History of lions in Europe

Short description: Lions in prehistoric and historic Europe

Skeleton of a cave lion (P. spelaea) in the Natural History Museum, Vienna

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 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; it eventually evolved into the smaller, modern lion-sized cave lion (Panthera spelaea),[3] which is widely depicted in Palaeolithic European cave painting.[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 Asiatic lions colonised Southeast Europe 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]

## Contents

* 1 Distribution
* 1.1 Pleistocene records
* 1.2 Holocene records
* 1.3 Historic range of Panthera leo
* 2 In culture
* 3 See also
* 4 References
* 5 External links

## Distribution

Records of modern lion in southeastern Europe
Red: excavated lion remains[2][7][12][13]
Orange: locations mentioned by ancient Greek authors[14][15][16][17][8][18]
Yellow: locations mentioned in Greek legends

Karanovo, Nova Zagora Municipality in Bulgaria, where the fragments of a lion's tooth about 6,000 years old were found[19]

### Pleistocene records

Lions seem to have first evolved in east Africa around the Plio-Peistocene boundary (ca. 2 million years ago).[20] 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.[21][22] These fossils are classified as Panthera fossilis and Panthera (spelaea) fossilis. Some specimens have been estimated at around 500 kg (1,100 lb) in life, which would make Panthera fossilis one of the largest felids that ever lived.[21][22] 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.[23][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 extinctions and the collapse of the mammoth steppe.[24]

### Holocene records

Although there are some claimed records of early Holocene cave lion fossils from Italy, the dating of these fossils are uncertain.[25] The oldest confirmed remains of modern lion in Europe date to the early Holocene, around 8,000-6,000 years ago. European Holocene lions are suggested to represent part of the same population as living Asiatic lions.[6]

A Neolithic lion tooth fragment representing the Atlantic Period was found in Karanovo, Bulgaria, and is estimated 6,000 years old.[19] 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.[26] Remains were also found in Romania and European Turkey.[13] Interestingly a human skeleton found near the Kozareva mound (Eastern Bulgaria) and dating to ca. 4600–4200 BC, bears lesions that indicate a big carnivore attack, most likely done by a lion.[27]

### Historic range of Panthera leo

A Pannonian 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.[28][29][6] It survived in Bulgaria until the 4th or 3rd century BC.[30][31] Around 1000 BC, it became extinct 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.[28][29][2][32][18] 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] 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.[30]

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.[33]

## In culture

The anthropomorphic Löwenmensch (lion-man) figurine 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.[25] 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.[34]

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.[35] Homer mentioned lions 45 times in his poems, but this could have been due to his experience in Asia Minor.[7] Phalaecus, a tyrant of Amvrakia (modern-day Arta), was allegedly killed by a female lion due to him holding a newborn lion cub, after finding it on a hunting expedition.[36] Conon refers to the myth of how the city of Olynthus got its name around the period of the Trojan War, when Olynthos, son of King Strymon, was killed by a lion during a lion hunt.[37] 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.[31] Xenophon stated around 400 BC that lions were hunted around Mount Kissos, Pangaio, the Pindus mountains and elsewhere.[16] 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.[15] Pliny the Elder mentions that European lions were stronger compared to those from Syria and Africa.[38] 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; the legend of Mermerus who was killed during a hunt by a lioness, opposite Corfu; 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.[17]

The Romans used Barbary lions from North Africa for lion-baiting,[39] and lions from Greece for gladiatorial games.[8][18]

* Upper Paleolithic cave painting depicting cave lions, found in the Chauvet Cave, France[40]

* Heracles and the Nemean lion, c. 540 BC, Boeotia, Greece

* Lion sculpture, 4th century BC, Koropi, Greece

* Depiction of a hunting scene on a dagger found in Mycenae, Greece, 16th century BC

* Marble lion from Greece, mid-4th century BC

* Depiction of the lion from the 4th century BC, Greece

* Silver stater struck in Velia 334-300 BC depicting Athena wearing a Phrygian helmet decorated with a centaur and lion devouring prey

* Stone relief of a lion with a polychrome decorations, Zhaba Mogila, Strelcha, Bulgaria, 5th century BC.

* Mosaic from Pella (ancient Macedonia), late 4th century BC, depicting Alexander the Great and Craterus.

## See also

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

## References

1. ↑ Diedrich, C.G. (2011). "The largest European lion Panthera leo spelaea (Goldfuss 1810) population from the Zoolithen Cave, Germany: specialised cave bear predators of Europe". Historical Biology 23 (2–3): 271–311. doi:10.1080/08912963.2010.546529. Bibcode: 2011HBio...23..271D.
2. ↑ 2.0 2.1 2.2 2.3 2.4 Bartosiewicz, L. (2009). "A Lion's share of attention: Archaeozoology and the historical record". Acta Archaeologica Academiae Scientiarum Hungaricae 60 (1): 759–773. doi:10.1556/AArch.59.2008.2.28.
3. ↑ 3.0 3.1 3.2 Marciszak, A.; Gornig, W. (2024). "From giant to dwarf: A trend of decreasing size in Panthera spelaea (Goldfuss, 1810) and its likely implications". Earth History and Biodiversity 1. doi:10.1016/j.hisbio.2024.100007.
4. ↑ Russo, G.; Milks, A.; Leder, D.; Koddenberg, T.; Starkovich, B. M.; Duval, M.; Zhao, J.-X.; Darga, R. et al. (2023). "First direct evidence of lion hunting and the early use of a lion pelt by Neanderthals". Scientific Reports 13 (1): 16405. doi:10.1038/s41598-023-42764-0. PMID 37828055. Bibcode: 2023NatSR..1316405R.
5. ↑ Stuart, A. J.; Lister, A. M. (2011). "Extinction chronology of the cave lion Panthera spelaea". Quaternary Science Reviews 30 (17): 2329–2340. doi:10.1016/j.quascirev.2010.04.023. Bibcode: 2011QSRv...30.2329S.
6. ↑ 6.0 6.1 6.2 6.3 6.4 Marciszak, A.; Ivanoff, D. V.; Semenov, Y. A.; Talamo, S.; Ridush, B.; Stupak, A.; Yanish, Y.; Kovalchuk, O. (2022). "The Quaternary lions of Ukraine and a trend of decreasing size in Panthera spelaea". Journal of Mammalian Evolution 30 (1): 109–135. doi:10.1007/s10914-022-09635-3.
7. ↑ 7.0 7.1 7.2 7.3 Schnitzler, A. E. (2011). "Past and present distribution of the North African-Asian lion subgroup: a review". Mammal Review 41 (3): 220–243. doi:10.1111/j.1365-2907.2010.00181.x. Bibcode: 2011MamRv..41..220S.
8. ↑ 8.0 8.1 8.2 Alden, M. (2005). "Lions in paradise: Lion similes in the Iliad and the Lion Cubs of IL. 18.318-22". The Classical Quarterly (55): 335–342. doi:10.1093/cq/bmi035.
9. ↑ Hermary, Antoine (2020-12-01). "Sculptures archaïques de Délos: deux lions, une sirène et deux oiseaux" (in fr). Bulletin de correspondance hellénique 144 (1). doi:10.4000/bch.1045. ISSN 0007-4217. https://journals.openedition.org/bch/1045. Retrieved 2024-08-17.
10. ↑ Epplett, W. C. (2009). Animal spectacula of the Roman Empire (Thesis). University Of British Columbia. University of British Columbia. doi:10.14288/1.0090535.
11. ↑ Barnett, R., Yamaguchi, N., Shapiro, B. & Sabin, R. (2008). Ancient DNA analysis indicates the first English lions originated from North Africa. Contributions to Zoology, 77(1), 7–16.
12. ↑ Thomas, N.R. (2004). "The Early Mycenaean Lion up to Date". ΧΑΡΙΣ: Essays in Honor of Sara A. Immerwahr. Hesperia Supplements. 33. The American School of Classical Studies at Athens. pp. 161−206.
13. ↑ 13.0 13.1 Thomas, N.R. (2014). Touchais, G.; Laffineur, R.; Rougemeont, F.. eds. "A lion's eye view of the Greek Bronze Age". Aegaeum. Annales Liégeoises et PASPiennes d'Archéologie égéenne (37): 375−92.
14. ↑ Beloe, W. (1830). "Book VII. Polymnia". Herodotus translated from the Greek. London: Jones and Co.. pp. 321−380. https://archive.org/stream/herodotustrfrom00hero?ref=ol#page/320/mode/2up.
15. ↑ 15.0 15.1 Αριστοτέλης, 4th century BC: Των περί τα ζώα ιστοριών.
16. ↑ 16.0 16.1 Ξενοφών, 5th-4th century BC: Κυνηγετικός
17. ↑ 17.0 17.1 Frazer, J. G., ed (1898). Pausanias's Description of Greece. London: Macmillan and Co.. https://archive.org/stream/pausaniassdescri01pausuoft/pausaniassdescri01pausuoft#page/n5/mode/2up.
18. ↑ 18.0 18.1 18.2 Uhm, D.P. van (2016). The Illegal Wildlife Trade: Inside the World of Poachers, Smugglers and Traders. Switzerland: Springer International Publishing.
19. ↑ 19.0 19.1 Bökönyi, S. (1989). "Erster vorläufiger Bericht über Tierknochenfunde der Karanovo-V-Besiedlung in Drama". Berichte der Römisch-Germanischen Kommission (70): 123–127.
20. ↑ Yamaguchi, N.; Cooper, A.; Werdelin, L.; Macdonald, D. W. (2004). "Evolution of the mane and group-living in the lion (Panthera leo): a review" (in en). Journal of Zoology 263 (4): 329–342. doi:10.1017/S0952836904005242. ISSN 1469-7998. https://zslpublications.onlinelibrary.wiley.com/doi/10.1017/S0952836904005242.
21. ↑ 21.0 21.1 Marciszak, A.; Lipecki, G.; Pawłowska, K.; Jakubowski, G.; Ratajczak-Skrzatek, U.; Zarzecka-Szubińska, K.; Nadachowski, A. (2021). "The Pleistocene lion Panthera spelaea (Goldfuss, 1810) from Poland – A review". Quaternary International. The Quaternary of Europe and Adjacent Areas: Stratigraphical Perspectives and Tools for Correlations – SEQS-2019 605-606: 213–240. doi:10.1016/j.quaint.2020.12.018. ISSN 1040-6182. Bibcode: 2021QuInt.605..213M.
22. ↑ 22.0 22.1 Madurell-Malapeira, J.; Prat-Vericat, M.; Bartolini-Lucenti, S.; Faggi, A.; Fidalgo, D.; Marciszak, A.; Rook, L. (2024). "A Review on the Latest Early Pleistocene Carnivoran Guild from the Vallparadís Section (NE Iberia)". Quaternary 7 (3): 40. doi:10.3390/quat7030040. Bibcode: 2024Quat....7...40M.
23. ↑ Marciszak, A.; Ivanoff, D. V.; Semenov, Y. A.; Talamo, S.; Ridush, B.; Stupak, A.; Yanish, Y.; Kovalchuk, O. (2023). "The Quaternary lions of Ukraine and a trend of decreasing size in Panthera spelaea". Journal of Mammalian Evolution 30 (1): 109–135. doi:10.1007/s10914-022-09635-3.
24. ↑ Stuart, A. J.; Lister, A. M. (2011). "Extinction chronology of the cave lion Panthera spelaea". Quaternary Science Reviews 30 (17): 2329–2340. doi:10.1016/j.quascirev.2010.04.023. Bibcode: 2011QSRv...30.2329S.
25. ↑ 25.0 25.1 Sigari, D.; Bourdier, C.; Conti, C.; Conti, J.; Forti, L.; García-Diez, M.; Lai, G.; Mazzini, I. et al. (June 2024). "The last cave lion of the late Upper Palaeolithic: The engraved feline of Grotta Romanelli (southern Italy)" (in en). Quaternary Science Reviews 334. doi:10.1016/j.quascirev.2024.108670. Bibcode: 2024QSRv..33408670S. https://linkinghub.elsevier.com/retrieve/pii/S0277379124001719.
26. ↑ Sommer, R. S.; Benecke, N. (2006). "Late Pleistocene and Holocene development of the felid fauna (Felidae) of Europe: A review". Journal of Zoology 269 (1): 7–19. doi:10.1111/j.1469-7998.2005.00040.x.
27. ↑ Karastoyanova, N., Russeva, V., Georgieva, P. & V. Danov 2026: Bones, bites, and burials: investigating a skeleton from eneolithic necropolis for evidence of probable lion attack in Bulgaria. Journal of Archaeological Science: Reports 69: 105526.
28. ↑ 28.0 28.1 Douglas, N. (1927). Birds and Beasts of Greek Anthology. Florence: Norman Douglas. http://gutenberg.net.au/ebooks03/0300611h.html. Retrieved 2018-08-31.
29. ↑ 29.0 29.1 Alden, M. (2005). "Lions in paradise: Lion Similes in the Iliad and the Lion Cubs of IL. 18.318-22". The Classical Quarterly (55): 335–342. doi:10.1093/cq/bmi035.
30. ↑ 30.0 30.1 Masseti, M.; Mazza, P.P.A. (2013). "Western European Quaternary lions: new working hypotheses". Biological Journal of the Linnean Society 109 (109): 66–77. doi:10.1111/bij.12032.
31. ↑ 31.0 31.1 Guggisberg, C. A. W. (1975). "Lion Panthera leo (Linnaeus, 1758)". Wild Cats of the World. New York: Taplinger Publishing. pp. 138–179. ISBN 978-0-8008-8324-9. https://archive.org/details/wildcatsofworld00gugg/page/138.
32. ↑ Cohen, A. (2010). "Master of Lions (and other Animals)". Art in the era of Alexander the Great: Paradigms of manhood and their cultural traditions. Cambridge: Cambridge University Press. pp. 64–118. ISBN 978-0-5217-6904-4. https://books.google.com/books?id=nX8F_ZV83vUC&pg=PA64.
33. ↑ Heptner, V. G.; Sludskiy, A. A. (1992). "Lion". Mlekopitajuščie Sovetskogo Soiuza. Moskva: Vysšaia Škola. Washington DC: Smithsonian Institution and the National Science Foundation. pp. 83–95. ISBN 978-90-04-08876-4. https://archive.org/stream/mammalsofsov221992gept#page/82/mode/2up.
34. ↑ Russo, G.; Milks, A.; Leder, D.; Koddenberg, T.; Starkovich, B. M.; Duval, M.; Zhao, J.-X.; Darga, R. et al. (2023). "First direct evidence of lion hunting and the early use of a lion pelt by Neanderthals". Scientific Reports 13 (1): 16405. doi:10.1038/s41598-023-42764-0. PMID 37828055. Bibcode: 2023NatSR..1316405R.
35. ↑ Hard, R. (2003). The Routledge Handbook of Greek Mythology: Based on HJ Rose's Handbook of Greek Mythology. London and New York: Routledge.
36. ↑ Stephens, S. Callimachus (2015). Aetia. Dickinson College Commentaries. [1]
37. ↑ Gedoyn, A. 1738: 186 Récits de Conon.
38. ↑ Боев, З. 2016. Левът наш балкански. – Списание Осем, 6: 98-105.
39. ↑ Nowell, K.; Jackson, P. (1996). "Panthera leo". Wild Cats: Status Survey and Conservation Action Plan. Gland, Switzerland: IUCN/SSC Cat Specialist Group. pp. 17–21, 37–41. ISBN 978-2-8317-0045-8. http://carnivoractionplans1.free.fr/wildcats.pdf. Retrieved 2018-02-02.
40. ↑ Chauvet, J.-M.; Brunel, D. E.; Hillaire, C. (1996). Dawn of Art: The Chauvet Cave. The oldest known paintings in the world. New York: Harry N. Abrams. https://archive.org/details/dawnofartchauvet0000chau.

## External links

* Panthera leo europaea
* The Last European Lion

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