306:
264:
125:
294:, seems to indicate that titanopterans did not utilize stridulation (unlike modern orthopterans), but rather used flashes of light from wing displays and crepitation, moving their wings to produce sound. The authors argue that stridulation, crepitation, castanet signaling or light flash alone do not fully explains the diversity of structures observed in Titanoptera, and note that both sexes seem to have the fluted region on the forewing.
380:
1883:
1034:
102:
443:
was originally attributed to
Titanoptera, but as fossil does not show the venational structures of the order Titanoptera, and it is even possible to be misidentification of plant fossil, possibly
638:
Schubnel, Thomas; Legendre, Frédéric; Roques, Patrick; Garrouste, Romain; Cornette, Raphaël; Perreau, Michel; Perreau, Naïl; Desutter-Grandcolas, Laure; Nel, André (2021-07-08).
255:
periods. Titanopterans were very large in comparison with modern insects, some having wingspans of up to 36 centimetres (14 in) or even 40 centimetres (16 in).
1972:
869:"The first representative of the suborder Mesotitanina from the Paleozoic and notes on the system and evolution of the order Titanoptera (Insecta: Polyneoptera)"
937:
409:. But later study considered that the relationships between Titanoptera and Tcholmanvissiidae remain controversial. Three genera known from
279:, but were much larger, had proportionally weaker hindlegs that could not allow the animals to leap, and grasping forelegs and elongated
709:
930:
2041:
283:. Another distinctive feature was the presence of prominent fluted regions on the forewings, which may have been used in
305:
391:
There is controversy regarding the classification of
Titanoptera. Titanoptera was previously thought to be related to
124:
923:
2036:
2000:
852:
401:), but it is no longer supported. Béthoux (2007) considered that genera in Titanoptera should be included in
1567:
1912:
949:
2046:
2008:
768:
1818:
263:
1760:
1740:
406:
325:
280:
1833:
591:
228:
223:
557:
520:
495:
458:
340:
is larger in volume. All known hind wings of
Titanoptera, whatever their sizes, have quite reduced
218:
213:
1848:
439:
427:
1858:
1780:
1755:
896:
790:
437:
originally belonged to
Mesotitanidae, but it is later considered to belong to Tcholmanvissiidae.
119:
2013:
598:
548:
511:
503:
478:
421:
415:
1775:
574:
535:
527:
487:
433:
392:
1985:
1959:
1808:
888:
705:
679:
661:
1990:
1950:
582:
1977:
1750:
1601:
1287:
880:
847:
780:
747:
669:
651:
1828:
1745:
1735:
1712:
1279:
287:. The general shape and anatomy of the titanopterans suggests that they were predators.
1887:
1843:
1838:
1823:
1770:
1373:
674:
639:
302:
would be the oldest known insect with a wing structure specialized for communication.
2030:
1798:
1790:
1016:
1008:
835:
812:
785:
248:
59:
900:
794:
379:
1964:
1730:
1503:
1480:
1472:
1414:
1369:
1201:
1144:
973:
368:
284:
186:
101:
1894:
1944:
1785:
1717:
1672:
1618:
1519:
1449:
1300:
1238:
1114:
465:
360:
341:
34:
1935:
656:
1803:
1527:
1337:
1246:
1230:
1214:
1042:
884:
402:
352:
331:
276:
79:
44:
892:
769:"How Gerarus lost its head: stem-group Orthoptera and Paraneoptera revisited"
665:
1765:
1609:
1569:
1544:
1397:
1308:
1267:
1254:
1085:
1072:
1033:
967:
868:
566:
356:
336:
156:
136:
111:
84:
683:
1929:
1813:
1722:
1511:
1381:
1173:
979:
298:
is 50 Ma older than the previous oldest species of
Titanoptera, and thus
252:
241:
176:
74:
69:
54:
49:
39:
1882:
752:
731:
1152:
410:
397:
244:
89:
64:
319:
Some titanopterans may have been able to only glide, not fly, such as
1489:
1389:
1316:
1222:
1131:
1123:
961:
946:
364:
166:
146:
1906:
640:"Sound vs. light: wing-based communication in Carboniferous insects"
304:
262:
1577:
915:
444:
1910:
1697:
992:
919:
1587:
813:"Cladotypic taxonomy applied: titanopterans are orthopterans"
290:
An examination of a fossil of the oldest titanopteran genus,
853:
10.1665/1082-6467(2002)011[0223:NDOTP]2.0.CO;2
312:, modern orthopteran with hind wing area close to that of
323:. The hind wing area of it is almost the same as that of
730:
Park, T.-Y.S.; Kim, D.-Y.; Nam, G.-S.; Lee, M. (2022).
431:
344:, while most extant flying Orthoptera have large ones.
1530:(net-winged insects: lacewings, mantidflies, antlions)
704:(2nd ed.). Oxford University Press. p. 322.
836:"New data on Tcholmanvissiidae (Orthoptera; Permian)"
700:
Hoell, H.V.; Doyen, J.T. & Purcell, A.H. (1998).
1919:
1600:
1558:
1543:
1502:
1471:
1462:
1413:
1367:
1336:
1299:
1266:
1199:
1172:
1143:
1122:
1113:
1084:
1041:
1007:
351:, reliable records of titanopterans are known from
736:n. gen. n. sp. from southwestern Korean Peninsula"
767:Béthoux, Olivier; Briggs, Derek E. G. (2008).
405:, and divided from extinct orthopteran family
931:
367:as well, titanopterans possibly had a circum-
8:
702:Introduction to Insect Biology and Diversity
482:(Some question its affinity as titanopteran)
1907:
1705:
1694:
1555:
1468:
1441:
1364:
1360:
1207:
1196:
1192:
1119:
1110:
1106:
1065:
1004:
1000:
989:
938:
924:
916:
725:
723:
721:
100:
20:
851:
784:
751:
673:
655:
363:. Considering some possible records from
1659:Four most speciose orders are marked in
1249:(crickets, wetas, grasshoppers, locusts)
378:
613:
695:
693:
817:Arthropod Systematics & Phylogeny
469:(Originally considered as synonym of
7:
1522:(alderflies, dobsonflies, fishflies)
806:
804:
633:
631:
629:
627:
625:
623:
621:
619:
617:
275:Titanopterans are related to modern
473:but some propose to keep the genus)
14:
1881:
1032:
786:10.1111/j.1365-3113.2008.00419.x
123:
1998:Orthoptera Species File (old):
834:Béthoux, Olivier (2009-01-20).
811:Béthoux, Olivier (2007-01-01).
867:Gorochov, A. V. (2007-10-01).
840:Journal of Orthoptera Research
1:
1452:(sawflies, wasps, ants, bees)
1400:(cicadas, aphids, true bugs)
330:, one of the largest modern
2063:
1590:(gnats, mosquitoes, flies)
1483:(twisted-winged parasites)
1155:(dragonflies, damselflies)
657:10.1038/s42003-021-02281-0
310:Pseudophyllanax imperialis
1875:
1708:
1704:
1693:
1655:
1444:
1363:
1210:
1195:
1109:
1068:
1030:
1003:
999:
988:
956:
885:10.1134/S0031030107060056
210:
205:
120:Scientific classification
118:
108:
99:
23:
1241:(stick and leaf insects)
334:, and a poor flier, but
271:, showing its large size
16:Extinct order of insects
1668:are paraphyletic groups
1311:(cockroaches, termites)
1075:(silverfish, firebrats)
873:Paleontological Journal
740:Journal of Paleontology
240:is an extinct order of
1019:(jumping bristletails)
644:Communications Biology
388:
316:
272:
2042:Extinct insect orders
2009:Paleobiology Database
773:Systematic Entomology
385:Clatrotitan andersoni
382:
308:
266:
1621:(moths, butterflies)
734:Magnatitan jongheoni
732:"A new titanopteran
753:10.1017/jpa.2022.30
1888:Insects portal
1859:Triadophlebioptera
389:
321:Gigatitan vulgaris
317:
273:
269:Gigatitan vulgaris
267:Reconstruction of
109:Reconstruction of
30:Moscovian–Triassic
2024:
2023:
1986:Open Tree of Life
1913:Taxon identifiers
1904:
1903:
1871:
1870:
1867:
1866:
1809:Palaeodictyoptera
1794:
1726:
1689:
1688:
1685:
1684:
1651:
1650:
1647:
1646:
1643:
1642:
1639:
1638:
1635:
1634:
1631:
1630:
1622:
1613:
1591:
1581:
1573:
1551:
1539:
1538:
1531:
1523:
1515:
1493:
1484:
1453:
1409:
1408:
1401:
1393:
1385:
1332:
1331:
1328:
1327:
1320:
1312:
1291:
1288:Mantophasmatodea
1283:
1272:
1258:
1250:
1242:
1234:
1226:
1218:
1168:
1167:
1164:
1163:
1156:
1135:
1124:Ephemeropteroidea
1076:
1028:
1027:
1020:
235:
234:
2054:
2037:Triassic insects
2017:
2016:
2004:
2003:
1994:
1993:
1981:
1980:
1968:
1967:
1955:
1954:
1953:
1940:
1939:
1938:
1908:
1886:
1885:
1819:Permoplectoptera
1792:
1751:Diaphanopterodea
1724:
1706:
1695:
1620:
1611:
1602:Amphiesmenoptera
1589:
1583:
1579:
1571:
1561:
1556:
1550:
1547:
1529:
1521:
1513:
1491:
1482:
1469:
1465:
1451:
1442:
1399:
1391:
1384:(barklice, lice)
1383:
1375:
1365:
1361:
1318:
1310:
1289:
1281:
1280:Grylloblattodea
1270:
1256:
1248:
1240:
1232:
1224:
1216:
1208:
1204:
1197:
1193:
1154:
1133:
1120:
1111:
1107:
1074:
1066:
1036:
1018:
1005:
1001:
990:
940:
933:
926:
917:
905:
904:
864:
858:
857:
855:
831:
825:
824:
808:
799:
798:
788:
764:
758:
757:
755:
746:(5): 1111–1118.
727:
716:
715:
697:
688:
687:
677:
659:
635:
198:
128:
127:
104:
94:
31:
27:Temporal range:
21:
2062:
2061:
2057:
2056:
2055:
2053:
2052:
2051:
2027:
2026:
2025:
2020:
2012:
2007:
1999:
1997:
1989:
1984:
1976:
1971:
1963:
1958:
1949:
1948:
1943:
1934:
1933:
1928:
1915:
1905:
1900:
1880:
1863:
1829:Protelytroptera
1761:Eudiaphanoptera
1746:Coxoplectoptera
1741:Carbotriplurida
1736:Campylopteridae
1713:Aethiocarenodea
1700:
1681:
1627:
1596:
1572:(scorpionflies)
1559:
1548:
1546:
1535:
1498:
1463:
1458:
1436:
1434:
1432:
1430:
1428:
1426:
1424:
1422:
1420:
1418:
1416:
1405:
1355:
1353:
1351:
1349:
1347:
1345:
1343:
1341:
1339:
1324:
1295:
1269:
1262:
1257:(angel insects)
1200:
1187:
1185:
1183:
1181:
1179:
1177:
1175:
1160:
1139:
1101:
1099:
1097:
1095:
1093:
1091:
1089:
1087:
1080:
1060:
1058:
1056:
1054:
1052:
1050:
1048:
1046:
1044:
1037:
1024:
995:
984:
952:
944:
914:
909:
908:
866:
865:
861:
833:
832:
828:
810:
809:
802:
766:
765:
761:
729:
728:
719:
712:
699:
698:
691:
637:
636:
615:
610:
407:Tchomanvissidae
377:
326:Pseudophyllanax
261:
226:
221:
216:
196:
122:
95:
93:
92:
87:
82:
77:
72:
67:
62:
57:
52:
47:
42:
37:
29:
28:
25:
17:
12:
11:
5:
2060:
2058:
2050:
2049:
2044:
2039:
2029:
2028:
2022:
2021:
2019:
2018:
2005:
1995:
1982:
1969:
1956:
1941:
1925:
1923:
1917:
1916:
1911:
1902:
1901:
1899:
1898:
1891:
1876:
1873:
1872:
1869:
1868:
1865:
1864:
1862:
1861:
1856:
1851:
1846:
1844:Protozygoptera
1841:
1839:Protorthoptera
1836:
1834:Protephemerida
1831:
1826:
1824:Protanisoptera
1821:
1816:
1811:
1806:
1801:
1796:
1793:(griffinflies)
1791:Meganisoptera
1788:
1783:
1778:
1773:
1771:Glosselytrodea
1768:
1763:
1758:
1753:
1748:
1743:
1738:
1733:
1728:
1720:
1715:
1709:
1702:
1701:
1698:
1691:
1690:
1687:
1686:
1683:
1682:
1680:
1679:
1669:
1663:
1656:
1653:
1652:
1649:
1648:
1645:
1644:
1641:
1640:
1637:
1636:
1633:
1632:
1629:
1628:
1626:
1625:
1615:
1606:
1604:
1598:
1597:
1595:
1594:
1584:
1564:
1562:
1553:
1541:
1540:
1537:
1536:
1534:
1533:
1525:
1517:
1512:Raphidioptera
1508:
1506:
1500:
1499:
1497:
1496:
1486:
1477:
1475:
1466:
1464:Neuropteroidea
1460:
1459:
1457:
1456:
1445:
1439:
1411:
1410:
1407:
1406:
1404:
1403:
1395:
1387:
1378:
1376:
1358:
1334:
1333:
1330:
1329:
1326:
1325:
1323:
1322:
1314:
1305:
1303:
1297:
1296:
1294:
1293:
1285:
1282:(ice-crawlers)
1276:
1274:
1264:
1263:
1261:
1260:
1252:
1244:
1236:
1228:
1220:
1211:
1205:
1190:
1170:
1169:
1166:
1165:
1162:
1161:
1159:
1158:
1149:
1147:
1141:
1140:
1138:
1137:
1132:Ephemeroptera
1128:
1126:
1117:
1104:
1082:
1081:
1079:
1078:
1069:
1063:
1039:
1038:
1031:
1029:
1026:
1025:
1023:
1022:
1017:Archaeognatha
1013:
1011:
997:
996:
993:
986:
985:
983:
982:
976:
970:
964:
957:
954:
953:
945:
943:
942:
935:
928:
920:
913:
912:External links
910:
907:
906:
879:(6): 621–625.
859:
846:(2): 223–235.
826:
800:
779:(3): 529–547.
759:
717:
710:
689:
612:
611:
609:
606:
605:
604:
603:
602:
592:Theiatitanidae
588:
587:
586:
578:
570:
554:
553:
552:
544:
531:
517:
516:
515:
507:
499:
491:
483:
474:
376:
375:Classification
373:
371:distribution.
260:
257:
233:
232:
229:Theiatitanidae
224:Gigantitanidae
208:
207:
203:
202:
194:
190:
189:
184:
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164:
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6:
4:
3:
2:
2059:
2048:
2045:
2043:
2040:
2038:
2035:
2034:
2032:
2015:
2010:
2006:
2002:
1996:
1992:
1987:
1983:
1979:
1974:
1970:
1966:
1961:
1957:
1952:
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1931:
1927:
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1896:
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1884:
1878:
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1860:
1857:
1855:
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1845:
1842:
1840:
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1835:
1832:
1830:
1827:
1825:
1822:
1820:
1817:
1815:
1812:
1810:
1807:
1805:
1802:
1800:
1799:Megasecoptera
1797:
1795:
1789:
1787:
1784:
1782:
1779:
1777:
1774:
1772:
1769:
1767:
1764:
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1759:
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1749:
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1714:
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1710:
1707:
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1696:
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1678:
1676:
1670:
1667:
1664:
1662:
1658:
1657:
1654:
1624:
1623:
1616:
1614:
1612:(caddisflies)
1608:
1607:
1605:
1603:
1599:
1593:
1592:
1585:
1582:
1578:Siphonaptera
1575:
1574:
1566:
1565:
1563:
1557:
1554:
1552:
1549:(Mecopterida)
1542:
1532:
1526:
1524:
1518:
1516:
1510:
1509:
1507:
1505:
1501:
1495:
1494:
1487:
1485:
1481:Strepsiptera
1479:
1478:
1476:
1474:
1470:
1467:
1461:
1455:
1454:
1447:
1446:
1443:
1440:
1438:
1412:
1402:
1396:
1394:
1390:Thysanoptera
1388:
1386:
1380:
1379:
1377:
1372:
1371:
1366:
1362:
1359:
1357:
1335:
1321:
1315:
1313:
1307:
1306:
1304:
1302:
1298:
1292:
1286:
1284:
1278:
1277:
1275:
1273:
1265:
1259:
1253:
1251:
1245:
1243:
1237:
1235:
1233:(webspinners)
1229:
1227:
1221:
1219:
1213:
1212:
1209:
1206:
1203:
1198:
1194:
1191:
1189:
1171:
1157:
1151:
1150:
1148:
1146:
1142:
1136:
1130:
1129:
1127:
1125:
1121:
1118:
1116:
1112:
1108:
1105:
1103:
1083:
1077:
1071:
1070:
1067:
1064:
1062:
1040:
1035:
1021:
1015:
1014:
1012:
1010:
1006:
1002:
998:
991:
987:
981:
977:
975:
971:
969:
965:
963:
959:
958:
955:
951:
948:
941:
936:
934:
929:
927:
922:
921:
918:
911:
902:
898:
894:
890:
886:
882:
878:
874:
870:
863:
860:
854:
849:
845:
841:
837:
830:
827:
822:
818:
814:
807:
805:
801:
796:
792:
787:
782:
778:
774:
770:
763:
760:
754:
749:
745:
741:
737:
735:
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724:
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718:
713:
711:0-19-510033-6
707:
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584:
579:
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569:
568:
563:
562:
561:
559:
558:Gigatitanidae
555:
551:
550:
545:
542:
538:
537:
532:
530:
529:
524:
523:
522:
521:Paratitanidae
518:
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492:
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484:
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459:Mesotitanidae
456:
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258:
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249:Carboniferous
246:
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219:Paratitanidae
215:
214:Mesotitanidae
209:
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91:
86:
81:
76:
71:
66:
61:
56:
51:
46:
41:
36:
22:
19:
2047:Orthopterida
1920:
1893:
1879:
1853:
1849:Syntonoptera
1731:Caloneurodea
1723:Blattoptera
1674:
1665:
1660:
1619:Lepidoptera
1617:
1610:Trichoptera
1586:
1568:
1520:Megaloptera
1514:(snakeflies)
1504:Neuropterida
1488:
1473:Coleopterida
1450:Hymenoptera
1448:
1370:Paraneoptera
1368:
1290:(gladiators)
1239:Phasmatodea
1217:(stoneflies)
1202:Polyneoptera
1145:Odonatoptera
1009:Monocondylia
974:Pancrustacea
972:(unranked):
876:
872:
862:
843:
839:
829:
820:
816:
776:
772:
762:
743:
739:
733:
701:
647:
643:
597:
581:
573:
565:
560:
547:
540:
534:
526:
510:
502:
494:
486:
477:
470:
464:
451:
449:
440:Steinhardtia
438:
432:
428:Monstrotitan
426:
420:
414:
396:
390:
384:
383:Forewing of
348:
346:
335:
324:
320:
318:
313:
309:
299:
295:
291:
289:
285:stridulation
277:grasshoppers
274:
268:
237:
236:
211:
199:
187:Polyneoptera
173:Infraclass:
110:
18:
1951:Titanoptera
1945:Wikispecies
1921:Titanoptera
1895:Wikispecies
1854:Titanoptera
1786:Lapeyriidae
1781:Hypoperlida
1756:Eoblattodea
1725:(roachoids)
1718:Archodonata
1560:Antliophora
1528:Neuroptera
1490:Coleoptera
1301:Dictyoptera
1271:(Xenonomia)
1247:Orthoptera
1231:Embioptera
1223:Dermaptera
1215:Plecoptera
1115:Palaeoptera
978:Subphylum:
466:Clatrotitan
452:Titanoptera
395:(including
361:South Korea
347:Other than
259:Description
238:Titanoptera
200:Titanoptera
24:Titanoptera
2031:Categories
1804:Miomoptera
1570:Mecoptera
1398:Hemiptera
1319:(mantises)
1309:Blattodea
1255:Zoraptera
1134:(mayflies)
1073:Zygentoma
968:Arthropoda
823:: 135–156.
650:(1): 794.
608:References
599:Theiatitan
549:Magnatitan
541:Microtitan
512:Ultratitan
504:Prototitan
479:Deinotitan
422:Deinotitan
416:Permotitan
403:Orthoptera
353:Kyrgyzstan
349:Theiatitan
332:Orthoptera
328:imperialis
300:Theiatitan
292:Theiatitan
247:from late
157:Arthropoda
1776:Heraridea
1766:Geroptera
1671:Based on
1545:Panorpida
1492:(beetles)
1382:Psocodea
1317:Mantodea
1268:Notoptera
1225:(earwigs)
960:Kingdom:
893:1555-6174
666:2399-3642
575:Nanotitan
567:Gigatitan
536:Minititan
528:Paratitan
488:Mesotitan
471:Mesotitan
434:Jubilaeus
393:Geraridae
357:Australia
337:Gigatitan
314:Gigatitan
296:Theiatian
281:mandibles
242:neopteran
206:Families
143:Kingdom:
137:Eukaryota
112:Gigatitan
1936:Q4458210
1930:Wikidata
1814:Paoliida
1588:Diptera
1392:(thrips)
1153:Odonata
980:Hexapoda
966:Phylum:
962:Animalia
901:85364942
795:84577837
684:34239029
590:Family †
556:Family †
519:Family †
457:Family †
253:Triassic
183:Cohort:
177:Neoptera
153:Phylum:
147:Animalia
133:Domain:
2001:1143980
1991:5320676
1699:Extinct
1673:Sasaki
1580:(fleas)
1374: *
675:8266802
596:Genus †
583:Ootitan
580:Genus †
572:Genus †
564:Genus †
546:Genus †
533:Genus †
525:Genus †
509:Genus †
501:Genus †
493:Genus †
485:Genus †
476:Genus †
463:Genus †
411:Permian
398:Gerarus
245:insects
193:Order:
167:Insecta
163:Class:
2014:224437
1677:(2013)
1675:et al.
1666:Italic
994:Extant
950:orders
947:Insect
899:
891:
793:
708:
682:
672:
664:
450:Order
369:Tethys
365:Russia
342:vannus
1978:11529
1973:IRMNG
1965:8NKHF
897:S2CID
791:S2CID
1661:bold
889:ISSN
706:ISBN
680:PMID
662:ISSN
445:fern
359:and
35:PreꞒ
1960:CoL
881:doi
848:doi
781:doi
748:doi
670:PMC
652:doi
251:to
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2011::
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1975::
1962::
1947::
1932::
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877:41
875:.
871:.
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744:96
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720:^
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616:^
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1188:a
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1174:N
1102:a
1100:t
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1094:y
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1061:a
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227:†
222:†
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