European leopard | |
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Skeleton at the Museo di Storia Naturale di Firenze | |
Scientific classification ![]() | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Mammalia |
Order: | Carnivora |
Suborder: | Feliformia |
Family: | Felidae |
Subfamily: | Pantherinae |
Genus: | Panthera |
Species: | P. pardus |
Population: | †European leopard |
Leopards have a long history in Europe, spanning from the Early-Middle Pleistocene transition, around 1.2-0.6 million years ago, until the end of the Late Pleistocene, around 12,000 years ago, and possibly later into the early Holocene. Remains of leopards have been found across Europe, from the Iberian Peninsula to the Caucasus.
The proposed Late Pleistocene European leopard subspecies Panthera pardus spelaea was first described as Felis pardus spelaea by Emil Bächler in 1936. [1]
Several fossil bones from the Early, Middle and Late Pleistocene were described and have been proposed as different leopard subspecies:
Analysis of mitochondrial genomes from Late Pleistocene European leopard specimens found in Baumannshöhle in Germany, dating to around 40,000 years ago, found that they shared a mitochondrial genome lineage that diverged from the common ancestor of living Asian leopards around 500,000 years ago, leading the authors of the study to propose that European leopards represented a distinct population from living Asian leopards. [6] However, later analysis of two mitochondrial genomes obtained from sedimentary layers of El Miron cave in Spain (dating to approximately 46,890–33,160 and 22,980–22,240 calibrated years Before Present respectively) found that these sequences were each others closest relatives, but they did not group with those from Germany, and were instead within the living diversity of Asian leopard mitochondrial DNA sequences, and in particular shared a close common ancestry with a 35,000 year old leopard mitochondrial DNA sequence obtained from Mezmaiskaya cave in the northwest Caucasus. [7]
The skulls of Late Pleistocene European leopards are medium-long, and their characteristics are closest to the Panthera pardus tulliana subspecies. An apparent depiction of a leopard in the Chauvet Cave shows a coat pattern similar to that of modern leopards but with an unspotted belly, presumably white. Like other mammals, leopards from the cold glacial periods of the Late Pleistocene are usually larger than those from the warm interglacial phases. As in modern leopards, there was a strong sexual dimorphism, with males being larger than females. [8] Remains from Equi cave in Italy exhibit considerable size variabilty, but are generally large in comparison to modern leopards. The muzzles of skulls from this locality are generally shorter than those of living leopards. [9] While Dietrich (2013), argued that earlier Middle Pleistocene European leopards differed in a number of morphological features of the skull and teeth from Late Pleistocene European leopards, [8] other authors have argued that no such trends exist, and that this simply reflects intraspecific variability and that some Late Pleistocene European leopards have similar skull and teeth features to those of earlier Middle Pleistocene European leopards. [9]
The timing of arrival of leopards in Europe is disputed. Some authors have posited that they arrived in Europe during the late Early Pleistocene around 1.2-1.1 million years ago. [10] while others have suggested that they arrived during the early Middle Pleistocene, around 600,000 years ago. [6] While initially very rare, records of leopards become more common and widely distributed from the late Middle Pleistocene onwards, following the extinction of the "European jaguar" Panthera gombaszoegensis, [10] though their fossil record in Europe is still scarce overall. [11] Northwards, leopards ranged to Great Britain, but their records here are rare and only recorded during Marine Isotope Stage 7 ~225,000 years ago. [12] During the Late Pleistocene, their northern limit was around Berlin in northern Germany. [9]
During the Last Glacial Maximum, leopards persisted in relatively temperate glacial refugia in the Iberian, Italian and Balkan Peninsulas. [13] Bone fragments of P. p. spelaea have been excavated in Switzerland, Italy, Spain, Germany, Poland and Greece. [1] [9] [14] [15] [16] Leopard fossils dating to ~43,000 BP were found in the Radochowska Cave in Poland. [10] The most complete skeleton of P. p. spelaea is known from Vjetrenica Cave in southern Bosnia and Herzegovina, where four leopard fossils were found. These are dated to the end of the Late Pleistocene, about 29,000–37,000 years ago. A cave painting of a leopard in the Chauvet Cave in southern France is dated to about 25,000–37,500 years old. The last leopards vanished from most parts of Europe about 24,000 years ago, just before the Last Glacial Maximum. [8]
The cave site of Equi in northwestern Italy, dating to the Last Glacial Period (MIS 3, ~53-27,000 years ago) represents the richest concentration of leopard remains from Pleistocene Europe, with some 200 bones of leopards having been excavated from the locality, including 5 well preserved skulls. The remains of cubs found in the cave suggests that leopards used it to give birth and rest. [11] The youngest reliable records for leopards outside of eastern Europe are from the Iberian Peninsula, around 17-11,000 years ago, with records in the Iberian Peninsula possibly extending into the early Holocene, during the Mesolithic. [10] Modern (Asian-type) leopards are still found on the fringes of Europe in the North Caucasus. [17]
Fossils of leopards in Europe are sometimes found in caves, where they apparently sought shelter or hid their prey. They generally preferred smaller caves, most likely because larger caves were usually occupied by larger predators such as cave bears, cave lions (P. spelaea), or humans. In European Ice Age caves, leopard bones are far rarer than those of lions, and all currently known fossils belong to adults, suggesting that they rarely, if ever, raised their cubs in caves. Where leopard remains are found in larger caves, they are often found in the cave's deeper recesses, as in Baumann's and Zoolithen Cave in Germany. It is not precisely known which prey species these leopards hunted, although they may have been similar to modern snow leopards, which prey on ibex, deer and wild boar. It is likely that leopards scavenged or occasionally killed cave bears during hibernation in their dens. During the cold phases, European leopards occurred mainly in mountain or alpine boreal forests or in mountains above the treeline, and were not usually found in the lowland mammoth steppes. [8]
The aurochs is an extinct species of bovine, considered to be the wild ancestor of modern domestic cattle. With a shoulder height of up to 180 cm (71 in) in bulls and 155 cm (61 in) in cows, it was one of the largest herbivores in the Holocene; it had massive elongated and broad horns that reached 80 cm (31 in) in length.
The leopard is one of the five extant cat species in the genus Panthera. It has a pale yellowish to dark golden fur with dark spots grouped in rosettes. Its body is slender and muscular reaching a length of 92–183 cm (36–72 in) with a 66–102 cm (26–40 in) long tail and a shoulder height of 60–70 cm (24–28 in). Males typically weigh 30.9–72 kg (68–159 lb), and females 20.5–43 kg (45–95 lb).
Panthera is a genus within the family Felidae, and one of two extant genera in the subfamily Pantherinae. It contains the largest living members of the cat family. There are five living species: the jaguar, leopard, lion, snow leopard and tiger, as well as a number of extinct species, including the cave lion and American lion.
Dinofelis is an extinct genus of machairodontine, usually classified in the tribe Metailurini. It was widespread in Europe, Asia, Africa and North America from 5 million to about 1.2 million years ago. Fossils very similar to Dinofelis from Lothagam range back to around 8 million years ago, in the Late Miocene.
Homotherium is an extinct genus of scimitar-toothed cat belonging to the extinct subfamily Machairodontinae that inhabited North America, Eurasia, and Africa during the Pliocene and Pleistocene epochs from around 4 million to 12,000 years ago. It was one of the last surviving members of the subfamily alongside the more famous sabertooth Smilodon, to which it was not particularly closely related. It was a large cat, comparable in size to a lion, functioning as an apex predator in the ecosystems it inhabited. In comparison to Smilodon, the canines of Homotherium were shorter, and it is suggested to have had a different ecology from Smilodon as a pursuit predator adapted to running down large prey in open habitats, with Homotherium also proposed to have likely engaged in cooperative hunting.
The woolly rhinoceros is an extinct species of rhinoceros that inhabited northern Eurasia during the Pleistocene epoch. The woolly rhinoceros was a member of the Pleistocene megafauna. The woolly rhinoceros was covered with long, thick hair that allowed it to survive in the extremely cold, harsh mammoth steppe. It had a massive hump reaching from its shoulder and fed mainly on herbaceous plants that grew in the steppe. Mummified carcasses preserved in permafrost and many bone remains of woolly rhinoceroses have been found. Images of woolly rhinoceroses are found among cave paintings in Europe and Asia. The range of the woolly rhinoceros contracted towards Siberia beginning around 17,000 years ago, with the youngest known records being around 14,000 years old in northeast Siberia, coinciding with the Bølling–Allerød warming, which likely disrupted its habitat, with environmental DNA records possibly extending the range of the species around 9,800 years ago. Its closest living relative is the Sumatran rhinoceros.
Palaeoloxodon is an extinct genus of elephant. The genus originated in Africa during the Early Pleistocene, and expanded into Eurasia at the beginning of the Middle Pleistocene. The genus contains the largest known species of elephants, over 4 metres (13 ft) tall at the shoulders and over 13 tonnes (29,000 lb) in weight, representing among the largest land mammals ever, including the African Palaeoloxodon recki, the European straight-tusked elephant and the South Asian Palaeoloxodon namadicus. P. namadicus has been suggested to be the largest known land mammal by some authors based on extrapolation from fragmentary remains, though these estimates are highly speculative. In contrast, the genus also contains many species of dwarf elephants that evolved via insular dwarfism on islands in the Mediterranean, some like Palaeoloxodon falconeri less than 1 metre (3.3 ft) in shoulder height as fully grown adults, making them the smallest elephants known. The genus has a long and complex taxonomic history, and at various times, it has been considered to belong to Loxodonta or Elephas, but today is usually considered a valid and separate genus in its own right.
Panthera spelaea, commonly known as the cave lion, is an extinct Panthera species that was native to Eurasia and northwest North America during the Pleistocene epoch. Genetic analysis of ancient DNA has revealed that while closely related, it was a distinct species genetically isolated from the modern lion, with the genetic divergence between the two species estimated at around 500,000 years ago. The earliest fossils of the P. spelaea lineage in Eurasia date to around 700,000 years ago. It is closely related and probably ancestral to the American lion. The species ranged from Western Europe to eastern Beringia in North America, and was a prominent member of the mammoth steppe fauna, and an important apex predator across its range. It became extinct about 13,000 years ago. It closely resembled living lions with a coat of yellowish-grey fur though unlike extant lions, males appear to have lacked manes.
The American lion, with the species name meaning "savage" or "cruel", also called the North American lion) is an extinct pantherine cat native to North America during the Late Pleistocene from around 130,000 to 12,800 years ago. Genetic evidence suggests that its closest living relative is the lion, with the American lion representing an offshoot from the lineage of the largely Eurasian cave lion, from which it is suggested to have split around 165,000 years ago. Its fossils have been found across North America, from Canada to Mexico. It was about 25% larger than the modern lion, making it one of the largest known felids to ever exist, and an important apex predator.
The Pantherinae is a subfamily of the Felidae; it was named and first described by Reginald Innes Pocock in 1917 as only including the Panthera species, but later also came to include the clouded leopards. The Pantherinae genetically diverged from a common ancestor between 9.32 to 4.47 million years ago and 10.67 to 3.76 million years ago.
The history of lions in Europe is part of the wider history of the lion species complex. The rediscovery and confirmation of their presence in Europe, already known by myths, historical accounts and ancient art, was made possible by the finds of fossils of Pleistocene, Holocene and Ancient lions excavated in Europe since the early 19th century. The first excavated lion fossil was found in southern Germany, and described by Georg August Goldfuss using the scientific name Felis spelaea. It probably dates to the Würm glaciation, and is 191,000 to 57,000 years old. Since then, older lion skull fragments were excavated in Germany and in other parts of Europe, including in Western Europe. Some of them were described by Wilhelm von Reichenau under Felis fossilis in 1906.
The European wild ass or hydruntine is an extinct equine from the Middle Pleistocene to Late Holocene of Europe and West Asia, and possibly North Africa. It is a member of the subgenus Asinus, and closely related to the living Asiatic wild ass. The specific epithet, hydruntinus, means from Otranto.
The straight-tusked elephant is an extinct species of elephant that inhabited Europe and Western Asia during the Middle and Late Pleistocene. One of the largest known elephant species, mature fully grown bulls on average had a shoulder height of 4 metres (13 ft) and a weight of 13 tonnes (29,000 lb). Straight-tusked elephants likely lived very similarly to modern elephants, with herds of adult females and juveniles and solitary adult males. The species was primarily associated with temperate and Mediterranean woodland and forest habitats, flourishing during interglacial periods, when its range would extend across Europe as far north as Great Britain and Denmark and eastwards into Russia, while persisting in southern Europe during glacial periods. Skeletons found in association with stone tools and in one case, a wooden spear, suggest they were scavenged and hunted by early humans, including Homo heidelbergensis and their Neanderthal successors.
Panthera gombaszoegensis, also known as the European jaguar, is a Panthera species that lived from about 2.0 to 0.35 million years ago in Europe, as well as likely elsewhere in Eurasia. The first fossils were excavated in 1938 in Gombasek Cave, Slovakia. P. gombaszoegensis was a medium-large sized species that formed an important part of the European carnivore guild for a period of over a million years. Many authors have posited that it is the ancestor of the American jaguar, with some authors considering it the subspecies Panthera onca gombaszoegensis, though the close relationship between the two species has been questioned by some authors.
Panthera fossilis is an extinct species of cat belonging to the genus Panthera, known from remains found in Eurasia spanning the Middle Pleistocene and possibly into the Early Pleistocene.
Cave hyena are extinct species or subspecies of hyena known from Eurasia, which ranged from Western Europe to eastern Asia and Siberia during the Pleistocene epoch. It is well represented in many European caves, primarily dating to the Last Glacial Period. It was an apex predator that preyed on large mammals, and was responsible for the accumulation of hundreds of large Pleistocene mammal bones in areas including horizontal caves, sinkholes, mud pits, and muddy areas along rivers.
Stephanorhinus is an extinct genus of two-horned rhinoceros native to Eurasia and North Africa that lived during the Late Pliocene to Late Pleistocene. Species of Stephanorhinus were the predominant and often only species of rhinoceros in much of temperate Eurasia, especially Europe, for most of the Pleistocene. The last two species of Stephanorhinus – Merck's rhinoceros and the narrow-nosed rhinoceros – went extinct during the last glacial period.
The narrow-nosed rhinoceros, also known as the steppe rhinoceros is an extinct species of rhinoceros belonging to the genus Stephanorhinus that lived in western Eurasia, including Europe, as well as North Africa during the Pleistocene. It first appeared in Europe around 500,000 years ago during the Middle Pleistocene and survived there until at least 34,000 years Before Present. It was native to temperate and Mediterranean environments, where it fed on low growing plants and to a lesser extent woody plants. Evidence has been found that it was exploited for food by archaic humans, including Neanderthals.
Ursus ingressus is an extinct species of the family Ursidae that lived in Central Europe during the Late Pleistocene. It is named after the Gamssulzen Cave in Austria, where the holotype of this species was found.
Panthera uncia pyrenaica, also known as the Arago snow leopard or the European snow leopard, is a subspecies of snow leopard that lived during the Middle Pleistocene in southern France.