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The present is a palaeobiological and taphonomic analysis of a Marine Isotope Stages (MIS) 4–3 (Late Pleistocene) assemblage of animal remains and hominin artefacts from layers 7–5 of Biśnik Cave, Czestochowa Upland, Poland. The analysis indicates that the bone assemblage is the result of a time-averaged palimpsest of both biotic and episodic abiotic events, the former consisting of many successive generations of animals and hominins that frequented the cave, and the latter including hydraulic winnowing. In fact, the taphonomic history of the fossil assemblage from Biśnik Cave's layers 7–5 is partially obscured by the overprint of hydraulic winnowing, which purportedly removed a certain amount of the original specimens. Besides evidence of cave bear deaths from non-violent, hibernation-related mortality and of occupation by generations of denning wolves and hyaenas, there is a wealth of flint artefacts, alongside remains of a few fireplaces and of a structure built in the cave by hominins to partition the cave chambers. The studied layer contains an impressive number of shed antlers, primarily of the red deer Cervus elaphus. Crocuta crocuta spelaea is normally held responsible for such accumulations of shed antlers in various European caves; Biśnik Cave's layers 7–5 will therefore simply add to the list. However, the role of accumulator of shed antlers attributed to the Pleistocene spotted hyena does not match the behaviour of its modern counterpart and seems not accounted for metabolically. The only reasonable alternative is that the antlers were collected by hominins. From this alternative perspective the cave would have functioned as a warehouse, where naturally shed antlers were stored as raw material, potentially to be shaped into tools and/or employed as tools to make other tools. The palaeobiological and taphonomic analysis presented here provides new insights into the succession of pre- and postdepositional events that involved the bone remains accumulated in the cave, as well as into the interactions between the animals and hominins of the time. More importantly, if hominins, and not hyaenas, were responsible for the amassment of the shed antlers in Biśnik Cave, this study raises doubts as to the hyaenid or human origin of other similar cave accumulations of shed antlers throughout Europe.

Panthera spelaea was recorded in Poland from 18 open-air and 42 cave sites dated in the range 750–28 ka. Most of these records are located in southern Poland (Silesia) and neighbouring areas. Among them, the find of Panthera spelaea in Kozi Grzbiet mentioned here for the first time is one of the oldest European records of the species. Most of the obtained AMS dates are concentrated on the second half of MIS 3 and showed that the Pleistocene lion disappeared more or less synchronously across Poland. The findings of its remains from open-air sites are mostly accidental discoveries during field works, while those in caves are mostly connected with archaeological research and the exploitation of cave sediments for producing fertilizers. Aside of a few juvenile bones and milk teeth, the remains of adult individuals, mostly males, predominate in the studied sample. Their behavior to hunting cave bear during the times, when the other, more typical food sources were scarce, tendention to hunt very big preys and went into the conflicts with other carnivores than females do, resulted in higher injuries and mortalities. Within the species, three chronosubspecies were recognized, with P. s. fossilis recorded from 7 sites, dated between 750 and 240 ka and P. s. intermedia only partially recognized in Wierzchowska Gorna Cave. The most numerous are the remains of P. s. spelaea which were found in 48 localities dated between 150 and 28 ka. With the exception of four caves, the Polish finds of P. spelaea are generally less abundant at sites where they occur. Older P. s. fossilis, dated on MIS 19–12, was a large form with robust stature and broad cheek teeth. Younger P. s. fossilis dated on MIS 11–9 was represented by a large and massive specimens, but with more advanced morphological dental features. Polish individuals of P. s. spelaea were slightly smaller and less massive than P. s. fossilis, with narrower cheek teeth. Since MIS 3, a dwarf and gracile specimens appeared, which is correlated with the genetic turnover ca. 48–45 ka. The Pleistocene lion was one of the earliest disappeared large carnivores predated only by a cave hyena. There are no direct evidences of P. spelaea encounters with humans. The extinction of P. spelaea is broadly correlated with the general collapse of the “mammoth steppe” ecosystem and was resulted of multiple reasons like climatic changes, re-building of herbivore guilds, competition and human pressure. Wolf and, to a lesser extent, bears were the main competitors for the Pleistocene lion in Poland.

The Sobrarbe-Pirineos UNESCO Global Geopark shows an extremely well-developed underground karst relief as a result of the great abundance and thickness of its limestone formations. The most important Pleistocene vertebrate site within the Geopark is Coro Tracito Cave at Tella. The fossil association is made up exclusively of bones belonging to Ursus spelaeus from the upper Pleistocene, accumulated over several thousand years. Based on scientific analysis of the fossil bones, an interesting public outreach project has been organized, involving the refurbishment of the site within the cave and the creation of a permanent exhibition called the Tella Cave Bear Museum. These two infrastructures are visited by thousands of tourists each year and constitute the main geoscientific tourist attraction of the Sobrarbe-Pirineos Geopark.

The lower molar (m1) of cave bears from Late Pleistocene localities of the Urals was studied employing the methods of traditional morphometry and geometric morphometrics. On the basis of the size and shape variation of m1, the small cave bear (Ursus ex gr. savini-rossicus) was found to have been a part of the faunas from the caves Skazka, Viasher, Dynamitnaya, Chudesnitsa, and Chernye Kosti. The small cave bear presence in faunas from the Medvezhya, Makhnevskaya Ledyanaya, Asha 1, Ignat’evskaya, and Barsuchii Dol caves was confirmed as well. The species range of the small cave bear encompassed the Northern, Middle, and Southern Urals in the Late Pleistocene. The ranges of the small cave bear and cave bear (Ursus kanivetz) overlapped from the beginning (marine isotope stage 5e) to the middle (middle marine isotope stage 3) of the Late Pleistocene.

International audience; The Chauvet-Pont-d'Arc Cave (Ardeche, France) contains some of the oldest Paleolithic paintings recorded to date, as well as thousands of bones of the extinct cave bear, and some remains and footprints of other animals. As part of the interdisciplinary research project devoted to this reference cave site, we analyzed a coprolite collected within the deep cave. AMS radiocarbon dating of bone fragments from the coprolite yielded an age of 30,450 +/- 550 RC yr. BP (AAR-19656; 36,150-34,000cal BP), similar to ages assigned to Paleolithic artwork and cave bear remains from the same cave sector. Using high-throughput shotgun DNA sequencing, we demonstrated a high abundance of canid DNA and lesser amounts of DNA from the extinct cave bear. We interpret the sample as feces from a canid that had consumed cave bear tissue. The high amount of canid DNA allowed us to reconstruct a complete canid mitochondrial genome sequence (average coverage: 83x) belonging to a deeply divergent Glade of extinct mitochondrial wolf lineages that are most closely related to coeval (-35 ka) Belgian wolves. Analysis of the nuclear genome yielded a similar coverage for the X chromosome (2.4x) and the autosomes (range: 2.3-3.2x), indicating that the Chauvet canid was a female. Comparing the relationship of the nuclear genome of this specimen with that of a variety of canids, we found it more closely related to gray wolves' genomes than to other wild canid or dog genomes, especially wolf genomes from Europe and the Middle East. We conclude that the coprolite is feces from an animal within an extinct wolf lineage. The consumption of cave bear by this wolf likely explains its intrusion into the dark cave sectors and sheds new light on the paleoecology of a major cave site.

The first paleoparasitological analysis of coprolites of a cave bear (Ursus kanivetz Vereshchagin, 1973) was performed. Material for research was obtained from different deeps at the excavation of Ignatievskaya Cave (Chelyabinsk Region, Southern Urals, Russia, 54°53’N 57°46’E). Fossil coprolites were find from the excavations N 1-2, situated et the 120 meters from the entry of cave. The depth of excavations was 1.9 m. Radiocarbon analysis show that the age of cave bear bones was more than 27500 years. Coprolites were analyzed by combine method and sedimentation. Slides were viewed under the Meiji microscope with Х100 and Х400 magnification and made a pictures by the Vision camera. Eggs were defined using scientific descriptions available in the national and foreign literature. As a result from the paleoparasitological analysis, the presence Nematodes eggs was detected. Eggs are belonged to Baylisascaris transfuga Rudolphi, 1819 – specific parasite of Ursidae. Obviously, that helminth infected different species of bears already at the Late Pleistocene period and had a wide spreading. This is the first finding of the nematode B. transfuga of Pleistocene age and the first finding of parasites in the cave bear coprolites.

Imanay Cave is located in the Southern Urals (53°02' N, 56°26' E), at 420 m.a.s.l. A 9.5 m2 trench was exca-vated in the grotto in the inner part of the cave to examine the sediments. The accretion thickness was 1.2 m. The taphocoenosis of the Imanay Cave is of the Pleistocene age and contains about 10,000 specimens of bone re-mains of large mammals. They mostly belong to small cave bear (U. ex gr. savini-rossicus), and the remaining bones — to species of the mammoth faunal complex (Lepus sp., Castor fiber, Marmota bobak, Canis lupus, Cuon alpinus, Vulpes vulpes, V. corsac, Meles sp., Gulo gulo, Martes sp., Mustela sp., Ursus kanivetz, U. arctos, U. thibetanus, Panthera ex gr. fossilis-spelaea, Mammuthus primigenius, Equus ferus, Coelodonta antiquitatis, Alces alces, Bison priscus, Saiga tatarica, Ovis ammon). In the layer with the bones, Middle Paleolithic stone artifacts were found, including several bifacial points. These tools have analogies in the Middle Paleolithic sites of the Caucasus region and Crimea. During excavations of the cave, the skull of a cave bear with artificial damage was found. The study of the artificial perforation on the skull was the purpose of the present paper. On the basis of dimensional and morphological features, it was established that the skull belongs to a small cave bear (U. ex gr. savini-rossicus). The skull was directly AMS radiocarbon dated to 34 940 ± 140 BP, IGANAMS-5652. Analysis of the growth layers in the teeth revealed that the animal died in winter at an age of 9-10 years. Trace evidence analysis showed, that the hole in the parietal region of the skull was made by a sharp bifacial flint point similar to the Middle Paleolithic points found in the cultural layer of the cave. The animal was killed during winter hiberna-tion, most probably by stabbing with a spear. This is the first direct evidence of human hunting of a small cave bear. With the abundance of cave bear bones, the skull with the hole in it is the only evidence of human impact on this animal. There are no bones with traces of butchering and harvesting of the bone marrow.

Introduction. Imanai-1 Cave is a new monument of the Middle Paleolithic in the Southern Urals. It was discovered by the authors in 2009 and is located in the west of the Ural mountain system, in the interfluve of the Belaya and Nugush Rivers, on the border of the mountain-forest and steppe zones. Goals. The paper aims to introduce preliminary results of archaeological investigations into scientific discourse. Results. The cave is of a tunnel type, its 70 m long passage ending with a far hall which contained bones of a small cave bear and a cave lion. The monument is multi-layered. The first cultural horizon contained 399 items of stone and bone. Tools make up to 60 % of all stone products, while cores and scales are absent, therefore, primary and secondary processing was carried out outside the far hall. The stone industry is characterized by the use of shards and amorphous flint chips. The working areas were made out with monofacial and bifacial retouching, incisal cleavage. The tools are of the following types: 3 Mousterian bifacial points, 4 convergent side-scrapers with bifacial processing, butt knives, some with bifacial processing ― 6 items, carvers on fragments and amorphous chips ― 229 items (59 %), points ― 19 items (5 %), tools with a thorn ― 13 items (3 %), incisors ― 21 items (5 %). At the base of the first cultural horizon, a skull of a small cave bear with an artificial hole made with a stone spearhead was found. The industry of the site has numerous analogies at the Ilskaya-1 site in the North Caucasus and in the materials of the upper layer of the Kiik-Koba grotto in the Crimea, as well as at other sites of the Middle Paleolithic of the Tayacian tradition. Three uncalibrated dates show the interval from 26 to 42 thousand years. This indicates the finale of the Mousterian era.

The variability of cheek teeth over time was studied with the use of univariate and multivatiate statistical analyses for 1370 tooth specimens of the Kudaro cave bear (Ursus kudarensis) (Baryshnikov 1985) collected in the Kudaro 1 and Kudaro 3 caves in the Caucasus. The material examined included two chronosubspecies: U. k. praekudarensis (middle part of the Middle Pleistocene, nearly 350 000 years ago) and U. k. kudarensis (Late Pleistocene, nearly 120 000–40 000 years ago), as well as a sample of intermediate stratigraphic position referred to as a “transitional form” (the end of the Middle Pleistocene, nearly 250 000 years ago). Our study detected no single pathway of evolutionary change in the cheek teeth of U. kudarensis, instead showing specific modifications for the premolars, molars, and even individual teeth (e.g., m3) in the context of high individual variability. In size, the teeth of U. k. praekudarensis occupied an intermediate position between the more robust teeth of the “transitional form” and the smaller teeth of U. k. kudarensis. By the range of morphometric characters (size and crown proportions), the “transitional form” was more closely related to U. k. kudarensis than to U. k. praekudarensis. At the same time, the parameters of allometric variation of the “transitional form” either related it to U. k. praekudarensis or were of the original nature. The calculated rate of evolution of the metric characters was found to be dissimilar not only for different teeth but also for different measurements of the same tooth crown, and these modifications themselves were oppositely directed. A comparison with the main evolutionary trends in European cave bears revealed the unique, independent character of evolution of the cheek teeth in the Kudaro cave bear, which is expressed to different degrees for the upper and lower rows of teeth.

Abstract Remains of a cave bear were studied from a new locality in the Prokoshev Cave in the Middle Urals (58°13´ N, 58°12´ E). Bones from all regions of the skeleton are present, bones are intact and without traces of human or animal activity. They all belong to the cave bear (Ursus kanivetz Vereshchagin, 1973). An AMS radiocarbon date of 53 375 ± 765 BP, IGANAMS–8632, was obtained from an adult mandible. The bones belonged to at least 18 individuals, including 4 individuals aged about one year, 1 aged about two years, 1 aged about three years, and 12 individuals over four years of age. Three skulls belonged to males and seven skulls belonged to females. The analysis has shown that the taphonomic type of this locality is a “mass burial.” This is the first “mass burial” of the cave bear in the Urals, found in situ, untouched by humans.

The aim of this study is to understand the feeding habits of the cave bear, Ursus spelaeus by investigating the dental microwear patterns of 106 molars from six Late Pleistocene caves in Catalonia (Spain): Ermitons Cave, Arbreda Cave, Mollet Cave, Llenes Cave, Toll Cave, and Teixoneres Cave. Dental microwear patterns of U. spelaeus were compared with a reference collection of extant ursid species. The results show an omnivorous and carnivorous diet in all sites analyzed with both intra- and inter-site pattern variability. Unlike previous studies, here dental microwear identified more carnivorous habits for the herbivorous cave bear during the days/weeks before death. More varied and higher energy items would help to cope with the hibernation period. The variability between the samples could be due to the characteristic climatic shifts of the Late Pleistocene and to the corresponding differences in the availability of resources.

Grotte de la Carriere is one of the secondary caves with paleontological record within the Reseau Lachambre karstic complex, an important network located in the Tet Valley at Villefranche-de-Conflent (Occitaine, France). After the discovery of dozens of cave bear remains in 2012, four systematical excavations in different parts of the cave allowed to recover a noteworthy Pleistocene mammal assemblage. Here we describe the dentognathic specimens of ursids coming from the lower layers of the first performed sondage. Based on morphological and morphometric comparisons with other late Early to Late Pleistocene European fossil material, the studied specimens were referred to the Middle Pleistocene species Ursus deningeri. Due to the scantiness of well-preserved and chronologically well-constrained U. deningeri remains, this study represents an important contribution to acquire knowledge on this Middle Pleistocene bear.

The composition of large mammal fauna from the Imanay Cave in the Southern Urals (53°02' N, 56°26' E) is described in the present study. The paper aims to provide an preliminary description of the large mammal remains from the Imanay Cave in order to establish their taxonomic status, geological age and to detect the factors that led to accumulation of the bone remains in the cave. An analysis of species composition of vertebrate fauna, archaeological finds and radiocarbon dates has shown that accumulation of fossils in the Imanay Cave spanned the whole period of the Late Pleistocene and Holocene. Remains of the small cave bear (Ursus “savini”) and cave lion (Panthera ex gr. fossilis-spelaea) are prevalent in the Pleistocene groups: 9414 (93%) and 536 (5%) bone specimens, respectively. The cave is the only site in the world where mass bone assemblages of two large terrestrial Carnivora species - small cave bear and cave lion – were found simultaneously. The composition of the skeletal samples for both species was described and the age and sex distributions were determined. Though Mousterian stone tools were found in the cave, no trace of the hunt of the bears and lions by humans was detected. The period during which the accumulation of the fossils took place can be broadly determined as the first half of the Late Pleistocene (MIS5–3).

In this article a third upper incisor of a bear, found by the author on the beach of Maasvlakte 2 (North Sea coast, Port of Rotterdam, The Netherlands), is described, analyzed and identified. On morphological grounds other large carnivores, and subsequently Ursus arctos and Ursus maritimus, are rejected as possible species determinations. A morphometric analysis strongly suggests Ursus deningeri, but intraspecific variation makes the identification of a loose find on that basis problematic. A detailed morphological analysis, however, shows that the incisor is characterized by a combination of arctoid and speleoid features, which is typical of Middle-Pleistocene cave bears, traditionally subsumed under the name Ursus deningeri. Thus the incisor can be assigned, with some caution, to that 'species', leading to the determination Ursus cf. U. deningeri. This article underlines the importance of studying not just (cave) bears' (pre)molars, but also their incisors.

[Abstract] Cova dos Santos is a karstic cavity in Abadín (Lugo), in a hitherto unexplored area that may have been the natural route between the well-known Quaternary faunas of the Cantabrian Mountain Range and those located further south in Galicia, such as in the Serra do Courel. The surface surveys carried out during the topographic layout revealed the presence of deposits of bone remains, usually extremely fragmented, of medium and large vertebrates. Due to the nature of these remains, different molecular techniques (ZooMS, stable isotopes), radiocarbon dating, and morphological and metric analysis were used to characterise the remains present at the site. Combining these methods, it has been possible to identify different taxa such as Ursus speleaeus, Ursus arctos, Panthera pardus, Cervus elaphus, Rhinocerotidae, and to confirm the occupation of this cave since at least 43000 years ago calBP. The presence of domestic species, such as Ovis aries, Equus sp. and Gallus gallus, also shows the use of this cave in more recent times. [Resumen] Cova dos Santos es una cavidad kárstica en Abadín (Lugo), en una zona hasta ahora inexplorada que podría haber sido la ruta natural entre las conocidas faunas cuaternarias de la cordillera Cantábrica y las situadas más al sur de Galicia, como en la Serra do Courel. La prospección de superficie realizada durante los trabajos de trazado topográfico reveló la presencia de depósitos de restos óseos, generalmente muy fragmentados, pertenecientes a vertebrados de mediano y gran tamaño. Debido al carácter de estos restos, se emplearon distintas técnicas moleculares (ZooMS, isótopos estables), dataciones por radiocarbono, y análisis morfológico y métrico para poder caracterizar los restos presentes en el yacimiento. Combinando estos métodos, se han podido identificar distintos taxones como Ursus speleaeus, Ursus arctos, Panthera pardus, Cervus elaphus, Rhinocerotidae, y constatar la ocupación de esta cueva desde hace al menos 43.000 años calBP. La presencia de especies domésticas, como Ovis aries, Equus sp. o Gallus gallus, también pone de manifiesto el uso de esta cavidad en épocas más recientes. This research was supported by financial assistance from the Sociedad Española de Paleontología ID AJISEP-2019-07121 and funds from the Instituto Universitario de Xeoloxía of the UDC. A. García-Vázquez is supported by a postdoctoral ICUB Fellowship for Young Researchers of the University of Bucharest Sociedad Española de Paleontología; AJISEP-2019-07121

The cave bear was a prominent member of the Upper Pleistocene fauna in Eurasia. While breakthroughs were recently achieved with respect to its phylogeny using ancient DNA techniques, it is still challenging to date cave bear fossils beyond the radiocarbon age range. Without an accurate and precise chronological framework, however, key questions regarding the palaeoecology cannot be addressed, such as the extent to which large climate swings during the last glacial affected the habitat and possibly even conditioned the final extinction of this mammal. Key to constraining the age of cave bear fossils older than the lower limit of radiocarbon dating is to date interlayered speleothems using 230Th/U. Here we report new results from one such site in the Eastern European Alps (Schwabenreith Cave), which yielded the highest density of bones of cave bear (Ursus spelaeus eremus). Although dating of the flowstones overlying this fossiliferous succession was partly compromised by diagenetic alteration, the 230Th/U dates indicate that the bear hibernated in this cave after about 113 ka and before about 109 ka. This time interval coincides with the equivalent of Greenland Stadial 25, suggesting possible climate control on the cave bear's habitat and behaviour.

Morphometric and morphotypic variability of the cave bear upper incisors from two different geographic regions (Caucasus and Urals), different stratigraphic periods (middle and late Pleistocene), and bearing different mitochondrial haplogroups (kudarensis and ingressus) was studied. The specific diet of the cave bears, i.e. hard vegetables, led to noticeable differences between their incisors and the incisors of the brown bear (Ursus arctos). It was found that the upper incisors of the Caucasian cave bears from different stratigraphic periods demonstrate consistent development of their morphology. The late Pleistocene cave bears from the Urals show a greater similarity to the Caucasian cave bears from earlier periods than with the cave bears from later periods. Our results suggest that the incisor morphology has evolved independently in the Caucasian and Ural cave bears as they belong to different phylogenetic lineages and display different ways of adaptation to local environmental conditions.

In this issue, we cover an exceptional topic in Vertebrate Paleobiology that has been an enjoyable challenge for scientists and the popular media alike: the life and death of the Pleistocene cave b...

The diet of the cave bear (Ursus spelaeus) is a controversial topic, as different paleobiological approaches (e.g. dental wear, isotopic biochemistry, skull morphometrics) result in differe...

This article analyzes the making of a novel consciousness of historicity in Germany around 1800, one that regarded mountains as vaults of a shared and palpable past. Revisiting a paleontological debate about the origin of large mammal bones found in caves, it reads the science of Johann Christian Rosenmuller (1771-1820) as a social and political accomplishment. By attributing the fossils to an indigenous "cave bear," and communicating an elite scientific debate to a lay audience, Rosenmuller presented an account of Germany's primordial past that fed seamlessly into its present, nurturing an idea of nationhood grounded in the (sub)soil.

Palaeogenomes provide the potential to study evolutionary processes in real time, but this potential is limited by our ability to recover genetic data over extended timescales.1 As a consequence, most studies so far have focused on samples of Late Pleistocene or Holocene age, which covers only a small part of the history of many clades and species. Here, we report the recovery of a low coverage palaeogenome from the petrous bone of a ∼360,000 year old cave bear from Kudaro 1 cave in the Caucasus Mountains. Analysis of this genome alongside those of several Late Pleistocene cave bears reveals widespread mito-nuclear discordance in this group. Using the time interval between Middle and Late Pleistocene cave bear genomes, we directly estimate ursid nuclear and mitochondrial substitution rates to calibrate their respective phylogenies. This reveals post-divergence mitochondrial transfer as the dominant factor explaining their mito-nuclear discordance. Interestingly, these transfer events were not accompanied by large-scale nuclear introgression. However, we do detect additional instances of nuclear admixture among other cave bear lineages, and between cave bears and brown bears, which are not associated with mitochondrial exchange. Genomic data obtained from the Middle Pleistocene cave bear petrous bone has thus facilitated a revised evolutionary history of this extinct megafaunal group. Moreover, it suggests that petrous bones may provide a means of extending both the magnitude and time depth of palaeogenome retrieval over substantial portions of the evolutionary histories of many mammalian clades.

During the Last Glacial Maximum (LGM, 30 to 16.5 ka ago), the Great Adriatic-Po Region (GAPR) was deeply affected by the spread of glaciers from the Alps to the southern foreland and by the dropping of the sea level to ~ -120 m amsl. The combination of these two events triggered the aggradation of the Great Po Plain (GPP), a vast flat area between the Alpine chain, the Italian Peninsula and the north-western Balkan Peninsula, physically and ecologically featured through a range of palaeogeographic and palaeoecological conditions. The low-elevated Prealpine sectors and the Alpine foothills supported more extensive forest stands, due to increased orographic rainfall. These were open boreal forests which persisted throughout the LGM, while open woodlands, steppes, semideserts and wetlands occupied the lowlands. A complex ecogradient, including both an Alpine and a continental timberline, is documented by the fossil records at the NE Alpine border, with a larch-pine forest-steppe belt, in contact with steppes and loess areas extending in the plain, on the dry extreme of the gradient. Still, edaphic wetlands occupied the waterlogged silty soils in the lowlands. Other areas, marked by active geodynamic processes, supported semideserts, i.e. grooves of xerophytic herbs and shrubs. Enhanced aridity and the development of deflation areas, prompted the accretion of loess cover at the northern and southern margins of the GPP. Fauna recorded the gradual disappearance of mammoth, woolly rhino and giant deer, together with cave bear. Gravettian and Epigravettian hunter-gatherer groups inhabited the GPP, although their presence and settlement dynamics at the margins and across this region has long been questioned. As a matter of fact, a handful of archaeological sites composes a patchy record of the peopling of the plain itself. At the northern rim of the GAPR, characterized by a well-developed karst region, several caves and rock shelters record the presence of hunters of bisons and horses at the margins of the GPP and ibexes and cave bears in some hilly landscapes. Nonetheless, evidence of contacts across this area is provided by the exploitation of chert sources and by stylistic and technical similarities in the lithic industries. The work resumes the currently available multidisciplinary data and adds new petroarchaeological evidence for reconstructing the settlement dynamics of the Gravettian - Epigravettian hunter-gatherers in this vast region up to the early Late Glacial, when the Prealpine and the Apennine foothills, along with the Dinarids, were persistently settled.

The Basura Cave (Toirano, Savona, NW Italy) hosts important cave bear bone assemblages and a numerous and varied, tracks and traces record left by humans and other producers. An outstanding element of the analysed material is represented by fossil bear fur fragments, which were found in the inner deposits of the cave, and that, to date, are virtually unknown in the cave global record. After analysing and discussing micromorphological features of the inedited material, we integrate and interpret new radiocarbon data, along with taphonomic, sedimentological, geochemical and mineralogical evidences, with the aim of improving our understanding about the nature and chronology of the bear fur-bearing deposit. The bear fur fragments are included in a stratigraphic succession corresponding to a secondary deposit, formed after the dismantling, reworking and redeposition of a former bear-bearing deposit, as a result of short but intensive flooding events that most probably took place at the end of the Last Glacial Maximum. After sediments redeposition, important diagenetic changes have occurred and probably driven by guano deposits, whose pre-existence, in absence of record, is inferred from corrosion features, nutrient concentrations, mineral species identified (REE bearing hydroxyapatite), and claw traces left by bats on the cave ceiling and walls. Diagenetic imprint derived by guano deposits caused mineralization of bear fur fragments by replacement with apatite, which faithfully copied the form and structure of hairs but also of vegetal tissues, phytoliths and pollen found within them. Our study demonstrates for the first time that the bear fur is one of the main vectors in introducing botanical microremains into the interior of the “Old World” caves.

The cave bear ( Ursus spelaeus s.l .) was an iconic extinct bear that inhabited the Pleistocene of Eurasia. The cause of extinction of this species is unclear and to identify the actual factors, it is crucial to understand its feeding preferences. Here, we quantified the shape descriptor metrics in three-dimensional (3D) models of the upper teeth (P 4 –M 2 ) of the cave bear to make inferences about its controversial feeding behaviour. We used comparative samples, including representatives of all living bear species with known diets, as a template. Our topographic analyses show that the complexity of upper tooth rows in living bears is more clearly associated with the mechanical properties of the items consumed than with the type of food. Cave bears exhibit intermediate values on topographic metrics compared with the bamboo-feeder giant panda ( Ailuropoda melanoleuca ) and specialists in hard mast consumption ( Ursus arctos and Ursus thibetanus ). The crown topography of cave bear upper teeth suggests that they could chew on tough vegetal resources of low quality with high efficiency, a characteristic that no living bear currently displays. Our results align with a climate-driven hypothesis to explain the extinction of cave bear populations during the Late Pleistocene.

a loose/broken stalagmite containing small fragments of cemented bones were collected from the Postojna Cave to investigate whether deoxyribonucleic acid (DNA) can be determined. The study is complemented by the Fourier-transform infrared spectroscopy - attenuated total reflectance (FTIR-ATR) analysis in order to determine the alteration of the bones and to test whether this analysis can be used as an indicator of possible DNA preservation. In addition, geochemical analyses were conducted in order to determine whether the associated flowstone/stalagmite is suitable for elucidating the timing of bone thanatocoenosis and further palaeoenvironmental analyses. The organic matter (collagen) is poorly preserved. However, we succeeded in amplifying a 94 bp long fragment of the cytochrome b (Cyt b) gene of mitochondrial DNA (mtDNA) in polymerase chain reaction (PCR) for one sample, and in sequencing the amto that of the Cyt b of the cave bear (Ursus deningeri or Ursus spelaeus sensu lato). The uranium-thorium dating of the speleothem covering the bones revealed its thanatocoenosis occurred prior to 55 ka, most likely in the late marine isotope stage 4 or early marine isotope stage 3. High porosity and recrystallisation of the flowstone/stalagmite at this part of the cave prevent high-resolution palaeoclimatic interpretation; however, low-resolution stable isotope geochemistry suggests a steppe-like environment during the subsequent growth of the speleothem.

The article presents the findings of the paleoparasitological analysis of coprolites from the cave bear (Ursus kanivetz Vereshchagin, 1973). The material for research was obtained during excavations in the Ignatievskaya Cave (Southern Urals, Russia; 54°53' N 57°46' E). The deposits with coprolites date back to the middle of the Late Pleistocene (90 000-30 000 years ago). On the basis of the paleoparasitological analysis, eggs of the nematode characteristic of representatives of Ursidae, namely, Baylisascaris transfuga Rudolphi, 1819, have been established to be present in the coprolites. The eggs are well preserved and have not lost their morphological features. The large cave bear was infested by the nematode B. transfuga. This is the first find of the nematode B. transfuga in Pleistocene-aged sediments and the first find of parasites in coprolites of the cave bear.

Caves often hold valuable palaeoclimate archives including speleothems, fossil remains, and clastic sediments that complement each other. This paper presents a multi-archive interdisciplinary study of an extensive deposit of fossil mammals from the scientific reserve in the Muierilor Cave, Southern Carpathians, Romania. We present two new palaeontological excavations that indicate a high abundance and diversity of MIS 3–2 fossil mammals (carnivores, omnivores and herbivores) synchronous with the early modern humans known from this cave. Using geochronological and sedimentological methods, we present a general reconstruction of the cave evolution between ~120 kyr B.P. and the Holocene. The study is based on a combination of geochronological tools including OSL dating of sediments, U/Th dating of speleothems, and radiocarbon dating of fossil remains, with a total of 54 ages. Based on U/Th dating of speleothems from stratigraphically-relevant positions, we show that the MIS 3 assemblage of fossil mammals were massively reworked and deposited during the post-LGM deglaciation, slightly earlier than previously known for the Southern Carpathians. On the other hand, several young radiocarbon ages of cave bear samples suggest that the Southern Carpathians might have been functioning as a glacial refuge for this species as late as ~22 kyr B.P.

The paper is a critical review of different evidence for the interpretation of an extremely important archaeological find, which is marked by some doubt. The unique find, a multiple perforated cave bear femur diaphysis, from the Divje babe I cave (Slovenia), divided the opinions of experts, between those who advocate the explanation that the find is a musical instrument made by a Neanderthal, and those who deny it. Ever since the discovery, a debate has been running on the basis of this division, which could only be closed by similar new finds with comparable context, and defined relative and absolute chronology.

Los resultados de ambos capítulos demuestran que los osos de las cavernas eran más herbívoros que el oso pardo actual (Ursus arctos) y se propone que probablemente se alimentaran de algún tipo de recurso altamente fibroso y de baja energía que estuviera presente en las regiones alpinas en las que habitaron. Ahora bien, la hipótesis ‘climática’ explica el declive de los osos de las cavernas no únicamente por ser herbívoros, sino también por su falta de capacidad de alimentarse de cualquier tipo de recurso en función de su disponibilidad, sobre todo en los momentos más infortunados de enfriamiento climático con una baja productividad primaria. De hecho, distintos estudios biogeoquímicos demuestran que los osos de las cavernas no cambiaron su dieta durante gran parte de su existencia, desde ≈100,000 a ≈20,000 años (Bocherens 2019; Terlatto et al. 2019). En consecuencia, los resultados obtenidos en el Capítulo 4.3 sugieren que los osos de las cavernas tenían una alimentación restringida al consumo de material vegetal. Además, se obtiene que esta falta de flexibilidad en la dieta sea una consecuencia del desarrollo exagerado de los senos paranasales. Puesto que en mamíferos actuales los senos están involucrados en la disminución de la tasa metabólica basal, se plantea la hipótesis de que este desarrollo de los senos en los osos de las cavernas pudiera deberse a sus largos períodos de hibernación necesarios para sobrellevar los inviernos más largos y más fríos de finales del Pleistoceno. Se concluye que dicho enfriamiento climático fue, probablemente, el agente selectivo para aumentar el tamaño de los senos a lo largo de la historia evolutiva del linaje. Por tanto, el estudio realizado en el capítulo 4.3 propone que dichos senos paranasales tan desarrollados en los osos de las cavernas y que conforman el típico domo frontal que caracteriza al linaje “espeloide” fueron necesarios para sobrellevar los largos períodos de hibernación por su función a la hora de disminuir la tasa metabólica basal. Sin embargo, dichos senos a su vez conllevaron una restricción biomecánica craneal que impidió a los osos de las cavernas alimentarse de cualquier tipo de recurso en función de su disponibilidad, siendo por tanto estos úrsidos, unos herbívoros obligados por la necesidad de poseer largos periodos de hibernación. En consecuencia, los cambios climáticos de finales del Pleistoceno podrían haber influido directamente en la capacidad de algunos miembros de la megafauna a la hora de obtener los nutrientes adecuados y sobrevivir con éxito durante las condiciones ecológicas extremas de los meses más fríos. Tales hallazgos plantean otra pregunta paleobiológica crucial sobre la evolución de la megafauna del Pleistoceno: ¿Cómo afectó el enfriamiento climático a otras especies de la megafauna? Para responder a esta pregunta, en el capítulo 4.5, se investiga la dieta de una especie de oso icónico del continente norteamericano: el oso de cara corta (A. simus). Este capítulo se centra en investigar si una serie de patologías de la dentición presentes en los dientes conservados en el yacimiento clásico de Rancho La Brea (California, EE. UU.) de finales del Pleistoceno obedecen a un proceso de caries. Los resultados confirman preferencias de alimentación variables entre diferentes poblaciones de A. simus de América del Norte como consecuencia del enfriamiento climático y la competencia de especies. Fecha de lectura de Tesis Doctoral: 17-1-2020 En esta tesis doctoral se usan diferentes métodos de análisis tridimensional con un potencial único para descifrar aspectos ecomorfológicos y evolutivos en diferentes grupos de vertebrados. Tales métodos están basados en la captación tridimensional tanto de las superficies externas del objeto como de las estructuras externas e internas mediante el uso de Tomografías Axiales Computarizadas (TACs). Específicamente, se investiga cómo tales técnicas permiten ir más allá del estado de la cuestión en la paleobiología de dos especies de úrsidos (Mammalia, Carnivora, Ursidae) emblemáticos de finales del Pleistoceno: el oso de las cavernas del continente europeo (Ursus spelaeus sensu lato) y el oso de cara corta del continente norteamericano (Arctodus simus). Tal y como se detalla en el siguiente apartado, la paleobiología de tales úrsidos es ciertamente controvertida en la literatura, siendo necesario, por tanto, la aplicación de nuevas técnicas analíticas que ofrezcan nuevas perspectivas para clarificar su paleobiología. Tales estudios han sido realizados mediante la aplicación de metodologías de última generación que nunca habían sido aplicados a la problemática en cuestión. Con respecto al oso de las cavernas, tales metodologías se aplicaron para responder una pregunta fundamental de su paleobiología que continua siendo objeto de discusión: ¿Por qué se extinguió el oso de las cavernas? Tal y como se ha comentado en la introducción, la respuesta a esta pregunta exige técnicas analíticas innovadoras que ofrezcan nuevas perspectivas para descifrar el comportamiento de alimentación de los osos de las cavernas y por ende de su extinción. Así, en los capítulos 4.1 y 4.2 se utilizan nuevas técnicas ecomorfológicas en tres dimensiones para investigar el comportamiento de alimentación del oso de las cavernas.

In this article a third upper incisor of a bear, found by the author on the beach of Maasvlakte 2 (North Sea coast, Port of Rotterdam, The Netherlands), is described, analyzed and identified. On morphological grounds other large carnivores, and subsequently Ursus arctos and Ursus maritimus, are rejected as possible species determinations. A morphometric analysis strongly suggests Ursus deningeri, but intraspecific variation makes the identification of a loose find on that basis problematic. A detailed morphological analysis, however, shows that the incisor is characterized by a combination of arctoid and speleoid features, which is typical of Middle-Pleistocene cave bears, traditionally subsumed under the name Ursus deningeri. Thus the incisor can be assigned, with some caution, to that 'species', leading to the determination Ursus cf. U. deningeri. This article underlines the importance of studying not just (cave) bears' (pre)molars, but also their incisors.

The Middle to Upper Palaeolithic transition in Europe witnessed the replacement and partial absorption of local Neanderthal populations by Homo sapiens populations of African origin1. However, this process probably varied across regions and its details remain largely unknown. In particular, the duration of chronological overlap between the two groups is much debated, as are the implications of this overlap for the nature of the biological and cultural interactions between Neanderthals and H. sapiens. Here we report the discovery and direct dating of human remains found in association with Initial Upper Palaeolithic artefacts2, from excavations at Bacho Kiro Cave (Bulgaria). Morphological analysis of a tooth and mitochondrial DNA from several hominin bone fragments, identified through proteomic screening, assign these finds to H. sapiens and link the expansion of Initial Upper Palaeolithic technologies with the spread of H. sapiens into the mid-latitudes of Eurasia before 45 thousand years ago3. The excavations yielded a wealth of bone artefacts, including pendants manufactured from cave bear teeth that are reminiscent of those later produced by the last Neanderthals of western Europe4–6. These finds are consistent with models based on the arrival of multiple waves of H. sapiens into Europe coming into contact with declining Neanderthal populations7,8. Direct dates for human remains found in association with Initial Upper Palaeolithic artefacts at Bacho Kiro Cave (Bulgaria) demonstrate the presence of Homo sapiens in the mid-latitudes of Europe before 45 thousand years ago.

In this article a third upper incisor of a bear, found by the author on the beach of Maasvlakte 2 (North Sea coast, Port of Rotterdam, The Netherlands), is described, analyzed and identified. On morphological grounds other large carnivores, and subsequently Ursus arctos and Ursus maritimus, are rejected as possible species determinations. A morphometric analysis strongly suggests Ursus deningeri, but intraspecific variation makes the identification of a loose find on that basis problematic. A detailed morphological analysis, however, shows that the incisor is characterized by a combination of arctoid and speleoid features, which is typical of Middle-Pleistocene cave bears, traditionally subsumed under the name Ursus deningeri. Thus the incisor can be assigned, with some caution, to that 'species', leading to the determination Ursus cf. U. deningeri. This article underlines the importance of studying not just (cave) bears' (pre)molars, but also their incisors.

This study is focused on cave bears from the Grotta del Bandito (Cuneo Province, Piedmont Region, Northern Italy), especially their teeth. The morphodynamic analyses have been elaborated and compared with other cave bears from Northern Italy, and other cave bear taxa from particular European sites. The Grotta del Bandito bears are similar in size to other cave bears from Italy, and no differences were observed with the other cave bear species or subspecies (e.g. U. ingressus, U. s. ladinicus, U. s. eremus). Thus their inclusion in the U. gr. spelaeus is the most appropriate classification, and they show a simpler dental surface than the “modern” cave bears (i.e. ingressus). The question of whether a single species of spelaeus lived in North Italy, is also introduced. In fact, the differences in size are probably the result of population variability, and no clear evidence of other spelaeoid species and/or subspecies, i.e. ingressus, s. ladinicus and s. eremus, was discovered.

Palaeogenomes provide the potential to study evolutionary processes in real time⁠, but this potential is limited by our ability to recover genetic data over extended timescales. The Late Pleistocene currently represents the age-limit for temperate-zone palaeogenome retrieval. Consequently, large portions of the evolutionary histories of many organismal clades remain inaccessible for direct genomic investigation. Here, we present the palaeogenome of a 367,000 year old cave bear from Kudaro 1 cave in the Caucasus. Analysis of this genome alongside those of Late Pleistocene representatives almost completely overturns the established view of cave bear evolution based on mitochondrial DNA. The Middle Pleistocene genome further allows us to directly estimate the ursine substitution rate to calibrate our revised nuclear genome phylogeny of Palaearctic bears. This reveals the Middle Pleistocene Transition as a key period in Palaearctic bear evolution, associated with the divergence of the major cave bear lineages, the divergence of brown and polar bears, and the end of gene flow between cave bears and brown bears. With an age encompassing an estimated 23% of the entire evolutionary history of cave bears, the Kudaro 1 genome extends the limit for temperate-zone palaeogenome retrieval well into the Middle Pleistocene.

Biomechanical simulations show a “trade-off” between hibernation length and a restricted herbivorous diet for European cave bears., The cave bear is one of the best known extinct large mammals that inhabited Europe during the “Ice Age,” becoming extinct ≈24,000 years ago along with other members of the Pleistocene megafauna. Long-standing hypotheses speculate that many cave bears died during their long hibernation periods, which were necessary to overcome the severe and prolonged winters of the Last Glacial. Here, we investigate how long hibernation periods in cave bears would have directly affected their feeding biomechanics using CT-based biomechanical simulations of skulls of cave and extant bears. Our results demonstrate that although large paranasal sinuses were necessary for, and consistent with, long hibernation periods, trade-offs in sinus-associated cranial biomechanical traits restricted cave bears to feed exclusively on low energetic vegetal resources during the predormancy period. This biomechanical trade-off constitutes a new key factor to mechanistically explain the demise of this dominant Pleistocene megafaunal species as a direct consequence of climate cooling.

In this article a third upper incisor of a bear, found by the author on the beach of Maasvlakte 2 (North Sea coast, Port of Rotterdam, The Netherlands), is described, analyzed and identified. On morphological grounds other large carnivores, and subsequently Ursus arctos and Ursus maritimus, are rejected as possible species determinations. A morphometric analysis strongly suggests Ursus deningeri, but intraspecific variation makes the identification of a loose find on that basis problematic. A detailed morphological analysis, however, shows that the incisor is characterized by a combination of arctoid and speleoid features, which is typical of Middle-Pleistocene cave bears, traditionally subsumed under the name Ursus deningeri. Thus the incisor can be assigned, with some caution, to that 'species', leading to the determination Ursus cf. U. deningeri. This article underlines the importance of studying not just (cave) bears' (pre)molars, but also their incisors.

The morphology of both crowns and tooth-roots reflects dietary specialisation in mammalian carnivores. In this article, we analyse the tooth-root morphology of maxillary teeth from CT scans of living bears (Ursus arctos, Ursus americanus, Ursus maritimus, Ursus thibetanus, Melursus ursinus, Helarctos malayanus, Tremarctos ornatus and Ailuropoda melanoleuca) in order to make inferences about the diet and feeding behaviour of the extinct cave bear (Ursus spelaeus sensu lato). Specifically, we investigate two major mitochondrial clades of extinct cave bears recognized by previous authors: Ursus ingressus and Ursus spelaeus (U. spelaeus spelaeus, U. spelaeus ladinicus, U. spelaeus eremus). Our results indicate a close association between tooth-root surface area and feeding behaviour in all living bear species. Tooth-root surface area values of cave bears suggest that they relied more on vegetative matter than living brown bears (Ursus arctos) but subtle differences between these species/subspecies could also indicate different feeding strategies among the members of cave bear complex.

In 1995, an unusually perforated femur of a juvenile cave bear was found in the Divje babe I Palaeolithic cave site in Slovenia. The supposition that it could be a flute led to heated debates. According to its archaeological context and chronostratigraphic position, if made by humans, it could only be attributed to Neanderthals. The crucial question was related to the origin of the holes. These could only have been made either by a carnivore or by human intervention. Results of experimental testing of both hypotheses do not support a carnivore origin of the holes. Furthermore, the method of artificial creation of the holes, which left no conventional traces of manufacture, was defined. Computed tomography revealed traces, which could be the result of human agency and called into serious question the origin of some features previously declared to be solely of carnivore origin. Recent musical experiments performed on a replica of the reconstructed musical instrument revealed its great musical capability. Together with some other findings from Divje babe I, the Mousterian musical instrument offers a unique insight into the Neanderthals’ symbolic behaviour and their cognitive abilities. The multidisciplinary results of comprehensive analyses of this exceptional find are first presented here together with its chronostratigraphic, palaeo-environmental, and archaeological contexts.

The cave bear (Ursus spelaeus) is one of the Late Pleistocene megafauna species that faced extinction at the end of the last ice age. Although it is represented by one of the largest fossil records in Europe and has been subject to several interdisciplinary studies including palaeogenetic research, its fate remains highly controversial. Here, we used a combination of hybridisation capture and next generation sequencing to reconstruct 59 new complete cave bear mitochondrial genomes (mtDNA) from 14 sites in Western, Central and Eastern Europe. In a Bayesian phylogenetic analysis, we compared them to 64 published cave bear mtDNA sequences to reconstruct the population dynamics and phylogeography during the Late Pleistocene. We found five major mitochondrial DNA lineages resulting in a noticeably more complex biogeography of the European lineages during the last 50,000 years than previously assumed. Furthermore, our calculated effective female population sizes suggest a drastic cave bear population decline starting around 40,000 years ago at the onset of the Aurignacian, coinciding with the spread of anatomically modern humans in Europe. Thus, our study supports a potential significant human role in the general extinction and local extirpation of the European cave bear and illuminates the fate of this megafauna species. © 2019, The Author(s). Introduction Results - Sample collection and processing - Phylogenetic analysis - Population size analysis Discussion - Phylogeography and population dynamics - Extinction Material and Methods - Sample collection - DNA extraction - Sequence processing - Data from other studies - Alignment and model selection - Inferring phylogenetic relationships - Comparison of D-Loop and Mitogenome Tree Topology Differences - Bayesian phylogenetic inference and demographic reconstruction - Accession numbers

Although many large mammal species went extinct at the end of the Pleistocene epoch, their DNA may persist due to past episodes of interspecies admixture. However, direct empirical evidence of the persistence of ancient alleles remains scarce. Here, we present multifold coverage genomic data from four Late Pleistocene cave bears (Ursus spelaeus complex) and show that cave bears hybridized with brown bears (Ursus arctos) during the Pleistocene. We develop an approach to assess both the directionality and relative timing of gene flow. We find that segments of cave bear DNA still persist in the genomes of living brown bears, with cave bears contributing 0.9 to 2.4% of the genomes of all brown bears investigated. Our results show that even though extinction is typically considered as absolute, following admixture, fragments of the gene pool of extinct species can survive for tens of thousands of years in the genomes of extant recipient species.

When studying an extinct species such as the cave bear (Ursus spelaeus ROSENMÜLLER 1794), it is possible to apply a variety of molecular biology techniques such as the study of stable isotopes or mitochondrial DNA (mDNA) to infer patterns of behaviour or physiology that would otherwise remain concealed. Throughout Europe and along time, differences in the isotopic values (δ13C and δ15N) of cave bears arise from environmental differences and the Pleistocene climatic evolution. The climate determines the hibernation length, during which the cave bears undergo a particular physiology that can be related to an increase in δ15N during climate cooling. In order to verify whether hibernation affected the isotopic values, we compared cave bears in different ontogenetic stages. The results show that perinatal values reflect the values for mothers during hibernation, while juveniles show differences in maternal investment. A previous study in the literature based on complete mitochondrial DNA sequences of several individuals collected from closely situated caves showed that each cave housed, almost exclusively, a single lineage of haplotypes. This pattern suggests extreme fidelity to the birth site, or homing behaviour, and that cave bears formed stable maternal social groups, at least for the purpose of hibernation. Studies of this type offer unexpected data on the palaeobiology of this extinct animal.

More than 300 cave bear bones from all over Europe have carbon and nitrogen isotopic composition that match overwhelmingly a diet based on plants, except for samples from two caves in Romania, for ...

The palaeoichnology of the cave bear, Ursus spelaeus , was studied at Ursilor Cave, Apuseni Mountains, Romania, to acquire new data about the ethology and the metrics of this extinct species. Eight stations containing footprints, scratch marks and nests were surveyed using laser scanning to build tridimensional models for several cave bear nests, wall scratch marks and footprints. Morphological observations were made using the models, and precise measurements of the breadth or depth of the scratch marks were performed. In addition, >100 cave bear nests and 10 footprints were measured and analyzed. The palaeoichnogical data were corroborated with previously obtained information on the taphonomy, geomorphology, sedimentology and geochronology from this site. Several bioglyphs (e.g., scratch marks, footprints) were indirectly dated to Marine Isotope Stage (MIS) 3 by identifying and assigning them to co-occurring cave bear skeletons. As a consequence, we stress the need for a multidisciplinary approach in order to assess the palaeoichnology of cave bear sites. Findings fill a gap in our knowledge because cave bear bioglyphs are one of the less studied branches of cave bear research due to the scarcity and fragility of ichnological features.