Bison Priscus Research Articles

Early Pleistocene forerunners of Mammuthus-Coelodonta Faunal Complex in Nihewan Basin, North China

The typical Mammuthus-Coelodonta Faunal Complex dominated the mid- and high latitudes of continental Eurasia during the Mid-Late Pleistocene; the dominant taxa include woolly mammoth (Mammuthus primigenius), woolly rhino (Coelodonta antiquitatis), steppe bison (Bison priscus) and diverse horses (Equus spp.), etc. Recent excavations at the Early Pleistocene site Shanshenmiaozui in Nihewan Basin of North China resulted in the discoveries of rich mammalian fossils which include steppe mammoth (Mammuthus trogontherii), Nihowan woolly rhino (Coelodonta nihowanensis), archaic Chinese bison (Bison (Eobison) palaeosinensis) and rich collections of early horse fossil. In Nihewan Basin, a couple of sites (Xiashagou, Shanshenmiaozui and Yeniupo) yielded the Mammuthus-Coelodonta-Bison assemblage, which can be regarded as the closest ancestors of the Mammuthus - Coelodonta Faunal Complex or the forerunners of the later Mammuthus - Coelodonta Faunal Complex. Nihewan Basin bears the richest and most complete fossils of Early Pleistocene Coelodonta and Bison to date, and the juvenile specimens of M. trogontherii also represent the richest collection for its kind and its geologic age.

Was the steppe bison a grazing beast in Pleistocene landscapes?

The history and palaeoecology of the steppe bison (Bison priscus) remain incompletely understood despite its widespread distribution. Using dental microwear textural analysis (DMTA) and vegetation modelling, we reconstructed the diet and assessed the habitat of steppe bison inhabiting Eurasia and Alaska since the Middle Pleistocene. During the Late Pleistocene, steppe bison occupied a variety of biome types: from the mosaic of temperate summergreen forest and steppe/temperate grassland (Serbia) to the tundra biomes (Siberia and Alaska). Despite the differences in the identified biome types, the diet of steppe bison did not differ significantly among populations in Eurasia. DMTA classified it as a mixed forager in all populations studied. The DMTA of Bb1 bison-a recently identified genetically extinct sister-clade of Bison bonasus-was typical of a highly grazing bovid species and differed from all B. priscus populations. The results of the study temper the common perception that steppe bison were grazers in steppe habitats. The dietary plasticity of the steppe bison was lower when compared with modern European bison and may have played an important role in its extinction, even in the stable tundra biome of eastern Siberia, where it has survived the longest in all of Eurasia.

Morphological peculiarity of the fossil bison (<i>Bison priscus</i>) from Yakutia

The steppe bison (Bison priscus) is one of the most widespread and numerous representatives of the mammoth fauna. However, the features of the extinction of steppe bison at the end of the Pleistocene and their replacement by modern forms of the American bison (Bison bison) and European bison (Bison bonasus) have not yet been fully studied. Thus, complete skulls of fossil steppe bison are of great interest for studies on taxonomy and kinship relations in the genus Bison. This article presents the results of a study of the finely preserved skull of a fossil bison found in the Verkhoyansk district (Yakutia). This skull was compared by its morphological and osteometric features with other bison skulls, which were found at different times and locations in Upper Pleistocene deposits, as well as with the skulls of modern American bison Bison bison from the collection of the Zoological Museum of the Lomonosov Moscow State University. Standard morphometric indicators of bison skulls and horns were obtained using the Skinner and Kaisen measurement methods. Analysis showed that skulls of Bison priscus from Late Pleistocene sediments of Yakutia, according to most parameters, correspond to the maximum size of modern American bison Bison bison or even larger than the latter. The horns of bison from the Late Pleistocene are 1.5–2 times larger in size than the horns of modern bison and are comparable to those of the Middle Pleistocene long-horned bison B. p. crassicornis Richardson (=longicornis). The disproportion in the size of the skull and horns of the Late Pleistocene bison is due to their habitat in the open landscapes of the Arctic steppe. The morphological uniqueness of the steppe bison of the Late Pleistocene in Yakutia has prospects for further research, because it deserves to be identified as an independent taxon (subspecies), which is relevant for regional paleontology and stratigraphy.

New data on the mammoth fauna in the Amga River basin

Many unique finds of Pleistocene animals have been made on the territory of Yakutia, including whole frozen bodies. However, the majority of the finds come from the northern regions. On the territory of Central Yakutia, in contrast to Northern Yakutia, there are much fewer finds. This is due to the smaller number of preserved relict ice deposits of the Upper Pleistocene ("Yedoma") and the lack of systematic, licensed collection of mammoth ivory and other remains of the mammoth fauna due to their unsatisfactory preservation, which is caused by the stronger thawing of the permafrost in summer in Central Yakutia. There are relatively few published scientific papers on the mammoth fauna of the Amgar River basin. Therefore, new data from this area are of scientific interest. We have studied new material on the mammoth fauna from the "Mount Korolenko" locality, located on the left bank of the Amgar River near the village of the same name. The material was collected during school expeditions organised by the Amga Lyceum and the Russian Geographical Society in 2022-2023. It shows that at this site bone remains are washed out by the river on the right bank of the Amgas. Scattered bones were found on both the left and right banks, but the right bank is richer in palaeontological material. The bone remains are represented by fragments and whole bones of the postcranial skeleton of various animals. The age of most of the finds probably corresponds to the Karga interstadial. The taxonomic identification of the bones showed the presence of the following species: woolly mammoth, woolly rhinoceros, Lena horse, steppe bison, reindeer, red deer. There is a fragment of the humerus of a roe deer, possibly of late Neopleistocene age. A morphometric study of the bone remains was carried out and comparisons made with literature data. The list of large mammals that lived in the Amgas basin in the late Neoproterozoic has been completed.

Fossil large mammals from Wielkopolska: a state of knowledge

A number of 125 open-air localities from Wielkopolska documented presence of 18 species (3 carnivores, 4 proboscideans, 3 perissodactyls and 8 artiodactyls). Most of species are cold-adapted members of mammoth fauna from the Late Pleistocene, such as Ursus arctos priscus, Mammuthus primigenius, Coelodonta antiquitatis, Equus ferus, Rangifer tarandus, and Bison priscus. The few species like Ursus arctos taubachensis, Palaeoloxodon antiquus and Stephanorhinus kirchbergensis represent an older, thermophilic fauna dated to MIS 5e. The vast majority of bones are accidental finds, without a stratigraphic context. Most artiodactyls were found in alluvial sediments, in bogs or swamps, while carnivores are represented only by isolated remains.

Arctic Quaternary Mammal Collections in the Museums of Yakutsk (Yakutia, East Siberia, Russia)

Many Ice Age fossil mammals remains are found in Yakutia, which occupies most of Eastern Siberia. In the city of Yakutsk, various museum and scientific institutions hold collections and exhibits of Quaternary mammals collected at different times in different locations in the Arctic zone of Yakutia. These collections mostly represents the Late Pleistocene Mammoth Fauna, and are housed in the Emelyan Yarosslavsky’s Yakutsk State Museum of History and Culture of the Peoples of the North, the Mammoth Museum, the Geological Museum of Diamond and Precious Metals Geology Institute, Siberian Branch of Russian Academy of Sciences, and the Department of the Mammoth Fauna study, Academy of Sciences of the Sakha (Yakutia) Republic. These collections include woolly mammoth skeletons, other fragments of mammoths, woolly rhinoceros and horse carcasses, whole carcasses of steppe bison, cave lion cubs, and other fossil mammals, many bones and teeth of various mammals of the Mammoth Fauna. The study of these fossil mammal remains allow identification of features of their anatomy and morphology, as well as reconstruction of the ecological conditions at the time they were alive.

A long-term perspective on Neanderthal environment and subsistence: Insights from the dental microwear texture analysis of hunted ungulates at Combe-Grenal (Dordogne, France).

Large bovids and cervids constituted major components of the European Middle Palaeolithic faunas and hence a key resource for Neanderthal populations. In paleoenvironmental reconstructions, red deer (Cervus elaphus) occurrence is classically considered as a tree-cover indicator while Bovinae (Bison priscus and Bos primigenius) and reindeer (Rangifer tarandus) occurrences are typically associated with open landscapes. However, insights into the ecology of extant ungulate populations show a more complex reality. Exploring the diet of past ungulates allows to better comprehend the hunting strategies of Palaeolithic populations and to reconstruct the modifications through time of past landscapes. By reflecting what animals have eaten during the last days or weeks of their life, dental microwear textures of herbivores link a population and its environment. Here we analyzed, via Dental Microwear Texture Analysis (DMTA), the diet of 50 Bos/Bison, 202 R. tarandus and 116 C. elaphus preyed upon by the Neanderthals that occupied Combe-Grenal rock-shelter, one of the most important Mousterian archaeo-sequences in southwestern France considering its long stratigraphy, abundance of faunal remains and the variations perceptible in Palaeolithic material culture. Grazers and mixed-feeders are the most represented dietary categories among Combe-Grenal's guild of herbivores, highlighting the availability, along the sequence, of open landscapes. The absence of clear changes in the use of plant resources by hunted ungulates through time, even though palaeoenvironmental changes were well-documented by previous studies along the sequence, is interpreted as resulting from the hunting of non-randomly selected prey by Neanderthals, preferentially in open environments. Thus, these results provide further insight into the hunting strategies of Neanderthals and modify our perception of potential links between subsistence and material culture. Combe-Grenal hunters "stayed in the open" through millennia, and were not forced to switch to hunting tactics and material technology adapted to close encounters in forested environments.

Bovid Bone Accumulation in Late Middle Palaeolithic Poland

ABSTRACT The hunting activities of Neanderthals inhabiting the European Lowlands during the Weichsel glaciation are poorly understood due to the scarcity of faunal remains. This work concerns the puzzling accumulation of mammalian remains at the Middle Palaeolithic site Haller Av. in Wrocław, southwestern Poland. The site yielded lithic artifacts in two levels and numerous bone remains typical for steppe-tundra fauna, dominated by steppe bison (Bison priscus). As the site was transformed by fluvial processes, the question arose whether the accumulation of faunal bones was the result of human activity. To resolve this question, we used a multiproxy approach, including spatial analysis with GIS, as well as taphonomic and paleozoological analyses. It was found that the accumulation of bone remains was multi-stage and involved a large area. Only the bison remains were spatially correlated with the traces of human activity. In our opinion, these data indicate the presence of a hunting site of monospecific fauna, demonstrating the profound knowledge of Neanderthals about the ecosystem of which they were a part.

Range and extinction dynamics of the steppe bison in Siberia: A pattern‐oriented modelling approach

AbstractAimTo determine the ecological processes and drivers of range collapse, population decline and eventual extinction of the steppe bison in Eurasia.LocationSiberia.Time periodPleistocene and Holocene.Major taxa studiedSteppe bison (Bison priscus).MethodsWe configured 110,000 spatially explicit population models (SEPMs) of climate–human–steppe bison interactions in Siberia, which we ran at generational time steps from 50,000 years before present. We used pattern‐oriented modelling (POM) and fossil‐based inferences of distribution and demographic change of steppe bison to identify which SEPMs adequately simulated important interactions between ecological processes and biological threats. These “best models” were then used to disentangle the mechanisms that were integral in the population decline and later extinction of the steppe bison in its last stronghold in Eurasia.ResultsOur continuous reconstructions of the range and extinction dynamics of steppe bison were able to reconcile inferences of spatio‐temporal occurrence and the timing and location of extinction in Siberia based on hundreds of radiocarbon‐dated steppe bison fossils. We showed that simulating the ecological pathway to extinction for steppe bison in Siberia in the early Holocene required very specific ecological niche constraints, demographic processes and a constrained synergy of climate and human hunting dynamics during the Pleistocene–Holocene transition.Main conclusionsEcological processes and drivers that caused ancient population declines of species can be reconstructed at high spatio‐temporal resolutions using SEPMs and POM. Using this approach, we found that climatic change and hunting by humans are likely to have interacted with key ecological processes to cause the extinction of the steppe bison in its last refuge in Eurasia.

Paleogenomes Reveal a Complex Evolutionary History of Late Pleistocene Bison in Northeastern China.

Steppe bison are a typical representative of the Mid-Late Pleistocene steppes of the northern hemisphere. Despite the abundance of fossil remains, many questions related to their genetic diversity, population structure and dispersal route are still elusive. Here, we present both near-complete and partial mitochondrial genomes, as well as a partial nuclear genome from fossil bison samples excavated from Late Pleistocene strata in northeastern China. Maximum-likelihood and Bayesian trees both suggest the bison clade are divided into three maternal haplogroups (A, B and C), and Chinese individuals fall in two of them. Bayesian analysis shows that the split between haplogroup C and the ancestor of haplogroups A and B dates at 326 ky BP (95% HPD: 397-264 ky BP). In addition, our nuclear phylogenomic tree also supports a basal position for the individual carrying haplogroup C. Admixture analyses suggest that CADG467 (haplogroup C) has a similar genetic structure to steppe bison from Siberia (haplogroup B). Our new findings indicate that the genetic diversity of Pleistocene bison was probably even higher than previously thought and that northeastern Chinese populations of several mammalian species, including Pleistocene bison, were genetically distinct.

Funeral Rites of the Greater Mammals

Funeral Rites of the Greater Mammals Colin Pope (bio) Thirty yards from the water, it lies wreathed by joggers,beachcombers, a pair of hand-holding lovers,a pectoral fin pointing at the sky in accusation."Jesus," Katie says, "how does this happen?"Thunderheads flash the horizon.The surf mumbles its eulogy as the authoritiesfumble and attempt: it's a long exchange, tying a limp monster by the tail. They are amateurs in a small townand so unspindle a steel chain from its gashed flukes,unfurl a spiral of manila rope instead, like a teenagerin a den with the phone cord stretched. I feel constantly like I should rush forth, like we need to waitto see which ghosts wriggle free its blowhole. "No hurry,"one of the workers says. "Gonna be two days beforethe tugboat." This is what I always forget about loss:just when you think it can't get any deeper,that the anchor is scraping your abyss, that's whenyou whiff the seaweed and plankton-eating bacteria, the rank tunnels fish have carved inside youand realize your immensity and hownothing hurts at all, it never did, that conclusionis a new problem which we will, like it or not, be forced to haul away. + + + Two days somehow the biorhythmically allottedcross-species timeframe forfeited to losing one's mindwith grief: after her babywas whipped around like a flywhisk by a lovesick male, [End Page 286] Gracie the chimp carried its corpse for two daysbefore the wardens of the L.A. Zoo could pry it freeand sneak the child from the paddock like an armloadof wet laundry. "They're just quiet": how the zookeeper describedthe toddler's family in mourning. Same with the fallen matriarch of the elephant herdin a South African preserve, its calfwailing and weeping in black streamsas the family tried to raise mama's enormous bulk with their trunks.They rumbled, quieted, covered her with dirt and twigs and stood vigil forty-eight hours, no sound on the savannabut a child's sorrow. Western lowland gorillas, too:a mother is observed cradling her stillborn son, grooming him with slow, steady fingers, two days straight.And the dolphins raising a whitened podmateand the giraffe licking the fur from its friend and the zebrasnudging at a chalky, dry muzzle, always, always two days, inexplicably, as though one sun, one moonweren't enough for even the souls of beaststo decipher what it means to go on. + + + "No one knows why they do it," Katie says, staring down into her phone. "They just end up on the beach."Entire fleets of cetaceans sometimes, lined across the shorelike a photograph from Vietnam, somewhose jackfruit-sized brains went haywireat signals from subs and sonar. Some done in by bad math in the shallows. Some photos of toothed whalesyou can almost see yourself inside, not as a pilotbut actually being, crooning, listening with your jawboneswhen a lifelong partner calls from up above, having literally [End Page 287] landed itself in trouble. Swimming to its aid, failing,then calling for more help untilyou're all gone. Tell me, thinkers, philosophers,tell me what of these strandings could faithfully be categorizeda natural phenomenon? + + + It gets so you stop wondering, so it's no big whoopwhen you traipse through the woods for your seventy years on Earthand never stumble upon a dead lynx or Kodiak, a bull mooselying in state, unbloodied, pure. Blue Babe, the 36,000-year-old steppe bison mummified in vivianite:even the Alaskan gold miner who unearthed it with a pressure hosecould tell the poor darling got caught by a sudden blizzardand so, we can cross it off the list, its termination a so-calledaccident: what if nothing dies the way humans do? Of biologic exhaustion, of spending ourselveslike produce in an open truck? What is it about two daysthat matters? You are shy enough to fall in love with someone who charms, who on a rocky outcropping by the seagets down on...

Aurochs roamed along the SW coast of Andalusia (Spain) during Late Pleistocene

In the Iberian Peninsula the fossil record of artiodactyls spans over 53 million years. During the Pleistocene, wild cattle species such as Bison and especially Bos became common. In Late Pleistocene, the aurochs (Bos primigenius) was widespread and the only bovine living along the large river valleys of southern Iberia. Although commonly found in fossil sites and especially in cave bone assemblages, the trace fossil record of aurochs was known worldwide only from the Holocene. Large bovine and roe deer/caprine tracks were found in at least five horizons of the early Late Pleistocene (MIS 5) beach and eolian deposits of Cape Trafalgar (Cadiz Province, South of Spain). The large bovine tracks are formally described as Bovinichnus uripeda igen. et isp. nov. and compared with the record of aurochs tracks, large red deer tracks and steppe bison biogeographical distribution in Iberia. Aurochs were the most likely producers of the newly described Trafalgar Trampled Surface (TTS) and some of the large artiodactyl tracks in the Matalascañas Trampled Surface, representing the oldest aurochs track record known. This new evidence, together with comparisons with the record of possible aurochs tracks in the Mid-Late Pleistocene coastal deposits from the Asperillo cliff section in Matalascañas (Huelva Province, SW Spain) and bone assemblages known in Gibraltar, point to a recurrent use of the coastal habitat by these large artiodactyls in SW Iberia.

Mammoth Fauna Remains From Late Pleistocene Deposits of the Dmitry Laptev Strait South Coast (Northern Yakutia, Russia)

The Yedoma Ice Complex in northern Yakutia provides perfect preservation conditions for frozen remains of vertebrate animals. Even complete mummified specimens of the late Pleistocene Beringian Mammoth fauna such as woolly mammoth, woolly rhinoceros, horse, and bison are occasionally found in permafrost deposits across eastern Siberia, i.e., in West Beringia, although bones are much more commonly found. The present study characterizes mammal bones from late Pleistocene and Holocene permafrost deposits exposed on the Oyogos Yar coast, part of the southern shore of the Dmitry Laptev Strait that connects the Laptev and East Siberian seas. The study applies a method to characterize fossil bone samples by the location of their discovery and by the accuracy of their relation to a depositional horizon. We analyzed a total of 38 finite radiocarbon ages of bone material from mammoth, horse, and musk ox, spanning from about 48.8 to 4.5 ka BP and including both our own data and data from the literature, in addition to previous publications that reported numerous bones with infinite ages from the Oyogos Yar coast. The distribution of bones and tooth along the coastal permafrost exposure is not uniform; it depends upon whether the material was found in situ, on thermo-terraces, or on the shore. The overall bone collection consists of 13 species of which Mammuthus primigenius (woolly mammoth, 41%), Bison priscus (bison, 19%), Equus ex gr., caballus (horse, 19%), and Rangifer tarandus (reindeer, 16%) predominate. The fossil bone species distribution is similar to those of other prominent Yedoma outcrops in the region, i.e., on Bykovsky Peninsula and on Bol’shoy Lyakhovsky Island. Correlation analysis shows that the Oyogos Yar bone sampling sites of different geomorphological settings are similar to each other but not to all sampling sites within the other two locations on Bykovsky Peninsula and on Bol’shoy Lykahovsky Island. High similarities in terms of correlation coefficients between specific sampling sites are often not represented in the cluster analysis.

MIS 3–1 fauna from Krosinko: Implications for the past biogeography, chronology and palaeoenvironments of Poland

A faunal assemblage was recently recovered from the Krosinko site in Poland, marking a new site in the Warsaw–Berlin ice-marginal valley. Analysis and radiocarbon dating of the remains revealed the presence of two subassemblages: one Late Pleistocene (at least MIS 3–2) and one Holocene (MIS 1), with the former being taxonomically highly diverse. By referring to previous Polish studies, the Pleistocene mammal bone assemblage allowed us to reconstruct the biogeography of its individual components: woolly mammoths, woolly rhinoceri, horses, giant deer, red deer, fallow deer, reindeer, steppe bison, aurochses, bears, and cave lions. The biogeography was later combined with radiocarbon chronology using 21 dates from Krosinko. The wide range of radiocarbon dates (48.4–26.4 ka BP) probably reflects different chronological distributions of species resulting from the changing climate and environment as well as the diverse original sources of the bones. The remains, which constituted a single, compact geological horizon, was possibly deposited in a single geological event around 26 ka BP, as inferred from the most recently produced radiocarbon date. Radiocarbon datings of woolly rhinoceros remains have shown the species to have been present around 38 ka BP, a time that it had previously been considered to have been non-existent in Europe (with no evidence having been discovered for the period 40–34 ka BP). This result alters the stratigraphic gap range for the woolly rhinoceros in Europe to 40–38 ka BP. The clear dominance of woolly mammoth and woolly rhinoceros specimens reflects the structure of the Mammuthus–Coelodonta faunal complex. The main representatives of this complex preferred the dry and cold conditions that characterized the palaeoenvironmental conditions in Krosinko in MIS 2 and thus in Central Europe, as shown by the integration of the results of the fauna and those of the sediments in which they were deposited. These palaeoenvironmental features are related to the mammoth steppe, which was a long-lived biome in Eurasia and whose diversity influenced the faunal distribution. Thus, the species-by-species review of taxa performed in this paper, against the background of previous results from Poland, illuminates the individual responses of taxa to changing environmental conditions in response to climatic ones. These changes led to the disappearance of the mammoth steppe and near or total extinction of the fauna species inhabiting it.

Upper Pleistocene remains of Bison priscus Bojanus, 1827 from Biśnik Cave (Middle Palaeolithic) and their significance for stratigraphy and palaeoecology

Remains of the steppe bison Bison priscus Bojanus, 1827 from the Upper Pleistocene layers of Biśnik Cave were morphometrically analysed and compared to those from other localities in Eurasia. The remains occurred in accumulations with archaeological artefacts. They bore traces of gnawing by carnivores. Morphometric analyses of the skull and post-cranial skeleton point to the small form of bison. The skull belonged to a small male while the bones of post-cranial skeleton indicate the occurrence of both males and females. Here we present all available and newly obtained calibrated 14C dates of the bison remains from this site, supplemented with new data for five specimens from Biśnik Cave, thus confirming the presence of the species during MIS 3 in Poland. The results are relevant to tracing the migration paths of the bison in the Upper Pleistocene during MIS 4-3.

Pleistocene mitogenomes reconstructed from the environmental DNA of permafrost sediments

Traditionally, paleontologists have relied on the morphological features of bones and teeth to reconstruct the evolutionary relationships of extinct animals.1 In recent decades, the analysis of ancient DNA recovered from macrofossils has provided a powerful means to evaluate these hypotheses and develop novel phylogenetic models.2 Although a great deal of life history data can be extracted from bones, their scarcity and associated biases limit their information potential. The paleontological record of Beringia3-the unglaciated areas and former land bridge between northeast Eurasia and northwest North America-is relatively robust thanks to its perennially frozen ground favoring fossil preservation.4,5 However, even here, the macrofossil record is significantly lacking in small-bodied fauna (e.g., rodents and birds), whereas questions related to migration and extirpation, even among well-studied taxa, remain crudely resolved. The growing sophistication of ancient environmental DNA (eDNA) methods have allowed for the identification of species within terrestrial/aquatic ecosystems,6-12 in paleodietary reconstructions,13-19 and facilitated genomic reconstructions from cave contexts.8,20-22 Murchie etal.6,23 used a capture enrichment approach to sequence a diverse range of faunal and floral DNA from permafrost silts deposited during the Pleistocene-Holocene transition.24 Here, we expand on their work with the mitogenomic assembly and phylogenetic placement of Equus caballus (caballine horse), Bison priscus (steppe bison), Mammuthus primigenius (woolly mammoth), and Lagopus lagopus (willow ptarmigan) eDNA from multiple permafrost cores spanning the last 40,000 years. We identify a diverse metagenomic spectra of Pleistocene fauna and identify the eDNA co-occurrence of distinct Eurasian and American mitogenomic lineages.

Remains of the Baikal yak (Poehpagus mutus baikalensis N. Verestchagin, 1954) from Late Pleistocene localities of Southern Siberia

136 bones of the postcranial skeleton and a part of the skull of a female Baikal yak (Poephagus mutus baikalensis N. Verestcthagin, 1954), originating from 18 cave locations and open-type Paleolithic sites in Altai-Sayan, Transbaikalia and Central Mongolia were examined. The material includes 38 metacarpals and 9 metatarsals of the yak. Morphometric differences in the structure of the postcranial skeleton of the yak and the bison (Bison priscus Bojanus, 1827) were revealed. The body size of the Baikal yak significantly exceeded that of the wild Tibetan yak (Poephagus mutus Przewalski, 1883). The largest representatives of P. m. baikalensis inhabited the Altai Mountains. In most of the sites, located in the mid-mountain landscapes of Southern Siberia (with absolute heights of 500–700 m), only a few remains of the Baikal yak were found, accounting for 0.01% to 1–2% of the number of megafauna remains. Most likely, herds of yaks did not live here permanently, but appeared only sporadically, during seasonal migrations. In higher mountainous areas (from 1000–1500 m) of Gorny Altai and Khangai Mountains in Central Mongolia, the proportion of the remains of the Baikal yak increases significantly – up to 16–22%. Like the contemporary P. mutus, the Pleistocene yak found its ecological optimum in the high-mountainous parts of ridges and mountain plateaus, dominated by cold, dry mountain-steppe landscapes with herb-grass vegetation and a small amount of snow. During the periods of cryochrones, the area of P. m. baikalensis apparently expanded significantly, incorporating the adjacent foothill territories. During the periods of thermochrones, it was most likely limited to the high-mountainous areas of the mountain uplifts of Southern Siberia.

Paleontological monument of nature “Mamontova Gora” exposure (Yakutia, Eastern Siberia, Russia)

The Mamontova Gora (“Mammoth Mountain”) exposure is the Neogene – Pleistocene key section of Siberia. This outcrop is located in the lower reaches of the Aldan River, 325 km above its mouth and extends for almost 12 km. It consists of an 80-meter structural plateau (80-meter terrace), 50- and 30-meter alluvial terraces. Sediments from the Middle Miocene (16-10 Ma) to the Upper Pleistocene are exposed on the 80-meter terrace. The basement of the 50-meter terrace is composed of Middle Miocene sediments, overlain by Pleistocene sediments. On a younger 30 m terrace, the deposits are dated from the Upper Pliocene to the Upper Pleistocene. The Mamontova Gora outcrop is one of the richest localities of the Neogene flora of Eurasia. There are numerous finds of remains of Miocene evergreen and thermophilic plants (tree stumps, leaf imprints, cones, nuts, seeds). More than 250 genera of fossil plants have been found on Mamontova Gora. This outcrop is also well known to paleontologists due to the abundance of bone remains of mammals of the Middle Pleistocene (early type mammoth, eastern horse, broad-fronted moose, long-horned bison) and Late Pleistocene (representatives of the mammoth fauna: woolly mammoth, woolly rhinoceros, Lena horse, reindeer, saiga-antelope, steppe bison, Arctic fox, wolverine, cave lion, etc.). It was revealed that the ancient frozen sediments on the Mamontova Gora outcrop abound with viable microorganisms and traces of their vital activity. A strain of microbe Bacillus sp. was isolated from ~ 2 - 3 Ma permafrost layers of this outcrop. A large group of microorganisms including fungi was isolated from the ancient ice wedge. Pleistocene permafrost deposits contain invertase, urease, catalase and dehydrogenase enzimes. Mamontova Gora is a unique geological object in Russia. By the decree of the Council of Ministers of the Yakutia Autonomous Soviet Socialist Republic of 18.02.1987 No. 56 Mamontova Gora was given the status of a “natural monument” and a specially protected natural area of regional significance. The article presents the main results of studies of ancient flora and faunas of Mamontova Gora.

Multiproxy analysis of permafrost preserved faeces provides an unprecedented insight into the diets and habitats of extinct and extant megafauna

The study of faecal samples to reconstruct the diets and habitats of extinct megafauna has traditionally relied on pollen and macrofossil analysis. DNA metabarcoding has emerged as a valuable tool to complement and refine these proxies. While published studies have compared the results of these three proxies for sediments, this comparison is currently lacking for permafrost preserved mammal faeces. Moreover, most metabarcoding studies have focused on a single plant-specific DNA marker region. In this study, we target both the commonly used chloroplast trnL P6 loop as well as nuclear ribosomal ITS (nrITS). The latter can increase taxonomic resolution of plant identifications but requires DNA to be relatively well preserved because of the target length (∼300–500 bp). We compare DNA results to pollen and macrofossil analyses from permafrost and ice-preserved faeces of Pleistocene and Holocene megafauna. Samples include woolly mammoth, horse, steppe bison as well as Holocene and extant caribou. Most plant identifications were found using DNA, likely because the studied faeces contained many vegetative remains that could not be identified using macrofossils or pollen. Several taxa were, however, identified to lower taxonomic levels uniquely with macrofossil and pollen analysis. The nrITS marker provides species level taxonomic resolution for commonly encountered plant families that are hard to distinguish using the other proxies (e.g. Asteraceae, Cyperaceae and Poaceae). Integrating the results from all proxies, we are able to accurately reconstruct known diets and habitats of the extant caribou. Applying this approach to the extinct mammals, we find that the Holocene horse and steppe bison were not strict grazers but mixed feeders living in a marshy wetland environment. The mammoths showed highly varying diets from different non-analogous habitats. This confirms the presence of a mosaic of habitats in the Pleistocene ‘mammoth steppe’ that mammoths could fully exploit due to their flexibility in food choice.

Small cave bear (U. ex gr. savini-rossicus) as a game species of prehistoric man

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.