History of lions in Europe

{{multiple image |align=right |direction=vertical |image1=Modern lion locations in southeastern Europe except Caucasus.png |caption1=Records of modern lion in southeastern Europe

Red: excavated lion remains{{cite book |last=Thomas |first=N.R. |year=2004 |title=ΧΑΡΙΣ: Essays in Honor of Sara A. Immerwahr |chapter=The Early Mycenaean Lion up to Date |jstor=1354068 |series=Hesperia Supplements |volume=33 |number=33 |pages=161−206 |publisher=The American School of Classical Studies at Athens}}

Orange: locations mentioned by ancient Greek authors{{Cite book |author=Beloe, W. |year=1830 |title=Herodotus translated from the Greek |chapter=Book VII. Polymnia |publisher=Jones and Co. |location=London |chapter-url=https://archive.org/stream/herodotustrfrom00hero?ref=ol#page/320/mode/2up |pages=321−380}}
Yellow: locations mentioned in Greek legends |image2=Eastern mound Karanovo-2011-03.jpg |caption2=Karanovo, Nova Zagora Municipality in Bulgaria, where the fragments of a lion's tooth about 6,000 years old were found}}

Lions seem to have first evolved in east Africa around the Plio-Peistocene boundary (ca. 2 million years ago).{{Cite journal |last1=Yamaguchi |first1=N. |last2=Cooper |first2=A. |last3=Werdelin |first3=L. |last4=Macdonald |first4=D. W. |date=2004 |title=Evolution of the mane and group-living in the lion (Panthera leo): a review |url=https://zslpublications.onlinelibrary.wiley.com/doi/10.1017/S0952836904005242 |journal=Journal of Zoology |language=en |volume=263 |issue=4 |pages=329–342 |doi=10.1017/S0952836904005242 |issn=1469-7998|url-access=subscription }} 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.{{Cite journal |last1=Marciszak |first1=A. |last2=Lipecki |first2=G. |last3=Pawłowska |first3=K. |last4=Jakubowski |first4=G. |last5=Ratajczak-Skrzatek |first5=U. |last6=Zarzecka-Szubińska |first6=K. |last7=Nadachowski |first7=A. |date=2021 |title=The Pleistocene lion Panthera spelaea (Goldfuss, 1810) from Poland – A review |journal=Quaternary International |series=The Quaternary of Europe and Adjacent Areas: Stratigraphical Perspectives and Tools for Correlations – SEQS-2019 |volume=605-606 |pages=213–240 |doi=10.1016/j.quaint.2020.12.018 |bibcode=2021QuInt.605..213M |issn=1040-6182}}{{Cite journal |last1=Madurell-Malapeira |first1=J. |last2=Prat-Vericat |first2=M. |last3=Bartolini-Lucenti |first3=S. |last4=Faggi |first4=A. |last5=Fidalgo |first5=D. |last6=Marciszak |first6=A. |last7=Rook |first7=L. |date=2024 |title=A Review on the Latest Early Pleistocene Carnivoran Guild from the Vallparadís Section (NE Iberia) |journal=Quaternary |volume=7 |issue=3 |pages=40 |doi=10.3390/quat7030040 |doi-access=free |bibcode=2024Quat....7...40M}} These fossils are classified as Panthera fossilis and Panthera (spelaea) fossilis. Some specimens have been estimated at around {{cvt|500|kg}} in life, which would make Panthera fossilis one of the largest felids that ever lived. This lion was widespread in Europe and across Asia, eventually giving rise to the true cave lion (Panthera 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.{{Cite journal |last1=Marciszak |first1=A. |last2=Ivanoff |first2=D. V. |last3=Semenov |first3=Y. A. |last4=Talamo |first4=S. |last5=Ridush |first5=B. |last6=Stupak |first6=A. |last7=Yanish |first7=Y. |last8=Kovalchuk |first8=O. |date=2023 |title=The Quaternary lions of Ukraine and a trend of decreasing size in Panthera spelaea |journal=Journal of Mammalian Evolution |volume=30 |issue=1 |pages=109–135 |doi=10.1007/s10914-022-09635-3|hdl=11585/903022 |hdl-access=free }} 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 extinctions and the collapse of the mammoth steppe.{{cite journal |author1=Stuart, A. J. |author2=Lister, A. M. |year=2011 |title=Extinction chronology of the cave lion Panthera spelaea |journal=Quaternary Science Reviews |volume=30 |issue=17 |pages=2329–2340 |bibcode=2011QSRv...30.2329S |doi=10.1016/j.quascirev.2010.04.023}}

Although there are some claimed records of early Holocene cave lion fossils from Italy, the dating of these fossils are uncertain.{{Cite journal |last1=Sigari |first1=D. |last2=Bourdier |first2=C. |last3=Conti |first3=C. |last4=Conti |first4=J. |last5=Forti |first5=L. |last6=García-Diez |first6=M. |last7=Lai |first7=G. |last8=Mazzini |first8=I. |last9=Pieruccini |first9=P. |last10=Sardella |first10=R. |date=June 2024 |title=The last cave lion of the late Upper Palaeolithic: The engraved feline of Grotta Romanelli (southern Italy) |url=https://linkinghub.elsevier.com/retrieve/pii/S0277379124001719 |journal=Quaternary Science Reviews |language=en |volume=334 |pages=108670 |doi=10.1016/j.quascirev.2024.108670|bibcode=2024QSRv..33408670S }} The oldest confirmed remains of modern lion in Europe date to the early Holocene, around 8,000-6,000 years ago.

A Neolithic lion tooth fragment representing the Atlantic Period was found in Karanovo, Bulgaria, and is estimated 6,000 years old.{{cite journal | last = Bökönyi | first = S. |year=1989 |title= Erster vorläufiger Bericht über Tierknochenfunde der Karanovo-V-Besiedlung in Drama |journal=Berichte der Römisch-Germanischen Kommission |issue=70 |pages= 123–127}} In Greece, lions first appeared around 6,500–6,000 years ago as indicated by a front leg bone found in Philippi. 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.{{cite journal | last1 =Sommer | first1 = R. S. |last2 =Benecke | first2 = N. |doi=10.1111/j.1469-7998.2005.00040.x |title=Late Pleistocene and Holocene development of the felid fauna (Felidae) of Europe: A review |year=2006 |journal= Journal of Zoology |volume=269 |issue=1 |pages= 7–19}} Remains were also found in Romania and European Turkey.{{cite journal | last = Thomas | first = N.R. |year=2014 | title =A lion's eye view of the Greek Bronze Age |pages=375−92 | number = 37 | journal = Aegaeum. Annales Liégeoises et PASPiennes d'Archéologie égéenne |editor1-last =Touchais | editor1-first = G. | editor2-last = Laffineur | editor2-first = R. | editor3-last = Rougemeont | editor3-first = F. }}

File:Pannonian_Basin_early-mid_Holocene.jpg early-mid Holocene scene, featuring, from left to right, Asiatic lions, European wild ass, Pannonian sousliks, European rollers, great bustards and a lesser grey shrike.]]

In Southeast Europe, the modern lion (Panthera leo) inhabited part of the Balkan Peninsula as well as adjacent areas, ranging northwestwards to Hungary and eastwards Ukraine during the Neolithic period,{{cite book |author=Douglas, N. |year=1927 |title=Birds and Beasts of Greek Anthology |location=Florence |publisher=Norman Douglas |url=http://gutenberg.net.au/ebooks03/0300611h.html |access-date=2018-08-31 |archive-date=2024-08-17 |archive-url=https://web.archive.org/web/20240817150805/http://gutenberg.net.au/ebooks03/0300611h.html |url-status=live}}{{cite journal |author=Alden, M. |year=2005 |title=Lions in paradise: Lion Similes in the Iliad and the Lion Cubs of IL. 18.318-22 |journal=The Classical Quarterly |issue=55 |pages=335–342 |doi=10.1093/cq/bmi035}} It survived in Bulgaria until the 4th or 3rd century BC.{{cite journal |last1=Masseti |first1=M. |last2=Mazza |first2=P.P.A. |year=2013 |title=Western European Quaternary lions: new working hypotheses |journal=Biological Journal of the Linnean Society |volume=109 |issue=109 |pages=66–77 |doi=10.1111/bij.12032 |doi-access=free}} Around 1000 BC, it became extinct in the Peloponnese.{{cite journal |author=Schnitzler, A. E. |year=2011 |title=Past and present distribution of the North African-Asian lion subgroup: a review |journal=Mammal Review |volume=41 |issue=3 |pages=220–243 |doi=10.1111/j.1365-2907.2010.00181.x |bibcode=2011MamRv..41..220S}} 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.{{cite book |last=Cohen |first=A. |year=2010 |title=Art in the era of Alexander the Great: Paradigms of manhood and their cultural traditions |publisher=Cambridge University Press |location=Cambridge |isbn=978-0-5217-6904-4 |chapter=Master of Lions (and other Animals) |pages=64–118 |chapter-url=https://books.google.com/books?id=nX8F_ZV83vUC&pg=PA64}}{{cite book |author=Uhm, D.P. van |year=2016 |title=The Illegal Wildlife Trade: Inside the World of Poachers, Smugglers and Traders |location=Switzerland |publisher=Springer International Publishing}} 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. Some records potentially belonging to Panthera leo have been reported from the early Holocene of the northern Iberian Peninsula and Northern Italy, including Cueva La Riera in Spain and Grotta all’Onda in Italy, though these fossils are poorly dated and fragmentary, making it impossible from morphology to determine whether they represent modern P. leo or cave lions.

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.{{Cite book |last1=Heptner |first1=V. G. |title=Mlekopitajuščie Sovetskogo Soiuza. Moskva: Vysšaia Škola |last2=Sludskiy |first2=A. A. |publisher=Smithsonian Institution and the National Science Foundation |year=1992 |isbn=978-90-04-08876-4 |location=Washington DC |pages=83–95 |trans-title=Mammals of the Soviet Union, Volume II, Part 2 |chapter=Lion |orig-year=1972 |chapter-url=https://archive.org/stream/mammalsofsov221992gept#page/82/mode/2up}}

{{see also|Damnatio ad bestias|Gulf of Lion|Lion Gate}}

File:Loewenmensch2.jpg found in Germany has been dated to the Upper Paleolithic during the Pleistocene, about 35,000 to 40,000 years ago]]

Cave lions feature in a number of works of Palaeolithic art, though depictions are comparatively rare. 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.{{Cite journal |last1=Russo |first1=G. |last2=Milks |first2=A. |last3=Leder |first3=D. |last4=Koddenberg |first4=T. |last5=Starkovich |first5=B. M. |last6=Duval |first6=M. |last7=Zhao |first7=J.-X. |last8=Darga |first8=R. |last9=Rosendahl |first9=W. |last10=Terberger |first10=T. |date=2023 |title=First direct evidence of lion hunting and the early use of a lion pelt by Neanderthals |journal=Scientific Reports |volume=13 |issue=1 |page=16405 |bibcode=2023NatSR..1316405R |doi=10.1038/s41598-023-42764-0 |pmc=10570355 |pmid=37828055}}

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.{{cite book |author=Hard, R. |year=2003 |title=The Routledge Handbook of Greek Mythology: Based on HJ Rose's Handbook of Greek Mythology |location=London and New York |publisher=Routledge}} Homer mentioned lions 45 times in his poems, but this could have been due to his experience in Asia Minor.

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.Stephens, S. Callimachus (2015). Aetia. Dickinson College Commentaries. [https://web.archive.org/web/20180117070329/https://ds-drupal.haverford.edu/dcc/pl/callimachus-aetia/book-3/phalaecus-ambracia] 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.Gedoyn, A. 1738: 186 Récits de Conon. 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.{{Cite book |last=Guggisberg |first=C. A. W. |chapter=Lion Panthera leo (Linnaeus, 1758) |title=Wild Cats of the World |year=1975 |publisher=Taplinger Publishing |location=New York |isbn=978-0-8008-8324-9 |pages=[https://archive.org/details/wildcatsofworld00gugg/page/138 138–179] |chapter-url-access=registration |chapter-url=https://archive.org/details/wildcatsofworld00gugg/page/138 }} Xenophon stated around 400 BC that lions were hunted around Mount Kissos, Pangaio, the Pindus mountains and elsewhere.Ξενοφών, 5th-4th century BC: Κυνηγετικός 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.Αριστοτέλης, 4th century BC: Των περί τα ζώα ιστοριών. Pliny the Elder mentions that European lions were stronger compared to those from Syria and Africa.Боев, З. 2016. Левът наш балкански. – Списание Осем, 6: 98-105. 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.{{cite book |editor=Frazer, J. G. |year=1898 |title=Pausanias's Description of Greece |location=London |publisher=Macmillan and Co. |url=https://archive.org/stream/pausaniassdescri01pausuoft/pausaniassdescri01pausuoft#page/n5/mode/2up}}

The Romans used Barbary lions from North Africa for lion-baiting,{{Cite book |last1=Nowell |first1=K. |last2=Jackson |first2=P. |title=Wild Cats: Status Survey and Conservation Action Plan |chapter-url=http://carnivoractionplans1.free.fr/wildcats.pdf |year=1996 |publisher=IUCN/SSC Cat Specialist Group |location=Gland, Switzerland |isbn=978-2-8317-0045-8 |pages=17–21, 37–41 |chapter=Panthera leo |access-date=2018-02-02 |archive-date=2005-05-29 |archive-url=https://web.archive.org/web/20050529182212/http://carnivoractionplans1.free.fr/wildcats.pdf |url-status=live}} and lions from Greece for gladiatorial games.

File:Lions painting, Chauvet Cave (museum replica).jpg|Upper Paleolithic cave painting depicting cave lions, found in the Chauvet Cave, France{{cite book |author=Chauvet, J.-M. |author2=Brunel, D. E. |author3=Hillaire, C. |year=1996 |title=Dawn of Art: The Chauvet Cave. The oldest known paintings in the world |url=https://archive.org/details/dawnofartchauvet0000chau |url-access=registration |location=New York |publisher=Harry N. Abrams}}

File:Herakles lion Louvre CA1340.jpg|Heracles and the Nemean lion, c. 540 BC, Boeotia, Greece

File:Brauron - Marble Slab with a Lion.jpg|Lion sculpture, 4th century BC, Koropi, Greece

File:Hunting Mycenaean Dagger.jpg|Depiction of a hunting scene on a dagger found in Mycenae, Greece, 16th century BC

File:Met, greek, attic, marble lion, mid 4th century BC.JPG|Marble lion from Greece, mid-4th century BC

File:7704 - Piraeus Arch. Museum, Athens - Lion from the grave for Pelthinikos - Photo by Giovanni Dall'Orto,.jpg|Depiction of the lion from the 4th century BC, Greece

File:AR nomos of Velia.jpg|Silver stater struck in Velia 334-300 BC depicting Athena wearing a Phrygian helmet decorated with a centaur and lion devouring prey

File:NHMB-Zhaba-mogila-Strelcha-relief-of-lion-polychrome-decoration-5-4-centuryBC.jpg|Stone relief of a lion with a polychrome decorations, Zhaba Mogila, Strelcha, Bulgaria, 5th century BC.

File:Lion hunt mosaic from Pella.jpg|Mosaic from Pella (ancient Macedonia), late 4th century BC, depicting Alexander the Great and Craterus.

• Panthera leo leo
• Asiatic lion
• Cape lion
• Barbary lion
• Panthera atrox
• Cultural depictions of lions

{{Reflist}}

• [http://tolweb.org/treehouses/?treehouse_id=4732 Panthera leo europaea]
• [http://www.strangehistory.net/2013/06/29/the-last-european-lion/ The Last European Lion]

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