Cave Lion Research Articles

The first radiocarbon and stable isotope data for megafauna remains in Kaliningrad Province, Russia

ABSTRACTThe first 14C dates for representatives of the Late Pleistocene – Early Holocene megafauna are reported from sites in Kaliningrad Province, Russia. The dates for woolly mammoth and cave lion are older than ca. 45 000 bp, and could have corresponded to the Middle Weichselian interglacial (Marine Isotope Stage 3). The age of reindeer is ca. 10 000 bp, and of aurochs ca. 7000 bp. This shows that megafauna existed in this part of the Eastern Baltic region prior to the Last Glacial Maximum (LGM). After the LGM, Kaliningrad Province was recolonized by reindeer.

Habitation conditions of the cave lion in the middle Indigirka River basin, North-East Siberia, during the Middle and Late Pleniglacial: Reconstruction based on palynological data

Pollen analyses of the sediment from skulls cavities of two cave lions Panthera spelaea from the middle Indigirka River basin, North-East Siberia, were performed to reconstruct the habitat conditions of the animals and to estimate changes in vegetation and climatic conditions in the region. Composition of the pollen and non-pollen palynomorphs indicates that the predators most likely died on the river floodplain or near the shore of the thermokarst lake, where grass/sedge meadows and shrub thickets, providing rich grazing grounds for large herbivores, existed. Palynological data indicate that both in the early Middle Pleniglacial and in the LGM the regional vegetation included forest, shrub tundra, meadow, mire and steppe communities similar to modern cold steppes in central Yakutia. During early Marine Isotope Stage (MIS) 3, the participation of arboreal vegetation was greater than in the Last Glacial Maximum (LGM) due to higher summer temperatures. In MIS 2, the role of open herbaceous cryoxerotic communities was greater, which is associated with a higher continentality of climate and a decrease in winter temperatures. This reconstruction is consistent with the general sequence of landscape and climatic changes in the Arctic and Subarctic regions of Siberia in the Middle and Late Pleniglacial, based on paleobotanical studies of the locations of mammoth fauna and multidisciplinary studies of the Late Pleniglacial sediments from a broader area.

About the origin and environment of Panthera spelaea (Goldfuss, 1810)

Current phylogenies focus on the phyletic relationships between sister taxa of species, but none point to an ancestral species that gave rise to Panthera spelaea. The knowledge of its origins is progressing, but the question remains of whether this ancestral form is Eurasian or African, and at what period in the Plio-Pleistocene the divergence may have taken place. In this collective work on Panthera spelaea, the Eurasian species par excellence, the question of whether Panthera gombaszogensis may have evolved into Panthera spelaea is once again raised and examined in the light of current data. In the absence of decisive new data, it may be useful to reverse the problem and start from hypotheses. The environment plays a fundamental role in the evolution of species, and the hypothesis adopted must necessarily take this into account. Panthera gombaszogensis morphology, extensive distribution, and great adaptability to a variety of ecosystems suggest that it is a potential Eurasian ancestor of the cave lion. Chronology, whose role remains decisive, makes this possible.

Comparative analysis of hair microstructure in the cave lion (Panthera spelaea): A review

Original studies of the fur of adult males and heterosexual cubs of the cave and African lions and generalization of literature data showed that the hair colour of the cave lion was similar, but not identical to that of the African lion. The fur of Panthera spelaea is distinguished by the presence of a thicker and denser undercoat, consisting of numerous tightly closed and compressed wavy downy hairs with a medulla that is not usually characteristic of the undercoat of most mammalian species. The microstructure of the medulla, cortex, and cuticle patterns of the cave and African lions are similar, but the cuticle scales along the stem are higher in the cave lion. In lion cubs of both species, the differences are most pronounced, but in case of them, the fur is not as polymorphic as in adults. The differences between the fur of cave and African lions may be related to the adaptation of the first species to the conditions of the ice age, and the second to a hot climate. Comparison of the hair microstructure of the cave lion with other pantherines and felines revealed a similar specific structure of the medulla and cuticle patterns at the base of the guard hairs in many of the compared species, which confirms the systematic closeness of these taxa.

Remains of cave hyenas Crocuta spelaea (Hyaenidae) from Late Pleistocene cave and alluvial localities in southern Western Siberia

The results of studying 562 bone remains of Crocuta spelaea (Goldfuss, 1823) from 12 cave sites in Khakassia and Altai and 4 alluvial sites in the upper Ob River are presented. In the Fanatics cave in Khakassia, 3 almost complete skeletons of cave hyenas and one partially preserved were found. In this trap cave, 37–42 thousand years ago, hyenas died falling into a deep karst well. Morphometric analysis of the skulls and bones of the postcranial skeleton showed that the cave hyenas of the Late Pleistocene of southern Western Siberia did not differ in body size from the hyenas of Western Europe. In the alluvial localities of the Upper Ob region, the remains of C. spelaea are extremely rare, averaging 0.07% of the number of megafauna remains, which is several times less than that of the cave lion. In the Altai and Khakassia mountains, the relative abundance of cave hyenas may not have been inferior to that of Panthera leo spelaea (Goldfuss, 1810). During the glaciation periods, when cold and arid mountain-steppe landscapes dominated here, inhabited by diverse and numerous megafauna, the abundance of C. spelaea was the highest. During the interglacial period, against the background of widespread forest vegetation, the abundance of herd-hoofed animals decreased, which led to a several-fold decrease in the number of cave hyena populations. The most recent remains of C. spelaea from the Chik River are dated to the age of 33.4 thousand years ago.

Distribution history of the cave lion (Panthera spelaea (Goldfuss, 1810))

The paper provides an overview of the distribution of the cave lion (Panthera spelaea) in Eurasia and North America, spanning the period from the first documented appearance of the species in the fossil record at the end of the Early Pleistocene to its extinction at the end of the Late Pleistocene. A summary of the paleontological materials obtained over the last 50 years, after N.K. Vereshchagin review, is presented. The database utilised in this study encompasses 1044 palaeontological sites and 1506 localities. The hypotheses regarding the migrations (dispersal) of the species, including the migration from Asia to North America, and the timing of extinction in different parts of the range are discussed. The paper presents a synthesis of current research into the distribution of the cave lion, establishing a foundation for further investigation of this distinctive representative of the Pleistocene fauna of the Holarctic.

From giant to dwarf: A trend of decreasing size in Panthera spelaea (Goldfuss, 1810) and its likely implications

The Eurasian fossil record of Panthera spelaea shows a gradual decrease in body size. The process peaking just before the extinction of the species at the end of the Late Pleistocene. The temporal trend of decreasing size particularly accelerating during MIS 2, as exemplified by the extremely small female skull from Kryshtaleva Cave. Some palaeoecological aspects of the evolutionary history and eventual extinction of Panthera spelaea are discussed. Special discussion is given on how reducing body size affects potential interactions with other carnivores. It is also an attempt to answer the question whether, did indeed this trend negatively impact the cave lion ability to survive. The conclusions were drawn on the basis of correlation with observations of contemporary related forms.

A history of research on the cave lion Panthera spelaea (Goldfuss, 1810)

In 1810, G.A. Goldfuss was the first to identify the bones of a cave lion discovered in the Zoolithenhöhle cave (Germany), naming it Felis spelaea. But it was prehistoric man who first described it in numerous parietal and movable representations. Since the beginning of the 19th century, scientific knowledge of the cave lion Panthera (Leo) spelaea has progressed steadily, establishing its leonine character. Numerous discoveries led to paleontological studies of deposits throughout Eurasia, often in caves and linked to prehistoric sites. In the 20th century, this research benefited from the emergence of new dating methods, the development of palynology and the isotopic biochemistry of collagen. As the species' range became better defined, it expanded to include the whole of Eurasia and even North America. Finally, the 21st century is marked by the development of paleogenetics and the multiplication of synthetic studies, in many countries, while benefiting from a diffusion largely facilitated by Internet exchanges and the use of English. Phylogeny of the Felidae and understanding of the origins of Panthera spelaea are progressing rapidly. Today's knowledge makes it possible to envisage a true paleobiology of this extinct species.

From Caves to the Savannah, the Mitogenome History of Modern Lions (Panthera leo) and Their Ancestors.

Lions (Panthera leo) play a crucial ecological role in shaping and maintaining fragile ecosystems within Africa. Conservation efforts should focus on genetic variability within wild populations when considering reintroduction attempts. We studied two groups of lions from two conservation sites located in Zambia and Zimbabwe to determine their genetic make-up, information that is usually unknown to the sites. In this study, we analysed 17 specimens for cytb and seven microsatellite markers to ascertain family relationships and genetic diversity previously obtained by observational studies. We then produced a standardised haplogroup phylogeny using all available entire mitogenomes, as well as calculating a revised molecular clock. The modern lion lineage diverged ~151 kya and was divided into two subspecies, both containing three distinct haplogroups. We confirm that Panthera leo persica is not a subspecies, but rather a haplogroup of the northern P.l. leo that exited Africa at least ~31 kya. The progenitor to all lions existed ~1.2 Mya, possibly in SE Africa, and later exited Africa and split into the two cave lion lineages ~175 kya. Species demography is correlated to major climactic events. We now have a detailed phylogeny of lion evolution and an idea of their conservation status given the threat of climate change.

The last cave lion of the late Upper Palaeolithic: The engraved feline of Grotta Romanelli (southern Italy)

On the occasion of the review of the portable art of Grotta Romanelli, a decorated stone with a feline figure was object of an interdisciplinary study. The analysis considered different approaches so to: characterise the stratigraphic setting of the finding, the rock support, look into the techniques used to decorate the stone, elaborate a graphic documentation (photographs, 3D models and tracings), relate the symbolic production with the environmental context, and consider the motifs into the wider late Upper Palaeolithic (LUP) art production.The work allows confirming that the represented subject corresponds to a Panthera spelaea, and fixing some issues concerning the variability of the decorating activity, which is in line with the graphic tradition of the European LUP. Style and formal variable features of the figure might have responded to specific social conventions or to single author's skills, tracing new investigation lines about the cultural behaviour and the decorating activity: from the collection of the raw material and the preliminary modelling of the support, to the different artistic techniques (engraving and painting), from the use of the object to the definition of possible local artistic variations and/or inspiration at large scale. Moreover, it questions the thematic aspect in relation to the local fauna and its influence in the symbolic production, highlighting the importance of this stone in the wider debate about the extinction of the cave lions. Indeed, the Romanelli lion may represent the last evidence of this animal in Europe.

First direct evidence of lion hunting and the early use of a lion pelt by Neanderthals

During the Upper Paleolithic, lions become an important theme in Paleolithic art and are more frequent in anthropogenic faunal assemblages. However, the relationship between hominins and lions in earlier periods is poorly known and primarily interpreted as interspecies competition. Here we present new evidence for Neanderthal-cave lion interactions during the Middle Paleolithic. We report new evidence of hunting lesions on the 48,000 years old cave lion skeleton found at Siegsdorf (Germany) that attest to the earliest direct instance of a large predator kill in human history. A comparative analysis of a partial puncture to a rib suggests that the fatal stab was delivered with a wooden thrusting spear. We also present the discovery of distal lion phalanges of at least 190,000 years old from Einhornhöhle (Germany), representing the earliest example of the use of cave lion skin by Neanderthals in Central Europe. Our study provides novel evidence on a new dimension of Neanderthal behavioral complexity.

Geographical, temporal variability and sexual size dimorphism of mandible in cave lion (Panthera spelaea) across Northern Eurasia

ABSTRACT The mandible size variability including sexual size dimorphism (SSD) of the cave lion (Panthera spelaea) in the Pleistocene of Northern Eurasia time has been studied basing on the univariate and multivariate statistical analyses of 76 specimens from 43 localities. It has been confirmed that SSD in adult P. spelaea, as well as in extant P. leo, is the dominant form of group variability of mandible and, to somewhat lesser extent, of variability of cheek teeth. The lowest degree of SSD has been found in a relatively small Beringian subspecies, P. spelaea vereshchagini. The geographical variability of the cave lion from Western to Eastern Eurasia was not a cline at least in the Late Pleistocene. Our results confirm a decrease in overall mandible size from the Middle Pleistocene to the Late Pleistocene over a large part of the species’ range. We speculate that evolutionary changes in mandible size, lower canines or cheek teeth may have proceeded at different rates, allometry and probable heterochrony. Our results have added a new evidence to support the subspecies status of the Beringian lion (P. spelaea vereshchagini), but they are insufficient for other decisions on cave lion taxonomy at the subspecies level.

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.

New Data on the Neopleistocene Mammal Fauna from the Region of the Natural Park “Lenskie Stolby” (Middle Lena River Basin, Yakutia)

The data on new finds of the remains of Mammoth fauna mammals in the basin of Middle Lena River (Yakutia, the area of natural park “Lenskiye Stolby (Lena Pillars)” and adjacent territories) is represented in the article. Based on these data, the species composition of the mammoth fauna list of large mammalian species from this region was specified. 11 species of large mammals were identified, among which, for the first time for this region, the presence of a cave lion was established. A series of new radiocarbon datings allowing to note that mass burials of the Pleistocene mammalian remains were mainly formed in this region during the Karginian interstadial (60 (55)–24 thousand years ago, MIS-3). The obtained radiocarbon dates and species composition are consistent with the data that the Karginian interstadial was the most favorable for the distribution of the mammoth fauna in the territory of Yakutia. The results of the studies show that in the Late Pleistocene, the ecological preferences of Ovis nivicola differed from the modern ones. For the first time, the distribution of Panthera spelaea is revealed in the region of the middle reaches of the Lena River.

НАХОДКА ИСКОПАЕМОГО ПЕЩЕРНОГО ЛЬВА PANTHERA SPELAEA НА ТЕРРИТОРИИ КАЗАХСТАНА

В статье рассмотрены вопросы обитания пещерного льва Panthera spelaea на территории Казахстана и сопредельных территорий в плейстоцене. Пещерный лев являлся одним из наиболее крупных представителей семейства кошачьих, остатки которого известны из отложений позднего неоплейстоцена в сопровождении представителей мамонтовой фауны. Анализ радиоуглеродных данных других находок пещерного льва указывает на диапазон его обитания от 50 000 до 14 000 лет назад. Описана находка плечевой кости пещерного льва (вымерший подвид львов, населявший в эпоху плейстоцена Европу и Сибирь) этого периода из местонахождения оз. Маралды (Павлодарская область).

The Quaternary lions of Ukraine and a trend of decreasing size in Panthera spelaea

The fossil record of the cave lion, Panthera spelaea, suggests a gradual decrease in body size, the process peaking just before the extinction of the species at the end of the Late Pleistocene. Such an evolutionary trend appears rather unusual for a large felid species and requires further investigation. This study reviews the cave lions of Ukraine, whose fossils are known from 46 localities dated from 800 kyr to 18–17 kyr ago, with a special emphasis on size changes through time. We describe several important finds including those of Panthera spelaea fossilis from Sambir, Panthera spelaea ssp. from Bilykh Stin Cave and Panthera spelaea spelaea from Kryshtaleva Cave. We make subspecific identifications of specimens from the region and focus on their size characteristics. Our analysis of Ukrainian cave lions agrees with the temporal trend of decreasing size, particularly accelerating during MIS 2, as exemplified by the extremely small female skull from Kryshtaleva Cave. We provide a direct AMS date for this specimen (22.0–21.5 cal kyr BP), which suggests that the Kryshtaleva lioness must have belonged to a Panthera spelaea spelaea population forced south by the spreading ice sheet. We discuss some palaeoecological aspects of the evolutionary history and eventual extinction of the cave lion. Finally, we review the subfossil records of the extant lion Panthera leo known from several Ukrainian sites archaeologically dated to 6.4–2.0 kyr BP. These finds most probably represent the Persian lion Panthera leo persica.

North-Eastern Caucasus in the Bakunian Stage (Middle Pleistocene): Paleogeographic Conditions and Archaeological Industries

Purpose. Complete review of current status of research in paleogeographic settings and evolution of the archaeological industries in the North-Eastern Caucasus during the Bakunian stage of the Caspian Sea Quaternary history.Results. The Bakunian stage is recognized to be a part of the Middle Pleistocene in the Quaternary chronostratigraphy of the Caspian Sea region and dated 0.8–0.4 Ma. In the early stages of the Bakunian major transformation in the composition of landscapes and animal communities occurred, being associated with general cooling resulting an increase in natural land zoning. Further progressive climate change towards cooling led to the disappearance of a large number of thermophilic species from the fauna and flora. This time in the North-Eastern Caucasus corresponds to the Tiraspol faunistic complex, and in the later stages a complex of cave predator fauna (cave lion, cave hyena, etc.) emerges in the region. Archaeological industries of that time (Darvagchay-1, Darvagchay-Zaliv-4, etc.) demonstrates the eradication of the small-tool tradition and the reorientation of stone tool processing towards systemic knapping. More diverse usage of raw materials is identified in the archaeological record. In primary technology, these assemblages are characterized by parallel unifacial knapping and the presence of the few radial/discoid cores. Tool kits include bifaces and choppers, with predominance of single-edged side-scrapers on flakes and pebbles, and denticulate, notched and spikelike tools. There are practically no pointes. Through time a significant percentage of the small-tool component remains, but its contribution gradually decreases. These industries should be considered as a local type in the stone tool production evolution, which is in the general Caucasian trend of the spread of Early Paleolithic industries with bifaces.Conclusion. The Early Paleolithic of Dagestan from the first half of the Middle Pleistocene is identified as a local variant of the final Early Paleolithic industries of the Caucasus, bringing together the Early Paleolithic small-tool industry with the Acheulian complexes of the western part of Eurasia and combining their individual elements.

Evidence of cave lion (Panthera spelaea) from Pleistocene Northeast China

ABSTRACT The cave lion (Panthera spelaea) was one of the most widespread carnivores in the Pleistocene. The species ranged throughout Eurasia and even into North America as a member of the Mammuthus-Coelodonta fauna. However, the species has not previously been reported from the Mammuthus-Coelodonta fauna from the Pleistocene assemblages of the Songhua River. This study presents evidence of two Panthera spelaea specimens from the Middle and Late Pleistocene Songhua River fossil assemblages (MIS 10–8 and MIS 5–3). A combination of lower population due to vegetation shifts in the region and misidentification of specimens as Panthera tigris may explain why Panthera spelaea has not previously been reported in the region. The Middle Pleistocene specimen displays traits regarded as primitive in European cave lions. The Late Pleistocene specimen displays a mixture of primitive and advanced traits. These traits indicate a morphological transition within the species and also a possible ancestral relationship to Panthera atrox.

Middle Pleistocene Steppe Lion Remains from Grotte de la Carrière (Têt Valley, Eastern Pyrenees)

Late Pleistocene cave lions are one of the most iconic species of Northern Hemisphere Quaternary taphocoenoses. Despite their often-scarce record in cave environments, their ubiquitous distribution across Eurasia and North America assemblages attests to their position as top ice-age predators. Nevertheless, the origins of these former large felids, their distribution during the Middle Pleistocene, and their paleoecology during co-existence with the scimitar-toothed cat Homotherium remain debated. Here we describe for the first time an abundant collection of large-sized and stout felid remains from the recently discovered site of Grotte de la Carrière in Eastern Pyrenees, with an estimated age corresponding to MIS 9. Our results highlight the larger size of Middle Pleistocene lions compared to Late Pleistocene ones as well as a trend of decreasing in size, which has been previously stated by other authors. Grotte de la Carrière steppe lions have similar morphological and biometrical parameters to those of other samples from MIS 11–9, being larger and stouter than younger latest Middle Pleistocene-Late Pleistocene forms and slightly smaller than older MIS 15–12 forms.

Stephen Bishop: The Celebrated Guide of Mammoth Cave

Stephen Bishop: The Celebrated Guide of Mammoth Cave Angelo I. George (bio) and Gary A. O’Dell (bio) An Extraordinary Man It was a pleasant afternoon in Kentucky in the spring of 1844.1 A few guests lounged on the veranda of the Mammoth Cave Hotel, some strolled about the grounds admiring the verdant scenery and clusters of wildflowers, while others consulted with their host about the evening menu or their plans for underground adventures. They were completely unaware that, deep in the earth below this tranquil setting, a life-and-death struggle was taking place. Here, where the Jordan River ran deep and cold through Mammoth Cave, a tiny group of terrified souls huddled together in neck-deep water in an [End Page 99] absolute and fearful darkness. Torches extinguished by the sudden plunge into the chilly waters, they were unable to discern the features of the nearest person. What began as a thrilling honeymoon adventure for two newlyweds and a pair of their closest friends had turned into a fight for survival when their boat capsized while traversing the subterranean river. They clung to each other and grasped at projections on the mud-slick walls of the passage, convinced of impending death.2 Earlier that day, the wedding party of four from Bowling Green entered the cave, led by African American guide Stephen Bishop. Subsequent events suggest that some in the party became quite inebriated after a rest break in a location near the termination of Cleave-land’s Avenue, also called the Dining Table or Cornelia’s Table. As part of the caving experience, the guides would take a basket full of food and alcoholic beverages to be served at this location. Numerous published accounts record that after the meal, visitors often became tipsy from “choice old sherry.” The wedding party may have been celebrating with alcoholic refreshment even prior to this point. Bishop led his charges into the lowest level of the cave where the Jordan River had to be traversed by boat for nearly a mile, filling the passage from wall to wall and nine feet deep in places. After disembarking at a side avenue and exploring, they came back to the river and boarded the boat for a return to the surface. According to one account, “some of the party, who were in high glee, got into a romp, and overturned the boat. Their lights were extinguished, their matches wet, the boat filled with water and sank immediately . . . [they were] up to their chins in water.”3 Despite the harrowing circumstances, Bishop maintained composure and calmed the frightened tourists struggling in the cold water. [End Page 100] During his independent explorations of the cave, he had on many occasions been forced to traverse passages while neck-deep in water. He knew the present situation was potentially quite perilous and that the visitors could drown if allowed to panic. He cautioned them to “remain perfectly still, for if they moved a single step they might get out of their depth in water, and swimming would not avail them, for they could not see where to swim to.” Moving about could plunge them unexpectedly into water over their heads. When the wedding party failed to return to the surface at the designated time, a search and rescue team from the hotel was dispatched. Mat Bransford, another Black guide, led the rescue party and located the missing tourists when he heard Stephen “cheering them, and directing his movements while swimming, by the sound of their voices, which were raised, one and all, in prayer and supplication for deliverance.” Everyone was returned safely.4 Stephen L. Bishop (1820–1857) was the foremost guide at Mammoth Cave during the antebellum period whose daring exploits, personal magnetism, and knowledgeable discourses on the science of caves brought him international celebrity. Newspaper accounts and guidebooks of the period heralded Bishop as the “Columbus of the Cave;” his enslaver, Franklin Gorin, described him as a “fine genius;” English journalist Thomas Gunn referred to him as the “Cave Lion;” and newspaperman Bayard Taylor reported that Bishop was the “ruler of his realm.” One tourist praised “Stephen the illustrious . . . guide [End Page 101] Click for...