471:
121:
139:
517:, and is curled and notably shriveled. However, Cooper refrained from describing this individual in 1964 due to his inability to reliably determine its anatomical details. New imaging techniques developed since then allowed for later researchers to describe the smaller specimen in detail. It was named in 2024 as the new genus
461:
The legs are presumably telescopically retracted and each carry four unequally long claws arranged asymmetrically with respect to the middle of the leg. Other features, such as the structure of the stiletto apparatus or the mucous musculature, can not be recognized; Also the position of the sex
446:
The third and fourth regions bear the second and third pairs of legs. Here there are also shorter transverse furrows on the back, which run out laterally to the legs. They are shifted forward against the center line of the
535:
with the tardigrades is certain. Moreover, the species can even be assigned to one of the three modern classes - the absence of head structures such as cirri and clavae and back armoring suggests a placement within the
431:
The first or prostomial region forms the head on which there are no structures, such as filiform cirri or clavae, which are used in some modern modes of sensory perception. Eyes can not be identified. The
420:
is enclosed in honey-colored amber. The animal itself is 0.3 millimeters long and 0.08 millimeters wide. As is the case with modern tardigrades, the cylindrical body is flattened on the ventral side.
427:
is one of the "naked" tardigrades. A total of four furrows extend transversely to the longitudinal axis of the body around the whole body, which divide these superficially into five regions:
400:
refers to Cooper's student
William M. Legg who collected the amber in the summer of 1940 in which the holotype was found. He died in 1953 before finishing his thesis at
1698:
1564:
1417:
462:
opening (gonopore) with respect to the anus, which could have given further indications of the class belonging to the species, can not be ascertained.
1685:
1551:
1404:
1711:
1577:
1430:
544:
was originally placed in a separate family
Beornidae. However, later studies based on more detailed imageing reinterpreted it as belonging to the
1793:
1300:
1150:
1124:
1788:
1750:
1616:
1482:
1238:
374:
The species was first described in 1964 by
Kenneth W. Cooper. The type specimen is now in the Museum of Comparative Zoology at
1778:
371:'s history, and its largely modern appearance suggests that tardigrades must have diversified considerably before this time.
1716:
1582:
1435:
1076:
490:
The origin of the present amber specimen is unknown. It was part of secondary sediment deposits on the lake shore of
423:
The cuticle is smooth, and slightly thickened on the backside, but does not harden to platelets (sclerites), making
1783:
1798:
470:
1232:
Mapalo, Marc A.; Robin, Ninon; Boudinot, Brendon E.; Ortega-Hernández, Javier; Barden, Phillip (2021-10-13).
138:
1803:
443:. A short transverse furrow is slightly displaced towards the back, opposite the center line of the segment.
1318:
1179:
190:
1737:
1603:
1469:
906:
491:
244:
1755:
1621:
1487:
1391:
1365:
663:
401:
325:
221:
889:
642:
627:
120:
1448:
947:
923:
851:
773:
568:
495:
375:
133:
1742:
1608:
1474:
1422:
1142:
Animal biodiversity: An outline of higher-level classification and survey of taxonomic richness
1724:
1672:
1590:
1569:
1538:
1456:
1378:
1296:
1273:
1255:
1214:
1196:
1146:
1120:
1114:
1095:
793:
348:
1729:
1703:
1595:
1461:
1356:
1140:
1263:
1247:
1204:
1188:
1085:
475:
393:
330:
302:
287:
41:
1663:
567:
sequences. They recovered these taxa as closely related members of the tardigrade family
1209:
1174:
1268:
1233:
1772:
749:
560:
78:
1067:
1677:
865:
701:
537:
396:, which can occur both in the shape of a man and that of a bear. The specific name
180:
1543:
1383:
1370:
1648:
1529:
1514:
1657:
1523:
1443:
1350:
545:
316:
200:
53:
1192:
1175:"Cretaceous amber inclusions illuminate the evolutionary origin of tardigrades"
725:
677:
388:
356:
320:
276:
170:
98:
63:
1341:
1259:
1200:
1099:
436:
region is inconspicuous and does not have warren-like projections (papillae).
597:
572:
150:
103:
47:
1277:
1251:
1218:
17:
1173:
Mapalo, Marc A.; Wolfe, Joanna M.; Ortega-Hernández, Javier (2024-08-06).
1642:
1508:
1335:
1090:
987:
519:
499:
417:
360:
335:
294:
93:
88:
73:
68:
58:
1116:
Thorp and Covich's
Freshwater Invertebrates: Ecology and General Biology
458:
The length ratio of the segments 2 to 5 is indicated as 1: 1.3: 1.3: 2.
1556:
1409:
451:
344:
108:
83:
509:
Another tardigrade specimen was identified in the same amber piece as
311:. It was originally classified as the only member of its family, the
298:
160:
1690:
1312:
469:
433:
383:
368:
290:
1396:
564:
440:
1316:
343:. The only other confidently known fossil tardigrade is the
1113:
James H. Thorp; D. Christopher Rogers (6 September 2014).
450:
In the fifth region there is a transverse furrow on the
1239:
Proceedings of the Royal
Society B: Biological Sciences
367:
is an example of the early existence of tardigrades in
319:. It is one of three fossil tardigrades known from the
575:
below, with extinct species designated with a dagger (
454:, which runs out slightly laterally on the back line.
404:. Cooper, Legg's friend and scientific mentor, named
983:
963:
789:
576:
1632:
1498:
1325:
128:Holotype specimen under various imaging techniques
279:and the first known fossil tardigrade, discovered
315:, but was later reclassified as belonging to the
1295:. Cambridge University Press. pp. 96–97.
1291:Grimaldi, David A.; Engel, Michael S. (2005).
559:, Mapalo, Wolfe & Ortega-Hernández (2024)
439:The second region contains the first pair of
8:
563:a combination of morphological features and
506:may have been a swamp-like wetland biotope.
502:. Observations suggest that the habitat of
1313:
1168:
1166:
1164:
1162:
119:
31:
1267:
1208:
1089:
355:In addition to some other finds from the
1061:
1059:
1057:
1053:
571:. These results are displayed in the
382:was chosen by Cooper in reference to
7:
1145:. Magnolia Press. 2011. p. 96.
939:
881:
857:
765:
741:
717:
693:
669:
620:
613:
589:
582:
498:outlet in the Canadian province of
551:To determine the relationships of
25:
1234:"A tardigrade in Dominican amber"
1066:Cooper, Kenneth W. (1964-01-01).
137:
1077:Psyche: A Journal of Entomology
1072:Cooper, from Cretaceous amber"
1068:"The first fossil tardigrade:
1:
1794:Fossil taxa described in 1964
352:from the Dominican Republic.
339:from the same amber piece as
280:
384:a character of the same name
27:Extinct genus of tardigrades
1789:Monotypic protostome genera
286:and described in 1964 from
1820:
1193:10.1038/s42003-024-06643-2
513:. It is much smaller than
1119:. Elsevier. p. 351.
981:
961:
944:
937:
920:
903:
886:
879:
866:Ramazzottius oberhaeuseri
862:
855:
787:
770:
763:
746:
739:
722:
715:
698:
691:
674:
667:
640:
625:
618:
611:
594:
587:
561:phylogenetically analyzed
386:from the children's book
250:
243:
134:Scientific classification
132:
127:
118:
34:
1293:Evolution of the Insects
890:Microhypsibius truncatus
702:Isohypsibius prosostomus
301:. The genus contains a
275:is an extinct genus of
1779:Parachela (tardigrade)
1252:10.1098/rspb.2021.1760
1180:Communications Biology
750:Doryphoribius macrodon
487:
1738:Paleobiology Database
1604:Paleobiology Database
1470:Paleobiology Database
924:Calohypsibius ornatus
907:Acutuncus antarcticus
774:Hexapodibius micronyx
540:. Within this class,
473:
402:Princeton University
326:Milnesium swolenskyi
948:Hypsibius dujardini
678:Ramajendas frigidus
531:The affiliation of
494:, not far from the
378:. The generic name
323:, the others being
1091:10.1155/1964/48418
726:Halobiotus crispae
496:Saskatchewan River
488:
376:Harvard University
329:from the Turonian
1784:Tardigrade genera
1766:
1765:
1725:Open Tree of Life
1591:Open Tree of Life
1457:Open Tree of Life
1319:Taxon identifiers
1302:978-0-521-82149-0
1152:978-1-86977-849-1
1126:978-0-12-385027-0
1044:
1043:
1035:
1034:
1026:
1025:
1017:
1016:
1008:
1007:
999:
998:
841:
840:
832:
831:
823:
822:
814:
813:
805:
804:
794:Paradoryphoribius
653:
652:
349:Paradoryphoribius
268:
267:
224:
16:(Redirected from
1811:
1799:Campanian genera
1759:
1758:
1746:
1745:
1733:
1732:
1720:
1719:
1707:
1706:
1694:
1693:
1681:
1680:
1668:
1667:
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1653:
1652:
1651:
1625:
1624:
1612:
1611:
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1560:
1559:
1547:
1546:
1534:
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1519:
1518:
1517:
1491:
1490:
1478:
1477:
1465:
1464:
1452:
1451:
1439:
1438:
1426:
1425:
1413:
1412:
1400:
1399:
1387:
1386:
1374:
1373:
1361:
1360:
1359:
1346:
1345:
1344:
1314:
1307:
1306:
1288:
1282:
1281:
1271:
1229:
1223:
1222:
1212:
1170:
1157:
1156:
1137:
1131:
1130:
1110:
1104:
1103:
1093:
1063:
985:
965:
940:
882:
858:
791:
766:
742:
718:
694:
670:
621:
614:
590:
583:
578:
476:life restoration
394:J. R. R. Tolkien
331:New Jersey amber
285:
282:
259:
255:
219:
212:
142:
141:
123:
113:
50:
40:Temporal range:
32:
21:
1819:
1818:
1814:
1813:
1812:
1810:
1809:
1808:
1769:
1768:
1767:
1762:
1754:
1749:
1741:
1736:
1728:
1723:
1715:
1710:
1702:
1697:
1689:
1684:
1676:
1671:
1662:
1661:
1656:
1647:
1646:
1641:
1628:
1620:
1615:
1607:
1602:
1594:
1589:
1581:
1576:
1568:
1563:
1555:
1550:
1542:
1537:
1528:
1527:
1522:
1513:
1512:
1507:
1494:
1486:
1481:
1473:
1468:
1460:
1455:
1447:
1442:
1434:
1429:
1421:
1416:
1408:
1403:
1395:
1390:
1382:
1377:
1369:
1364:
1355:
1354:
1349:
1340:
1339:
1334:
1321:
1311:
1310:
1303:
1290:
1289:
1285:
1231:
1230:
1226:
1172:
1171:
1160:
1153:
1139:
1138:
1134:
1127:
1112:
1111:
1107:
1065:
1064:
1055:
1050:
1045:
1036:
1027:
1018:
1009:
1000:
842:
833:
824:
815:
806:
664:Isohypsibioidea
654:
529:
468:
414:
412:Characteristics
288:Late Cretaceous
283:
264:
261:
253:
252:
239:
218:
210:
136:
114:
112:
111:
106:
101:
96:
91:
86:
81:
76:
71:
66:
61:
56:
46:
45:
42:Late Cretaceous
38:
28:
23:
22:
15:
12:
11:
5:
1817:
1815:
1807:
1806:
1804:Canadian amber
1801:
1796:
1791:
1786:
1781:
1771:
1770:
1764:
1763:
1761:
1760:
1747:
1734:
1721:
1708:
1695:
1682:
1669:
1654:
1638:
1636:
1630:
1629:
1627:
1626:
1613:
1600:
1587:
1574:
1561:
1548:
1535:
1520:
1504:
1502:
1496:
1495:
1493:
1492:
1479:
1466:
1453:
1440:
1427:
1414:
1401:
1388:
1375:
1362:
1347:
1331:
1329:
1323:
1322:
1317:
1309:
1308:
1301:
1283:
1224:
1158:
1151:
1132:
1125:
1105:
1052:
1051:
1049:
1046:
1042:
1041:
1038:
1037:
1033:
1032:
1029:
1028:
1024:
1023:
1020:
1019:
1015:
1014:
1011:
1010:
1006:
1005:
1002:
1001:
997:
996:
993:
992:
980:
977:
976:
973:
972:
960:
957:
956:
953:
952:
943:
938:
936:
933:
932:
929:
928:
919:
916:
915:
912:
911:
902:
899:
898:
895:
894:
885:
880:
878:
875:
874:
871:
870:
861:
856:
854:
848:
847:
844:
843:
839:
838:
835:
834:
830:
829:
826:
825:
821:
820:
817:
816:
812:
811:
808:
807:
803:
802:
799:
798:
786:
783:
782:
779:
778:
769:
764:
762:
759:
758:
755:
754:
745:
740:
738:
735:
734:
731:
730:
721:
716:
714:
711:
710:
707:
706:
697:
692:
690:
687:
686:
683:
682:
673:
668:
666:
660:
659:
656:
655:
651:
650:
647:
646:
643:Macrobiotoidea
639:
636:
635:
632:
631:
628:Eohypsibioidea
624:
619:
617:
612:
610:
607:
606:
603:
602:
593:
588:
586:
581:
528:
527:Classification
525:
478:of the coeval
467:
464:
456:
455:
448:
444:
437:
413:
410:
303:single species
266:
265:
262:
248:
247:
241:
240:
232:
230:
226:
225:
208:
204:
203:
198:
194:
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188:
184:
183:
178:
174:
173:
168:
164:
163:
158:
154:
153:
148:
144:
143:
130:
129:
125:
124:
116:
115:
107:
102:
97:
92:
87:
82:
77:
72:
67:
62:
57:
52:
51:
39:
26:
24:
14:
13:
10:
9:
6:
4:
3:
2:
1816:
1805:
1802:
1800:
1797:
1795:
1792:
1790:
1787:
1785:
1782:
1780:
1777:
1776:
1774:
1757:
1752:
1748:
1744:
1739:
1735:
1731:
1726:
1722:
1718:
1713:
1709:
1705:
1700:
1696:
1692:
1687:
1683:
1679:
1674:
1670:
1665:
1659:
1655:
1650:
1644:
1640:
1639:
1637:
1635:
1631:
1623:
1618:
1614:
1610:
1605:
1601:
1597:
1592:
1588:
1584:
1579:
1575:
1571:
1566:
1562:
1558:
1553:
1549:
1545:
1540:
1536:
1531:
1525:
1521:
1516:
1510:
1506:
1505:
1503:
1501:
1497:
1489:
1484:
1480:
1476:
1471:
1467:
1463:
1458:
1454:
1450:
1445:
1441:
1437:
1432:
1428:
1424:
1419:
1415:
1411:
1406:
1402:
1398:
1393:
1389:
1385:
1380:
1376:
1372:
1367:
1363:
1358:
1352:
1348:
1343:
1337:
1333:
1332:
1330:
1328:
1324:
1320:
1315:
1304:
1298:
1294:
1287:
1284:
1279:
1275:
1270:
1265:
1261:
1257:
1253:
1249:
1245:
1241:
1240:
1235:
1228:
1225:
1220:
1216:
1211:
1206:
1202:
1198:
1194:
1190:
1186:
1182:
1181:
1176:
1169:
1167:
1165:
1163:
1159:
1154:
1148:
1144:
1143:
1136:
1133:
1128:
1122:
1118:
1117:
1109:
1106:
1101:
1097:
1092:
1087:
1083:
1079:
1078:
1073:
1071:
1062:
1060:
1058:
1054:
1047:
1040:
1039:
1031:
1030:
1022:
1021:
1013:
1012:
1004:
1003:
995:
994:
991:
990:
989:
979:
978:
975:
974:
971:
970:
969:
959:
958:
955:
954:
951:
950:
949:
942:
941:
935:
934:
931:
930:
927:
926:
925:
918:
917:
914:
913:
910:
909:
908:
901:
900:
897:
896:
893:
892:
891:
884:
883:
877:
876:
873:
872:
869:
868:
867:
860:
859:
853:
850:
849:
846:
845:
837:
836:
828:
827:
819:
818:
810:
809:
801:
800:
797:
796:
795:
785:
784:
781:
780:
777:
776:
775:
768:
767:
761:
760:
757:
756:
753:
752:
751:
744:
743:
737:
736:
733:
732:
729:
728:
727:
720:
719:
713:
712:
709:
708:
705:
704:
703:
696:
695:
689:
688:
685:
684:
681:
680:
679:
672:
671:
665:
662:
661:
658:
657:
649:
648:
645:
644:
638:
637:
634:
633:
630:
629:
623:
622:
616:
615:
609:
608:
605:
604:
601:
600:
599:
592:
591:
585:
584:
580:
574:
570:
566:
562:
558:
554:
549:
547:
543:
539:
534:
526:
524:
522:
521:
516:
512:
507:
505:
501:
497:
493:
485:
481:
477:
474:Hypothetical
472:
465:
463:
459:
453:
449:
445:
442:
438:
435:
430:
429:
428:
426:
421:
419:
411:
409:
407:
403:
399:
395:
391:
390:
385:
381:
377:
372:
370:
366:
362:
358:
353:
351:
350:
346:
342:
338:
337:
332:
328:
327:
322:
318:
314:
310:
309:
304:
300:
296:
292:
289:
278:
274:
273:
260:
258:
249:
246:
245:Binomial name
242:
238:
237:
236:B. leggi
231:
228:
227:
223:
217:
216:
209:
206:
205:
202:
199:
196:
195:
192:
189:
186:
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175:
172:
169:
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162:
159:
156:
155:
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149:
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145:
140:
135:
131:
126:
122:
117:
110:
105:
100:
95:
90:
85:
80:
75:
70:
65:
60:
55:
49:
43:
37:
33:
30:
19:
1633:
1499:
1326:
1292:
1286:
1243:
1237:
1227:
1184:
1178:
1141:
1135:
1115:
1108:
1084:(2): 41–48.
1081:
1075:
1069:
986:
982:
967:
966:
962:
946:
945:
922:
921:
905:
904:
888:
887:
864:
863:
852:Hypsibioidea
792:
788:
772:
771:
748:
747:
724:
723:
700:
699:
676:
675:
641:
626:
596:
595:
569:Hypsibioidea
556:
552:
550:
541:
538:Eutardigrada
532:
530:
518:
514:
510:
508:
503:
489:
483:
479:
460:
457:
424:
422:
415:
405:
397:
387:
379:
373:
364:
354:
347:
340:
334:
324:
312:
307:
306:
271:
270:
269:
263:Cooper, 1964
256:
251:
235:
234:
214:
213:
181:Eutardigrada
35:
29:
1658:Wikispecies
1524:Wikispecies
1444:NatureServe
1357:Beorn leggi
1351:Wikispecies
1327:Beorn leggi
1070:Beorn leggi
546:Hypsibiidae
542:Beorn leggi
408:after him.
406:Beorn leggi
317:Hypsibiidae
284: 1940
257:Beorn leggi
201:Hypsibiidae
18:Beorn leggi
1773:Categories
1187:(1): 953.
1048:References
492:Cedar Lake
482:(top) and
389:The Hobbit
357:Cretaceous
321:Cretaceous
277:tardigrade
171:Tardigrada
1664:Beornidae
1649:Q20020090
1634:Beornidae
1515:Q18520266
1449:2.1202122
1371:Beornidae
1260:0962-8452
1201:2399-3642
1100:0033-2615
598:Milnesium
573:cladogram
313:Beornidae
229:Species:
191:Parachela
157:Kingdom:
151:Eukaryota
48:Campanian
1643:Wikidata
1509:Wikidata
1423:11656441
1336:Wikidata
1278:34610770
1246:(1960).
1219:39107512
1210:11303527
988:Aerobius
553:Aerobius
520:Aerobius
500:Manitoba
486:(bottom)
484:Aerobius
447:segment.
418:holotype
363:period,
361:Cambrian
359:and the
336:Aerobius
308:B. leggi
295:Manitoba
220:Cooper,
197:Family:
167:Phylum:
161:Animalia
147:Domain:
1730:5004771
1596:5004772
1570:1035029
1557:2254046
1462:5004773
1410:2254047
1397:1048082
1342:Q818582
1269:8493197
466:Habitat
452:abdomen
345:Miocene
254:†
233:†
207:Genus:
187:Order:
177:Class:
1756:955448
1743:347395
1717:710176
1704:104066
1622:955449
1609:347397
1583:710257
1488:956693
1475:347398
1436:710543
1299:
1276:
1266:
1258:
1217:
1207:
1199:
1149:
1123:
1098:
299:Canada
1751:WoRMS
1699:IRMNG
1678:622FB
1617:WoRMS
1565:IRMNG
1530:Beorn
1500:Beorn
1483:WoRMS
1418:IRMNG
968:Beorn
557:Beorn
533:Beorn
515:Beorn
511:Beorn
504:Beorn
480:Beorn
434:mouth
425:Beorn
398:leggi
380:Beorn
369:earth
365:Beorn
341:Beorn
293:from
291:amber
272:Beorn
215:Beorn
36:Beorn
1712:ITIS
1691:3127
1686:GBIF
1578:ITIS
1552:GBIF
1544:39KT
1431:ITIS
1405:GBIF
1384:LJ7D
1297:ISBN
1274:PMID
1256:ISSN
1215:PMID
1197:ISSN
1147:ISBN
1121:ISBN
1096:ISSN
565:rRNA
555:and
441:legs
416:The
333:and
222:1964
54:PreꞒ
1673:CoL
1539:CoL
1392:EoL
1379:CoL
1366:ADW
1264:PMC
1248:doi
1244:288
1205:PMC
1189:doi
1086:doi
579:):
392:by
1775::
1753::
1740::
1727::
1714::
1701::
1688::
1675::
1660::
1645::
1619::
1606::
1593::
1580::
1567::
1554::
1541::
1526::
1511::
1485::
1472::
1459::
1446::
1433::
1420::
1407::
1394::
1381::
1368::
1353::
1338::
1272:.
1262:.
1254:.
1242:.
1236:.
1213:.
1203:.
1195:.
1183:.
1177:.
1161:^
1094:.
1082:71
1080:.
1074:.
1056:^
548:.
523:.
305:,
297:,
281:c.
104:Pg
44:,
1305:.
1280:.
1250::
1221:.
1191::
1185:7
1155:.
1129:.
1102:.
1088::
984:†
964:†
790:†
577:†
211:†
109:N
99:K
94:J
89:T
84:P
79:C
74:D
69:S
64:O
59:Ꞓ
20:)
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