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Zokor

The zokor is a for in the Myospalacinae of the family , comprising the genera Myospalax and Eospalax, which are highly adapted subterranean herbivores endemic to central and eastern . These medium-sized mammals typically measure 15–28 cm in body length and weigh 150–600 grams, featuring cylindrical bodies covered in dense, moisture-resistant gray or brown fur, tiny vestigial eyes, short external ears, powerful forelimbs with long curved claws for excavation, and prominent incisors for loosening soil. Native to regions including northern and , southern , , and , zokors inhabit diverse environments such as temperate grasslands, steppes, river valleys, woodlands, and agricultural fields, often at elevations from up to 4,200 meters, where they prefer loose, nutrient-rich soils conducive to burrowing. Zokors lead a predominantly fossorial lifestyle, spending 85–90% of their time in complex systems that can extend up to 100 meters in length, with chambers for nesting, food storage, and waste; these are solitary, territorial, and primarily nocturnal or crepuscular, using vocalizations like grunts and head-banging to communicate and defend territories. Their diet consists mainly of parts such as roots, bulbs, tubers, rhizomes, and seeds, supplemented occasionally by and , which they gather and in burrows; this behavior aerates , enhances , and influences local communities, though zokors are often considered agricultural pests due to crop damage. Reproduction occurs seasonally, typically in , with females giving birth to litters of 2–6 young after a period of about 30–38 days; are altricial, remaining in the for several weeks before dispersing, and zokors generally live 3–5 years in . Most of the approximately 10 recognized are classified as Least Concern by the IUCN, though some face threats from habitat loss, , and traditional uses in bone trade; high-altitude like the plateau zokor (Eospalax baileyi) exhibit unique physiological adaptations, such as elevated levels and tolerance, for life in oxygen-poor environments.

Taxonomy

Classification

Zokors are classified within the subfamily Myospalacinae, established by Lilljeborg in 1866, as part of the family , which encompasses mole-rats and related burrowing . This placement reflects their shared morphological and ecological traits with other subterranean spalacids, such as adaptations. The subfamily Myospalacinae includes two extant genera: Eospalax (plateau zokors) and Myospalax (prairie zokors), distinguished primarily by cranial features like occiput shape. An extinct genus, †Pliosiphneus, is also recognized within the subfamily, representing early evolutionary forms with rooted molars. Molecular phylogenetic analyses, based on complete mitochondrial genomes and data, confirm Myospalacinae as a distinct within , closely allied with the blind mole-rats of Spalacinae and the of Rhizomyinae. These studies indicate that zokors diverged earlier from Rhizomyinae than from Spalacinae, supporting a shared ancestry among these subterranean lineages. As Asiatic burrowing , zokors exhibit an evolutionary history tied to the uplift of the Qinghai-Tibet Plateau, with the earliest fossils of an extinct myospalacine to the middle in . Major diversification within the genera occurred during the , approximately 4.68 million years ago, coinciding with environmental changes that promoted subterranean lifestyles.

Species

The subfamily Myospalacinae, known as zokors, includes approximately 10-12 recognized depending on taxonomic classifications, primarily divided between the genera Eospalax and Myospalax, with some debated as full . The genus Eospalax, endemic to high-altitude regions of , encompasses seven recognized adapted to plateau environments. Eospalax baileyi (plateau zokor), elevated to full status in 2014, inhabits the . Eospalax cansus (Gansu zokor) is restricted to loess plateau habitats in northwestern , with its status supported by morphological and genetic distinctions. Eospalax fontanierii (Chinese zokor) is the most widespread at lower elevations. Eospalax rothschildi (Rothschild's zokor) occupies mountainous areas in central . Additional include Eospalax rufescens, Eospalax smithii (Smith's zokor), and Eospalax muliensis (described in 2022 from the ). In contrast, the genus Myospalax consists of five species primarily adapted to lowland steppes and grasslands across northern , including parts of , , and . Myospalax psilurus (Manchurian zokor) ranges through northeastern grasslands. Myospalax aspalax (false zokor or Asiatic least zokor) inhabits steppe regions in southern and . Myospalax myospalax (Siberian zokor or Baikal zokor) is found in forested steppes near . Myospalax armandi and Myospalax epsilanus are also recognized in some classifications, contributing to taxonomic debates.

Description

Physical characteristics

Zokors, belonging to the subfamily Myospalacinae, exhibit a compact, cylindrical body adapted to subterranean life, with head and body lengths ranging from 15 to 27 cm and weights between 150 and 560 grams, though these measurements vary by species and sex, with males generally larger than females. Their is short, typically measuring 2 to 6 cm, and is sparsely haired, aiding in balance during underground movement. Externally, zokors possess soft, velvety that ranges from brown to gray dorsally, fading to paler shades ventrally, which helps them in environments. They have small, tiny eyes often covered by fur, rendering limited, and lack external pinnae, with auditory structures internalized to suit burrowing. Short vibrissae are present on the head for tactile navigation in dark tunnels. The limbs are short and robust, featuring wide, strong forefeet equipped with powerful, curved claws—the third claw on the forefoot being the longest and strongest—for primary excavation, distinguishing zokor claw-based digging from the incisor-digging of related spalacids. Hindlimbs are similarly short but provide propulsion during movement. The skull is robust and reinforced, supporting a dental formula of I 1/1, C 0/0, P 0/0, M 3/3, with large, orthodont incisors that are more robust than those of surface-dwelling rodents like rats, facilitating occasional soil manipulation despite claws being the main digging tool. The molars are hypsodont, with high crowns suited to their diet, and sexual dimorphism is evident in skull size, with males exhibiting larger dimensions.

Adaptations for burrowing

Zokors exhibit specialized sensory adaptations suited to their subterranean , where is absent and relies on non-visual cues. Their eyes are tiny and often covered by , rendering minimal but sufficient for detecting light gradients upon surfacing. External ears are absent, further emphasizing reliance on other senses, while keen hearing and aid in detecting predators and locating underground. Facial vibrissae, though short, provide tactile for maneuvering in confined, dark tunnels. Physiologically, zokors are adapted to the low-oxygen, high-carbon dioxide conditions within sealed burrows through enhanced oxygen transport mechanisms. They possess elevated levels of red cells, , and compared to related species like pikas and rats, allowing oxygen to be approximately 1.5 times higher despite venous being notably lower. This supports efficient gaseous in hypoxic environments, complemented by positively selected genes such as SFTPA2 and CSF2RB involved in respiratory function. Muscularly, their forelimbs are robust, with strong, curved s—particularly the elongated third —enabling powerful scratch- motions, while dense and musculature facilitates displacement. Lower resting rates and optimized aerobic further conserve during prolonged digging efforts. The skin and pelage of zokors are adapted to withstand and in soil-filled burrows. Their is soft, thick, and dense, ranging from gray to with paler underparts, providing and against particles during excavation. This velvety coat resists matting and wear, essential for their lifestyle. Unlike chisel-tooth digging spalacines (such as blind mole-rats in the genus ), which primarily use enlarged to excavate hard soils and experience significant tooth wear, zokors employ a claw-dominant scratch-digging strategy with their forepaws. This reduces incisor , as their teeth serve mainly to sever roots rather than bear the brunt of digging, allowing for less extreme dental while maintaining effective burrowing efficiency.

Distribution and habitat

Geographic range

Zokors, belonging to the subfamily Myospalacinae, are primarily distributed across Central and Northern Asia, with their core range encompassing northern and , , , and southern in . This distribution centers on the , particularly the vast steppes and high plateaus of the region, where the two main genera—Myospalax and Eospalax—occupy distinct but overlapping areas. The genus Eospalax, often associated with plateau environments, extends to high elevations on the and surrounding areas, reaching up to 4,300 meters in species like Eospalax baileyi. In contrast, Myospalax species are more prevalent in lowland to mid-elevation grasslands, typically up to 2,750 meters, as seen in Myospalax myospalax across the and adjacent lowlands. These altitudinal variations reflect adaptations to diverse topographic features within the broader Asian continental landscape. The geographic range of zokors has remained relatively stable historically, with no major contractions documented prior to 2025, though human activities such as agriculture and urbanization have led to in some areas. Overall, zokors exhibit high endemicity to the steppes and plateaus of , with most species confined to these biomes and limited dispersal beyond their native continental boundaries. For instance, certain Eospalax species are restricted to specific highland locales in , while Myospalax extends into transboundary regions of and .

Habitat preferences

Zokors of both genera Myospalax and Eospalax exhibit a strong preference for loose, friable soils that facilitate burrowing, such as deposits with high content and good air permeability, commonly found in grasslands, steppes, meadows, and valleys. Myospalax species favor these substrates in lowland to mid-elevation areas, while Eospalax species, including the Gansu zokor (Eospalax cansus), are adapted to thicker layers in the Chinese Loess Plateau, up to 350 meters deep, avoiding compact or rocky terrains that hinder excavation. Vegetation associations are closely tied to areas with dense root systems, including grasses and shrubs that provide structural support for burrows. Myospalax habitats include meadow steppes dominated by species like Stipa baicalensis and Leymus chinensis, while Eospalax prefers alpine meadows with such as Kobresia species on the . Zokors avoid waterlogged or heavily rocky soils, opting instead for environments where vegetation cover supports stable microhabitats, such as natural grasslands transitioning to forested edges. Microhabitat features include burrows constructed in pastures, old agricultural fields, and vegetable gardens, often on gentle slopes (10°–40°) with convex at slope tops to minimize risks. These tolerate a range of climates from semi-arid steppes to temperate and alpine meadows, typically at elevations of 900–4,300 meters, with Myospalax below 2,750 m and Eospalax from 2,700 m upward. Seasonal shifts in habitat use reflect climatic adaptations, with surface activity peaking in non-freezing seasons like and autumn when is workable, allowing for expansion near the surface (around 50 cm deep). In winter, zokors retreat to deeper burrows, up to 2 meters, where stable temperatures (1–10°C) are maintained in nesting chambers, reducing exposure to freezing surface conditions. This pattern ensures year-round subterranean living while optimizing energy for burrowing in favorable seasonal windows.

Behavior

Burrowing and activity patterns

Zokors construct extensive, multi-level systems that serve as their primary , consisting of interconnected tunnels, nesting chambers, and waste areas. These systems typically span lengths of 100 to 225 meters per individual, with home ranges varying from 10 to 1,500 square meters, allowing for solitary occupation during non-breeding periods. The tunnels are often angled between 2° and 30° to minimize light penetration, featuring branches, transportation passages, and deeper nesting chambers where individuals spend 85-90% of their time. Waste areas are designated within the system to manage refuse, while entrances can be rapidly sealed following damage to maintain burrow integrity. Digging occurs year-round using strong, claw-based excavation, where zokors loosen with their forelimbs and push it rearward for ejection. This displaces soil to the surface, forming characteristic mounds similar to those of voles but larger in scale, with an average volume of 0.007 cubic meters per zokor daily during active periods. These mounds result from continuous reshaping of the while accessing underground resources, contributing to the maintenance and expansion of the tunnel network. Zokors exhibit predominantly nocturnal and crepuscular activity patterns, with daily peaks typically between 3:00-7:00 a.m. and 5:00-11:00 p.m., though they remain active for about 50% of the day overall. Each individual occupies a single system solitarily outside of times, patrolling and repairing it as needed. Seasonally, activity intensifies in and summer for increased surface and expansion, while in winter, entrances are sealed to conserve energy and protect against harsh conditions. and resource accumulation phases, such as mid-May to late May and late August, each last 20-30 days and drive heightened digging efforts.

Social structure

Zokors are predominantly solitary , with each adult maintaining an exclusive system that serves as its primary , minimizing interactions with conspecifics outside of the breeding season. This solitary lifestyle reduces competition for resources in their subterranean habitats, where systems function as self-contained units for , nesting, and waste management. Overlap between systems is rare and typically limited to brief encounters during the period, after which individuals resume isolation to avoid conflict. Territorial behaviors in zokors are pronounced, involving scent marking with and to communicate ownership and deter intruders, as well as aggressive vocalizations and physical defense when territories are threatened. These exhibit high aggression toward same-sex conspecifics, particularly in defending entrances and core areas, which helps maintain spatial separation. Home ranges are dynamic, shrinking considerably during non-breeding periods for , but expanding significantly during breeding; for instance, in plateau zokors (Eospalax baileyi), male home ranges average 327 m²—about 6.5 times larger than females' 50 m²—to facilitate mate searching. Zokors also use seismic signaling to detect and occupy empty neighboring tunnels, thereby extending their territory without direct confrontation. Intraspecific interactions are infrequent and often tense, with rare aggression between males and females except during , where temporary range overlaps occur but quickly resolve post-mating. remain with the through and summer for and initial , dispersing in autumn after to establish independent burrows and avoid familial competition. Gender differences in territoriality are evident during the breeding season, when males become more exploratory and risk-prone to compete for access to females' burrows, while females maintain stable, smaller ranges focused on defense and . This pattern ensures efficient resource use in their isolated underground environments.

Ecology

Diet and foraging

Zokors are strictly herbivorous, with their diet consisting primarily of underground plant parts such as , bulbs, rhizomes, and tubers, supplemented by , grasses, and occasional above-ground like leaves and shoots. Forbs, particularly from families like (e.g., Echinops and ) and (e.g., ), dominate the diet, often comprising over 90% of consumed biomass, while grasses contribute significantly but less preferentially. This subterranean strategy minimizes exposure to predators, as zokors pull downward into their burrows using powerful incisors and forelimbs, targeting nutrient-rich perennial and even tree near habitats. Foraging activity peaks in autumn for high-altitude species like the plateau zokor, when they collect and food in specialized burrow chambers to sustain them through the harsh winter, relying heavily on stored tubers and during periods of cover when fresh is limited. In summer, they incorporate more fresh greens and aerial parts, including high-fat for reproductive demands, shifting from the tuber-heavy winter reliance. Each burrow system typically contains 1–4 caches, with individual caches holding up to approximately 1.45 of biomass across 18–28 plant species, primarily forbs like Polygonum viviparum and Stellera chamaejasme, ensuring a diverse, high-energy reserve. Zokors exhibit nutritional adaptations suited to their high-fiber, low-quality , functioning as hindgut fermenters with a well-developed that hosts microbial communities for breakdown and short-chain production. This fermentation process in the enhances digestibility of fibrous roots and tubers, supporting their energy-intensive burrowing lifestyle while maintaining efficient extraction from material. Traits such as composition and caching may vary among adapted to different elevations and climates.

Reproduction and

Zokors exhibit a seasonal reproductive cycle, breeding once annually, with timing varying by and , often during the months of to May in many populations. This timing aligns with increased food availability in their and habitats, facilitating the energy demands of . Mating occurs when males extend their systems to intersect with those of females, allowing access without cohabitation. Gestation lasts approximately 38–50 days, depending on the , after which females give birth to a single in a protected chamber within the system. Litter sizes typically range from 1 to 6 young, with an average of about 3 pups per litter across such as Eospalax baileyi and Myospalax fontanierii. Newborns are altricial, born hairless, blind, and weighing around 7 g, requiring intensive maternal care. Parental care is provided solely by the female, who nurses the young in the nest chamber. persists for about 50 days, during which the pups remain dependent on maternal milk and protection. Males play no role in rearing offspring, consistent with the solitary lifestyle of zokors outside the brief period. After , the young disperse to establish independent burrows, typically within 4–7 weeks of birth. Sexual maturity is attained at 10–12 months of , allowing individuals to participate in breeding during their second year. In the wild, zokors have a lifespan of 2–5 years, though some may reach 3–4 years on average due to predation and environmental stresses; captive individuals can live longer, up to 5–6 years. Reproductive traits may vary among species, with high-altitude forms exhibiting adaptations to shorter growing seasons.

Relationship with humans

Agricultural impact

Zokors, particularly species such as the Manchurian zokor (Myospalax psilurus) and zokor (Eospalax cansus), are recognized as significant agricultural pests in meadow steppes and areas of and , where their burrowing and root-feeding behaviors cause substantial damage to crops and pastures. These primarily consume underground parts of , including of perennial herbs from families like , , and , as well as tree in plantations, leading to reduced plant vigor and widespread die-off. In regions, their mound-building disrupts , creates bare patches that promote invasive weeds, and diminishes quality for . For instance, in Northeast 's meadow steppes, zokor activity affects approximately 38,019 s of grazing land annually, with mound densities exceeding 255 per in triggering sharp increases in loss. The economic repercussions are pronounced, particularly in agricultural zones of , , and provinces, where zokors contribute to reduced productivity in pastures and newly established forests. In afforestation efforts on the , zokors damage up to 30% of Chinese pine (Pinus tabuliformis) saplings, resulting in significant replanting costs and hindering initiatives. Across China's , small mammals including zokors are estimated to cause annual forage losses equivalent to 13.884 billion kilograms, valued at roughly CNY 2.85 billion (approximately USD 400 million). In Mongolia's Gannan Prefecture, plateau zokors (Eospalax baileyi) degrade about 1.28 million hectares of , halving carrying capacity in severely affected areas through mound coverage and plant mortality. Each zokor can store 20–30 kg of roots and hay underground, directly competing with human and exacerbating yield reductions in root-dependent crops and pastures. Management efforts focus on mitigating these impacts through a combination of lethal and ecological control methods, though challenges persist due to high costs and incomplete coverage. Common approaches include with arrow devices, using types C and D, and application of rodenticides like and anticoagulants, with poisoning campaigns covering 31,151 km² in targeted regions during the and reducing zokor populations by up to 31.6% in some areas between 1990 and 2000. Biological controls, such as installing perches to attract predators like upland buzzards, are increasingly promoted to minimize non-target effects. Only about 25% of affected grasslands receive , limited by labor intensity and expenses estimated at USD 100–120 million annually for similar issues. While zokors provide ecological benefits such as soil aeration and enhanced water infiltration—reducing runoff by 21–88% on slopes—these advantages are often outweighed by agricultural losses in high-density areas, where risk escalates beyond mound thresholds of 123–434 per . Conflicts have intensified since the mid-20th century, coinciding with and grassland conversion post-1949, when systematic programs shifted from traditional traps to industrialized rodenticides, reflecting broader efforts to protect expanding farmlands and pastures. Early on zokor prevention dates to 1982, underscoring the ongoing tension between their role in soil turnover and their status as pests.

Conservation status

The majority of zokor species are assessed as Least Concern by the International Union for Conservation of Nature (IUCN), indicating stable populations across their core ranges in . For instance, the zokor (Myospalax psilurus) is classified as Least Concern due to its wide distribution and tolerance to habitat modifications. Similarly, Rothschild's zokor (Eospalax rothschildi) holds a Least Concern status, though it faces localized pressures from habitat alterations. The zokor (Eospalax fontanierii), previously considered Vulnerable in , was reclassified as Least Concern following population recovery and expanded assessments. Key threats to zokor populations include driven by agricultural expansion and , which disrupts burrowing networks in and . Poisoning campaigns targeted at as agricultural pests have led to significant local declines, particularly in China's Province, where eradication efforts in the 1990s reduced populations to less than one-third of prior levels. exacerbates these issues on high-elevation plateaus by altering composition and increasing degradation risks, potentially shifting suitable habitats northward. Population estimates for zokors remain challenging due to their subterranean lifestyle, but densities in intact habitats typically range from 15 to 50 individuals per , suggesting stability in core areas like the . However, fragmented landscapes show notable declines, with genetic studies indicating reduced diversity and smaller effective population sizes in isolated patches, such as those in the for the Gansu zokor (Eospalax cansus). Conservation measures in emphasize protected areas, where several zokor populations, including Altai zokors (Myospalax myospalax), benefit from nature reserves that safeguard alpine meadows and steppes. Post-2020 research has advanced non-lethal control strategies, such as fertility inhibitors like and quinestrol mixtures, which reduce in field trials without broad environmental harm, promoting coexistence with . These efforts, combined with habitat restoration, aim to mitigate declines in fragmented regions.

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