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Rhyniophyte

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In 2013, Hao and Xue returned to the earlier definition. Their class Rhyniopsida (rhyniopsids) is defined by the presence of sporangia that terminate isotomous branching systems (i.e. the plants have branching patterns in which the branches are equally sized, rather than one branch dominating, like
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in 1968 under the name Rhyniophytina. The original definition was: "plants with naked (lacking emergences), dichotomizing axes bearing sporangia that are terminal, usually fusiform and may dehisce longitudinally; they are diminutive plants and, in so far as is known, have a small terete xylem strand
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since some of its members are now known to lack vascular tissue. The definition that seems to be used most often now is that of D. Edwards and D.S. Edwards: "plants with smooth axes, lacking well-defined spines or leaves, showing a variety of branching patterns that may be isotomous, anisotomous,
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pseudomonopodial or adventitious. Elongate to globose sporangia were terminal on main axes or on lateral systems showing limited branching. It seems probable that the xylem, comprising a solid strand of tracheids, was centrarch." However, Edwards and Edwards also decided to include
436: 276:, since when there have been many redefinitions, including by Banks himself. "As a result, the Rhyniophytina have slowly dissolved into a heterogeneous collection of plants ... the group contains only one species on which all authors agree: the type species 280:". When defined very broadly, the group consists of plants with dichotomously branched, naked aerial axes ("stems") with terminal spore-bearing structures (sporangia). The rhyniophytes are considered to be stem group tracheophytes (vascular plants). 358:
use Rhyniophyta as a formal taxon, but with a loose definition: plants "characterized by dichotomously branched, naked aerial axes with terminal sporangia". They thus include under "other rhyniophytes" plants apparently without vascular tissue.
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for the polysporangiophytes in which the Rhyniaceae are shown as the sister group of all other tracheophytes (vascular plants). Some other former "rhyniophytes", such as
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Evolution and Environment: A Symposium Presented on the Occasion of the 100th Anniversary of the Foundation of Peabody Museum of Natural History at Yale University
339:, plants which "look like rhyniophytes, but cannot be assigned unequivocally to that group because of inadequate anatomical preservation", but exclude plants like 1546: 354:
In 1966, slightly before Banks created the subdivision, the group was treated as a division under the name Rhyniophyta. Taylor et al. in their book
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The Early Devonian Posongchong Flora of Yunnan – A Contribution to an Understanding of the Evolution and Early Diversification of Vascular Plants
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There is no agreement on the formal classification to be used for the rhyniophytes. The following are some of the names which may be used:
382:, had radially symmetrical sporangia that were longer than wide and possessed vascular tissue with S-type tracheids. Cooksonioids, such as 370:
the trunk of a tree). The shape and symmetry of the sporangia was then used to divide up the group. Rhynialeans (order Rhyniales), such as
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Kidston, R. & Lang, W.H. (1920), "On Old Red Sandstone plants showing structure, from the Rhynie Chert Bed, Aberdeenshire. Part III.
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https://web.archive.org/web/20070423195457/http://www2.ncsu.edu/unity/lockers/project/dendrology/index/plantae/vascular/vascular.html
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and renalioids. Genera included by Hao and Xue are listed below, with assignments to their three subgroups where these are given.
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have been tentatively classified as tracheophytes in at least one recent cladistic analysis of Early Devonian land plants.
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Edwards, D.; Edwards, D.S. (1986), "A reconsideration of the Rhyniophytina Banks", in Spicer, R.A.; Thomas, B.A. (eds.),
1388:"On Old Red Sandstone plants showing structure, from the Rhynie Chert Bed, Aberdeenshire. Part II. Additional notes on 394:, had radially symmetrical or trumpet-shaped sporangia, without clear evidence of vascular tissue. Renalioids, such as 481:, are placed outside the tracheophyte clade, as they did not possess true vascular tissue (in particular did not have 1634: 1367:
Němejc, F. (1950), "The natural systematic of plants in the light of the present palaeontological documents",
17: 135: 992:), may also be a rhyniophyte. Others have placed some of these genera in different groups. For example, 738: 737:
The taxon and informal terms corresponding to it have been used in different ways. Hao and Xue in 2013
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Niklas, K.J. (1979), "An Assessment of Chemical Features for the Classification of Plant Fossils",
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https://web.archive.org/web/20071028153446/http://www.palaeos.com/Plants/Rhyniophytes/index.html
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Cronquist, A.; Takhtajan, A. & Zimmermann, W. (1966), "On the higher Taxa of Embryobionta",
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Gonez, P. & Gerrienne, P. (2010a), "A New Definition and a Lectotypification of the Genus
1593: 1354:Учебник палеонтологии (Uchebnik paleontologii; Textbook of Palaeontology, 3rd edition only) 8: 1075: 323:. Informally, they are often called rhyniophytes or, as mentioned below, rhyniophytoids. 303: 273: 366:
was suggested, to distinguish such plants from 'true' tracheophytes or eutracheophytes.
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Crane, P.R.; Herendeen, P. & Friis, E.M. (2004), "Fossils and plant phylogeny",
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Banks, H.P. (1968), "The early history of land plants", in Drake, E.T. (ed.),
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Partial cladogram by Crane et al. including the more certain rhyniophytes:
347: 299: 272:. The first definition of the group, under the name Rhyniophytina, was by 1062: 1056: 987: 915: 892: 789: 759: 539: 341: 251: 47: 114: 1490: 1265: 1158: 751: 407: 316: 312: 92: 57: 1106:
The Origin and Early Diversification of Land Plants: A Cladistic Study
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sites with similar flora. Used in this way, these terms refer to a
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their Rhyniopsida quite broadly, dividing it into rhynialeans,
600: 241: 1047: 1023: 147: 1243: 1060:, are also common; there are also more complex plants, like 1046:
assemblage of more or less related early land plants, not a
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with a central protoxylem." With this definition, they are
1212:, New Haven, Conn.: Yale University Press, pp. 73–107 1182:
Paleobotany, The Biology and Evolution of Fossil Plants
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Sborník Národního Muzea V Praze: Řada B, Přírodni Vědy
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Andrews, H.N.; Kasper, A. & Mencher, E. (1968), "
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Taylor, T.N.; Taylor, E.L. & Krings, M. (2009).
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Systematic and taxonomic approaches in palaeobotany
1184:(2nd ed.). Amsterdam; Boston: Academic Press. 1108:, Washington, D.C.: Smithsonian Institution Press, 260:used for this group, some treating it as the class 1421: 1271: 1179: 1626: 1239: 1237: 975:It has been suggested that the poorly preserved 1277: 239:that are considered to be similar to the genus 1509:Transactions of the Royal Society of Edinburgh 1404:Transactions of the Royal Society of Edinburgh 1225: 1221: 1219: 1099: 1097: 1095: 1093: 1091: 1234: 1137:, a New Devonian Plant from Northern Maine", 1128: 1126: 1124: 1379: 1203: 1201: 1104:Kenrick, Paul & Crane, Peter R. (1997), 1103: 1502: 1496: 1385: 1315:Hao, Shougang & Xue, Jinzhuang (2013), 1230:, Oxford: Clarendon Press, pp. 199–218 1216: 1088: 351:which definitely do not possess tracheids. 1462: 1310: 1308: 1306: 1304: 1121: 1022:" is sometimes used for the assemblage of 406:had bilaterally symmetrical sporangia and 113: 1446: 1392:, Kidston and Lang; with descriptions of 1198: 1066:, which has a very early form of leaves. 1351: 1175: 1173: 1171: 1169: 1167: 1301: 1284:International Journal of Plant Sciences 14: 1627: 1468: 1366: 1314: 424:Cronq., Takht. & Zimmermann (1966) 219:Cronq., Takht. & Zimmermann (1966) 1560: 1559: 1386:Kidston, R. & Lang, W.H. (1920), 1207: 1164: 1139:Bulletin of the Torrey Botanical Club 729:article for the expanded cladogram.) 315:consisted of branched stems bearing 24: 469:In 2004, Crane et al. published a 290:Polysporangiophyte § Taxonomy 25: 1656: 1551:Crane, Herendeen & Friis 2004 1530: 1640:Early Devonian first appearances 330:, the group was found not to be 256:). Sources vary in the name and 134: 45: 1360: 1352:Yakovlev, Nikolai, ed. (1925). 1003: 1340: 283: 13: 1: 1081: 294:The group was described as a 235:are a group of extinct early 996:has been considered to be a 464: 264:, others as the subdivision 7: 1069: 778:(cooksonioids + renalioids) 413: 10: 1661: 1645:Early Devonian extinctions 1435:American Journal of Botany 1321:, Beijing: Science Press, 1026:found in the Rhynie chert 1007: 428:Subdivision Rhyniophytina 400:Cooksonia crassiparietilis 287: 1568: 1521:10.1017/S0080456800004506 1416:10.1017/S0080456800004488 732: 673: 657: 650: 614: 597: 590: 573: 566: 556: 536: 529: 514: 507: 451:Kidston & Lang (1920) 202: 195: 131:Scientific classification 129: 121: 112: 34: 985:age (Llandovery, around 981:, found in deposits of 372:Rhynia gwynne-vaughanii 326:However, as originally 278:Rhynia gwynne-vaughanii 124:Rhynia gwynne-vaughanii 27:Extinct group of plants 1507:, Kidston and Lang.", 1448:10.3732/ajb.91.10.1683 1390:Rhynia Gwynne-Vaughani 1030:- rich fossil beds in 1635:Early Devonian plants 1594:Paleobiology Database 422:Division Rhyniophyta 392:C. hemisphaerica 1135:Psilophyton forbesii 988:440 to 430 252:419 to 393 1505:Asteroxylon mackiei 1076:Polysporangiophytes 504:polysporangiophytes 309:polysporangiophytes 304:Harlan Parker Banks 161:Polysporangiophytes 1014:The general term " 727:Polysporangiophyte 449:Family Rhyniaceae 434:Class Rhyniopsida 388:C. paranensis 362:In 2010, the name 122:Reconstruction of 1622: 1621: 1562:Taxon identifiers 1328:978-7-03-036616-0 1191:978-0-12-373972-8 1115:978-1-56098-730-7 990:million years ago 722: 721: 713: 712: 704: 703: 695: 694: 686: 685: 638: 637: 629: 628: 485:). However, both 452: 446: 440: 431: 384:Cooksonia pertoni 364:paratracheophytes 254:million years ago 229: 228: 223:paratracheophytes 220: 214: 208: 16:(Redirected from 1652: 1615: 1614: 1602: 1601: 1589: 1588: 1587: 1557: 1556: 1524: 1523: 1500: 1494: 1493: 1477:(5/6): 505–516, 1466: 1460: 1459: 1450: 1430: 1419: 1418: 1383: 1377: 1376: 1364: 1358: 1357: 1344: 1338: 1337: 1336: 1335: 1312: 1299: 1298: 1275: 1269: 1268: 1241: 1232: 1231: 1223: 1214: 1213: 1205: 1196: 1195: 1177: 1162: 1161: 1130: 1119: 1118: 1101: 991: 662:and stem groups 660:Lycopodiophytina 653: 652: 648:Eutracheophytes 593: 592: 569: 568: 559: 558: 532: 531: 517:Horneophytopsida 510: 509: 500: 499: 450: 444: 443:Order Rhyniales 435: 429: 425: 298:of the division 268:or the division 255: 218: 212: 206: 188: 139: 138: 117: 107: 44: 38:Temporal range: 32: 31: 21: 1660: 1659: 1655: 1654: 1653: 1651: 1650: 1649: 1625: 1624: 1623: 1618: 1610: 1605: 1597: 1592: 1583: 1582: 1577: 1564: 1533: 1528: 1527: 1501: 1497: 1483:10.2307/1219787 1467: 1463: 1441:(10): 1683–99, 1431: 1422: 1398:Hornea Lignieri 1384: 1380: 1365: 1361: 1345: 1341: 1333: 1331: 1329: 1313: 1302: 1276: 1272: 1258:10.2307/1217531 1242: 1235: 1224: 1217: 1206: 1199: 1192: 1178: 1165: 1151:10.2307/2483801 1131: 1122: 1116: 1102: 1089: 1084: 1072: 1012: 1006: 986: 973: 797:) (rhynialeans) 735: 723: 714: 705: 696: 687: 676:Euphyllophytina 639: 630: 467: 423: 416: 321:vascular tissue 292: 286: 250: 245:, found in the 237:vascular plants 186: 133: 108: 106: 105: 100: 95: 90: 85: 80: 75: 70: 65: 60: 55: 50: 40: 39: 36: 28: 23: 22: 15: 12: 11: 5: 1658: 1648: 1647: 1642: 1637: 1620: 1619: 1617: 1616: 1603: 1590: 1574: 1572: 1566: 1565: 1554: 1553: 1544: 1539: 1532: 1531:External links 1529: 1526: 1525: 1515:(3): 643–680, 1495: 1461: 1420: 1410:(3): 603–627, 1400:, n.g., n.sp." 1378: 1359: 1339: 1327: 1300: 1296:10.1086/648988 1290:(2): 199–215, 1270: 1252:(4): 129–134, 1233: 1215: 1197: 1190: 1163: 1120: 1114: 1086: 1085: 1083: 1080: 1079: 1078: 1071: 1068: 1038:, and roughly 1020:rhyniophytoids 1008:Main article: 1005: 1002: 983:Early Silurian 972: 971: 963: 956: 955:(rhynialeans?) 948: 940: 933: 930:Sporathylacium 926: 919: 912: 911:(rhynialeans?) 904: 896: 889: 881: 874: 858: 850: 843: 842:) (renalioids) 831: 824: 821:Fusiformitheca 812: 805: 798: 786: 779: 771: 764: 756: 747: 734: 731: 720: 719: 716: 715: 711: 710: 707: 706: 702: 701: 698: 697: 693: 692: 689: 688: 684: 683: 680: 679: 672: 669: 668: 665: 664: 656: 651: 649: 645: 644: 641: 640: 636: 635: 632: 631: 627: 626: 623: 622: 613: 610: 609: 606: 605: 596: 591: 589: 586: 585: 582: 581: 572: 567: 565: 557: 555: 549: 548: 545: 544: 535: 530: 528: 525: 524: 521: 520: 513: 508: 506: 498: 466: 463: 462: 461: 460: 459: 458: 457: 456: 455: 454: 453: 415: 412: 337:rhyniophytoids 311:, since their 285: 282: 247:Early Devonian 227: 226: 225: 224: 221: 215: 211:Rhyniophytina 209: 207:Krysht. (1925) 200: 199: 193: 192: 184: 177: 176: 171: 164: 163: 158: 151: 150: 145: 141: 140: 127: 126: 119: 118: 110: 109: 101: 96: 91: 86: 81: 76: 71: 66: 61: 56: 51: 46: 42:Early Devonian 37: 26: 9: 6: 4: 3: 2: 1657: 1646: 1643: 1641: 1638: 1636: 1633: 1632: 1630: 1613: 1608: 1604: 1600: 1595: 1591: 1586: 1580: 1576: 1575: 1573: 1571: 1567: 1563: 1558: 1552: 1548: 1545: 1543: 1540: 1538: 1535: 1534: 1522: 1518: 1514: 1510: 1506: 1499: 1492: 1488: 1484: 1480: 1476: 1472: 1465: 1458: 1454: 1449: 1444: 1440: 1436: 1429: 1427: 1425: 1417: 1413: 1409: 1405: 1401: 1399: 1396:, n.sp., and 1395: 1391: 1382: 1374: 1370: 1363: 1355: 1349: 1343: 1330: 1324: 1320: 1319: 1311: 1309: 1307: 1305: 1297: 1293: 1289: 1285: 1281: 1274: 1267: 1263: 1259: 1255: 1251: 1247: 1240: 1238: 1229: 1222: 1220: 1211: 1204: 1202: 1193: 1187: 1183: 1176: 1174: 1172: 1170: 1168: 1160: 1156: 1152: 1148: 1144: 1140: 1136: 1129: 1127: 1125: 1117: 1111: 1107: 1100: 1098: 1096: 1094: 1092: 1087: 1077: 1074: 1073: 1067: 1065: 1064: 1059: 1058: 1053: 1049: 1045: 1041: 1037: 1033: 1032:Aberdeenshire 1029: 1025: 1021: 1017: 1011: 1001: 999: 995: 994:Tortilicaulis 989: 984: 980: 979: 970:(rhynialeans) 969: 968: 964: 962: 961: 960:Tortilicaulis 957: 954: 953: 949: 947:(rhynialeans) 946: 945: 944:Stockmansella 941: 939: 938: 934: 932: 931: 927: 925: 924: 920: 918: 917: 913: 910: 909: 905: 903:(rhynialeans) 902: 901: 897: 895: 894: 890: 887: 886: 882: 880: 879: 875: 872: 868: 864: 863: 859: 857:(rhynialeans) 856: 855: 851: 849: 848: 844: 841: 837: 836: 832: 830: 829: 828:Grisellatheca 825: 822: 818: 817: 813: 811: 810: 809:Filiformorama 806: 804: 803: 799: 796: 792: 791: 787: 785: 784: 780: 777: 776: 772: 770: 769: 765: 763:(rhynialeans) 762: 761: 757: 754: 753: 749: 748: 746: 744: 740: 739:circumscribed 730: 728: 718: 717: 709: 708: 700: 699: 691: 690: 682: 681: 678: 677: 671: 670: 667: 666: 663: 661: 655: 654: 647: 646: 643: 642: 634: 633: 625: 624: 621: 620: 619: 618:Stockmansella 612: 611: 608: 607: 604: 603: 602: 595: 594: 588: 587: 584: 583: 580: 579: 578: 571: 570: 564: 561: 560: 554: 551: 550: 547: 546: 543: 542: 541: 534: 533: 527: 526: 523: 522: 519: 518: 512: 511: 505: 502: 501: 497: 494: 492: 488: 484: 480: 476: 472: 448: 447: 445:Němejc (1950) 442: 441: 438: 437:Kryshtofovich 433: 432: 427: 426: 421: 420: 419: 411: 409: 405: 401: 397: 393: 389: 385: 381: 377: 376:Stockmansella 373: 367: 365: 360: 357: 352: 350: 349: 344: 343: 338: 333: 329: 328:circumscribed 324: 322: 318: 314: 310: 305: 301: 297: 291: 281: 279: 275: 271: 267: 266:Rhyniophytina 263: 259: 253: 248: 244: 243: 238: 234: 222: 216: 210: 205:Rhyniopsida 204: 203: 201: 198: 194: 191: 185: 182: 179: 178: 175: 174:Tracheophytes 172: 169: 166: 165: 162: 159: 156: 153: 152: 149: 146: 143: 142: 137: 132: 128: 125: 120: 116: 111: 104: 99: 94: 89: 84: 79: 74: 69: 64: 59: 54: 49: 43: 33: 30: 19: 18:Rhyniophytina 1569: 1512: 1508: 1504: 1498: 1474: 1470: 1464: 1438: 1434: 1407: 1403: 1397: 1394:Rhynia major 1393: 1389: 1381: 1372: 1368: 1362: 1353: 1347: 1342: 1332:, retrieved 1317: 1287: 1283: 1282:Lang 1937", 1279: 1273: 1249: 1245: 1227: 1209: 1181: 1142: 1138: 1134: 1105: 1061: 1055: 1019: 1016:rhyniophytes 1015: 1013: 1010:Rhynie chert 1004:Rhynie flora 993: 978:Eohostimella 976: 974: 965: 958: 950: 942: 937:Steganotheca 935: 928: 921: 914: 906: 898: 891: 888:(renalioids) 883: 876: 870: 866: 860: 852: 845: 839: 833: 826: 820: 814: 807: 800: 794: 788: 783:Culullitheca 781: 773: 766: 758: 755:(renalioids) 750: 743:cooksonioids 736: 724: 674: 658: 616: 615: 599: 598: 575: 574: 562: 553:Tracheophyta 538: 537: 515: 495: 490: 487:Horneophyton 486: 478: 475:Horneophyton 474: 468: 430:Banks (1968) 417: 403: 399: 395: 391: 387: 383: 379: 375: 371: 368: 363: 361: 355: 353: 348:Horneophyton 346: 340: 336: 332:monophyletic 325: 300:Tracheophyta 293: 277: 269: 265: 261: 240: 233:rhyniophytes 232: 230: 217:Rhyniophyta 213:Banks (1968) 190:Rhyniophytes 189: 180: 167: 154: 123: 29: 1570:Rhyniophyta 1145:(1): 1–11, 1063:Asteroxylon 1057:Aglaophyton 1028:Lagerstätte 998:horneophyte 923:Sennicaulis 916:Sartilmania 893:Resilitheca 871:Eocooksonia 867:Cooksonella 790:Eogaspesiea 760:Aglaophyton 540:Aglaophyton 491:Aglaophyton 479:Aglaophyton 408:protosteles 356:Paleobotany 342:Aglaophyton 313:sporophytes 296:subdivision 284:Definitions 270:Rhyniophyta 262:Rhyniopsida 35:Rhyniophyte 1629:Categories 1334:2019-10-25 1082:References 878:Pertonella 795:Eogaspesia 752:Aberlemnia 563:Rhyniaceae 396:Aberlemnia 288:See also: 181:Stem group 1612:100474404 1547:Cladogram 1348:Rhyniales 1280:Cooksonia 1044:floristic 952:Tarrantia 908:Salopella 862:Junggaria 816:Fusitheca 775:Cooksonia 725:(See the 483:tracheids 471:cladogram 465:Phylogeny 317:sporangia 144:Kingdom: 1607:Tropicos 1579:Wikidata 1457:21652317 1070:See also 1036:Scotland 967:Uskiella 802:Eorhynia 414:Taxonomy 249:(around 197:Synonyms 1585:Q133366 1491:1219787 1375:(3): 55 1266:1217531 1159:2483801 1054:, like 1052:anatomy 885:Renalia 854:Huvenia 577:Huvenia 404:Renalia 380:Huvenia 148:Plantae 1599:250240 1489:  1455:  1325:  1264:  1188:  1157:  1112:  1040:coeval 1024:plants 1018:" or " 900:Rhynia 733:Genera 601:Rhynia 439:(1925) 242:Rhynia 1549:from 1487:JSTOR 1471:Taxon 1262:JSTOR 1246:Taxon 1155:JSTOR 1048:taxon 274:Banks 168:Clade 155:Clade 1453:PMID 1350:in: 1323:ISBN 1186:ISBN 1110:ISBN 847:Huia 840:Hsüa 835:Hsua 768:Caia 489:and 477:and 402:and 390:and 378:and 345:and 258:rank 231:The 48:PreꞒ 1517:doi 1479:doi 1443:doi 1412:doi 1346:As 1292:doi 1288:171 1254:doi 1147:doi 865:(= 819:(= 793:(= 302:by 1631:: 1609:: 1596:: 1581:: 1513:52 1511:, 1485:, 1475:28 1473:, 1451:, 1439:91 1437:, 1423:^ 1408:52 1406:, 1402:, 1371:, 1303:^ 1286:, 1260:, 1248:, 1236:^ 1218:^ 1200:^ 1166:^ 1153:, 1143:95 1141:, 1123:^ 1090:^ 1034:, 1000:. 869:, 838:(= 410:. 398:, 386:, 374:, 183:: 170:: 157:: 98:Pg 1519:: 1481:: 1445:: 1414:: 1373:6 1294:: 1256:: 1250:2 1194:. 1149:: 873:) 823:) 187:† 103:N 93:K 88:J 83:T 78:P 73:C 68:D 63:S 58:O 53:Ꞓ 20:)

Index

Rhyniophytina
Early Devonian
PreꞒ

O
S
D
C
P
T
J
K
Pg
N

Scientific classification
Edit this classification
Plantae
Polysporangiophytes
Tracheophytes
Rhyniophytes
Synonyms
vascular plants
Rhynia
Early Devonian
419 to 393
rank
Banks
Polysporangiophyte § Taxonomy
subdivision

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