
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] These findings have confirmed that lions have been present more or less continuously in Europe up to historic times, starting in the early Pleistocene.
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 felids to have ever existed, with this species eventually evolving into the smaller cave lion (Panthera spelaea),[3] which is widely depicted in Palaeolithic European cave art.[4] Remains of P. fossilis and P. spelaea are known from across Europe.[3] Cave lions became extinct around 14,000 years ago at the end of the Pleistocene.[5] During the early-middle Holocene (from around 8,000-6,000 years ago) modern lions colonised Southwest, Southeast and parts of Central and Eastern Europe,[6] before becoming extinct in Europe likely during classical times[7] (or perhaps as late as the Middle Ages).[6]
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.[8] 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.[9]
During the Roman Republic and later Roman Empire, using lions in gladiatorial games and public spectacles was a prized endeavor. [10] North African Barbary lions were imported into Europe during the Middle Ages.[11]
Distribution
Pleistocene records
Lions seem to have first evolved in east Africa around the Plio-Peistocene boundary (ca. 2 million years ago).[21] The earliest remains of lions in Eurasia have been dated to around 1 million years ago, and lions became ubiquitous in Europe around 700-600,000 years ago.[22][23] These fossils are classified as Panthera (spelaea) fossilis. Some specimens have been estimated at around 500 kg in life, which would make Panthera fossilis one of the largest felids that ever lived.[22][23] This lion was widespread in Europe and across Asia, eventually giving rise to the true cave lion (Panthera (spelaea) spelaea) of the later Pleistocene, which was still large, but experienced a drastic size reduction during the late Pleistocene (around 50 to 45,000 years ago).[24][3] The range of the cave lion seems to have collapsed across Eurasia around 14,000 years ago, and its extinction around the same time is to be considered within the broader context of the late Pleistocene megafauna extinctions and the collapse of the mammoth steppe.[25]
Holocene records
Although there are some tentative records of early Holocene cave lion remains from Italy, their dating is considered uncertain.[26] The earliest European remains of modern lions are from the Iberian Peninsula, and have been dated to the earliest Holocene (Boreal/Subboreal). The Balkans were most likely colonised from Asia, across the Bosporus, in the early Holocene, around 6,000 years ago.[6] The earliest record here is a Neolithic tooth fragment from the Atlantic Period found in Karanovo, Bulgaria.[20] 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 2,000 years old.[27][28] Remains were also found in Romania and European Turkey.[14]
Historic range of Panthera leo
In Southeast Europe, the lion inhabited part of the Balkan Peninsula as well as adjacent areas, ranging northwestwards to Hungary and eastwards Ukraine during the Neolithic period,[29][30][6] In the Iberian Peninsula, lions were present until at least the early Holocene, however, it is unknown how these relate to the better-known remains from the Balkans. Potentially, however, the lion remains from the Iberian Peninsula may represent an earlier colonisation event.[27] In the Balkan Peninsula, the lion survived in Bulgaria until the 4th or 3rd century BC,[31][32] and until around 1000 BC in the Peloponnese.[2][7] It disappeared from Macedonia around the first century AD, from Western Thrace not before the 2nd century AD and from Thessaly possibly in the 4th century CE; Themistius regretted that no more lions could be furnished for beast-shows.[29][30][2][33][19] Some authors have argued that the lion may have survived in Ukraine as late as the High Middle Ages, based on a report of the 12th century Kievan Rus' prince Vladimir Monomakh encountering a ‘fierce beast’, which some have conjectured to be a lion.[6]
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.[34]
In culture

Cave lions feature in a number of works of Palaeolithic art, though depictions are comparatively rare.[26] 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.[35]
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.[36] Homer mentioned lions 45 times in his poems, but this could have been due to his experience in Asia Minor.[12] 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.[37] 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.[38] 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.[32] Xenophon stated around 400 BC that lions were hunted around Mount Kissos, Pangaio, the Pindus mountains and elsewhere.[17] Aristotle in the 4th century BC 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.[16] Pliny the Elder mentions that European lions were stronger compared to those from Syria and Africa.[39] In the 2nd century AD, 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 BC, 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.[18]
The Romans used Barbary lions from North Africa for lion-baiting,[40] and lions from Greece for gladiatorial games.[8][19]
-
Lion sculpture, 4th century BC, Koropi, Greece
-
Depiction of a hunting scene on a dagger found in Mycenae, Greece, 16th century BC
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Marble lion from Greece, mid-4th century BC
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Depiction of the lion from the 4th century BC, Greece
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Silver stater struck in Velia 334-300 BC depicting Athena wearing a Phrygian helmet decorated with a centaur and lion devouring prey
-
Mosaic from Pella (ancient Macedonia), late 4th century BC, depicting Alexander the Great and Craterus.
See also
References
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- ^ a b c Marciszak, Adrian; Gornig, Wiktoria (September 2024). "From giant to dwarf: A trend of decreasing size in Panthera spelaea (Goldfuss, 1810) and its likely implications". Earth History and Biodiversity. 1: 100007. doi:10.1016/j.hisbio.2024.100007.
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- ^ Stuart, A. J. & Lister, A. M. (2011). "Extinction chronology of the cave lion Panthera spelaea". Quaternary Science Reviews. 30 (17): 2329–2340. Bibcode:2011QSRv...30.2329S. doi:10.1016/j.quascirev.2010.04.023.
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