1107:
121:
1598:
1688:
1604:
405:, and though both genera/species are the only currently diagnosable lithornithids several other contemporary species probably belong to separate species. Comparisons with other palaeognaths render the possibility of both birds being different sexes of the same species unlikely, and both
361:
with long primaries and remiges. In AMNH 30578 most of it is damaged due to post-mortem decomposition, with disorganised patches in the pectoral and pelvic region and the left wing traces and impressions being damaged, but the right wing is mostly intact, even showing evidence of
458:
Nesbitt, Sterling J.; Clarke, Julia A., The anatomy and taxonomy of the exquisitely preserved Green River
Formation (early Eocene) lithornithids (Aves) and the relationships of Lithornithidae. (Bulletin of the American Museum of Natural History, no. 406),
299:, dating to around 51 million years ago. It is an exceptionally well preserved specimen, preserving most of the post-cranial skeleton aside from the femur and associated pelvic region, as well as soft-tissue impressions of feathers, skin,
366:; in AMNH AMNH 30560 a wing is similarly well preserved. It is unclear if it had a tail, as the left wing feathers block the caudal region in AMNH 30578, but other lithornithids lack tail feathers.
430:
vindicates previous statements of competent flying in lithornithids, indicating that not only they were capable of sustained flapping but also migratory behaviour.
1106:
1806:
1755:
528:
1811:
418:
As mentioned above, skull and limb proportions are different among both species, implying that they occupied different ecological niches.
1801:
1816:
1597:
521:
120:
599:
1796:
1708:
514:
288:
1768:
1691:
392:
292:
268:
1149:
936:
883:
653:
340:
1006:
1603:
992:
919:
439:
1668:
1633:
1346:
891:
307:, was found in the vicinity, composed of a similarly complete most mostly disarticulate skeleton.
626:
115:
1773:
1661:
1326:
999:
1209:
1013:
1654:
1125:
1073:
968:
905:
492:
232:
1640:
1288:
961:
834:
482:
1166:
982:
898:
818:
584:
411:
401:
379:
335:
326:
316:
1746:
1519:
781:
198:
185:
391:
Studies suggest that there was bare minimum of two species-level lithornithids in the
1790:
1492:
1428:
1336:
1046:
1024:
811:
723:
710:
665:
537:
350:
172:
64:
1647:
1450:
1366:
876:
866:
300:
256:
271:, its are exceptionally well preserved, bearing impressions of skin and feathers.
1731:
1740:
1547:
1535:
1508:
1468:
1459:
1407:
1373:
1281:
1216:
1158:
1036:
744:
638:
260:
39:
1621:
1478:
1421:
1391:
1295:
1254:
1189:
1137:
797:
755:
311:
310:
It is diagnosed by several features of the skull and pelvis as well as having
84:
49:
1556:
1501:
1435:
1357:
951:
929:
851:
804:
766:
611:
132:
89:
33:
496:
1725:
1573:
1563:
1400:
1274:
1247:
1199:
562:
331:
152:
79:
74:
59:
54:
44:
1760:
1414:
1380:
1310:
1265:
1177:
1054:
363:
358:
345:
296:
264:
252:
94:
69:
314:
IV and II subequal in distal extent. It differs from the contemporary
556:
506:
487:
470:
248:
142:
107:
1702:
324:
the skull is shorter than the humerus, while in inverse happens in
469:
Torres, Christopher R.; Norell, Mark A.; Clarke, Julia A. (2019).
568:
162:
1706:
1619:
1123:
708:
549:
510:
471:"Estimating Flight Style of Early Eocene Stem Palaeognath Bird
1234:
1569:
1531:
1489:
1474:
1342:
1332:
1322:
1231:
1205:
1195:
1185:
1146:
1042:
1032:
979:
948:
916:
863:
847:
778:
762:
751:
740:
395:
at a given time, suggesting a high local diversity.
1715:
1546:
1518:
1488:
1458:
1449:
1390:
1356:
1309:
1264:
1230:
1176:
1145:
1136:
1072:
1023:
978:
947:
915:
862:
832:
777:
721:
664:
637:
610:
583:
409:specimens belong to adult birds, not juveniles of
354:in a less curved and more gracile scapular blade.
383:, as sister-taxa to all other lithornithids.
522:
8:
1703:
1616:
1524:
1455:
1315:
1239:
1142:
1133:
1120:
1069:
840:
733:
729:
718:
705:
643:
616:
589:
580:
546:
529:
515:
507:
20:
486:
399:indeed co-existed directly at least with
330:-, as well as a proportionally narrower
451:
247:is an extinct genus of bird from the
7:
303:and claw sheaths. Another specimen,
267:. Like many other fossils from the
259:, a member of a lineage of flying
14:
377:is recovered as a sister taxa to
1807:Paleogene birds of North America
1687:
1686:
1602:
1596:
1105:
119:
357:Feather imprints show abundant
263:, distantly related to modern
1:
1812:Fossil taxa described in 2016
1833:
1682:
1628:
1615:
1593:
1527:
1318:
1242:
1132:
1119:
1102:
1068:
843:
736:
732:
717:
704:
646:
619:
592:
579:
545:
279:The original specimen of
275:Discovery and description
228:
223:
116:Scientific classification
114:
23:
1802:Eocene genus extinctions
475:grandei(Lithornithidae)"
320:by a shorter skull - in
1817:Paleontology in Wyoming
16:Extinct genus of birds
1769:Paleobiology Database
479:The Anatomical Record
393:Green River Formation
293:Green River Formation
269:Green River Formation
341:Lithornis promiscuus
440:Dinosaur coloration
289:Fossil Butte Member
287:, was found in the
218:Nesbitt et al. 2016
426:A recent study on
344:in aspects of the
1784:
1783:
1709:Taxon identifiers
1700:
1699:
1678:
1677:
1611:
1610:
1591:
1590:
1587:
1586:
1583:
1582:
1445:
1444:
1305:
1304:
1226:
1225:
1150:Opisthodactylidae
1126:Notopalaeognathae
1115:
1114:
1100:
1099:
1096:
1095:
1092:
1091:
1074:Notopalaeognathae
1064:
1063:
828:
827:
700:
699:
696:
695:
692:
691:
688:
687:
684:
683:
334:shaft and longer
240:
239:
233:Calciavis grandei
219:
186:Lithornithiformes
1824:
1777:
1776:
1764:
1763:
1751:
1750:
1749:
1736:
1735:
1734:
1704:
1690:
1689:
1641:Incognitoolithus
1617:
1606:
1601:
1600:
1571:
1533:
1525:
1491:
1476:
1456:
1344:
1334:
1324:
1316:
1240:
1235:Dinornithiformes
1233:
1207:
1197:
1187:
1148:
1143:
1134:
1121:
1110:
1109:
1070:
1044:
1034:
981:
950:
918:
865:
849:
841:
835:Struthioniformes
780:
764:
753:
742:
734:
730:
719:
706:
644:
617:
590:
581:
574:
573:
547:
531:
524:
517:
508:
501:
500:
490:
488:10.1002/ar.24207
466:
460:
456:
217:
210:
197:
184:
124:
123:
106:
36:
29:Temporal range:
21:
1832:
1831:
1827:
1826:
1825:
1823:
1822:
1821:
1787:
1786:
1785:
1780:
1772:
1767:
1759:
1754:
1745:
1744:
1739:
1730:
1729:
1724:
1711:
1701:
1696:
1674:
1624:
1607:
1595:
1579:
1542:
1514:
1484:
1441:
1386:
1352:
1301:
1260:
1222:
1172:
1167:Opisthodactylus
1128:
1111:
1104:
1088:
1060:
1019:
983:Ergilornithidae
974:
943:
937:Galligeranoides
911:
899:Palaeophasianus
884:Galligeranoides
858:
837:
824:
819:Pseudocrypturus
773:
726:
713:
680:
660:
633:
606:
600:Avemetatarsalia
585:Avemetatarsalia
575:
552:
551:
541:
535:
505:
504:
468:
467:
463:
457:
453:
448:
436:
424:
412:Pseudocrypturus
402:Pseudocrypturus
389:
380:Pseudocrypturus
372:
336:tarsometatarsus
327:Pseudocrypturus
317:Pseudocrypturus
277:
216:
208:
195:
182:
118:
110:
105:
104:
103:
102:
97:
92:
87:
82:
77:
72:
67:
62:
57:
52:
47:
42:
31:
30:
27:
17:
12:
11:
5:
1830:
1828:
1820:
1819:
1814:
1809:
1804:
1799:
1797:Lithornithidae
1789:
1788:
1782:
1781:
1779:
1778:
1765:
1752:
1737:
1721:
1719:
1713:
1712:
1707:
1698:
1697:
1695:
1694:
1683:
1680:
1679:
1676:
1675:
1673:
1672:
1665:
1658:
1651:
1644:
1637:
1629:
1626:
1625:
1620:
1613:
1612:
1609:
1608:
1594:
1592:
1589:
1588:
1585:
1584:
1581:
1580:
1578:
1577:
1567:
1560:
1552:
1550:
1544:
1543:
1541:
1540:
1528:
1522:
1520:Casuariiformes
1516:
1515:
1513:
1512:
1505:
1497:
1495:
1493:Aepyornithidae
1486:
1485:
1483:
1482:
1472:
1464:
1462:
1453:
1447:
1446:
1443:
1442:
1440:
1439:
1432:
1425:
1418:
1411:
1404:
1396:
1394:
1388:
1387:
1385:
1384:
1377:
1370:
1362:
1360:
1354:
1353:
1351:
1350:
1340:
1330:
1319:
1313:
1307:
1306:
1303:
1302:
1300:
1299:
1292:
1285:
1278:
1270:
1268:
1262:
1261:
1259:
1258:
1251:
1243:
1237:
1228:
1227:
1224:
1223:
1221:
1220:
1213:
1203:
1193:
1182:
1180:
1174:
1173:
1171:
1170:
1163:
1154:
1152:
1140:
1130:
1129:
1124:
1117:
1116:
1113:
1112:
1103:
1101:
1098:
1097:
1094:
1093:
1090:
1089:
1087:
1086:
1085:
1084:
1078:
1076:
1066:
1065:
1062:
1061:
1059:
1058:
1051:
1040:
1029:
1027:
1021:
1020:
1018:
1017:
1010:
1007:Sinoergilornis
1003:
996:
988:
986:
976:
975:
973:
972:
965:
957:
955:
945:
944:
942:
941:
933:
925:
923:
913:
912:
910:
909:
902:
895:
888:
880:
872:
870:
860:
859:
857:
856:
844:
838:
833:
830:
829:
826:
825:
823:
822:
815:
808:
801:
794:
786:
784:
782:Lithornithidae
775:
774:
772:
771:
760:
749:
737:
727:
722:
715:
714:
709:
702:
701:
698:
697:
694:
693:
690:
689:
686:
685:
682:
681:
679:
678:
677:
676:
670:
668:
662:
661:
659:
658:
657:
656:
647:
641:
635:
634:
632:
631:
630:
629:
620:
614:
608:
607:
605:
604:
603:
602:
593:
587:
577:
576:
572:
571:
565:
559:
550:
543:
542:
536:
534:
533:
526:
519:
511:
503:
502:
461:
450:
449:
447:
444:
443:
442:
435:
432:
423:
420:
388:
385:
371:
368:
276:
273:
238:
237:
226:
225:
221:
220:
206:
202:
201:
199:Lithornithidae
193:
189:
188:
180:
176:
175:
170:
166:
165:
160:
156:
155:
150:
146:
145:
140:
136:
135:
130:
126:
125:
112:
111:
100:
99:
98:
93:
88:
83:
78:
73:
68:
63:
58:
53:
48:
43:
38:
37:
28:
15:
13:
10:
9:
6:
4:
3:
2:
1829:
1818:
1815:
1813:
1810:
1808:
1805:
1803:
1800:
1798:
1795:
1794:
1792:
1775:
1770:
1766:
1762:
1757:
1753:
1748:
1742:
1738:
1733:
1727:
1723:
1722:
1720:
1718:
1714:
1710:
1705:
1693:
1685:
1684:
1681:
1671:
1670:
1666:
1664:
1663:
1659:
1657:
1656:
1652:
1650:
1649:
1645:
1643:
1642:
1638:
1636:
1635:
1631:
1630:
1627:
1623:
1618:
1614:
1605:
1599:
1576:
1575:
1568:
1566:
1565:
1561:
1559:
1558:
1554:
1553:
1551:
1549:
1545:
1538:
1537:
1530:
1529:
1526:
1523:
1521:
1517:
1511:
1510:
1506:
1504:
1503:
1499:
1498:
1496:
1494:
1487:
1481:
1480:
1473:
1471:
1470:
1466:
1465:
1463:
1461:
1457:
1454:
1452:
1448:
1438:
1437:
1433:
1431:
1430:
1426:
1424:
1423:
1419:
1417:
1416:
1412:
1410:
1409:
1405:
1403:
1402:
1398:
1397:
1395:
1393:
1389:
1383:
1382:
1378:
1376:
1375:
1371:
1369:
1368:
1364:
1363:
1361:
1359:
1355:
1349:
1348:
1341:
1339:
1338:
1337:Querandiornis
1331:
1329:
1328:
1321:
1320:
1317:
1314:
1312:
1308:
1298:
1297:
1293:
1291:
1290:
1286:
1284:
1283:
1279:
1277:
1276:
1275:Anomalopteryx
1272:
1271:
1269:
1267:
1263:
1257:
1256:
1252:
1250:
1249:
1245:
1244:
1241:
1238:
1236:
1229:
1219:
1218:
1214:
1212:
1211:
1204:
1202:
1201:
1194:
1192:
1191:
1184:
1183:
1181:
1179:
1175:
1169:
1168:
1164:
1161:
1160:
1156:
1155:
1153:
1151:
1144:
1141:
1139:
1135:
1131:
1127:
1122:
1118:
1108:
1082:
1081:
1080:
1079:
1077:
1075:
1071:
1067:
1057:
1056:
1052:
1049:
1048:
1047:Pachystruthio
1041:
1039:
1038:
1031:
1030:
1028:
1026:
1025:Struthionidae
1022:
1016:
1015:
1011:
1009:
1008:
1004:
1002:
1001:
997:
995:
994:
993:Amphipelargus
990:
989:
987:
984:
977:
971:
970:
966:
964:
963:
959:
958:
956:
953:
946:
939:
938:
934:
932:
931:
927:
926:
924:
921:
920:Palaeotididae
914:
908:
907:
903:
901:
900:
896:
894:
893:
889:
886:
885:
881:
879:
878:
874:
873:
871:
868:
861:
854:
853:
846:
845:
842:
839:
836:
831:
821:
820:
816:
814:
813:
812:Paracathartes
809:
807:
806:
802:
800:
799:
795:
793:
792:
788:
787:
785:
783:
776:
769:
768:
761:
758:
757:
750:
747:
746:
739:
738:
735:
731:
728:
725:
724:Palaeognathae
720:
716:
712:
711:Palaeognathae
707:
703:
674:
673:
672:
671:
669:
667:
666:Palaeognathae
663:
655:
651:
650:
649:
648:
645:
642:
640:
636:
628:
624:
623:
622:
621:
618:
615:
613:
609:
601:
597:
596:
595:
594:
591:
588:
586:
582:
578:
570:
566:
564:
560:
558:
554:
553:
548:
544:
539:
532:
527:
525:
520:
518:
513:
512:
509:
498:
494:
489:
484:
480:
476:
474:
465:
462:
455:
452:
445:
441:
438:
437:
433:
431:
429:
421:
419:
416:
414:
413:
408:
404:
403:
398:
394:
386:
384:
382:
381:
376:
369:
367:
365:
360:
355:
353:
352:
351:Paracathartes
347:
343:
342:
337:
333:
329:
328:
323:
319:
318:
313:
308:
306:
302:
298:
294:
290:
286:
282:
274:
272:
270:
266:
262:
258:
254:
250:
246:
245:
236:
235:
234:
227:
222:
215:
214:
207:
204:
203:
200:
194:
191:
190:
187:
181:
178:
177:
174:
173:Palaeognathae
171:
168:
167:
164:
161:
158:
157:
154:
151:
148:
147:
144:
141:
138:
137:
134:
131:
128:
127:
122:
117:
113:
109:
96:
91:
86:
81:
76:
71:
66:
61:
56:
51:
46:
41:
35:
26:
22:
19:
1716:
1669:Tsondabornis
1667:
1660:
1653:
1648:Medioolithus
1646:
1639:
1634:Diamantornis
1632:
1572:
1562:
1555:
1534:
1507:
1500:
1477:
1467:
1451:Novaeratitae
1434:
1427:
1420:
1413:
1406:
1399:
1379:
1372:
1367:Crypturellus
1365:
1347:Roveretornis
1345:
1335:
1325:
1294:
1287:
1280:
1273:
1255:Megalapteryx
1253:
1246:
1215:
1208:
1198:
1188:
1165:
1157:
1053:
1045:
1035:
1012:
1005:
998:
991:
967:
960:
935:
928:
904:
897:
892:Geranodornis
890:
882:
877:Eogeranoides
875:
867:Geranoididae
850:
817:
810:
803:
796:
790:
789:
765:
754:
743:
478:
472:
464:
454:
427:
425:
417:
410:
406:
400:
396:
390:
378:
374:
373:
356:
349:
339:
325:
321:
315:
309:
304:
284:
280:
278:
261:palaeognaths
257:lithornithid
243:
242:
241:
231:
229:
212:
211:
169:Infraclass:
24:
18:
1741:Wikispecies
1548:Casuariidae
1536:Diogenornis
1509:Mullerornis
1460:Apterygidae
1408:Nothoprocta
1374:Nothocercus
1289:Euryapteryx
1159:Diogenornis
1037:Orientornis
745:Asteriornis
654:Maniraptora
639:Maniraptora
538:Palaeognath
348:, and from
312:metatarsals
305:AMNH 30560
301:foot scales
1791:Categories
1662:Psammornis
1479:Proapteryx
1422:Rhynchotus
1392:Nothurinae
1327:Miniothura
1296:Pachyornis
1190:Heterorhea
1138:Rheiformes
1083:see below↓
1000:Ergilornis
798:Fissuravis
756:Eremopezus
675:see below↓
459:2016-06-30
446:References
285:AMNH 30578
255:. It is a
1747:Calciavis
1732:Q27990781
1717:Calciavis
1557:Casuarius
1502:Aepyornis
1436:Tinamotis
1429:Taoniscus
1358:Tinaminae
1311:Tinamidae
1210:Protorhea
1014:Urmiornis
952:Eogruidae
930:Palaeotis
852:Remiornis
805:Lithornis
791:Calciavis
767:Limenavis
627:Theropoda
612:Theropoda
555:Kingdom:
473:Calciavis
428:Calciavis
407:Calciavis
397:Calciavis
375:Calciavis
370:Phylogeny
322:Calciavis
281:Calciavis
244:Calciavis
213:Calciavis
139:Kingdom:
133:Eukaryota
25:Calciavis
1761:11162362
1726:Wikidata
1692:Category
1655:Namornis
1574:Emuarius
1564:Dromaius
1401:Eudromia
1248:Dinornis
1200:Hinasuri
1055:Struthio
969:Sonogrus
906:Paragrus
563:Chordata
561:Phylum:
557:Animalia
497:31313482
434:See also
364:barbules
332:coracoid
224:Species
192:Family:
153:Chordata
149:Phylum:
143:Animalia
129:Domain:
32:51
1469:Apteryx
1415:Nothura
1381:Tinamus
1266:Emeidae
1178:Rheidae
567:Class:
387:Ecology
359:plumage
346:ischium
338:, from
297:Wyoming
291:of the
265:ratites
253:Wyoming
205:Genus:
179:Order:
159:Class:
101:↓
1774:379323
1622:Ootaxa
962:Eogrus
540:genera
495:
422:Flight
249:Eocene
108:Eocene
1282:Emeus
1756:GBIF
1217:Rhea
652:see
625:see
598:see
569:Aves
493:PMID
163:Aves
40:PreꞒ
483:doi
251:of
1793::
1771::
1758::
1743::
1728::
491:.
481:.
477:.
415:.
295:,
283:,
90:Pg
34:Ma
1570:†
1539:?
1532:†
1490:†
1475:†
1343:†
1333:†
1323:†
1232:†
1206:†
1196:†
1186:†
1162:?
1147:†
1050:?
1043:†
1033:†
985:?
980:†
954:?
949:†
940:?
922:?
917:†
887:?
869:?
864:†
855:?
848:†
779:†
770:?
763:†
759:?
752:†
748:?
741:†
530:e
523:t
516:v
499:.
485::
230:†
209:†
196:†
183:†
95:N
85:K
80:J
75:T
70:P
65:C
60:D
55:S
50:O
45:Ꞓ
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