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Research Article Palaeopopulations of Late Pleistocene Top Predators in Europe: Ice Age Spotted Hyenas and Steppe Lions in Battle and Competition About Prey

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Research Article Palaeopopulations of Late Pleistocene Top Predators in Europe: Ice Age Spotted Hyenas and Steppe Lions in Battle and Competition About Prey Hindawi Publishing Corporation Paleontology Journal Volume 2014, Article ID 106203, 34 pages http://dx.doi.org/10.1155/2014/106203 Research Article Palaeopopulations of Late Pleistocene Top Predators in Europe: Ice Age Spotted Hyenas and Steppe Lions in Battle and Competition about Prey Cajus G. Diedrich Private Research Institute Paleologic, Petra Bezruce 96,26751 Zdice, Czech Republic Correspondence should be addressed to Cajus G. Diedrich; [email protected] Received 21March 2013; Accepted 11June 2013 Academic Editor: Vlad Codrea Copyright © 2014 Cajus G. Diedrich. Tis is an open access article distributed under the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. Late Pleistocene spotted hyena Crocuta crocuta spelaea (Goldfuss, 1823) and steppe lion Panthera leo spelaea (Goldfuss, 1810) were top predators in Central Europe. Te fossil record (2.303 hyena/1.373 lion bones = ratio 3/1) from 106 cave and open air sites demonstrates comparable associations to modern African hyenas/lions resulting in competition about prey and territory. Cannibalism within extinct spotted hyenas is well documented, including two individual skeletons. Tose hyenas produced bone accumulations at dens. Feeding specializations on diferent megamammal groups are demonstrated for Late Pleistocene hyenas whose prey partly overlaps (e.g., cave bears) with those of lions and wolves. At most fossil sites, 1–3% of the lion remains indicate scavenging of lions by hyenas. Te larger Late Pleistocene felids focussed on cervids (reindeers specialization during the high glacial = LGM), on bovids (steppe bison/aurochs), and possibly on saiga antelope and on the cave bear, hunting deep in caves during their hibernations and targeting cubs. Te cave bear feeding was the target of all three top predators (lions, hyenas, and wolves) in the Late Pleistocene boreal forests which caused deathly conficts in caves between them, especially with lions/hyenas and herbivorous cave bears that have no modern analogue. 1. Introduction more detail afer their rediscovery in 2009 [8]. Finally, also the “cave wolf Canis spelaeus”wasdescribedin1823basedon Pleistocene top predator research started in Europe with the a cub skull found also in the Zoolithen Cave [7]. frst discoveries of “foreign animal fnds” in the Zoolithen Te modern era of “cave hyena” den research started in Cave (Germany) in 1774 when the German Priest Esper 1823, when the English Geologist Buckland published his discovered some hyena (Figures 1–3), lion (Figures 4–6), “Reliquiae Diluvianae” [9]. Modern hyena den interpreta- wolf, and cave bear remains and explained them resulting tion from historic times was based on mainly the famous from the “great deluge” [1]. Rosenmuller¨ collected a large Kent’s Caverns (E) and the Konig-Ludwigs¨ Cave (D, there amount of cave bear remains including the Ursus spelaeus mentioned as “Kuhstall or Rabenstein Cave”) [9].Te Ger- holotype skull [2]andthelargestamountoflionmaterialin man Biologist Giebel excavated hyena [10]andotherfaunal the same cave [3].Tis collection survived until today [2]. remains from Perick Caves (= Sundwig Cave, Sauerland Te French zoologist Cuvier interpreted in 1805 [4]some Karst) in 1849–1852 in Germany and hyena and prey fauna skull fragments from the German Zoolithen Cave as “hyena”. remains from the Sewecken-Berge and Westeregeln (D) [11] It was the German Paleontologist Goldfuss who described open air gypsum karst areas of northern-central Germany. In and named the frst top predator of the Pleistocene with the 1863, Dawkins discovered the hyena bone-rich (467remains, holotype skull of the “cave lion Felis spelaea”in1810[5, 6]. mainly teeth) and Neanderthal artefact bearing (e.g., bifaces) In 1823 he published on the second large predator with the Wookey Hole Cave hyena den and overlapping Pleistocene holotype skull of the “cave hyena Hyaena spelaea”[7]. Both Neanderthal human camp site (E) [12, 13]. Den research of these skulls from the Zoolithen Cave were described in continued with the descriptions of the faunas of famous hyena 2 Paleontology Journal (a) Late Pleistocene (113,000-24,000 BP) Crocuta crocuta spelaea Crocuta crocuta crocuta Flat Sligth convex Strong convex (b)Glacier Glaciers (c) Glacier Hyena absence- small cave bears Oslo Oslo Skull shape types Crocuta crocuta spelaea Moskow Skeletons, skeleton remains Pin Hole Tiede Cave (GB) Bottrop gypsum karst (D) Pin Hole Open air (D) Königsaue Cave (GB) Westeregeln open air (D) Sandfort Hill gypsum karst (D) Westeregeln Perick Cave (GB) Warsaw Hohenmölsen Wookey Hole Perick (D) Caves (D) open air (D) Berlin Lindenthal Berlin Cave (GB) Caves (D) Cave (D) Warzaw London Amsterdam Pössneck Bleadon Amsterdam Teufelskammer Cavity (D) Cave (GB) Cave (D) Townton London Trmice open air (CZ) Cave (GB) Srbsko Cave (CZ) Bad Wildungen Hutton open air (D) Sloup Cave (CZ) Cave (GB) Sophie's Výpustek Cave (CZ) Sandfort Hill Cave (D) Sloup Cave (CZ) Cave (GB) Zoolithen Výpustek Cave (CZ) Carpathians Cave (D) Carpathians Zoolithen Koneprusy Srbsko Brühl-Spielewiesen Paris Cave (D) Cave (CZ) Cave (CZ) Paris open air (D) Javorka Cave (CZ) Ketsch-Kreuzwiese Munich Munich open air (D) Main study area Irpfel Budapest Erkenbrechtsweiler Plumettes limestone karst (D) Cave (D) Budapest Cave (F) Badel Cave (A) Rochelot Alps Oase Alps Cave (F) Cave (Ro) Oase Cave (Ro) Bukarest Bukarest Trou du Cluzeau Cave (F) Dinarids Dinarids Agios Georgios Cave (Gr) Rome Rome Barcelona Barcelona Madrid Madrid Mountains with few vegetation Boreal conifere forests Taiga steppe Mixed forests Athene Athene Tundra Temperate broadleaf forests San Teodoro Cave (It) Forest tundra Mediterranean forests 500 km 500 km (d) Výpustek Cave (CZ) Srbsko Chlum-Komín Cave (CZ) Koneprusy Cave (CZ) Figure 1: (a) Spotted hyenas globally during the Late Pleistocene. ((b) and (c)) Ice Age spotted hyena Crocuta crocuta spelaea (Goldfuss, 1823) sites in Europe during the Late Pleistocene before the Last Glacial Maximum (113.000–24.000 BP) with absence in alpine regions (= red areas). (d) Skeletons from Europe: adult individual skeleton from Vypustek´ Cave (Moravian Karst, CZ, coll. NHMW); composed cub skeleton from Srbsko Chlum-Kom´ın Cave (Bohemian Karst, coll. NHMP); adult individual skeleton from Koneprusy˘ Cave (Bohemian Karst, CZ, coll. NHMP). Paleontology Journal 3 den cave sites such as the Lindental Cave Gera (D) in 1867, (teeth and crushed longbone fragments) were found within where phosphatic excrement layers were reported and where the bovid/equid/suid dominated prey bone accumulation fractured rhinoceros bones resulting from hyenas have been [34]. Te most eastern known Late Pleistocene hyena den misidentifed as “Pleistocene Neanderthal tools” [14].Tis was reported in 2000 from the Prolom II Cave [35]in small cave also had artefacts from Neanderthals, which lead the Crimea, where there are hyena dens overlapping with to historical conclusions that these nontool bones must be human camp sites at cave entrances/portals. Te overlapping the “kitchen rubbish” of Pleistocene humans who “chewed on ofen does not enable clear attribution to hyena prey or the bones” leaving bite marks (= “Osteodontokeratic culture” human kitchen rubbish. Interestingly, other hyena dens with [15]). Nehring continued the German Late Pleistocene hyena possibly overlapping wolf dens were published for some Late research and collected some skulls and several postcranial Pleistocene Italian caves in 2004 [36]. Most probably instead hyena remains and prey bones from a new open air gypsum the wolves were also prey and imported by hyenas, which karst den site Tiede (D) [16]andattheRosenbeck¨ Cave (D) is unsolved yet. Te San Teodoro Cave on Sicily (I) was [17]. In 1892 Haarlestartedthe“hyenaden”researchinSE-´ then presented in 2011, demonstrating the low marine sea France [18]. Reynolds published in 1902 [19]byfarthebest level during the glacial of the Late Pleistocene which allowed monograph on the English Late Pleistocene hyena remains the migration of hyenas to the island of present day [37]. of its time, but he confused some material as “individual Currant compiled several forgotten and new hyena dens skeletons” which are actually from at least two diferent caves, from England in 2004, including the rediscovered Reynolds Wookey Hole [19, 20]andSandfortHill[19, 20]Caves(E). collections from World War II describing further caves as Two rediscovered skulls [17]fromthesetwolocalitiesanda hyena dens such as Bleadon, Cavern, or Hutton Cavern large collection which was hidden during the second World [20]. A skull from Pin Hole [17] was added for English War under coals in the Somerset Museum [20]havenot sites. In 2008 Polish hyena remains were compiled [38]. In been analysed yet at present. Te Teufelslucken Cave (A) the Czech Republic in 2004, the author’s “European Late was another hyena cave den and the frst described from Pleistocene spotted hyena project” provided an overview of Austria in 1937 [21]. Zapfe presented in 1939 a paper on Late Pleistocene hyena den types, mainly caves, which was damaged bones by hyenas from diferent cave and open air presented for the Bohemian Karst mountainous region (= sites, especially from Austria [22]. Afer the Second World between 150 and 650 a.s.l.) near Prague [39]. In addition War,newLatePleistocenehyenadenswerediscoveredin1957 the most famous Czech Late Pleistocene caves and hyena with the Torbryan Cave and Tonewton Cave (E) [20, 23].
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