpine belts. Jumping mice (genus Zapus) inhabit wooded areas, grassy fields, and alpine meadows. They are common in the thick vegetation bordering streams, ponds, and marshes. The woodland jumping mouse (N. insignis) inhabits sprucefir and hemlock-hardwood forests where it prefers moist, cool places with dense vegetation. It is common along streams in bogs and swamps. The Chinese jumping mouse (E. setchuanus) is found near stream banks in cool forests at high mountain elevations of 9,840-–13,120 ft (3,000–4,000 m). Jerboas occupy all type of desert habitats from moving sands to clay depressions and rocky-gravel plateau and dry mountain slopes. Up to five species can be seen in one habitat, and the local fauna can account for eight species.

Many species are habitat specific while others are less selective. Some selective species include Paradipus ctenodactylus that inhabits top ridges of moving sands (barkhans) in the deserts of Turkmenistan and Uzbekistan; Allactodipus bobrinskii that inhabits clay and rocky-clay plains covered with the thin sandy-gravel wash; and Pygeretmus pumilio, an inhabitant of clay depressions with hard surface covered by sparse vegetation of succulents. In general, sandy habitats are more common for tree-toed jerboas, whereas five-toed jerboas usually inhabit areas with hard clay soils. Jerboas that are adapted for locomotion on soft sandy substrate have brushes of long hard hairs on sides of their toes to stabilize the foot on the soft sand. When the three-toed jerboas run, the distal part (all toes) of their hind limbs contacts the substrate surface. In contrast, when five-toed jerboas run, they touch the hard substrate only by the tall compact callus on top of the central toe. The second and the fourth toes are shorter and mostly act as shock absorbers (bumpers).

Behavior

Locomotion

Birch mice and jumping mice are quadrupeds. During ground travel, they often climb tussock grass, shrubs, and bushes, curling the tail for additional support around branches and twigs. Jumping mice hop up to 3.3 ft (1 m) high and 6 ft (1.8 m) long. They are also a good swimmers (Zapus, Napaeozapus). All jerboas are bipedal, but differ in capacity to run fast for a long distance. The best runners among such five-toed jerboas as Allactaga major or A. severtzovi, can maintain a speed of 25 mph (40 kph). Pygmy jerboas do not increase their speed over 6 mph (9 kph). Normally, the birch mice and jumping mice use a quadrupedal walk when moving slowly. When startled, however, they often take several leaps, using their powerful hind legs for propulsion; the forelimbs are used just for support when the body contacts the substrate

(quadrupedal hop). The jumping mouse can make three to four jumps without touching the substrate with their forelimbs. While the pygmy jerboa moves slowly with short hops, it also can touch the substrate with forelimbs. When it increases its speed, the hind feet work simultaneously, and the forelimbs never touch the substrate. Pygmy jerboas cannot run long distances. When chased they demonstrate cryptic behavior and stretch out on the substrate in a shadow of small shrub or hollow. They never use shelter burrows to escape danger.

When running quickly, five-toed and three-toed jerboas push off the substrate with one hind foot after another. Such non-synchronous propulsion provides additional acceleration to the bipedal gallop. Large-sized and fast species escape danger by running for a distance of hundreds of feet (meters) (Allactaga major, A. severtzovi, Paradipus ctenodactylus, Dipus sagitta). The large home range of Paradipus ctenodactylus in the Karakum desert of Turkmenistan may reach 1.2 mi (2 km) in diameter. Smaller forms with relatively weak hind feet range in a diameter often less than of 328 ft (100 m) (Pygeretmus pumilio, P. shitkovi, Stylodipus telum). When moving slowly with short jumps, jerboas are unable to keep position of the body axis stable. Shifts of the body's center of gravity force it to make deep bows after each jump. Because bipedal hopping at low speeds is considered energetically expensive, the best runners among five-toed jerboas and all three-toed jerboas use another gait for slow movements—when foraging, they walk on hind feet as humans do.

Shelters and burrowing activity

Birch mice and jumping mice rarely dig. For shelters they often use space under logs, boards, or roots, or even clumps of vegetation where they build nests of dry leaves and woven grass or moss. The birch mouse (Sicista betulina) often uses hollows in old wood. Sicista also can dig shallow burrows or use burrows built by other species. The woodland jumping mouse (Napaeozapus insignis) also can dig simple burrows, but prefers to occupy old burrows left by other rodents. All species of jerboas dig very well and construct three types of burrows: day (summer) burrows used to sleep during the day time, shelter burrows used to escape predatory attacks in the night time, and wintering burrows used for hibernation. The first and last burrow types, including day burrows where females bear their pups, are similarly constructed and differ mostly in depth and length. Five-toed jerboas dig in a hard soil with their incisors and excavate soil with the help of forelimbs, pulling soil out of the tunnel by the snout. Jerboas start with the digging of the main inclined tunnel. From a point close to the end of the main tunnel chamber it digs a second, steep and narrow tunnel up to the surface, sometimes with one to three cavities and only one outside opening. It is indistinct and usually closed by a plug when the jerboa is in the burrow. Jerboa closes the initial tunnel by the soil dug out from the second hole.

Day burrows of three-toed jerboas are more diverse in construction, but similar in building techniques. In contrast to five-toed jerboas, forelimbs play the predominant role in digging sandy soils by the three-toed jerboas. Three-toed jerboas can build simple daily burrows with only one tunnel in a soft sand. Shelter burrows of five-toed and some three-toed jerboas look like simple short tunnels with open holes used to escape predatory attacks. Burrows contain nest chambers

loosely constructed from dry vegetation or, in the case of the desert jerboa, camel hair.

Hibernation

Birch mice, jumping mice, and most species of jerboas hibernate. Duration of hibernation is taxon dependent and also dependent on geography. Birch mice hibernate up to six, or seven months. The weight loss during the half year hibernation in S. betulina reaches 50%. The body temperature in a hibernating birch mouse can drop to as low as 36°F (2°C). Among the jerboa species breeding only one time in a year, the five-toed pygmy jerboa (Cardiocranius paradoxus) goes into winter sleep in Mongolia from the end of August up to May of the next year. Species with the spring and fall breeding cycle have shorter hibernation periods. Jerboas inhabiting subtropical deserts do not hibernate, but a state of torpor may continue for several days.

Daily activity

All groups of Dipodidae are nocturnal rodents, although the birch mice and jumping mice can be active in the day time (mostly in the morning or evening). Jerboas are strictly nocturnal and sleep in individual burrows with the entrance closed by a soil plug during the daytime. Pygmy jerboas build plugs with their tail, whereas all other jerboa species use the muzzle. Only in spring is it possible to observe jerboas feeding in dusk before sunrise or immediately after sunset.

Territoriality

Dipodidae are solitary—as a rule, each individual uses its own burrow to sleep in the daytime and to hibernate. At the same time, antagonistic behavior is taxon and dependent upon location. Very little is known about home ranges and social relationships among species of birch mice. In the woodland jumping mouse (Napaeozapus insignis) individuals are highly tolerant of one another. Their home ranges vary from 1 to 9 acres (0.4–3.6 ha), and overlap. For Zapus princeps in the Colorado mountains, Stinson in 1977 reported very small home ranges (0.4 acres [0.17 ha] for males and 0.2 acres [0.1 ha] for females). Males had overlapping home ranges and were tolerant of one another, whereas females had more exclusive territories. Neighboring individuals tend to share the shelter burrows, although their home ranges can overlap considerably. In many jerboa species, individuals avoid contact with each other within overlapping areas. Only pygmy jerboas, especially Salpingotus crassicauda, seem to be aggressive and territorial.

Jerboas have no well-developed, specific skin glands for scent marking. Specific marking behavior (touching substrate by ano-genital area) is identified in the long-eared jerboa (Euchoreutes naso). Pygmy jerboas often bathe in sand, and sites of sand bathing can play the role of chemical marks.

Feeding ecology and diet

Birch mice and jumping mice feed on fungus, nuts, berries, fruits, and arthropods. They, as well jerboas, do not store food. Jerboas represent a diverse spectra of feeding adaptations from carnivory of Euchoreutes naso and Salpingotus kozlovi

to granivory-insectivory of Salpingotus crassicauda, granivory of Cardiocranius paradoxus, granivory-folivory of Dipus sagitta and Stylodipus telum, Eremodipus lichtensteini, A. elater and folivory of Paradipus ctenodactylus and of jerboas of genus Pygeretmus. Some species are omnivorous, feeding on green parts of plants, roots, bulbs, seeds, fruits, and arthropods (Allactaga major, A. sibirica, A. severtzovi, etc.). Based on the data on feeding of birch mice, jumping mice, and pygmy jerboas as well as on the data on teeth structure of extinct and recent Dipodidae, it is plausible that distribution of feeding resources and predation risk are predominant factors in the evolution of bipedal locomotion. In the open, desert-like habitat, the rodent that looks for sparsely distributed energy-rich food items becomes exposed to the predator much more often than in a sheltered, densely vegetated habitat. In such situations, natural selection may support an increase in acceleration of the escape flight at its initial stage, determining the vector of morphological innovations.

Foraging strategies among jerboas are intriguing, particularly among food specialists. For instance, the long-eared jerboa (Euchoreutes naso) can actively locate flying insects by their extremely large, flexible auricles that are close in size to the body length. The long-eared jerboa catches moths by making vertical rushes quick as lightning. The pygmy jerboa (Salpingotus kozlovi) forages at sand dune slopes. Its muzzle has a snout surrounded by a field of dense, long vibrissa oriented to the side. Digging in the sand, it makes small sand-slips and detects arthropods and their larva with both the nose and vibrissa field. Its very large middle ear cavities probably facilitate hearing ground-born sounds too. The comb-toed jerboa (Paradipus ctenodactylus) feeds on young green twigs of succulent bushes that grow on slopes of sand dunes. It makes vertical

jumps to cut off a twig and runs with it to the top of the sand dune (barkhan) to feed. Extremely acute hearing and strong hind feet protect this jerboa from predators while in open areas.

Reproductive biology

Birch mice have a single litter per year with a gestation period of four to five weeks and the duration of parental care four weeks—that is rather long for a very small rodent. Birch mice attain sexual maturity only after hibernation. In Sicista betulina, a single litter contains three to 11 pups. Among jumping mice, Zapus hudsonius is known to produce at least two litters, occasionally three, per year, but Z. princeps and Z. trinotatus have only a single annual litter. The gestation period is 17–23 days, and litter size is two to nine pups with an average of four to six young. Young females of only Z. hudsonius may be able to produce their own litters in the same season of their birth. In Napaeozapus insignis, females in southern populations give birth twice during the breeding season from early May to early September; in the north one litter per year is more common. The litter size varies from two to seven, averaging approximately 4.5 young. The gestation period is about 23 days, and the newborn pups are weaned at 34 days. Both males and females start to breed only after the first hibernation. Very little is known about reproductive biology of the Chinese jumping mouse (Eozapus setchuanus).

Birch mice and jumping mice are relatively long-lived creatures. The maximum reported lifetime in nature was four years for Sicista, Zapus, and Napaeozapus. In respect to breeding biology, jerboas can be classified into two groups. The first includes species that breed only one time during the warm season (in spring or summer). Two consecutive litters in a season is rare. This group includes Euchoreutes naso, Cardiocranius paradoxus, Stylodipus telum, S. andrewsi, Allactaga major, A. sibirica, A. balicunica, and Pygeretmus platyurus. The second group includes species that breed in the spring and fall, and can produce three litters in a year: Dipus sagitta, Allactaga severtzovi, Allactaga elater, Allactodipus bobrinskii, Pygeretmus pumilio, and P. shitkovi. Juveniles of this group of species, born in spring, can participate in the fall breeding. Juveniles of the first group of species start to breed only in the next year. The litter size of the species that breed only once per year is larger (from two to nine) than in the group that can produce multiple litters annually (from one to eight). Jerboas are polygamous. Gestation is described only for several species, and is rather long, varying from 19–20 days in Stylodipus telum to 25–30 days in Dipus sagitta, and to 28–30 days in Allactaga severtzovi. In many species, pups stay from five to six weeks in the nest burrow before emergence. The reasonable explanation of such a long duration of juveniles' postnatal development is that the motor skills needed for bipedal locomotion must be completely developed before the juvenile leaves the burrow. Soon after emergence pups become independent of their mother. Only in the first nights after they first go out juveniles follow the female. Some species of three-toed jerboas follow each other tail to tail and move in a linear fashion.

Conservation status

Among birch mice, the IUCN classified Sicista armenica as Critically Endangered. S. caudata is classified as Vulnerable because of progressive loss of its preferred habitat. Among jumping mice, the IUCN designates Eozapus setchuanus as Vulnerable because of destruction of mountain forest habitats. There are two subspecies of Zapus hudsonius with conservation status designated as Endangered (Z. h. preblei) and Vulnerable (Z. g. campestris). Zapus trinotatus orarius was designated by IUCN as Conservation Dependent, being restricted to a very small area on the northern California coast. Among jerboas, the IUCN declared special conservation status for 11 species. Euchoreutes naso and Allactaga tetradactyla were classified as Endangered; A. firouzi as Critically Endangered; Cardiocranius paradoxus and Salpingotus crassicauda as Vulnerable; and S. kozlovi, S. heptneri, Jaculus orientalis, Allactaga euphratica, Pygeretmus shitkovi, and Allactaga bullata as Lower Risk/Near Threatened.

The recent data from Mongolia suggest that Cardiocranius paradoxus and Allactaga bullata are common, widely distributed species in Mongolia. Another Mongolian species, Euchoreutes naso is characterized by a low but a stable population density in southern Gobi, where human impact is minimal (Rogovin and Shenbrot, 1995; Sokolov et al., 1996). On the other hand, Salpingotus heptneri must be considered critically endangered. Its range is limited to two small areas of in the northern Kyzylkum desert of Uzbekistan and Kazakhstan. The species has not been observed since the beginning of 1980. Now, due to the catastrophic desiccation of the Aral Sea and changes of the climate in the northern Kyzylkum, the future of this species is unknown and special research is needed.

Two subspecies of Jaculus blanfordi (J. b. turcmenicus and J. b. margianus) are also Endangered—they have small ranges restricted to northern and southern Turkmenistan and to the Kyzylkum Desert in Uzbekistan. Many areas of their typical habitat, the clay depressions and dry river streams between stabilized and semi-stabilized sand ridges, were plowed during 1970–1980. Salpingotus pallidus and Allactaga vinogradovi should be considered vulnerable because of their limited and fragmented ranges. Very little is known about Salpingotus michaelis and S. thomasi from Iran and Afganistan. New species of genus Salpingotus are still possible in Iran, Afganistan and northwestern China.

Significance to humans

Birch and jumping mice and jerboas have little significance to humans. In a plague, some jerboas (Stylodipus telum and Allactaga elater) can be invoked into epizootic process. Jerboas play a significant role in desert ecosystems. The density of most abundant species such as Stylodipus telum, Allactaga elater, or Pygeretmus pumilio can reach 40–50 individuals per 2.5 acres (1 ha).

Species accounts

List of Species

Southern birch mouse

Sicista subtilis

subfamily

Smintinae

taxonomy

Sicista subtilis (Pallas, 1773), Tobol River Valley, Kurgan Oblast, Russia. Four subspecies.

other common names

English: Steppe sicista; German: (Steppen-) Streifenmaus, (Steppen-) Birkenmaus; French: Siciste des steppes; Russian: Stepnaya myshovka.

physical characteristics

Head and body length 2.4–3 in (61–73 mm); tail 3–3.2 in (75–82 mm); body mass 0.2–0.4 oz (6–10 g). Upperparts striped light tan and brownish and paler underneath.

distribution

From eastern Austria to lake Baikal in south-central Siberia.

habitat

Flatland steppes of different types, fields, meadows at the southern age of forest zones.

behavior

Solitary. In captivity, females chase away the males before the birth of pups. Active in dusk and night, and sometimes in the day time.

feeding ecology and diet

Arthropods, seeds, roots, bulbs, and plant greens.

reproductive biology

Polygamous. Usually produces one litter of two to eight pups in May to beginning of June. Young from early litters can

breed in the same birth year. Gestation period is not less than 25 days, lactation 33–35 days. Pups stay in the burrow 27–34 days.

conservation status

Not threatened; a common species.

significance to humans

None known.

Long-eared jerboa

Euchoreutes naso

subfamily

Euchoreutinae

taxonomy

Euchoreutes naso Sclater, 1891, Xinjiang, China.

other common names

German: Langohrspringer, Riesenohrspringmaus; Russian: Dlinnoukhii tushkanchik.

physical characteristics

Head and body length 3–4.3 in (74–110 mm); tail 6–7.3 in (152–185 mm); body mass 1–1.6 oz (23–45g). Upperparts reddish yellow to pale russset and white underneath.

distribution

Deserts of China (Inner Mongolia and Xinjiang) and South of Mongolia (Trans-Altai Gobi).

habitat

Dry, sandy gravel watercourses, stony plain watersheds, sandy-gravel plateau in deserts with very sparse vegetation.

behavior

Solitary, but non-aggressive territoriality. Very large, highly overlapping home ranges. Often leaves scent marks with a strong smell.

feeding ecology and diet

Diet includes arthropods and even lizards. Often hunts flying insects.

reproductive biology

Polygamous. In Mongolia produces one litter of two to six pups in June. Young start to breed at the next year. Gestation period is unknown. Pups stay in the burrow not less than one month.

conservation status

It is considered to be a rare species due to permanently low densities, and is classified by IUCN as Endangered. Included in the Red Data Book of Mongolia.

significance to humans

None known.

Five-toed pygmy jerboa

Cardiocranius paradoxus

subfamily

Cardiocraniinae

taxonomy

Cardiocranius paradoxus Satunin, 1903, China, Gansu.

other common names

English: Satunin's five-toed pygmy jerboa, five-toed dwarf jerboa; German: Fünfzehige Zwergspringmaus; Russian: Patipalyi karlikovyi tushkanchik.

physical characteristics

Head and body length 2–3 in (52–68 mm); tail 3–4 in (68–91 mm); body mass 0.5–0.7 oz (13–19 g). Upperparts grayish buff and white underneath.

distribution

Deserts and semi-deserts of western and southern Mongolia, northwestern and northern China and eastern Kazakhstan. Range includes four isolated areas. Species range has at least four isolated portions.

habitat

Flat and slightly inclined mountain bases and foot hills with sandy to clay/gravel soils and a predominance of grass of genus Stipa in plant communities.

behavior

Solitary, but with overlapping home ranges that vary from 1.5 to 7.4 acres (0.6–3 ha) in size.

feeding ecology and diet

Feeds mostly on seeds and fruits. Green vegetation present in the diet only during spring.

reproductive biology

Polygamous. Breeds in summer, and has only one litter of one to five pups annually. Young breed in second year of life. Duration of gestation and lactation are unknown.

conservation status

Traditionally, it is considered rare, and it is included in the Red Data Books of Russia and Kazakhstan. In Mongolia, however, it is a rather common jerboa, although is still not well studied. There is no information on effects of habitat loss as a result of overgrazing. IUCN classifies it as Vulnerable.

significance to humans

None known.

Comb-toed jerboa

Paradipus ctenodactylus

subfamily

Dipodinae (Paradipodinae)

taxonomy

Paradipus ctenodactylus (Vinogradov, 1929) Turkmenistan.

other common names

German: Kammzehen-Springmaus; Russian: Grebnepalyi tushkanchik.

physical characteristics

Head and body length 5.5–6.5 in (140–165 mm); tail 7–9 in (180–225 mm); body mass 4–6.5 oz (112–185 g). Upperparts hazel to pinkish cinnamon and white undernath.

distribution

Deserts of Turkmenistan, Uzbekistan, and northern Iran.

habitat

Inhabitant of moving sands (barkhans) with very sparse vegetation.

behavior

Solitary with vast home ranges of up to 1.2 mi (2 km) oriented along sand ridges.

feeding ecology and diet

Feeds on plant greens of a few species of perennial sand desert plants. Seeds appear in a diet only at the end of summer and in fall. Forages at slopes of dunes, but transports food to the top ridges to ingest.

reproductive biology

Polygamous. In Karakum desert breeds one time in spring, in western Kyzylkum two times, once in the spring and summer. Only two- to three-year old females participate in the second wave of breeding. Litter size is small and stable with two to three pups. Duration of gestation and lactation periods are unknown.

conservation status

Common. Density is always low, but stable from one to five individuals per 2.5 acres (1 ha). Arrangements for stabilization of moving sands negatively effect this jerboa.

significance to humans

None known.

Hairy-footed jerboa

Dipus sagitta

subfamily

Dipodinae

taxonomy

Dipus sagitta (Pallas, 1773), Pavlodarskaya Oblast, Kazakhstan. Ten subspecies.

other common names

English: Rough-legged jerboa, feather-footed jerboa, northern three-toed jerboa; German: Rauhfub-Springmaus, Raufübige Wüstenspringmaus, Pfeilspringmaus; Russian: Mokhnonogii tushkanchik.

physical characteristics

Head and body length 4.5–6 in (115–145 mm); tail 7–7.1 in (175–180 mm); body mass 2.4–4 oz (69–104 g). Upperparts orangish and black during winter and pale sandy buff during summer. White underneath.

distribution

Sand deserts of northern Iran, Middle Asia (Turkmenistan, Uzbekistan, southwestern Kazakhstan), Kazakhstan, Central Asia (Mongolia, China), southeast of European Russia, and south of Altai Republic (RF). There are 10 large isolated range fragments and several small ranges.

habitat

Inhabitant of sandy habitats found in steppe, semi-desert and desert zones. In Central Asia (Mongolia, China), the hairy-footed jerboa also lives in areas with hard, rocky/gravel surfaces. Vegetation is always sparse. At the most northern part of the range inhabits pine forests.

behavior

Solitary with overlapping home ranges. In Mongolia, non-aggressive contacts dominate in nature. In captivity, hairy-footed jerboas easily form pairs and the male and female sleep in one nest.

feeding ecology and diet

Feeds mainly on seeds and desert plant greens. Arthropods sporadically supplement diet.

reproductive biology

Polygamous. Adult female bears two litters in spring and fall. Juveniles of first litters become mature at the age of 2.5 to three months, and participate in the fall breeding. Pregnancy continues not less than 35 days, lactation 40–45 days. Litter size varies from one to eight pups, commonly three to five offspring. In spring, lactation overlaps with gestation.

conservation status

Common, not threatened. One subspecies D. sagitta nogai is considered vulnerable because of steppe expansion in Southeastern European Russia beginning in 1990 and resulting in overgrowth of open sand dunes.

significance to humans

None known.

Little five-toed jerboa

Allactaga elater

subfamily

Allactaginae

taxonomy

Allactaga elater (Lichtenstein, 1828), Kyrgiz steppe, Kazakhstan. Seven subspecies.

other common names

English: Small five-toed jerboa; French: Petite jerboa; German: Zwerg-Pferdespringer, Kleiner Erdhase, Kleine Springmaus; Russian: Malyi tushkanchik.

physical characteristics

Head and body length 3.5–5 in (90–120 mm); tail 5.5–7.5 in (140–190 mm); body mass 1–3 oz (32–77 g). Upperparts sandy colored and paler underneath.

distribution

Deserts and semi-deserts of southeastern Europe, Kazakhstan, Middle Asia, Iran, northeastern Turkey, Afghanistan, western Pakistan, northwestern China, and southwest Mongolia. Species range appears uniform with two relatively small isolated pockets.

habitat

Inhabitant of diverse spectra of desert and semidesert habitats with patchy vegetation of small perennial shrubs, herbs, and succulents. Soils vary from hard clay to sometimes stony and also sandy.

behavior

Solitary with overlapping home ranges. Avoidance of contact.

feeding ecology and diet

Feeds mainly on seeds and desert plant greens. Arthropods and plant roots are rarely ingested.

reproductive biology

Polygamous. Adult female can bare two litters in spring and fall. Juveniles born in spring participate in the fall breeding. Duration of pregnancy is unclear. Juveniles emerged from the maternal burrow 35–40 days of life. In spring, lactation over-laps with gestation. Litter size varies from one to nine pups, commonly three to six offspring.

conservation status

Common, not threatened. Ranges of two subspecies, A. elater caucasicus (Nehring, 1900) and A. elater aralychensis (Satunin, 1901) are small.

significance to humans

Can be implicated in epizootic spread of plague. In some regions, the little five-toed jerboa is considered an agricultural pest.

Bobrinski's jerboa

Allactodipus bobrinskii

subfamily

Allactaginae

taxonomy

Allactodipus bobrinskii Kolesnikov, 1937, Uzbekistan.

other common names

German: Bobrinski's Pferde-Springer; Russian: Tushkanchik Bobrinskogo.

physical characteristics

Head and body length 4–5 in (110–135 mm); tail 7–8 in (170–200 mm); body mass 2–3 oz (52–77 g). Upperparts reddish brown and black to sandy and grayish buff; whitish underneath.

distribution

Endemic to central Turan lowland. Range consists of six isolated portions, three in Turkmenistan and three in Uzbekistan.

habitat

Inhabitant of clay and clay-stony plains with a thin sand-gravel sheet. Vegetation is sparse and represented by small succulent shrubs from the Chenopodiaceae family.

behavior

Solitary with overlapping home ranges of 2.5–6 acres (1–2.5 ha). Two to three daily burrows are used in rotation. Contact between neighbors is avoided.

feeding ecology and diet

Feeds mainly on vegetative parts of plants. Seeds and insects are ingested to a lesser degree.

reproductive biology

Little information is available. Assumed polygamous. Adult females can produce two litters in spring and fall. There is no exact data on the duration of gestation and lactation. Juveniles emerge from the maternal burrow at the age of 40–45 days. In spring, lactation overlaps with gestation. Litter size vary from two to seven pups, commonly four to six offspring.

conservation status

A fairly common species with a restricted and fragmented range, and a relatively low population density that varies from 0.5 to 3 individuals per 2.5 acres (1 ha) in optimal habitats.

significance to humans

None known.

Resources

Books

Corbet, G. B., and J. E. Hill. A World List of Mammalian Species. Oxford: Oxford University Press, 1991.

Fokin, I. M. Jerboas. St. Petersburg: Izdatelstvo LGU, 1978. (In Russian).

——. Locomotion and Morphology of Locomotive Organs of Jerboas. St. Petersburg: Nauka Press, 1978. (In Russian).

Gambarian, P. P. "Superfamily Groups of Rodents." In Rodents: Materialy VI Vsesoyusnogo soveshaniya. St. Petersburg: Nauka Press, 1983.

Nowak, Robert M. Walker Mammals of the World. Baltimore: Johns Hopkins University Press, 1999.

Ognev, S. I. "Jerboas." In Mammals of the USSR and Neighboring Countries. Moscow and St. Petersburg: Izdatelstvo AN SSSR, 1948. (In Russian).

Shenbrot, G. I., V. E. Sokolov, V. G. Heptner, and Yu. M. Kovalskaya. "Dipodoidea." In Mammals of Russia and Neighboring Regions. Moscow: Nauka Press, 1995. (In Russian).

Sokolov V. E., V. S. Lobachev, and V. N. Orlov. The Mammals of Mongolia. Family Didodidae: Euchoreutinae, Cardiocraniinae, Dipodinae. Moscow: Nauka Press, 1996 (In Russian).

Vinogradov, B. S. "Jerboas." In Fauna of the USSR. Mammals. Moscow-St. Petersburg: Izdatelstvo AN SSSR, 1937. (In Russian).

Periodicals

Rogovin, K. A., and G. I. Shenbrot. "Geographical Ecology of Monagolian Desert Rodents." Journal of Biogeography 22 (1995): 111–128.

Simpson, G. G. "The Principles of Classification and Classification of Mammals." Bulletin of American Museum of Natural History 85 (1945): 1–350.

Stinson, N. "Home Range of the Western Jumping Mouse, Zapus princeps, in the Colorado Rocky Mountains." Great Basin Naturalist 37 (1977): 87–90

Vorontsov, N. N., N. A. Malygina, and S. I. Radjabli. "Chromosomes of Jerboas (Rodentia, Dipodidae)." Zoologicheskii zjurnal 50 (1971): 1853–1860. (In Russian)

Organizations

The American Society of Mammalogists. Web site: <http://www.mammalsociety.org/>

IUCN—The World Conservation Union. Rue Mauverney 28, Gland, 1196 Switzerland. Phone: +41 (22) 999 0000. Fax: +41 (22) 999 0002. E-mail: mail@hq.iucn.org Web site: <http://www.iucn.org>

United States Fish and Wildlife Service. Web site: <http://www.fws.gov/>

Konstantin A. Rogovin, PhD