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. [1] [2] 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. [3] 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. [4]
The oldest remains of lions in Europe, assigned to the species Panthera fossilis , are over 600,000 years old. This species represents one of the largest known felines to have ever existed, with this species eventually evolving into the smaller (around the size of a modern lion) cave lion (Panthera spelaea), [5] which is widely depicted in Palaeolithic European art like cave paintings. [6] Remains of P. fossilis and P. spelaea are known from across Europe. [5] Cave lions became extinct around 14,000 years ago at the end of the Pleistocene. [7] During the early-middle Holocene (from around 8,000-6,000 years ago) modern lions colonised Southeast and parts of Central and Eastern Europe, [8] before becoming extinct in Europe likely during classical times [9] (or perhaps as late as the Middle Ages [8] ).
Lions have appeared in European literature since the times of Ancient Greece, such as in the Iliad , or with the story of the Nemean lion. [10] Ancient Greeks also depicted them in sculpture, such as with the Lion Gate of Mycenae or in the island-sanctuary of Delos, where various sculptures of lions survive to this day. [11]
During the Roman Republic and later Roman Empire, using lions in gladiatorial games and public spectacles was a prized endeavor. [12] North African Barbary lions were imported into Europe during the Middle Ages. [13]
Bone fragments of fossil spelaea lions indicate that they were bigger than the modern lion and had less specialized lower teeth, reduced lower premolars and smaller incisors. [14]
As indicated by numerous artistic depictions, modern lions in the Balkans had less developed manes, and lacked abdominal and lateral manes as well as limb hair. Οn the other hand, lions from Transcaucasia exhibited all these features. [15]
The oldest fossils excavated near Pakefield in the United Kingdom are estimated at 680,000 years old, and represent Panthera fossilis . [25] Lion fossils were excavated in Spain, Portugal, Italy, Belgium, France, United Kingdom, Germany, Poland, Czech Republic, Slovakia, Hungary and Russia. [14] [26] [27] [28] Panthera fossilis later evolved in to the cave lion (Panthera spelaea), which is known from fossils across Europe during the Late Pleistocene. [5]
The youngest unambiguous and well dated remains of cave lions are around 14,000 years old. [29]
Although there are some claimed records of early Holocene lion fossils from Italy, the dating of these fossils are uncertain. [30] The oldest confirmed remains of modern lion in Europe date to the early Holocene, around 8,000-6,000 years ago. [8]
A Neolithic lion tooth fragment representing the Atlantic Period was found in Karanovo, Bulgaria, and is estimated 6,000 years old. [24] In Greece, lions first appeared around 6,500–6,000 years ago as indicated by a front leg bone found in Philippi. [2] Bone fragments of the modern lion were excavated in Hungary and in Ukraine's Black Sea region, which are estimated at around 5,500 to 3,000 years old. [31] Remains were also found in Romania and European Turkey. [18]
In Southeast Europe, the lion inhabited part of the Balkan Peninsula, up to Hungary and Ukraine during the Neolithic period, [32] [33] (though some authors have argued that lions may have survived in Ukraine until the Middle Ages). [8] It survived in Bulgaria until the 4th or 3rd century BCE. [34] [35] Around 1000 BCE, it became extinct in the Peloponnese. [2] [9] It disappeared from Macedonia around the first century CE, from Western Thrace not before the 2nd century CE and from Thessaly possibly in the 4th century CE; Themistius regretted that no more lions could be furnished for beast-shows. [32] [33] [2] [36] [23]
In Transcaucasia, the lion was present until the 10th century. The peak of its historic range covered all of the plains and foothills of eastern Transcaucasia, westward almost to Tbilisi in modern Georgia. Northwards, its range extended through the eastern Caucasus, from the Apsheron Peninsula to the mouth of the Samur River near the current Azerbaijan-Russia border, extending to the Araks river. From there, the boundary of its range narrowly turned east to Yerevan in modern Armenia, with its northern boundary then extending westward to Turkey. [15]
Cave lions feature in a number of works of Palaeolithic art, though depictions are comparatively rare. [30] These include cave paintings, engravings, and sculptures, notably including the famous anthropomorphic lion-man figurine with a human-like body and a cave lion head. [37]
Lions feature in ancient Greek mythology and writings, including the myth of the Nemean lion, which was believed to be a supernatural lion that occupied the sacred town of Nemea in the Peloponnese. [38] Homer mentioned lions 45 times in his poems, but this could have been due to his experience in Asia Minor. [16] Phalaecus, a tyrant of Amvrakia (modern-day Arta), was allegedly killed by a female lion due to his holding a newborn lion cub, after finding it on a hunting expedition. [39] Conon refers to the myth of how Olynthus city got its name, when during around the period of the Trojan War, son of Strymon, Olynthos during a lion hunt was killed by a lion. [40] According to Herodotus lions occurred between Achelous river and Nestus, being plentiful between Akanthos and Thermi. When Xerxes advanced near Echedorus in 480 BC, the troops' camels were attacked by lions. [35] Xenophon stated around 400 BCE that lions were hunted around Mount Kissos, Pangaio, the Pindus mountains and elsewhere. [21] Aristotle in the 4th century BCE provided some data on lion distribution, behaviour, breeding and also anatomy. According to him, lions were more numerous in North Africa than in Europe; they had approached towns, and attacked people only if they were old, or had poor dental health. [20] Pliny the Elder mentions that European lions were stronger compared to those from Syria and Africa. [41] In the 2nd century CE, Pausanias referred to lion presence east of Nestus in Thrace, in the area of Abdera. He also referred to a story about Polydamas of Skotoussa, an Olympic winner in the 5th century BCE, who allegedly used his bare hands to kill a lion on Thessalian part of Mount Olympus; and to one about Caranus of Macedon who according to the Macedonians, raised a trophy that was thrown down and destroyed by a lion that was rushing down from Mount Olympus. [22]
The Romans used Barbary lions from North Africa for lion-baiting, [42] and lions from Greece for gladiatorial games. [10] [23]
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.
Felidae is the family of mammals in the order Carnivora colloquially referred to as cats. A member of this family is also called a felid.
The lion is a large cat of the genus Panthera, native to Africa and India. It has a muscular, broad-chested body; a short, rounded head; round ears; and a dark, hairy tuft at the tip of its tail. It is sexually dimorphic; adult male lions are larger than females and have a prominent mane. It is a social species, forming groups called prides. A lion's pride consists of a few adult males, related females, and cubs. Groups of female lions usually hunt together, preying mostly on medium-sized and large ungulates. The lion is an apex and keystone predator; although some lions scavenge when opportunities occur and have been known to hunt humans, lions typically do not actively seek out and prey on humans.
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.
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 a had 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 Barbary lion was a population of the lion subspecies Panthera leo leo. It was also called North African lion, Atlas lion and Egyptian lion. It lived in the mountains and deserts of the Maghreb of North Africa from Morocco to Egypt. It was eradicated following the spread of firearms and bounties for shooting lions. A comprehensive review of hunting and sighting records revealed that small groups of lions may have survived in Algeria until the early 1960s, and in Morocco until the mid-1960s. Today, it is locally extinct in this region. Fossils of the Barbary lion dating to between 100,000 and 110,000 years were found in the cave of Bizmoune near Essaouira.
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 Asiatic lion is a lion population of the subspecies Panthera leo leo. Until the 19th century, it occurred in Saudi Arabia, eastern Turkey, Iran, Mesopotamia, and from east of the Indus River in Pakistan to the Bengal region and the Narmada River in Central India. Since the turn of the 20th century, its range has been restricted to Gir National Park and the surrounding areas in the Indian state of Gujarat. The first scientific description of the Asiatic lion was published in 1826 by the Austrian zoologist Johann N. Meyer, who named it Felis leo persicus.
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.
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. The first fossils were excavated in 1938 in Gombasek Cave, Slovakia. Some records were also reported from Africa and Asia. 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.
Panthera youngi is a fossil cat species that was described in 1934; fossil remains of this cat were excavated in a Homo erectus formation in Choukoutien, northeastern China. Upper and lower jaws excavated in Japan's Yamaguchi Prefecture were also attributed to this species. It is estimated to have lived about 350,000 years ago in the Pleistocene epoch. It was suggested that it was conspecific with Panthera atrox and P. spelaea due to their extensive similarities. Some dental similarities were also noted with the older P. fossilis, however, Panthera youngi showed more derived features.
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 lions are large extinct carnivorous felids that are classified either as subspecies of the lion, or as distinct but closely related species, depending on the authority.
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.
The Petralona Cave a karst formation, is located at 300 m (984 ft) above sea-level on the western foot of Mount Katsika, about 1 km (0.62 mi) east of the village of Petralona, about 35 km (22 mi) south-east of Thessaloniki city on the Chalkidiki peninsula, Greece. The site came to public attention when in 1960 a fossilized archaic human skull was found. The cave had been discovered accidentally only a year earlier (1959) after erosion had left clefts in the rock. "Bejeweled" with impressive stalactite and stalagmite formations and holding an abundance of fossils, the cave soon attracted geologists and paleontologists. After decades of excavations the cave is open to the public and scientific work is documented and presented in an adjacent archaeological museum.
Panthera pardus spelaea, also known as the European Ice Age leopard or the cave leopard, is a fossil leopard subspecies which roamed Europe in the Late Pleistocene and possibly the Holocene.
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 shawi is an extinct prehistoric cat, of which a single canine tooth was excavated in Sterkfontein cave in South Africa by Robert Broom in the 1940s. It is thought to be one of the oldest known Panthera species in Africa.
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