Extinction chronology of the cave lion Panthera spelaea

The cave lion, Panthera spelaea, was widespread across northern Eurasia and Alaska/Yukon during the Late Pleistocene. Both morphology and DNA indicate an animal distinct from modern lions (probably at the species level) so that its disappearance in the Late Pleistocene should be treated as a true extinction. New AMS radiocarbon dates directly on cave lion from across its range, together with published dates from other studies – totalling 111 dates – indicate extinction across Eurasia in the interval ca. 14–14.5 cal ka BP, and in Alaska/Yukon about a thousand years later. It is likely that its extinction occurred directly or indirectly in response to the climatic warming that occurred ca. 14.7 cal ka BP at the onset of Greenland Interstadial 1, accompanied by a spread of shrubs and trees and reduction in open habitats. Possibly there was also a concomitant reduction in abundance of available prey, although most of its probable prey species survived substantially later. At present it is unclear whether human expansion in the Lateglacial might have played a role in cave lion extinction. Gaps in the temporal pattern of dates suggest earlier temporary contractions of range, ca. 40–35 cal ka BP in Siberia (during MIS 3) and ca. 25–20 cal ka BP in Europe (during the ‘Last Glacial Maximum’), but further dates are required to corroborate these. The Holocene expansion of modern lion (Panthera leo) into south-west Asia and south-east Europe re-occupied part of the former range of P. spelaea, but the Late Pleistocene temporal and geographical relationships of the two species are unknown.     

Late Pleistocene steppe lion Panthera leo spelaea (Goldfuss, 1810) footprints and bone records from open air sites in northern Germany – Evidence of hyena-lion antagonism and scavenging in Europe

Bone remains and a trackway of Pantheraichnus bottropensis nov. ichg. ichnsp. of the Late Pleistocene lion Panthera leo spelaea (Goldfuss, 1810) have been recovered from Bottrop and other open air sites in northern Germany. Some of these bones are from open air hyena den sites. A relative high proportion of lion bones (20%) exhibit bite, chew or nibble marks, or bone crushing and nibbling caused by a large carnivore. Repeated patterns of similar bone damage have been compared to bone remains found at hyena dens in gypsum karst areas and cave sites in northern Germany. Ice Age spotted hyenas have been the main antagonists and the main scavengers on lion carcasses. The remains appear to have been imported often by hyenas into their communal dens, supporting the theory of strong hyena-lion antagonism, similar to the well documented antagonism between modern African lions and spotted hyenas. Most of the lion bones from the open air hyena den at Bottrop are probably a result of such antagonism, as are the rare remains of these carnivores found within large hyena prey bone accumulations along the Pleistocene rivers. The Emscher River terrace also has the largest quantity of hyena remains from open air river terrace sites in northern Germany. Their cub remains, and incomplete chewed prey bones from mammoths and woolly rhinoceroses, typical of hyena activity, underline the character of these sites as cub-raising and communal dens, where their prey was accumulated along the riverbanks in a similar manner to modern African hyenas.     

Steppe lion remains imported by Ice Age spotted hyenas into the Late Pleistocene Perick Caves hyena den in northern Germany

Upper Pleistocene remains of the Ice Age steppe lion Panthera leo spelaea (Goldfuss, 1810) have been found in the Perick Caves, Sauerland Karst, NW Germany. Bones from many hyenas and their imported prey dating from the Lower to Middle Weichselian have also been recovered from the Perick Cave hyena den. These are commonly cracked or exhibit deep chew marks. The absence of lion cub bones, in contrast to hyena and cave bear cub remains in the Perick Caves, and other caves of northern Germany, excludes the possibility that P. leo spelaea used the cave for raising cubs. Only in the Wilhelms Cave was a single skeleton of a cub found in a hyena den. Evidence of the chewing, nibbling and cracking of lion bones and crania must have resulted from the importation and destruction of lion carcasses (4% of the prey fauna). Similar evidence was preserved at other hyena den caves and open air sites in Germany. The bone material from the Perick and other Central European caves points to antagonistic hyena and lion conflicts, similar to clashes of their modern African relatives.     

The first find of the cave hyena (<Emphasis Type="Italic">Crocuta crocuta spelea</Emphasis>) on the Oka plateau (Eastern Sayan)

These data indicate that the last representatives of Hyaenidae on the Oka plateau, i.e., cave hyenas (C. c. spelaea), are direct descendants of the hyenas that lived in the Altai–Sayan and Baikal mountain country from the Late Miocene to the end of the Pleistocene. Their continued existence over a long time undoubtedly indicates the absence of repeated glaciation during the Pleistocene. Cave hyenas are relics of the late Tertiary fauna in the area of Lake Baikal; they became extinct due to gradual cooling, which peaked at the end of the Late Pleistocene.     

Co-existence of scimitar-toothed cats,lions and hominins in the European Pleistocene. Implications of the post-cranial anatomy of Homotherium latidens (Owen) for comparative palaeoecology

Human occupants of Europe shared food resources with a number of larger Carnivora, and their coexistence with two lion-sized felids, the lion and the scimitar-toothed machairodont Homotherium latidens, poses intriguing paleoecological problems. We investigate the ecology of Homotherium latidens using an exceptional sample of postcrania from the Spanish Early Pleistocene site of Incarcal, making comparisons with modern cats and with other machairodont species. Evidence of cursorial adaptations in Homotherium suggests a hunting technique different from modern cats or smilodontine sabre-tooths. Some, like reduction of the claws, would have limited the ability of individual homotheres to bring down large prey, implying group action. Homotherium would also have been disadvantaged in direct confrontation with Pleistocene lions by smaller body mass, reduced forepaw muscle strength, smaller claws and more fragile dentition. Its hunting technique would have worked best in more open habitats, but competition from lions would have forced it to seek moderate cover. Among factors that could de-stabilise coexistence of the two big cat species in Pleistocene Europe we invoke a decrease in environmental mosaicism associated with stepped climatic change over the last million years, and the increased importance of humans within the larger predator guild.     

Age of Middle Pleistocene fauna and Lower Palaeolithic industries from Kent's Cavern,Devon

Kent's Cavern has long been known as potentially among the oldest Palaeolithic sites in the country, with the basal Breccia deposit containing a sparse Lower Palaeolithic industry. The sediment consists of a chaotic clayey conglomerate emplaced as a series of debris flows, which entered the cave via blocked entrances at its southwest end. The Breccia contains a fauna dominated by the bear Ursus deningeri, with lion Felis leo and the voles Arvicola cantiana and Microtus oeconomus, establishing a late Cromerian age for the deposit. The artefacts comprise an industry of crudely manufactured handaxes and flakes, and show damage suggesting that they were brought into the cave by the debris flows, and may thus predate the sediment and fauna. We demonstrate an age of >340 ka for the Breccia using two independant dating methods, consistent with existing models of the age of the British Middle Pleistocene sequence.     

The first record of “spelaeoid” bears in Arctic Siberia

For a long time, “spelaeoid” (cave-bear-like) bears, Ursus (Spelearctos) spp., were believed to be almost purely European animals. Their geographic range has recently been extended to the east, in southern Siberia, Transbaikalia, Kirghizia, Mongolia and Korea. Two unexpected new findings, presented here in detail, significantly change existing views on the distribution of cave bears; both were found in North-Eastern Siberia, far beyond the Arctic Circle, more than 1500 km to the north-east of the previously accepted range.One of the fossils is a mandible, found near the town of Cherskiy at 68.73°N, 161.38°E. The analysis of local geology and accompanying mammal fossils suggests that it comes from the Olyorian Fauna (Early to early Middle Pleistocene). Morphologically, the Cherskiy mandible is closest to Ursus savini, a small middle Pleistocene cave bear from the British Cromer Forest-bed Formation, but differs in having a slightly more advanced dentition, and thus it is described as a new subspecies Ursus savini nordostensis. Another newly recognized fossil of the “spelaeoid” bear is an astragalus found at the Oskhordokh site at 67.54°N, 135.67°E, on a large gravel bar on the right bank of the Adycha River. This specimen is attributed to Ursus cf. deningeri.The paper also presents an interesting example of the interaction between classical and “molecular” palaeontology.The new finds significantly change existing ideas on the ecology and evolution of cave bears, some of the most remarkable members of the extinct Pleistocene megafauna.     

Pengelly's legacy reconsidered: a GIS approach to spatial analysis of palaeontological and archaeological collections from Kents Cavern, England

Tens of thousands of palaeontological and archaeological remains were collected by William Pengelly during 19th century excavations of Kents Cavern, but are now widely dispersed between museums. This has previously precluded spatial analysis. We have now assembled available museum records into a single database, and, using our previously-reconstructed Pengelly excavation map as a base, we have been able to exploit the unique Pengelly location code to set up a GIS mapping system. This allows, for the first time, the analysis of spatial patterns. In addition, the GIS serves to highlight potential problems of recording or curation in the original data. Here we report on the construction of the GIS system and its first use in the analysis of spatial distribution of bear remains. The maps demonstrate that Ursus deningeri entered the cave through a now-sealed High Level Chamber entrance at the back of the cave in the middle Pleistocene, whereas Ursus arctos accessed the cave in the late Pleistocene through the now-sealed Northeast Gallery entrance. The denning areas are reconstructed as Labyrinth/Bear's Den for U. deningeri and Vestibule/Great Chamber for U. arctos. Considerable post-mortem re-distribution of the remains of both species is indicated.     

First record of the Eurasian jaguar in southern Asia and a review of dental differences between pantherine cats

Felid material from the middle Early Pleistocene Haro River quarry, northern Pakistan, is described. Most of the material is referred to the Eurasian jaguar, Panthera gombaszogensis georgica, representing the first record of the jaguar in southern Asia. A systematic review of dental morphology of all living species of Pantherinae is made to provide a better background for studying the evolution of pantherine cats. A renewed dispersal scenario of the jaguar in Eurasia is provided. Panthera gombaszogensis georgica is suggested to be a distinct subspecies distributed in central, western and southern Asia during the Early Pleistocene, characterized by relatively small body size and slender dentition. The Eurasian jaguar does not possess the derived dental characters of the living jaguar Panthera onca (e.g. weak or no canine vertical groove, robust premolars), and presumably did not specialize on hard-shelled prey, and therefore should be viewed as a distinct species from the living P. onca. Two additional felids from the Haro River Quarry, the large sabre-toothed felid? Megantereon sp. and a small undetermined feline, are also reported. The Felidae faunal composition of southern Asia during the Early Pleistocene is generally similar to that of northern Eurasia, and the paleoenvironment is probably slightly more closed.     

Paleobiological Implications of the Isotopic Signatures (C,N) of Fossil Mammal Collagen in Scladina Cave (Sclayn, Belgium)

An isotopic investigation of upper Pleistocene mammal bones and teeth from Scladina cave (Sclayn, Belgium) demonstrated the very good quality of collagen preservation. A preliminary screening of the samples used the amount of nitrogen in whole bone and dentine in order to estimate the preserved amount of collagen before starting the extraction process. The isotopic abundances of fossil specimens from still-extant species are consistent with their trophic position. Moreover, the¹⁵N isotopic abundance is higher in dentine than in bone in bears and hyenas, a phenomenon already observed in modern specimens. These results demonstrate that the isotopic compositions of samples from Scladina cave can be interpreted in ecological terms. Mammoths exhibit a high¹⁵N isotopic abundance relative to other herbivores, as was the case in Siberian and Alaskan samples. These results suggest distinctive dietary adaptations in herbivores living in the mammoth steppe. Cave bears are clearly isotopically different from coeval brown bears, suggesting an ecological separation between species, with a pure vegetarian diet for cave bear and an omnivorous diet for brown bear.     

New palaeontological evidence suggests an early Middle Pleistocene age for the lower levels of Sun Hole Cave,Cheddar, Somerset,UK

Sun Hole, a small fissure cave on the north side of Cheddar Gorge, Somerset, is best known as the site where late Upper Palaeolithic artefacts are associated with human and other mammal remains that immediately pre-date the Lateglacial interstadial. These remains from the upper levels (Unit I) overlie sediments (Unit II) that are thought to have accumulated during a full glacial period, below which are sediments (Unit III) attributed to an interglacial stage. The extinct land snail Retinella (Lyrodiscus) sp., which in Britain was only known from the Hoxnian Stage, had previously been recovered from Unit III. The interglacial at the base of the sequence was therefore assigned to the Hoxnian Stage (Marine Isotope Stage (MIS) 11, 424–374 kyr). Here we provide additional analyses of samples from Unit III, which as well as containing further Retinella (Lyrodiscus), also yielded a specimen of Monachoides incarnatus, its first record from the British Pleistocene. An associated assemblage of small mammals included a few that are rare in the British Pleistocene, such as birch mouse (Sicista cf. betulina) and garden dormouse (Eliomys quercinus). The vole Lasiopodomys gregaloides and the shrew Sorex (Drepanosorex) savini do not occur as late as the Hoxnian and therefore suggest an older age in the early Middle Pleistocene. The record from Sun Hole therefore has parallels with the well-known sequence from the upper Calcareous Member in the nearby cavern system at Westbury-sub-Mendip.     

Late Quaternary environmental and human events at En Gedi, reflected by the geology and archaeology of the Moringa Cave (Dead Sea area, Israel)

The Moringa Cave within Pleistocene sediments in the En Gedi area of the Dead Sea Fault Escarpment contains a sequence of various Pleistocene lacustrine deposits associated with higher-than-today lake levels at the Dead Sea basin. In addition it contains Chalcolithic remains and 5th century BC burials attributed to the Persian period, cemented and covered by Late Holocene travertine flowstone. These deposits represent a chain of Late Pleistocene and Holocene interconnected environmental and human events, echoing broader scale regional and global climate events. A major shift between depositional environments is associated with the rapid fall of Lake Lisan level during the latest Pleistocene. This exposed the sediments, providing for cave formation processes sometime between the latest Pleistocene (ca. 15 ka) and the Middle Holocene (ca. 4500 BC), eventually leading to human use of the cave. The Chalcolithic use of the cave can be related to a relatively moist desert environment, probably related to a shift in the location of the northern boundary of the Saharo-Arabian desert belt. The travertine layer was U-Th dated 2.46 ± 0.10 to 2.10 ± 0.04 ka, in agreement with the archaeological finds from the Persian period. Together with the inner consistency of the dating results, this strongly supports the reliability of the radiometric ages. The 2.46-2.10 ka travertine deposition within the presently dry cave suggests a higher recharge of the Judean Desert aquifer, correlative to a rising Dead Sea towards the end of the 1st millennium BC. This suggests a relatively moist local and regional climate facilitating human habitation of the desert.     

Late Quaternary environments and biogeography in the Great Basin

Plant and animal remains found in packrat (Neotoma spp.) middens and cave fill from the eastern and southern Great Basin region reveal the presence of subalpine conifers and boreal mammals at relatively low elevations during the Late Wisconsin. Limber pine (Pinus flexilis) and bristlecone pine (P. longaeva) were important in the late Pleistocene plant communities throughout this region. Spruce (Picea cf. engelmannii) and common juniper (Juniperus communis) were present in some of the more northerly localities, and Douglas fir (Pseudotsuga menziesii) and white fir (Abies concolor) were present in southern and eastern localities. Single needle pinyon pine (Pinus monophylla), common across this region today, was apparently not present north of the Sheep Range of southern Nevada during the Late Wisconsin. Pikas (Ochotona cf. princeps), small boreal mammals present in only a few Great Basin mountain ranges today, were common throughout the region. Heather voles (Phenacomys cf. intermedius) have been found in two cave fill deposits in Nevada, though they are unknown in the Great Basin today. Limber and bristlecone pines are generally restricted to rocky substrates in modern subalpine habitats in the Great Basin, and this may also have been the case when these plants grew at lower elevations during the Late Wisconsin. Subalpine conifers were present on the rock outcrops sampled by the packrat middens, but shrub communities, perhaps dominated by sagebrush (Artemisia spp.), may have been present on alluvial valley-bottom substrates. Forested habitats would thus have been isolated habitat islands, as they are today. Boreal small mammals, including pikas and heather voles, were able to colonize the Great Basin mountain ranges during the late Pleistocene. We suggest that these mammals were able to survive in the intervening valley-bottoms under a cool-summer climatic regime, and that continuous forest or woodland corridors were not necessary for migration.     

广西巴马县水晶宫洞穴沉积物特征及其沉积环境

巴马县水晶宫洞内次生化学沉积物景观丰富,类型多样,有石钟乳、石幔、石瀑布、石笋、石柱、石盾等特色景观;尤其是近期仍在发育的洞顶鹅管、卷曲石、石花等是国内外较为罕见的洞穴奇景。据沉积物的沉积类型、物质成分、沉积序次及测年等资料研究表明,水晶宫洞穴沉积物形成于晚第三纪以来。晚第三纪和第四纪早更新世早期,洞内沉积主要为外源水携带的硅质岩、砂岩类砾石沉积形成的钙华砾石层。早更新世中期—中更新世气候温暖湿润,是水晶宫洞穴主要景观的形成期,沉积物景观宏大而壮观,230Th-U系测年得出其分别形成于100～35万年前。晚更新世早期12.5～9万年的末次间冰期主要为温暖气候环境,洞内仅有少量的石笋沉积,且极少保存;9万年后气候环境发生突变,更不利沉积物发育和保存;自4万年后气候环境开始表现为温暖湿润,化学沉积活跃,洞穴沉积物主要由高约50～150 cm的纯白色石笋群和钟乳石组成,为洞穴奇景的形成期,230Th-U系测年得出其分别形成于40 ka BP和7 500 a BP以及1 450 a BP。晚更新世晚期以来形成的洞穴沉积物景观,不仅为水晶宫提供了丰富的旅游资源和价值,同时也为广西重建古气候环境提供了重要的信息载体。      

周口店地区第四纪哺乳动物群的演化与环境变迁探讨

讨论了Ａｌｉｌｅｐｕｓａｎｎｅｃｔｅｎｓ、ＨｙｐｏｌａｇｕｓｂｒａｃｈｙｐｕｓＬｅｐｕｓｓｐ．、Ｓｃｉｕｒｏｔａｍｉａｓｐｒａｅｃｏｘ、ＡｌｌｏｃｒｉｃｅｔｕｓｔｅｉｌｈａｒｄｉＣｒｉｃｅｔｉｎｕｓｖａｒｉａｎｓ、Ｈｙｐｅｒａｃｒｉｕｓｙｅｎｓｈａｎｅｎｓｉｓ、Ａｌｌｏｐｈａｉｏｍｙｓｃｆ．ｐｌｉｏｃａｅｎｉｃｕｓ、Ｐｒｏｅｄｒｏｍｙｓｃｆ．ｂｅｄｆｏｒｄｉ、Ｃｈａｒｄｉｎｏｍｙｓｎｉｈｅｗａｎｉｃｕｓ、Ｍｕｓｃｆ．ｍｕｓｃｕｌｕｓ和Ａｐｏｄｅｍｕｓｄｏｍｉｎａｎｓ的地史分布。根据哺乳动物群的分析与对比，得出第１８地点为早更新世早期，第１２地点、东洞地点为早更新世中期，第９地点、西洞地点为早更新世晚期，第１３地点为早更新世末期至中更新世初期，第１、２、５、６、１１、１０、２１、２３及上店洞地点为中更新世，新洞为中更新世晚期至晚更新世早期，东岭子洞为晚更新世，山顶洞为晚更新世晚期。还讨论了周口店地区一些主要第四纪哺乳动物群的生态类型特点，第四纪的气候波动及生态环境变迁。     

Proneofiber, a new genus of vole (Cricetidae: Rodentia) from the Pleistocene Seymour Formation of Texas,and its evolutionary and stratigraphic significance

A new genus and species of a microtine rodent, Proneofiber guildayi, is named on the basis of rooted teeth with cement from the warm Gilliland local fauna, Seymour Formation of Pleistocene age, Knox County, Texas. Proneofiber guildayi is considered to be ancestral to Neofiber, the round-tailed water rat, now living in the extreme southeastern United States. The presence of Pleistocene faunas prior to the first continental glaciation and the existence of three Pearlette-like volcanic ashes in the Plains Region suggest a revision of the age assignment of Pleistocene deposits from the nonglaciated region.     

流入型洞穴——山东沂源九天洞初步研究

九天洞是一个典型的直接由外源水补给形成的流入型洞穴。洞内大片分布的石花起源于四大古水池沉积物的存在。九天洞发育于第四纪早更新世晚期至中更新世早期,经历过成洞期、残余水流(古水池)活动期(中更新世中、晚期)石花发育期(晚更新世以来)和主要滴石、流石(含浅水池)沉积期(晚更新世晚期至今)等发育阶段。今日的九天洞又是一个残余洞穴,已被完全蚀去的近流入口段的洞穴长度不小于200m,约占洞穴全长的37%。九天洞及其洞穴群的景观资源价值和科学价值,在我国北方喀斯特洞穴中占有重要的地位。      

秦岭中更新世以来抬升的新资料及认识

秦岭北坡的陕西省洛南县南洛河北岸发育高度不同的6层溶蚀洞穴,其中5层溶蚀洞穴中发现沉积物和古生物化石。对这些洞穴进行了考察和发掘,并研究了洞穴内沉积物的化石。另外对洞穴外的河流阶地进行了高程测量和对比分析,用选频光释光法测定各洞穴底部沉积物和洞穴外河流阶地沉积物的年龄,确定了各洞穴大致的形成时代,并首次得到秦岭中更新世以来的抬升资料。其抬升过程中速度不均一性、时代越新抬升速率越强的总趋势等状况与青藏高原相似     

The northernmost record of Catagonus stenocephalus (Lund in Reinhardt, 1880) (Mammalia,Cetartiodactyla) and its palaeoenvironmental and palaeobiogeographical significance

During fieldwork carried out in January 2009 at Aurora do Tocantins (Tocantins State, northern Brazil), we recovered a fragmentary right maxilla (UNIRIO-PM 1006) of Catagonus stenocephalus from a sedimentary deposit of presumed late Pleistocene age in a karstic cave. This paper aims to: (1) provide the first record of C. stenocephalus in the northern region of Brazil (and consequently, also the northernmost one); (2) update the geographic distribution of C. stenocephalus; (3) present a date for the specimen; and (4) discuss the palaeoenvironmental and palaeobiogeographical implications of the finding. The species C. stenocephalus (Lund) is known from the Bonaerian (middle Pleistocene) and Lujanian (late Pleistocene to earliest Holocene) ages in Argentina, Uruguay, Brazil and Bolivia. The new record presented here extends the geographical distribution of C. stenocephalus more than 1000 km north from the former northernmost record (caves of Lagoa Santa region). Peccaries of the genus Catagonus have several morphological features associated with cursorial habits in relatively open and dry environments. The new distributional range of C. stenocephalus is coincident with the Chacoan subregion, characterized by dry climates and open areas. As the studied material comes from the top of the carbonate layer, this may suggest that the deposition of the C. stenocephalus remains described here is synchronous with the onset of a wetter climate phase. This argument is also in accordance with the datation results, around 20 ky BP, just after the last glacial maximum. This increasingly wet climate, which may also be related to the climatic changes that occurred during the late Pleistocene/early Holocene, could be a factor in the extinction of C. stenocephalus in South America.     

Paleoenvironmental implications of the Quaternary distribution of the eastern chipmunk (Tamias striatus) in central Texas

Four Quaternary cave sites in central Texas demonstrate that the eastern chipmunk (Tamias striatus) ranged more than 600 km southwest of its modern distribution. Climatographs suggest that the late Pleistocene/early Holocene summer climates were either 7.5°C cooler and 120 mm moister than today or 300 mm moister, if temperature remained unchanged. The distribution of T. striatus also implies that mixed deciduous forest existed on the eastern Edwards Plateau at this time.