Panthera Leo Spelaea Research Articles

Morphometric comparison of Panthera spelaea (Goldfuss, 1810) from Poland with the lion remains from Eurasia over the last 700 ka

The Pleistocene lion Panthera spelaea was not a homogenous taxon; it was represented by three chronoforms. The oldest one, from the Early and Middle Pleistocene, was P. s. fossilis, the youngest was the Late Pleistocene cave lion P. s. spelaea. The intermediate form between them was P. s. intermedia. The scarcity of stratigraphically old lion made it difficult to understand the evolution of this carnivore. Therefore, we revised the fossil material of the Pleistocene lion from Polish excavation sites. The analysis revealed the presence of P. s. fossilis in 7 sites: Tunel Wielki, Południowa, Wschodnia, Draby, Biśnik, Deszczowa and Wierzchowska Górna caves. The remains from these localities significantly differed in their average size and teeth proportions from the nominate younger chronoform, i.e. the cave lion. Using this material and all other available data, we compared the lion remains grouped into four palaeoclimate periods: MIS 17-12, MIS 11-9, MIS 8-6 and MIS 5-2. We found significant time-related trends in many morphometric features of dentition and limb bones. Between the first two periods, the lion became larger but from MIS 8, it gradually decreased in size. The teeth showed a much greater rate of changes than the metapodial bones, which in turn were characterised by more heterogeneous and mosaic evolution. The changes observed in the fossil material can result from in situ evolution of lion populations occupying Europe or immigration and replacement of the native forms by those from other regions, for example Asia.

Montane record of the late Pleistocene Panthera spelaea (Goldfuss, 1810) from the Západné Tatry Mountains (northern Slovakia)

ABSTRACT A minimum of four adult cave lions (Panthera spelaea) from the late Pleistocene (MIS 3) are reported from Medvedia Cave in the Západné Tatry Mountains, the mountainous region of northern Slovakia in central Europe. A well-preserved fully adult male skeleton is the altitudinally highest recorded skeleton of this species and belongs to one of the largest reported cave lions. This specimen and a previous reported cave lion female from the same site exemplify the evident sexual dimorphism in this species. The fossils document a cave lion range extension into mountain areas during the Last Glacial, most likely in search of hibernating cave bear prey. Although it is generally assumed that the social behavior pattern of cave lions was similar to that of extant lions, we hypothesize that, at least in mountain environments, a solitary or paired existence was more likely for these large extinct felids. SUPPLEMENTAL DATA—Supplemental materials are available for this article for free at www.tandfonline.com/UJVP Citation for this article: Sabol, M., J. Gullár, and J. Horvát. 2018. Montane record of the late Pleistocene Panthera spelaea (Goldfuss, 1810) from the Západné Tatry Mountains (northern Slovakia). Journal of Vertebrate Paleontology. DOI: 10.1080/02724634.2018.1467921.

Gigantic lion, Panthera leo, from the Pleistocene of Natodomeri, eastern Africa

AbstractThe partial skull of a lion from Natodomeri, northwest Kenya is described. The Natodomeri sites are correlated with Member I of the Kibish Formation, dated to between 195 ka and ca. 205 ka. The skull is remarkable for its very great size, equivalent to the largest cave lions (Panthera spelaea [Goldfuss, 1810]) of Pleistocene Eurasia and much larger than any previously known lion from Africa, living or fossil. We hypothesize that this individual represents a previously unknown population or subspecies of lion present in the late Middle and Late Pleistocene of eastern Africa rather than being an indication of climate-driven size increase in lions of that time. This raises questions regarding the extent of our understanding of the pattern and causes of lion evolution in the Late Pleistocene.

Under the Skin of a Lion: Unique Evidence of Upper Paleolithic Exploitation and Use of Cave Lion (Panthera spelaea) from the Lower Gallery of La Garma (Spain).

Pleistocene skinning and exploitation of carnivore furs have been previously inferred from archaeological evidence. Nevertheless, the evidence of skinning and fur processing tends to be weak and the interpretations are not strongly sustained by the archaeological record. In the present paper, we analyze unique evidence of patterned anthropic modification and skeletal representation of fossil remains of cave lion (Panthera spelaea) from the Lower Gallery of La Garma (Cantabria, Spain). This site is one of the few that provides Pleistocene examples of lion exploitation by humans. Our archaeozoological study suggests that lion-specialized pelt exploitation and use might have been related to ritual activities during the Middle Magdalenian period (ca. 14800 cal BC). Moreover, the specimens also represent the southernmost European and the latest evidence of cave lion exploitation in Iberia. Therefore, the study seeks to provide alternative explanations for lion extinction in Eurasia and argues for a role of hunting as a factor to take into account.

Mitogenomics of the Extinct Cave Lion, Panthera spelaea (Goldfuss, 1810), Resolve its Position within the Panthera Cats

The extinct cave lion ( Panthera spelaea ) was an apex predator of the Pleistocene, and one of the largest felid species ever to exist. We report the first mitochondrial genome sequences for this species, derived from two Beringian specimens, one of which has been radiocarbon dated to 29,860 ± 210 14C a BP. Phylogenetic analysis confirms the placement of the cave lion as the sister taxon to populations of the modern lion ( P. leo ). Using newly recovered stem pantherine fossils to calibrate a molecular clock, we estimate that P. spelaea and P. leo diverged about 1.89 million years ago (95% credibility interval: 1.23–2.93 million years), highlighting the likely position of this extinct carnivore as a distinct species.

Morphological and genetic identification and isotopic study of the hair of a cave lion (Panthera spelaea Goldfuss, 1810) from the Malyi Anyui River (Chukotka, Russia)

We present the first detailed analyses of the preserved hair of a cave lion (Panthera spelaea Goldfuss, 1810). The hair was found in association with a skeleton that was recovered recently from perennially frozen Pleistocene sediments in the lower reaches of the Malyi Anyui River (Chukotka, Russia). We extract mitochondrial DNA from the hair to confirm its taxonomic identity, and perform detailed morphological analyses of the color and structure of the hair using light optical microscopy and SEM. In addition, we compare the cave lion hair to hair taken from the back and mane of an African lion. We find that cave lion hair is similar but not identical to that of the present-day lion. In addition to slightly different coloration, cave lions had a very thick and dense undercoat comprising closed and compressed wavy downy hair with a medulla. In addition, while the microstructures of the medulla and cortex of cave lion hair are similar in extinct and living lions, the cuticular scales of cave lion hair are higher than those in living lions, suggesting that cave lion hair is stronger and more robust than that of living lions. We hypothesize that the differences between cave lion hair and present-day lion hair may be due to adaptations of cave lions to the harsh climatic and environmental conditions of the Pleistocene Ice Ages.

Small and large mammals from the Ciota Ciara cave (Borgosesia, Vercelli, Italy): An Isotope Stage 5 assemblage

The Ciota Ciara cave is located within the Monte Fenera karst system (Borgosesia, Vercelli, Italy) at 670m a.s.l. The cave entrance presents a deposit with Mousterian quartz and flint industry. The faunal remains from Stratigraphic Units 13, 14 and 103 are the subject of this work and are presented here as a whole for the first time. The large mammal assemblage is dominated by Ursus spelaeus. In addition, a few remains of carnivores such as Panthera leo spelaea, Panthera pardus, Meles meles have been found together with ungulates such as Cervus elaphus and Rupicapra rupicapra. The small mammal assemblage is characterized by a high biodiversity, especially in bats, by the dominance of Clethrionomys glareolus and by a relatively large number of Pliomys coronensis, a species that seems to disappear from the Italian Peninsula at the end of Marine Isotope Stage 5 or during the beginning of MIS 4. The changes in frequency of the small and large mammals between the two S.U. suggest a change from a relatively cold-humid (S.U. 14) to a warmer-still humid climate (S.U. 13). Although no radiometric dates are available yet, the small and large mammal assemblages, the presence of Pliomys coronensis and the climate change inferred by the variation of the small mammal frequencies allow us to correlate these two units of the Ciota Ciara cave to a relatively warm moment of Marine Isotope Stage 5.

On the discovery of a cave lion from the Malyi Anyui River (Chukotka, Russia)

An incomplete postcranial skeleton (67 elements) of a cave lion, a lower jaw and a bundle of fine yellowish hair were found by a local resident in 2008 and 2009 washed out from the perennially frozen Pleistocene sediments in the lower reaches of the Malyi Anyui River (western Chukotka). This is the first skeleton of a cave lion (Panthera spelaea Goldfuss) to be found in Russia. The bone sizes are similar to finds of cave lion bones known from N–E Russia, but larger than East Beringian and smaller than West European ones. The remains have been studied using a variety of methods, including morphology, morphometry, SEM-examination, AMS-dating, and isotopic study, which included examination of over 100 samples of various members of the mammoth faunal assemblage (mammoth, wooly rhinoceros, bison, horse, bear, etc.). The results showed that the northeastern Asian cave lion hunted mainly bison and horses, but not reindeer, unlike its Western Europe counterpart. Bone and claw sheath dating showed an unexpectedly old geochronological age of over 61,000 years (OZQ290, OZQ291), while the hair was dated 28,690 ± 130 (OZQ292), which makes its affinity with the same individual as the skeleton questionable. Further studies to investigate possible unremoved contamination and obtain more reliable date are planned.

Population Demography and Genetic Diversity in the Pleistocene Cave Lion

With a range that covered most of northern Eurasia and parts of North America, the cave lion (Panthera spelaea) was one of the most widespread carnivores of the Late Pleistocene. Earlier ancient DN ...

Large mammals from the Upper Neopleistocene reference sections in the Tunka rift valley, southwestern Baikal Region

This work presents the data on new finds of fossil macrotheriofauna in the reference sections of the Upper Neopleistocene sediments in the Tunka rift valley (southwestern Baikal Region). The osteological material of a number of Late Neopleistocene mammals including extinct species rare for the Baikal region such as Crocuta spelaea, Panthera spelaea, and Spirocerus kiakhtensis (?) was directly dated with a radiocarbon (AMS) method. The obtained 14C data (18000–35000 years) allow one to rejuvenate significantly the upper limit of the common age interval of habitat of these animals in southern part of Eastern Siberia. Cave hyena and spiral-horned antelope lived in the Tunka rift valley in the Baikal region in Late Kargino time (37–24 ka), and cave lion survived the maximum in the Sartan cryochron in the region (21–20 ka). The study of collected paleontological collections provides a basis for selection of independent Kargino (MIS 3) faunal assemblages to use them for regional biostratigraphic analysis of Pleistocene deposits. Radiocarbon age dating of samples allows one to attribute confidently all paleofaunal remains available to the second half of the Late Pleistocene.

The remarkable Panthera pardus (Felidae, Mammalia) record from Equi (Massa, Italy): taphonomy, morphology, and paleoecology

Leopard remains are rare in the European fossil record, probably a consequence of its solitary and elusive habits. Equi, dating back to Marine Isotope Stage 3 (MIS3), represents a rich and outstanding exception. Historical excavations (from 1911 to about 1917) at Equi allowed more than 200 remains to be recovered of this large felid, a sample attesting the richest leopard-bearing site in the Pleistocene of Europe. The Equi site testifies the survival of a prosperous leopard population in northwestern Italy (the Apuan Alps area) during the pre Last Glacial Maximum Late Pleistocene.Almost all skeletal elements are represented in the collection (except ribs, and most of axial skeleton). At Equi, leopard is represented by both young and adult animals and, up to now, it seems the first and unique Pleistocene evidence of cave usage for cubs raised in Europe.This study focuses on the whole record of leopard from Equi, housed in the Museum of Natural History of Florence, and is aimed to analyze the taphonomical assemblage, to provide an anatomical description of the remains and the comparison to the European evidences. Moreover, we infer the paleoecology of leopards, and their relationships with other large carnivores from the Equi guild (Ursus spelaeus, Canis lupus, and Panthera leo spelaea). The site was probably an occasional large carnivores den, used over time by different large carnivore taxa, perhaps in different seasons or times (mutual avoiding).Our morphological analysis allows us to recognize sexual dimorphic characters in the cranium, and a large morphological variability in body size.

Middle Pleistocene stratigraphy of the deposits in Za Hájovnou Cave (Javoříčko Karst, northern Moravia, Czech Republic)

Middle Pleistocene stratigraphy of the deposits in Za Hájovnou Cave (Javoříčko Karst, northern Moravia, Czech Republic)

First Asian record of Panthera (Leo) fossilis (Mammalia, Carnivora, Felidae) in the Early Pleistocene of Western Siberia, Russia

A lion-like pantherine felid is described as Panthera (Leo) fossilis from the late Early Pleistocene sediments of the Kuznetsk Basin (Western Siberia, Russia). The find of P. fossilis first recorded in Asia considerably extends the current notion of the eastward expansion of the most ancient lions. The Siberian lion is geologically the oldest form and is dimensionally among the largest members of the group of fossil lions on the Eurasian continent. Although known by mandibular remains only, it is readily distinguished from Panthera (Leo) spelaea by a heavy built mandibular corpus with rectangular profile in the cheek teeth area, a deep, well-outlined and narrow anterior section of the masseteric fossa, and a large p4 supported by a big unreduced anterior root. The Siberian lion shares these features with the European Middle Pleistocene P. fossilis and the American Late Pleistocene P. (Leo) atrox, which suggests their close relationship. P. atrox originated from P. fossilis and was isolated in North America south of the Late Pleistocene ice sheets. This explains why the American lion has retained more primitive features than the coeval Eurasian cave lion P. (L.) spelaea.

Palaeopopulations of Late Pleistocene Top Predators in Europe: Ice Age Spotted Hyenas and Steppe Lions in Battle and Competition about Prey

Late Pleistocene spotted hyenaCrocuta crocuta spelaea(Goldfuss, 1823) and steppe lionPanthera leo spelaea(Goldfuss, 1810) were top predators in Central Europe. The fossil record (2.303 hyena/1.373 lion bones = ratio 3/1) from 106 cave and open air sites demonstrates comparable associations to modern African hyenas/lions resulting in competition about prey and territory. Cannibalism within extinct spotted hyenas is well documented, including two individual skeletons. Those hyenas produced bone accumulations at dens. Feeding specializations on different megamammal groups are demonstrated for Late Pleistocene hyenas whose prey partly overlaps (e.g., cave bears) with those of lions and wolves. At most fossil sites, 1–3% of the lion remains indicate scavenging of lions by hyenas. The larger Late Pleistocene felids focussed on cervids (reindeers specialization during the high glacial = LGM), on bovids (steppe bison/aurochs), and possibly on saiga antelope and on the cave bear, hunting deep in caves during their hibernations and targeting cubs. The cave bear feeding was the target of all three top predators (lions, hyenas, and wolves) in the Late Pleistocene boreal forests which caused deathly conflicts in caves between them, especially with lions/hyenas and herbivorous cave bears that have no modern analogue.

Decreasing size process in the cave (Pleistocene) lion Panthera spelaea (Goldfuss, 1810) evolution – A review

This study analyzed size changes in the course of the time among European Middle and Late Pleistocene lions. The general decreasing size as a tendency in cave lion evolution is confirmed. The whole time span of lion presence in Europe was divided into four main periods. The first period covers Middle and late Middle Pleistocene (MIS 19-17 to MIS 7-6) and is characterized by the presence of very large individuals with somewhat primitive morphology. It might be cautiously said that lions in that time were comparatively large throughout the Middle and late Middle Pleistocene. The second period included the last part of the late Middle and the beginning of the Late Pleistocene (MIS 6-5). This period is documented by a mixed presence of large individuals with somewhat primitive features together with smaller, evolutionarily more advanced dentition lions. The third period covers the first part of the last glacial, from MIS 5 to the beginning of MIS 3. For this period, the occurrence of moderate sized specimens with evolutionarily advanced dentition is documented. The last period began with dramatic population decline and genetic variability reduction. It lasted from MIS 3 and 2, and within it size drops sharply. Among moderate sized specimens, very small lions with somewhat lesser posture also appeared. Asian or African affinities as a result of a new migration event for those dwarf lions could not be ruled out. Size decrease showed that Panthera spelaea was a dynamically evolved species. Thus, the size does not seem to be a reliable criterion in determining the age of cave lion findings. The general overview is complicated by the great sexual dimorphism, local evolution and extinction, climatic conditions, and migrations. Size changes in P. spelaea evolution cannot be used as a useful biochronological tool.

Environment during the Middle to Late Palaeolithic transition in southern France: The archaeological sequence of Tournal Cave (Bize-Minervois, France)

Late Pleistocene deposits of Tournal Cave, southwestern France, provided several human occupations attributed to Mousterian, Aurignacian, and Magdalenian cultures. Some human remains (Homo neanderthalensis and Homo sapiens) were identified in each unit. This study presents a detailed temporal reconstruction of habitats surrounding the cave, using various proxies based on rich large mammal assemblages and related to biological activities. The levels correlated to MIS 3 and MIS 2 are characterized by alternating carnivore, Neanderthal, and Anatomically Modern Human (AMH) occupations. Faunal exploitation was mainly oriented towards Equus caballus and Rangifer tarandus, with a significant increase in the latter prey from the last Aurignacian level onwards. The results exhibit geographic and temporal variations of reindeer and horse body size. Reindeer can be used as a suitable ecological marker, as rapid changes are correlated to environmental turnover, whereas the horse presents a different pattern and is an accurate chronological estimator. The palaeoecological results indicate a major climate change between the two Aurignacian levels. The Mousterian and the first Aurignacian levels, documenting the Middle to Late Palaeolithic transition, show high indices of specific richness and diversity of large mammals in relation with a periglacial moderate cold and wet climate. The region developed a mixed landscape and displayed a non-analogue fauna with E. caballus, R. tarandus, Cervus elaphus, Megaloceros giganteus, Bison priscus, Bos primigenius, Capra caucasica praepyrenaica, Sus scrofa and Coelodonta antiquitatis for the ungulates and Ursus spelaeus, Ursus arctos, Crocuta crocuta spelaea, Panthera leo spelaea, Panthera pardus, Lynx spelaeus, Canis lupus and Vulpes vulpes for the carnivores. The breakdown occurring in the last Aurignacian level is characterized by a change in ecological settings, with a colder and drier climate and opening of the landscape, involving shifts in seasonality, plant phenology, reindeer body size and animal population densities. This impacted habitat fragmentation and geographic distribution of populations, implying various selective pressures that were reflected in human meat procurement and dispersal events. At Tournal Cave, the late Middle/early Late Palaeolithic transition is not directly marked by significant differences in terms of faunal exploitation, due to local climate and site function, but the major environmental shift recorded is delayed. The results indicate that abrupt climate oscillations during MIS 3 contributed to the decline of Neanderthal populations, notably because of habitat fragmentation. AMHs could have generated additional stresses.

Late Pleistocene leopards across Europe – northernmost European German population, highest elevated records in the Swiss Alps, complete skeletons in the Bosnia Herzegowina Dinarids and comparison to the Ice Age cave art

European leopard sites in Europe demonstrate Early/Middle Pleistocene out of Africa lowland, and Late Pleistocene Asian alpine migrations being driven by climatic changes. Four different European Pleistocene subspecies are known. The final European Late Pleistocene “Ice Age leopard” Panthera pardus spelaea (Bächler, 1936) is validated taxonomically. The skull shows heavy signs of sexual dimorphism with closest cranial characters to the Caucasian Panthera pardus ciscaucasica (Persian leopard). Late Pleistocene leopards were distributed northernmost, up to S-England with the youngest stratigraphic records by skeletons and cave art in the MIS 2/3 (about 32,000–26,000 BP). The oldest leopard painting left by Late Palaeolithics (Aurignacians/Gravettians) in the Chauvet Cave (S-France) allows the reconstruction of the Ice Age leopard fur spot pattern being close to the snow or Caucasian leopards. The last Ice Age glacial leopard habitat was the mountain/alpine boreal forest (not mammoth steppe lowland), where those hunted even larger prey such as alpine game (Ibex, Chamois). Into some lairs, those imported their prey by short-term cave dwelling (e.g. Baumann's Cave, Harz Mountains, Germany). Only Eurasian Ice Age leopards specialized, similar as other Late Pleistocene large felids (steppe lions), on cave bear predation/scavenging partly very deep in caves. In Vjetrenica Cave (Dinarid Mountains, Bosnia Herzegovina), four adult leopards (two males/two females) of the MIS 3 were found about two km deep from the entrance in a cave bear den, near to one cave bear skeleton, that remained articulated in its nest. Leopards died there, partly being trapped by raising water levels of an active ponor stream, but seem to have been killed possibly either, similar as for lions known, in battles with cave bears in several cave bear den sites of Europe (e.g. Baumann's Cave, Wildkirchli Cave, Vjetrenica Cave). At other large cave sites, with overlap of hyena, wolf and dhole dens at the cave entrances, leopard bones with bite damages indicate their remains to have been imported and consumed by predators in alpine regions due to reduced prey availability. The best models for the competition/taphonomy of large predators – felids, hyenids, canids – within large cave bear dens of Europe is represented in combination of the Zoolithen Cave and Vjetrenica Cave taphonomy.

Late Pleistocene Eemian hyena and steppe lion feeding strategies on their largest prey—Palaeoloxodon antiquus Falconer and Cautley 1845 at the straight-tusked elephant graveyard and Neanderthal site Neumark-Nord Lake 1, Central Germany

The scavenging activities of hyenas and lions at the Neumark-Nord Lake 1 (Central Germany) site on straight tusk elephant carcasses is commonly documented by slight bone damages and initial skeleton partitioning. Canine bite marks are found at several joints of elephant long bones, as well as on the vertebrae and skulls. Deep bite and scratch marks from 5 to 7 mm in width made by large carnivore canine teeth (Panthera leo spelaea, Crocuta crocuta spelaea) on the ventral vertebral columns and pelvises of two Palaeoloxodon antiquus straight-tusked elephant skeletons suggest that the intestines and inner organs were consumed by both large lion/hyena predators as is commonly observed when modern African lions and spotted hyenas feed on contemporary elephant carcasses. A scavenging strategy can be demonstrated using 24 partly preserved Palaeoloxodon skeletons, where Ice Age spotted hyenas appear to be the main scavenger (95 %) and sole skeleton decompositor such as bone destructor. While Crocuta may feed anywhere on an elephant carcass, they are known to specialize to eat the complete feet, whereas scavenging from the anus into the body cavity (intestine/internal organ feeding) is more typical of lions. Shallow water at Neumark-Nord Lake 1 prevented hyenas from completely scavenging elephants; thus, bone damage at this site was constrained to initial stages of destruction and final carcass partitioning by high water stands and floods.

Were bears or lions involved in salmon accumulation in the Middle Palaeolithic of the Caucasus? An isotopic investigation in Kudaro 3 cave

Bone fragments of large anadromous salmon in the Middle Palaeolithic archaeological layers of Kudaro 3 cave (Caucasus) suggested fish consumption by archaic Hominins, such as Neandertals. However, large carnivores such as Asiatic cave bears (Ursus kudarensis) and cave lions (Panthera spelaea) were also found in the cave and could have been responsible for such an accumulation. The diet of these carnivores was evaluated using carbon, nitrogen and sulphur isotopes in faunal bone collagen. The results suggest that anadromous fish were neither part of the diet of either cave bear (vegetarian) or cave lion (predators of herbivores from arid areas) and therefore provide indirect support to the idea that Middle Palaeolithic Hominins, probably Neandertals, were able to consume fish when it was available.

The Janda cavity at Fruška Gora, the first cave assemblage from the southeast Pannonian lowland (Vojvodina, Serbia)

The recently discovered Janda cavity yielded the first large mammal fossil assemblage of the Late Pleistocene age in the southeast Pannonian lowland (Vojvodina), outside the karst region of Serbia. The cavity is formed in the Badenian reef limestone on the northern hillside of the Fruška Gora mountain. The paper provides a paleontological description and taphonomic data on mammal remains collected from the loose sediments or scree. Herbivores are represented by Bison priscus (Bojanus, 1827), which is the most abundant species, followed by Megaloceros giganteus (Blumenbach, 1803) and Equus germanicus Nehring, 1884, with several remains of Mammuthus sp. and Coelodonta antiquitatis (Blumenbach, 1799). Carnivores are represented by Ursus spelaeus ingressus (Rabeder et al., 2004), Crocuta crocuta spelaea (Goldfuss, 1823), Panthera leo spelaea (Goldfuss, 1810), as well as Canis lupus Linnaeus, 1758. Wolf remains are characterized by their large size, almost reaching the size of hyenas, which is in contrast with the finds of rather small Pleistocene wolves from the caves in the mountainous part of Serbia. The faunal composition and the presence of mostly grazing forms among the herbivores indicate glacial conditions and an open landscape in the surroundings of the site during the formation of the Janda cavity infill. Taphonomic observations indicate the cave hyena as a dominant bone accumulating agent. In contrast to other cave hyena sites in Serbia, where mostly small and medium sized prey remains were accumulated, prey preference for large and extra large mammals is noted. It is supposed that this is the consequence of the differences between mountainous and lowland landscapes and environments, climatic conditions, different composition of animal and plant associations, and also in different modes of competitions between predators.