穴狮

穴狮; 化石时期：中－晚更新世 0.6–0.013 Ma PreЄ Є O S D C P T J K Pg N ▼
	全身骨架，展于维也纳自然史博物馆
科学分类
界：	动物界 Animalia
门：	脊索动物门 Chordata
纲：	哺乳纲 Mammalia
目：	食肉目 Carnivora
科：	猫科 Felidae
属：	豹属 Panthera
种：	†穴狮 P. spelaea
二名法
	†Panthera spelaea; Goldfuss, 1810
亚种
	白令穴狮 Panthera spelaea vereshchagini (Baryshnikov & Boeskorov, 2001); 欧亚穴狮 Panthera spelaea spelaea Goldfuss, 1810;
	穴狮分布区域（红色）; 美洲拟狮分布区域（蓝色）; 狮分布区域（绿色）
异名
	Panthera leo spelaea M. Boule & L. De Villeneuve, 1927;

穴狮（学名：Panthera spelaea）又称洞狮、洞穴狮，是一种已经灭绝的豹属物种，生存于更新世时期的欧亚大陆至北美洲西北部，在第三次克罗默尔间冰期（英语：Cromerian interglacial）于欧洲出现，少于 600,000 年前。它们与现代狮子拥有同样的祖先，是栖息于一些开阔的环境如猛犸草原（英语：mammoth steppe）或是北方针叶林的肉食性猛兽，主要以驯鹿及其他草食动物为食。穴狮在1万3千年前灭绝^[1]。

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分类

穴狮（Panthera spelaea）本来被视为是现代狮的亚种，因此原被命名为 Panthera leo spelaea^[2]^[3]^[4]^[5]。然而，颅骨与齿式解剖学上的差异足以将穴狮独立为一新种^[6]^[7]系统发生树上的研究成果同样也支持这项理论^[8]^[9]^[10]。

演化

穴狮在莎勒冰期（英语：Saalian glaciation）晚期或艾木间冰期（英语：Eemian）早期时演化自原始狮（原始狮则是出现于 700,000 年前的欧亚大陆）^[5]。在波兰发现的穴狮骨头碎片经放射性碳定年法判定年代大约于魏奇塞冰期（英语：Weichselian glaciation）的早期至晚期，大约是 109,000 至 57,000 年前^[12]。欧亚大陆上的穴狮大约于 14,900 至 14,100 年前绝种。白令陆桥上的穴狮则在 13,800 至 13,300 年前魏奇塞冰期结束时绝种^[1]。

由化石残骸上的线粒体DNA分析显示美洲拟狮为穴狮的姐妹群，约由 340,000 年前被困在柯迪勒拉冰盖（英语：Cordilleran Ice Sheet）南方的穴狮族群分支演化而来^[9]。下面的支序分类表示了穴狮与其他豹属物种的关系^[10]：

猫 Felis silvestris catus

云豹 Neofelis nebulosa

	苏门答腊虎 Panthera tigris sumatrae

	孟加拉虎 Panthera tigris tigris

美洲豹 Panthera onca

雪豹 Panthera uncia

	豹 Panthera pardus

	远东豹 Panthera pardus japonensis

	狮 Panthera leo

	亚洲狮 Panthera leo persica

	†穴狮 Panthera spelaea

	†美洲拟狮 Panthera atrox

描述

穴狮时在旧石器时代常以石洞壁画、象牙雕刻、黏土塑像的形式呈现，这些遗物代表着穴狮也可能出现于旧石器时代的宗教仪式当中^[13]^[14]。在法国肖维岩洞（约 30,000 年前）描绘了一对走在一起的穴狮，其中一只的体型略小于另一只，而体型较大的个体则绘有阴囊，但不具有鬃毛^[15]。这代表穴狮和现代狮子不一样，牠们可能不具有明显的鬃毛^[1]。

在被独立为一种时，穴狮曾是狮子所有亚种中体格最大的之一，1985 年在德国锡格斯多夫附近发现的一只成年雄性洞穴狮化石表明，其及肩身高约1.2米（3.9英尺），身长2.1米（6.9英尺）（不含尾巴），这大致相当于一头体型相当大的现代狮子。实际上，这只洞穴狮并不是同类中最大的，因此可以推断，洞穴狮要比现代狮子大5%到10%，但比较早期的原始狮（Panthera fossilis）还有之后的美洲拟狮小^[16]。和现代狮子一样，雌狮的体型较雄狮小^[17]

古生态学

穴狮生活在一些开阔的环境如猛犸草原（英语：mammoth steppe）或是北方针叶林。牠们的骨骼多半与洞鬣狗、洞熊和旧石器时代文物一同在洞穴中发现因而得名^[19]^[20]。穴狮可能为独居性的掠食者^[21]。

从欧洲^[22]^[23]及白令陆桥东部^[24]所发现化石中胶原蛋白样本的同位素分析显示，在末次冰期时，驯鹿是穴狮的主要食物来源^[23]。而当有机会时，牠们也会狩猎捕食洞熊的幼崽^[23]^[22]，其他可能的猎物包括大角鹿、欧洲马鹿、野马、麝牛、原牛、欧洲野牛、西伯利亚野牛、幼年披毛犀以及幼年猛犸象。与穴狮共同竞争这些猎物的掠食者可能包括欧亚大陆上的穴花豹（英语：Panthera pardus spelaea）、洞鬣狗、棕熊与灰狼^[25]；以及白令陆桥地区的短面熊、似剑齿虎与白令狼（英语：Beringian wolf）^[23]。

参见

参考来源

[1]
Stuart, A. J., Lister, A. M. Extinction chronology of the cave lion Panthera spelaea. Quaternary Science Reviews. 2011, 30 (17): 2329–2340. doi:10.1016/j.quascirev.2010.04.023.
[2]
Kurtén, B., 1968. Pleistocene Mammals of Europe. Weidenfeld and Nicolson, London.
[3]
Hemmer, H. Untersuchungen zur Stammesgeschichte der Pantherkatzen (Pantherinae) Teil 3. Zur Artgeschichte des Löwen Panthera (Panthera) leo (Linnaeus, 1758). Veröffentlichungen der Zoologischen Staatssammlung 17. 1974: 167–280.
[4]
Turner, A. Dental sex dimorphism in European lions (Panthera leo L.) of the Upper Pleistocene: palaeoecological and palaeoethological implications. Annales Zoologici Fennici. 1984, 21: 1–8.
[5]
Burger, J.; Rosendahl, W.; Loreille, O.; Hemmer, H.; Eriksson, T.; Götherström, A.; Hiller, J.; Collins, M. J.; Wess, T.; Alt, K. W. Molecular phylogeny of the extinct cave lion Panthera leo spelaea. Molecular Phylogenetics and Evolution. 2004, 30 (3): 841–849 [2011-12-17]. PMID 15012963. doi:10.1016/j.ympev.2003.07.020. （原始内容存档于2019-06-26）.
[6]
Spassov, N., Iliev, N. Animal remains from the submerged Late Eneolithic – early Bronze Age settlements in Sozopol (South Bulgarian Black Sea Coast). Proceedings of the International Symposium VI. Thracia Pontica. 1994: 287–314.
[7]
Sotnikova, M., Nikolskiy, P. Systematic position of the cave lion Panthera spelaea (Goldfuss) based on cranial and dental characters. Quaternary International. 2006,. 142-143: 218–228. doi:10.1016/j.quaint.2005.03.019.
[8]
Christiansen, P. Phylogeny of the great cats (Felidae: Pantherinae), and the influence of fossil taxa and missing characters. Cladistics. 2008, 24 (6): 977–992. doi:10.1111/j.1096-0031.2008.00226.x.
[9]
Barnett, R., Shapiro, B., Barnes, I. A. N., Ho, S. Y., Burger, J., Yamaguchi, N., Higham, T. F., Wheeler, H., Rosendahl, W., Sher, A. V., Sotnikova, M. Phylogeography of lions (Panthera leo ssp.) reveals three distinct taxa and a late Pleistocene reduction in genetic diversity (PDF). Molecular Ecology. 2009, 18 (8): 1668–1677. PMID 19302360. doi:10.1111/j.1365-294X.2009.04134.x. （原始内容 (PDF)存档于21 January 2012）.
[10]
Barnett, R.; Mendoza, M. L. Z.; Soares, A. E. R.; Ho, S. Y. W.; Zazula, G.; Yamaguchi, N.; Shapiro, B.; Kirillova, I. V.; Larson, G.; Gilbert, M. T. P. Mitogenomics of the Extinct Cave Lion, Panthera spelaea (Goldfuss, 1810), Resolve its Position within the Panthera Cats. Open Quaternary. 2016, 2: 4. doi:10.5334/oq.24.
[11]
Chauvet, J.-M.; Brunel, D. E.; Hillaire, C. Dawn of Art: The Chauvet Cave. The oldest known paintings in the world. New York: Harry N. Abrams. 1996.
[12]
Marciszak, A., and Stefaniak, K. Two forms of cave lion: Middle Pleistocene Panthera spelaea fossilis Reichenau, 1906 and Upper Pleistocene Panthera spelaea spelaea Goldfuss, 1810 from the Bisnik Cave, Poland. Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen. 2010, 258 (3): 339–351. doi:10.1127/0077-7749/2010/0117.
[13]
Bahn, P. G., Vertut, J. Journey Through the Ice Age. London: Weidenfeld & Nicolson. 1997.
[14]
Guthrie, R. D. The Nature of Paleolithic Art. Chicago: University of Chicago Press. 2005.
[15]
Yamaguchi, N.; Cooper, A.; Werdelin, L.; MacDonald, D. W. Evolution of the mane and group-living in the lion (Panthera leo): a review. Journal of Zoology. 2004, 263 (4): 329–342. doi:10.1017/S0952836904005242.
[16]
Koenigswald, W. v. (2002). Lebendige Eiszeit. Darmstadt: Theiss-Wissenschaftliche Buchgemeinschaft. ISBN 3-8062-1734-3
[17]
Diedrich, C. G. Late Pleistocene Panthera leo spelaea (Goldfuss, 1810) skeletons from the Czech Republic (central Europe); their pathological cranial features and injuries resulting from intraspecific fights, conflicts with hyenas, and attacks on cave bears. Bulletin of Geosciences. 2011, 86 (4): 817–840. doi:10.3140/bull.geosci.1263.
[18]
Bölsche, W. & Harder, H. Tiere der Urwelt. Serie III. Wandsbek-Hamburg: Verlag der Kakao-Compagnie Theodor Reichardt. 1900.
[19]
Diedrich, C. G. The largest European lion Panthera leo spelaea (Goldfuss 1810) population from the Zoolithen Cave, Germany: specialised cave bear predators of Europe. Historical Biology. 2011, 23 (2–3): 271–311. doi:10.1080/08912963.2010.546529.
[20]
Diedrich, C. G. Pleistocene Panthera leo spelaea (Goldfuss 1810) remains from the Balve cave (NW Germany) - a cave bear, hyena den and middle palaeolithic human cave – and review of the Sauerland Karst lion cave sites. Quaternaire. 2011, 22 (2): 105–127.
[21]
Bocherens, H. Isotopic tracking of large carnivore palaeoecology in the mammoth steppe. Quaternary Science Reviews. 2015, 117: 42–71. doi:10.1016/j.quascirev.2015.03.018.
[22]
Bocherens, H.; Drucker, D. G.; Bonjean, D.; Bridault, A.; Conard, N. J.; Cupillard, C.; Germonpré, M.; Höneisen, M.; Münzel, S. C.; Napierala, H. & Patou-Mathis, M. Isotopic evidence for dietary ecology of cave lion (Panthera spelaea) in North-Western Europe: prey choice, competition and implications for extinction (PDF). Quaternary International. 2011, 245 (2): 249–261 [18 September 2020]. Bibcode:2011QuInt.245..249B. S2CID 129706533. doi:10.1016/j.quaint.2011.02.023. （原始内容存档 (PDF)于11 May 2020）.
[23]
Bocherens, H. Isotopic tracking of large carnivore palaeoecology in the mammoth steppe. Quaternary Science Reviews. 2015, 117: 42–71. Bibcode:2015QSRv..117...42B. doi:10.1016/j.quascirev.2015.03.018.
[24]
Drucker, Dorothée G. The Isotopic Ecology of the Mammoth Steppe. Annual Review of Earth and Planetary Sciences. 2022-05-31, 50 (1): 395–418. ISSN 0084-6597. doi:10.1146/annurev-earth-100821-081832 （英语）.
[25]
Diedrich, C. G. Late Pleistocene leopards across Europe – northernmost European German population, highest elevated records in the Swiss Alps, complete skeletons in the Bosnia Herzegowina Dinarids and comparison to the Ice Age cave art. Quaternary Science Reviews. 2013, 76: 167–193. Bibcode:2013QSRv...76..167D. doi:10.1016/j.quascirev.2013.05.009.

外部链接

[Stuart2011-1] [1]
Stuart, A. J., Lister, A. M. Extinction chronology of the cave lion Panthera spelaea. Quaternary Science Reviews. 2011, 30 (17): 2329–2340. doi:10.1016/j.quascirev.2010.04.023.

[2] [2]
Kurtén, B., 1968. Pleistocene Mammals of Europe. Weidenfeld and Nicolson, London.

[Hemmer-3] [3]
Hemmer, H. Untersuchungen zur Stammesgeschichte der Pantherkatzen (Pantherinae) Teil 3. Zur Artgeschichte des Löwen Panthera (Panthera) leo (Linnaeus, 1758). Veröffentlichungen der Zoologischen Staatssammlung 17. 1974: 167–280.

[4] [4]
Turner, A. Dental sex dimorphism in European lions (Panthera leo L.) of the Upper Pleistocene: palaeoecological and palaeoethological implications. Annales Zoologici Fennici. 1984, 21: 1–8.

[Burger-5] [5]
Burger, J.; Rosendahl, W.; Loreille, O.; Hemmer, H.; Eriksson, T.; Götherström, A.; Hiller, J.; Collins, M. J.; Wess, T.; Alt, K. W. Molecular phylogeny of the extinct cave lion Panthera leo spelaea. Molecular Phylogenetics and Evolution. 2004, 30 (3): 841–849 [2011-12-17]. PMID 15012963. doi:10.1016/j.ympev.2003.07.020. （原始内容存档于2019-06-26）.

[6] [6]
Spassov, N., Iliev, N. Animal remains from the submerged Late Eneolithic – early Bronze Age settlements in Sozopol (South Bulgarian Black Sea Coast). Proceedings of the International Symposium VI. Thracia Pontica. 1994: 287–314.

[7] [7]
Sotnikova, M., Nikolskiy, P. Systematic position of the cave lion Panthera spelaea (Goldfuss) based on cranial and dental characters. Quaternary International. 2006,. 142-143: 218–228. doi:10.1016/j.quaint.2005.03.019.

[8] [8]
Christiansen, P. Phylogeny of the great cats (Felidae: Pantherinae), and the influence of fossil taxa and missing characters. Cladistics. 2008, 24 (6): 977–992. doi:10.1111/j.1096-0031.2008.00226.x.

[Barnett2009-9] [9]
Barnett, R., Shapiro, B., Barnes, I. A. N., Ho, S. Y., Burger, J., Yamaguchi, N., Higham, T. F., Wheeler, H., Rosendahl, W., Sher, A. V., Sotnikova, M. Phylogeography of lions (Panthera leo ssp.) reveals three distinct taxa and a late Pleistocene reduction in genetic diversity (PDF). Molecular Ecology. 2009, 18 (8): 1668–1677. PMID 19302360. doi:10.1111/j.1365-294X.2009.04134.x. （原始内容 (PDF)存档于21 January 2012）.

[Barnett2016-10] [10]
Barnett, R.; Mendoza, M. L. Z.; Soares, A. E. R.; Ho, S. Y. W.; Zazula, G.; Yamaguchi, N.; Shapiro, B.; Kirillova, I. V.; Larson, G.; Gilbert, M. T. P. Mitogenomics of the Extinct Cave Lion, Panthera spelaea (Goldfuss, 1810), Resolve its Position within the Panthera Cats. Open Quaternary. 2016, 2: 4. doi:10.5334/oq.24.

[Chauvet1996-11] [11]
Chauvet, J.-M.; Brunel, D. E.; Hillaire, C. Dawn of Art: The Chauvet Cave. The oldest known paintings in the world. New York: Harry N. Abrams. 1996.

[Marciszak2010-12] [12]
Marciszak, A., and Stefaniak, K. Two forms of cave lion: Middle Pleistocene Panthera spelaea fossilis Reichenau, 1906 and Upper Pleistocene Panthera spelaea spelaea Goldfuss, 1810 from the Bisnik Cave, Poland. Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen. 2010, 258 (3): 339–351. doi:10.1127/0077-7749/2010/0117.

[13] [13]
Bahn, P. G., Vertut, J. Journey Through the Ice Age. London: Weidenfeld & Nicolson. 1997.

[14] [14]
Guthrie, R. D. The Nature of Paleolithic Art. Chicago: University of Chicago Press. 2005.

[Yamaguchietal2004-15] [15]
Yamaguchi, N.; Cooper, A.; Werdelin, L.; MacDonald, D. W. Evolution of the mane and group-living in the lion (Panthera leo): a review. Journal of Zoology. 2004, 263 (4): 329–342. doi:10.1017/S0952836904005242.

[16] [16]
Koenigswald, W. v. (2002). Lebendige Eiszeit. Darmstadt: Theiss-Wissenschaftliche Buchgemeinschaft. ISBN 3-8062-1734-3

[17] [17]
Diedrich, C. G. Late Pleistocene Panthera leo spelaea (Goldfuss, 1810) skeletons from the Czech Republic (central Europe); their pathological cranial features and injuries resulting from intraspecific fights, conflicts with hyenas, and attacks on cave bears. Bulletin of Geosciences. 2011, 86 (4): 817–840. doi:10.3140/bull.geosci.1263.

[18] [18]
Bölsche, W. & Harder, H. Tiere der Urwelt. Serie III. Wandsbek-Hamburg: Verlag der Kakao-Compagnie Theodor Reichardt. 1900.

[19] [19]
Diedrich, C. G. The largest European lion Panthera leo spelaea (Goldfuss 1810) population from the Zoolithen Cave, Germany: specialised cave bear predators of Europe. Historical Biology. 2011, 23 (2–3): 271–311. doi:10.1080/08912963.2010.546529.

[20] [20]
Diedrich, C. G. Pleistocene Panthera leo spelaea (Goldfuss 1810) remains from the Balve cave (NW Germany) - a cave bear, hyena den and middle palaeolithic human cave – and review of the Sauerland Karst lion cave sites. Quaternaire. 2011, 22 (2): 105–127.

[21] [21]
Bocherens, H. Isotopic tracking of large carnivore palaeoecology in the mammoth steppe. Quaternary Science Reviews. 2015, 117: 42–71. doi:10.1016/j.quascirev.2015.03.018.

[Bocherens-22] [22]
Bocherens, H.; Drucker, D. G.; Bonjean, D.; Bridault, A.; Conard, N. J.; Cupillard, C.; Germonpré, M.; Höneisen, M.; Münzel, S. C.; Napierala, H. & Patou-Mathis, M. Isotopic evidence for dietary ecology of cave lion (Panthera spelaea) in North-Western Europe: prey choice, competition and implications for extinction (PDF). Quaternary International. 2011, 245 (2): 249–261 [18 September 2020]. Bibcode:2011QuInt.245..249B. S2CID 129706533. doi:10.1016/j.quaint.2011.02.023. （原始内容存档 (PDF)于11 May 2020）.

[Bocherens2015-23] [23]
Bocherens, H. Isotopic tracking of large carnivore palaeoecology in the mammoth steppe. Quaternary Science Reviews. 2015, 117: 42–71. Bibcode:2015QSRv..117...42B. doi:10.1016/j.quascirev.2015.03.018.

[24] [24]
Drucker, Dorothée G. The Isotopic Ecology of the Mammoth Steppe. Annual Review of Earth and Planetary Sciences. 2022-05-31, 50 (1): 395–418. ISSN 0084-6597. doi:10.1146/annurev-earth-100821-081832 （英语）.

[Diedrich_2013-25] [25]
Diedrich, C. G. Late Pleistocene leopards across Europe – northernmost European German population, highest elevated records in the Swiss Alps, complete skeletons in the Bosnia Herzegowina Dinarids and comparison to the Ice Age cave art. Quaternary Science Reviews. 2013, 76: 167–193. Bibcode:2013QSRv...76..167D. doi:10.1016/j.quascirev.2013.05.009.

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