Panthera Leo Spelaea Research Articles

Upper Pleistocene Panthera leo spelaea (Goldfuss, 1810) remains from the Bilstein Caves (Sauerland Karst) and contribution to the steppe lion taphonomy, palaeobiology and sexual dimorphism

Remains of the steppe lion Panthera leo spelaea (Goldfuss) from historical digs in the Bilstein Caves of Warstein (Sauerland, NW Germany) are described. Their age seems to be from the Early Weichselian periods (Upper Pleistocene). Whereas the Bilstein cave was inhabited by cave bears at that time only a few hyena prey remains, were most likely imported into the cave entrance by hyenas. Bite and crush marks on a few bones of Bison priscus, Bos primigenius, Cervus elaphus, a rhinoceros Coelodonta antiquitatis vertebra and even several chewed cave bear bones prove the hyena presence which is similar to other caves in the Sauerland hyena den cave rich region. Additionally some larger wolves subspecies Canis lupus spelaeus bones were found, but only few Crocuta crocuta spelaea remains are present. After taphonomic comparisons to six other hyena and cave bear den caves of northern Germany, this cave can be classified as a cave bear den, which was briefly used by hyenas only for food storage or commuting or cave bear predation site in one part of the Cave. The lion material refers at least to one young adult lioness, one more adult female and two male lions; therefore, at minimum, the remains of four adult individuals are represented. The absence of juvenile lion material, in contrast to cave bear cub remains in the Bilstein Caves, proves that P. leo spelaea did not use this and all other caves in the region to raise their cubs. The bone material from the Bilstein Caves would prove the same hyena-lion antagonism conflict being recently proven for the Perick Caves, Balve Cave or Martins Cave well. Other situations in caves such as the Keppler Cave and the Bilstein Cave initially show the more complex taphonomic situation of lion remains in European caves, especially in cave bear dens, where they seem to have hunted periodically cave bears, such as it is already proven for hyenas in the Sauerland Karst and other caves of Europe.

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.

A lion’s share of attention: Archaeozoology and the historical record

This is a review worth starting with Aesop (?620–564 BC). Eight classical Greek texts written in the Age of Homer (8–7th c. BC) reveal that lion was most frequently mentioned among wild animals, surpassed only by the six common domesticates that have directly surrounded people.1 References to this beast far outnumbered those to comparably large, dangerous and highly valued carnivores (Fig. 1). This is in sharp contrast with the relative underrepresentation of lion bones in archaeozoological assemblages. As A. Meyer noted with immense foresight over a century ago, “From the philological side there is evidently no basis to doubt... that lions indeed lived in Europe... around 500 BC... [Although] until recently no lion bones have been found in Greece, one should consider how few excavations took place in those lands”.2 Only a quarter of a century later, ecavations at the 5th–2nd century BC site of Ol’bia, a Greek colony on the Black Sea coast in modern-day Ukraine indeed yielded the fi rst bones of large Felids,3 attributed at the time to animals kept in a menagerie or the import of precious pelts.4 Although these possibilities must always be taken into consideration, reconfi rming Meyer’s optimistic view large faunal assemblages recovered in the Balkans during the last three decades have produced increasing numbers of lion remains from prehistoric as well as early historical times. This study seeks reconciling the remianing discrepancies between the historical record and archaeozoological evidence. As for the zoological aspect of this work, there are fundamental diffi culties in distinguishing between the bones of Late-Glacial and Holocene lions in Europe. Morphometric differences between the Pleistocene cave lion (Panthera leo spelaea Goldfuss, 1810) and the recent lion (Panthera leo Linnaeus, 1758) are negligible. While the bones of cave lion tend to be larger, precise identifi cation is often impossible using only sporadically occurring remains. Ancient DNA sequences indicate that cave lion from Austria and Southern Germany represent lineages that diverged from lions in Africa and Asia about 600,000 BP, and became extinct without mitochondrial descendants in those two continents. Depending on convention, even this ancient, distant form, however, may fi t within a single species nomenclature.5 Modern lions are closely related. Craniological traits of the now extinct North African Barbary lion (Panthera leo leo Linnaeus, 1758) and Asiatic lions (Panthera leo persica Meyer, 1826) are very similar. There must have been a contiguous population inhabiting North Africa and Asia. Moreover, the population in South-East Europe possibly once linked lions in the Middle East and North Africa, although the European form

Cave lion (Panthera spelaea) from the Pleistocene of Chuvashiya, European Russia

Cave lion (Panthera spelaea) from the Pleistocene of Chuvashiya, European Russia

The holotypes of the upper PleistoceneCrocuta crocuta spelaea(Goldfuss, 1823: Hyaenidae) andPanthera leo spelaea(Goldfuss, 1810: Felidae) of the Zoolithen Cave hyena den (South Germany) and their palaeo-ecological interpretation

The rediscovered holotype skulls of Late Pleistocene Panthera leo spelaea (Goldfuss, 1810) (Felidae) and Crocuta crocuta spelaea (Goldfuss, 1823) (Hyaenidae) from the Zoolithen Cave at Burggeilenreuth, southern Germany, are discussed. The cave became famous mainly due to its rich cave bear bone remains from the late Saalian (OIS 6–8) to Eemian/Weichselian (OIS 3–6) including additionally a third holotype of Ursus spelaeus Rosenmuller, 1794 (Ursidae). The ‘Felis spelaea’ holotype represents an adult male with a strong bite mark on the saggital crest, which was in an early stage of healing. Compared with other European Late Pleistocene lion skulls and skeletons, and with modern African lions, it provides evidence of intraspecific conflict between male Ice Age lions. The holotype of ‘Hyaena spelaea’ is one of several hundred hyena remains from a well-frequented hyena den cave. The cave was used intensively by Late Pleistocene hyena clans, for collecting lion carcasses in addition to their accustomed prey, as happened in many caves throughout Europe. Ice Age spotted hyena clans might have killed Ice Age steppe lions for many reasons, such as fights over prey and territory, and the protection of cubs, but they did not always scavenge on their carcasses. © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 154, 822–831.

Upper Pleistocene Panthera leo spelaea (Goldfuss, 1810) skeleton remains from Praha-Podbaba and other lion finds from loess and river terrace sites in Central Bohemia (Czech Republic)

Upper Pleistocene Panthera leo spelaea (Goldfuss, 1810) skeleton remains from Praha-Podbaba and other lion finds from loess and river terrace sites in Central Bohemia (Czech Republic)

Upper Pleistocene Panthera leo spelaea (Goldfuss, 1810) skeleton remains from Praha-Podbaba and other lion finds from loess and river terrace sites in Central Bohemia (Czech Republic)

Upper Pleistocene Panthera leo spelaea (Goldfuss, 1810) skeleton remains from Praha-Podbaba and other lion finds from loess and river terrace sites in Central Bohemia (Czech Republic)

SYSTEMATIC POSITION OF A COMPLETE LION-LIKE CAT SKULL FROM THE EEMIAN OSSIFEROUS RUBBLE NEAR ZANDOBBIO (BERGAMO, NORTH ITALY)

Morphologic and morphometric data of a lion-like cat skull found in the Zandobbio (Lombardy -Italy) Eemian ossiferous rubble and stored in the Civic Museum of Natural History E. Caffi of Bergamo are presented. The skull shows the typical lion morphology and its relatively small dimensions suggest that it belonged to a female individual. Carnassial tooth analysis underlines advanced lion characters already recognized in Italy during the Eemian. According to skull and teeth characters it is possible to ascribe the specimen to the group of Upper Pleistocene lion-like cats Panthera leo spelaea (Goldfuss, 1810). The presence of P. leo spelaea in the Quaternary deposit of Zandobbio has remarkable importance not only locally. In fact, besides being the first report of this great feline at Zandobbio, it is the second report from Lombardy and, the oldest certain P. leo spelaea finding and the oldest complete P. leo spelaea skull known from Italy to date. The advanced P. leo spelaea characters were already present in Italy during the Eemian. SHORT NOTE

Late Pleistocene and Holocene development of the felid fauna (Felidae) of Europe: a review

AbstractIn the fossil deposits of the Late Pleistocene (c. 115 000–11 500 years ago), five felid species are recorded in Europe: the wildcat Felis silvestris, Eurasian lynx Lynx lynx, Iberian lynx Lynx pardinus, leopard Panthera pardus and cave lion Panthera leo spelaea. In the Holocene, Europe was colonized by F. silvestris, L. lynx and L. pardinus as well as the lion Panthera leo. The status of P. pardus in post‐glacial Europe is unknown. So far, only sparse records indicate that P. pardus survived into the early Holocene. During the Late Glacial, both L. lynx and L. pardinus occurred on the Iberian Peninsula. However, from the Holocene, only the Iberian lynx is recorded in this region. There are subfossil records that indicate that L. pardinus also occurred in central and western France until c. 3000 years ago. Surprisingly, with reservations on the determination of the bones (by J. Altuna), both lions and cave lions seem to be recorded in the Iberian Peninsula in the Late Glacial. There are published records of the lion P. leo in the northern Iberian Peninsula from the early Holocene. However, its presence in Europe on the basis of subfossil records was proven initially from the Atlantic period. In Ponto‐Mediterranean regions of Europe, the lion is recorded from the Atlantic to the younger sub‐Atlantic.

Systematic position of the cave lion Panthera spelaea (Goldfuss) based on cranial and dental characters

Well-preserved skulls of Late Pleistocene Panthera spelaea are described from two geographically distant regions of Russia situated in the Russian Plain and Northern Siberia. The resemblance of the studied material with contemporaneous fossil lions from Alaska is established. The cranial morphology of P. spelaea confirms its phylogenetic position within the lion group, whereas many of the characters supporting the cave lion's relationship with the tiger are primitive. Both living and cave lions are advanced relative to the Middle Pleistocene P. fossilis in having larger upper incisors and more inflated bullae. P. spelaea also possesses the following advanced characters in common with the living lion: widened muzzle in the canine and P2 areas, wide nasal bones, relatively short pre-orbital part of the skull, laterally widened mastoid area, V-shaped form of the posterior tip of the frontal process of maxillary bone, and reduced cusp on the protocone bulge of P4. These conditions establish P. spelaea as a distinct species.

Molecular phylogeny of the extinct cave lion Panthera leo spelaea

To reconstruct the phylogenetic position of the extinct cave lion ( Panthera leo spelaea), we sequenced 1 kb of the mitochondrial cytochrome b gene from two Pleistocene cave lion DNA samples (47 and 32 ky B.P.). Phylogenetic analysis shows that the ancient sequences form a clade that is most closely related to the extant lions from Africa and Asia; at the same time, cave lions appear to be highly distinct from their living relatives. Our data show that these cave lion sequences represent lineages that were isolated from lions in Africa and Asia since their dispersal over Europe about 600 ky B.P., as they are not found among our sample of extant populations. The cave lion lineages presented here went extinct without mitochondrial descendants on other continents. The high sequence divergence in the cytochrome b gene between cave and modern lions is notable.

Cranial pathology ofUrsus spelaeus ROSENMÜLLER & HEINROTH from Chateau Pignon, Basque Territories (Spain)

A complex lesion, on a skull of a female cave bear (Ursus spelaeus) associated with perforations of the frontal sinus is described. Paleopathological analysis suggests that the lesion is attributable to complex cranial trauma, probably complicated by an infection which appears to have occurred following a fight, most likely with another bear or, possibly, a cave lion (Panthera leo spelaea); at least one of the perforations observed on the skull may have been produced by the canine of a large carnivore. This hypothesis is supported by the frequent intraspecific aggressiveness documented in modern bears, and by evidence suggesting interspecific fights between cave bears and cave lions. The case is discussed in the light of other evidence reported in the literature. © 1998 John Wiley & Sons, Ltd.

Cranial pathology of Ursus spelaeus Rosenmller & Heinroth from Chateau Pignon, Basque Territories (Spain)

A complex lesion, on a skull of a female cave bear (Ursus spelaeus) associated with perforations of the frontal sinus is described. Paleopathological analysis suggests that the lesion is attributable to complex cranial trauma, probably complicated by an infection which appears to have occurred following a fight, most likely with another bear or, possibly, a cave lion (Panthera leo spelaea); at least one of the perforations observed on the skull may have been produced by the canine of a large carnivore. This hypothesis is supported by the frequent intraspecific aggressiveness documented in modern bears, and by evidence suggesting interspecific fights between cave bears and cave lions. The case is discussed in the light of other evidence reported in the literature. © 1998 John Wiley & Sons, Ltd.

Panthera leo fossilis (v. Reich., 1906) (Felidae) de Isernia la Pineta (Pléistocène moyen inférieur d'Italie)

An upper carnassial of lion is been found at Isernia la Pineta (Molise, Southern Italy), a prehistoric site of the early Middle Pleistocene. The tooth is been compared to eight P/4 of Panthera leo spelaea of late Middle or Upper Pleistocene deposits. Morphological differences come out, once noticed by G. Schütt (1969) in the subspecies Panthera leo fossilis (v. Reichenau, 1906). It is assumed that the specimen of Isernia belongs to this subspecies and, moreover, that the area of distribution of this ancient lion extended from the Northern Europe to South Italy, in the Mediterranean belt.

Pleistocene remains of the lion-like cat (Panthera atrox) from the Yukon Territory and northern Alaska

Skull and forelimb fragments of the large, extinct cat (Panthera atrox) from Pleistocene sediments in the Dawson area, Yukon Territory, are the first records of the species for Canada. A further specimen from the Kaolak River, near the arctic coast of Alaska, significantly extends the known northward range of this cat. The species has been reported from approximately 26 localities from Alaska to Peru, and may have migrated from Eurasia just prior to the Sangamon interglacial. Data reviewed suggest that Panthera atrox from North America, the Eurasian "cave lion" Panthera leo spelaea, and the large Chinese cat Panthera youngi are conspecific.

INTERGLACIAL DEPOSITS AT TRAFALGAR SQUARE, LONDON

SummaryFossiliferous gravels were discovered during recent building operations on the south side of Trafalgar Square. At a later date organic deposits were exposed and an investigation of their plant remains was undertaken by the writer.A similar organic deposit had previously been described from the site of the Admiralty Offices (Abbott, 1892). In that work the following plants were recorded: Amblystegium ripanum Schimp., Hypninn stramineum Dicks., H. fluitans Hedw., Potamogeton sp., Lemna cf. polyrhiza, Ceratophyllum demersnm L., Polygonum sp., Rumex sp., Carex sp., Scirpus sp., and Betula nana L. With the exception of B. nana all the vascular plants recorded above have been also recognized in the present study. It seems probable that the record of B. nana was due to mis‐identification, since the general aspect of the flora is not one of cold or open conditions.The mammalian and molluscan remains from the present site have been studied by Dr. A. J. Sutcliffe and Mr. M. P. Kerney and will be published in detail later. The list of mammals forms an interesting counterpart to the plant determinations and is as follows: Panthera spelaea (Goldfuss), cave lion; Rhinoceros sp. indet., rhinoceros; Palaeoloxodon antiquas (Falconer), straight‐tusked elephant; Bos primigenius Bojanus, wild ox; Hippopotamus amphibius Linn., hippopotamus; Cervus elephas Linn., red deer; Dama cf. dama (Linn.), fallow deer. In addition a coprolite suggests that hyaenas may have been present and remains of bears not represented at this site have been found on two previous occasions during nearby excavations in other parts of the same terrace.