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Alpine marmot

The Alpine marmot (Marmota marmota) is a robust, ground-dwelling rodent belonging to the squirrel family Sciuridae, characterized by its plump body, short legs, and dense fur ranging from blond to reddish-dark gray, with adults typically weighing 3–4.5 kg and a head-body length of 50–60 cm and a tail length of 13–15 cm. Native to the high-altitude grasslands and rocky slopes of the European Alps, this diurnal species is renowned for its extensive hibernation periods lasting from October to May, during which it retreats to complex burrow systems to survive harsh winter conditions.^[1] Primarily distributed across central and southern Europe, including France, Italy, Switzerland, Austria, and Germany, the Alpine marmot inhabits elevations between 800 and 3,000 meters above sea level, preferring open meadows with short vegetation and suitable burrowing substrates like gravelly or frozen soil.^[2] Through successful reintroduction programs since the mid-20th century, populations have been established in non-native areas such as the Pyrenees, Romanian Carpathians, and the Apennines, expanding its range and aiding ecological restoration in alpine ecosystems.^[3] These efforts have contributed to stable or increasing numbers, with the species classified as Least Concern on the IUCN Red List due to its wide distribution and lack of major threats.^[4] Alpine marmots exhibit highly social behavior, living in monogamous family groups of up to 15–20 individuals comprising a dominant breeding pair, subadults, and offspring, where they engage in cooperative activities such as territorial defense and vigilance against predators like eagles and foxes.^[1] Their diet is primarily herbivorous, consisting mainly of grasses, flowers, seeds, and roots, which they forage during the short active season from spring to autumn, storing fat reserves for hibernation.^[1] Reproduction occurs annually in May following emergence from hibernation, with females giving birth to litters of 1–7 young after a 33–34 day gestation, and juveniles reaching sexual maturity at around two years of age.^[1] With a lifespan of 15–18 years in the wild, these marmots play a key role in alpine environments by aerating soil through burrowing and serving as prey for larger carnivores.^[1]

Taxonomy and evolution

Taxonomy

The Alpine marmot (Marmota marmota) is a species of large ground squirrel first described by Carl Linnaeus in his Systema Naturae (10th edition), where it was initially classified as Mus marmota.^[5] It belongs to the family Sciuridae, which encompasses squirrels, chipmunks, and prairie dogs, and is placed in the genus Marmota, a group of robust, hibernating rodents adapted to montane and steppe environments.^[4] The genus Marmota includes 15 extant species, with approximately six native to North America and nine to Eurasia, reflecting a Holarctic distribution shaped by historical migrations across Beringia.^[6] The name "marmot" originates from Romance languages of the Alps, deriving from the Romansch marmott or similar terms, which trace back to Latin murem montis ("mountain mouse"), alluding to the animal's alpine habitat and rodent-like appearance.^[7] Linnaeus's binomial nomenclature formalized this in 1758, drawing on earlier European descriptions of the species as a distinctive high-elevation dweller. Within the genus, the Alpine marmot occupies the subgenus Marmota, distinguishing it from the North American subgenus Petromarmota. Two subspecies of M. marmota are currently recognized, varying primarily in body size, skull morphology, and pelage coloration due to geographic isolation in distinct mountain ranges; these differences are subtle but include larger body sizes in northern populations and paler fur in southern ones. Key subspecies include:

M. m. marmota (Linnaeus, 1758): The nominate subspecies, distributed across the central and western European Alps (e.g., Switzerland, Austria, France, Italy, Germany); it exhibits the typical robust build with head-body length of 50–60 cm and tawny-brown fur.^[8]
M. m. latirostris (Kratochvíl, 1961): Endemic to the High Tatras (Slovakia and Poland), this subspecies has a broader muzzle and shorter limbs, reflecting local adaptation to rocky subalpine terrain; it is considered a distinct evolutionary lineage from post-glacial isolation.^[9]

Phylogenetically, the Alpine marmot clusters within the Eurasian clade of Marmota, with close relatives including Asian species like the bobak marmot (M. bobak) and tarbagan marmot (M. sibirica), as well as North American forms such as the yellow-bellied marmot (M. flaviventris), based on molecular analyses of mitochondrial and nuclear DNA that highlight divergence during the Pliocene.^[10]

Evolutionary history

The genus Marmota, to which the Alpine marmot (Marmota marmota) belongs, originated in western North America during the late Miocene, approximately 7–5 million years ago, from ground squirrel ancestors within the tribe Marmotini, with molecular estimates placing the crown group divergence between 10 and 7 million years ago.^[6] Dispersal to Eurasia occurred via the Bering Land Bridge during the Pliocene, around 3–4 million years ago, marking the establishment of Eurasian lineages that would give rise to species adapted to continental climates.^[11] The Alpine marmot's specific evolutionary trajectory in Europe began in the Early Pleistocene, with the oldest known fossils dating to about 0.8 million years ago from the Gran Dolina site in Sierra de Atapuerca, Spain, indicating an initial presence in southern European landscapes.^[12] The fossil record of the Alpine marmot becomes more abundant in the Middle and Late Pleistocene, spanning 0.78 million to 11,700 years ago, with remains commonly found in cave deposits across Europe, such as those in the East Brina Cave in Croatia, reflecting its adaptation to periglacial steppe environments during glacial advances.^[13] During the Pleistocene's cold phases, including the Last Glacial Maximum around 21,000 years ago, the species' range contracted significantly, with populations likely surviving in southern European refugia such as the Iberian Peninsula and Balkans, while being extirpated from glaciated alpine regions.^[6] Fossil evidence from northern European plains during interglacials and periglacial zones underscores its resilience in open, cold-adapted habitats, peaking in the late Pleistocene.^[14] Key adaptations to Ice Age climates included the evolution of prolonged hibernation, lasting up to eight months, facilitated by genetic changes in lipid metabolism and immune response genes, such as elevated unsaturation in adipose tissues for cold tolerance and rapid evolution in cytokines like Interleukin 4 (dN/dS ratio of 2.31).^[10] These traits emerged as responses to the Pleistocene's seasonal extremes, enabling survival in resource-scarce, freezing environments. Genetic studies, including a 2019 whole-genome analysis of 11 individuals, reveal that such adaptations coincided with severe population bottlenecks, resulting in extremely low genetic diversity, with heterozygosity at just 0.12% per kilobase—one of the lowest among mammals—and impaired purifying selection against deleterious mutations.^[10] Following the end of the Last Glacial Maximum, the Alpine marmot recolonized the Alps around 12,000 years ago, expanding eastward from western refugia in a pattern driven by post-glacial warming and habitat availability in high-altitude meadows.^[10] This expansion contributed to the broader radiation of marmot species globally, influenced by Quaternary climate oscillations that promoted altitudinal shifts and periglacial niche exploitation.^[6] Recent genetic insights highlight persistent low heterozygosity stemming from historical isolation and bottlenecks during the Pleistocene, with modern populations showing no significant recovery in diversity despite numerical abundance, raising concerns for adaptability to ongoing climate change.^[10] This genomic stasis, characterized by a slowed evolutionary rate and high molecular conservation, underscores the long-term legacy of Ice Age perturbations on the species' resilience.^[10]

Physical characteristics

Morphology

The Alpine marmot (Marmota marmota) possesses a stocky, robust body adapted to a fossorial lifestyle, with short legs, a blunt muzzle, small rounded ears, and prominent claws on the forefeet specialized for digging. The thumb has a nail rather than a claw, aiding in burrowing.^[1] Adults exhibit a head-body length of 50–60 cm and a tail length of 13–16 cm, yielding a total length of 63–76 cm.^[15]^[1] Body mass ranges from 2.5 to 7 kg, with seasonal variation leading to higher weights in autumn; males are approximately 5% heavier than females, reflecting minimal sexual dimorphism in size, while other external traits show minimal differences.^[1] Skeletally, the species features a thick, broad skull and strong, muscular limbs that facilitate burrowing through rocky substrates.^[1] The dental formula is I 1/1, C 0/0, P 2/1, M 3/3 (total 22), consisting of continuously growing incisors and high-crowned molars suited for grinding plant material.^[16]

Adaptations

The Alpine marmot (Marmota marmota) possesses a thick, double-layered coat adapted for insulation in high-altitude cold environments, consisting of a dense underfur that traps air close to the body and coarser guard hairs that provide protection from wind and moisture.^[17] This pelage undergoes an annual molt after emergence from hibernation, with fur coloration generally tawny-brown with yellowish underparts and darker markings on the head and tail, providing crypsis in rocky, grassy highland habitats between 800 and 3,000 meters elevation.^[18] Sensory adaptations prioritize non-visual cues suited to burrow-dwelling and vigilance in open meadows; eyesight is relatively poor, but hearing is acute for detecting predator calls or alarm whistles from conspecifics, while olfaction aids in foraging and social recognition, supplemented by vibrissae (whiskers) for tactile navigation in dark burrows.^[19] Physiologically, the Alpine marmot exhibits a markedly elevated metabolic rate during the short summer active period, enabling rapid fat accumulation supporting up to ~70% weight gain to fuel extended hibernation without feeding.^[20] This hyperphagia-driven storage, peaking at around 300 grams of abdominal white adipose tissue by late summer, supports survival in oxygen-scarce, cold conditions.^[20]

Distribution and habitat

Geographic range

The Alpine marmot (Marmota marmota) is native to high-altitude regions of central and southern Europe, with its core distribution centered in the Alps, which span France, Switzerland, Italy, Austria, Germany, and Slovenia. Populations also occur in the Pyrenees along the France-Spain border, the Northern Apennines in Italy, the Carpathian Mountains across Romania, Slovakia, Poland, and Ukraine, and the Tatra Mountains in Slovakia and Poland. These areas represent the species' natural and reintroduced ranges in mountainous terrain above the tree line.^[21]^[22]^[23] The species inhabits elevations ranging from 800 to 3,200 meters above sea level, with the majority of colonies found between 1,500 and 2,500 meters; it is absent from lowland and forested regions below 800 meters. Historically, during the Pleistocene epoch, the Alpine marmot's range extended more broadly across Europe but contracted significantly to the higher elevations of the Alps following post-glacial warming and reforestation around 10,000 years ago. This restricted distribution persisted until the 19th and 20th centuries, when human-led reintroduction efforts expanded its presence; notable examples include the 1948 releases in the Pyrenees from stock originating in the French Alps (such as Vanoise and Mercantour), which established viable populations, and subsequent programs in the Carpathians starting in 1973.^[23]^[24]^[25]^[22]^[3]^[26] Population estimates exceed 100,000 individuals across its range as of 1994, with local densities in optimal alpine meadows reaching up to 1 individual per hectare, though varying widely based on habitat quality and human intervention. In the Romanian Carpathians, for instance, the population numbers approximately 815 individuals as of 2025. Recent habitat modeling studies from the 2020s document that the elevational optimum of colonies in the Central Alps increased by approximately 86 meters from 1982 to 2022 in response to warming temperatures, with no detected shifts in range margins.^[22]^[27]^[28]^[29]^[30]^[3]

Habitat requirements

The Alpine marmot (Marmota marmota) primarily inhabits alpine meadows, subalpine grasslands, and rocky slopes above the tree line, where open terrain facilitates foraging and vigilance against predators.^[31] These environments typically occur at elevations between 1,500 and 3,000 m, with a preference for south- or east-facing slopes that receive ample sunlight for thermoregulation and earlier snowmelt in spring.^[32] The species avoids dense forests and very steep cliffs, favoring areas with intermediate slopes (less than 45 degrees) that allow for effective burrowing and escape routes.^[33] Suitable soils are well-drained and gravelly, often consisting of friable glacial sediments or moraines that support extensive burrow networks without excessive waterlogging.^[31] The climate in these habitats features cool summers with average temperatures around 8–15°C and cold winters averaging -5 to -10°C, enabling the marmot's hibernation strategy from October to April.^[27] Annual precipitation ranges from 800 to 1,500 mm, predominantly as snow in winter, which provides insulation for burrows but requires sufficient summer melt for vegetation growth.^[31] Microhabitats center on burrow colonies housing 10–20 individuals in family groups, with main burrows featuring multiple entrances and tunnels for hibernation and shelter, supplemented by satellite burrows for foraging and escape.^[34] These systems are often positioned near water sources like streams or snowmelt areas to support hydration and maintain moist soils for digging.^[32] Vegetation in these microhabitats includes moderate cover of grasses and forbs, such as those in Juncion trifidi or Festucion picturatae communities, providing both food and camouflage.^[31] Alpine marmots show tolerance for human-modified habitats like moderately grazed pastures, which mimic natural meadows and enhance forage availability through regrowth.^[35] However, they avoid heavily grazed or urbanized areas where soil compaction, overgrazing, or disturbance disrupts burrowing and increases predation risk.^[36]

Ecology

Diet

The Alpine marmot is primarily herbivorous, consuming a diet dominated by plant material such as leaves, flowers, fruits, and seeds from grasses and forbs, with dicotyledonous forbs comprising over 50% of vegetative fragments due to their higher nutritional value.^[37] Preferred plant families include Leguminosae (e.g., Anthyllis and Medicago species), Compositae, Liliaceae, Plantaginaceae, and Umbelliferae, while monocotyledonous grasses like Festuca species and Nardus stricta are eaten less frequently despite their abundance.^[37] Flowering plants often represent 70-80% of the diet depending on the site and season, providing essential polyunsaturated fatty acids such as linoleic acid for fat storage.^[38] Occasional consumption of invertebrates (about 2.3% on average), lichens, or fungi occurs, particularly post-hibernation, but these do not significantly contribute to the overall herbivorous regimen.^[37] Seasonal variations in diet reflect the marmot's need to support growth, reproduction, and pre-hibernation fattening during its short active period from May to September. In spring (May-June), following emergence from hibernation, the diet shifts toward roots and higher proportions of animal prey (up to 7.5%) and grasses (e.g., 28% Gramineae in June) to replenish nutrients quickly, with emerging flowers adding protein-rich greens.^[37] During summer (July-August), foraging intensifies on high-protein forbs and a peak in flowers and fruits (22-26%), aiding juvenile growth and adult maintenance, while vegetative parts remain dominant at 74-78%.^[37] In autumn (September), the focus turns to high-carbohydrate storage foods like sedges, clover, and seeds to facilitate fat accumulation, enabling adults to gain approximately 30% body weight and pups up to 180% before hibernation.^[30]^[39] Foraging occurs primarily during daylight hours, with marmots dedicating time to selective grazing on nutrient-rich plants near burrows, often in family groups, though midday retreats to burrows reduce activity in hot weather.^[40] This behavior enhances intake during peak summer months, when food consumption more than doubles compared to other seasons to support energy demands.^[39] Nutritional adaptations include preferential selection of forbs for elevated phosphorus, calcium, and water content, which bolsters digestive efficiency via seasonal enlargement of the gastrointestinal tract.^[37]^[20] The diet's linoleic acid content is crucial for converting fats into phospholipids that maintain cardiac function during torpor.^[40] Selective feeding by Alpine marmots alters local plant communities by promoting nutrient cycling through burrowing and grazing.^[41] Studies on reintroduced populations show marmot grazing enhances vegetation equitability and heterogeneity without major compositional shifts.^[41]

Predators and parasites

The primary predators of the Alpine marmot (Marmota marmota) include birds of prey such as the golden eagle (Aquila chrysaetos) and mammalian carnivores like the red fox (Vulpes vulpes). These predators target marmots in open alpine meadows where visibility is high, with eagles posing aerial threats and foxes hunting on the ground. Other potential predators include the peregrine falcon (Falco peregrinus), Eurasian lynx (Lynx lynx), and brown bear (Ursus arctos). Juveniles are particularly vulnerable due to their smaller size and inexperience, experiencing higher predation pressure during the active season outside burrows.^[42]^[29]^[43] Predation contributes significantly to juvenile mortality, with rates estimated at around 14% during the summer active period from predator attacks, compared to 24% from overwinter causes in some populations. Alarm calls emitted by vigilant individuals can reduce predation risk by alerting the group to flee to burrows, allowing up to 90% of dispersing juveniles to survive initial encounters. Overall, predation accounts for a substantial portion of summer mortality, though exact rates vary by habitat and predator density.^[44]^[45]^[46] Alpine marmots host a range of ectoparasites, including mites such as Echinonyssus blanchardi and ticks like Ixodes ricinus, which can transmit pathogens but are less prevalent at higher elevations due to harsher conditions. Endoparasites are more common, encompassing helminths (e.g., Ctenotaenia marmotae, Ascaris laevis, Citellina alpina) and protozoans, with gastrointestinal infections showing higher prevalence at lower altitudes where warmer temperatures may favor parasite transmission. Hibernation periods help mitigate some infestations through natural grooming behaviors that dislodge ectoparasites.^[47]^[48] Disease outbreaks occasionally affect populations, though blood parasites and many vector-borne pathogens appear rare or absent in high-alpine environments. As of 2017, higher parasite prevalence at lower elevations suggests potential constraints on downward shifts, and a 2025 study indicates climate-driven upward elevational shifts in distribution, which may reduce infection risks in some populations. Defensive strategies such as burrowing for refuge and heightened vigilance during foraging further limit exposure to both predators and parasites.^[47]^[49]^[30]

Behavior and reproduction

Alpine marmots live in family-based colonies typically consisting of 10 to 20 individuals, including a dominant breeding pair, subordinate adults, and offspring from previous years.^[43] These groups are highly cohesive, with members sharing a common territory that ranges from 0.5 to 3 hectares in size, defended through scent marking primarily via cheek-rubbing on burrow entrances and prominent objects.^[50] The dominant pair maintains control over the colony, exhibiting monogamous pairing that structures the social unit around cooperative defense and resource access.^[51] Within the hierarchy, subordinates—often philopatric offspring aged 2 years or older—assist the dominants by contributing to vigilance against predators and territorial intruders, enhancing group survival through collective monitoring.^[43] Dispersal typically occurs at around 2 years of age, when young marmots leave to form or join new colonies, reducing intraspecific competition while allowing experienced subordinates to delay departure for added benefits like increased foraging efficiency.^[52] This cooperative dynamic fosters a stable social order, where non-breeding adults tolerate hierarchy enforcement to gain indirect fitness advantages. Communication in Alpine marmot colonies is multifaceted, relying on vocalizations, olfactory cues, and visual signals to maintain cohesion and alert members to threats. Alarm calls are particularly prominent, with distinct whistle types signaling specific dangers: a single high-pitched whistle often indicates aerial predators, while a series of shorter whistles warns of terrestrial threats, demonstrating semantic elements in their signaling system.^[53] Affiliative interactions include allogrooming to strengthen bonds and reduce tension, as well as submissive postures like lowered heads or averted gazes during encounters with dominants, which help regulate social harmony.^[54] Daily routines are diurnal and often bimodal, with peaks in activity during morning and late afternoon foraging sessions, interspersed with midday rests in burrows to avoid peak heat.^[55] Juveniles engage in play behaviors such as chasing and wrestling, which develop motor skills and social bonds, while adults frequently sunbathe on rocks to thermoregulate, a passive activity that also facilitates vigilance.^[43] Regarding interspecific interactions, Alpine marmots generally tolerate cohabitation with other herbivores like chamois in shared alpine meadows, showing minimal aggression toward non-threatening species, but respond aggressively to conspecific intruders through chases and vocal displays to protect territory boundaries.^[43]

Reproduction

The Alpine marmot (Marmota marmota) exhibits a primarily monogamous mating system, in which the dominant breeding pair within each social group reproduces annually.^[56] Only the dominant female typically breeds successfully due to reproductive suppression of subordinates, mediated by increased agonistic interactions from the dominant that elevate stress hormones (glucocorticoids) in subordinates, thereby inhibiting their progesterone production and disrupting the hypothalamic-pituitary-gonadal axis.^[57] This suppression is more pronounced toward unrelated subordinates, while related subordinates (e.g., daughters) may receive amicable treatment and gain indirect fitness benefits by aiding in offspring care.^[57] Breeding takes place shortly after emergence from hibernation, typically from March to May depending on altitude and snowmelt timing.^[40] Litters range in size from 1 to 7 young, with an average of 3–4 pups, though litter sizes have been decreasing since 1990 in response to climate change;^[58] gestation lasts 30–37 days, and births occur in a dedicated chamber within the burrow system.^[59]^[60] Offspring are altricial at birth, emerging blind, hairless, and weighing approximately 30 g.^[61] They remain in the natal burrow for about 40 days until weaning and first emerge aboveground around 50 days post-birth, typically between mid-June and mid-July, though emergence dates have been advancing in recent years due to climate change (as of 2025).^[40]^[62]^[63] Sexual maturity is attained at 2–3 years of age, after the second hibernation.^[40] Parental care is biparental, with the dominant male and female both contributing to protection and territory defense, while subordinate group members provide alloparental assistance, such as huddling to warm pups during early development.^[40] Infanticide is rare but documented, occasionally perpetrated by intruding males to bring females into estrus or by subordinate females to reduce competition.

Hibernation and life cycle

Hibernation process

Alpine marmots enter hibernation in late September or October, a process triggered by declining ambient temperatures and the accumulation of sufficient fat reserves to sustain the long torpor period. This hibernation lasts approximately 7-8 months, concluding with emergence in April or May, allowing the animals to endure the harsh alpine winter without feeding.^[38]^[64]^[20] Prior to hibernation, marmots engage in hyperphagia during autumn, substantially increasing their food intake to build fat stores, with body mass rising by up to 68% from spring to fall as white adipose tissue expands to support the energy demands of torpor. Family groups prepare by retreating to deep winter burrows, which they seal using a combination of soil, vegetation, and feces to insulate against cold and maintain stable internal conditions.^[20]^[65] During hibernation, profound physiological adaptations occur to minimize energy expenditure. Body temperature drops from normothermic levels of 38-40°C to as low as 1.5-5°C, closely tracking but often defending against burrow ambient temperatures below 5°C. Heart rate decreases dramatically from around 200 beats per minute in active states to 9 beats per minute or lower during torpor bouts. Metabolic rate is suppressed by approximately 95%, enabling survival on stored fat with minimal oxygen consumption of about 13.6 ml O₂·kg⁻¹·h⁻¹.^[38]^[66]^[64] Hibernation involves periodic arousals, occurring roughly every 12-13 days and lasting 8-12 hours, during which marmots rewarm to euthermic temperatures around 37°C for maintenance processes before returning to torpor. These arousals are synchronized within family units that huddle together in burrows extending 2-3 meters deep, where shared body heat helps stabilize temperatures and reduces individual energy costs by facilitating passive warming and limiting metabolic demands.^[20]^[67]^[68] Emergence from hibernation is gradual, often beginning with dominant males surveying the surroundings before females and juveniles follow, marking the transition to the active season. Recent analyses of energy budgets, including a 2022 study on free-living populations, confirm near-total metabolic suppression during torpor phases, with overall energy savings approaching 95-99% relative to active metabolism, underscoring the efficiency of this strategy for overwinter survival.^[20]

Life history traits

The Alpine marmot (Marmota marmota) typically lives 15–18 years in the wild, though individuals have been observed reaching up to 16 years in long-term studies.^[69]^[62] In captivity, lifespan can extend to 17–20 years, reflecting reduced environmental stressors and predation.^[69] Juvenile mortality is particularly high, with approximately 38% of young dying in their first year (survival rate of 0.62) due to factors like harsh weather and limited foraging experience.^[70] Growth occurs rapidly during the active summer period, enabling marmots to build substantial fat reserves for hibernation; adults reach peak body mass of around 8 kg just before entering torpor.^[20] Over the hibernation period, they lose about 30% of this mass, primarily from fat catabolism, resulting in a pronounced annual weight cycle that underscores their adaptation to seasonal resource scarcity. Population dynamics in core alpine ranges remain generally stable, supported by territorial behaviors that limit overcrowding.^[71] Density-dependent regulation occurs primarily through dispersal, where subordinates, often juveniles or young adults, emigrate to new areas when local group sizes increase, preventing resource depletion and inbreeding.^[71]^[72] Senescence manifests as declining reproductive output after approximately 8 years, with litter sizes and breeding success decreasing due to reduced body condition and dominance challenges.^[73] Unlike humans, Alpine marmots exhibit no menopause, continuing to attempt breeding into old age despite lower success rates.^[74]

Conservation and human interactions

Conservation status

The Alpine marmot (Marmota marmota) is classified as Least Concern on the IUCN Red List, a status it has held since its assessment in 2016, reflecting its overall stable core populations across the European Alps despite some local declines due to habitat fragmentation and environmental pressures.^[75] Population estimates for the species in the Alps range from 100,000 to 500,000 individuals, with stable or increasing numbers in protected areas, though monitoring efforts using capture-mark-recapture techniques reveal variability in density, such as approximately 0.9 individuals per hectare in certain Italian sites.^[22]^[27] Core populations remain abundant in suitable alpine meadows, but peripheral ranges show slower growth or contraction in response to changing conditions. Major threats to the Alpine marmot include habitat loss from residential development and ski infrastructure expansion, which fragment grasslands and increase human disturbance in high-elevation zones.^[76] Climate change exacerbates these issues by driving upward range shifts, with studies documenting an elevational optimum increase of about 86 meters over four decades in Swiss valleys, equivalent to roughly 22 meters per decade, potentially limiting access to suitable habitat as treelines advance.^[29] These shifts, combined with altered snow cover and vegetation patterns, pose risks to hibernation sites and foraging areas, though the species' adaptability has so far buffered major declines. Reintroduction programs have bolstered populations outside the native Alpine range, demonstrating success through careful site selection and acclimation protocols. In the Pyrenees, approximately 400 individuals were released between 1948 and 1988, leading to a current population exceeding 10,000 across suitable subalpine habitats.^[77] Similarly, in the Romanian Carpathians, initial releases of around 50 marmots in 1973 have expanded to over 800 individuals by 2025, supported by soft-release methods involving pre-acclimation enclosures to enhance survival and establishment.^[36] Reintroductions have also succeeded in the Apennines, where populations established since the 1960s now number several thousand, contributing to biodiversity restoration. Recent research in 2025 highlights the importance of incorporating genetic diversity in reintroduction strategies to mitigate inbreeding depression in isolated colonies, recommending the addition of 30–40 individuals from diverse Western European sources to Romanian populations to sustain long-term viability.^[36] Ongoing monitoring, including radio-collaring in select studies to track dispersal and habitat use, informs adaptive management to address these genetic and environmental challenges.^[78]

Human interactions

The Alpine marmot has long been intertwined with human activities in the European Alps, particularly through historical exploitation. In the 19th century and earlier, these marmots were widely hunted for their fur, which was used in clothing and accessories, and for their meat as a food source in mountainous regions. Their fat was also harvested for folk medicine, believed to alleviate rheumatism when applied topically. This hunting pressure contributed to local population declines across parts of their range until protective measures were implemented in the late 20th century.^[79]^[80] In modern times, interactions often involve conflicts, though these are relatively rare compared to historical levels. Alpine marmots occasionally damage pastures and crops by grazing on grasses and forbs in lowland areas near human settlements, leading some farmers to view them as pests, particularly in reintroduction zones where populations have expanded. Burrowing activities can also disrupt ski slopes and hiking trails in high-tourism areas, prompting localized management efforts.^[81]^[82] Despite these tensions, marmots play positive roles in human-altered landscapes. They attract ecotourists to alpine regions, where visitors observe colonies sunning themselves or whistling alarms from rocky outcrops, boosting local economies through guided wildlife watching in places like the Swiss and French Alps. Additionally, their foraging and burrowing behaviors aid seed dispersal, enhancing plant diversity and grassland health by transporting seeds via fur and creating nutrient-rich microsites that benefit vegetation recovery.^[83] Alpine marmots serve as valuable models in scientific research, especially for hibernation studies. Their ability to enter deep torpor, reducing metabolic rates dramatically during winter, has informed medical investigations into induced hypothermia for organ preservation and treatments for metabolic disorders like obesity and diabetes. Keeping them as pets is illegal in many areas due to their protected status under European wildlife laws, which prohibit capture and trade to prevent disease spread and habitat disruption.^[20]^[84] Culturally, the Alpine marmot holds symbolic importance in Alpine folklore, serving as the mascot for chimney sweepers in historical Savoyard traditions, where it represented industriousness and mountain life. Today, increased tourism brings both opportunities and challenges, as frequent human presence near burrows can stress colonies, altering vigilance behaviors and potentially reducing reproductive success in heavily visited sites.^[80]^[85]^[82]

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[13]
Ultraconserved elements improve resolution of marmot phylogeny ...
Marmots are members of the rodent family Sciuridae, a clade that emerged sometime in the Eocene but whose geographic origins are unknown (Heissig, 2003).Missing: position closest
[14]
The oldest European marmots: Metrical study of the Marmota fossils ...
May 6, 2022 · The oldest European remains of marmots (Genus Marmota) are 0.8 my old and come from the site of Gran Dolina, Atapuerca.
[15]
Taphonomy of alpine marmot ( Marmota marmota ) remains from the ...
Jun 30, 2025 · The Late Pleistocene deposits of the East and West Brina Caves near Drniš have yielded one of the largest collections of alpine marmot ...
[16]
the periglacial expansion of marmots (marmota) in - jstor
Marmota marmota, which presently inhabits three small areas in the Alps and Carpathians, occupied a vast territory, both lowland and mon- tane, in the ...
[17]
Alpine Marmot - Comparative Mammalian Brain Collections
Body length is 50-60 cm and the tail length is 13-16 cm. It generally lives in open areas such as meadows or forest edge. Marmots spend most of their lives in ...
[18]
Postweaning mass gain in juvenile alpine marmots Marmota marmota
A body mass sexual dimorphism of juveniles occurred at emergence, suggesting that mothers may provide more care for their male than their female offspring. ...Missing: morphology weight dental formula skeletal adaptations<|control11|><|separator|>
[19]
[PDF] Marmota caligata (Rodentia: Sciuridae)
Sep 26, 2011 · It is distributed in western North America from Alaska and Canada south to Washington and Montana and is found at elevations ranging from sea ...<|separator|>
[20]
Sciuridae (squirrels) | INFORMATION - Animal Diversity Web
... ( Marmota marmota , 3 to 8 kg). They also vary substantially in fur color ... The dental formula of sciurids is 1/1, 0/0, 1-2/1, 3/3 = 20-22. Other ...<|separator|>
[21]
[PDF] MAMMALIAN SPECIES No. 135, pp. 1-8, 5 figs. - Marmota flaviventris.
Apr 15, 1980 · 3); dental formula i 1/1, c 0/0, p 2/1, m 3/3, total 22; cheek teeth high crowned; anterior face of incisors yellowish-white. Measurements.
[22]
review of molt in mammals, with an emphasis on marmots (Rodentia
Jun 27, 2024 · In this pattern, the large follicle grows a guard hair and the 2 others grow underfur or wool. However, the terms “guard hair” and “underfur ...
[23]
[PDF] Fur color diversity in marmots
Some populations of this species are considerably lighter and the large subspecies, M. ... First international symposium on alpine marmot (Marmota marmota) and on ...
[24]
Do animals with fur get skin damage from sunlight like humans do ...
Jul 13, 2019 · So at least among mammals with a decent amount of fur, these pigments presumably go a long way towards UV protection. This has in fact been ...Missing: Alpine marmot
[25]
The allometry and scaling of the size of vertebrate eyes
On the other hand, rodents have highly developed senses of olfaction and hearing, senses which at times are better suited than vision to a nocturnal lifestyle.
[26]
Energy expenditure and body composition in a hibernator, the ... - NIH
Alpine marmots prefer relatively cool areas in alpine regions (Arnold 1999). Apparently, their body shape and insulation are optimal for activity in late winter ...
[27]
Comparative genomics reveals putative evidence for high-elevation ...
We found 141 enriched terms with many related to hypoxia, metabolism, mitochondrial function/development, and DNA repair.
[28]
1. Map of the geographic ranges of the living marmot species.
The alpine marmot (Marmota marmota L.) is a key species of high-altitude habitats in Europe, with populations found in the Pyrenees, Alps, and Carpathian ...Missing: subspecies list descriptions
[29]
Who and Where are we? - The Alpine Marmot Project
Since 1948, the marmot was introduced and not reintroduced, to the Pyrenees and now occupies many areas (Herrero et al. 1988). The marmot was also introduced ...
[30]
Alpine Marmot (Megafauna Parks (W Palearctic)) - iNaturalist
Alpine marmots live at heights between 800 and 3,200 metres in the Alps, Carpathians, Tatras, the Pyrenees and Northern Apennines in Italy. They were ...
[31]
Altitudinal Distribution of Alpine Marmot (Marmota marmota) in the ...
May 3, 2018 · The alpine marmot occupies areas ranging in altitude from 1300 to 2800 m in the Pyrenees, although 67% of the colonies are found between 1800 ...Missing: geographic | Show results with:geographic<|control11|><|separator|>
[32]
Climate change and the conservation of marmots
The currently recognized 15 species of marmots (Table 1) are restricted to the northern hemisphere [1]. The habitats occupied by marmots range from small, ...<|control11|><|separator|>
[33]
A double observer approach for tracking the abundance of Alpine ...
Jul 1, 2025 · We monitored a population of Alpine marmots (Marmota marmota) in Stelvio National Park (Italian Alps) over an 8-year period, from 2017 to 2024.
[34]
https://esj-journals.onlinelibrary.wiley.com/doi/full/10.1002/1438-390X.12104
[35]
Shifting Heights? A 40‐Year Resurvey of Alpine Marmot Distribution ...
Jul 20, 2025 · The Alpine marmot is particularly sensitive to high temperatures and highly territorial (Arnold 1999).Missing: behavior | Show results with:behavior
[36]
Shifting Heights? A 40‐Year Resurvey of Alpine Marmot Distribution ...
Jul 20, 2025 · This study examines the elevational changes in Alpine marmot distribution over the past 40 years in a Central Alps Mountain valley. Based on ...Missing: reintroductions | Show results with:reintroductions
[37]
Full article: Microhabitat Utilization of the Tatra Marmot (Marmota ...
Jan 5, 2018 · The function of Alpine marmot subspecies burrows and factors influencing their distribution were investigated at 17 sites in the Tatra ...Missing: adaptations | Show results with:adaptations
[38]
Habitat preferences of alpine marmots, Marmota marmota
Marmots preferred sites with a southern or eastern aspect (where snow melts relatively early), intermediate slopes, moderate plant cover, and a low level of ...Missing: soil terrain paper
[39]
(PDF) Habitat preferences of alpine marmots, Marmota marmota
Aug 6, 2025 · In the Alps, marmots are typically found between 1200 and 2700 m asl., but up to 3000 m asl., avoiding the forest belt and favouring grasslands ...Missing: soil | Show results with:soil
[40]
Marginal habitats provide unexpected survival benefits to the alpine ...
Jan 6, 2022 · The alpine marmot (Marmota marmota) is a ground-dwelling rodent which preferably inhabits high elevation meadows. This habitat is currently ...
[41]
(PDF) The successful introduction of the alpine marmot Marmota ...
Aug 5, 2025 · The alpine marmot is widely distributed and, apparently, well established in the Pyrenees. Population structure and demographic parameters are ...
[42]
Population Dynamics and Reintroduction Strategies for the Alpine ...
Aug 25, 2025 · This study aims to evaluate the current status of the alpine marmot population and assess the potential for reintroducing the species into three ...
[43]
[PDF] Diet selection of the alpine Marmot (Marmota m. marmota L ... - HAL
We studied the diet composition and selection of plants in the Alpine marmot Marmota m. marmota of two family groups in the Western pyrenees ...
[44]
Hibernation | The Alpine Marmot Project - WordPress.com
At the entry of hibernation, this fat storage reaches one third of an individual body mass (Körtner et Heldmaier 1995). To increase the energy of these reserves ...
[45]
Body Weight Cycles and Energy Balance in the Alpine Marmot ...
Body weight gain in alpine marmots was achieved by a more than 100% enhanced food intake during the summer months.
[46]
https://thealpinemarmotproject.files.wordpress.com/2019/01/dupont-2017-the-influence-of-sociality-on-population-dynamics-in-the-alpine-marmot-annotated.pdf
[47]
Ecosystem engineers in the extreme: The modest impact of marmots ...
Mar 27, 2023 · The presence of marmots and their burrows has surprisingly little effect on plant nutrient content and on vegetation cover in an extremely arid, cold mountain ...Missing: paper | Show results with:paper
[48]
How much do ecosystem engineers contribute to landscape ...
Alpine marmot (Marmota marmota) is the largest and most widespread burrowing animal occurring in European alpine meadows. This species may have direct local ...Missing: slope | Show results with:slope
[49]
Extrinsic and intrinsic factors affecting the activity budget of alpine ...
Mar 15, 2022 · Alpine marmots had a higher probability of being observed foraging on cloudy days compared to variable and sunny days.Missing: intake | Show results with:intake
[50]
[PDF] Personality and vigilance behaviour in alpine marmot (Marmota ...
Alpine marmots represent a good mode! for the study of antipredator strategies and risk taking behaviour; they live in social groups that occupy a burrow ...
[51]
[PDF] Population Size and Life-History Traits of the Alpine Marmot ... - Parcs
Jul 21, 2009 · The size of alpine marmots differs greatly between adults and juveniles. This difference makes it easy to visually estimate the age distribution ...<|separator|>
[52]
Model Complexity and Population Predictions. The Alpine Marmot ...
Twenty animals (c. 90%) survived the dangers of predation immediately on departing the natal neighbourhood, and the remainder of mortality occurred after the ...
[53]
[PDF] The Influence of sociality on population dynamics in the Alpine Marmot
predation for marmots by showing that summer mortality was mainly driven by predation in yellow-bellied marmots (Van vuren 2001), Vancouver Island marmots.
[54]
Absence of blood parasites and other vector-borne pathogens in ...
Jan 16, 2021 · At present, there is virtually no information regarding blood parasites and other VBPs in Alpine marmots. The aim of this study was to ...
[55]
Species-level microbiota of ticks and fleas from Marmota ... - Frontiers
Jun 20, 2023 · In the present study, we described the microbiota community of ticks and fleas sampling from marmots in Qinghai-Tibet Plateau, China. We aimed ...
[56]
(PDF) Parasite Adaptations to Hibernation in Alpine Marmots ...
There are three helminths common to Alpine marmots, Ctenotaenia marmotae, Ascaris laevis and Citellina alpina. The marmot's intestine greatly atrophies during ...Missing: papers | Show results with:papers
[57]
Higher risk of gastrointestinal parasite infection at lower elevation ...
Higher risk of gastrointestinal parasite infection at lower elevation suggests possible constraints in the distributional niche of Alpine marmots. Stefania ...
[58]
Population Dynamics of Enteric Parasites in the Endangered ...
Jul 22, 2024 · Alpine marmots (Marmota marmota) had a higher risk of enteric parasite infections at lower elevations, suggesting that parasite burden may ...
[59]
Social Signals Involved in Scent-Marking Behavior by Cheek ...
The Alpine marmot Marmota marmota is a territorial rodent. Resident adults regularly scent-mark their territory by cheek-rubbing, mainly on burrow entrance.Missing: size | Show results with:size
[60]
The evolution of marmot sociality: I. Why disperse late?
Aug 10, 2025 · Offspring did not disperse before sexual maturity at age 2 (Fig. ... Hence, hypothesis 1 is not supported, at least not for adult-sized, > 2 years ...
[61]
The evolution of marmot sociality: I. Why disperse late?
Offspring did not disperse before sexual maturity at age 2 (Fig. 1). Only territorial females bred, whereas territorial males were not able to monopolize ...
[62]
Alarm calling in Alpine marmot (Marmota marmota L.) - ResearchGate
Aug 10, 2025 · Many species produce shorter alarm calls to aerial predators and longer calls to terrestrial predators (e.g., Heard 1973, Dunford 1977 ...
[63]
(PDF) Sweet mama: affiliative interactions are related to the ...
Oct 16, 2024 · Here we investigate how sociality traits are associated to reproductive success in female Alpine marmots (Marmota marmota), a highly social and ...
[64]
Thermoregulation as a limit to habitat use in alpine marmots ...
We suggest that the alpine marmots' preference for south oriented slopes is due to the better conditions for hibernation there, the microclimate during summer ...
[65]
Sociality, mating system and reproductive skew in marmots
Most marmot species are monogamous with suppressed subordinate female reproduction, while the yellow-bellied marmot is polygynous with more even reproduction.Missing: biparental | Show results with:biparental
[66]
Reproductive suppression in female Alpine marmots, Marmota ...
Population dynamics, mating system and philopatry in a high altitude colony of alpine marmots (Marmota marmota L.) ... Reproductive suppression among female ...Missing: dimorphism size
[67]
Population dynamics, mating system and philopatry in a high ...
Population dynamics, mating system and philopatry in a high altitude colony of alpine marmots (Marmota marmota L.) ... Mean litter size was 4 ± 0.65, and the ...Missing: average | Show results with:average
[68]
Hormonal and behavioural changes during the mating season and ...
Dec 1, 2003 · ... Marmota marmota ranges from 34 (Psenner, 1957) to 37 days and is followed by 45 days of lactation. The average litter size is 2.4 pups ( ...Missing: care | Show results with:care
[69]
Annual and Life Cycle | The Alpine Marmot Project - WordPress.com
Gestation lasts 33 to 35 days (Psenner 1957). Birthing of the young occurs in a special birthing chamber within the burrow. A few days before giving birth, the ...
[70]
Age-specific survival in the socially monogamous alpine marmot ...
Mar 12, 2016 · The altricial offspring stay in the natal burrow during 40 days and once weaned emerge above ground between mid-June and mid-July ( Psenner ...
[71]
Regulation of body temperature and energy requirements of ...
Alpine marmots experience temperatures that do not allow minimal metabolism over the whole hibernation season because Ta in natural hibernacula decreases ...
[72]
Alpine marmot | Wildnispark Zürich
Marmota marmota. Did you know ... that the marmot family includes monkeys ... Marmots spend about 90 per cent of their lives underground and their burrows can ...<|control11|><|separator|>
[73]
Seasonal variation of cardiovascular function in the marmot ...
During hibernation heart rate fell to 9 ± 1.1 B/min. Mean arterial ... (Marmota marmota). 2003, Journal of Wildlife Diseases. View all citing articles ...
[74]
Rewarming rates of two large hibernators - ScienceDirect.com
In both species hibernation is broken every 13 days on average by rewarming to euthermic temperatures. We found that the time course of a rewarming could be ...
[75]
Social thermoregulation during hibernation in alpine marmots ...
Body temperature (T b ) of socially hibernating alpine marmots, a pair and two family groups, was monitored continuously from October to March.
[76]
Alpine marmot (Marmota marmota) longevity, ageing, and life history
Taxonomy: Kingdom: Animalia Phylum: Chordata Class: Mammalia (Taxon entry) Order: Rodentia Family: Sciuridae Genus: Marmota ; Kingdom: Animalia ; Phylum: ChordataMissing: binomial | Show results with:binomial
[77]
Variation in survival rates for the alpine marmot (Marmota marmota)
Survival rates for young of the year (YOY, from weaning to first birthday) were, on average, lower than in the older age class. In the older age class, annual ...
[78]
Model complexity and population predictions. The alpine marmot as ...
Aug 5, 2025 · All models were also used to predict potential density-dependent effects on alpine marmot population growth. In this regard, the models ...
[79]
Restricted dispersal reduces the strength of spatial density ... - NIH
We conclude that restricted dispersal can modify spatial density dependence in the wild, which has implications for the way population dynamics are likely to ...
[80]
Age-specific survival in the socially monogamous alpine marmot ...
We investigated age-specific variation in survival of dominant individuals in a long-lived and monogamous mammal, the alpine marmot (Marmota marmota), ...
[81]
The role of menopause and reproductive senescence in a long-lived ...
Feb 3, 2009 · Our work supports previous research showing that menopause and long post-reproductive lifespans are not a human phenomenon.
[82]
Threats to Alpine nature - | WWF
Habitat loss and fragmentation are the major threats to Alpine biodiversity. Spreading settlements, unsustainable farming, road networks, and river dams are ...<|separator|>
[83]
Explaining the successful introduction of the alpine marmot in the ...
Aug 5, 2025 · Our results reveal that the Alpine marmots reintroduced in the Pyrenees ... The study proposes a phased reintroduction strategy, combining ...Missing: longevity | Show results with:longevity
[84]
Effects of radio-collars on behaviour of Alpine chamois Rupicapra ...
Aug 10, 2025 · We observed in 30-minute focal animal sampling periods the behaviour of nine female alpine chamois carrying a radio-collar for > 2 years and 40 ...
[85]
[PDF] Hunting, but not outdoor recreation, modulates behavioural ...
Dec 16, 2021 · Alpine marmots frequently use human-modified landscapes, such as alpine pastures, and habitats used for human recreation (Bätzing 2002).Missing: grazing | Show results with:grazing
[86]
Marmots in human culture: from folklore to research (Chapter 1)
Marmots have been hunted for fur and meat in the western Alps and the southern Jura since the Late Pleistocene (Tomé and Chaix Reference Tomé, Chaix ...
[87]
Hunting, but not outdoor recreation, modulates behavioural ...
May 15, 2025 · Alpine marmots Marmota marmota are large, diurnal, and ground-dwelling rodents, inhabiting high-elevation alpine and subalpine grasslands in ...
[88]
Everything you need to know about Alpine Marmots ... - YouTube
Jun 21, 2025 · ... line. Marmots build extensive burrows for their communal family group and feed on alpine vegetation. Marmots in this video were filmed in near ...Missing: water | Show results with:water
[89]
Alpine marmot – Marmota marmota – Marmota alpina
Mar 1, 2023 · The Alpine marmot became extinct in the Pyrenees more than 15,000 years ago. Between 1948 and 1988 around 400 individuals were reintroduced in ...
[90]
The Alpine Marmot: A Remarkable Mountain Dweller by Franka ...
Alpine Marmots play a crucial role in their ecosystem by aerating the soil and facilitating seed dispersal through their burrowing activities.
[91]
The Torpid State: Recent Advances in Metabolic Adaptations and ...
This study conducted in marmots (M. flaviventris) showed that adiponectin is low at the beginning of fat accumulation and elevated during hibernation (Florant ...
[92]
Symbolic entities in the European Alps: Perception and use of a ...
Alpine ibex, chamois, marmot, and golden eagle are perceived as symbolic animals. Edelweiss, gentian, pine trees, and alpenrose are perceived as symbolic ...