48:
187:
173:
35:
403:. The sequences indicate the dire wolf to be a highly divergent lineage which last shared a most recent common ancestor with the wolf-like canines 5.7 million years ago, with morphological similarity to the grey wolf being a result of convergent evolution. The study's findings are consistent with the previously proposed taxonomic classification of the dire wolf as genus
518:
by
Christopher J. Bell and Ernest L. Lundelius Jr., Anthony D. Barnosky, Russell W. Graham, Everett H. Lindsay, Dennis R. Ruez Jr., Holmes A. Semken Jr., S. David Webb, and Richard J. Zakrzewski. January 2004 in the book: Late Cretaceous and Cenozoic Mammals of North America: Biostratigraphy and
639:
Perri, Angela R.; Mitchell, Kieren J.; Mouton, Alice; Álvarez-Carretero, Sandra; Hulme-Beaman, Ardern; Haile, James; Jamieson, Alexandra; Meachen, Julie; Lin, Audrey T.; Schubert, Blaine W.; Ameen, Carly; Antipina, Ekaterina E.; Bover, Pere; Brace, Selina; Carmagnini, Alberto (2021-03-04).
584:
Berta, A. 1995. Fossil carnivores from the Leisey Shell Pits, Hillsborough County, Florida. In R.C. Hulbert, Jr., G.S. Morgan, and S.D. Webb (editors), Paleontology and geology of the Leisey Shell Pits, early
Pleistocene of Florida. Bulletin of the Florida Museum of Natural History 37:
413:
ostensibly evolved in
Eurasia and expanded into North America more recently, during the Late Pleistocene, and did not experience admixture with the dire wolf. This long-term isolation of the dire wolf lineage implies that other American fossil canines, including
407:. The study proposes an early origin of the dire wolf lineage in the Americas, and that this geographic isolation also resulted in reproductive isolation, driving their divergence 5.7 million years ago. Concurrently, Coyotes, dholes, gray wolves, and the extinct
608:
Martin, R. A., and S. D. Webb (1974). Late
Pleistocene mammals from the Devil's Den fauna, Levy County p114-45 in S. D. Webb (ed.), Pleistocene Mammals of Florida, Gainesville: University Presses of Florida
1976:
775:
1907:
1986:
515:
620:
Wang, Xiaoming; Tedford, Richard H.; Dogs: Their Fossil
Relatives and Evolutionary History. New York: Columbia University Press, 2008.
768:
569:
531:
R. M. Nowak. 1979. North
American Quaternary Canis. Monograph of the Museum of Natural History, University of Kansas 6:1-154 LINK:
1971:
231:. It is notable because it is proposed as the ancestor of one of the most famous prehistoric carnivores in North America, the
361:
629:
Nowak, R. M. and
Federoff, N. E. (2002). The systematic status of the Italian wolf Canis lupus. Acta theriol. 47(3): 333-338
1966:
1859:
1830:
761:
262:, Maryland, in an Irvingtonian terrestrial horizon. Fossil distribution is widespread throughout the United States.
542:
594:
47:
1961:
1760:
703:
Bartolini-Lucenti, S.; Cirilli, O.; Melchionna, M.; Raia, P.; Tseng, Z. J.; Flynn, J. J.; Rook, L. (2024).
1875:
1800:
1750:
1292:
265:
In the Middle
Pleistocene of North America, wolf species became larger, with some studies concluding that
519:
Geochronology. Chapter: 7. Publisher: Columbia
University Press; Editors: Michael O. Woodburne. pp274-276
1981:
1933:
560:
R.M. Nowak (2003). "Chapter 9 - Wolf evolution and taxonomy". In Mech, L. David; Boitani, Luigi (eds.).
143:
475:
1282:
1272:
1700:
1446:
653:
290:
259:
1590:
1436:
1312:
532:
392:
around central
America in open terrain habitats, with later eastward expansion and displacement of
276:
1740:
1675:
1471:
1335:
1302:
685:
498:
310:
251:
42:
1938:
1920:
1820:
1720:
1710:
1620:
1490:
740:
677:
669:
565:
544:
Preliminary report on a recently discovered Pleistocene cave deposit near Cumberland, Maryland
1925:
372:, with Nowak stating that there were specimens from Cumberland Cave, Maryland that indicated
1650:
1345:
730:
720:
661:
490:
186:
1506:
1355:
911:
860:
247:
753:
735:
704:
657:
172:
1640:
1630:
1610:
1561:
1366:
1066:
846:
439:
420:
321:
315:
267:
228:
34:
1955:
1790:
1780:
1770:
1730:
1154:
1118:
1100:
1045:
689:
212:
641:
502:
476:"Phylogenetic Systematics of the North American Fossil Caninae (Carnivora: Canidae)"
1411:
1375:
1073:
1000:
991:
946:
890:
389:
381:
216:
162:
709:
type (Canidae, Mammalia) from the Upper Valdarno Basin (Italy, Early Pleistocene)"
1247:
1168:
1145:
1059:
1018:
959:
932:
904:
869:
220:
1898:
725:
665:
325:
share dental and cranial similarities developed for hypercarnivory, suggesting
1810:
1662:
1182:
1091:
1080:
1052:
918:
883:
824:
784:
399:
In 2021, researchers sequenced the nuclear DNA (from the cell nucleus) of the
673:
1526:
1423:
1401:
1227:
1217:
1161:
1134:
1127:
1107:
1038:
1031:
1007:
980:
973:
966:
925:
897:
818:
400:
232:
99:
59:
744:
681:
1892:
1687:
1536:
1458:
1391:
1237:
876:
812:
806:
433:
409:
342:
285:
79:
1912:
1259:
1201:
1175:
788:
208:
109:
1516:
1323:
800:
444:
89:
69:
1869:
1573:
1548:
494:
431:
In a 2024 study, Armbruster's wolf was considered as a species of
224:
185:
119:
642:"Dire wolves were the last of an ancient New World canid lineage"
1873:
757:
474:
Tedford, Richard H.; Wang, Xiaoming; Taylor, Beryl E. (2009).
516:
The _Blancan, Irvingtonian and Rancholabrean Mammal Ages
180:
Range of Armbruster's wolf based on fossil distribution
364:, R. H. Tedford and R. M. Nowak have all proposed that
313:, on the other hand, suggested that the South American
1826:
1816:
1806:
1796:
1786:
1776:
1766:
1756:
1746:
1736:
1726:
1716:
1706:
1696:
1671:
1646:
1636:
1626:
1616:
1606:
1596:
1586:
1557:
1532:
1522:
1512:
1502:
1467:
1442:
1432:
1407:
1397:
1387:
1351:
1341:
1331:
1308:
1298:
1288:
1278:
1268:
1243:
1233:
1223:
1213:
943:
857:
842:
1882:
1685:
1660:
1571:
1546:
1489:
1456:
1421:
1374:
1365:
1322:
1257:
1200:
1144:
1117:
1090:
1017:
990:
942:
856:
833:
27:
Temporal range: Middle Pleistocene-Late Pleistocene
546:, JW Gidley - 1913 - US Government Printing Office
483:Bulletin of the American Museum of Natural History
443:was considered more closely related to the modern
380:. The two taxa share a number of characteristics (
258:at that time. The first fossils were uncovered at
564:. University of Chicago Press. pp. 239–258.
769:
469:
467:
465:
463:
461:
459:
250:in 1913. It appeared in North America in the
8:
562:Wolves: Behaviour, Ecology and Conservation
555:
553:
527:
525:
357:before their migration into North America.
1870:
1579:
1495:
1486:
1380:
1371:
1206:
1023:
951:
776:
762:
754:
616:
614:
254:, and is a wolf-like form larger than any
171:
33:
20:
734:
724:
455:
305:, positing that this then gave rise to
341:. Based on morphology from China, the
7:
1977:Prehistoric mammals of North America
424:, may belong to the same lineage as
349:may have been the ancestor for both
14:
46:
705:"Virtual reconstruction of the
1:
1987:Fossil taxa described in 1913
279:disagreed, and believed that
271:diverged into the large wolf
194:in red. (Tedford & Wang)
16:Extinct species of carnivore
384:), suggesting an origin of
329:was the common ancestor of
2003:
726:10.1038/s41598-024-53073-5
666:10.1038/s41586-020-03082-x
275:based on tooth specimens.
1856:
1582:
1498:
1383:
1209:
1026:
954:
795:
223:epoch, spanning from 1.9
179:
170:
149:
142:
43:Scientific classification
41:
32:
23:
190:Timeline of canids with
1972:Pleistocene carnivorans
787:members of the family
293:proposed instead that
195:
1934:Paleobiology Database
235:, which replaced it.
215:and lived during the
189:
1562:C. alpinus europaeus
595:Fossilworks website
360:The paleontologists
260:Cumberland Bone Cave
211:that was endemic to
1967:Prehistoric canines
1801:C. l. mogollonensis
658:2021Natur.591...87P
297:was not related to
135:C. armbrusteri
1811:C. l. monstrabilis
713:Scientific Reports
311:Richard H. Tedford
252:Middle Pleistocene
196:
1949:
1948:
1921:Open Tree of Life
1884:Canis armbrusteri
1876:Taxon identifiers
1867:
1866:
1852:
1851:
1848:
1847:
1844:
1843:
1840:
1839:
1771:C. l. griseoalbus
1485:
1484:
1481:
1480:
1196:
1195:
1192:
1191:
597:Canis armbrusteri
368:had evolved from
244:Canis armbrusteri
204:Canis armbrusteri
199:Armbruster's wolf
192:Canis armbrusteri
184:
183:
156:Canis armbrusteri
24:Armbruster's wolf
1994:
1942:
1941:
1929:
1928:
1916:
1915:
1903:
1902:
1901:
1871:
1828:
1818:
1808:
1798:
1791:C. l. hodophilax
1788:
1778:
1768:
1758:
1748:
1741:C. l. floridanus
1738:
1731:C. l. cristaldii
1728:
1718:
1708:
1698:
1673:
1648:
1638:
1628:
1618:
1608:
1598:
1591:C. apolloniensis
1588:
1580:
1559:
1534:
1524:
1514:
1504:
1496:
1487:
1469:
1444:
1434:
1409:
1399:
1389:
1381:
1372:
1353:
1343:
1333:
1310:
1300:
1290:
1280:
1270:
1245:
1235:
1225:
1215:
1207:
1024:
952:
945:
859:
844:
778:
771:
764:
755:
749:
748:
738:
728:
700:
694:
693:
636:
630:
627:
621:
618:
609:
606:
600:
592:
586:
582:
576:
575:
557:
548:
540:
534:
529:
520:
513:
507:
506:
480:
471:
303:C. priscolatrans
268:C. priscolatrans
207:) is an extinct
175:
158:
154:
51:
50:
37:
21:
2002:
2001:
1997:
1996:
1995:
1993:
1992:
1991:
1952:
1951:
1950:
1945:
1937:
1932:
1924:
1919:
1911:
1906:
1897:
1896:
1891:
1878:
1868:
1863:
1836:
1711:C. l. beothucus
1681:
1656:
1567:
1542:
1477:
1452:
1447:L. ensenadensis
1417:
1361:
1318:
1293:U. minicephalus
1253:
1188:
1140:
1113:
1086:
1013:
986:
938:
912:Paraenhydrocyon
861:Hesperocyoninae
852:
829:
791:
782:
752:
702:
701:
697:
652:(7848): 87–91.
638:
637:
633:
628:
624:
619:
612:
607:
603:
593:
589:
583:
579:
572:
559:
558:
551:
541:
537:
530:
523:
514:
510:
478:
473:
472:
457:
453:
291:Ronald M. Nowak
248:James W. Gidley
241:
227:—250,000 years
166:
160:
152:
151:
138:
45:
28:
25:
17:
12:
11:
5:
2000:
1998:
1990:
1989:
1984:
1979:
1974:
1969:
1964:
1962:Extinct wolves
1954:
1953:
1947:
1946:
1944:
1943:
1930:
1917:
1904:
1888:
1886:
1880:
1879:
1874:
1865:
1864:
1857:
1854:
1853:
1850:
1849:
1846:
1845:
1842:
1841:
1838:
1837:
1835:
1834:
1824:
1814:
1804:
1794:
1784:
1774:
1764:
1761:C. l. gregoryi
1754:
1744:
1734:
1724:
1721:C. l. bernardi
1714:
1704:
1693:
1691:
1683:
1682:
1680:
1679:
1668:
1666:
1658:
1657:
1655:
1654:
1644:
1634:
1624:
1621:C. cedazoensis
1614:
1604:
1601:C. armbrusteri
1594:
1583:
1577:
1569:
1568:
1566:
1565:
1554:
1552:
1544:
1543:
1541:
1540:
1530:
1520:
1510:
1499:
1493:
1483:
1482:
1479:
1478:
1476:
1475:
1464:
1462:
1454:
1453:
1451:
1450:
1440:
1429:
1427:
1419:
1418:
1416:
1415:
1405:
1395:
1384:
1378:
1369:
1363:
1362:
1360:
1359:
1349:
1339:
1336:V. qiuzhudingi
1328:
1326:
1320:
1319:
1317:
1316:
1306:
1296:
1286:
1276:
1265:
1263:
1255:
1254:
1252:
1251:
1241:
1231:
1221:
1210:
1204:
1198:
1197:
1194:
1193:
1190:
1189:
1187:
1186:
1179:
1172:
1165:
1158:
1150:
1148:
1142:
1141:
1139:
1138:
1131:
1123:
1121:
1115:
1114:
1112:
1111:
1104:
1096:
1094:
1088:
1087:
1085:
1084:
1077:
1070:
1067:Microtomarctus
1063:
1056:
1049:
1042:
1035:
1027:
1021:
1015:
1014:
1012:
1011:
1004:
996:
994:
988:
987:
985:
984:
977:
970:
963:
955:
949:
940:
939:
937:
936:
929:
922:
915:
908:
901:
894:
887:
880:
873:
865:
863:
854:
853:
851:
850:
847:Prohesperocyon
839:
837:
835:incertae sedis
831:
830:
828:
827:
821:
815:
809:
803:
796:
793:
792:
783:
781:
780:
773:
766:
758:
751:
750:
707:Canis arnensis
695:
631:
622:
610:
601:
587:
577:
570:
549:
535:
521:
508:
454:
452:
449:
440:Canis edwardii
416:C. armbrusteri
394:C. armbrusteri
376:diverged into
374:C. armbrusteri
370:C. armbrusteri
351:C. armbrusteri
347:C. chihliensis
327:C. armbrusteri
295:C. armbrusteri
281:C. armbrusteri
273:C. armbrusteri
240:
237:
182:
181:
177:
176:
168:
167:
161:
147:
146:
140:
139:
131:
129:
125:
124:
117:
113:
112:
107:
103:
102:
97:
93:
92:
87:
83:
82:
77:
73:
72:
67:
63:
62:
57:
53:
52:
39:
38:
30:
29:
26:
15:
13:
10:
9:
6:
4:
3:
2:
1999:
1988:
1985:
1983:
1980:
1978:
1975:
1973:
1970:
1968:
1965:
1963:
1960:
1959:
1957:
1940:
1935:
1931:
1927:
1922:
1918:
1914:
1909:
1905:
1900:
1894:
1890:
1889:
1887:
1885:
1881:
1877:
1872:
1862:
1861:
1855:
1833:
1832:
1825:
1823:
1822:
1821:C. l. nubilus
1815:
1813:
1812:
1805:
1803:
1802:
1795:
1793:
1792:
1785:
1783:
1782:
1775:
1773:
1772:
1765:
1763:
1762:
1755:
1753:
1752:
1745:
1743:
1742:
1735:
1733:
1732:
1725:
1723:
1722:
1715:
1713:
1712:
1705:
1703:
1702:
1695:
1694:
1692:
1690:
1689:
1684:
1678:
1677:
1676:C. l. orcutti
1670:
1669:
1667:
1665:
1664:
1659:
1653:
1652:
1651:C. lepophagus
1645:
1643:
1642:
1635:
1633:
1632:
1625:
1623:
1622:
1615:
1613:
1612:
1605:
1603:
1602:
1595:
1593:
1592:
1585:
1584:
1581:
1578:
1576:
1575:
1570:
1564:
1563:
1556:
1555:
1553:
1551:
1550:
1545:
1539:
1538:
1531:
1529:
1528:
1521:
1519:
1518:
1511:
1509:
1508:
1501:
1500:
1497:
1494:
1492:
1488:
1474:
1473:
1466:
1465:
1463:
1461:
1460:
1455:
1449:
1448:
1441:
1439:
1438:
1437:L. cultridens
1431:
1430:
1428:
1426:
1425:
1420:
1414:
1413:
1406:
1404:
1403:
1396:
1394:
1393:
1386:
1385:
1382:
1379:
1377:
1373:
1370:
1368:
1364:
1358:
1357:
1350:
1348:
1347:
1340:
1338:
1337:
1330:
1329:
1327:
1325:
1321:
1315:
1314:
1307:
1305:
1304:
1303:U. progressus
1297:
1295:
1294:
1287:
1285:
1284:
1277:
1275:
1274:
1267:
1266:
1264:
1262:
1261:
1256:
1250:
1249:
1242:
1240:
1239:
1232:
1230:
1229:
1222:
1220:
1219:
1212:
1211:
1208:
1205:
1203:
1199:
1185:
1184:
1180:
1178:
1177:
1173:
1171:
1170:
1166:
1164:
1163:
1159:
1157:
1156:
1155:Paratomarctus
1152:
1151:
1149:
1147:
1143:
1137:
1136:
1132:
1130:
1129:
1125:
1124:
1122:
1120:
1119:Aelurodontina
1116:
1110:
1109:
1105:
1103:
1102:
1101:Paracynarctus
1098:
1097:
1095:
1093:
1089:
1083:
1082:
1078:
1076:
1075:
1071:
1069:
1068:
1064:
1062:
1061:
1057:
1055:
1054:
1050:
1048:
1047:
1046:Metatomarctus
1043:
1041:
1040:
1036:
1034:
1033:
1029:
1028:
1025:
1022:
1020:
1016:
1010:
1009:
1005:
1003:
1002:
998:
997:
995:
993:
989:
983:
982:
978:
976:
975:
971:
969:
968:
964:
962:
961:
957:
956:
953:
950:
948:
941:
935:
934:
930:
928:
927:
923:
921:
920:
916:
914:
913:
909:
907:
906:
902:
900:
899:
895:
893:
892:
888:
886:
885:
881:
879:
878:
874:
872:
871:
867:
866:
864:
862:
855:
849:
848:
841:
840:
838:
836:
832:
826:
822:
820:
816:
814:
810:
808:
804:
802:
798:
797:
794:
790:
786:
779:
774:
772:
767:
765:
760:
759:
756:
746:
742:
737:
732:
727:
722:
718:
714:
710:
708:
699:
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31:
22:
19:
1982:Irvingtonian
1883:
1858:
1831:C. l. youngi
1829:
1819:
1809:
1799:
1789:
1781:C. l. hattai
1779:
1769:
1759:
1751:C. l. fuscus
1749:
1739:
1729:
1719:
1709:
1699:
1686:
1674:
1661:
1649:
1639:
1629:
1619:
1609:
1600:
1599:
1589:
1572:
1560:
1547:
1535:
1525:
1515:
1505:
1472:S. pacivorus
1470:
1457:
1445:
1435:
1422:
1412:Theriodictis
1410:
1400:
1390:
1376:Cerdocyonina
1354:
1346:V. riffautae
1344:
1334:
1311:
1301:
1291:
1281:
1271:
1258:
1246:
1236:
1226:
1216:
1181:
1174:
1167:
1160:
1153:
1133:
1126:
1106:
1099:
1079:
1074:Protomarctus
1072:
1065:
1058:
1051:
1044:
1037:
1030:
1006:
1001:Cynarctoides
999:
992:Phlaocyonini
979:
972:
965:
958:
947:Borophaginae
931:
924:
917:
910:
903:
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891:Sunkahetanka
889:
882:
875:
868:
845:
834:
716:
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390:Irvingtonian
388:in the late
385:
382:synapomorphy
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277:R. A. Martin
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217:Irvingtonian
203:
202:
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197:
191:
163:J. W. Gidley
155:
150:
134:
133:
120:
18:
1701:C. l. alces
1641:C. etruscus
1631:C. edwardii
1611:C. arnensis
1507:Cynotherium
1356:V. skinneri
1283:U. galushai
1273:U. citrinus
1248:Prototocyon
1169:Protepicyon
1146:Borophagina
1060:Psalidocyon
1019:Borophagini
960:Archaeocyon
933:Ectopocynus
905:Enhydrocyon
870:Hesperocyon
719:(1). 8303.
421:C. edwardii
335:C. nehringi
322:C. nehringi
289:. However,
221:Pleistocene
1956:Categories
1663:C. latrans
1183:Borophagus
1092:Cynarctina
1081:Tephrocyon
1053:Euoplocyon
919:Osbornodon
884:Cynodesmus
825:Caniformia
823:Suborder:
451:References
1424:Lycalopex
1402:Protocyon
1228:Metalopex
1218:Leptocyon
1162:Carpocyon
1135:Aelurodon
1128:Tomarctus
1108:Cynarctus
1039:Desmocyon
1032:Cormocyon
1008:Phlaocyon
981:Rhizocyon
974:Otarocyon
967:Oxetocyon
926:Caedocyon
898:Philotrox
819:Carnivora
799:Kingdom:
690:231604957
674:0028-0836
489:: 1–218.
401:dire wolf
233:dire wolf
128:Species:
100:Carnivora
66:Kingdom:
60:Eukaryota
1899:Q3884275
1893:Wikidata
1860:Category
1688:C. lupus
1537:Xenocyon
1527:Aenocyon
1459:Speothos
1392:Dusicyon
1313:U. webbi
1238:Nurocyon
877:Mesocyon
813:Mammalia
807:Chordata
805:Phylum:
801:Animalia
745:38594298
736:11004169
682:33442059
585:463–499.
503:83594819
437:, while
434:Aenocyon
426:Aenocyon
410:Xenocyon
405:Aenocyon
386:C. dirus
378:C. dirus
366:C. dirus
355:C. lupus
343:Pliocene
339:C. dirus
307:C. dirus
299:C. lupus
286:C. lupus
239:Taxonomy
106:Family:
90:Mammalia
80:Chordata
76:Phylum:
70:Animalia
56:Domain:
1926:3612502
1913:5936115
1260:Urocyon
1202:Caninae
1176:Epicyon
817:Order:
811:Class:
789:Canidae
785:Extinct
654:Bibcode
362:X. Wang
331:C. gezi
316:C. gezi
209:species
153:†
132:†
116:Genus:
110:Canidae
96:Order:
86:Class:
1517:Eucyon
1491:Canina
1367:Canini
1324:Vulpes
743:
733:
688:
680:
672:
646:Nature
568:
501:
445:coyote
301:, but
165:, 1913
1939:44827
1574:Canis
686:S2CID
499:S2CID
479:(PDF)
345:wolf
256:Canis
121:Canis
1908:GBIF
1549:Cuon
741:PMID
678:PMID
670:ISSN
566:ISBN
418:and
353:and
337:and
319:and
283:was
731:PMC
721:doi
662:doi
650:591
491:doi
487:325
396:.
225:Mya
1958::
1936::
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309:.
229:BP
1827:†
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843:†
777:e
770:t
763:v
747:.
723::
692:.
664::
656::
574:.
505:.
493::
201:(
Text is available under the Creative Commons Attribution-ShareAlike License. Additional terms may apply.
