459:
386:
1890:. Felids invaded the continent from Eurasia about 20 million years ago and were better ambush predators, in part due to their retractable claws. Climate change, which led to the replacement of North American forests with grasslands, may also have been a factor; borophagines were less suited to running down prey than canines. Although these specific species developed powerful limbs that are capable of cracking bones in vicious pounce attacks, other carnivorous species that rely on quick ambush for catching prey were likely more successful than the slower Borophaginae.
336:
148:
633:
366:
125:
625:
432:, which is a small fox-sized animal mostly found in the fossil beds in western North America. The borophagines soon diversified into several major groups. They evolved to become considerably larger than their predecessors, and filled a wide range of niches in late
578:, showing that the group was highly diverse in its heyday. All Borophaginae had a small fifth toe on their rear feet (similar to the toes that bear dew claws on the front feet), where as all modern
458:
499:
methods, with
Hesperocyoninae as an archaic group of canids, as the outgroup. Aside from some transitional forms, Borophaginae can be organized into four major clades:
2205:
2381:
613:, the Borophaginae played broad ecological roles that are performed by at least three living carnivoran families, Canidae (which they were a part of),
3579:
2350:
2329:
1882:, the decline of borophagines from a diversity of about 30 species 15 million years ago to extinction was largely driven by competition with
3569:
1948:
2374:
2106:
2027:
Wang, Xiaoming; Tedford, Richard; Taylor, Beryl (1999-11-17). "Phylogenetic systematics of the
Borophaginae (Carnivora, Canidae)".
3574:
2277:"In the Pursuit of the Predatory Behavior of Borophagines (Mammalia, Carnivora, Canidae): Inferences from Forelimb Morphology"
2071:
610:
3564:
2193:
3465:
3436:
2367:
147:
3559:
515:(all erected as new tribes or subtribes). The Borophaginae begins with a group of small fox-sized genera, such as
385:
1981:
1963:
3366:
3481:
3406:
3356:
2898:
1907:
227:
3554:
3526:
1995:
1969:
483:
There are 66 identified borophagine species, including 18 newly identified species that range from the
2888:
2878:
3513:
3306:
3206:
3052:
2239:
2146:
3196:
3042:
2918:
3346:
3281:
3077:
2941:
2908:
2296:
142:
2063:
335:
3531:
3426:
3326:
3316:
3226:
3096:
2346:
2325:
2276:
2257:
2174:
2112:
2102:
2077:
2067:
1944:
1938:
547:
stages. These canids reached their maximum diversity of species around 28 million years ago.
3256:
2951:
2288:
2247:
2164:
2154:
2055:
2036:
1919:
624:
3112:
2961:
2517:
2466:
1108:
632:
423:
331:
262:
2359:
2243:
2150:
3246:
3236:
3216:
3167:
2972:
2672:
2452:
2169:
2134:
2056:
1479:
859:
365:
3548:
3396:
3386:
3376:
3336:
2760:
2724:
2706:
2651:
2300:
2201:
1607:
1540:
1407:
1357:
1088:
Cladogram showing borophagine interrelationships, following Wang et al., figure 141:
1000:
948:
917:
814:
508:
274:
73:
3017:
2981:
2679:
2606:
2597:
2496:
2194:"Smarts, stealth and retractable claws: Science confirms cats are better than dogs"
1503:
1246:
1232:
874:
739:
725:
567:
500:
492:
124:
2340:
2319:
2096:
17:
2853:
2774:
2751:
2665:
2624:
2565:
2538:
2510:
2475:
1990:
1655:
1593:
1455:
1285:
1154:
1030:
993:
844:
770:
666:
618:
517:
512:
428:
114:
48:
3504:
2228:"Habitat changes and changing predatory habits in North American fossil canids"
2135:"The role of clade competition in the diversification of North American canids"
574:. Their good fossil record has also allowed a detailed reconstruction of their
3416:
3268:
2788:
2697:
2686:
2658:
2524:
2489:
2430:
2390:
2342:
The
Skeleton of the Borophaginae (Carnivora, Canidae): Morphology and Function
2292:
1696:
1527:
1431:
1343:
1066:
1060:
903:
889:
829:
605:
599:
566:
are abundant and widespread; in all likelihood, they were probably one of the
544:
504:
463:
93:
58:
2116:
3132:
3029:
3007:
2833:
2823:
2767:
2740:
2733:
2713:
2644:
2637:
2613:
2586:
2579:
2572:
2531:
2503:
2424:
2159:
2081:
1910:(1945). "The principles of classification and a classification of mammals".
1879:
1631:
1571:
1554:
1374:
1323:
1299:
1263:
1212:
1188:
1171:
1015:
977:
962:
932:
799:
784:
754:
711:
696:
681:
614:
587:
575:
571:
559:
535:
529:
523:
475:
469:
278:
254:
199:
159:
133:
110:
98:
42:
2261:
2226:
Figueirido, B.; MartΓn-Serra, A.; Tseng, Z. J.; Janis, C. M. (2015-08-18).
2178:
1036:
895:
865:
850:
1943:. Oxford Biology (2 ed.). Oxford University Press. pp. 103β107.
1006:
3498:
3293:
3142:
3064:
2997:
2843:
2482:
2418:
2412:
649:
Borophagine taxonomy, following Wang et al. (million years=in existence)
550:
Often generically referred to as "bone-crushing dogs" for their powerful
437:
433:
282:
179:
88:
83:
68:
63:
53:
287:
2865:
2807:
2781:
2394:
2252:
2227:
1887:
1878:
According to an analysis of the fossil record of North
American fossil
1679:
1130:
1095:
1051:
1045:
983:
968:
938:
923:
908:
880:
835:
820:
805:
790:
760:
745:
730:
717:
702:
687:
672:
654:
637:
593:
579:
540:
488:
484:
446:
381:
314:
270:
266:
258:
209:
103:
78:
1072:
1021:
953:
775:
3122:
2929:
2406:
2040:
1923:
563:
189:
169:
3475:
3179:
3154:
2275:
MartΓn-Serra, Alberto; Figueirido, Borja; Palmqvist, Paul (2016).
2133:
Silvestro, D.; Antonelli, A.; Salamin, N.; Quental, T. B. (2015).
1883:
631:
623:
555:
551:
496:
457:
3518:
441:
3479:
2363:
285:
and lived roughly 34β2.5 million years ago, existing for about
2098:
Dogs : their fossil relatives and evolutionary history
426:. The earliest and most primitive borophagine is the genus
261:
family. The other two canid subfamilies are the extinct
2321:
Dogs: Their Fossil
Relatives and Evolutionary History
3432:
3422:
3412:
3402:
3392:
3382:
3372:
3362:
3352:
3342:
3332:
3322:
3312:
3302:
3277:
3252:
3242:
3232:
3222:
3212:
3202:
3192:
3163:
3138:
3128:
3118:
3108:
3073:
3048:
3038:
3013:
3003:
2993:
2957:
2947:
2937:
2914:
2904:
2894:
2884:
2874:
2849:
2839:
2829:
2819:
2549:
2463:
2448:
358:
328:
3488:
3291:
3266:
3177:
3152:
3095:
3062:
3027:
2980:
2971:
2928:
2863:
2806:
2750:
2723:
2696:
2623:
2596:
2548:
2462:
2439:
2062:. Washington, D.C.: National Geographic. pp.
2029:Bulletin of the American Museum of Natural History
1912:Bulletin of the American Museum of Natural History
269:. Borophaginae, called "bone-crushing dogs", were
2139:Proceedings of the National Academy of Sciences
2128:
2126:
628:Phylogeny of borophagines by R.L. Tedford, 1977
562:was more primitive than that of hyenas), their
422:The Borophaginae descended from the subfamily
2375:
8:
2318:Wang, X.; Tedford, R. H.; AntΓ³n, M. (2010).
2095:Wang, Xiaoming; Tedford, Richard H. (2008).
495:analysis of the species was conducted using
3476:
3185:
3101:
3092:
2986:
2977:
2812:
2629:
2557:
2382:
2368:
2360:
2022:
2020:
2018:
2016:
2014:
2012:
123:
31:
2251:
2168:
2158:
2324:. New York: Columbia University Press.
2101:. New York: Columbia University Press.
1940:Dog Behaviour, Evolution, and Cognition
1899:
7:
585:Noteworthy genera in this group are
1671:
1647:
1623:
1599:
1546:
1519:
1495:
1471:
1447:
1423:
1399:
1349:
1339:
1315:
1291:
1238:
1228:
1204:
1146:
1123:
1101:
1091:
351:
321:
309:
2345:. University of California Press.
25:
384:
364:
334:
146:
2208:from the original on 2015-08-18
558:-like features (although their
27:Extinct subfamily of carnivores
3580:Prehistoric animal subfamilies
2281:Journal of Mammalian Evolution
636:Fluctuation of species within
582:have only four toes normally.
1:
3570:Oligocene first appearances
444:-sized carnivores, such as
3596:
436:North America, from small
3462:
3188:
3104:
2989:
2815:
2632:
2560:
2401:
2293:10.1007/s10914-016-9321-5
1693:
1676:
1669:
1652:
1645:
1628:
1621:
1604:
1597:
1568:
1551:
1544:
1524:
1517:
1500:
1493:
1476:
1469:
1452:
1445:
1428:
1421:
1404:
1397:
1371:
1354:
1347:
1337:
1320:
1313:
1296:
1289:
1260:
1243:
1236:
1226:
1209:
1202:
1185:
1168:
1151:
1144:
1128:
1121:
1106:
1099:
379:
356:
349:
326:
319:
240:
235:
143:Scientific classification
141:
131:
122:
34:
2339:Kathleen Munthe (1989).
2192:Perry, D. (2015-08-14).
2160:10.1073/pnas.1502803112
660:Subfamily β Borophaginae
3575:Taxa described in 1945
2393:members of the family
1937:Miklosi, Adam (2015).
641:
629:
480:
476:Tomarctus brevirostris
3527:Paleobiology Database
2232:Nature Communications
2054:Turner, Alan (2004).
1996:Paleobiology Database
1970:Paleobiology Database
640:over 40 million years
635:
627:
461:
3565:Pliocene extinctions
3168:C. alpinus europaeus
1070:) 12β5 Ma, existing
115:Earliest Pleistocene
3407:C. l. mogollonensis
2244:2015NatCo...6.7976F
2151:2015PNAS..112.8684S
2058:Prehistoric Mammals
1049:12β10 Ma, existing
1034:16β12 Ma, existing
981:16β12 Ma, existing
966:23β16 Ma, existing
936:16β12 Ma, existing
921:19β16 Ma, existing
906:20β10 Ma, existing
893:16β14 Ma, existing
878:20β16 Ma, existing
863:21β13 Ma, existing
848:16β13 Ma, existing
833:18β16 Ma, existing
818:19β16 Ma, existing
803:25β16 Ma, existing
788:30β20 Ma, existing
743:30β18 Ma, existing
728:33β13 Ma, existing
715:33β26 Ma, existing
700:34β30 Ma, existing
685:33β28 Ma, existing
670:33β26 Ma, existing
464:Borophagus secundus
3560:Mammal subfamilies
3417:C. l. monstrabilis
2253:10.1038/ncomms8976
1019:16β5 Ma, existing
1004:16β5 Ma, existing
951:20β5 Ma, existing
773:30β3 Ma, existing
642:
630:
481:
470:Aelurodon taxoides
303:Canid subfamilies
253:form one of three
3542:
3541:
3482:Taxon identifiers
3473:
3472:
3458:
3457:
3454:
3453:
3450:
3449:
3446:
3445:
3377:C. l. griseoalbus
3091:
3090:
3087:
3086:
2802:
2801:
2798:
2797:
2352:978-0-520-09724-7
2331:978-0-231-13529-0
2145:(28): 8684β8689.
1994:. Gateway to the
1870:
1869:
1861:
1860:
1852:
1851:
1843:
1842:
1834:
1833:
1825:
1824:
1816:
1815:
1807:
1806:
1798:
1797:
1789:
1788:
1780:
1779:
1771:
1770:
1762:
1761:
1753:
1752:
1744:
1743:
1735:
1734:
1726:
1725:
1717:
1716:
1708:
1707:
1583:
1582:
1386:
1385:
1275:
1274:
420:
419:
415:
414:
406:
405:
397:
396:
257:found within the
247:
246:
231:
117:
18:Bone-crushing dog
16:(Redirected from
3587:
3535:
3534:
3522:
3521:
3509:
3508:
3507:
3477:
3434:
3424:
3414:
3404:
3397:C. l. hodophilax
3394:
3384:
3374:
3364:
3354:
3347:C. l. floridanus
3344:
3337:C. l. cristaldii
3334:
3324:
3314:
3304:
3279:
3254:
3244:
3234:
3224:
3214:
3204:
3197:C. apolloniensis
3194:
3186:
3165:
3140:
3130:
3120:
3110:
3102:
3093:
3075:
3050:
3040:
3015:
3005:
2995:
2987:
2978:
2959:
2949:
2939:
2916:
2906:
2896:
2886:
2876:
2851:
2841:
2831:
2821:
2813:
2630:
2558:
2551:
2465:
2450:
2384:
2377:
2370:
2361:
2356:
2335:
2305:
2304:
2272:
2266:
2265:
2255:
2223:
2217:
2216:
2214:
2213:
2189:
2183:
2182:
2172:
2162:
2130:
2121:
2120:
2092:
2086:
2085:
2061:
2051:
2045:
2044:
2024:
2007:
2006:
2004:
2002:
1978:
1972:
1961:
1955:
1954:
1934:
1928:
1927:
1918:: iβxvi, 1β350.
1904:
1672:
1648:
1624:
1600:
1547:
1520:
1496:
1472:
1448:
1424:
1400:
1350:
1340:
1316:
1292:
1239:
1229:
1205:
1147:
1124:
1102:
1092:
1076:
1055:
1040:
1025:
1010:
987:
972:
957:
942:
927:
912:
899:
884:
869:
854:
839:
824:
809:
794:
779:
764:
749:
734:
721:
706:
691:
676:
609:). According to
388:
368:
360:
352:
338:
330:
322:
310:
300:
299:
291:
226:
221:
151:
150:
127:
109:
108:
45:
38:Temporal range:
32:
21:
3595:
3594:
3590:
3589:
3588:
3586:
3585:
3584:
3545:
3544:
3543:
3538:
3530:
3525:
3517:
3512:
3503:
3502:
3497:
3484:
3474:
3469:
3442:
3317:C. l. beothucus
3287:
3262:
3173:
3148:
3083:
3058:
3053:L. ensenadensis
3023:
2967:
2924:
2899:U. minicephalus
2859:
2794:
2746:
2719:
2692:
2619:
2592:
2544:
2518:Paraenhydrocyon
2467:Hesperocyoninae
2458:
2435:
2397:
2388:
2353:
2338:
2332:
2317:
2314:
2312:Further reading
2309:
2308:
2274:
2273:
2269:
2225:
2224:
2220:
2211:
2209:
2191:
2190:
2186:
2132:
2131:
2124:
2109:
2094:
2093:
2089:
2074:
2053:
2052:
2048:
2026:
2025:
2010:
2000:
1998:
1980:
1979:
1975:
1962:
1958:
1951:
1936:
1935:
1931:
1906:
1905:
1901:
1896:
1876:
1871:
1862:
1853:
1844:
1835:
1826:
1817:
1808:
1799:
1790:
1781:
1772:
1763:
1754:
1745:
1736:
1727:
1718:
1709:
1584:
1387:
1276:
1109:Hesperocyoninae
1071:
1050:
1035:
1020:
1005:
982:
967:
952:
937:
922:
907:
894:
879:
864:
849:
834:
819:
804:
789:
774:
759:
744:
729:
716:
701:
686:
671:
647:
456:
424:Hesperocyoninae
416:
407:
398:
332:Hesperocyoninae
298:
286:
263:Hesperocyoninae
225:
219:
145:
118:
107:
106:
101:
96:
91:
86:
81:
76:
71:
66:
61:
56:
51:
40:
39:
36:
28:
23:
22:
15:
12:
11:
5:
3593:
3591:
3583:
3582:
3577:
3572:
3567:
3562:
3557:
3547:
3546:
3540:
3539:
3537:
3536:
3523:
3510:
3494:
3492:
3486:
3485:
3480:
3471:
3470:
3463:
3460:
3459:
3456:
3455:
3452:
3451:
3448:
3447:
3444:
3443:
3441:
3440:
3430:
3420:
3410:
3400:
3390:
3380:
3370:
3367:C. l. gregoryi
3360:
3350:
3340:
3330:
3327:C. l. bernardi
3320:
3310:
3299:
3297:
3289:
3288:
3286:
3285:
3274:
3272:
3264:
3263:
3261:
3260:
3250:
3240:
3230:
3227:C. cedazoensis
3220:
3210:
3207:C. armbrusteri
3200:
3189:
3183:
3175:
3174:
3172:
3171:
3160:
3158:
3150:
3149:
3147:
3146:
3136:
3126:
3116:
3105:
3099:
3089:
3088:
3085:
3084:
3082:
3081:
3070:
3068:
3060:
3059:
3057:
3056:
3046:
3035:
3033:
3025:
3024:
3022:
3021:
3011:
3001:
2990:
2984:
2975:
2969:
2968:
2966:
2965:
2955:
2945:
2942:V. qiuzhudingi
2934:
2932:
2926:
2925:
2923:
2922:
2912:
2902:
2892:
2882:
2871:
2869:
2861:
2860:
2858:
2857:
2847:
2837:
2827:
2816:
2810:
2804:
2803:
2800:
2799:
2796:
2795:
2793:
2792:
2785:
2778:
2771:
2764:
2756:
2754:
2748:
2747:
2745:
2744:
2737:
2729:
2727:
2721:
2720:
2718:
2717:
2710:
2702:
2700:
2694:
2693:
2691:
2690:
2683:
2676:
2673:Microtomarctus
2669:
2662:
2655:
2648:
2641:
2633:
2627:
2621:
2620:
2618:
2617:
2610:
2602:
2600:
2594:
2593:
2591:
2590:
2583:
2576:
2569:
2561:
2555:
2546:
2545:
2543:
2542:
2535:
2528:
2521:
2514:
2507:
2500:
2493:
2486:
2479:
2471:
2469:
2460:
2459:
2457:
2456:
2453:Prohesperocyon
2445:
2443:
2441:incertae sedis
2437:
2436:
2434:
2433:
2427:
2421:
2415:
2409:
2402:
2399:
2398:
2389:
2387:
2386:
2379:
2372:
2364:
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2287:(3): 237β249.
2267:
2218:
2198:OregonLive.com
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1956:
1950:978-0199545667
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1513:
1512:
1509:
1508:
1499:
1494:
1492:
1489:
1488:
1485:
1484:
1480:Microtomarctus
1475:
1470:
1468:
1465:
1464:
1461:
1460:
1451:
1446:
1444:
1441:
1440:
1437:
1436:
1427:
1422:
1420:
1417:
1416:
1413:
1412:
1403:
1398:
1396:
1393:
1392:
1389:
1388:
1384:
1383:
1380:
1379:
1370:
1367:
1366:
1363:
1362:
1353:
1348:
1346:
1338:
1336:
1333:
1332:
1329:
1328:
1319:
1314:
1312:
1309:
1308:
1305:
1304:
1295:
1290:
1288:
1282:
1281:
1278:
1277:
1273:
1272:
1269:
1268:
1259:
1256:
1255:
1252:
1251:
1242:
1237:
1235:
1227:
1225:
1222:
1221:
1218:
1217:
1208:
1203:
1201:
1198:
1197:
1194:
1193:
1184:
1181:
1180:
1177:
1176:
1167:
1164:
1163:
1160:
1159:
1150:
1145:
1143:
1139:
1138:
1135:
1134:
1127:
1122:
1120:
1117:
1116:
1113:
1112:
1105:
1100:
1098:
1090:
1086:
1085:
1084:
1083:
1082:
1081:
1080:
1079:
1078:
1077:
1056:
1041:
1026:
1011:
990:
989:
988:
973:
945:
944:
943:
928:
900:
885:
870:
860:Microtomarctus
855:
840:
825:
810:
795:
767:
766:
765:
750:
722:
707:
692:
677:
646:
645:Classification
643:
554:and jaws, and
543:through early
455:
452:
418:
417:
413:
412:
409:
408:
404:
403:
400:
399:
395:
394:
391:
390:
378:
375:
374:
371:
370:
355:
350:
348:
345:
344:
341:
340:
325:
320:
318:
308:
305:
304:
297:
294:
245:
244:
238:
237:
233:
232:
217:
213:
212:
207:
203:
202:
197:
193:
192:
187:
183:
182:
177:
173:
172:
167:
163:
162:
157:
153:
152:
139:
138:
129:
128:
120:
119:
102:
97:
92:
87:
82:
77:
72:
67:
62:
57:
52:
47:
46:
37:
26:
24:
14:
13:
10:
9:
6:
4:
3:
2:
3592:
3581:
3578:
3576:
3573:
3571:
3568:
3566:
3563:
3561:
3558:
3556:
3553:
3552:
3550:
3533:
3528:
3524:
3520:
3515:
3511:
3506:
3500:
3496:
3495:
3493:
3491:
3487:
3483:
3478:
3468:
3467:
3461:
3439:
3438:
3431:
3429:
3428:
3427:C. l. nubilus
3421:
3419:
3418:
3411:
3409:
3408:
3401:
3399:
3398:
3391:
3389:
3388:
3381:
3379:
3378:
3371:
3369:
3368:
3361:
3359:
3358:
3351:
3349:
3348:
3341:
3339:
3338:
3331:
3329:
3328:
3321:
3319:
3318:
3311:
3309:
3308:
3301:
3300:
3298:
3296:
3295:
3290:
3284:
3283:
3282:C. l. orcutti
3276:
3275:
3273:
3271:
3270:
3265:
3259:
3258:
3257:C. lepophagus
3251:
3249:
3248:
3241:
3239:
3238:
3231:
3229:
3228:
3221:
3219:
3218:
3211:
3209:
3208:
3201:
3199:
3198:
3191:
3190:
3187:
3184:
3182:
3181:
3176:
3170:
3169:
3162:
3161:
3159:
3157:
3156:
3151:
3145:
3144:
3137:
3135:
3134:
3127:
3125:
3124:
3117:
3115:
3114:
3107:
3106:
3103:
3100:
3098:
3094:
3080:
3079:
3072:
3071:
3069:
3067:
3066:
3061:
3055:
3054:
3047:
3045:
3044:
3043:L. cultridens
3037:
3036:
3034:
3032:
3031:
3026:
3020:
3019:
3012:
3010:
3009:
3002:
3000:
2999:
2992:
2991:
2988:
2985:
2983:
2979:
2976:
2974:
2970:
2964:
2963:
2956:
2954:
2953:
2946:
2944:
2943:
2936:
2935:
2933:
2931:
2927:
2921:
2920:
2913:
2911:
2910:
2909:U. progressus
2903:
2901:
2900:
2893:
2891:
2890:
2883:
2881:
2880:
2873:
2872:
2870:
2868:
2867:
2862:
2856:
2855:
2848:
2846:
2845:
2838:
2836:
2835:
2828:
2826:
2825:
2818:
2817:
2814:
2811:
2809:
2805:
2791:
2790:
2786:
2784:
2783:
2779:
2777:
2776:
2772:
2770:
2769:
2765:
2763:
2762:
2761:Paratomarctus
2758:
2757:
2755:
2753:
2749:
2743:
2742:
2738:
2736:
2735:
2731:
2730:
2728:
2726:
2725:Aelurodontina
2722:
2716:
2715:
2711:
2709:
2708:
2707:Paracynarctus
2704:
2703:
2701:
2699:
2695:
2689:
2688:
2684:
2682:
2681:
2677:
2675:
2674:
2670:
2668:
2667:
2663:
2661:
2660:
2656:
2654:
2653:
2652:Metatomarctus
2649:
2647:
2646:
2642:
2640:
2639:
2635:
2634:
2631:
2628:
2626:
2622:
2616:
2615:
2611:
2609:
2608:
2604:
2603:
2601:
2599:
2595:
2589:
2588:
2584:
2582:
2581:
2577:
2575:
2574:
2570:
2568:
2567:
2563:
2562:
2559:
2556:
2554:
2547:
2541:
2540:
2536:
2534:
2533:
2529:
2527:
2526:
2522:
2520:
2519:
2515:
2513:
2512:
2508:
2506:
2505:
2501:
2499:
2498:
2494:
2492:
2491:
2487:
2485:
2484:
2480:
2478:
2477:
2473:
2472:
2470:
2468:
2461:
2455:
2454:
2447:
2446:
2444:
2442:
2438:
2432:
2428:
2426:
2422:
2420:
2416:
2414:
2410:
2408:
2404:
2403:
2400:
2396:
2392:
2385:
2380:
2378:
2373:
2371:
2366:
2365:
2362:
2354:
2348:
2344:
2343:
2337:
2333:
2327:
2323:
2322:
2316:
2315:
2311:
2302:
2298:
2294:
2290:
2286:
2282:
2278:
2271:
2268:
2263:
2259:
2254:
2249:
2245:
2241:
2237:
2233:
2229:
2222:
2219:
2207:
2203:
2202:The Oregonian
2199:
2195:
2188:
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2180:
2176:
2171:
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2156:
2152:
2148:
2144:
2140:
2136:
2129:
2127:
2123:
2118:
2114:
2110:
2108:9780231135283
2104:
2100:
2099:
2091:
2088:
2083:
2079:
2075:
2069:
2065:
2060:
2059:
2050:
2047:
2042:
2038:
2034:
2030:
2023:
2021:
2019:
2017:
2015:
2013:
2009:
1997:
1993:
1992:
1987:
1985:
1977:
1974:
1971:
1967:
1966:
1960:
1957:
1952:
1946:
1942:
1941:
1933:
1930:
1925:
1921:
1917:
1913:
1909:
1908:Simpson, G.G.
1903:
1900:
1893:
1891:
1889:
1885:
1881:
1873:
1866:
1865:
1857:
1856:
1848:
1847:
1839:
1838:
1830:
1829:
1821:
1820:
1812:
1811:
1803:
1802:
1794:
1793:
1785:
1784:
1776:
1775:
1767:
1766:
1758:
1757:
1749:
1748:
1740:
1739:
1731:
1730:
1722:
1721:
1713:
1712:
1704:
1703:
1700:
1699:
1698:
1691:
1690:
1687:
1686:
1683:
1682:
1681:
1674:
1673:
1667:
1666:
1663:
1662:
1659:
1658:
1657:
1650:
1649:
1643:
1642:
1639:
1638:
1635:
1634:
1633:
1626:
1625:
1619:
1618:
1615:
1614:
1611:
1610:
1609:
1608:Paratomarctus
1602:
1601:
1595:
1592:
1591:
1588:
1587:
1579:
1578:
1575:
1574:
1573:
1566:
1565:
1562:
1561:
1558:
1557:
1556:
1549:
1548:
1542:
1541:Aelurodontina
1539:
1538:
1535:
1534:
1531:
1530:
1529:
1522:
1521:
1515:
1514:
1511:
1510:
1507:
1506:
1505:
1498:
1497:
1491:
1490:
1487:
1486:
1483:
1482:
1481:
1474:
1473:
1467:
1466:
1463:
1462:
1459:
1458:
1457:
1450:
1449:
1443:
1442:
1439:
1438:
1435:
1434:
1433:
1426:
1425:
1419:
1418:
1415:
1414:
1411:
1410:
1409:
1408:Metatomarctus
1402:
1401:
1395:
1394:
1391:
1390:
1382:
1381:
1378:
1377:
1376:
1369:
1368:
1365:
1364:
1361:
1360:
1359:
1358:Paracynarctus
1352:
1351:
1345:
1342:
1341:
1335:
1334:
1331:
1330:
1327:
1326:
1325:
1318:
1317:
1311:
1310:
1307:
1306:
1303:
1302:
1301:
1294:
1293:
1287:
1284:
1283:
1280:
1279:
1271:
1270:
1267:
1266:
1265:
1258:
1257:
1254:
1253:
1250:
1249:
1248:
1241:
1240:
1234:
1231:
1230:
1224:
1223:
1220:
1219:
1216:
1215:
1214:
1207:
1206:
1200:
1199:
1196:
1195:
1192:
1191:
1190:
1183:
1182:
1179:
1178:
1175:
1174:
1173:
1166:
1165:
1162:
1161:
1158:
1157:
1156:
1149:
1148:
1142:Borophaginae
1141:
1140:
1137:
1136:
1133:
1132:
1126:
1125:
1119:
1118:
1115:
1114:
1111:
1110:
1104:
1103:
1097:
1094:
1093:
1089:
1075:million years
1074:
1069:
1068:
1063:
1062:
1057:
1054:million years
1053:
1048:
1047:
1042:
1039:million years
1038:
1033:
1032:
1027:
1024:million years
1023:
1018:
1017:
1012:
1009:million years
1008:
1003:
1002:
1001:Paratomarctus
997:
996:
995:
991:
986:million years
985:
980:
979:
974:
971:million years
970:
965:
964:
959:
958:
956:million years
955:
950:
949:Aelurodontina
946:
941:million years
940:
935:
934:
929:
926:million years
925:
920:
919:
918:Paracynarctus
914:
913:
911:million years
910:
905:
901:
898:million years
897:
892:
891:
886:
883:million years
882:
877:
876:
871:
868:million years
867:
862:
861:
856:
853:million years
852:
847:
846:
841:
838:million years
837:
832:
831:
826:
823:million years
822:
817:
816:
815:Metatomarctus
811:
808:million years
807:
802:
801:
796:
793:million years
792:
787:
786:
781:
780:
778:million years
777:
772:
768:
763:million years
762:
757:
756:
751:
748:million years
747:
742:
741:
736:
735:
733:million years
732:
727:
723:
720:million years
719:
714:
713:
708:
705:million years
704:
699:
698:
693:
690:million years
689:
684:
683:
678:
675:million years
674:
669:
668:
663:
662:
661:
658:
657:
656:
652:
651:
650:
644:
639:
634:
626:
622:
620:
616:
612:
611:Xiaoming Wang
608:
607:
602:
601:
596:
595:
590:
589:
583:
581:
577:
573:
569:
568:top predators
565:
561:
557:
553:
548:
546:
542:
538:
537:
532:
531:
526:
525:
520:
519:
514:
510:
509:Aelurodontina
506:
502:
498:
494:
490:
486:
478:
477:
472:
471:
466:
465:
460:
453:
451:
449:
448:
443:
440:to powerful,
439:
435:
431:
430:
425:
411:
410:
402:
401:
393:
392:
389:
387:
383:
377:
376:
373:
372:
369:
367:
363:
354:
353:
347:
346:
343:
342:
339:
337:
333:
324:
323:
316:
312:
311:
307:
306:
302:
301:
295:
293:
290:million years
289:
284:
280:
276:
275:North America
272:
268:
264:
260:
256:
252:
243:
239:
234:
229:
224:
218:
215:
214:
211:
208:
205:
204:
201:
198:
195:
194:
191:
188:
185:
184:
181:
178:
175:
174:
171:
168:
165:
164:
161:
158:
155:
154:
149:
144:
140:
137:
135:
130:
126:
121:
116:
112:
105:
100:
95:
90:
85:
80:
75:
70:
65:
60:
55:
50:
44:
33:
30:
19:
3555:Borophagines
3490:Borophaginae
3489:
3464:
3437:C. l. youngi
3435:
3425:
3415:
3405:
3395:
3387:C. l. hattai
3385:
3375:
3365:
3357:C. l. fuscus
3355:
3345:
3335:
3325:
3315:
3305:
3292:
3280:
3267:
3255:
3245:
3235:
3225:
3215:
3205:
3195:
3178:
3166:
3153:
3141:
3131:
3121:
3111:
3078:S. pacivorus
3076:
3063:
3051:
3041:
3028:
3018:Theriodictis
3016:
3006:
2996:
2982:Cerdocyonina
2960:
2952:V. riffautae
2950:
2940:
2917:
2907:
2897:
2887:
2877:
2864:
2852:
2842:
2832:
2822:
2787:
2780:
2773:
2766:
2759:
2739:
2732:
2712:
2705:
2685:
2680:Protomarctus
2678:
2671:
2664:
2657:
2650:
2643:
2636:
2612:
2607:Cynarctoides
2605:
2598:Phlaocyonini
2585:
2578:
2571:
2564:
2553:Borophaginae
2552:
2537:
2530:
2523:
2516:
2509:
2502:
2497:Sunkahetanka
2495:
2488:
2481:
2474:
2451:
2440:
2341:
2320:
2284:
2280:
2270:
2235:
2231:
2221:
2210:. Retrieved
2197:
2187:
2142:
2138:
2097:
2090:
2057:
2049:
2032:
2028:
1999:. Retrieved
1989:
1984:Borophaginae
1983:
1976:
1965:Borophaginae
1964:
1959:
1939:
1932:
1915:
1911:
1902:
1877:
1695:
1694:
1678:
1677:
1654:
1653:
1630:
1629:
1606:
1605:
1570:
1569:
1553:
1552:
1526:
1525:
1504:Protomarctus
1502:
1501:
1478:
1477:
1454:
1453:
1430:
1429:
1406:
1405:
1373:
1372:
1356:
1355:
1322:
1321:
1298:
1297:
1262:
1261:
1247:Cynarctoides
1245:
1244:
1233:Phlaocyonini
1211:
1210:
1187:
1186:
1170:
1169:
1153:
1152:
1129:
1107:
1087:
1065:
1059:
1044:
1029:
1014:
999:
976:
961:
931:
916:
888:
875:Protomarctus
873:
858:
843:
828:
813:
798:
783:
753:
740:Cynarctoides
738:
726:Phlaocyonini
710:
695:
680:
665:
659:
648:
604:
598:
592:
586:
584:
549:
534:
528:
522:
516:
501:Phlaocyonini
493:phylogenetic
482:
474:
468:
462:
445:
427:
421:
380:
362:Borophaginae
361:
357:
327:
251:Borophaginae
250:
249:The extinct
248:
241:
223:Borophaginae
222:
132:
41:34β2.5
35:Borophaginae
29:
3307:C. l. alces
3247:C. etruscus
3237:C. edwardii
3217:C. arnensis
3113:Cynotherium
2962:V. skinneri
2889:U. galushai
2879:U. citrinus
2854:Prototocyon
2775:Protepicyon
2752:Borophagina
2666:Psalidocyon
2625:Borophagini
2566:Archaeocyon
2539:Ectopocynus
2511:Enhydrocyon
2476:Hesperocyon
1991:Fossilworks
1880:carnivorans
1656:Protepicyon
1594:Borophagina
1456:Psalidocyon
1286:Borophagini
1155:Archaeocyon
1031:Protepicyon
994:Borophagina
845:Psalidocyon
771:Borophagini
667:Archaeocyon
619:Procyonidae
518:Archaeocyon
513:Borophagina
429:Archaeocyon
277:during the
265:and extant
255:subfamilies
216:Subfamily:
3549:Categories
3269:C. latrans
2789:Borophagus
2698:Cynarctina
2687:Tephrocyon
2659:Euoplocyon
2525:Osbornodon
2490:Cynodesmus
2431:Caniformia
2429:Suborder:
2212:2016-09-03
2073:0792271343
1894:References
1874:Extinction
1697:Borophagus
1528:Tephrocyon
1432:Euoplocyon
1344:Cynarctina
1067:Osteoborus
1061:Borophagus
992:Subtribe β
947:Subtribe β
904:Cynarctina
902:Subtribe β
890:Tephrocyon
830:Euoplocyon
606:Osteoborus
600:Borophagus
572:ecosystems
545:Arikareean
505:Cynarctina
473:(middle),
3030:Lycalopex
3008:Protocyon
2834:Metalopex
2824:Leptocyon
2768:Carpocyon
2741:Aelurodon
2734:Tomarctus
2714:Cynarctus
2645:Desmocyon
2638:Cormocyon
2614:Phlaocyon
2587:Rhizocyon
2580:Otarocyon
2573:Oxetocyon
2532:Caedocyon
2504:Philotrox
2425:Carnivora
2405:Kingdom:
2301:254698991
2117:185095648
2041:2246/1588
2001:29 August
1924:2246/1104
1632:Carpocyon
1572:Aelurodon
1555:Tomarctus
1375:Cynarctus
1324:Desmocyon
1300:Cormocyon
1264:Phlaocyon
1213:Rhizocyon
1189:Otarocyon
1172:Oxetocyon
1016:Carpocyon
978:Aelurodon
963:Tomarctus
933:Cynarctus
800:Desmocyon
785:Cormocyon
758:30β19 Ma,
755:Phlaocyon
712:Rhizocyon
697:Otarocyon
682:Oxetocyon
615:Hyaenidae
588:Aelurodon
576:phylogeny
570:of their
560:dentition
539:, in the
536:Rhizocyon
530:Otarocyon
524:Oxetocyon
497:cladistic
479:(bottom).
438:omnivores
279:Oligocene
242:See text
200:Carnivora
166:Kingdom:
160:Eukaryota
134:Aelurodon
111:Oligocene
3519:47438146
3505:Q2729984
3499:Wikidata
3466:Category
3294:C. lupus
3143:Xenocyon
3133:Aenocyon
3065:Speothos
2998:Dusicyon
2919:U. webbi
2844:Nurocyon
2483:Mesocyon
2419:Mammalia
2413:Chordata
2411:Phylum:
2407:Animalia
2262:26285033
2238:: 7976.
2206:Archived
2179:26124128
2082:54079923
491:ages. A
434:Cenozoic
283:Pliocene
206:Family:
190:Mammalia
180:Chordata
176:Phylum:
170:Animalia
156:Domain:
136:stirtoni
2866:Urocyon
2808:Caninae
2782:Epicyon
2423:Order:
2417:Class:
2395:Canidae
2391:Extinct
2240:Bibcode
2170:4507235
2147:Bibcode
1968:in the
1888:canines
1680:Epicyon
1131:Caninae
1096:Canidae
1046:Epicyon
769:Tribe β
724:Tribe β
655:Canidae
653:Family
638:Canidae
594:Epicyon
580:Caninae
564:fossils
541:Orellan
489:Blancan
485:Orellan
467:(top),
454:Species
447:Epicyon
382:Caninae
317:
315:Canidae
271:endemic
267:Caninae
236:Genera
228:Simpson
210:Canidae
196:Order:
186:Class:
3123:Eucyon
3097:Canina
2973:Canini
2930:Vulpes
2349:
2328:
2299:
2260:
2177:
2167:
2115:
2105:
2080:
2070:
2066:β114.
1947:
1884:felids
617:, and
597:, and
533:, and
511:, and
313:
296:Origin
230:(1945)
3532:83323
3180:Canis
2297:S2CID
556:hyena
552:teeth
259:canid
3155:Cuon
2347:ISBN
2326:ISBN
2258:PMID
2175:PMID
2113:OCLC
2103:ISBN
2078:OCLC
2068:ISBN
2003:2018
1945:ISBN
1886:and
442:bear
288:33.5
49:Preκ
3514:EoL
2289:doi
2248:doi
2165:PMC
2155:doi
2143:112
2064:112
2037:hdl
2033:243
1920:hdl
1037:2.7
896:2.7
487:to
281:to
273:to
3551::
3529::
3516::
3501::
2295:.
2285:23
2283:.
2279:.
2256:.
2246:.
2234:.
2230:.
2204:.
2200:.
2196:.
2173:.
2163:.
2153:.
2141:.
2137:.
2125:^
2111:.
2076:.
2035:.
2031:.
2011:^
1988:.
1916:85
1914:.
1064:(=
1022:11
1007:11
954:15
909:10
791:10
776:27
761:11
746:12
731:20
621:.
603:(=
591:,
527:,
521:,
507:,
503:,
450:.
292:.
99:Pg
43:Ma
3433:β
3423:β
3413:β
3403:β
3393:β
3383:β
3373:β
3363:β
3353:β
3343:β
3333:β
3323:β
3313:β
3303:β
3278:β
3253:β
3243:β
3233:β
3223:β
3213:β
3203:β
3193:β
3164:β
3139:β
3129:β
3119:β
3109:β
3074:β
3049:β
3039:β
3014:β
3004:β
2994:β
2958:β
2948:β
2938:β
2915:β
2905:β
2895:β
2885:β
2875:β
2850:β
2840:β
2830:β
2820:β
2550:β
2464:β
2449:β
2383:e
2376:t
2369:v
2355:.
2334:.
2303:.
2291::
2264:.
2250::
2242::
2236:6
2215:.
2181:.
2157::
2149::
2119:.
2084:.
2043:.
2039::
2005:.
1986:"
1982:"
1953:.
1926:.
1922::
1073:7
1058:β
1052:2
1043:β
1028:β
1013:β
998:β
984:4
975:β
969:7
960:β
939:4
930:β
924:3
915:β
887:β
881:4
872:β
866:7
857:β
851:7
842:β
836:2
827:β
821:3
812:β
806:9
797:β
782:β
752:β
737:β
718:4
709:β
703:4
694:β
688:4
679:β
673:7
664:β
359:β
329:β
220:β
113:-
104:N
94:K
89:J
84:T
79:P
74:C
69:D
64:S
59:O
54:κ
20:)
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