Research Article

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

Table 1

2.303 hyena and 1.373 lion bones of Late Pleistocene age from 105 studied cave and open air localities in Central Europe with Ice Age spotted hyena and steppe lion remains (details about the material, locality positions, and housing in collections can be found in the references).

LocalitySite typeAgeHyenas
(NISP)
Lions
(NISP)
Bone assemblage (NISP)References

Ahlen (D)Open air loess, mammoth skeleton scavenging siteWeichselian (glacial)Indirect by scavenging marks0Mammoth skeleton scavenging site[55]
Alfter (D)Open air loess on river terraceWeichselian (glacial)02? unclear[61]
Altlussheim, Almendwiesen (D)Open air loess on river terraceWeichselian (glacial)11562[61]
Altlussheim, Eichelgarten (D)Open air loess on river terraceWeichselian (glacial)01? unclear[61]
Altlussheim, Silzwiesen (D)Open air loess on river terraceWeichselian (glacial)01? unclear[61]
Altrip, Neuhofener Altrhein (D)Open air loess on river terraceWeichselian (glacial)01? unclear[61]
Bad Köstritz (D)Zechstein gypsum karst hyena den
(prey depot and communal den)
Weichselian (glacial)24? unclear[61]
Bad Kösen (D)Open air loessWeichselian (glacial)01? unclear[61]
Bad Lauchstädt (D)Open air loessWeichselian (glacial)06 (from an adult female individual skeleton)? unclear[61]
Bad Wildungen (D)Open air loess on river terraceWeichselian (glacial)11 (and 15 coprolites)0233[49]
Balve Cave (D)Carboniferous limestone cave (cave bear den, hyena cub raising and communal den/overlapping Neanderthal, and Aurignacian camp site)Eemian-Weichselian (interglacial, mainly glacial)3456 (2 skulls)? unclear[42, 65]
Baumann’s Cave (D)Carboniferous limestone cave (cave bear den, short time hyena den, ? temporary wolf den)? Saalian-Eemian, mainly-Weichselian (interglacial, mainly glacial)10148? unclear, half Million cave bear bonesUnpublished
Berlin-Kreuzberg (D)Open air river terraceWeichselian (glacial)01? unclear[61]
Beroun (CZ)Open air river terraceWeichselian (glacial)01 (skull)? unclear[80]
Bilstein Cave (D)Carboniferous limestone cave (cave bear den, short time hyena den, ? temporary wolf den)Weichselian (glacial)239? unclear, many cave bear remains[82]
Brühl (Koller), Schlangenwinkel (D)Open air river terraceWeichselian (glacial)012 bones of a male skeleton? unclear[85]
Brühl, Spieswiesen-Ost (D)Open air river terraceWeichselian (glacial)01? unclear[61]
Brühl, Spieswiesen-West (D)Open air river terraceWeichselian (glacial)8 (2 skulls)41.255[61]
Brühl (Koller), Rheingewann (D)Open air river terraceWeichselian (glacial)13511[61]
Burgtonna (D)TravertinEemian (interglacial)1 (unclear amount)3 (unclear amount)? unclear[61]
Bochum (D)? Open air river terrace? Weichselian (glacial)01 (skull)? unclear[61]
Bottrop (D)Open air river terrace (hyena communal den, partly cub raising den and prey depot)Weichselian (glacial)5026 (and one trackway)?, about 3.000 bones, many of hyena den origin (especially woolly rhinoceros)[48, 61]
Deutmecker Cave (D)Carboniferous limestone cave (hyena den)Weichselian (glacial)20? unclear[65]
Edingen (Brühl), Edinger Ried (D)Open air river terraceWeichselian (glacial)2 (skulls)2 and 8 bones of a male skeleton, including skull1.111[85]
Flörsheim (D)Open air loess on river terraceWeichselian (glacial)10? unclearUnpublished
Fuchsluken Cavities Saalfeld (D)Zechstein limestone karst cavities hyena den (cub raising and communal den)Eemian-Weichselian (interglacial, mainly glacial)21891.035[61]
Freiburg a. d. U. (D)Open air loess on river terrace, most probably Middle Palaeolithic siteWeichselian (glacial)013 (from an adolescent individual skeleton)?[63]
Geddin (D)Open air river terraceWeichselian (glacial)01? unclear[61]
Genther Cave (D)Upper Jurassic dolomite cave (cave bear den)Weichselian (glacial)01 (skull)? unclear[71]
Göttingen (D)Open air river terraceWeichselian (glacial)01? unclear[61]
Grürmanns Cave (D)Carboniferous limestone cave (hyena den)Weichselian (glacial)111? unclear[65]
Halle/Saale (D)Open air loessWeichselian (glacial)01? unclear[50]
Haltern (D)Open air river terraceWeichselian (glacial)01? unclear[61]
Heddesheim, Neuwiesen (D)Open air river terraceWeichselian (glacial)02? unclear[61]
Hermann’s Cave (D)Carboniferous limestone cave (cave bear den)Weichselian (glacial)047 (one skull, two individual skeleton remains)? unclear, many cave bear bones[76]
Herne (D)Open air river terraceWeichselian (glacial)10? unclear[47]
Herten-Stuckenbusch (D)Open air river terraceWeichselian (glacial)02? unclear[61]
Hohenmölsen (D)Open air loessWeichselian (glacial)1 (skull)0? unclear[50]
Hohle Stein Cave (D)Carboniferous limestone cave (hyena prey depot)Weichselian (glacial)1135[65]
Holedeč near Žatec (CZ)?Weichselian (glacial)02? unclear[80]
Hostím (CZ)Open air river terraceWeichselian (glacial)04? unclear[80]
Huttenheim-Huttenheimer Kammer (D)Open air river terraceWeichselian (glacial)03 and 37 bones of one lioness individual skeleton including skull? unclear[85]
Huttenheim, Sandfeld (D)Open air river terraceWeichselian (glacial)01? unclear[61]
Johannes Cave (D)Carboniferous limestone cave (hyena prey depot)Weichselian (glacial)3231[65]
Karlsruhe, Neureut (D)Open air river terraceWeichselian (glacial)01? unclear[61]
Keppler Cave (D)Carboniferous limestone cave (cave bear den)Weichselian (glacial)033? unclear, many cave bear remains[65]
Ketsch, Kreuzwiese (D)Open air river terraceWeichselian (glacial)2 (skulls)1447[61]
Ketsch, Hohwiesen (D)Open air river terraceWeichselian (glacial)02? unclear[61]
Kleinbesten (D)Open air river terraceWeichselian (glacial)01? unclear[61]
Königsaue (D)Open air river terrace (hyena den/overlapping Neanderthal camp site)Weichselian (glacial)13 (skull)2? unclear[50, 61]
Körbisdorf (D)Open air river terraceWeichselian (glacial)04? unclear[61]
Körbiskrug (D)Open air river terraceWeichselian (glacial)01? unclear[61]
Konĕprusy Cave (CZ)Carboniferous limestone cave (hyena prey depot den)Weichselian (glacial)12 (25 coprolites, 114 bones from individual skeleton)23611[38]
Kreuz Cave (D)Carboniferous limestone cave (unclear site, ? cave bear den)? Weichselian (glacial)02? unclear[65]
Lahntal Cave (D)Karst cave (cub raising den)Weichselian (glacial)82013[65]
Lampertheim, In der Tanne (D)Open air river terraceWeichselian (glacial)02? unclear[61]
Lampertheim, Lüderitzbucht (D)Open air river terraceWeichselian (glacial)01? unclear[61]
Lichtenau, Hasenkopf (D)Open air river terraceWeichselian (glacial)01? unclear[61]
Lipperode (D)Open air river terraceWeichselian (glacial)10? unclear[47]
Martins Cave (D)Carboniferous limestone cave (cave bear den, hyena cub raising and communal den, wolf den)Weichselian (glacial)1471716 (real amount unclear), many cave bear remains[65]
Medvedia Cave near Liptovský (Sk)Limestone cave (cave bear den)Weichselian (glacial)0Skeleton of a subadult? unclear[86]
Minice (CZ)?Weichselian (glacial)01? unclear[80]
Mücheln (D)Lake? Eemian/
Weichselian
01? unclear[61]
Neumark-Nord Lake 1 (D)Lake, forest elephants at a graveyard surrounded by Middle Palaeolithic campsEemian (interglacial)6 (4 coprolites)4 (and 172 bones from ill lioness individual skeleton)Many hyena/lion scavenged elephant carcasses[56, 84]
Niederlehme (D)Open air river terraceWeichselian (glacial)17? unclear[61]
Oberneissen Cave (D)Karst cave (cub raising den)Weichselian (glacial)140? unclearUnpublished
Oberröblingen (D)Open air loessWeichselian (glacial)01? unclear[61]
Oberrohn (D)Open air loessWeichselian (glacial)01? unclear[61]
Otterstadt, Altrhein-Süd (D)Open air river terraceWeichselian (glacial)02? unclear[61]
Otterstadt, Altrhein-Nord (D)Open air river terraceWeichselian (glacial)03? unclear[61]
Otterstadt, Waldwiesen (D)Open air river terraceWeichselian (glacial)03? unclear[61]
Osteroden (D)Zechstein gypsum karst hyena den (type unclear)? Weichselian (glacial)10? unclear[50]
Perick Caves (D)Carboniferous limestone cave (hyena cub raising and communal den)Weichselian (glacial)147 (5 skulls)59373[10, 62]
Petershagen (D)Open air river terraceWeichselian (glacial)01Woolly rhinoceros skeleton, unscavanged[61, 88]
Pfefferburg Cave (D)Carboniferous limestone cave (hyena den, cave bear den, ? temporary wolf den)Weichselian (glacial)37 (and 8 coprolites)231 (real amount ? unclear)[65]
Phoeben (D)Open air river terraceWeichselian (glacial)01? unclear[61]
Praha-Podbaba (CZ)Open air river terraceWeichselian (glacial)016 (from two individual skeletons)? unclear[80]
Praha-Košíře (CZ)Open air river terraceWeichselian (glacial)01? unclear[80]
Praha-Libeň (CZ)Open air river terraceWeichselian (glacial)06? unclear[80]
Questenberg (D)? unclear? Weichselian (glacial)10? unclear[50]
Rösenbeck Cave (D)Carboniferous limestone cave (hyena communal and cave bear den)Weichselian (glacial)83 (5 skulls)0? unclear, many cave bear remains[17]
Selm-Ternsche (D)Open air river terraceWeichselian (glacial)30? unclear[47]
Siegsdorf (D)River branch, river terrace siteWeichselian (glacial)(3 coprolites)62 bones of one male individual skeleton including skullMammoth bull carcass scavenging site[75]
Speyer, Deutschof (D)Open air river terraceWeichselian (glacial)01? unclear[61]
Svobodné Dvory near Hradec Králové (CZ)? open airWeichselian (glacial)01? unclear[80]
Sewecken-Berge Quedlinburg (D)Triassic gypsum karst hyena den
(prey depot and communal den)
Eemian-Weichselian (interglacial, mainly glacial)115 (1 skull of cub)17660[52, 61]
Senzig (D)Open air river terrace? Weichselian (glacial)01? unclear[61]
Sloup Cave (CZ)Carboniferous limestone cave (hyena den, cave bear den, ? wolf den)Weichselian (glacial)85 (2 skulls)452[40]
Sophie’s Cave (D)Jurassic limestone cave (hyena cub raising and communal, cave bar, and wolf den)Weichselian (glacial)126, many cave bear remains (often carnivore damaged)[45]
Speyer, Binsfeld-Südost (D)Open air river terraceWeichselian (glacial)01? unclear[61]
Srbsko Chlum-Komín 
Cave (CZ)
Carboniferous limestone cave (hyena prey depot and commuting den)Weichselian (glacial)366 (from 6 individual skeletons, including cubs, 4 coprolites)Two individual skeletons: 149 bones from an adolescent ill lioness, and 107 bones of a juvenile2.947 (mainly Przewalski horse remains)[39]
Teufelskammer Cave (D)Carboniferous limestone cave (hyena den, cave bear den, ? temporary wolf den)Weichselian (glacial)71197[41]
Thiede (D)Zechstein gypsum karst hyena den
(prey depot and communal den)
Weichselian (glacial)30 (1 skull of adult)14120 (even more, unclear amount at this stage)[67]
Türmitz (CZ)Open air loessWeichselian (glacial)2 (1 skull)2? unclear[17]
stí nad Labem (CZ)Open air river terraceWeichselian (glacial)01? unclear[80]
Výpustek Cave (CZ)Carboniferous limestone cave (hyena den, cave bear den, ? temporary wolf den)? Eemian/
Weichselian (interglacial/
glacial)
236 (3 skulls, one individual skeleton)36? unclearUnpublished, skeleton in [39]
Wanne (D)Open air river terraceWeichselian (glacial)01? unclear[61]
Westeregeln (D)Zechstein gypsum karst hyena den
(prey depot and communal den) and overlapping Neanderthal camp site
Weichselian (glacial)84 (10 skulls, and 20 coprolites)12488[51, 61]
Wildkirchli Cave (Ch)Limestone cave (cave bear den)Weichselian (glacial)023?, many cave bear remainsUnpublished
Wiesental, Allmendweg (D)Open air river terraceWeichselian (glacial)01? unclear[61]
Wiesental, Viehweg (D)Open air river terraceWeichselian (glacial)01? unclear[61]
Wilhelms Cave (D)Carboniferous limestone cave (hyena cub raising and communal den)Weichselian (glacial)16915 (sibling skeleton remain)126[65]
Wörth, Geisbögel (D)Open air river terraceWeichselian (glacial)01? unclear[61]
Wörth, Rheinanlagen (D)Open air river terraceWeichselian (glacial)01? unclear[61]
Wüste Scheuer Abri (D)Zechstein limestone abri/small cavityWeichselian (glacial)27 (1 skull)061[54]
Zechovice near Volyně (CZ)? open air LoessWeichselian (glacial)01? unclear[80]
Zoolithen Cave (D)Jurassic limestone cave (hyena cub raising and communal, cave bar, and wolf den)Saalian to Eemian/
Weichselian (interglacial/
glacial)
207 (2 skulls)229 (9 skulls)Few bones, mainly damaged cave bear bones in unclear amount (about half Million cave bear bones)[43, 44]