782:
845:
814:
132:
931:
714:
373:
857:
108:
900:
869:
748:
1474:
Zamora, Samuel; Lefebvre, Bertrand; Javier Álvaro, J.; Clausen, Sébastien; Elicki, Olaf; Fatka, Oldrich; Jell, Peter; Kouchinsky, Artem; Lin, Jih-Pai; Nardin, Elise; Parsley, Ronald; Rozhnov, Sergei; Sprinkle, James; Sumrall, Colin D.; Vizcaïno, Daniel; Smith, Andrew B. (2013). "Cambrian echinoderm
951:
Ctenocystoids were widespread during the Middle
Cambrian, and have been found in the United States, the United Kingdom, Australia, France, the Czech Republic, Poland, Spain, and Morocco. The earliest ctenocystoids date to the beginning of Stage 5 of the Cambrian, now known as the
1549:
Zhao, Yuanlong; Yuan, Jinliang; Babcock, Loren E.; Guo, Qingjun; Peng, Jin; Yin, Leiming; Yang, Xinglian; Peng, Shanchi; Wang, Chunjiang; Gaines, Robert R.; Esteve, Jorge; Tai, Tongsu; Yang, Ruidong; Wang, Yue; Sun, Haijing; Yang, Yuning (2019).
417:
All ctenocystoids had a ctenoid apparatus, a comb-like arrangement of movable plates at the anterior end of the animal. Beneath the ctenoid apparatus was a mouth, bordered on each side by a food groove.
1214:
336:, or were only very slightly asymmetrical. They are believed to be one of the earliest-diverging branches of echinoderms, with their bilateral symmetry a trait shared with other
1552:"Global Standard Stratotype-Section and Point (GSSP) for the conterminous base of the Miaolingian Series and Wuliuan Stage (Cambrian) at Balang, Jianhe, Guizhou, China"
356:. Homalozoa is now recognized as a polyphyletic group of echinoderms without radial symmetry. Ctenocystoids were geographically widespread during the
643:. The presence of stereom plates indicates that they most likely belong to the echinoderm total group, rendering a hemichordate affinity unlikely.
269:
1699:
1350:
David, Bruno; Lefebvre, Bertrand; Mooi, Rich; Parsley, Ronald (2000). "Are homalozoans echinoderms? An answer from the extraxial-axial theory".
299:
249:
259:
289:
1158:"The youngest ctenocystoids from the Upper Ordovician of the United Kingdom and the evolution of the bilateral body plan in echinoderms"
279:
239:
414:
as in other echinoderms. It was located at the posterior end, defining a clear anterior-posterior body axis unlike other echinoderms.
1030:
Rahman, Imran A.; Clausen, Sébastien (2009). "Re-evaluating the palaeobiology and affinities of the
Ctenocystoidea (Echinodermata)".
388:
plates. Ctenocystoids had near bilateral symmetry, with some species exhibiting slightly asymmetric plate shapes. Like cinctans and
1512:"Laurentian origin of solutan echinoderms: new evidence from the Guzhangian (Cambrian Series 3) Weeks Formation of Utah, USA"
1511:
988:
The class
Ctenocystoidea was named in 1969 by Richard A. Robison and James Sprinkle. It originally contained one species,
1758:
119:
1334:
635:
Ctenocystoids are likely among the most basal stem-group echinoderms. They have also been interpreted as aberrant
1371:
1326:
Sistemática, anatomía, estructura y función de
Ctenocystoidea (Echinodermata, Carpoidea) del Paleozoico Inferior
1753:
661:
may be closely related to each other, as both have a reduced marginal frame compared to other ctenocystoids.
394:, most ctenocystoids had large marginal plates surrounding the sides of their body, but unlike cinctans and
131:
1000:
994:
1585:"The oldest echinoderm faunas from Gondwana show that echinoderm body plan diversification was rapid"
1241:
1184:
1680:
398:, which had only one row of marginal plates, most ctenocystoids had two rows of marginal plates.
1730:
1642:
1725:
1717:
1704:
1596:
1523:
1414:
1359:
1278:
1269:
Robison, R. A.; Sprinkle, J. (1969). "Ctenocystoidea: New Class of
Primitive Echinoderms".
1091:
1039:
410:
lost the marginal plates entirely. The anus of ctenocystoids was surrounded by a pyramidal
8:
349:
1600:
1527:
1418:
1363:
1282:
1095:
1043:
1763:
1492:
1386:
1302:
1157:
1122:
1079:
1055:
333:
126:
1712:
1622:
1614:
1568:
1551:
1496:
1453:
1426:
1330:
1294:
1246:
1189:
1127:
1109:
1059:
1394:. Symposium of Vertebrate Palaeontology and Comparative Anatomy. London. p. 30.
1306:
1604:
1563:
1531:
1484:
1422:
1388:
A cladogram for the
Deuterostomia based on molecular-biological and fossil evidence
1367:
1286:
1236:
1226:
1179:
1169:
1117:
1099:
1047:
957:
1290:
1104:
774:
361:
357:
1080:"Plated Cambrian bilaterians reveal the earliest stages of echinoderm evolution"
1665:
1440:
Domínguez Alnoso, Patricio; Jefferies, Richard P. S.; Gil Cid, Dolores (2002).
936:
819:
719:
649:
was probably the most basal ctenocystoid, given its anatomical similarities to
390:
1535:
1051:
1747:
1618:
1457:
1250:
1193:
1113:
787:
640:
65:
1626:
1298:
1174:
1131:
337:
176:
163:
1691:
1674:
40:
1004:, meaning "sac". It was originally assigned to the echinoderm subphylum
1609:
1584:
1231:
969:
353:
329:
321:
210:
186:
85:
50:
1215:"Deciphering the early evolution of echinoderms with Cambrian fossils"
968:. The geologically youngest ctenocystoid, the only one known from the
372:
1005:
850:
636:
411:
341:
143:
90:
1636:
1659:
1488:
977:
965:
706:
325:
107:
80:
75:
60:
55:
45:
34:
1473:
1385:
Dominguez-Alonso, Patricio; Jefferies, Richard (September 2005).
1078:
Zamora, Samuel; Rahman, Imran A.; Smith, Andrew B. (2012-06-06).
961:
953:
862:
740:
385:
314:
95:
70:
30:
1439:
1441:
1324:
905:
874:
753:
345:
153:
384:
Like other echinoderms, ctenocystoids had a skeleton made of
376:
A selection of early echinoderms, including the ctenocystoid
317:
1583:
Smith, Andrew B.; Zamora, Samuel; Álvaro, J. Javier (2013).
1156:
Rahman, Imran A.; Stewart, Sarah E.; Zamora, Samuel (2015).
1442:"An annotated check-list of genera and species of carpoids"
1372:
10.1666/0094-8373(2000)026<0529:AHEAAF>2.0.CO;2
1384:
1349:
669:
The following genera of ctenocystoids have been named:
402:
had only one row of marginal plates, like cinctans and
1509:
1405:
Rahman, Imran A. (2009). "Making sense of carpoids".
332:
periods. Unlike other echinoderms, ctenocystoids had
295:
285:
275:
265:
255:
245:
235:
1155:
1548:
340:. Ctenocystoids were once classified in the taxon
1582:
1510:Lefebvre, Bertrand; Lerosey-Aubril, Rudy (2018).
1077:
629:Phylogenetic relationships within Ctenocystoidea
1745:
1322:
1318:
1316:
1268:
1469:
1467:
1433:
1264:
1262:
1260:
1029:
1503:
1378:
1313:
1212:
1151:
1149:
1147:
1145:
1143:
1141:
1073:
1071:
1069:
1025:
1023:
1021:
956:age, or possibly slightly earlier, in late
1464:
1398:
1329:(PhD). Universidad Complutense de Madrid.
1257:
106:
1608:
1576:
1567:
1242:1983/951cc086-7ed1-469b-98ac-2cc338c4e823
1240:
1230:
1213:Zamora, Samuel; Rahman, Imran A. (2015).
1206:
1185:1983/6531d6a6-2b00-43d0-8f2d-0a8d5df836d0
1183:
1173:
1138:
1121:
1103:
1066:
1018:
1542:
1343:
371:
960:. Most species date to the Wuliuan and
1746:
1404:
1090:(6). Keith A. Crandall (ed.): –38296.
992:. The name comes from the Greek words
360:, with one species surviving into the
1641:
1640:
924:
893:
838:
807:
773:
739:
705:
344:, also known as Carpoidea, alongside
16:Extinct clade of marine invertebrates
1225:(6). Andrew Smith (ed.): 1105–1119.
1477:Geological Society, London, Memoirs
1475:diversity and palaeobiogeography".
1323:Domínguez Alonso, Patricio (1999).
1032:Journal of Systematic Palaeontology
13:
120:National Museum of Natural History
14:
1775:
421:
1427:10.1111/j.1365-2451.2009.00703.x
929:
898:
867:
855:
843:
812:
780:
746:
724:The geologically youngest genus
712:
426:
130:
38:
946:
367:
1:
1291:10.1126/science.166.3912.1512
1162:Acta Palaeontologica Polonica
1011:
1569:10.18814/epiiugs/2019/019013
1105:10.1371/journal.pone.0038296
222:Robison & Sprinkle, 1969
204:Robison & Sprinkle, 1969
7:
792:The first genus discovered
671:
10:
1780:
1759:Prehistoric animal classes
1446:Coloquios de Paleontología
983:
1649:
1536:10.1017/S0016756817000152
1052:10.1017/S1477201909990046
664:
574:
559:
552:
537:
530:
515:
508:
484:
469:
462:
455:
440:
324:, which lived during the
232:
227:
216:
209:
127:Scientific classification
125:
114:
105:
23:
980:of the United Kingdom.
768:Robison & Sprinkle
1175:10.4202/app.00048.2013
998:, meaning "comb", and
381:
1754:Paleozoic echinoderms
1726:Paleobiology Database
1589:Nature Communications
990:Ctenocystis utahensis
758:The most basal genus
375:
116:Ctenocystis utahensis
888:Džik & Orłowski
802:Fatka & Kordule
1601:2013NatCo...4.1385S
1528:2018GeoM..155.1190L
1516:Geological Magazine
1419:2009GeolT..25...34R
1364:2000Pbio...26..529D
1283:1969Sci...166.1512R
1277:(3912): 1512–1514.
1096:2012PLoSO...738296Z
1044:2009JSPal...7..413R
1610:10.1038/ncomms2391
1232:10.1111/pala.12138
639:and as stem-group
382:
334:bilateral symmetry
1741:
1740:
1713:Open Tree of Life
1643:Taxon identifiers
944:
943:
919:Domínguez Alonso
833:Domínguez Alonso
734:Domínguez Alonso
700:Domínguez Alonso
633:
632:
623:
622:
614:
613:
605:
604:
596:
595:
587:
586:
497:
496:
308:
307:
205:
1771:
1734:
1733:
1721:
1720:
1708:
1707:
1695:
1694:
1685:
1684:
1683:
1670:
1669:
1668:
1638:
1637:
1631:
1630:
1612:
1580:
1574:
1573:
1571:
1546:
1540:
1539:
1522:(5): 1190–1204.
1507:
1501:
1500:
1471:
1462:
1461:
1437:
1431:
1430:
1402:
1396:
1395:
1393:
1382:
1376:
1375:
1347:
1341:
1340:
1320:
1311:
1310:
1266:
1255:
1254:
1244:
1234:
1210:
1204:
1203:
1201:
1200:
1187:
1177:
1153:
1136:
1135:
1125:
1107:
1075:
1064:
1063:
1027:
958:Cambrian Stage 4
935:
933:
932:
925:Middle Cambrian
904:
902:
901:
894:Middle Cambrian
873:
871:
870:
861:
859:
858:
849:
847:
846:
839:Middle Cambrian
818:
816:
815:
808:Middle Cambrian
786:
784:
783:
752:
750:
749:
718:
716:
715:
672:
555:
554:
533:
532:
511:
510:
465:
464:
458:
457:
436:
435:
431:
430:
297:
287:
277:
267:
257:
247:
237:
203:
198:
135:
134:
110:
100:
37:
27:Temporal range:
21:
20:
1779:
1778:
1774:
1773:
1772:
1770:
1769:
1768:
1744:
1743:
1742:
1737:
1729:
1724:
1716:
1711:
1703:
1698:
1690:
1688:
1679:
1678:
1673:
1664:
1663:
1658:
1645:
1635:
1634:
1581:
1577:
1547:
1543:
1508:
1504:
1472:
1465:
1438:
1434:
1403:
1399:
1391:
1383:
1379:
1348:
1344:
1337:
1321:
1314:
1267:
1258:
1211:
1207:
1198:
1196:
1154:
1139:
1076:
1067:
1028:
1019:
1014:
986:
949:
930:
928:
899:
897:
868:
866:
856:
854:
844:
842:
813:
811:
781:
779:
775:Middle Cambrian
747:
745:
713:
711:
667:
624:
615:
606:
597:
588:
498:
429:
424:
370:
362:Late Ordovician
358:Middle Cambrian
223:
220:
202:
196:
129:
101:
99:
98:
93:
88:
83:
78:
73:
68:
63:
58:
53:
48:
43:
29:
28:
25:
17:
12:
11:
5:
1777:
1767:
1766:
1761:
1756:
1739:
1738:
1736:
1735:
1722:
1709:
1696:
1686:
1681:Ctenocystoidea
1671:
1655:
1653:
1651:Ctenocystoidea
1647:
1646:
1633:
1632:
1575:
1562:(2): 165–184.
1541:
1502:
1489:10.1144/M38.13
1483:(1): 157–171.
1463:
1432:
1397:
1377:
1358:(4): 529–555.
1342:
1335:
1312:
1256:
1205:
1137:
1065:
1038:(4): 413–426.
1016:
1015:
1013:
1010:
985:
982:
948:
945:
942:
941:
939:
937:United Kingdom
926:
923:
920:
917:
911:
910:
908:
895:
892:
889:
886:
880:
879:
877:
840:
837:
834:
831:
825:
824:
822:
820:Czech Republic
809:
806:
803:
800:
798:Etoctenocystis
794:
793:
790:
777:
772:
769:
766:
760:
759:
756:
743:
738:
735:
732:
726:
725:
722:
720:United Kingdom
709:
704:
701:
698:
692:
691:
688:
685:
682:
679:
676:
666:
663:
653:and cinctans.
651:Ctenoimbricata
631:
630:
626:
625:
621:
620:
617:
616:
612:
611:
608:
607:
603:
602:
599:
598:
594:
593:
590:
589:
585:
584:
581:
580:
573:
570:
569:
566:
565:
558:
553:
551:
548:
547:
544:
543:
540:Etoctenocystis
536:
531:
529:
526:
525:
522:
521:
514:
509:
507:
504:
503:
500:
499:
495:
494:
491:
490:
483:
480:
479:
476:
475:
468:
463:
461:
456:
454:
451:
450:
447:
446:
439:
434:
428:
425:
423:
422:Classification
420:
404:Ctenoimbricata
396:Ctenoimbricata
391:Ctenoimbricata
369:
366:
311:Ctenocystoidea
306:
305:
304:
303:
293:
283:
273:
270:Etoctenocystis
263:
253:
243:
230:
229:
225:
224:
221:
214:
213:
207:
206:
200:Ctenocystoidea
194:
190:
189:
184:
180:
179:
174:
167:
166:
161:
157:
156:
151:
147:
146:
141:
137:
136:
123:
122:
112:
111:
103:
102:
94:
89:
84:
79:
74:
69:
64:
59:
54:
49:
44:
39:
26:
24:Ctenocystoidea
15:
9:
6:
4:
3:
2:
1776:
1765:
1762:
1760:
1757:
1755:
1752:
1751:
1749:
1732:
1727:
1723:
1719:
1714:
1710:
1706:
1701:
1697:
1693:
1687:
1682:
1676:
1672:
1667:
1661:
1657:
1656:
1654:
1652:
1648:
1644:
1639:
1628:
1624:
1620:
1616:
1611:
1606:
1602:
1598:
1594:
1590:
1586:
1579:
1570:
1565:
1561:
1557:
1553:
1545:
1537:
1533:
1529:
1525:
1521:
1517:
1513:
1506:
1498:
1494:
1490:
1486:
1482:
1478:
1470:
1468:
1459:
1455:
1451:
1447:
1443:
1436:
1428:
1424:
1420:
1416:
1412:
1408:
1407:Geology Today
1401:
1390:
1389:
1381:
1373:
1369:
1365:
1361:
1357:
1353:
1346:
1338:
1336:84-669-2421-3
1332:
1328:
1327:
1319:
1317:
1308:
1304:
1300:
1296:
1292:
1288:
1284:
1280:
1276:
1272:
1265:
1263:
1261:
1252:
1248:
1243:
1238:
1233:
1228:
1224:
1220:
1219:Palaeontology
1216:
1209:
1195:
1191:
1186:
1181:
1176:
1171:
1167:
1163:
1159:
1152:
1150:
1148:
1146:
1144:
1142:
1133:
1129:
1124:
1119:
1115:
1111:
1106:
1101:
1097:
1093:
1089:
1085:
1081:
1074:
1072:
1070:
1061:
1057:
1053:
1049:
1045:
1041:
1037:
1033:
1026:
1024:
1022:
1017:
1009:
1007:
1003:
1002:
997:
996:
991:
981:
979:
975:
971:
967:
963:
959:
955:
940:
938:
927:
921:
918:
916:
913:
912:
909:
907:
896:
890:
887:
885:
882:
881:
878:
876:
864:
852:
841:
835:
832:
830:
827:
826:
823:
821:
810:
804:
801:
799:
796:
795:
791:
789:
788:United States
778:
776:
770:
767:
765:
762:
761:
757:
755:
744:
742:
736:
733:
731:
728:
727:
723:
721:
710:
708:
702:
699:
697:
694:
693:
689:
686:
683:
680:
677:
674:
673:
670:
662:
660:
656:
652:
648:
644:
642:
641:hemichordates
638:
628:
627:
619:
618:
610:
609:
601:
600:
592:
591:
583:
582:
579:
578:
572:
571:
568:
567:
564:
563:
557:
556:
550:
549:
546:
545:
542:
541:
535:
534:
528:
527:
524:
523:
520:
519:
513:
512:
506:
505:
502:
501:
493:
492:
489:
488:
482:
481:
478:
477:
474:
473:
467:
466:
460:
459:
453:
452:
449:
448:
445:
444:
438:
437:
433:
432:
427:Relationships
419:
415:
413:
409:
405:
401:
397:
393:
392:
387:
379:
374:
365:
363:
359:
355:
351:
347:
343:
339:
338:deuterostomes
335:
331:
327:
323:
319:
316:
312:
302:
301:
294:
292:
291:
284:
282:
281:
274:
272:
271:
264:
262:
261:
254:
252:
251:
244:
242:
241:
234:
233:
231:
226:
219:
215:
212:
208:
201:
195:
192:
191:
188:
187:Echinodermata
185:
182:
181:
178:
175:
172:
169:
168:
165:
164:Deuterostomia
162:
160:Superphylum:
159:
158:
155:
152:
149:
148:
145:
142:
139:
138:
133:
128:
124:
121:
117:
113:
109:
104:
97:
92:
87:
82:
77:
72:
67:
62:
57:
52:
47:
42:
36:
32:
22:
19:
1650:
1592:
1588:
1578:
1559:
1555:
1544:
1519:
1515:
1505:
1480:
1476:
1449:
1445:
1435:
1413:(1): 34–38.
1410:
1406:
1400:
1387:
1380:
1355:
1352:Paleobiology
1351:
1345:
1325:
1274:
1270:
1222:
1218:
1208:
1197:. Retrieved
1168:(1): 39–48.
1165:
1161:
1087:
1083:
1035:
1031:
999:
993:
989:
987:
973:
964:ages of the
950:
947:Distribution
915:Pembrocystis
914:
883:
828:
797:
763:
730:Courtessolea
729:
695:
668:
658:
654:
650:
647:Courtessolea
646:
645:
634:
576:
575:
561:
560:
539:
538:
518:Pembrocystis
517:
516:
486:
485:
471:
470:
443:Courtessolea
442:
441:
416:
407:
403:
400:Courtessolea
399:
395:
389:
383:
377:
354:stylophorans
310:
309:
300:Pembrocystis
298:
288:
278:
268:
258:
250:Courtessolea
248:
238:
217:
199:
177:Ambulacraria
170:
115:
18:
1675:Wikispecies
1595:(1): 1385.
976:, from the
764:Ctenocystis
678:Discoverers
637:blastozoans
577:Ctenocystis
378:Ctenocystis
368:Description
322:echinoderms
260:Ctenocystis
218:Ctenocystis
1748:Categories
1199:2020-04-02
1012:References
970:Ordovician
884:Jugoszovia
659:Jugoszovia
472:Jugoszovia
406:, whereas
330:Ordovician
290:Jugoszovia
211:Type genus
1764:Homalozoa
1619:2041-1723
1497:130481550
1458:1132-1660
1452:: 33–68.
1251:0031-0239
1194:0567-7920
1114:1932-6203
1060:129093037
1006:Homalozoa
851:Australia
412:periproct
380:(middle).
342:Homalozoa
150:Kingdom:
144:Eukaryota
1689:BioLib:
1666:Q5792856
1660:Wikidata
1627:23340425
1556:Episodes
1307:26730941
1299:17742852
1132:22701623
1084:PLOS ONE
978:Sandbian
974:Conollia
966:Cambrian
829:Gilcidia
707:Sandbian
696:Conollia
687:Location
655:Conollia
562:Gilcidia
487:Conollia
408:Conollia
346:cinctans
326:Cambrian
280:Gilcidia
240:Conollia
183:Phylum:
154:Animalia
140:Domain:
118:fossil,
35:Sandbian
1718:5292828
1597:Bibcode
1524:Bibcode
1415:Bibcode
1360:Bibcode
1279:Bibcode
1271:Science
1123:3368939
1092:Bibcode
1040:Bibcode
984:History
962:Drumian
954:Wuliuan
863:Denmark
741:Wuliuan
386:stereom
350:solutes
315:extinct
228:Genera
193:Class:
31:Wuliuan
1692:917648
1625:
1617:
1495:
1456:
1333:
1305:
1297:
1249:
1192:
1130:
1120:
1112:
1058:
1001:kystis
995:ktenos
934:
906:Poland
903:
875:France
872:
860:
848:
817:
785:
754:France
751:
717:
690:Notes
665:Genera
352:, and
313:is an
1731:31127
1700:IRMNG
1493:S2CID
1392:(PDF)
1303:S2CID
1056:S2CID
972:, is
922:1999
891:1995
836:1999
805:1985
771:1969
737:1999
703:1999
681:Named
675:Genus
318:clade
171:Clade
1705:1382
1623:PMID
1615:ISSN
1454:ISSN
1331:ISBN
1295:PMID
1247:ISSN
1190:ISSN
1128:PMID
1110:ISSN
657:and
328:and
41:PreꞒ
1605:doi
1564:doi
1532:doi
1520:155
1485:doi
1423:doi
1368:doi
1287:doi
1275:166
1237:hdl
1227:doi
1180:hdl
1170:doi
1118:PMC
1100:doi
1048:doi
684:Age
320:of
1750::
1728::
1715::
1702::
1677::
1662::
1621:.
1613:.
1603:.
1591:.
1587:.
1560:42
1558:.
1554:.
1530:.
1518:.
1514:.
1491:.
1481:38
1479:.
1466:^
1450:53
1448:.
1444:.
1421:.
1411:25
1409:.
1366:.
1356:26
1354:.
1315:^
1301:.
1293:.
1285:.
1273:.
1259:^
1245:.
1235:.
1223:57
1221:.
1217:.
1188:.
1178:.
1166:60
1164:.
1160:.
1140:^
1126:.
1116:.
1108:.
1098:.
1086:.
1082:.
1068:^
1054:.
1046:.
1034:.
1020:^
1008:.
865:,
853:,
364:.
348:,
173::
91:Pg
1629:.
1607::
1599::
1593:4
1572:.
1566::
1538:.
1534::
1526::
1499:.
1487::
1460:.
1429:.
1425::
1417::
1374:.
1370::
1362::
1339:.
1309:.
1289::
1281::
1253:.
1239::
1229::
1202:.
1182::
1172::
1134:.
1102::
1094::
1088:7
1062:.
1050::
1042::
1036:7
296:†
286:†
276:†
266:†
256:†
246:†
236:†
197:†
96:N
86:K
81:J
76:T
71:P
66:C
61:D
56:S
51:O
46:Ꞓ
33:–
Text is available under the Creative Commons Attribution-ShareAlike License. Additional terms may apply.
