631:
111:
149:
128:
651:, or humans. In European Ice Age caves, leopard bones are far rarer than those of lions, and all currently known fossils belong to adults, suggesting that they rarely, if ever, raised their cubs in caves. Where leopard remains are found in larger caves, they are often found in the cave's deeper recesses, as in Baumann's and
671:. It is likely that leopards scavenged or occasionally killed cave bears during hibernation in their dens. During the cold phases, European Ice Age leopards occurred mainly in mountain or alpine boreal forests or in mountains above the treeline, and were not usually found in the lowland mammoth steppes.
613:
The site of Equi in northwestern Italy represents the richest concentration of leopard remains from
Pleistocene Europe, with some 200 leopards having been excavated. The youngest reliable records for leopards outside of eastern Europe are from the Iberian Peninsula, around 17-11,000 years ago, with
638:
Fossils of
European Ice Age leopards in Europe are sometimes found in caves, where they apparently sought shelter or hid their prey. They generally preferred smaller caves, most likely because larger caves were usually occupied by larger predators such as
855:
Paijmans, Johanna L. A.; Barlow, Axel; Förster, Daniel W.; Henneberger, Kirstin; Meyer, Matthias; Nickel, Birgit; Nagel, Doris; Worsøe Havmøller, Rasmus; Baryshnikov, Gennady F.; Joger, Ulrich; Rosendahl, Wilfried; Hofreiter, Michael (December 2018).
712:
Diedrich, C. G. (2013). "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".
516:
The timing of arrival of leopards in Europe is disputed. Some authors have posited that they arrived in Europe during the late Early
Pleistocene around 1.2-1.1 million years ago. while others have suggested that they arrived during the early
469:
Mitochondrial genomes from Late
Pleistocene European leopard specimens found in Germany suggests that they represent a distinct group of extinct leopards that diveged from the common ancestor of Asian leopards (including those from the
1159:
Stein, A.B.; Athreya, V.; Gerngross, P.; Balme, G.; Henschel, P.; Karanth, U.; Miquelle, D.; Rostro-Garcia, S.; Kamler, J. F.; Laguardia, A.; Khorozyan, I. & Ghoddousi, A. (2020) .
602:
in southern France is dated to about 25,000–37,500 years old. The last
European Ice Age leopards vanished from most parts of Europe about 24,000 years ago, just before the
598:, where four leopard fossils were found. These are dated to the end of the Late Pleistocene, about 29,000–37,000 years ago. A cave painting of a leopard in the
1140:Τsoukala, Ε.; Bartsiokas, Α.; Chatzοpoulou, Κ.; Lazaridis, G. (2006). "Quaternary mammalian remains from the Kitseli Pothole (Alea, Nemea, Peloponnese)".
492:
shows a coat pattern similar to that of modern leopards but with a unspotted belly, presumably white. Like other mammals, leopards from the cold
1236:
1211:
769:
Recherches sur les ossemens fossiles ou l'on retablit les caractères de plusieurs animaux dont les revolutions du globe ont détruit les espèces
630:
110:
119:
987:
Sommer, R. S. & Benecke, N. (2006). "Late
Pleistocene and Holocene development of the felid fauna (Felidae) of Europe: a review".
1241:
1231:
655:
in
Germany. It is not precisely known which prey species these leopards hunted, although they may have been similar to modern
521:, around 600,000 years ago. While initially rare, leopards become much more common and widely distributed from the late
1216:
586:. Leopard fossils dating to ~43,000 BP were found in the Radochowska Cave in Poland. The most complete skeleton of
127:
1221:
148:
930:
Marciszak, A.; Lipecki, G.; Gornig, W.; Matyaszczyk, L.; Oszczepalińska, O.; Nowakowski, D.; Talamo, S. (2022).
555:
527:
484:
783:
Fraipont, C. (1923). "Crane de Panthère ou de Lynx géant provenent de la caverne de Trois-Frères (Ariège)".
607:
595:
280:
940:
1109:
Marciszak, A.; Krajcarz, M.T.; Krajcarz, M. & Stefaniak, K. (2011). "The first record of leopard
1063:
949:
869:
722:
603:
535:
1183:
1052:"Bone accumulation by Leopards in the Late Pleistocene in the Moncayo Massif (Zaragoza, NE Spain)"
989:
522:
518:
482:
The
European Ice Age leopard's skull was medium-long, and its characteristics are closest to the
459:
426:
309:
143:
1023:(Felidae, Mammalia) record from Equi (Massa, Italy): taphonomy, morphology, and paleoecology".
858:"Historical biogeography of the leopard (Panthera pardus) and its extinct Eurasian populations"
1226:
1091:
905:
887:
543:
539:
505:
422:
1178:
1122:
1081:
1071:
1032:
998:
967:
957:
895:
877:
834:
730:
685:
644:
497:
430:
399:
652:
767:
1067:
953:
873:
726:
614:
records in the
Iberian Peninsula possibly extending into the early Holocene, during the
1086:
1051:
900:
857:
619:
493:
1160:
838:
300:
1205:
1169:
1002:
575:
61:
1036:
734:
656:
599:
501:
489:
136:
1076:
559:
230:
36:
1050:
Sauqué, V.; Rabal-Garcés, R.; Sola-Almagro, C. & Cuenca-Bescós, G. (2014).
618:. Modern (Asian-type) leopards are still found on the fringes of Europe in the
1126:
882:
615:
591:
391:
210:
81:
46:
891:
668:
640:
200:
160:
86:
1095:
909:
471:
403:
240:
180:
76:
71:
56:
51:
41:
972:
962:
931:
680:
571:
547:
388:
254:
220:
91:
66:
583:
579:
395:
385:
190:
170:
538:, leopards persisted in relatively temperate glacial refugia in the
629:
567:
563:
664:
660:
825:
n. ssp., a new leopard from the Pleistocene of Vraona/Greece".
634:
Skull with a hole determined to have been caused by a lion bite
433:
were described and proposed as different leopard subspecies:
827:
Neues Jahrbuch für Geologie und Paläontologie - Monatshefte
606:. In Germany, the leopard survived at least into the early
525:
onwards, following the extinction of the "European jaguar"
488:
subspecies. An apparent depiction of this leopard in the
932:"The first radiocarbon-dated remains of the Leopard
267:
1115:Acta Zoologica Cracoviensia - Series A: Vertebrata
1019:Ghezzo, E. & Rook, L. (2015). "The remarkable
504:phases. As in modern leopards, there was a strong
1113:LINNAEUS, 1758 from the Pleistocene of Poland".
936:(Linnaeus, 1758) from the Pleistocene of Poland"
1184:10.2305/IUCN.UK.2020-1.RLTS.T15954A163991139.en
707:
705:
703:
701:
1142:Επιστημονική Επετηρίδα του Τμήματος Γεωλογίας
1014:
1012:
808:Neues Jahrbuch für Geologie und Paläontologie
746:
744:
8:
500:are usually larger than those from the warm
126:
109:
20:
1182:
1085:
1075:
971:
961:
899:
881:
508:, with males being larger than females.
697:
755:. St. Gallen: H. Tschudy. p. 254.
414:The subspecies was first described as
135:Rock art depiction of a leopard from
7:
925:
923:
921:
919:
850:
848:
1170:IUCN Red List of Threatened Species
806:n. subsp. Aus den Mauerer Sanden".
120:Museo di Storia Naturale di Firenze
14:
753:Das Wildkirchli: eine Monographie
1003:10.1111/j.1469-7998.2005.00040.x
147:
1037:10.1016/j.quascirev.2014.12.020
772:. Paris: Dufour et E. d'Ocagne.
735:10.1016/j.quascirev.2013.05.009
550:Peninsulas. Bone fragments of
421:Several fossil bones from the
16:Pleistocene leopard subspecies
1:
1237:Pleistocene mammals of Europe
1212:Fossil taxa described in 1936
1077:10.1371/journal.pone.0092144
474:) around 500,000 years ago.
839:10.1127/njgpm/1999/1999/129
804:Panthera pardus sickenbergi
351:Panthera pardus sickenbergi
1258:
1025:Quaternary Science Reviews
823:Panthera pardus vraonensis
715:Quaternary Science Reviews
359:Panthera pardus vraonensis
1127:10.3409/azc.54a_1-2.39-46
883:10.1186/s12862-018-1268-0
458:These are now considered
418:by Emil Bächler in 1936.
315:
308:
286:
279:
144:Scientific classification
142:
134:
125:
117:
108:
23:
862:BMC Evolutionary Biology
802:Schütt, Von G. (1969). "
528:Panthera gombaszoegensis
485:Panthera pardus tulliana
378:European Ice Age leopard
343:Panthera pardus begoueni
1242:Prehistoric pantherines
1232:Pleistocene carnivorans
373:Panthera pardus spelaea
335:Panthera pardus antiqua
293:Panthera pardus spelaea
25:Panthera pardus spelaea
1177:: e.T15954A163991139.
635:
608:Weichselian glaciation
596:Bosnia and Herzegovina
785:Revue d'Anthropologie
633:
751:Bächler, E. (1936).
604:Last Glacial Maximum
536:Last Glacial Maximum
416:Felis pardus spelaea
376:, also known as the
319:Felis pardus spelaea
1068:2014PLoSO...992144S
963:10.1017/RDC.2022.33
954:2022Radcb..64.1359M
874:2018BMCEE..18..156P
821:Nagel, D. (1999). "
766:Cuvier, G. (1835).
727:2013QSRv...76..167D
990:Journal of Zoology
681:Leopard subspecies
636:
523:Middle Pleistocene
519:Middle Pleistocene
272:P. p. spelaea
1217:Fossils of Serbia
594:Cave in southern
506:sexual dimorphism
448:P. p. sickenbergi
402:and possibly the
369:
368:
1249:
1222:Late Pleistocene
1196:
1195:
1193:
1191:
1186:
1156:
1150:
1149:
1137:
1131:
1130:
1106:
1100:
1099:
1089:
1079:
1047:
1041:
1040:
1016:
1007:
1006:
984:
978:
977:
975:
965:
948:(6): 1359–1372.
927:
914:
913:
903:
885:
852:
843:
842:
818:
812:
811:
799:
793:
792:
780:
774:
773:
763:
757:
756:
748:
739:
738:
709:
686:Zanzibar leopard
659:, which prey on
498:Late Pleistocene
453:P. p. vraonensis
431:Late Pleistocene
400:Late Pleistocene
363:
355:
347:
339:
331:
323:
295:
291:
269:
152:
151:
130:
118:Skeleton at the
113:
103:
33:
32:Late Pleistocene
29:Temporal range:
21:
1257:
1256:
1252:
1251:
1250:
1248:
1247:
1246:
1202:
1201:
1200:
1199:
1189:
1187:
1163:Panthera pardus
1158:
1157:
1153:
1139:
1138:
1134:
1111:Panthera pardus
1108:
1107:
1103:
1049:
1048:
1044:
1021:Panthera pardus
1018:
1017:
1010:
986:
985:
981:
934:Panthera pardus
929:
928:
917:
854:
853:
846:
820:
819:
815:
801:
800:
796:
782:
781:
777:
765:
764:
760:
750:
749:
742:
711:
710:
699:
694:
677:
628:
514:
494:glacial periods
480:
460:junior synonyms
412:
361:
353:
346:Fraipoint, 1923
345:
337:
329:
321:
304:
297:
289:
288:
275:
258:
146:
104:
102:
101:
100:
99:
94:
89:
84:
79:
74:
69:
64:
59:
54:
49:
44:
39:
31:
30:
27:
17:
12:
11:
5:
1255:
1253:
1245:
1244:
1239:
1234:
1229:
1224:
1219:
1214:
1204:
1203:
1198:
1197:
1151:
1132:
1121:(1–2): 39–46.
1101:
1042:
1008:
979:
915:
844:
833:(3): 129–150.
813:
794:
775:
758:
740:
696:
695:
693:
690:
689:
688:
683:
676:
673:
653:Zoolithen Cave
627:
624:
620:North Caucasus
590:is known from
513:
510:
479:
476:
456:
455:
450:
445:
443:P. p. begoueni
440:
411:
408:
367:
366:
365:
364:
356:
348:
340:
338:(Cuvier, 1835)
332:
327:Felis antiquus
324:
313:
312:
306:
305:
298:
284:
283:
281:Trinomial name
277:
276:
266:
264:
260:
259:
255:P. pardus
252:
250:
246:
245:
238:
234:
233:
228:
224:
223:
218:
214:
213:
208:
204:
203:
198:
194:
193:
188:
184:
183:
178:
174:
173:
168:
164:
163:
158:
154:
153:
140:
139:
132:
131:
123:
122:
115:
114:
106:
105:
97:
96:
95:
90:
85:
80:
75:
70:
65:
60:
55:
50:
45:
40:
35:
34:
28:
15:
13:
10:
9:
6:
4:
3:
2:
1254:
1243:
1240:
1238:
1235:
1233:
1230:
1228:
1225:
1223:
1220:
1218:
1215:
1213:
1210:
1209:
1207:
1185:
1180:
1176:
1172:
1171:
1166:
1164:
1155:
1152:
1147:
1143:
1136:
1133:
1128:
1124:
1120:
1116:
1112:
1105:
1102:
1097:
1093:
1088:
1083:
1078:
1073:
1069:
1065:
1062:(3): e92144.
1061:
1057:
1053:
1046:
1043:
1038:
1034:
1030:
1026:
1022:
1015:
1013:
1009:
1004:
1000:
996:
992:
991:
983:
980:
974:
969:
964:
959:
955:
951:
947:
943:
942:
937:
935:
926:
924:
922:
920:
916:
911:
907:
902:
897:
893:
889:
884:
879:
875:
871:
867:
863:
859:
851:
849:
845:
840:
836:
832:
828:
824:
817:
814:
809:
805:
798:
795:
790:
786:
779:
776:
771:
770:
762:
759:
754:
747:
745:
741:
736:
732:
728:
724:
720:
716:
708:
706:
704:
702:
698:
691:
687:
684:
682:
679:
678:
674:
672:
670:
666:
662:
658:
657:snow leopards
654:
650:
648:
642:
632:
626:Palaeobiology
625:
623:
621:
617:
611:
609:
605:
601:
597:
593:
589:
588:P. p. spelaea
585:
581:
577:
576:Great Britain
573:
569:
565:
561:
557:
553:
552:P. p. spelaea
549:
545:
541:
537:
532:
531:
529:
524:
520:
511:
509:
507:
503:
499:
495:
491:
487:
486:
477:
475:
473:
467:
465:
464:P. p. spelaea
461:
454:
451:
449:
446:
444:
441:
439:
438:P. p. antiqua
436:
435:
434:
432:
428:
424:
419:
417:
409:
407:
405:
401:
397:
394:which roamed
393:
390:
387:
383:
379:
375:
374:
360:
357:
352:
349:
344:
341:
336:
333:
328:
325:
322:Bächler, 1936
320:
317:
316:
314:
311:
307:
302:
296:
294:
285:
282:
278:
274:
273:
265:
262:
261:
257:
256:
251:
248:
247:
244:
243:
239:
236:
235:
232:
229:
226:
225:
222:
219:
216:
215:
212:
209:
206:
205:
202:
199:
196:
195:
192:
189:
186:
185:
182:
179:
176:
175:
172:
169:
166:
165:
162:
159:
156:
155:
150:
145:
141:
138:
133:
129:
124:
121:
116:
112:
107:
93:
88:
83:
78:
73:
68:
63:
58:
53:
48:
43:
38:
26:
22:
19:
1188:. Retrieved
1174:
1168:
1162:
1154:
1145:
1141:
1135:
1118:
1114:
1110:
1104:
1059:
1055:
1045:
1028:
1024:
1020:
994:
988:
982:
973:11585/887180
945:
939:
933:
865:
861:
830:
826:
822:
816:
807:
803:
797:
788:
784:
778:
768:
761:
752:
718:
714:
646:
645:cave lions (
637:
612:
600:Chauvet Cave
587:
551:
533:
526:
515:
512:Distribution
502:interglacial
490:Chauvet Cave
483:
481:
468:
463:
457:
452:
447:
442:
437:
420:
415:
413:
382:cave leopard
381:
377:
372:
371:
370:
358:
354:Schutt, 1969
350:
342:
334:
330:Cuvier, 1835
326:
318:
292:
287:
271:
270:
263:Subspecies:
253:
241:
137:Chauvet Cave
24:
18:
1031:: 131–151.
997:(1): 7–19.
941:Radiocarbon
721:: 167–193.
560:Switzerland
534:During the
478:Description
362:Nagel, 1999
231:Pantherinae
227:Subfamily:
1206:Categories
1190:15 January
1148:: 273–284.
868:(1): 156.
810:: 299–310.
692:References
647:P. spelaea
641:cave bears
616:Mesolithic
592:Vjetrenica
392:subspecies
211:Feliformia
207:Suborder:
892:1471-2148
669:wild boar
556:excavated
249:Species:
201:Carnivora
167:Kingdom:
161:Eukaryota
1227:Leopards
1096:24642667
1056:PLOS ONE
910:30348080
675:See also
472:Caucasus
410:Taxonomy
404:Holocene
310:Synonyms
242:Panthera
217:Family:
191:Mammalia
181:Chordata
177:Phylum:
171:Animalia
157:Domain:
1087:3958443
1064:Bibcode
950:Bibcode
901:6198532
870:Bibcode
723:Bibcode
572:Germany
544:Italian
540:Iberian
496:of the
398:in the
389:leopard
384:, is a
380:or the
303:, 1936)
301:Bächler
290:†
237:Genus:
221:Felidae
197:Order:
187:Class:
98:↓
1094:
1084:
908:
898:
890:
584:Greece
580:Poland
548:Balkan
427:Middle
396:Europe
386:fossil
791:: 42.
568:Spain
564:Italy
554:were
423:Early
1192:2022
1175:2020
1092:PMID
906:PMID
888:ISSN
831:1999
667:and
665:deer
661:ibex
582:and
546:and
429:and
37:PreꞒ
1179:doi
1123:doi
1082:PMC
1072:doi
1033:doi
1029:110
999:doi
995:269
968:hdl
958:doi
896:PMC
878:doi
835:doi
731:doi
558:in
462:of
1208::
1173:.
1167:.
1146:98
1144:.
1119:54
1117:.
1090:.
1080:.
1070:.
1058:.
1054:.
1027:.
1011:^
993:.
966:.
956:.
946:64
944:.
938:.
918:^
904:.
894:.
886:.
876:.
866:18
864:.
860:.
847:^
829:.
789:33
787:.
743:^
729:.
719:76
717:.
700:^
663:,
643:,
622:.
610:.
578:,
574:,
570:,
566:,
562:,
542:,
466:.
425:,
406:.
87:Pg
1194:.
1181::
1165:"
1161:"
1129:.
1125::
1098:.
1074::
1066::
1060:9
1039:.
1035::
1005:.
1001::
976:.
970::
960::
952::
912:.
880::
872::
841:.
837::
737:.
733::
725::
649:)
530:.
299:(
268:†
92:N
82:K
77:J
72:T
67:P
62:C
57:D
52:S
47:O
42:Ꞓ
Text is available under the Creative Commons Attribution-ShareAlike License. Additional terms may apply.
