Hubbry Logo
Cave bearCave bearMain
Open search
Cave bear
Community hub
Cave bear
logo
8 pages, 0 posts
0 subscribers
Be the first to start a discussion here.
Be the first to start a discussion here.
Cave bear
Cave bear
Cave bear
View on Wikipedia
from Wikipedia

Cave bear
Temporal range: Middle to Late Pleistocene, 0.25–0.024 Ma
↓
Mounted cave bear skeleton at Devil's Cave, Germany
Scientific classification Edit this classification
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Carnivora
Family: Ursidae
Subfamily: Ursinae
Genus: Ursus
Species:
†U. spelaeus
Binomial name
†Ursus spelaeus

The cave bear (Ursus spelaeus) is a prehistoric species of bear that lived in Europe and Asia during the Pleistocene and became extinct about 24,000 years ago during the Last Glacial Maximum.

The common and scientific name for the species comes from their remains having been largely been found caves. This reflects the views of experts that cave bears spent more time in caves than the brown bear, frequently using them to hibernate during the winter months. Unlike brown bears, cave bears are thought to have been almost entirely or exclusively herbivorous.

Cave bears exhibit a great degree of size, morphological and genetic variability, and Late Pleistocene cave bears are often (though not universally) considered to be species complex of up to 6 different species.[1][2]

Taxonomy

[edit]
Rearing Ursus spelaeus skeleton at the AMNH

Cave bear skeletons were first described in 1774 by Johann Friedrich Esper, in his book Newly Discovered Zoolites of Unknown Four Footed Animals. While scientists at the time considered that the skeletons could belong to apes, canids, felids, or even dragons or unicorns, Esper postulated that they actually belonged to polar bears. Twenty years later, Johann Christian Rosenmüller, an anatomist at Leipzig University, gave the species its binomial name. The bones were so numerous that most researchers had little regard for them. During World War I, with the scarcity of phosphate dung, earth from the caves where cave bear bones occurred was used as a source of phosphates. When the "dragon caves" in Austria’s Styria region were exploited for this purpose, only the skulls and leg bones were kept.[3]

Many caves in Central Europe have skeletons of cave bears inside, such as the Heinrichshöhle in Hemer and the Dechenhöhle in Iserlohn, Germany. A complete skeleton, five complete skulls, and 18 other bones were found inside Kletno Bear Cave, in 1966 in Poland.[4] In Romania, in a cave called Bears' Cave, 140 cave bear skeletons were discovered in 1983.[5][failed verification]

Remains assigned to "cave bears" sensu lato from the Late Pleistocene exhibit a strong degree of morphological and size variability, and have often been assigned to their own species, including Ursus rossicus (Eastern Europe, Central Asia and Siberia), Ursus ingressus (Central Europe to the Urals), Ursus kanivetz, (Urals) Ursus kudarensis (the Caucasus), Ursus eremus (Central Europe, possibly a subspecies of U. spelaeus s.s.) and Ursus spelaeus sensu stricto (Western, Central and Southeast Europe).[6][2] These populations/species show considerable genetic divergence from each other, (with genetic divergences estimated at hundreds of thousands to a million years)[2] though whether these species should be regarded as synonyms of U. spelaeus is debated.[6]

Evolution

[edit]

Both the cave bear and the brown bear are thought to be descended from the Early Pleistocene species Ursus etruscus.[7][8][9][10] The date of divergence between the ancestors of cave bears and brown bears has been estimated at around 1.2–1.5 million years ago.[11][2] The earliest remains of the cave bear lineage are assigned to the species Ursus deningeri (sometimes called Deninger's bear), which first appears in the fossil record at the end of the Early Pleistocene, around 1.2-0.8 million years ago.[12][6] U. deningeri is known from remains spanning from Europe to China.[13] The transition between Ursus deningeri and Ursus spelaeus is often given as the Last Interglacial (130-115,000 years ago), although the boundary between these forms is arbitrary, and intermediate or transitional taxa have been proposed, e.g. Ursus spelaeus deningeroides,[14] while other authorities consider both taxa to be chronological variants of the same species.[15]

Cave bears found anywhere will vary in age, thus facilitating investigations into evolutionary trends. The three anterior premolars were gradually reduced, then disappeared, possibly in response to a largely vegetarian diet. In a fourth of the skulls found in the Conturines, the third premolar is still present, while more derived specimens elsewhere lack it. The last remaining premolar became conjugated with the true molars, enlarging the crown and granting it more cusps and cutting borders. This phenomenon, called molarization, improved the mastication capacities of the molars, facilitating the processing of tough vegetation. This allowed the cave bear to gain more energy for hibernation, while eating less than its ancestors.[16]

In 2005, scientists recovered and sequenced the nuclear DNA of a cave bear that lived between 42,000 and 44,000 years ago. The procedure used genomic DNA extracted from one of the animal's teeth. Sequencing the DNA directly (rather than first replicating it with the polymerase chain reaction), the scientists recovered 21 cave bear genes from remains that did not yield significant amounts of DNA with traditional techniques.[17] This study confirmed and built on results from a previous study using mitochondrial DNA extracted from cave bear remains ranging from 20,000 to 130,000 years old.[11] Both show that the cave bear was more closely related to the brown bear and polar bear than it was to the American black bear, but had split from the brown bear lineage before the distinct eastern and western brown bear lineages diversified, and before the split of brown bears and polar bears..[11] However, a recent study showed that both species had some hybridization between them.[18]

Description

[edit]
Life restoration.

The cave bear had a very broad, domed skull with a steep forehead; its stout body had long thighs, massive shins and in-turning feet, making it similar in skeletal structure to the brown bear.[19] Cave bears were comparable in size to, or larger than, the largest modern-day bears, measuring up to 2 m (6.6 ft) in length.[20] The average weight for males was 350 to 600 kg (770 to 1,320 lb),[21] while females weighed 225 to 250 kg (495 to 550 lb).[21] Of cave bear skeletons in museums, 90% are classified as male due to a misconception that the female skeletons were merely "dwarfs". Cave bears grew larger during glaciations and smaller during interglacials, probably to adjust heat loss rate.[22]

The morphology of the mandible of cave bears varied depending on climate; individuals in colder, drier, and more seasonal climates possessed more slender mandibles and a dorsoventrally smaller mandibular ramus relative to individuals living in warmer and wetter conditions.[23] Cave bears of the last Ice Age lacked the usual two or three premolars present in other bears; to compensate, the last molar is very elongated, with supplementary cusps.[24] The humerus of the cave bear was similar in size to that of the polar bear, as were the femora of females. The femora of male cave bears, however, bore more similarities in size to those of Kodiak bears.[21]

Behaviour

[edit]

Dietary habits

[edit]
Ursus spelaeus lacked the usual two or three premolars present in other bear species.

Cave bear teeth were very large and show greater wear than most modern bear species, suggesting a diet of tough materials. However, tubers and other gritty food, which cause distinctive tooth wear in modern brown bears, do not appear to have constituted a major part of cave bears' diets on the basis of dental microwear analysis.[25] Seed fruits are documented to have been consumed by cave bears.[26] Cave bear dental microwear from the high altitude site of Ramesch-Knochenhöhle in the Totes Gebirge indicates that some cave bears living in mountains ingested large amounts of sand as a result of feeding on alpine vegetation. The results from Ramesch-Knochenhöhle also indicate two very distinct patterns of microwear among the same subspecies, which may potentially reflect differences in feeding behaviour between male and female cave bears.[27]

The morphological features of the cave bear chewing apparatus, including loss of premolars, have long been suggested to indicate their diets displayed a higher degree of herbivory than the Eurasian brown bear.[8] Indeed, a solely vegetarian diet has been inferred on the basis of tooth morphology.[9] Results obtained on the stable isotopes of cave bear bones also point to a largely vegetarian diet in having low levels of nitrogen-15 and carbon-13, which are accumulated at a faster rate by carnivores as opposed to herbivores.[28][29][30] Among Mediterranean cave bears such as those found in Toll Cave in northeastern Spain, the particularly low δ15N values may be a result of their high intake of nitrogen-fixing plants.[31]

Detail of the molars of the lower jaw

However, some evidence points toward the occasional inclusion of animal protein in cave bear diets. For example, toothmarks on cave bear remains in areas where cave bears are the only recorded potential carnivores suggests occasional cannibalistic scavenging,[32][33] possibly on individuals that died during hibernation, and dental microwear analysis indicates the cave bear may have fed on a greater quantity of bone than its contemporary, the smaller Eurasian brown bear.[34] The dental microwear patterns of cave bear molars from the northeastern Iberian Peninsula show that cave bears may have consumed more meat in the days and weeks leading up to hibernation.[35] Cave bear dental microwear from Belgium has also showed clear evidence of their omnivory during predormancy, with bone, insect, and mammal meat consumption being inferred from the microwear patterns.[36] Comparisons of the dental microwear and macrowear of brown bears and cave bears in northern Spain found that the latter were significantly more osteophagous than the former.[37] Additionally, cave bear remains from Peștera cu Oase in the southwestern tip of the Romanian part of the Carpathian Mountains had elevated levels of nitrogen-15 in their bones, indicative of omnivorous diets,[30][38][39] although the values are within the range of those found for the strictly herbivorous mammoth.[40] One stable isotopic study concluded that the degree of omnivory in cave bears was similar to that in modern brown bears.[41]

Although the current prevailing opinion concludes that cave bears were largely herbivorous, and more so than any modern species of the genus Ursus,[42] increasing evidence points to omnivorous diets, based both on regional variability of isotopic composition of bone remains indicative of dietary plasticity,[30][38] and on a recent re-evaluation of craniodental morphology that places the cave bear squarely among omnivorous modern bear species with respect to its skull and tooth shapes.[43]

Mortality

[edit]
Standing skeleton of juvenile cave bear

Death during hibernation was a common end for cave bears, mainly befalling specimens that failed ecologically during the summer season through inexperience, sickness or old age.[44] Some cave bear bones show signs of numerous ailments, including spinal fusion, bone tumours, cavities, tooth resorption, necrosis (particularly in younger specimens), osteomyelitis, periostitis, rickets and kidney stones.[19] There is also evidence that cave bears suffered from tuberculosis.[45] Male cave bear skeletons have been found with broken bacula, probably due to fighting during the breeding season.[44] Cave bear longevity is unknown, though it has been estimated that they seldom exceeded twenty years of age.[46] Paleontologists doubt adult cave bears had any natural predators, save for pack-hunting wolves and cave hyenas, which would probably have attacked sick or infirm individuals.[46] Cave hyenas are thought to be responsible for the disarticulation and destruction of some cave bear skeletons. Such large carcasses were an optimal food resource for the hyenas, especially at the end of the winter, when food was scarce.[47] The presence of fully articulated adult cave lion skeletons, deep in cave bear dens, indicates the lions may have occasionally entered dens to prey on hibernating cave bears, with some dying in the attempt.[48]

Range and habitat

[edit]

The cave bear's range stretched across Europe; from Spain[49] and the British Isles in the west,[42] Belgium,[50] Italy,[51] parts of Germany,[52] Poland,[53] the Czech Republic,[54][55] the Balkans,[56][57] Romania,[58][59] Georgia, and parts of Russia,[60] including the Caucasus; and northern Iran.[61] No traces of cave bears have been found in the northern British Isles, Scandinavia or the Baltic countries, which were all covered in extensive glaciers at the time. The largest numbers of cave bear remains have been found in Austria, Switzerland, northern Italy, northern Spain, southern France, and Romania, roughly corresponding with the Pyrenees, Alps, and Carpathians. The huge number of bones found in southern, central and eastern Europe has led some scientists to think Europe may have once had herds of cave bears. Others, however, point out that, though some caves have thousands of bones, they were accumulated over a period of 100,000 years or more, thus requiring only two deaths in a cave per year to account for the large numbers.[46]

The cave bear inhabited low mountainous areas, especially in regions rich in limestone caves. They seem to have avoided open plains, preferring forested or forest-edged terrains.[46]

Relationship with humans

[edit]
Cave bear (upper right) along with other animals depicted in rock art from the Les Combarelles cave

Cave bear and brown bear remains that are modified by Neanderthals to be used as tools have been found in various regions across Europe, indicating the exploitation of bear carcasses by Paleolithic humans.[62][63][64] Probable evidence of Neanderthals' opportunistic hunting and/or competition against cave bears and brown bears have also been reported.[65][66]

Between the years 1917 and 1923, the Drachenloch cave in Switzerland was excavated by Emil Bächler. The excavation uncovered more than 30,000 cave bear skeletons. It also uncovered a stone chest or cist, consisting of a low wall built from limestone slabs near a cave wall with a number of bear skulls inside it. A cave bear skull was also found with a femur bone from another bear stuck inside it. Scholars speculated that it was proof of prehistoric human religious rites involving the cave bear, or that the Drachenloch cave bears were hunted as part of a hunting ritual, or that the skulls were kept as trophies.[67] In Archaeology, Religion, Ritual (2004), archaeologist Timothy Insoll strongly questions whether the Drachenloch finds in the stone cist were the result of human interaction. Insoll states that the evidence for religious practices involving cave bears in this time period is "far from convincing". Insoll also states that comparisons with the religious practices involving bears that are known from historic times are invalid.[68]

A similar phenomenon was encountered in Regourdou, southern France. A rectangular pit contained the remains of at least twenty bears, covered by a massive stone slab. The remains of a Neanderthal lay nearby in another stone pit, with various objects, including a bear humerus, a scraper, a core, and some flakes, which were interpreted as grave offerings.[69]

An unusual discovery in a deep chamber of Basura Cave in Savona, Italy, is thought to be related to cave bear worship, because there is a vaguely zoomorphic stalagmite surrounded by clay pellets. It is thought to have been used by Neanderthals for a ceremony; bear bones scattered on the floor further suggests it was likely to have had some sort of ritual purpose.[70]

Extinction

[edit]
Skeleton of a cave bear in the '"Bears' Cave", Chișcău, Romania

Reassessment of fossils in 2019 indicate that the cave bear probably died out 24,000 years ago.[71] A complex set of factors, rather than a single factor, are suggested to have led to the extinction.[72]

Compared with other megafaunal species that also became extinct during the Last Glacial Maximum, the cave bear was believed to have had a more specialized diet of high-quality plants and a relatively restricted geographical range. This was suggested as an explanation as to why it died out so much earlier than the rest.[42] Some experts have disputed this claim, as the cave bear had survived multiple climate changes prior to extinction. Additionally, mitochondrial DNA research indicated that the genetic decline of the cave bear began long before it became extinct, demonstrating habitat loss due to climate change was not responsible.[72] Finally, high δ15N levels were found in cave bear bones from Romania, indicating wider dietary possibilities than previously believed.[30]

Life restoration by Charles R. Knight

Some evidence indicates that the cave bear used only caves for hibernation and was not inclined to use other locations, such as thickets, for this purpose, in contrast to the more versatile brown bear. This specialized hibernation behavior would have caused a high winter mortality rate for cave bears that failed to find available caves. Therefore, as human populations slowly increased, the cave bear faced a shrinking pool of suitable caves, and slowly faded away to extinction, as both Neanderthals and anatomically modern humans sought out caves as living quarters, depriving the cave bear of vital habitat. This hypothesis is being researched as of 2010. According to the research study, published in the journal Molecular Biology and Evolution, radiocarbon dating of the fossil remains shows that the cave bear ceased to be abundant in Central Europe around 35,000 years ago.[73]

In addition to environmental change, human hunting has also been implicated in the ultimate extinction of the cave bear.[74] In 2019, the results of a large scale study of 81 bone specimens (resulting in 59 new sequences) and 64 previously published complete mitochondrial genomes of cave bear mitochondrial DNA remains found in Switzerland, Poland, France, Spain, Germany, Italy and Serbia, indicated that the cave bear population drastically declined starting around 40,000 years ago at the onset of the Aurignacian, coinciding with the arrival of anatomically modern humans.[75][76] It was concluded that human hunting and/or competition played a major role in their decline and ultimate disappearance, and that climate change was not likely to have been the dominant factor.[76] In a study of Spanish cave bear mtDNA, each cave used by cave bears was found to contain almost exclusively a unique lineage of closely related haplotypes, indicating a homing behaviour for birthing and hibernation. The conclusion of this study is cave bears could not easily colonize new sites when in competition with humans for these resources.[77]

Overhunting by humans has been dismissed by some as human populations at the time were too small to pose a serious threat to the cave bear's survival. However, the two species may have competed for living space in caves.[46][72] The Chauvet Cave contains around 300 "bear hollows" created by cave bear hibernation.[78] Unlike brown bears, cave bears are seldom represented in cave paintings, leading some experts to believe the cave bear may have been avoided by human hunters[79] or their habitat preferences may not have overlapped. Paleontologist Björn Kurtén hypothesized cave bear populations were fragmented and under stress even before the advent of the glaciers.[46] Populations living south of the Alps possibly survived significantly longer.[42]

See also

[edit]

References

[edit]
[edit]
Revisions and contributorsEdit on WikipediaRead on Wikipedia

Cave bear

View on Grokipedia
from Grokipedia
The cave bear (Ursus spelaeus) was an extinct species of large ursid that lived during the Middle and epochs, approximately from 360,000 to 24,000 years ago, primarily in and western . Closely related to the modern (Ursus arctos), it is distinguished by its robust morphology, with adult males typically weighing 400–500 kg and females 250–300 kg, often exceeding the size of contemporary bear species. Known from abundant remains preserved in caves—where the bears hibernated and died—the species exhibited a predominantly herbivorous diet, relying on vegetation such as leaves, berries, roots, and possibly some scavenging or omnivory during pre-hibernation periods. This bear's anatomy featured a steep , large molars adapted for grinding plant matter, and a stocky build suited to forested and mountainous habitats across its range, from the to the . Genetic studies indicate that cave bears diverged from brown bears around 1.2 to 1.6 million years ago, evolving distinct adaptations like enhanced fat storage for prolonged , which may have contributed to their vulnerability during environmental shifts; also reveals some hybridization with brown bears, with traces in modern populations. Populations showed low , particularly in isolated refugia during glacial maxima, reflecting a specialized lifestyle tied to temperate woodlands that diminished with the onset of colder, open landscapes. The extinction of U. spelaeus during the around 24,000 years ago is attributed to a combination of climate cooling, habitat loss, and an inflexible herbivorous diet that limited adaptability to changing food availability, along with possible human impacts such as competition for shelters and . Unlike more versatile brown bears that survived by broadening their diets, cave bears' reliance on specific resources and high demands for their large body size likely accelerated their decline. Fossil evidence from sites like in highlights their cultural significance in art, where they were depicted as powerful symbols, underscoring their prominence in ecosystems.

Taxonomy and Phylogeny

Classification and Subspecies

The cave bear belongs to the family Ursidae within the order , and is placed in the genus Ursus as the species Ursus spelaeus, a binomial name formally established by German anatomist Johann Christian Rosenmüller in 1794 based on fossil remains recovered from caves in the Franken Mountains of Germany. Early taxonomic debates surrounded these fossils, with figures like Johann Friedrich Esper initially attributing them to polar bears (Ursus maritimus) transported by ancient floods, while Rosenmüller's detailed anatomical analysis confirmed them as a distinct extinct bear species rather than a variant of the extant (Ursus arctos). Older classifications sometimes subsumed U. spelaeus under the as a subspecies, such as Ursus arctos priscus, reflecting uncertainties in distinguishing it from Pleistocene forms until morphological and later molecular evidence supported its separation as a full species. Several subspecies of U. spelaeus are recognized, differentiated by regional adaptations and fossil distributions across Europe and western Asia during the Middle to Late Pleistocene. The nominate subspecies Ursus spelaeus spelaeus is associated with central European populations, characterized by typical cave bear traits in lowland to mid-altitude habitats. Ursus spelaeus eremus represents Mediterranean variants, often smaller in size and adapted to warmer, coastal environments in southern Europe. Ursus spelaeus ladinicus is identified from high-altitude Alpine sites, showing specialized features linked to montane isolation. Subspecies differentiation relies on morphological criteria, including variations in cranial architecture—such as neurocranium proportions, palate width, and zygomatic arch robustness—as well as tooth size and occlusal patterns that reflect dietary and environmental pressures. Geographic isolation further drove these distinctions, with populations in disparate regions like the Alps, Mediterranean basins, and eastern steppes exhibiting measurable divergences in skeletal metrics over time. These traits, analyzed through craniometrics and dental morphometrics, underscore the cave bear's adaptive radiation within its Pleistocene range.

Evolutionary Origins

The cave bear lineage traces its roots to the Late Miocene, approximately 10–12 million years ago, when early ursine bears like Indarctos arctoides emerged in Eurasia as part of the initial radiation of the Ursidae family. Fossils of I. arctoides, characterized by medium-sized builds and omnivorous dental features, have been recovered from sites across Europe and Asia, indicating a widespread ancestor that bridged earlier primitive bears such as Ursavus and more derived ursines. This period marked the development of key ursine traits, including enhanced masticatory adaptations that foreshadowed later specializations in the clade. By the Early Pleistocene, around 2 million years ago, the lineage had evolved into Ursus etruscus, a pivotal species regarded as the common ancestor of both cave bears and brown bears (Ursus arctos). U. etruscus exhibited transitional morphology, with fossils from Eurasian sites showing a shift toward larger body sizes and preliminary dental modifications for processing tougher vegetation amid cooling climates. This species dispersed across and , adapting to the onset of glacial cycles through increased reliance on forested and cave habitats. Key evidence comes from Early Pleistocene deposits, highlighting its role in the divergence of ursine sublineages. The split between the cave bear and lineages occurred approximately 1.2–1.6 million years ago, based on and genomic analyses that calibrate molecular clocks against Pleistocene fossils. This divergence likely happened in during the early Middle Pleistocene, with cave bears evolving distinct adaptations for colder, more seasonal environments as ice ages intensified. Early relatives, such as Ursus dolinensis from the Gran Dolina site in (dated to about 800,000–1,000,000 years ago), provide crucial fossil evidence of this transition, featuring primitive crania and that prefigure the speloid form. Over the Pleistocene, morphological evolution in the cave bear lineage emphasized herbivory through significant body size increases—reaching up to 1,000 kg in adults—and dental specializations for grinding, including enlarged molars with low-crowned, bunodont cusps suited for fibrous plants. These changes, evident in mandibular and cranial fossils from Middle Pleistocene strata, reflect selective pressures from glacial habitats favoring energy-efficient on during extended winters. Such adaptations distinguished cave bears from their more omnivorous relatives, solidifying their niche as predominantly herbivorous .

Genetic Insights

Genetic analyses of mitochondrial DNA (mtDNA) from cave bear remains have established that the species diverged from the brown bear (Ursus arctos) lineage approximately 1.2–1.6 million years ago, predating the diversification within brown bears. This split is supported by sequence data from multiple cave bear specimens dating between 130,000 and 20,000 years before present, revealing distinct phylogenetic separation with limited hybridization until late in the Pleistocene. Late Pleistocene populations exhibited notably low genetic diversity, with mtDNA haplotypes showing reduced variability compared to contemporaneous brown bears, indicative of isolation and demographic bottlenecks. Ancient DNA studies have further illuminated , demonstrating a prolonged decline in beginning around 25,000 years before the cave bear's approximately 24,000 years ago. This decline, tracked through mtDNA and nuclear markers from over 100 specimens across , contrasted with stable or fluctuating diversity in , suggesting cave bears faced unique pressures leading to reduced effective population sizes estimated in the low thousands. A 2019 mitogenomic analysis of 59 cave bear samples reinforced this, linking local extirpations and overall to human expansion during the , which disrupted between refugial populations and fragmented habitats. Genomic studies also indicate limited , with traces of cave bear DNA persisting in modern genomes at low levels (0.9–2.4%). Palaeoproteomic approaches have provided complementary insights by analyzing ancient proteins from fossils, bypassing DNA degradation issues in older specimens. In a 2025 study, from a ~1-million-year-old Ursus dolinensis from Gran Dolina, , confirmed its position as basal to the speloid cave bear lineage (Ursus spelaeus sensu lato), with phylogenetic trees derived from 147 protein sequences placing it as an early diverging relative to later cave bears and brown bears. This molecular evidence aligns with fossil records, highlighting the deep European roots of the cave bear radiation. Genome-wide sequencing of ancient cave bear DNA has revealed signatures of and persistently small effective population sizes, exacerbating vulnerability to environmental changes. Analysis of complete mitochondrial genomes and partial nuclear loci from late specimens showed elevated runs of homozygosity and distortions consistent with consanguineous mating in isolated groups, with effective population sizes (Ne) dropping below 1,000 individuals in terminal phases. These findings, corroborated by modeling, indicate that from low Ne contributed to the species' inability to adapt, distinct from the more resilient populations.

Physical Characteristics

Morphology and Size

The cave bear (Ursus spelaeus) possessed a robust, heavily built body with a barrel-shaped and relatively short limbs, reflecting adaptations to a primarily terrestrial existence in rugged, forested environments rather than arboreal activities. Its forelimbs were particularly strong and muscular, suited for digging into soil. Fossil evidence indicates significant size variation, with average adult males estimated at 400–500 kg in body mass and females at 225–250 kg, though exceptional individuals likely exceeded 700 kg based on femoral dimensions. When standing upright on their hind legs, adult males could attain heights of approximately 3–3.5 m, underscoring their imposing stature comparable to the largest modern bears. Body length in a quadrupedal posture typically ranged from 2 to 2.5 m for males. The species displayed pronounced , with s approximately 20–30% larger than females in linear skeletal measurements, a pattern evident in the greater size of canines and body mass reconstructions from femoral circumferences.00955-2) This dimorphism is further highlighted in cranial features, where skulls featured a more pronounced high-domed profile, shorter snout, and expansive for enhanced attachment to support powerful mastication.

Skeletal and Dental Adaptations

The cave bear (Ursus spelaeus) displayed cranial adaptations indicative of a primarily herbivorous diet, including reduced teeth and enlarged premolars suited for folivory, as evidenced by three-dimensional geometric morphometric analyses of crania from multiple European sites. These modifications shifted bite mechanics away from shearing toward grinding, with biomechanical simulations showing compromised skull strength due to expanded —an adaptation likely aiding heat conservation during prolonged periods. Dentally, the cave bear possessed a formula of I3/3, C1/1, P4/4, M2/3, featuring large molars with flat occlusal surfaces optimized for processing vegetation. Microwear analysis of these molars reveals high frequencies of scratches and pits consistent with an abrasive plant-based diet, including tough foliage and possibly grit-contaminated tubers, supporting predominantly herbivorous foraging with occasional omnivory. Recent studies highlight regional variations in microwear patterns. In the postcranial skeleton, the cave bear exhibited robust vertebral columns and reinforced rib cages, facilitating the curled posture adopted during hibernation and supporting the physiological demands of extended torpor. Skeletal remains frequently show pathologies, including healed fractures in long bones and ribs attributable to falls in karst environments or intraspecific aggression during mating seasons. These injuries often healed with minimal deformation, indicating extended lifespans and effective recovery mechanisms despite the bears' massive build exceeding 500 kg in adults.

Paleobiology

Diet and Foraging Behavior

The cave bear (Ursus spelaeus) exhibited a predominantly herbivorous diet, consisting primarily of plant matter such as grasses, herbs, tubers, fruits, and bark, with stable isotope analyses of bone collagen indicating that its dietary protein derived overwhelmingly from vegetable sources. Low δ¹³C values (typically -21 to -19‰) suggest consumption of C₃ from forested or shaded environments, while δ¹⁵N values (range 3.6–9.8‰ across European populations) indicate a primarily herbivorous with evidence of some omnivory in certain contexts. A 2025 study on specimens from Šalitrena Pećina in revealed regional variations, with δ¹³C values slightly lower than those of sympatric herbivores like (Cervus elaphus), pointing to a diet enriched in and occasional aquatic , but still overwhelmingly plant-based. Foraging strategies were adapted to seasonal availability in temperate Eurasian landscapes, involving in mixed forests and open meadows during warmer months to access fresh foliage and fruits, as evidenced by dental microwear textures showing from tough, fibrous . Microwear analysis from sites in South-Eastern Europe indicates locally adapted behaviors, such as increased consumption of hard-shelled nuts and seeds in forested areas during pre-hibernation hyperphagia, with patterns varying by site to exploit seasonal resources like spring herbs and autumn berries. Limited evidence from deposits further supports this, revealing undigested plant fibers and from grasses and forbs, consistent with opportunistic in meadows adjacent to habitats. The high-fiber nature of this diet necessitated physiological adaptations, including an enlarged gut capacity for microbial fermentation to extract nutrients from cellulose-rich vegetation, as inferred from skeletal proportions and isotopic signatures. During glacial maxima, such as the (ca. 26–19 ka BP), isotopic shifts toward more depleted δ¹³C values indicate reliance on coniferous bark and lichens in resource-scarce, open-steppe environments, reflecting dietary flexibility within herbivory constraints. Cave bears likely competed with ungulates like for shared in meadows and forest edges, as comparative δ¹⁵N profiles show overlapping resource use, potentially intensifying pressure during climatic fluctuations. Their dental structures, with low-crowned molars suited for grinding, facilitated efficient processing of this fibrous intake.

Hibernation and Physiology

The cave bear (Ursus spelaeus) underwent extended hibernation periods lasting up to 6-8 months annually, a adaptation suited to the prolonged cold seasons of the Pleistocene in Europe. During hibernation, individuals entered deep torpor states, reducing their metabolic rate by about 75% compared to active periods, which minimized energy expenditure while relying on accumulated body fat for sustenance. This physiological slowdown involved urea recycling, where nitrogenous waste was reincorporated into proteins, preventing muscle atrophy and supporting tissue maintenance despite immobility. Bone histology provides direct evidence of these cyclic physiological shifts, with lines of arrested growth (LAGs) in long bones and dental cementum indicating annual pauses in deposition corresponding to hibernation. These structures reflect slowed osteogenesis and metabolic activity during torpor, similar to patterns observed in modern hibernating bears. Fat accumulation, built primarily from a herbivorous diet rich in vegetation during active seasons, served as the primary energy source, enabling survival without foraging. Biomechanical analyses from a 2020 study reveal morphological adaptations enhancing efficiency, such as expanded for improved metabolic control during prolonged and conservation, but these may have reduced dietary versatility in responding to rapid fluctuations. Such traits optimized survival in stable cold environments but potentially constrained adaptability to environmental changes. The intensive use of caves as dens is evidenced by dense accumulations of skeletal remains, often representing mass mortality sites where bears succumbed during . These high demands during extended inactivity heightened vulnerability to , particularly if winters lengthened or food resources for pre- fattening diminished due to climatic shifts.

Reproduction and Growth

The of the cave bear (Ursus spelaeus) closely resembled that of its extant relatives, such as the (U. arctos), with mating occurring in late spring to early summer followed by delayed implantation of fertilized embryos. This adaptation allowed implantation to take place in the autumn, synchronizing birth with the onset of when females entered winter dens. Cubs, typically numbering 2 to 4 per , were born blind and altricial during mid-winter within these protected sites, weighing approximately 300–500 grams at birth—comparable to newborn brown bears. Evidence from assemblages, including associated mother-cub remains in caves, indicates that litters of at least two individuals were common, as seen in exceptionally preserved neonate skeletons from sites like Tecchia di Equi in . Births occurred after a period effectively extended by delayed implantation to about 7–8 months, aligning with the females' entry into . Parental care was extensive, with females hibernating alongside their cubs for the first year, providing and during this vulnerable period. Stable of reveals that cubs nursed for approximately 1.5 years, during which their δ¹⁵N values reflected the mother's physiological stress from , decreasing post-birth and increasing as solid foods were introduced after . This prolonged and denning strategy supported cub survival in harsh Pleistocene environments. Growth rates were rapid in early , with cubs achieving near-adult body size by 3–5 years, though skeletal maturity—marked by the completion of growth lines—was delayed until 10–14 years, later than in most modern . Tooth and wear patterns, analyzed via annuli, further indicate around 3–4 years for females and 4–6 years for males, aligning with the onset of breeding capability. Life expectancy in the wild is estimated at 19–25 years on average, inferred from cementum layers and growth mark counts in fossils, with exceptional individuals reaching up to 30–32 years before natural . High juvenile mortality likely limited many to shorter lifespans, but adults that survived early years benefited from the species' robust physiology.

Distribution and Environment

Geographic Range

The cave bear (Ursus spelaeus) primarily inhabited during the Middle and Late Pleistocene, with its range extending from the in the southwest to the in the east, and further into the and . The northern boundary of this distribution reached the and central regions of , where suitable forested and mountainous environments supported populations. Evidence indicates that the cave bear's presence also extended into Asia, particularly Siberia, where fossils have been documented in permafrost deposits; a notable 2025 discovery includes a 39,500-year-old specimen remarkably preserved with intact internal organs, highlighting the eastern limits of the species' range. The species' distribution fluctuated temporally in response to Pleistocene climatic cycles, with expansions across during warmer periods and contractions to southern refugia, such as the Mountains, during colder glacial phases. These shifts reflect adaptations to varying environmental conditions, influencing population connectivity and . Fossil records underscore the abundance of cave bears, with over 140,000 specimens documented from more than 140 sites across their range, the majority concentrated in karst cave systems that served as hibernation and breeding grounds. Several subspecies, such as U. s. spelaeus and U. s. ingressus, have been distinguished based on morphological variations tied to these regional populations.

Habitat Preferences

The cave bear (Ursus spelaeus) primarily inhabited forested uplands and regions across , where abundant formations provided natural systems for shelter, while largely avoiding open steppes and plains that lacked suitable cover and resources. These environments offered dense and hilly terrains that supported the bear's predominantly herbivorous diet, with landscapes being particularly favored due to their network of interconnected caves and stable geological features. Cave bears occupied a broad altitudinal range, from near in lowland areas to elevations up to 2,500 meters in mountainous uplands, allowing to varied topographic conditions within their preferred ecological niches. selection was influenced by proximity to resource-rich areas, such as meadows for foraging on and reliable water sources often accessible via systems or nearby in these regions. For denning, cave bears selected deep, interior cave chambers that provided thermal stability and protection from , as evidenced by large accumulations of skeletal remains in such sites across Eurasian formations, indicating repeated use over generations. These deep dens maintained consistent microclimates, essential for , and often preserved bone beds due to minimal disturbance and natural deposition.

Paleoenvironmental Context

The cave bear (Ursus spelaeus) existed during the Middle to , primarily from approximately 250,000 to 24,000 years ago, encompassing (MIS) 8 through 2, a period characterized by significant Pleistocene climate oscillations between glacial and phases. Fossils indicate that populations were most abundant during the relatively warmer conditions of MIS 3 (approximately 57,000 to 29,000 years ago), an interstadial phase within the that allowed for expanded habitats suitable for their herbivorous lifestyle. During colder stadials, such as MIS 4 and the onset of MIS 2, their presence became more restricted, reflecting sensitivity to broader climatic shifts. As a , the cave bear played a key role in Pleistocene ecosystems, particularly in mixed forest-tundra ecotones across , where it coexisted with other including woolly mammoths (Mammuthus primigenius) and (Rangifer tarandus). These environments featured diverse , from open grasslands to wooded areas, supporting the bear's reliance on plant matter such as herbs, roots, and fruits, which it consumed in large quantities to sustain its massive body size. Interactions within this megafaunal community likely influenced nutrient cycling and dynamics, with cave bears contributing to and soil disturbance through foraging activities. Glacial advances during colder phases, such as those in MIS 4 and leading into the , profoundly impacted cave bear habitats by reducing vegetation productivity through lowered temperatures and expanded ice sheets. This led to sparser plant cover in lowlands, compelling populations to undertake altitudinal migrations toward higher elevations where milder microclimates preserved more favorable grounds, as evidenced by concentrations in montane caves. Such shifts highlight the species' to fluctuating environmental pressures but also its to prolonged cooling. Proxy data from pollen preserved in cave sediments provide direct insights into local floral changes associated with cave bear occupations. For instance, analyses from sites in the Lombardian Pre-Alps reveal shifts from open steppe-like vegetation during colder intervals to more wooded assemblages during interstadials, correlating with bear population peaks. These records indicate that cave interiors trapped pollen from surrounding landscapes, documenting transitions in dominant plant taxa like grasses and shrubs that formed the base of the cave bear's diet.

Human Interactions

Archaeological Discoveries

The discovery of cave bear fossils has been pivotal in understanding their , with major sites revealing extensive accumulations of remains that illuminate behaviors and . In the Conturines Cave, located at an elevation of approximately 2,800 meters in the of , excavations since 1987 have uncovered a significant accumulation of Ursus ladinicus bones, representing dozens of individuals from over 50,000 years ago. These findings, preserved in a high-altitude environment, include skeletal elements indicating repeated use of the cave for prior to 50,000 years . Similarly, Peștera Urșilor () in western has yielded over 11,500 skeletal elements from at least 105 individuals, dating primarily to the , providing one of the largest assemblages in and evidence of dense local populations. A particularly remarkable recent find occurred in 2025, when a 39,500-year-old cave bear mummy was recovered from Siberian permafrost, featuring exceptionally preserved fur, soft tissues, and internal organs, marking the best-preserved Ursus spelaeus specimen to date. This discovery highlights the expanding range of cave bear fossils into northern Asia and underscores the role of thawing permafrost in exposing new material. Archaeological evidence also documents co-occurrence of cave bears with Neanderthals and early modern humans, as indicated by cut marks on bones from multiple sites. For instance, at sites like Schöningen in Germany, dated to around 320,000 years ago, cut marks on cave bear phalanges and crania suggest early hominins skinned bears for fur, potentially predating Neanderthal dominance in Europe. In southern Alpine caves, taphonomic analyses reveal percussion and cut marks on Ursus spelaeus remains, pointing to Neanderthal hunting or scavenging activities during the Middle Paleolithic. Later, in Romania's Peștera cu Oase, cave bear bones intermingle with early modern human fossils from approximately 40,000 years ago. Taphonomic studies differentiate natural accumulation from human-influenced deposits, aiding in interpreting site formation processes. Many cave bear bone beds, such as those in Conturines Cave, exhibit mortality profiles consistent with natural traps—vertical shafts or chambers where hibernating bears fell or died from stress, leading to attritional death assemblages dominated by prime-age adults without significant beyond self-induced damage. In contrast, human-accumulated sites show concentrated cut and percussion marks, burn traces, and association with lithic tools, as seen in Peștera Urșilor, where selective bone breakage indicates butchery rather than random predation or trampling. Cave bears themselves contributed to taphonomic signatures through intra-species scavenging and dismemberment of carcasses during hibernation awakenings. Preservation of cave bear fossils owes much to specialized environmental conditions, including and microclimates. In Siberian sites, 's sub-zero temperatures and low oxygen levels have enabled the rare mummification of soft tissues, as in the 2025 discovery, by halting bacterial and maintaining structural integrity. Within European karst caves like Conturines and Peștera Urșilor, stable microclimates—characterized by consistent humidity, minimal air circulation, and temperatures near freezing—facilitate and prevent post-depositional degradation, allowing long-term fossil stability despite exposure to minor fluctuations. These conditions contrast with surface sites, where would destroy remains, emphasizing and as critical taphonomic windows into Pleistocene .

Cultural and Symbolic Role

Depictions of cave bears in art are rare, with only about 60 representations identified across European sites, underscoring their limited role as prey but suggesting a special symbolic status among artists. In , , dated to around 36,000–30,000 years ago, several cave bear figures appear, including three outlined in red ochre near the entrance and shaded engravings in deeper chambers, often emphasizing the animal's distinctive high forehead and robust form. These artworks, rendered with exceptional detail, portray bears in profile or as isolated motifs, potentially evoking them as spirit animals or totems rather than everyday . Evidence for bear cults emerges from arranged bone deposits in Paleolithic sites, indicating ritualistic human interactions with cave bears during the . In , clusters of cave bear skulls and long bones were deliberately placed in niches or alcoves, separate from natural hibernation accumulations, suggesting ceremonial deposition rather than utilitarian use. Similar findings in central European caves, such as Drachenloch in , include bear skulls aligned in stone enclosures and thighbones inserted through cheekbones, interpreted by some scholars as totemic rituals honoring the bear's power or spiritual essence. These patterns, spanning and occupations from approximately 50,000 to 20,000 years ago, point to possible shamanistic practices where cave bears symbolized strength, as rebirth, or clan ancestry in . The cave bear's legacy persists in modern , where bears often embody ancestral figures or mythical progenitors, echoing prehistoric reverence. In Basque traditions, for instance, humans are mythically descended from bears, a motif preserved in the "" (ATU 301), which features hybrid bear-human offspring and reflects pan-European animist beliefs in ursine genealogy. This influence extends to scientific nomenclature, with the species named Ursus spelaeus in 1794 by Johann Christian Rosenmüller, deriving "spelaeus" from Greek for "cave" to denote the abundance of fossils unearthed in subterranean sites. Ethnographic parallels to these prehistoric practices appear in historic Eurasian cultures, where bear reverence manifests in elaborate rituals treating the animal as a sacred intermediary. Among the Sami of northern , the bear hunt concluded with feasts and "birching" ceremonies to honor the spirit and ensure its rebirth, mirroring potential totemic deposits. Similarly, the Ainu of and Gilyaks of raised bear cubs as divine guests before ritual , emphasizing reciprocity and mediation—patterns linked by archaeologists to Magdalenian-era (ca. 20,000 years ago) bear cults in Europe, including art and bone arrangements at sites like Trois-Frères Cave. These traditions highlight a continuum of bear symbolism across , from prehistoric symbolic roles to enduring cultural .

Extinction

Timeline and Patterns

The cave bear (Ursus spelaeus) flourished across and parts of from approximately 300,000 years ago through much of the , with abundant fossil records indicating stable and widespread populations until around 50,000 years (BP). Regional extinctions began emerging around 40,000 years BP, particularly in southern European regions such as the and the Mediterranean, where radiocarbon-dated fossils show a marked reduction in occurrences. The final remnants of cave bear populations persisted until approximately 24,000 years BP in refugial areas like the and the , as evidenced by direct of the latest fossils from these locales, which cluster between 26,000 and 24,000 years BP. These dates confirm that was not synchronous but varied regionally, with central and eastern European sites yielding some of the most recent remains. Population decline exhibited distinct patterns across the continent: gradual in , where records show a progressive thinning over millennia, contrasted with more abrupt disappearances in eastern regions, reflecting asynchronous regional dynamics. Demographic evidence from stratified sites indicates decreasing abundance after 30,000 , with fewer individuals per assemblage and reduced site occupancy signaling a broader contraction. Genetic analyses of late-phase remains reveal bottlenecks that further underscore the demographic stress during this period.

Proposed Causes

The extinction of the cave bear (Ursus spelaeus) has been attributed to a combination of environmental, ecological, and anthropogenic pressures, with multiple hypotheses emphasizing their interplay rather than a single dominant factor. One prominent explanation centers on during the (approximately 26,000 to 19,000 years ago), when severe cooling and led to widespread vegetation reduction across , diminishing food availability for the herbivorous cave bear. This period of intensified cold not only contracted suitable habitats but also exacerbated the bears' reliance on , as prolonged winters forced extended periods of with limited energy reserves. A 2020 biomechanical study highlighted how the cave bear's anatomical adaptations for prolonged —such as enlarged nasal sinuses that facilitated air warming and reduced heat loss—created a with their strictly herbivorous diet, rendering them particularly vulnerable to the nutritional scarcity induced by glacial cooling. These adaptations, while advantageous for surviving short-term cold snaps in earlier interstadials, became maladaptive during the sustained harsh conditions of the , as the bears struggled to accumulate sufficient fat from sparse herbaceous vegetation before entering . Dietary inflexibility further compounded these challenges, as stable isotope analyses of bone collagen have revealed that cave bears maintained a predominantly even under environmental stress, limiting their ability to switch to alternative food sources like or scavenged remains. A 2025 study on Serbian specimens from Šalitrena Pećina Cave showed varied δ¹³C and δ¹⁵N values consistent with a flexible across diverse s, suggesting adaptability within herbivory that nonetheless proved insufficient against broader climatic and habitat pressures. This contrasted with more opportunistic modern bears, contributing to population declines as ecosystems shifted toward less productive tundra-steppe biomes. Human activities also played a significant role, particularly through competition for denning sites and direct predation by early Homo sapiens. A 2019 mitogenomic analysis of over 100 ancient cave bear samples indicated stable population sizes from 200,000 to 50,000 years ago, followed by sharp declines around 40,000 years ago—coinciding with the arrival of modern humans in —who likely competed for resources and hunted bears, accelerating local extirpations. Evidence from archaeological sites shows overlapping use of caves, implying that human expansion fragmented bear habitats and increased mortality rates during vulnerable periods. Integrated models propose that low , evidenced by a 25,000-year decline in variability preceding full , amplified these stressors by reducing the population's resilience to combined climatic and anthropogenic pressures. This genetic bottleneck, likely initiated by during earlier Pleistocene fluctuations, made cave bear populations less capable of evolving behavioral or physiological adaptations, turning incremental environmental changes into existential threats. These hypotheses align with the observed timeline of regional declines, underscoring a multifaceted process rather than abrupt catastrophe.

Modern Research and Discoveries

In 2025, herders in discovered a remarkably intact cave bear (Ursus spelaeus) specimen preserved in , dated to approximately 39,500 years old, marking it as the best-preserved cave bear found to date. This mummified remains, including internal organs and soft tissues, have enabled unprecedented analyses, such as detection through metagenomic sequencing and dietary reconstruction via preserved gut contents, providing new insights into the and of late Pleistocene populations. Advancements in palaeoproteomics have further illuminated cave bear , with a 2025 study analyzing enamel proteins from Early to specimens across to construct a protein-based phylogeny. This approach resolved ambiguities in the early "speloid" lineage's divergence, revealing genetic adaptations to hibernal lifestyles and confirming the cave bear's deep roots in Eurasian ursine where preservation was insufficient. Recent stable isotope analyses of from Serbian cave bear sites, combined with dental microwear studies from South-Eastern European and Caucasian populations (2023–2025), have uncovered significant intra-population dietary variations, including shifts toward more herbaceous in response to local environmental changes. These findings highlight dietary flexibility within a herbivorous framework, challenging uniform models of specialization and underscoring population-specific vulnerabilities near the despite some adaptability. Ongoing genomic sequencing efforts, building on metagenomic libraries from Pleistocene remains, continue to map cave bear for potential applications in understanding resilience, while climate modeling projects integrate isotopic data with paleoenvironmental simulations to reconstruct habitat suitability and predict analogous threats to modern ursids under global warming.

References

  1. https://phys.org/news/2021-02-dna-year-old-bone-reveals-oldest.html
  2. https://www.sciencedirect.com/science/article/pii/S163106830900133X
  3. https://pmc.ncbi.nlm.nih.gov/articles/PMC2741261/
  4. https://digitalcommons.usf.edu/cgi/viewcontent.cgi?article=5525&context=kip_articles
  5. https://pmc.ncbi.nlm.nih.gov/articles/PMC2206582/
  6. https://www.sciencedirect.com/science/article/pii/S096098220100046X
  7. https://www.nationalgeographic.com/science/article/cave-bears-ancient-dna-paleogenomics-extinction
  8. https://academic.oup.com/mbe/article/27/5/975/1025434
  9. https://www.buffalo.edu/news/releases/2020/04/002.html
  10. https://archeologie.culture.gouv.fr/chauvet/en/cave-bears
  11. https://www.gbif.org/species/144099575
  12. https://www.zobodat.at/pdf/Abh-Naturhist-Ges-Nuernberg_45_0199-0214.pdf
  13. https://palaeo-electronica.org/content/2012-issue-2-articles/263-cave-bears-from-poland
  14. https://www.sciencedirect.com/science/article/abs/pii/S1040618210004945
  15. https://www.aaps-journal.org/pdf/JPS.C.21.0002.pdf
  16. https://www.sciencedirect.com/science/article/abs/pii/S1040618211001303
  17. https://www.researchgate.net/publication/310463283_On_Factors_that_Influence_the_Morphology_of_the_Cave_Bear_Dentition_and_a_Study_of_the_Geographical_Variation_in_the_Lower_Carnassial
  18. https://digitalcommons.usf.edu/kip_articles/3558/
  19. https://www.researchgate.net/publication/239750226_A_Review_of_Bear_Evolution
  20. https://www.biorxiv.org/content/10.1101/2025.10.05.680322v1.full.pdf
  21. https://www.tandfonline.com/doi/abs/10.1080/08912963.2022.2067993
  22. https://www.cell.com/current-biology/fulltext/S0960-9822(21)00138-X
  23. https://www.biorxiv.org/content/10.1101/2025.10.05.680322
  24. https://www.sciencedirect.com/science/article/abs/pii/S0031018221005721
  25. https://academic.oup.com/mbe/article/19/11/1920/1012476
  26. https://pubmed.ncbi.nlm.nih.gov/20335279/
  27. https://www.nature.com/articles/s41598-019-47073-z
  28. https://pmc.ncbi.nlm.nih.gov/articles/PMC6590514/
  29. https://www.biorxiv.org/content/10.1101/2025.10.05.680322v1
  30. https://royalsocietypublishing.org/doi/10.1098/rsbl.2015.0822
  31. https://www.researchgate.net/publication/233057513_Limb_bone_proportions_and_body_mass_of_the_cave_bear_Ursus_spelaeus
  32. https://www.prehistoric-wildlife.com/species/ursus-spelaeus-cave-bear/
  33. https://institutions.ville-geneve.ch/fileadmin/user_upload/mhn/documents/Museum/Revue_de_Paleo/379-393_Baryshnikov_etal.pdf
  34. https://www.researchgate.net/publication/285723380_Ursus_deningeri-spelaeus_group_from_Cere_Cave_Veneto_North_Italy_in_the_new_evolutionary_frame_of_the_cave_bear_Part_one_Skulls_and_mandibles
  35. https://www.researchgate.net/publication/259141358_Functional_morphology_of_the_cave_bear_Ursus_spelaeus_cranium_A_three-dimensional_geometric_morphometric_analysis
  36. https://www.science.org/doi/10.1126/sciadv.aay9462
  37. https://pmc.ncbi.nlm.nih.gov/articles/PMC7775984/
  38. https://www.researchgate.net/publication/324784576_Isotopic_insights_on_cave_bear_palaeodiet
  39. https://www.pnas.org/doi/10.1073/pnas.0711063105
  40. https://www.tandfonline.com/doi/full/10.1080/14614103.2025.2521944
  41. https://www.sciencedirect.com/science/article/pii/S0031018225004857
  42. https://www.researchgate.net/publication/394501105_Dental_microwear_of_cave_bear_Ursus_spelaeus_reveals_locally_adapted_foraging_strategies_in_South-Eastern_Europe_during_late_MIS_3
  43. https://www.sciencedirect.com/science/article/abs/pii/S1040618219307931
  44. https://www.nature.com/articles/s41598-020-62990-0
  45. https://ruc.udc.es/rest/api/core/bitstreams/6393ab6f-9f29-4e39-a2e0-27f3db3410ee/content
  46. https://www.pnas.org/doi/10.1073/pnas.0907373106
  47. https://www.pnas.org/doi/10.1073/pnas.0800915105
  48. https://frontiersinzoology.biomedcentral.com/articles/10.1186/s12983-019-0312-2
  49. https://journals.plos.org/plosone/article?id=10.1371/journal.pone.0206791
  50. https://mountainscholar.org/items/6cc3a9e7-d7d7-4bd7-a0a4-03bbf31be82b
  51. https://www.nature.com/articles/s41598-024-60222-3
  52. https://pmc.ncbi.nlm.nih.gov/articles/PMC11059340/
  53. https://www.researchgate.net/publication/287872358_The_cave_bear_Ursus_spelaeus_from_Goyet_Belgium_The_bear_den_in_Chamber_B_bone_horizon_4
  54. https://www.sciencedirect.com/science/article/abs/pii/S1040618210003848
  55. https://journals.openedition.org/quaternaire/20451
  56. https://pmc.ncbi.nlm.nih.gov/articles/PMC5460516/
  57. https://pmc.ncbi.nlm.nih.gov/articles/PMC6248942/
  58. https://bmcecolevol.biomedcentral.com/articles/10.1186/s12862-015-0521-z
  59. https://www.researchgate.net/publication/323284525_Longevity_and_life_history_of_cave_bears_-_a_review_and_novel_data_from_tooth_cementum_and_relative_emergence_of_permanent_dentition
  60. https://www.researchgate.net/figure/Map-of-cave-bear-distribution-orange-distribution-of-Ursus-spelaeus-sensu-lato_fig2_24024257
  61. https://www.sciencedirect.com/science/article/pii/S0031018224004061
  62. https://allthatsinteresting.com/siberia-mummified-cave-bear
  63. https://www.researchgate.net/publication/230049129_The_first_record_of_spelaeoid_bears_in_Arctic_Siberia
  64. https://pmc.ncbi.nlm.nih.gov/articles/PMC5059403/
  65. https://www.researchgate.net/publication/45098142_First_fossil_evidence_of_an_%27interglacial_refugium%27_in_the_Pyrenean_region
  66. https://www.sciencedirect.com/science/article/abs/pii/S1040618213002711
  67. https://link.springer.com/article/10.1007/s12520-022-01624-0
  68. https://journals.openedition.org/quaternaire/16605
  69. https://www.tandfonline.com/doi/full/10.1080/10256016.2024.2376730
  70. https://www.jstor.org/stable/23735450
  71. https://www.researchgate.net/publication/309134526_Retreat_and_extinction_of_the_Late_Pleistocene_cave_bear_Ursus_spelaeus_sensu_lato
  72. https://www.nature.com/articles/s41598-019-42152-7
  73. https://onlinelibrary.wiley.com/doi/abs/10.1111/j.1502-3885.2008.00071.x
  74. https://www.cambridge.org/core/journals/radiocarbon/article/radiocarbon-constraints-on-the-age-of-the-worlds-highestelevation-cavebear-population-conturines-cave-dolomites-northern-italy/EC95C294BFF5C5C5EAD40A0F419B3608
  75. https://www.researchgate.net/publication/396864209_The_Conturines_Cave_in_the_Dolomites_and_the_Alpine_climate_of_the_last_70000_years_Reconstruction_of_the_climate_based_on_fossilized_vertebrate_remains_from_caves
  76. https://www.researchgate.net/publication/313405854_SIZE_OF_THE_CAVE_BEAR_POPULATION_AND_SKELETAL_DISTRIBUTION_FROM_URSILOR_CAVE_ROMANIA
  77. https://www.sci.news/archaeology/schoningen-bear-skins-11508.html
  78. https://www.sciencedirect.com/science/article/abs/pii/S030544031730184X
  79. https://bioone.org/journals/acta-palaeontologica-polonica/volume-61/issue-2/app.00201.2015/Fossil-Population-Structure-and-Mortality-Analysis-of-the-Cave-Bears/10.4202/app.00201.2015.full
  80. https://www.researchgate.net/publication/235768881_New_insights_on_the_cave_bear_population_from_the_ursilor_cave_Romania
  81. https://www.persee.fr/doc/mhnly_1762-8024_2004_act_2_1_1411
  82. https://arstechnica.com/science/2020/09/beautifully-preserved-cave-bears-emerge-from-siberian-permafrost/
  83. https://onlinelibrary.wiley.com/doi/10.1002/jqs.3712
  84. https://artsandculture.google.com/story/the-bears-of-chauvet-cave-grotte-chauvet/owWRJA5ZuHCZJA?hl=en
  85. https://archeologie.culture.gouv.fr/chauvet/en/bears-recess
  86. https://www.researchgate.net/publication/358045071_The_cave_bear_in_Chauvet_Cave
  87. https://www.penn.museum/sites/expedition/the-cult-of-the-cave-bear/
  88. https://www.researchgate.net/publication/293076364_Cave_bear_worship_in_the_palaeolithic
  89. https://www.academia.edu/111431103/_The_Bears_Son_Tale_Traces_of_an_ursine_genealogy_and_bear_ceremonialism_in_a_pan_European_oral_tradition
  90. http://www.macroevolution.net/ursus-spelaeus.html
  91. https://www.laits.utexas.edu/sami/diehtu/siida/religion/bear.htm
  92. https://www.sciencedaily.com/releases/2017/08/170824151007.htm
  93. https://www.dw.com/en/genetic-research-blames-humans-for-prehistoric-bear-demise/a-50045046
  94. https://www.researchgate.net/figure/Radiocarbon-dates-of-the-latest-cave-bears-Ursus-spelaeus-in-Central-Europe-for-the_fig11_273001950
  95. https://www.britannica.com/animal/cave-bear
  96. https://pmc.ncbi.nlm.nih.gov/articles/PMC7112751/
  97. https://www.researchgate.net/publication/383183295_Dental_microwear_analysis_of_Kudaro_cave_bears_Insights_into_dietary_evolution_from_the_Middle_to_Late_Pleistocene
  98. https://phys.org/news/2025-08-lessons-cave-extinction-endangered.html
Add your contribution
Related Hubs
User Avatar
No comments yet.