Woolly Rhinoceros Research Articles

The woolly rhinoceros (Coelodonta antiquitatis) from Ondorkhaan, eastern Mongolia

The Mammoth Faunas, the famous cold‐adapted mammal assemblages, were distributed widely across northern Eurasia and North America during the Late Pleistocene. The now extinct woolly rhinoceros, Coelodonta antiquitatis, was a major component. Abundant fossil remains of this species with radiocarbon dates have been reported through almost all of northern Eurasia, but the fossil rhinoceroses of Mongolia are poorly known. Here, we describe a rhinocerotid skeleton from Ondorkhaan, eastern Mongolia, and compare it with four Late Pleistocene rhinoceros species of northern Eurasia (Elasmotherium sibiricum, Stephanorhinus hemitoechus, Stephanorhinus kirchbergensis and C. antiquitatis), resulting in its identification as a woolly rhinoceros (C. antiquitatis). Accelerator mass spectrometry radiocarbon dates obtained from two samples (ODK01 and ODK02) of the skeleton essentially agree, 42 160–40 040 cal. a BP and 42 105–39 955 cal. a BP, and the two samples had δ13C values of −19.5‰ and −20.2‰ SMMKW, respectively. This find suggests that the Mammoth Faunas were distributed in eastern Mongolia c. 45–40 ka during the period of climatic amelioration between Heinrich events 5 (H5: 46 000 cal. a BP) and 4 (H4: 39 000 cal. a BP).

Coelodonta antiquitatis in the Pleistocene of Bulgaria (Perissodactyla: Rhinocerotidae)

An analysis of the geographical and altitudinal distribution of remains of the woolly rhinoceros (Coelodonta antiquitatis) in Bulgaria is given. The paper summarizes all scattered data on the distribution of the species gathered in the last 114 years, only a part of them having been published before. Data on nine fossil localities (early–latest Pleistocene) from six provinces of Bulgaria are presented. The Fossil record proves the wide species distribution in the riverine foothills of mountains, plains, and lowlands of the country. Its Pleistocene localities were concentrated in the Danubian Plain, although the species was recorded also in the southernmost part of the country. About 90 percent of the localities were situated below 400 m a. s. l., although the species’ altitudinal distribution reached the maximum of 750 m a. s. l. All findings of the wooly rhinoceros in Bulgaria were made together with those of some species that now inhabit deciduous woodlands or rocky massifs of the temperate zone – Cervus elaphus, Capreolus capreolus, Rupicapra rupicapra, or Capra ibex, as well as extinct Bos primigenius. In Bulgaria, C. antiquitatis showed the same habitat preferences as in the core parts of the species range in Siberia. The Bulgarian localities of findings were both natural habitats and former human dwellings.

New Complex Investigations of the Novhorod-Siverskyi Upper Palaeolithic Site

In the article the materials of the Novhorod-Siverskyi site research both in the 1930's, and relatively recently, are analyzed. First of all, attention is paid to the faunal collection. The faunal associations from old excavations are coherent, typical of a cold steppe environment from the glacial period, near a riparian forest. However, it seems to result to a mix between natural taphonomic complexes and animal remains associated with human activity. Judging by the flint collection from the old excavations, the site was inhabited by representatives of the Pushkari type. New research yielded two archaeological layers that are very poor in flint artefacts. Concerning fauna, the faunal spectrum is restricted in both layers, with the woolly mammoth, the woolly rhinoceros, the horse and the reindeer in the lower and upper layers. In the upper layer there are also the bison, the fox and the hare. According to taphonomic observations, the bones had remained for a long time at an open air before being buried, in subsurface in a wet environment but few submitted to precipitations in link with permafrost activities. Particularly in the lower layer (2) some remains appear to be in place, while others seem to have been imported by hydraulic phenomena, either from the top of the promontory or from the Desna River. In the upper layer (1), some bones show possible anthropogenic impacts of breakage, linked to marrow recovery. In both layers we have some cranial and postcranial elements, mainly from adults sensu lato which could correspond to human predation. They could correspond to temporary camps of quite small human groups, potentially occupied at the end of the cold season/beginning of the warm season.

Rhinoceros exploitation at Érd (Hungary). What a place for the megaherbivores in the Neanderthal diet in Hungary?

The Hungarian Transdanubian site of Érd, where a Mousterian industry and abundant osteological material were discovered in the early 1960s is well known to prehistorians. The remains of megaherbivores (Mammuthus primigenius, Coelodonta antiquitatis) are re-examined here under the taphonomic and archaeozoological components in order to complete the Hungarian and European s.l. data and reassess the potential exploitation of these two pachyderms in the Neanderthal diet and economy. The cut marks, the intense activity of carnivores/hyenas and the skeletal profiles indicate a mixed origin of the carcasses. Mortality patterns of rhinoceros are characterized by the presence of young, subadult and adults, and suggest multiple acquisition by active scavenging and/or hunting with quick access. Skeletal profiles suggest a selective transport of rich/nutritive elements by humans to the site. The cut marks and fracturing of some elements (in situ butchery treatment) confirm that Neanderthals consumed these species on site and that they had at least partial primary access. The mode of acquisition seems active with rapid access for a young mammoth. Érd confirms the Neanderthal exploitation of rhinos and mammoths in their steppic environment during the Middle Palaeolithic. Érd is currently the only Hungarian Middle Palaeolithic site with a proven exploitation and consumption of these megaherbivores.

Nonpyrogenic charring of Late Pleistocene large mammal remains in northeastern Russia

Mammal remains preserved in the permafrost zone often bear traces of postmortem transformations, reflecting aspects of the palaeoenvironment and the processes that took place during the accumulation of host sediments. Multidisciplinary studies including radiocarbon dating, infrared spectroscopy, and microfossil analyses and grain size of infilling sediments from remains allow recognition of their stratigraphical and palaeogeographical origins and facilitate reconstructions of taphonomic pathways and the pre‐burial environments. Here, as exemplified by skulls of woolly rhinoceros, cave lion, and ancient bison, some distinct features of postmortem changes such as nonpyrogenic charring and vivianite encrustation indicate that the remains have undergone a complex range of burial processes in aerobic and anaerobic conditions in the Pleistocene landscapes of Arctic northeastern Russia. We hypothesize that these processes were mainly confined to the warmer intervals in the Late Pleistocene.

Prospects for new sites of wooly rhinoceros and mammoths on the territory of paleontological site in Starunia, Carpathian region

An international geopark “Park of the Ice Age” located near the Starunia village in Bohorodchany district of Ivano-Frankivsk region is world-famous for being a paleontological site of Pleistocene fauna of the woolly rhinoceroses and mammoth (the Eemian interglacial period, 46–10 000 years ago according to radiocarbon dating C14) and for its mud volcano which is unique in the Carpathian region. This is a geological nature monument with an area of 60 hectares where abandoned ozokerite mine and oil and gas exploratory boreholes are presented. The detailed investigations carried out in the years 2004–2009 by Ukrainian-Polish expeditions were focused on discoveries of possible new sites of fauna fossils and human remains of European early modem humans (EEMH).The future discoveries at the paleontological site of Starunia are extremely important. They intend to carry out ecological assessment of soil, surface and stream waters, atmospheric air, snow, ashes of meadow grasses, radiological screening, and geo-radar sensing to find new locations of extinct mammals. The Starunia area meets all the requirements of a geopark according to UNESCO standards and concepts. The first research findings (1907) are connected with the remains of woolly rhinoceros, mammoth, horse, roe deer and other Pleistocene mammals which had been found in the ozokerite mine at the depth of 12 m near Starunia (Bohorodchany district, Ivano-Frankivsk region). In 1914 the scientists from Lviv (Ukraine) and Krakow (Poland) fully appreciated these discoveries and published a set of articles and a monograph. In 1929 the members of the expedition of Skill Academy (Krakow, Poland) found the remains of three more woolly rhinoceroses in the ozokerite mine at the depth of 17 m. Numerous bones of vertebrates (rodents), remains of mussels, a lot of species of insects, beetles, parasitic worms, fleas, butterflies, spiders, snails, vascular plants, mosses, seeds and branches of dwarf birch (Betula Nana), alder (Alnus glutinosa), and other fossil of tundra flora were also discovered. Polish scientists organized detailed investigations of flora and fauna around Starunia, their results were published in articles but the World War II suspended this process. The investigations around Starunia intensified the comprehensive study of stratigraphy, paleontology, paleogeography, geochronology and other aspects of the Quaternary Period. In 1932 there was established the International Geological Congress (the International Union of Quaternary Study) by INQUA in Leningrad. The main methodological principals of this study dealt with the investigations around Starunia.

Paleo-Ecology of the Yedoma Ice Complex on Sobo-Sise Island (EasternLena Delta, Siberian Arctic)

Late Pleistocene permafrost of the Yedoma type constitutes a valuable paleo-environmental archive due to the presence of numerous and well-preserved floral and faunal fossils. The study of the fossil Yedoma inventory allows for qualitative and quantitative reconstructions of past ecosystem and climate conditions and variations over time. Here, we present the results of combined paleo-proxy studies including pollen, chironomid, diatom and mammal fossil analyses from a prominent Yedoma cliff on Sobo-Sise Island in the eastern Lena Delta, NE Siberia to complement previous and ongoing paleo-ecological research in western Beringia. The Yedoma Ice Complex (IC) cliff on Sobo-Sise Island (up to 28 m high, 1.7 km long) was continuously sampled at 0.5 m resolution. The entire sequence covers the last about 52 cal kyr BP, but is not continuous as it shows substantial hiatuses at 36–29 cal kyr BP, at 20–17 cal kyr BP and at 15–7 cal kyr BP. The Marine Isotope Stage (MIS) 3 Yedoma IC (52–28 cal kyr BP) pollen spectra show typical features of tundra–steppe vegetation. Green algae remains indicate freshwater conditions. The chironomid assemblages vary considerably in abundance and diversity. Chironomid-based TJuly reconstructions during MIS 3 reveal warmer-than-today TJuly at about 51 cal kyr BP, 46-44 and 41 cal kyr BP. The MIS 2 Yedoma IC (28–15 cal kyr BP) pollen spectra represent tundra-steppe vegetation as during MIS 3, but higher abundance of Artemisia and lower abundances of algae remains indicate drier summer conditions. The chironomid records are poor. The MIS 1 (7–0 cal kyr BP) pollen spectra indicate shrub-tundra vegetation. The chironomid fauna is sparse and not diverse. The chironomid-based TJuly reconstruction supports similar-as-today temperatures at 6.4–4.4 cal kyr BP. Diatoms were recorded only after about 6.4 cal kyr BP. The Sobo-Sise Yedoma record preserves traces of the West Beringian tundra-steppe that maintained the Mammoth fauna including rare evidence for woolly rhinoceros’ presence. Chironomid-based TJuly reconstructions complement previous plant-macrofossil based TJuly of regional MIS 3 records. Our study from the eastern Lena Delta fits into and extends previous paleo-ecological Yedoma studies to characterize Beringian paleo-environments in the Laptev Sea coastal region.

Late Pleistocene chronology and environment of woolly rhinoceros (Coelodonta antiquitatis (Blumenbach, 1799)) in Beringia

Uncertain chronology and data scarcity have impeded realistic reconstructions of megafauna extinctions in the Late Pleistocene in several key regions of the Northern Hemisphere such as Beringia. This region was a refuge for several plants, animals during the extremely cold period of the Late Pleistocene in high Arctic latitudes. The woolly rhinoceros was one of the most widespread members of the megafauna in the Asiatic part of the region (West Beringia) between ∼60 and 14 cal ka BP. This study is based on statistical analyses of 20 newly obtained and 110 previously published radiocarbon dates. We found three large “waves” in the woolly rhinoceros range changes separated between themselves by the cold climatic Heinrich events (H2 and H4). The chronology of the woolly rhinoceros was overlaid on data of environmental changes obtained basing on 504 generalized early published pollen spectra throughout the species range and, separately, outside the range – in the east of the West Beringia realm and in East Beringia. In general, milder environmental conditions of MIS3 (57–29 ka BP) were more favourable for the woolly rhinoceros than the harsh conditions of the Last Glacial Maximum (∼29–15 cal ka BP) in MIS2. We have concluded that the feed base was unlikely the main limiting factor in the distribution of woolly rhinoceros in Western Beringia, and other ecological factors (temperature/precipitation) determined the species range and its oscillation over the MIS3–MIS2 stages. Based on summarizing available data and this research, we have proposed that there were sets of different reasons that prevented the woolly rhinoceros migration to the east of Beringia in different periods of the Late Pleistocene. Abrupt woolly rhinoceros extinction in Beringia between 15 and 14 cal ka BP coincided with the Bölling warming and the Older Dryas cooling. The ecological situation just before the extinction, associated with climate warming, moisture increasing and shrub tundra expansion in West Beringia, was qualitatively different from previous cases of the species range degradations in the second half of the Late Pleistocene. This multi-proxy study of woolly rhinoceros chronology provide a new basis for further understanding of its population history, demography, and biology in Beringia before its extinction.

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.

Late Pleistocene paleoecology and phylogeography of woolly rhinoceroses

The woolly rhinoceros (Coelodonta antiquitatis) was a cold-adapted herbivore, widely distributed from western Europe to north-east Siberia during the Late Pleistocene. Previous studies have associated the extinction of the species ∼14,000 calendar years before present to climatic and vegetational changes, suggesting the later survival of populations in north-east Siberia may have related to the later persistence of open vegetation in the region. Here, we analyzed carbon (δ13C) and nitrogen (δ15N) stable isotopes and mitochondrial DNA sequences to elucidate the evolutionary ecology of the species. Our dataset comprised 286 woolly rhinoceros isotopic records, including 192 unpublished records, from across the species range, dating from >58,600 to 12,135 14C years before present (equivalent to 14,040 calendar years ago). Crucially, we present the first 71 isotopic records available to date of the 15,000 years preceding woolly rhinoceros extinction. The data revealed ecological flexibility and geographic variation in woolly rhinoceros stable isotope compositions across time. In north-east Siberia, we detected stability in δ15N through time, which could reflect long-term environmental stability, and may have enabled the later survival of the species in the region. To further investigate the paleoecology of woolly rhinoceroses, we compared their isotopic compositions with other contemporary herbivores. Our findings suggested isotopic similarities between woolly rhinoceros and both musk ox (Ovibos moschatus) and saiga (Saiga tatarica), albeit at varying points in time, and possible niche partitioning between woolly rhinoceros and both horse (Equus spp.) and woolly mammoth (Mammuthus primigenius). To provide phylogeographic context to the isotopic data, we compiled and analyzed the 61 published mitochondrial control region sequences. The genetic data showed a lack of geographic structuring; we found three haplogroups with overlapping distributions, all of which showed a signal of expansion during the Last Glacial Maximum. Furthermore, our genetic findings support the notion that environmental stability in Siberia influenced the paleoecology of woolly rhinoceroses in the region. Our study highlights the utility of combining stable isotopic records with ancient DNA to advance our knowledge of the evolutionary ecology of past populations and extinct species.

The mammoth's mysterious companion: the materials of the Republican ecological and paleontological camp for schoolchildren “Tusk,” years 1997–2013

This article deals with a unique discovery. In 2010, a school expedition found an obscure skull of a woolly rhinoceros on Mamontova Mountain in the Republic of Sakha (Yakutia). Young paleontologists with their leader P. D. Maksimov conducted an anatomical study of the woolly rhinoceros' fossil skull. During the entire period (years 1973–2021) of the camp's existence, young paleontologists have found the remains of 57 animal fossils. All the materials found during the expeditions have been stored in the museum-laboratory at the Republican Ecological-Paleontological Camp of Schoolchildren “Tusk.” Among the gathered paleontological finds, the young researchers were attracted to rhinoceros skulls found at different times near various excavations. The expedition participants have put forward the hypothesis that during the period of the flourishing of the mammoth fauna, there might have been some species or subspecies of woolly rhinoceros on the territory of Paleoyakutia besides the Lena species. The results of the research were highly appreciated by the experts of the republican scientific conferences.

Thick skin cutters of Siberian frozen mummies—The coevolutionary adaptation of Eurasian Ice Age spotted hyenas

AbstractSiberian extinct top predators—Ice Age spotted hyenas—are discovered in the permafrost of Yakutia in‐between frozen mammoth, rhinoceros, bison or horse carcasses, such as in Mongolia at new open‐air sites. Historic described European holotypes and new crania from Siberia, Mongolia, and Europe allow presenting the monophyly of Crocuta reaching back 2.53 Mio years (Late Pliocene). Spotted hyenas coevolved in dental change and body size to their largest guilt, thick‐skinned elephants and rhinoceroses. The Late Pleistocene (MIS 5d‐3, 113.00–25.000 BP) Crocuta crocuta spelaea (Goldfuss, 1823) is proven to have scavenged Siberian woolly rhinoceros Coeleodonta antiquitatis (Blumenbach, 1799) carcasses. Rhino mummy skins have scavenging signs attributable only to hyenas. Chew cuts of breaking‐cutting scissor dentition are found at mummy skins. The last Eurasian hyenas left always similar chew‐damaged woolly rhinoceroses’ crania with braincase openings and damaged mandibles, whereas their deep bite scratches are often found on the distal joints of long bones. Strongest rhino bone damage is found at natal den sites (cave or open air), to which hyenas imported guilt to their cubs. The recently known Ice Age spotted hyena palaeobiogeography overlaps exactly with those of woolly rhinoceros and mammoths. All reached Bering Straits.

A re-examination of the West European species of Boreonectes Angus, 2010, with particular reference to B. multilineatus (Falkenström, 1922) (Coleoptera, Dytiscidae).

The West European species of Boreonectes Angus, 2010 are reviewed. B. multilineatus (Falkenström, 1922) is shown to be widely distributed in the Pyrenees, where it is the only species known to occur. The chromosomes of all five west European species are found to have, in addition their different numbers of chromosomes, differences in the number and locations of secondary constrictions, and in some cases, the number of chromosomes with clear centromeric C-bands. The level of differences between the chromosomes of the species is in stark contrast with the very slight genetic (DNA) differences between them and this suggests that chromosome differentiation may have been a driver of speciation. Two of the species, B. griseostriatus (De Geer, 1774) and B. multilineatus, have distributions extending northwards as far as Arctic Scandinavia. It is pointed out that, while these northern areas now constitute the major portions of their ranges, they must be of fairly recent origins as most of the area would have been covered by ice sheets and therefore not habitable during the glacial maximum of the Last Glaciation. This contrasts with the situation in the area of the Central European mountains where fossil faunas, including Boreonectes, are known. B. griseostriatus, identifiable to species by its parameres, was present in the Woolly Rhinoceros site at Starunia in the Western Ukraine, and this fauna is discussed as well as an English fauna of similar age.

Исследовательские работы на верхнепалеолитической стоянке Актас (Северный Казахстан) в 2021 году

In May 2021, a joint expedition of the Institute of Archeology and Ethnography SB RAS and the National Museum of the Republic of Kazakhstan carried out archaeological survey at the Aktas site in northern Kazakhstan. The Aktas site is located on the flat top of a hill (420 m) with a quartzite rocky outcrop. The site was studied by V.N. Matvienko in 1982-1983. At that time, a representative collection of faunal remains (several thousand items) was recovered from the excavation area of 25 sq.m. It contains remains of Crocuta crocuta spelaea, Panthera leo spelaea, Coelodonta antiquitatis, Bos primigenius, Cervus elaphus. Occurrence of objects with traces of working and lithic products interpreted as tools allowed researchers to correlate the materials from this site with Upper Paleolithic human activities. The main task of the works in 2021 was to verify involvement of ancient humans information of the materials at the site and determination of its age. A new stratigraphic profile containing 6 lithological layers was established along the wall of the excavation trench of 1982-1983. Faunal remains were mostly collected from layers 3 (brownish-gray sandy loam) and 4 (detrital horizon). In total, 82 animal bones were recovered including remains of Coelodonta antiquitatis, Equus ferus and Caprinae gen. No lithic artifacts were found, however, solitary bones bearing traces of probable anthropogenic impact (splitting, polishing) were found. The Aktas location on a hill excludes the possibility of bone-bed formation due to natural geological processes. The derived information makes it possible to link the accumulated materials at the site to anthropogenic activity. Based on the composition of faunal remains, Aktas was aged to late MIS 3 - early MIS 2.

Taphonomy of Pleistocene large mammal remains in the deposits of river Raba, southern Poland

An assemblage of 120 mammal remains of Pleistocene age has been collected from the fluvial deposits of river Raba at a gravel pit in the village of Targowisko, 30 km east of Kraków, southern Poland. Nearly 100 remains represent woolly mammoth Mammuthus primigenius . Other remains belong to four or five such mammal species as horse Equus ferus , woolly rhinoceros Coelodonta antiquitatis, red deer Cervus elaphus and steppe bison Bison priscus or aurochs Bos primigenius . Pleistocene coarse-grained deposits containing isolated bones, teeth and tusks occur in the lowermost part of the fluvial succession in the open pit, presently inundated by groundwater. The surfaces of the majority of bones and teeth show abrasion damages by fluvial transport, including their rounding and smoothing as well as scratches and grooves. Traces of carnivore activity are visible on mammoth and horse bones. The location, dimension and shape of these marks suggest wolf or cave hyena gnawing.

The Mammuthus-Coelodonta Faunal Complex at its southeastern limit: A biogeochemical paleoecology investigation in Northeast Asia

During the past several decades, the paleoecology of the Mammuthus-Coelodonta Faunal Complex in the Palearctic has been thoroughly explored, especially using stable isotope analysis. Numerous studies have documented high ecological plasticity and regional heterogeneities for this fauna. However, very limited attention has focused on Northeast Asia, at the southeastern edge of the distribution of the mammoth steppe biome. In the present study, we undertook radiocarbon dating, zooarchaeology by mass spectrometry (ZooMS), and stable isotope analysis on the fossil faunas from Yanjiagang Paleolithic site, Northeast (NE) China, and from the Geographical Society Cave in the nearby Russian Far East, to explore the paleoecology of this fauna in this middle-latitude region. Isotopic (δ13C, δ15N) data from these two sites suggested that the woolly mammoth (Mammuthus primigenius) was a grazer feeding on grasses/sedges, while the woolly rhinoceros (Coelodonta antiquitatis) and steppe bison (Bison priscus) utilized a wider range of food resources. All megaherbivores exhibited some ecological plasticity. Meanwhile, the mammal remains from Geographical Society Cave prevalently exhibited lower δ15N values than those from China, indicating variable environments and vegetation in Northeast Asia during MIS 3. Interestingly, through reevaluating the diachronic mammal distribution and ecology based on direct radiocarbon dating and ZooMS, we report the credible presence of Bubalus, typically interpreted as a mild climate faunal element in the Pleistocene, within the Mammuthus-Coelodonta Faunal Complex in Yanjiagang. This emphasizes the unique scenario of this biome in Northeast Asia, where the fauna exhibited sensitivity to synergic effects of climatic oscillations and local geographic traits. After comparing isotopic data of mammoth from different subregions of Asia, we found that NE China displayed both highest δ13C and δ15N values, possibly related to the higher temperature compared to the Arctic regions. This study reveals the characteristics and complexity at the southeastern limit of the range of the mammoth steppe biome and urges more systematic studies within and outside this region.

Ecological conditions during the Middle to Upper Palaeolithic transition (MIS 3) in Iberia: the cold‐adapted faunal remains from Mainea, northern Iberian Peninsula

Cold‐adapted faunal populations shifted southwards during the coldest episodes of the Middle and Late Pleistocene, reaching the Iberian Peninsula. In this context, some of these species, such as Coelodonta antiquitatis and Mammuthus primigenius extended throughout the territory, leaving evidence in, at least, 35 and 29 sites, respectively; however, in most cases, the evidence is scarce. Here we present the fossil assemblage from Mainea cave (Uitzi, Navarre, northern Iberian Peninsula), one of the sites yielding cold‐adapted faunal remains closer to the Pyrenees. The cave possibly acted as a natural trap due to its external geomorphology (a sinkhole) and the absence of aerial exposure, anthropogenic marks or evidence compatible with this site being a hyena den. This site has yielded the most evidence of Coelodonta antiquitatis remains (NISP = 190) and one of the most abundant in terms of number of individuals (five, each with different ages at death) for the Iberian Peninsula, five remains attributed to a single Mammuthus primigenius and a single Cervus elaphus remain. The direct dating of a woolly rhinoceros tooth (46 354–44 428 2σ cal. a BP) places Mainea within MIS 3, between the H4 and H5 Heinrich Events, during the Middle to Upper Palaeolithic transition, within a context of changes in human behaviour and human species replacement in Europe. We have performed the first collagen isotopic analysis for a woolly rhinoceros from the Iberian Peninsula. The δ13C and δ15N values of the Coelodonta antiquitatis remains are similar to other European sites, thus suggesting the presence of Mammoth steppe environments at relatively high altitudes, which would compensate for the southern latitude. This would be consistent with the presence of open environments in lower altitudes. This site expands our understanding of the ecology of cold‐adapted faunas at their furthest southern range limit during the Middle to Upper Palaeolithic transition.

A New Glance at Old Samples: Remains of Freshwater Invertebrates Associated with Mummified Carcasses of Large Quaternary Mammals

Carcasses of mammals with well-preserved soft tissues from the permafrost of Siberia and northern North America are among the most important sources of our knowledge on Quaternary environments. However, the potential of this information source is far from being exhausted. The objective of this study is to reveal and describe the remains of freshwater invertebrates in the hair and gut contents of large Pleistocene mammals stored for a long time in museum collections. We have studied the gut contents of two mammoths, as well as the hair of two other mammoths, one woolly rhinoceros, four bison, and two horses. Remains of invertebrates were found in the gut contents of both mammoths, as well as in the hair of two mammoths and the woolly rhinoceros, while no remains were detected in the bison and horse hair. Among the remains of microscopic invertebrates, the most common are branchiopod crustaceans, namely the ephippia of Daphnia sp., Simocephalus sp., the head shields and valves of Chydoridae (Cladocera), and the mandibles of tadpole shrimps (Notostraca). Based on our current level of knowledge, we are able to identify only a limited number of Pleistocene branchiopods. Studies on Recent taxa with the aim to develop identification keys to the aforementioned groups are needed, as they could significantly expand the potential of palaeoecological analysis.

Quaternary megafauna from the Dnieper alluvium near Kaniv (central Ukraine): Implications for biostratigraphy

Subfossil remains of the Quaternary megafauna from two Ukrainian natural history collections stored at Kaniv Nature Reserve and the National Museum of Natural History NAS of Ukraine were examined. Most of the bones were collected in 1965–1966, in the building pit and the gateway of the Kaniv hydroelectric power plant. The fauna is represented at least by ten taxa (Gulo gulo, Mammuthus trogonterii chosaricus, Mammuthus intermedius, Coelodonta antiquitatis, Megaloceros giganteus, Alces alces, Bison priscus, Cervus elaphus, Capra cf. ibex, and Equus ferus). Skeletal parts are represented mainly by skulls, long bones, horns, and tusks. Based on the species composition of proboscideans, at least part of the faunal assemblage is dated by the end of the Middle Pleistocene (Dnieper Stage = Saale, Warta, MIS 6), but majority of bones could be older or younger (Late Pleistocene and Holocene age) due to the alluvial origin of accumulation.

Evolution: Untangling the Woolly Rhino’s Extinction

The woolly rhinoceros was a charismatic inhabitant of the frigid steppes of Pleistocene Eurasia. Now, the genome of an 18,500-year-old woolly rhino has been sequenced. It points to a thriving population less than 5000 years before the species disappeared.