820:
804:
150:
832:
111:
129:
488:
evolution were challenged by the work of Nancy Todd published in 2001 and 2005. Todd found that the different subspecies exhibited high variability in molar dimensions within subspecies, but low variability between the subspecies, and that not all of the subspecies clustered together with each other
1018:
Beden, M. 1987. Les faunes Plio-Pléistocène de la basse vallée de l’Omo (Éthiopie), Vol. 2: Les Eléphantidés (Mammalia-Proboscidea) (directed by Y. Coppens and F. C. Howell): 1-162. Cahiers de Paléontologie-Travaux de Paléontologie est-africaine. Centre
National de la Recherche Scientifique (CNRS).
764:-type stone tools, and the Olorgesailie Basin Member 1 Site 15 in Kenya, dated to 992–974,000 years ago, and the Nadung’a 4 site near Lake Turkana, Kenya, dating to approximately 700,000 years ago. The Barogali site in Djibouti, dating to 1.6-1.3 million years ago, where a disassociated specimen of
644:
individuals reaching 4 metres (13 ft) in length, and masses likely considerably in excess of 100 kilograms (220 lb). In 2024, Biswas, Chang and Tsai estimated its maximum shoulder height over 4.23 metres (13.9 ft), with the estimated body masses of 7 measured specimens ranging from
1076:
Stimpson, Christopher M.; Lister, Adrian; Parton, Ash; Clark-Balzan, Laine; Breeze, Paul S.; Drake, Nick A.; Groucutt, Huw S.; Jennings, Richard; Scerri, Eleanor M.L.; White, Tom S.; Zahir, Muhammad; Duval, Mathieu; Grün, Rainer; Al-Omari, Abdulaziz; Al
Murayyi, Khalid Sultan M. (July 2016).
759:
remains with cut marks and/or stone tools include Upper Bed II at the Bell's
Korongo site in Olduvai Gorge, dating to around 1.35 million years ago, which has been suggested to be the oldest site in the world with reliable evidence of elephant butchery, associated with
784:
and other bifaced tools, and displays cut marks and fracture marks indicative of butchery, though the fracturing of the skull, which has been suggested to be the result of an attempt to extract the brain, may alternatively be the result of postmortem
395:
The two deposits are not contemporaneous and the specimens from each locality are morphologically distinctive from each other, which has led to confusion about which locality represents the "typical" morphology of the species. The placement of
1032:
Larramendi, Asier; Zhang, Hanwen; Palombo, Maria Rita; Ferretti, Marco P. (February 2020). "The evolution of
Palaeoloxodon skull structure: Disentangling phylogenetic, sexually dimorphic, ontogenetic, and allometric morphological signals".
330:. During most of its existence, the species (in its broad sense) represented the dominant elephant species in East Africa. The species is divided into five roughly chronologically successive subspecies. While the type and latest subspecies
640:, Kenya, suggested to have been approximately 40 years old when it died, was estimated in a 2016 study to have measured 4.27 metres (14.0 ft) tall at the shoulder and weighed 12.3 tonnes (27,000 lb), with the tusks of some
1009:
Beden, M. 1983. Family
Elephantidae. In J. M. Harris (ed.), Koobi Fora Research Project. Vol. 2. The fossil Ungulates: Proboscidea, Perissodactyla, and Suidae: 40-129. Oxford: Clarendon Press.
751:
have been found associated with stone tools. In some cases like
Olduvai FLK, these are likely coincidental, but in others which bears cut marks, these likely represent evidence of butchery by
618:) in northern Israel, the Middle Pleistocene (c. 500,000 years ago) Ti's al Ghadah site in northern Saudi Arabia, and the late Middle Pleistocene Shishan Marsh site in Jordan, belongs to
819:
381:, a German paleontologist and geologist who had done the initial surveying of the gorge in 1913, and had collected a considerable number of fossils from the locality.
810:
803:
508:
and that the supposed subspecies substantially chronologically overlapped, though later work suggested that this chronological overlap was likely overstated.
1562:
1493:
1577:
1572:
1369:
981:
442:
primarily focusing on molar morphology identified five successive subspecies, from oldest to youngest with ages according to
Sanders (2023)
1567:
1557:
1000:
Dietrich, 1915 (Proboscidea, Elephantidae). Èvolution au cours du Plio-Pléistocène en
Afrique orientale. Geobios 13(6): 891-901. Lyon.
1128:"Faunal remains from recent excavations at Shishan Marsh 1 (SM1), a Late Lower Paleolithic open-air site in the Azraq Basin, Jordan"
559:, primarily known from the Late Pliocene-Early Pleistocene of the Indian subcontinent, and that this species should be placed into
831:
1552:
870:"Late Middle Pleistocene Elephants from Natodomeri, Kenya and the Disappearance of Elephas (Proboscidea, Mammalia) in Africa"
1079:"Middle Pleistocene vertebrate fossils from the Nefud Desert, Saudi Arabia: Implications for biogeography and palaeoecology"
772:) was found with numerous stone tools (probably Oldowan) created onsite, has also been suggested to be a butchery site. The
1127:
868:
Manthi, Fredrick Kyalo; Sanders, William J.; Plavcan, J. Michael; Cerling, Thure E.; Brown, Francis H. (September 2020).
662:
149:
391:
in
Ethiopia, and suggested that they were distinctive enough that they warranted being placed as the distinct species
1392:"Late Quaternary Proboscidean Sites in Africa and Eurasia with Possible or Probable Evidence for Hominin Involvement"
1126:
Pokines, James T.; Lister, Adrian M.; Ames, Christopher J. H.; Nowell, April; Cordova, Carlos E. (March 2019).
366:
838:
1471:
610:, including that from the earliest Middle Pleistocene (c. 780,000 years ago) Paleolithic archaeological site
513:
A 2020 PhD thesis by Steven Zhang, focusing primarily on skull morphology again challenged the monophyly of
653:
at the top of the skull is only weakly developed. The frons (forehead) is tall and biconvex. Like
Eurasian
1547:
1433:
711:
407:
1519:
732:
689:, with the molar teeth of later subspecies showing greater adaption to grazing than earlier subspecies.
362:
268:
248:
632:
Members of the species were larger than any living elephant. A large mostly complete male specimen of
349:
are uncertain, with it being suggested they are unrelated and should be elevated to separate species.
1480:
1306:
1257:
1210:
1139:
1090:
1042:
686:
1174:
1338:
1078:
1058:
921:
897:
716:
611:
555:
490:
327:
277:
144:
1524:
1506:
1413:
1365:
1330:
1322:
1275:
1226:
1155:
977:
889:
384:
323:
137:
1511:
1403:
1314:
1265:
1218:
1189:
1147:
1106:
1098:
1050:
969:
929:
881:
825:
Underside of the skull and lower jaws at the Gallery of Paleontology and Comparative Anatomy
720:
561:
589:
though he suggested that the two may be sister species), and questioned their placement in
701:
in Africa, at the end of the Early Pleistocene, around 800,000 years ago, a population of
606:
Outside its core East African distribution. It has also been suggested that material from
1246:"Land of the giants: Body mass estimates of Palaeoloxodon from the Pleistocene of Taiwan"
1310:
1261:
1211:"Estimating tusk masses in proboscideans: a comprehensive analysis and predictive model"
1143:
1094:
1046:
539:
subspecies were likely unrelated, and more closely related to true members of the genus
752:
546:
869:
1541:
1357:
1062:
901:
484:" as a succession of subspecies was accepted as orthodoxy. However, Beden's views of
416:
406:
was contested as early as 1942, when in a publication by Douglas Gordon MacInnes and
358:
340:
224:
61:
1295:"The role of behaviour in adaptive morphological evolution of African proboscideans"
1270:
1245:
1102:
1054:
963:
211:
110:
1342:
1222:
685:
regardless of true evolutionary relationships, are thought to have been dedicated
645:
4.5–15.3 tonnes (9,900–33,700 lb). In comparison to most Eurasian species of
933:
1465:
922:"Reanalysis of African Elephas recki: implications for time, space and taxonomy"
201:
36:
885:
670:
637:
388:
307:
128:
81:
46:
1456:
1417:
1326:
1279:
1230:
1159:
893:
786:
777:
607:
531:
378:
315:
161:
86:
1334:
1194:
1450:
1294:
1244:
Biswas, Deep Shubhra; Chang, Chun-Hsiang; Tsai, Cheng-Hsiu (2024-07-15).
1151:
319:
311:
181:
76:
71:
56:
51:
41:
1408:
1391:
1318:
1111:
569:
should be elevated to full species status as a true member of the genus
1498:
781:
761:
541:
402:
91:
66:
421:
though many later authors continued to place the species in the genus
410:'s posthumous monograph on fossil proboscideans it was suggested that
345:
the relationships of the other, chronologically earlier subspecies to
191:
171:
1427:
973:
529:
which genetic data shows is closely related to African elephants (
357:
The species was initially named from specimens found at Bed IV in
705:
migrated out of Africa, giving rise to the Eurasian radiation of
1485:
669:
show an increasing number of lamellae, and taller crown height (
1431:
1358:"Ecological Interactions of Elephantids in Pleistocene Eurasia"
493:, as would be expected if they formed a single species, with
387:
in 1942 described additional specimens of the species from
776:
specimen from Gesher Bnot Ya'akov is associated with an
1175:"Shoulder height, body mass and shape of proboscideans"
968:(1 ed.). Boca Raton: CRC Press. pp. 267–293.
452:
Beden, 1980 Early Pliocene-~3.2 million years ago (mya)
338:
are widely accepted to be closely related to Eurasian
480:
For decades after Beden's publications, his view of "
715:, is known from remains found across Africa of late
476:(Dietrich, 1916) 1.6/1.4 mya-late Middle Pleistocene
365:
in 1915, originally as a subspecies of the European
1440:
965:Evolution and Fossil Record of African Proboscidea
709:Its descendant taxon or last evolutionary stage,
438:Michel Beden's publications during the 1980s on
811:Gallery of Paleontology and Comparative Anatomy
573:. However, while Sanders (2023) accepted that
8:
1362:Human Paleoecology in the Levantine Corridor
118:Size comparison of a 40 year old adult male
747:At several sites across Africa, remains of
1428:
593:, while he considered the relationship of
127:
109:
20:
1407:
1269:
1193:
1110:
855:
799:
665:outwards. Over time the molar teeth of
525:has a close relationship with Eurasian
347:P. recki recki and P. recki ileretensis
577:was unlikely to be closely related to
7:
1027:
1025:
957:
955:
953:
951:
949:
947:
945:
943:
915:
913:
911:
863:
861:
859:
657:species, the premaxillary bones of
962:Sanders, William J. (2023-07-07).
14:
723:age. Following the extinction of
1293:Lister, Adrian M. (2013-06-26).
1209:Larramendi, Asier (2023-12-10).
830:
818:
802:
495:E. r. brumpti E. r. shungurensis
148:
32:Late Pliocene–Middle Pleistocene
1563:Pleistocene species extinctions
1364:, Oxbow Books, pp. 53–60,
1271:10.1016/j.quascirev.2024.108761
1103:10.1016/j.quascirev.2016.05.016
1055:10.1016/j.quascirev.2019.106090
920:Todd, Nancy E. (January 2005).
649:the parieto-occipital crest of
517:as a whole, finding that while
874:Journal of Mammalian Evolution
731:it was replaced by the modern
581:he rejected the synonymity of
1:
1578:Fossil taxa described in 1894
1573:Pleistocene mammals of Africa
1223:10.1080/08912963.2023.2286272
1182:Acta Palaeontologica Polonica
501:being placed separately from
300:, often known by the synonym
934:10.1016/j.quaint.2004.04.015
470:Beden, 1987 ~1.8-1.4/1.6 mya
464:Arambourg, 1947 ~2.3-1.8 mya
414:should instead be placed in
1390:Haynes, Gary (March 2022).
697:Following the emergence of
661:containing the tusks flare
545:(which contains the living
16:Extinct species of elephant
1594:
1568:Pliocene mammals of Africa
1356:Lister, Adrian M. (2004),
1250:Quaternary Science Reviews
1083:Quaternary Science Reviews
1035:Quaternary Science Reviews
886:10.1007/s10914-019-09474-9
120:Palaeoloxodon recki atavus
1558:Pleistocene proboscideans
614:(otherwise attributed to
334:as well as the preceding
283:
276:
254:
247:
145:Scientific classification
143:
135:
126:
117:
108:
23:
926:Quaternary International
743:Relationship with humans
681:All named subspecies of
549:). Zhang suggested that
458:Beden, 1980 ~3.2-2.3 mya
367:straight-tusked elephant
1173:Larramendi, A. (2016).
371:Palaeoloxodon antiquus,
1553:Pliocene proboscideans
1195:10.4202/app.00136.2014
712:Palaeoloxodon iolensis
587:"Elephas" planifrons (
408:Henry Fairfield Osborn
375:Elephas antiquus recki
1520:Paleobiology Database
839:Museum für Naturkunde
733:African bush elephant
503:E. r. ileretensis and
377:. It was named after
369:, what is now called
363:Wilhelm Otto Dietrich
314:native to Africa and
1152:10.1017/qua.2018.113
774:P. antiquus/P. recki
693:Evolutionary history
556:"Elephas" planifrons
553:was synonymous with
336:P. recki ileretensis
136:Life restoration by
1472:Palaeoloxodon recki
1409:10.3390/quat5010018
1319:10.1038/nature12275
1311:2013Natur.500..331L
1262:2024QSRv..33608761B
1144:2019QuRes..91..768P
1132:Quaternary Research
1095:2016QSRv..143...13S
1047:2020QSRv..22906090L
755:. Sites containing
612:Gesher Bnot Ya'akov
535:), the other named
297:Palaeoloxodon recki
261:Palaeoloxodon recki
25:Palaeoloxodon recki
1215:Historical Biology
928:. 126–128: 65–72.
737:Loxodonta africana
717:Middle Pleistocene
491:cladistic analysis
456:E. r. shungurensis
328:Middle Pleistocene
1535:
1534:
1507:Open Tree of Life
1434:Taxon identifiers
1371:978-1-78570-965-4
1305:(7462): 331–334.
983:978-1-315-11891-8
837:Lower jaw at the
768:(suggested to be
519:E. r. ileretensis
468:E. r. ileretensis
429:as a subgenus of
385:Camille Arambourg
324:Early Pleistocene
293:
292:
1585:
1528:
1527:
1515:
1514:
1502:
1501:
1489:
1488:
1476:
1475:
1474:
1461:
1460:
1459:
1429:
1422:
1421:
1411:
1387:
1381:
1380:
1379:
1378:
1353:
1347:
1346:
1290:
1284:
1283:
1273:
1241:
1235:
1234:
1206:
1200:
1199:
1197:
1179:
1170:
1164:
1163:
1123:
1117:
1116:
1114:
1073:
1067:
1066:
1029:
1020:
1016:
1010:
1007:
1001:
996:Beden, M. 1980.
994:
988:
987:
959:
938:
937:
917:
906:
905:
865:
834:
822:
806:
721:Late Pleistocene
601:to be uncertain.
591:Phanagoroloxodon
562:Phanagoroloxodon
263:
259:
223:
153:
152:
131:
122:from Koobi Fora
113:
103:
33:
29:Temporal range:
21:
1593:
1592:
1588:
1587:
1586:
1584:
1583:
1582:
1538:
1537:
1536:
1531:
1523:
1518:
1510:
1505:
1497:
1492:
1484:
1479:
1470:
1469:
1464:
1455:
1454:
1449:
1436:
1426:
1425:
1389:
1388:
1384:
1376:
1374:
1372:
1355:
1354:
1350:
1292:
1291:
1287:
1243:
1242:
1238:
1208:
1207:
1203:
1177:
1172:
1171:
1167:
1125:
1124:
1120:
1075:
1074:
1070:
1031:
1030:
1023:
1017:
1013:
1008:
1004:
995:
991:
984:
961:
960:
941:
919:
918:
909:
867:
866:
857:
852:
847:
846:
845:
842:
835:
826:
823:
814:
807:
795:
770:E. recki atavus
745:
695:
679:
634:P. recki atavus
630:
579:E. recki recki,
355:
288:Dietrich, 1894
272:
265:
257:
256:
243:
221:
147:
104:
102:
101:
100:
99:
94:
89:
84:
79:
74:
69:
64:
59:
54:
49:
44:
39:
31:
30:
27:
17:
12:
11:
5:
1591:
1589:
1581:
1580:
1575:
1570:
1565:
1560:
1555:
1550:
1540:
1539:
1533:
1532:
1530:
1529:
1516:
1503:
1490:
1477:
1462:
1446:
1444:
1438:
1437:
1432:
1424:
1423:
1382:
1370:
1348:
1285:
1236:
1201:
1165:
1138:(2): 768–791.
1118:
1068:
1021:
1011:
1002:
989:
982:
974:10.1201/b20016
939:
907:
880:(3): 483–495.
854:
853:
851:
848:
844:
843:
836:
829:
827:
824:
817:
815:
808:
801:
798:
797:
796:
794:
791:
753:archaic humans
744:
741:
707:Palaeoloxodon.
703:P. recki recki
694:
691:
678:
675:
659:P. recki recki
651:P. recki recki
647:Palaeoloxodon,
629:
626:
620:P. recki recki
599:E. recki recki
547:Asian elephant
527:Palaeoloxodon.
523:E. recki recki
478:
477:
471:
465:
459:
453:
361:, Tanzania by
354:
351:
332:P. recki recki
303:Elephas recki,
291:
290:
281:
280:
274:
273:
266:
252:
251:
245:
244:
236:
234:
230:
229:
219:
215:
214:
209:
205:
204:
199:
195:
194:
189:
185:
184:
179:
175:
174:
169:
165:
164:
159:
155:
154:
141:
140:
138:Mauricio Antón
133:
132:
124:
123:
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:
1590:
1579:
1576:
1574:
1571:
1569:
1566:
1564:
1561:
1559:
1556:
1554:
1551:
1549:
1548:Palaeoloxodon
1546:
1545:
1543:
1526:
1521:
1517:
1513:
1508:
1504:
1500:
1495:
1491:
1487:
1482:
1478:
1473:
1467:
1463:
1458:
1452:
1448:
1447:
1445:
1443:
1442:Elephas recki
1439:
1435:
1430:
1419:
1415:
1410:
1405:
1401:
1397:
1393:
1386:
1383:
1373:
1367:
1363:
1359:
1352:
1349:
1344:
1340:
1336:
1332:
1328:
1324:
1320:
1316:
1312:
1308:
1304:
1300:
1296:
1289:
1286:
1281:
1277:
1272:
1267:
1263:
1259:
1255:
1251:
1247:
1240:
1237:
1232:
1228:
1224:
1220:
1216:
1212:
1205:
1202:
1196:
1191:
1187:
1183:
1176:
1169:
1166:
1161:
1157:
1153:
1149:
1145:
1141:
1137:
1133:
1129:
1122:
1119:
1113:
1108:
1104:
1100:
1096:
1092:
1088:
1084:
1080:
1072:
1069:
1064:
1060:
1056:
1052:
1048:
1044:
1040:
1036:
1028:
1026:
1022:
1015:
1012:
1006:
1003:
999:
998:Elephas recki
993:
990:
985:
979:
975:
971:
967:
966:
958:
956:
954:
952:
950:
948:
946:
944:
940:
935:
931:
927:
923:
916:
914:
912:
908:
903:
899:
895:
891:
887:
883:
879:
875:
871:
864:
862:
860:
856:
849:
840:
833:
828:
821:
816:
812:
809:Skull at the
805:
800:
792:
790:
788:
783:
779:
775:
771:
767:
763:
758:
754:
750:
742:
740:
738:
734:
730:
726:
722:
718:
714:
713:
708:
704:
700:
692:
690:
688:
684:
676:
674:
672:
668:
664:
660:
656:
655:Palaeoloxodon
652:
648:
643:
639:
635:
627:
625:
624:
621:
617:
613:
609:
604:
603:
600:
596:
592:
588:
584:
583:E. r. brumpti
580:
576:
575:E. r. brumpti
572:
568:
564:
563:
558:
557:
552:
551:E. r. brumpti
548:
544:
543:
538:
534:
533:
528:
524:
520:
516:
515:Elephas recki
511:
510:
507:
504:
500:
496:
492:
487:
486:Elephas recki
483:
482:Elephas recki
475:
472:
469:
466:
463:
460:
457:
454:
451:
450:E. r. brumpti
448:
447:
446:
444:
441:
440:Elephas recki
436:
435:
432:
428:
427:Palaeoloxodon
424:
420:
418:
417:Palaeoloxodon
413:
412:Elephas recki
409:
405:
404:
400:in the genus
399:
394:
390:
386:
383:
380:
376:
372:
368:
364:
360:
359:Olduvai Gorge
352:
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286:Elephas recki
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249:Binomial name
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240:P. recki
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1112:10072/142575
1086:
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1034:
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212:Elephantidae
119:
24:
18:
1466:Wikispecies
683:P/E. recki,
628:Description
616:P. antiquus
474:E. r. recki
310:species of
202:Proboscidea
1542:Categories
1396:Quaternary
1377:2020-04-14
1256:: 108761.
1041:: 106090.
850:References
671:hypsodonty
638:Koobi Fora
389:Omo Valley
1418:2571-550X
1402:(1): 18.
1327:0028-0836
1280:0277-3791
1231:0891-2963
1160:0033-5894
1089:: 13–36.
1063:213676377
902:198190671
894:1064-7554
787:trampling
778:Acheulean
663:laterally
608:West Asia
532:Loxodonta
425:treating
393:E. recki.
379:Hans Reck
318:from the
316:West Asia
233:Species:
168:Kingdom:
162:Eukaryota
1451:Wikidata
1335:23803767
1217:: 1–14.
841:. Berlin
813:, France
766:P. recki
757:P. recki
749:P. recki
729:iolensis
699:P. recki
667:P. recki
642:P. recki
565:, while
537:E. recki
423:Elephas,
353:Taxonomy
320:Pliocene
312:elephant
278:Synonyms
269:Dietrich
208:Family:
192:Mammalia
182:Chordata
178:Phylum:
172:Animalia
158:Domain:
1512:3607199
1499:5936006
1486:4454723
1457:Q693352
1307:Bibcode
1258:Bibcode
1140:Bibcode
1091:Bibcode
1043:Bibcode
793:Gallery
782:handaxe
762:Oldowan
687:grazers
677:Ecology
571:Elephas
542:Elephas
431:Elephas
403:Elephas
326:to the
308:extinct
271:, 1894)
258:†
237:†
218:Genus:
198:Order:
188:Class:
98:↓
1525:159800
1416:
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1343:883007
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1299:Nature
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306:is an
1339:S2CID
1178:(PDF)
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978:ISBN
890:ISSN
521:and
497:and
37:PreꞒ
1481:EoL
1404:doi
1315:doi
1303:500
1266:doi
1254:336
1219:doi
1190:doi
1148:doi
1107:hdl
1099:doi
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