1583:
1733:
1650:
1703:
1466:
1672:
1378:
1328:
149:
1238:
1503:
118:
1268:
819:
1890:
1782:
1298:
1910:
1523:
958:
1933:
1358:
1563:
2575:
Elphick, Maurice R.; Thomas-Chollier, Morgane; Hankeln, Thomas; Mertes, Florian; Wallberg, Andreas; Rast, Jonathan P.; Copley, Richard R.; Martinez, Pedro; Telford, Maximilian J. (2019). "Mitigating
Anticipated Effects of Systematic Errors Supports Sister-Group Relationship between Xenacoelomorpha and Ambulacraria".
850:, in which the developmental fate of the cells in the developing embryo is not determined by the identity of the parent cell. Thus, if the first four cells are separated, each can develop into a complete small larva; and if a cell is removed from the blastula, the other cells will compensate. This is the source of
822:
Early development differences between deuterostomes versus protostomes. In deuterostomes, blastula divisions occur as radial cleavage because they occur parallel or perpendicular to the major polar axis. In protostomes the cleavage is spiral because division planes are oriented obliquely to the polar
940:
The defining characteristic of the deuterostome is the fact that the blastopore (the opening at the bottom of the forming gastrula) becomes the anus, whereas in protostomes the blastopore becomes the mouth. The deuterostome mouth develops at the opposite end of the embryo, from the blastopore, and a
584:
may also be deuterostomes, but molecular studies have placed them in the protostomes more often. Genetic studies have also revealed that deuterostomes have more than 30 genes not found in any other animal groups, but which yet are present in some marine algae and prokaryotes. This could mean they
944:
In many animals, these early development stages later evolved in ways that no longer reflect these original patterns. For instance, humans have already formed a gut tube at the time of formation of the mouth and anus. Then the mouth forms first, during the fourth week of development, and the anus
2574:
Philippe, HervĂ©; Poustka, Albert J.; Chiodin, Marta; Hoff, Katharina J.; Dessimoz, Christophe; Tomiczek, Bartlomiej; Schiffer, Philipp H.; MĂŒller, Steven; Domman, Daryl; Horn, Matthias; Kuhl, Heiner; Timmermann, Bernd; Satoh, Noriyuki; Hikosaka-Katayama, Tomoe; Nakano, Hiroaki; Rowe, Matthew L.;
2514:
Bourlat, Sarah J.; Juliusdottir, Thorhildur; Lowe, Christopher J.; Freeman, Robert; Aronowicz, Jochanan; Kirschner, Mark; Lander, Eric S.; Thorndyke, Michael; Nakano, Hiroaki; Kohn, Andrea B.; Heyland, Andreas; Moroz, Leonid L.; Copley, Richard R.; Telford, Maximilian J. (2006). "Deuterostome
1974:
Support for the clade
Deuterostomia is not unequivocal. In particular, the Ambulacraria are sometimes shown to be related to the Xenacoelomorpha. If true, this raises two possibilities: either the Ambulacraria are taken out of the deuterostome-protostome dichotomy (in which case the grouping
925:
The highly modified nervous system of echinoderms obscures much about their ancestry, but several facts suggest that all present deuterostomes evolved from a common ancestor that had pharyngeal gill slits, a hollow nerve cord, circular and longitudinal muscles and a segmented body.
3124:
Zamora, Samuel; Wright, David F.; Mooi, Rich; Lefebvre, Bertrand; Guensburg, Thomas E.; Gorzelak, PrzemysĆaw; David, Bruno; Sumrall, Colin D.; Cole, Selina R.; Hunter, Aaron W.; Sprinkle, James; Thompson, Jeffrey R.; Ewin, Timothy A. M.; Fatka, OldĆich; Nardin, Elise (2020-03-09).
1190:. The tree is based on 16S +18S rRNA sequence data and phylogenomic studies from multiple sources. The approximate dates for each radiation into a new clade are given in millions of years ago (Mya). Not all dates are consistent, as of date ranges only the center is given.
1170:, and various Jurassic-aged and Tertiary-aged spicules tentatively attributed to ascidians.. Fossils of Echinodermata remain very common after the Cambrian. Fossils of Hemichordata are less common, except for graptolites until the Lower-Carbonoferous.
1010:). It has been proposed that the ancestral deuterostome, before the chordate/ambulacrarian split, could have been a chordate-like animal with a terminal anus and pharyngeal openings but no gill slits, with active suspension feeding strategy.
3600:
Philippe, HervĂ©; Poustka, Albert J.; Chiodin, Marta; Hoff, Katharina J.; Dessimoz, Christophe; Tomiczek, Bartlomiej; Schiffer, Philipp H.; MĂŒller, Steven; Domman, Daryl; Horn, Matthias; Kuhl, Heiner (2019-06-03).
2396:
Marlétaz, Ferdinand; Martin, Elise; Perez, Yvan; Papillon, Daniel; Caubit, Xavier; Lowe, Christopher J.; Freeman, Bob; Fasano, Laurent; Dossat, Carole; Wincker, Patrick; Weissenbach, Jean (2006-08-01).
3488:
Kapli, Paschalia; Natsidis, Paschalis; Leite, Daniel J.; Fursman, Maximilian; Jeffrie, Nadia; Rahman, Imran A.; Philippe, Hervé; Copley, Richard R.; Telford, Maximilian J. (March 1, 2021).
3733:
Philippe, Hervé; Brinkmann, Henner; Copley, Richard R.; Moroz, Leonid L.; Nakano, Hiroaki; Poustka, Albert J.; Wallberg, Andreas; Peterson, Kevin J.; Telford, Maximilian J. (2011-02-10).
3798:
Edgecombe, Gregory D.; Giribet, Gonzalo; Dunn, Casey W.; Hejnol, Andreas; Kristensen, Reinhardt M.; Neves, Ricardo C.; Rouse, Greg W.; Worsaae, Katrine; SĂžrensen, Martin V. (June 2011).
1996:
Fedonkin, M. A.; Vickers-Rich, P.; Swalla, B. J.; Trusler, P.; Hall, M. (2012). "A new metazoan from the
Vendian of the White Sea, Russia, with possible affinities to the ascidians".
572:
based on morphological and embryological characteristics. However, Deuterostomia was redefined in 1995 based on DNA molecular sequence analyses, leading to the removal of the
2345:
Halanych, K.M.; Bacheller, J.; Liva, S.; Aguinaldo, A. A.; Hillis, D.M. & Lake, J.A. (17 March 1995). "18S rDNA evidence that the
Lophophorates are Protostome Animals".
441:, while the mouth is formed at a different site later on. This was initially the group's distinguishing characteristic, but deuterostomy has since been discovered among
2876:, the Vendian mollusc-like organism (White sea region, Russia): palaeoecological and evolutionary implications", in Vickers-Rich, Patricia; Komarower, Patricia (eds.),
823:
major axis. During gastrulation, deuterostome embryos' anus is given first by the blastopore while the mouth is formed secondarily, and vice versa for the protostomes
2753:
3191:
899:(in the larval stage). Some hemichordates also have a tubular nerve cord. In the early embryonic stage, it looks like the hollow nerve cord of chordates.
2122:
Han, Jian; Morris, Simon Conway; Ou, Qiang; Shu, Degan; Huang, Hai (2017). "Meiofaunal deuterostomes from the basal
Cambrian of Shaanxi (China)".
1029:. There are also a few earlier fossils that may represent deuterostomes, but these remain debated. The earliest of these disputed fossils are the
3892:
Cannon, Johanna Taylor; Vellutini, Bruno
Cossermelli; Smith III, Julian; Ronquist, Frederik; Jondelius, Ulf; Hejnol, Andreas (4 February 2016).
5466:
935:
3059:
Cracknell, Kelsie; GarcĂa-Bellido, Diego C.; Gehling, James G.; Ankor, Martin J.; Darroch, Simon A. F.; Rahman, Imran A. (2021-02-18).
3426:
Tassia, Michael G.; Cannon, Johanna T.; Konikoff, Charlotte E.; Shenkar, Noa; Halanych, Kenneth M.; Swalla, Billie J. (2016-10-04).
971:
and deuterostomes. Deuterostomes consist of chordates (which include the vertebrates) and ambulacrarians. It seems likely that the
5453:
3603:"Mitigating Anticipated Effects of Systematic Errors Supports Sister-Group Relationship between Xenacoelomorpha and Ambulacraria"
2502:
2095:
2893:
2763:
5458:
5512:
5522:
3020:
4100:
123:
1147:
is a acorn-worm from the
Burgess Shale, providing proof that all main lineages were already well established 508 mya.
3670:"The mitochondrial genomes of the acoelomorph worms Paratomella rubra, Isodiametra pulchra and Archaphanostoma ylvae"
918:, which contracts to pump blood. This suggests a presence in the deuterostome ancestor of the three groups, with the
3949:"Topology-dependent asymmetry in systematic errors affects phylogenetic placement of Ctenophora and Xenacoelomorpha"
2916:
Butterfield, N.J. (December 2006). "Hooking some stem-group "worms": fossil lophotrochozoans in the
Burgess Shale".
4137:
3668:
Robertson, Helen E.; Lapraz, François; Egger, Bernhard; Telford, Maximilian J.; Schiffer, Philipp H. (2017-05-12).
2043:; Ou, Qiang; Shu, Degan; Huang, Hai (2017). "Meiofaunal deuterostomes from the basal Cambrian of Shaanxi (China)".
1979:), or the Xenacoelomorpha are re-positioned next to Ambulacraria within the Deuterostomia as in the above diagram.
5471:
3549:; Ou, Qian; Shu, Degan; Huang, Hai (2017). "Meiofaunal deuterostomes from the basal Cambrian of Shaanxi (China)".
2454:
Marlétaz, Ferdinand; Peijnenburg, Katja T.C.A.; Goto, Taichiro; Satoh, Noriyuki; Rokhsar, Daniel S. (2019-01-21).
4132:
1182:
showing consensus relationships among deuterostome taxa. Phylogenomic evidence suggests the enteropneust family,
598:
129:
3218:
983:
was a member of the protostomes. That implies that the protostome and deuterostome lineages split long before
876:
Another feature present in both the
Hemichordata and Chordata is pharyngotremy â the presence of spiracles or
4216:
4206:
4189:
2959:
Li, Yujing; Dunn, Frances S.; Murdock, Duncan J.E.; Guo, Jin; Rahman, Imran A.; Cong, Peiyun (May 10, 2023).
128:
125:
127:
835:. In deuterostomes, the early divisions occur parallel or perpendicular to the polar axis. This is called
585:
are very ancient genes that were lost in other organisms, or that a common ancestor acquired them through
148:
3061:"Pentaradial eukaryote suggests expansion of suspension feeding in White Sea-aged Ediacaran communities"
2241:"Annelida and Arthropoda are Not Sister Taxa: A Phylogenetic Analysis of Spiralian Metazoan Morphology"
4072:
3045:
M. A. Fedonkin (1996). "Ausia as an ancestor of archeocyathans, and other sponge-like organisms". In:
1582:
1025:
Deuterostomes have a rich fossil record with thousands of fossil species being found throughout the
126:
124:
122:
586:
3127:"Re-evaluating the phylogenetic position of the enigmatic early Cambrian deuterostome Yanjiahella"
121:
5408:
4221:
1150:
On the other hand, fossils of early chordates are very rare, as non-vertebrate chordates have no
4012:"Deciphering deuterostome phylogeny: Molecular, morphological and palaeontological perspectives"
3837:
Rouse, Greg W.; Wilson, Nerida G.; Carvajal, Jose I.; Vriejenhoek, Robert C. (4 February 2016).
5484:
5370:
4093:
1133:
847:
5479:
4184:
2331:
1142:
522:
1154:
or teeth, and fossils of no Post-Cambrian non-vertebrate chordates are known aside from the
5507:
5440:
5417:
3960:
3905:
3850:
3746:
3681:
3614:
3558:
3501:
3439:
3361:
3314:
3259:
3246:; Han, J.; et al. (January 2003). "Head and backbone of the Early Cambrian vertebrate
3138:
3072:
2972:
2657:
2584:
2524:
2467:
2410:
2354:
2240:
2194:
2131:
2052:
2005:
1732:
2181:
MartĂn-DurĂĄn, JosĂ© M.; Passamaneck, Yale J.; Martindale, Mark Q.; Hejnol, Andreas (2016).
8:
5517:
3546:
3302:
3243:
2834:
2040:
1003:
946:
4059:
4016:
Philosophical
Transactions of the Royal Society of London. Series B, Biological Sciences
3964:
3909:
3854:
3750:
3685:
3618:
3562:
3505:
3443:
3365:
3318:
3263:
3142:
3076:
2976:
2661:
2588:
2528:
2471:
2414:
2358:
2198:
2135:
2056:
2009:
4649:
4201:
4179:
4167:
4142:
4036:
4011:
3983:
3948:
3929:
3874:
3819:
3775:
3734:
3710:
3669:
3582:
3522:
3489:
3462:
3427:
3412:
3377:
3330:
3283:
3167:
3126:
3101:
3060:
2941:
2807:
2780:
2729:
2702:
2626:
2556:
2436:
2378:
2218:
2182:
2163:
2100:
2076:
2021:
707:
536:
143:
5422:
3799:
1649:
5071:
4991:
4341:
4211:
4174:
4162:
4152:
4086:
4041:
3988:
3933:
3921:
3866:
3780:
3762:
3715:
3697:
3650:
3642:
3637:
3574:
3527:
3467:
3275:
3172:
3154:
3106:
3088:
2990:
2933:
2899:
2889:
2880:, Special publications, vol. 286, London: Geological Society, pp. 157â179,
2854:
2812:
2759:
2734:
2683:
2675:
2630:
2618:
2610:
2605:
2548:
2540:
2485:
2428:
2370:
2301:
2260:
2210:
2155:
2147:
2068:
2025:
1179:
604:
3823:
2945:
2440:
2382:
2324:
2183:"The developmental basis for the recurrent evolution of deuterostomy and protostomy"
4126:
4031:
4023:
3978:
3968:
3913:
3878:
3858:
3811:
3770:
3754:
3705:
3689:
3632:
3622:
3566:
3517:
3509:
3490:"Lack of support for Deuterostomia prompts reinterpretation of the first Bilateria"
3457:
3447:
3408:
3381:
3369:
3334:
3322:
3305:& Zhang, X.-L. (November 1999). "Lower Cambrian vertebrates from south China".
3287:
3267:
3210:
3162:
3146:
3096:
3080:
2980:
2925:
2881:
2846:
2802:
2792:
2724:
2714:
2665:
2600:
2592:
2560:
2532:
2475:
2418:
2362:
2344:
2291:
2252:
2202:
2139:
2060:
2013:
1041:
337:
4345:
4065:
3586:
2167:
2080:
1465:
5352:
5258:
5109:
5012:
4869:
4402:
3452:
2961:"Cambrian stem-group ambulacrarians and the nature of the ancestral deuterostome"
2280:"The Phylogenetic Position of Entoprocta, Ectoprocta, Phoronida, and Brachiopoda"
2222:
1797:
1233:
1183:
851:
836:
807:
803:
621:
525:
465:
261:
1671:
1377:
5017:
4995:
4813:
4725:
4703:
4666:
4157:
3693:
3150:
3084:
2456:"A New Spiralian Phylogeny Places the Enigmatic Arrow Worms among Gnathiferans"
1976:
1702:
1697:
1645:
1353:
1323:
1113:
1057:, and as such their true affinity remains uncertain. Another Ediacaran fossil,
1007:
667:
577:
576:
which was later combined with other protostome animals to form the superphylum
253:
3893:
3838:
3815:
3800:"Higher-level metazoan relationships: recent progress and remaining questions"
3627:
3602:
3214:
2985:
2960:
2670:
2645:
2596:
2480:
2455:
2423:
2398:
2256:
2017:
1049:
period. While these may in fact be tunicates, others have interpreted them as
5501:
5393:
5162:
5066:
4565:
3766:
3701:
3646:
3158:
3092:
3021:"Polypenartige Organismen aus dem Jung-PrÀkambrium (Nama-Gruppe) von Namibia"
2903:
2679:
2614:
2544:
2515:
phylogeny reveals monophyletic chordates and the new phylum Xenoturbellida".
2264:
2151:
1777:
1728:
1715:
1632:
1578:
1448:
1373:
1310:
1187:
1124:
1107:
840:
728:
720:
660:
396:
301:
277:
70:
2850:
2797:
2399:"Chaetognath phylogenomics: a protostome with deuterostome-like development"
2366:
2206:
1502:
1327:
5268:
5263:
5151:
4893:
4779:
4682:
4661:
4531:
4482:
4438:
4419:
4045:
4027:
3992:
3973:
3925:
3870:
3784:
3719:
3654:
3578:
3531:
3513:
3471:
3279:
3176:
3110:
3049:. Moscow, Paleontological Institute, Russian Academy of Sciences, p. 90-91.
3007:
2994:
2937:
2858:
2816:
2738:
2719:
2687:
2622:
2552:
2489:
2432:
2305:
2296:
2279:
2214:
2159:
2072:
1975:
Deuterostomia dissolves, with Chordata and Protostomia grouped together as
1666:
1622:
1438:
1340:
1237:
1101:
967:, one of the five major lineages of animals, is split into two groups; the
903:
787:
747:
701:
695:
674:
581:
573:
565:
529:
473:
412:
282:
272:
267:
224:
3839:"New deep-sea species of Xenoturbella and the position of Xenacoelomorpha"
2374:
5432:
5402:
5229:
5200:
5186:
5176:
4761:
4577:
4490:
4272:
2239:
Eernisse, Douglas J.; Albert, James S.; Anderson, Frank E. (1992-09-01).
1889:
1865:
1755:
1518:
1151:
1138:
1117:
from the Chengjiang biota, are the earliest bodyfossils of fish, whereas
1065:
1026:
919:
870:
862:
461:
454:
427:
416:
328:
245:
183:
45:
20:
3917:
3862:
3758:
3570:
3271:
2871:
2536:
2143:
2064:
1267:
5326:
5311:
5306:
5287:
5282:
5210:
5157:
5127:
5119:
5061:
4935:
4927:
4910:
4690:
4602:
4555:
4495:
4457:
4431:
4277:
4262:
3347:
3300:
2929:
2503:
Acorn worm genome reveals gill origins of human pharynx | Berkeley News
2180:
1926:
1558:
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1280:
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989:
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957:
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818:
781:
759:
713:
647:
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557:
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514:
509:
485:
481:
469:
442:
434:
408:
321:
285:
256:
248:
90:
55:
4078:
1781:
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5205:
5194:
5143:
5138:
5090:
5002:
4789:
4784:
4746:
4595:
4587:
4516:
4468:
4389:
4376:
4357:
4352:
4311:
3373:
3348:
Shu, D.-G.; Conway Morris, S. & Zhang, X.-L. (November 1996). "A
2885:
1885:
1545:
1485:
1461:
1297:
1200:
1094:
1070:
1050:
1046:
999:
973:
964:
877:
776:
756:
734:
569:
532:
489:
477:
209:
196:
160:
107:
95:
5364:
3946:
2703:"Developmental and evolutionary origins of the pharyngeal apparatus"
1909:
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5316:
5300:
5223:
5215:
5170:
5133:
5114:
5103:
5085:
5079:
4940:
4881:
4857:
4754:
4635:
4524:
4408:
4299:
4290:
4196:
3058:
1905:
1263:
1220:
1128:
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1030:
907:
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501:
420:
404:
240:
85:
80:
65:
60:
50:
39:
35:
3326:
3241:
2872:
Fedonkin, M.A.; Simonetta, A; Ivantsov, A.Y. (2007), "New data on
1522:
5276:
5252:
5244:
5181:
5095:
4915:
4902:
4835:
4826:
4771:
4696:
3735:"Acoelomorph flatworms are deuterostomes related to Xenoturbella"
2832:
1995:
1155:
1089:
1059:
1014:
889:
881:
769:
653:
561:
505:
497:
493:
100:
75:
5445:
3891:
2781:"Cambrian problematica and the diversification of deuterostomes"
1932:
802:
There is a possibility that Ambulacraria is the sister clade to
5321:
5055:
4255:
4242:
4113:
4110:
1119:
1092:), are quite common from the start of Stage 3 of the Cambrian,
1054:
866:
828:
450:
438:
400:
324:
170:
3836:
3008:
Connexins evolved after early chordates lost innexin diversity
2513:
1357:
352:
915:
911:
433:
In deuterostomes, the developing embryo's first opening (the
424:
392:
346:
3732:
2453:
941:
digestive tract develops in the middle, connecting the two.
3947:
Paschalia Kapli; Maximilian J. Telford (11 December 2020).
3667:
3425:
2395:
1562:
1013:
The last common ancestor of the deuterostomes had lost all
388:
379:
373:
367:
355:
3797:
3599:
2573:
987:
appeared, and hence well before the start of the Cambrian
343:
3487:
3190:
Bengtson, S. (2004). Lipps, J.H.; Waggoner, B.M. (eds.).
3047:
Enigmatic Organisms in Phylogeny and Evolution. Abstracts
895:
A hollow nerve cord is found in all chordates, including
3123:
2321:
5045:
3394:
2238:
839:, and also occurs in certain protostomes, such as the
831:
first develops into a hollow ball of cells, called a
468:
development is more varied. Protostomia includes the
361:
340:
1123:, discovered much earlier but from the Mid Cambrian
1105:. Two other Cambrian Stage 3 (521-514 mya) species,
1002:
Period (circa 635-539 Mya, around the end of global
364:
349:
2833:Erwin, Douglas H.; Eric H. Davidson (1 July 2002).
1127:, is now regarded as a primitive chordate. The Mid
1073:) period is another notable stem group echinoderm.
521:Deuterostomia and Protostomia, together with their
370:
358:
3894:"Xenacoelomorpha is the sister group to Nephrozoa"
3352:-like chordate from the Lower Cambrian of China".
3196:in NeoproterozoicâCambrian Biological Revolutions"
2323:
3399:, a Middle Cambrian rhabdopleurid hemichordate".
2915:
2646:"Zoology: Worming into the Origin of Bilaterians"
430:is also thought to be a member of Deuterostomia.
5499:
2958:
2778:
2700:
865:of the developed gut that pinch off to form the
2909:
2096:"This Prehistoric Human Ancestor Was All Mouth"
1063:, may represent the earliest echinoderm, while
884:, which is also found in some primitive fossil
120:
3544:
2234:
2232:
2121:
2038:
945:forms four weeks later, temporarily forming a
929:
4094:
3039:
2828:
2826:
1076:Fossils of one major deuterostome group, the
464:, animals that develop mouth first and whose
445:as well. The deuterostomes are also known as
3189:
3183:
2865:
2637:
2567:
2317:
2315:
556:Initially, Deuterostomia included the phyla
551:
19:For the cell biology protein structure, see
4481:
4009:
3395:Bengtson, S.; Urbanek, A. (October 2007). "
2507:
2229:
2115:
2087:
936:Embryological origins of the mouth and anus
376:
305:
4455:
4101:
4087:
2823:
2271:
813:
399:. Deuterostomia is further divided into 4
4648:
4035:
3982:
3972:
3774:
3709:
3636:
3626:
3521:
3461:
3451:
3166:
3100:
3018:
2984:
2806:
2796:
2751:
2728:
2718:
2669:
2604:
2479:
2422:
2312:
2295:
827:In both deuterostomes and protostomes, a
4515:
4439:Hemichordata (acorn worms and relatives)
4387:
3483:
3481:
2878:The Rise and Fall of the Ediacaran Biota
2643:
956:
952:
817:
4240:
4108:
2277:
5500:
4433:Echinodermata (starfish and relatives)
4403:Xenacoelomorpha (acoels and relatives)
3428:"The Global Diversity of Hemichordata"
2952:
5369:
5368:
4374:
4082:
3478:
2835:"The last common bilaterian ancestor"
2752:Valentine, James W. (June 18, 2004).
4704:Acanthocephala (thorny-headed worms)
4010:Swalla, B. J.; Smith, A. B. (2008).
3804:Organisms, Diversity & Evolution
2093:
387:), typically characterized by their
2322:Brusca, R.C.; Brusca, G.J. (1990).
2284:Integrative and Comparative Biology
13:
4738:
4544:
4301:Cnidaria (jellyfish and relatives)
4003:
3413:10.1111/j.1502-3931.1986.tb00743.x
2644:Marlétaz, Ferdinand (2019-06-17).
2094:Wade, Nicholas (30 January 2017).
603:This is the generally agreed upon
116:
14:
5534:
4060:Introduction to the Deuterostomia
4053:
2701:Graham, A; Richardson, J (2012).
5345:Phyla with â„1000 extant species
1931:
1908:
1888:
1780:
1731:
1701:
1670:
1648:
1581:
1561:
1521:
1501:
1464:
1376:
1356:
1326:
1296:
1266:
1236:
1020:
512:, etc.), as well as the extinct
460:Deuterostomia's sister clade is
336:
147:
43:
16:Superphylum of bilateral animals
3940:
3885:
3830:
3791:
3726:
3661:
3593:
3538:
3419:
3388:
3341:
3294:
3235:
3117:
3052:
3012:
3001:
2772:
2758:. University of Chicago Press.
2745:
2694:
2496:
2447:
2389:
998:during the earlier part of the
599:List of bilateral animal orders
4532:Nematomorpha (horsehair worms)
3203:Paleontological Society Papers
2338:
2330:. Sinauer Associates. p.
2187:Nature Ecology & Evolution
2174:
2032:
1989:
1080:(whose modern members include
546:
1:
4375:
3019:Hahn, G; Pflug, H. D (1985).
1982:
914:, homologous to the chordate
5144:Echinozoans inc. sea urchins
3453:10.1371/journal.pone.0162564
1173:
1099:starting with forms such as
922:having secondarily lost it.
7:
5513:Ediacaran first appearances
5201:Chelicerates inc. arachnids
4941:Phoronida (horseshoe worms)
4756:Platyhelminthes (flatworms)
4667:Gnathostomulida (jaw worms)
4556:Loricifera (corset animals)
930:Formation of mouth and anus
846:Most deuterostomes display
592:
138:Diversity of deuterostomes
10:
5539:
5523:Taxa named by Karl Grobben
5091:Medusozoans inc. jellyfish
4837:Annelida (segmented worms)
4662:Chaetognatha (arrow worms)
4578:Onychophora (velvet worms)
3694:10.1038/s41598-017-01608-4
3151:10.1038/s41467-020-14920-x
3085:10.1038/s41598-021-83452-1
1137:has been interpreted as a
933:
611:Superphylum Deuterostomia
596:
18:
5377:
5341:
5139:Asterozoans inc. starfish
5041:
5034:
4985:
4936:Brachiopoda (lamp shells)
4926:
4917:Ectoprocta (moss animals)
4901:
4892:
4877:
4868:
4853:
4846:
4822:
4811:
4770:
4762:Gastrotricha (hairybacks)
4745:
4734:
4723:
4675:
4657:
4644:
4633:
4586:
4573:
4564:
4551:
4540:
4508:
4491:Kinorhyncha (mud dragons)
4477:
4466:
4451:
4427:
4418:
4398:
4383:
4370:
4335:
4310:
4286:
4271:
4263:Ctenophora (comb jellies)
4251:
4236:
4120:
3816:10.1007/s13127-011-0044-4
3638:21.11116/0000-0004-DC4B-1
3628:10.1016/j.cub.2019.04.009
3215:10.1017/S1089332600002345
2986:10.1016/j.cub.2023.04.048
2779:Smith, Andrew B. (2012).
2768:– via Google Books.
2671:10.1016/j.cub.2019.05.006
2606:21.11116/0000-0004-DC4B-1
2597:10.1016/j.cub.2019.04.009
2481:10.1016/j.cub.2018.11.042
2424:10.1016/j.cub.2006.07.016
2278:Nielsen, C. (July 2002).
2018:10.1134/S0031030112010042
1923:
1903:
1883:
1876:
1869:
1795:
1775:
1768:
1753:
1746:
1726:
1719:
1694:
1663:
1643:
1636:
1626:
1576:
1556:
1549:
1516:
1496:
1489:
1479:
1459:
1452:
1442:
1371:
1351:
1344:
1321:
1314:
1291:
1284:
1261:
1254:
1231:
1224:
1214:
1204:
1069:from the early Cambrian (
910:have a thickening of the
869:. This process is called
552:History of classification
237:
232:
144:Scientific classification
142:
137:
115:
34:Temporal range: Earliest
30:
4697:Rotifera (wheel animals)
4496:Priapulida (penis worms)
587:horizontal gene transfer
4911:Entoprocta or Kamptozoa
4883:Nemertea (ribbon worms)
4604:Tardigrada (waterbears)
4597:Arthropoda (arthropods)
4073:EncyclopĂŠdia Britannica
2851:10.1242/dev.129.13.3021
2798:10.1186/1741-7007-10-79
2367:10.1126/science.7886451
2257:10.1093/sysbio/41.3.305
2207:10.1038/s41559-016-0005
1998:Paleontological Journal
1162:, trace fossils of the
1141:hemichordate., whereas
1097: million years ago
992: million years ago
814:Notable characteristics
528:, constitute the large
437:) becomes the anus and
5086:Anthozoans inc. corals
4785:Dicyemida or Rhombozoa
4028:10.1098/rstb.2007.2246
3974:10.1126/sciadv.abc5162
3514:10.1126/sciadv.abe2741
3194:Early Skeletal Fossils
2755:On the Origin of Phyla
2720:10.1186/2041-9139-3-24
1134:Rhabdotubus johanssoni
961:
848:indeterminate cleavage
824:
772:and extinct relatives)
677:(bony fish - includes
607:of the deuterostomes:
132:
5480:Paleobiology Database
5216:Hexapods inc. insects
4526:Nematoda (roundworms)
3131:Nature Communications
960:
953:Origins and evolution
857:In deuterostomes the
821:
806:, and could form the
423:fossils. The extinct
397:embryonic development
131:
4409:Chordata (chordates)
3613:(11): 1818â1826.e6.
3547:Morris, Simon Conway
2971:(12): 2359â2366.e2.
2583:(11): 1818â1826.e6.
2297:10.1093/icb/42.3.685
2041:Morris, Simon Conway
670:(cartilaginous fish)
535:, i.e. animals with
4859:Mollusca (molluscs)
4344:of the animal root
4022:(1496): 1557â1568.
3965:2020SciA....6.5162K
3918:10.1038/nature16520
3910:2016Natur.530...89C
3863:10.1038/nature16545
3855:2016Natur.530...94R
3759:10.1038/nature09676
3751:2011Natur.470..255P
3686:2017NatSR...7.1847R
3619:2019CBio...29E1818P
3571:10.1038/nature21072
3563:2017Natur.542..228H
3506:2021SciA....7.2741K
3444:2016PLoSO..1162564T
3366:1996Natur.384..157S
3319:1999Natur.402...42S
3272:10.1038/nature01264
3264:2003Natur.421..526S
3143:2020NatCo..11.1286Z
3077:2021NatSR..11.4121C
2977:2023CBio...33E2359L
2662:2019CBio...29.R577M
2589:2019CBio...29E1818P
2537:10.1038/nature05241
2529:2006Natur.444...85B
2472:2019CBio...29E.312M
2415:2006CBio...16.R577M
2359:1995Sci...267.1641H
2353:(5204): 1641â1643.
2199:2016NatEE...1....5M
2144:10.1038/nature21072
2136:2017Natur.542..228H
2065:10.1038/nature21072
2057:2017Natur.542..228H
2010:2012PalJ...46....1F
1004:Marinoan glaciation
391:forming before the
5072:Calcareous sponges
4257:Porifera (sponges)
3674:Scientific Reports
3065:Scientific Reports
2930:10.1002/bies.20507
2245:Systematic Biology
2101:The New York Times
1186:, fall within the
1166:colonial tunicate
1160:Paleobranchiostoma
962:
825:
708:Planctosphaeroidea
537:bilateral symmetry
415:, and the extinct
133:
5495:
5494:
5371:Taxon identifiers
5362:
5361:
5337:
5336:
5030:
5029:
5026:
5025:
4981:
4980:
4977:
4976:
4973:
4972:
4969:
4968:
4965:
4964:
4961:
4960:
4957:
4956:
4953:
4952:
4949:
4948:
4807:
4806:
4803:
4802:
4719:
4718:
4715:
4714:
4629:
4628:
4625:
4624:
4621:
4620:
4617:
4616:
4613:
4612:
4504:
4503:
4447:
4446:
4366:
4365:
4331:
4330:
4327:
4326:
3745:(7333): 255â258.
3557:(7640): 228â231.
3360:(6605): 157â158.
3303:Conway Morris, S.
3258:(6922): 526â529.
3244:Conway Morris, S.
2924:(12): 1161â1166.
2895:978-1-86239-233-5
2845:(13): 3021â3032.
2765:978-0-226-84548-7
2656:(12): R577âR579.
2466:(2): 312â318.e3.
2409:(15): R577âR578.
2130:(7640): 228â231.
2051:(7640): 228â231.
1971:
1970:
1962:
1961:
1953:
1952:
1944:
1943:
1855:
1854:
1845:
1844:
1835:
1834:
1826:
1825:
1817:
1816:
1808:
1807:
1683:
1682:
1612:
1611:
1603:
1602:
1594:
1593:
1534:
1533:
1428:
1427:
1419:
1418:
1409:
1408:
1399:
1398:
1389:
1388:
1180:phylogenetic tree
1033:-like organisms
453:develops through
295:
294:
228:
5530:
5488:
5487:
5475:
5474:
5462:
5461:
5449:
5448:
5436:
5435:
5426:
5425:
5413:
5412:
5411:
5398:
5397:
5396:
5366:
5365:
5043:
5042:
4999:
4899:
4898:
4875:
4874:
4851:
4850:
4820:
4819:
4816:
4743:
4742:
4736:
4735:
4732:
4731:
4728:
4655:
4654:
4646:
4645:
4642:
4641:
4638:
4571:
4570:
4549:
4548:
4542:
4541:
4513:
4512:
4479:
4478:
4475:
4474:
4471:
4464:
4463:
4460:
4453:
4452:
4425:
4424:
4396:
4395:
4392:
4385:
4384:
4372:
4371:
4349:
4284:
4283:
4249:
4248:
4245:
4238:
4237:
4149:
4129:
4103:
4096:
4089:
4080:
4079:
4049:
4039:
3997:
3996:
3986:
3976:
3959:(50): eabc5162.
3953:Science Advances
3944:
3938:
3937:
3889:
3883:
3882:
3834:
3828:
3827:
3795:
3789:
3788:
3778:
3730:
3724:
3723:
3713:
3665:
3659:
3658:
3640:
3630:
3597:
3591:
3590:
3542:
3536:
3535:
3525:
3500:(12): eabe2741.
3494:Science Advances
3485:
3476:
3475:
3465:
3455:
3438:(10): e0162564.
3423:
3417:
3416:
3392:
3386:
3385:
3374:10.1038/384157a0
3345:
3339:
3338:
3298:
3292:
3291:
3239:
3233:
3232:
3230:
3229:
3223:
3217:. Archived from
3200:
3187:
3181:
3180:
3170:
3121:
3115:
3114:
3104:
3056:
3050:
3043:
3037:
3036:
3034:
3032:
3016:
3010:
3005:
2999:
2998:
2988:
2956:
2950:
2949:
2913:
2907:
2906:
2886:10.1144/SP286.12
2869:
2863:
2862:
2830:
2821:
2820:
2810:
2800:
2776:
2770:
2769:
2749:
2743:
2742:
2732:
2722:
2698:
2692:
2691:
2673:
2641:
2635:
2634:
2608:
2571:
2565:
2564:
2511:
2505:
2500:
2494:
2493:
2483:
2451:
2445:
2444:
2426:
2393:
2387:
2386:
2342:
2336:
2335:
2329:
2319:
2310:
2309:
2299:
2275:
2269:
2268:
2236:
2227:
2226:
2178:
2172:
2171:
2119:
2113:
2112:
2110:
2108:
2091:
2085:
2084:
2036:
2030:
2029:
1993:
1935:
1912:
1892:
1879:
1878:
1872:
1871:
1784:
1771:
1770:
1749:
1748:
1735:
1722:
1721:
1705:
1674:
1652:
1639:
1638:
1629:
1628:
1585:
1565:
1552:
1551:
1525:
1505:
1492:
1491:
1482:
1481:
1468:
1455:
1454:
1445:
1444:
1380:
1360:
1347:
1346:
1330:
1317:
1316:
1300:
1287:
1286:
1270:
1257:
1256:
1240:
1227:
1226:
1217:
1216:
1207:
1206:
1197:
1196:
1098:
993:
977:
976:million year old
449:, because their
447:enterocoelomates
386:
385:
382:
381:
378:
375:
372:
369:
366:
363:
360:
357:
354:
351:
348:
345:
342:
319:
316:
313:
310:
307:
223:
152:
151:
119:
105:
28:
27:
5538:
5537:
5533:
5532:
5531:
5529:
5528:
5527:
5498:
5497:
5496:
5491:
5483:
5478:
5470:
5465:
5457:
5452:
5444:
5439:
5431:
5429:
5421:
5416:
5407:
5406:
5401:
5392:
5391:
5386:
5373:
5363:
5358:
5333:
5259:Phylactolaemata
5047:
5037:
5036:
5022:
5013:Xenambulacraria
4989:
4945:
4922:
4888:
4870:Kryptotrochozoa
4864:
4842:
4812:
4799:
4790:Monoblastozoa (
4766:
4724:
4711:
4683:Micrognathozoa
4671:
4634:
4609:
4582:
4560:
4536:
4500:
4467:
4456:
4443:
4414:
4388:
4379:
4362:
4339:
4323:
4306:
4275:
4267:
4241:
4232:
4231:
4147:
4125:
4116:
4107:
4056:
4006:
4004:Further reading
4001:
4000:
3945:
3941:
3890:
3886:
3835:
3831:
3796:
3792:
3731:
3727:
3666:
3662:
3607:Current Biology
3598:
3594:
3543:
3539:
3486:
3479:
3424:
3420:
3393:
3389:
3346:
3342:
3313:(6757): 42â46.
3299:
3295:
3240:
3236:
3227:
3225:
3221:
3198:
3188:
3184:
3122:
3118:
3057:
3053:
3044:
3040:
3030:
3028:
3017:
3013:
3006:
3002:
2965:Current Biology
2957:
2953:
2914:
2910:
2896:
2870:
2866:
2831:
2824:
2777:
2773:
2766:
2750:
2746:
2699:
2695:
2650:Current Biology
2642:
2638:
2577:Current Biology
2572:
2568:
2523:(7115): 85â88.
2512:
2508:
2501:
2497:
2460:Current Biology
2452:
2448:
2403:Current Biology
2394:
2390:
2343:
2339:
2320:
2313:
2276:
2272:
2237:
2230:
2179:
2175:
2120:
2116:
2106:
2104:
2092:
2088:
2037:
2033:
1994:
1990:
1985:
1972:
1963:
1954:
1945:
1856:
1846:
1836:
1827:
1818:
1809:
1798:Torquaratoridae
1684:
1613:
1604:
1595:
1535:
1429:
1420:
1410:
1400:
1390:
1234:Cephalochordata
1193:
1184:Torquaratoridae
1176:
1093:
1023:
988:
972:
955:
938:
932:
852:identical twins
837:radial cleavage
816:
808:Xenambulacraria
804:Xenacoelomorpha
790:(sea cucumbers)
750:(brittle stars)
622:Cephalochordata
601:
595:
554:
549:
526:Xenacoelomorpha
466:digestive tract
339:
335:
317:
314:
311:
308:
262:Xenacoelomorpha
222:
146:
130:
117:
111:
110:record, 557 Ma)
104:
103:
98:
93:
88:
83:
78:
73:
68:
63:
58:
53:
48:
42:
32:
24:
17:
12:
11:
5:
5536:
5526:
5525:
5520:
5515:
5510:
5493:
5492:
5490:
5489:
5476:
5463:
5450:
5437:
5427:
5414:
5399:
5383:
5381:
5375:
5374:
5360:
5359:
5357:
5356:
5349:
5342:
5339:
5338:
5335:
5334:
5332:
5331:
5330:
5329:
5324:
5319:
5314:
5309:
5297:
5296:
5295:
5290:
5285:
5273:
5272:
5271:
5266:
5261:
5249:
5248:
5247:
5242:
5237:
5232:
5224:Platyhelminths
5220:
5219:
5218:
5213:
5208:
5203:
5191:
5190:
5189:
5184:
5179:
5167:
5166:
5165:
5160:
5148:
5147:
5146:
5141:
5136:
5124:
5123:
5122:
5117:
5112:
5100:
5099:
5098:
5093:
5088:
5076:
5075:
5074:
5069:
5064:
5051:
5049:
5039:
5038:
5035:
5032:
5031:
5028:
5027:
5024:
5023:
5021:
5020:
5018:Centroneuralia
5015:
5010:
5005:
5000:
4986:
4983:
4982:
4979:
4978:
4975:
4974:
4971:
4970:
4967:
4966:
4963:
4962:
4959:
4958:
4955:
4954:
4951:
4950:
4947:
4946:
4944:
4943:
4938:
4932:
4930:
4924:
4923:
4921:
4920:
4913:
4907:
4905:
4896:
4890:
4889:
4887:
4886:
4878:
4872:
4866:
4865:
4863:
4862:
4854:
4848:
4844:
4843:
4841:
4840:
4833:
4823:
4817:
4814:Lophotrochozoa
4809:
4808:
4805:
4804:
4801:
4800:
4798:
4797:
4787:
4782:
4776:
4774:
4768:
4767:
4765:
4764:
4759:
4751:
4749:
4740:
4729:
4726:Platytrochozoa
4721:
4720:
4717:
4716:
4713:
4712:
4710:
4709:
4708:
4707:
4700:
4688:
4685:(Limnognathia)
4679:
4677:
4673:
4672:
4670:
4669:
4664:
4658:
4652:
4639:
4631:
4630:
4627:
4626:
4623:
4622:
4619:
4618:
4615:
4614:
4611:
4610:
4608:
4607:
4600:
4592:
4590:
4584:
4583:
4581:
4580:
4574:
4568:
4562:
4561:
4559:
4558:
4552:
4546:
4538:
4537:
4535:
4534:
4529:
4521:
4519:
4510:
4506:
4505:
4502:
4501:
4499:
4498:
4493:
4487:
4485:
4472:
4461:
4449:
4448:
4445:
4444:
4442:
4441:
4436:
4428:
4422:
4416:
4415:
4413:
4412:
4405:
4399:
4393:
4381:
4380:
4368:
4367:
4364:
4363:
4361:
4360:
4355:
4350:
4336:
4333:
4332:
4329:
4328:
4325:
4324:
4322:
4321:
4317:
4315:
4314:(Triploblasts)
4308:
4307:
4305:
4304:
4297:
4295:and relatives)
4287:
4281:
4269:
4268:
4266:
4265:
4260:
4252:
4246:
4234:
4233:
4230:
4229:
4228:
4227:
4226:
4225:
4222:Mesomycetozoea
4219:
4214:
4204:
4194:
4193:
4192:
4187:
4182:
4177:
4172:
4171:
4170:
4158:Diaphoretickes
4155:
4150:
4145:
4140:
4135:
4130:
4122:
4121:
4118:
4117:
4106:
4105:
4098:
4091:
4083:
4077:
4076:
4063:
4055:
4054:External links
4052:
4051:
4050:
4005:
4002:
3999:
3998:
3939:
3884:
3829:
3810:(2): 151â172.
3790:
3725:
3660:
3592:
3537:
3477:
3418:
3407:(4): 293â308.
3387:
3340:
3293:
3234:
3182:
3116:
3051:
3038:
3025:Pascal-Francis
3011:
3000:
2951:
2908:
2894:
2864:
2822:
2771:
2764:
2744:
2693:
2636:
2566:
2506:
2495:
2446:
2388:
2337:
2311:
2290:(3): 685â691.
2270:
2251:(3): 305â330.
2228:
2173:
2114:
2086:
2031:
1987:
1986:
1984:
1981:
1977:Centroneuralia
1969:
1968:
1965:
1964:
1960:
1959:
1956:
1955:
1951:
1950:
1947:
1946:
1942:
1941:
1938:
1937:
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1742:
1741:
1738:
1737:
1725:
1720:
1718:
1712:
1711:
1708:
1707:
1698:Planctosphaera
1693:
1690:
1689:
1686:
1685:
1681:
1680:
1677:
1676:
1662:
1659:
1658:
1655:
1654:
1646:Cephalodiscida
1642:
1637:
1635:
1627:
1625:
1619:
1618:
1615:
1614:
1610:
1609:
1606:
1605:
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1430:
1426:
1425:
1422:
1421:
1417:
1416:
1412:
1411:
1407:
1406:
1402:
1401:
1397:
1396:
1392:
1391:
1387:
1386:
1383:
1382:
1370:
1367:
1366:
1363:
1362:
1354:Actinopterygii
1350:
1345:
1343:
1337:
1336:
1333:
1332:
1324:Chondrichthyes
1320:
1315:
1313:
1307:
1306:
1303:
1302:
1290:
1285:
1283:
1277:
1276:
1273:
1272:
1260:
1255:
1253:
1247:
1246:
1243:
1242:
1230:
1225:
1223:
1215:
1213:
1205:
1203:
1195:
1175:
1172:
1144:Spartobranchus
1114:Myllokunmingia
1022:
1019:
1008:Neoproterozoic
954:
951:
934:Main article:
931:
928:
815:
812:
800:
799:
798:
797:
796:
795:
794:
793:
792:
791:
785:
773:
762:
753:
752:
751:
744:
725:
724:
723:
717:
710:
690:
689:
688:
687:
686:
685:
684:
683:
682:
671:
668:Chondrichthyes
657:
656:(jawless fish)
644:
629:
594:
591:
578:Lophotrochozoa
553:
550:
548:
545:
293:
292:
291:
290:
289:
288:
280:
275:
265:
259:
254:Vetulocystidae
251:
243:
235:
234:
230:
229:
217:
213:
212:
207:
200:
199:
194:
187:
186:
181:
174:
173:
168:
164:
163:
158:
154:
153:
140:
139:
135:
134:
113:
112:
99:
94:
89:
84:
79:
74:
69:
64:
59:
54:
49:
44:
33:
15:
9:
6:
4:
3:
2:
5535:
5524:
5521:
5519:
5516:
5514:
5511:
5509:
5506:
5505:
5503:
5486:
5481:
5477:
5473:
5468:
5464:
5460:
5455:
5451:
5447:
5442:
5438:
5434:
5428:
5424:
5423:Deuterostomia
5419:
5415:
5410:
5409:Deuterostomia
5404:
5400:
5395:
5389:
5385:
5384:
5382:
5380:
5379:Deuterostomia
5376:
5372:
5367:
5354:
5353:dubious phyla
5350:
5348:
5344:
5343:
5340:
5328:
5325:
5323:
5320:
5318:
5315:
5313:
5310:
5308:
5305:
5304:
5303:
5302:
5298:
5294:
5291:
5289:
5286:
5284:
5281:
5280:
5279:
5278:
5274:
5270:
5267:
5265:
5262:
5260:
5257:
5256:
5255:
5254:
5250:
5246:
5243:
5241:
5238:
5236:
5233:
5231:
5228:
5227:
5226:
5225:
5221:
5217:
5214:
5212:
5209:
5207:
5204:
5202:
5199:
5198:
5197:
5196:
5192:
5188:
5185:
5183:
5180:
5178:
5175:
5174:
5173:
5172:
5168:
5164:
5161:
5159:
5156:
5155:
5154:
5153:
5152:Hemichordates
5149:
5145:
5142:
5140:
5137:
5135:
5132:
5131:
5130:
5129:
5125:
5121:
5118:
5116:
5113:
5111:
5108:
5107:
5106:
5105:
5101:
5097:
5094:
5092:
5089:
5087:
5084:
5083:
5082:
5081:
5077:
5073:
5070:
5068:
5067:Glass sponges
5065:
5063:
5060:
5059:
5058:
5057:
5053:
5052:
5050:
5044:
5040:
5033:
5019:
5016:
5014:
5011:
5009:
5008:Deuterostomia
5006:
5004:
5001:
4997:
4994:of Bilateria
4993:
4988:
4987:
4984:
4942:
4939:
4937:
4934:
4933:
4931:
4929:
4925:
4919:
4918:
4914:
4912:
4909:
4908:
4906:
4904:
4900:
4897:
4895:
4891:
4885:
4884:
4880:
4879:
4876:
4873:
4871:
4867:
4861:
4860:
4856:
4855:
4852:
4849:
4845:
4839:
4838:
4834:
4832:
4830:
4827:Cycliophora (
4825:
4824:
4821:
4818:
4815:
4810:
4795:
4793:
4788:
4786:
4783:
4781:
4778:
4777:
4775:
4773:
4769:
4763:
4760:
4758:
4757:
4753:
4752:
4750:
4748:
4744:
4741:
4737:
4733:
4730:
4727:
4722:
4706:
4705:
4701:
4699:
4698:
4694:
4693:
4692:
4689:
4687:
4686:
4681:
4680:
4678:
4674:
4668:
4665:
4663:
4660:
4659:
4656:
4653:
4651:
4647:
4643:
4640:
4637:
4632:
4606:
4605:
4601:
4599:
4598:
4594:
4593:
4591:
4589:
4585:
4579:
4576:
4575:
4572:
4569:
4567:
4566:Panarthropoda
4563:
4557:
4554:
4553:
4550:
4547:
4543:
4539:
4533:
4530:
4528:
4527:
4523:
4522:
4520:
4518:
4514:
4511:
4507:
4497:
4494:
4492:
4489:
4488:
4486:
4484:
4480:
4476:
4473:
4470:
4465:
4462:
4459:
4454:
4450:
4440:
4437:
4435:
4434:
4430:
4429:
4426:
4423:
4421:
4417:
4411:
4410:
4406:
4404:
4401:
4400:
4397:
4394:
4391:
4386:
4382:
4378:
4373:
4369:
4359:
4356:
4354:
4351:
4347:
4343:
4338:
4337:
4334:
4319:
4318:
4316:
4313:
4309:
4303:
4302:
4298:
4296:
4294:
4289:
4288:
4285:
4282:
4279:
4274:
4270:
4264:
4261:
4259:
4258:
4254:
4253:
4250:
4247:
4244:
4239:
4235:
4223:
4220:
4218:
4215:
4213:
4210:
4209:
4208:
4205:
4203:
4200:
4199:
4198:
4195:
4191:
4188:
4186:
4185:Stramenopiles
4183:
4181:
4178:
4176:
4173:
4169:
4166:
4165:
4164:
4161:
4160:
4159:
4156:
4154:
4151:
4148:(major groups
4146:
4144:
4141:
4139:
4136:
4134:
4131:
4128:
4124:
4123:
4119:
4115:
4112:
4104:
4099:
4097:
4092:
4090:
4085:
4084:
4081:
4075:
4074:
4069:
4068:
4067:Deuterostomia
4064:
4061:
4058:
4057:
4047:
4043:
4038:
4033:
4029:
4025:
4021:
4017:
4013:
4008:
4007:
3994:
3990:
3985:
3980:
3975:
3970:
3966:
3962:
3958:
3954:
3950:
3943:
3935:
3931:
3927:
3923:
3919:
3915:
3911:
3907:
3903:
3899:
3895:
3888:
3880:
3876:
3872:
3868:
3864:
3860:
3856:
3852:
3848:
3844:
3840:
3833:
3825:
3821:
3817:
3813:
3809:
3805:
3801:
3794:
3786:
3782:
3777:
3772:
3768:
3764:
3760:
3756:
3752:
3748:
3744:
3740:
3736:
3729:
3721:
3717:
3712:
3707:
3703:
3699:
3695:
3691:
3687:
3683:
3679:
3675:
3671:
3664:
3656:
3652:
3648:
3644:
3639:
3634:
3629:
3624:
3620:
3616:
3612:
3608:
3604:
3596:
3588:
3584:
3580:
3576:
3572:
3568:
3564:
3560:
3556:
3552:
3548:
3541:
3533:
3529:
3524:
3519:
3515:
3511:
3507:
3503:
3499:
3495:
3491:
3484:
3482:
3473:
3469:
3464:
3459:
3454:
3449:
3445:
3441:
3437:
3433:
3429:
3422:
3414:
3410:
3406:
3402:
3398:
3391:
3383:
3379:
3375:
3371:
3367:
3363:
3359:
3355:
3351:
3344:
3336:
3332:
3328:
3327:10.1038/46965
3324:
3320:
3316:
3312:
3308:
3304:
3297:
3289:
3285:
3281:
3277:
3273:
3269:
3265:
3261:
3257:
3253:
3249:
3248:Haikouichthys
3245:
3238:
3224:on 2008-10-03
3220:
3216:
3212:
3208:
3204:
3197:
3195:
3186:
3178:
3174:
3169:
3164:
3160:
3156:
3152:
3148:
3144:
3140:
3136:
3132:
3128:
3120:
3112:
3108:
3103:
3098:
3094:
3090:
3086:
3082:
3078:
3074:
3070:
3066:
3062:
3055:
3048:
3042:
3026:
3022:
3015:
3009:
3004:
2996:
2992:
2987:
2982:
2978:
2974:
2970:
2966:
2962:
2955:
2947:
2943:
2939:
2935:
2931:
2927:
2923:
2919:
2912:
2905:
2901:
2897:
2891:
2887:
2883:
2879:
2875:
2868:
2860:
2856:
2852:
2848:
2844:
2840:
2836:
2829:
2827:
2818:
2814:
2809:
2804:
2799:
2794:
2790:
2786:
2782:
2775:
2767:
2761:
2757:
2756:
2748:
2740:
2736:
2731:
2726:
2721:
2716:
2712:
2708:
2704:
2697:
2689:
2685:
2681:
2677:
2672:
2667:
2663:
2659:
2655:
2651:
2647:
2640:
2632:
2628:
2624:
2620:
2616:
2612:
2607:
2602:
2598:
2594:
2590:
2586:
2582:
2578:
2570:
2562:
2558:
2554:
2550:
2546:
2542:
2538:
2534:
2530:
2526:
2522:
2518:
2510:
2504:
2499:
2491:
2487:
2482:
2477:
2473:
2469:
2465:
2461:
2457:
2450:
2442:
2438:
2434:
2430:
2425:
2420:
2416:
2412:
2408:
2404:
2400:
2392:
2384:
2380:
2376:
2372:
2368:
2364:
2360:
2356:
2352:
2348:
2341:
2333:
2328:
2327:
2326:Invertebrates
2318:
2316:
2307:
2303:
2298:
2293:
2289:
2285:
2281:
2274:
2266:
2262:
2258:
2254:
2250:
2246:
2242:
2235:
2233:
2224:
2220:
2216:
2212:
2208:
2204:
2200:
2196:
2192:
2188:
2184:
2177:
2169:
2165:
2161:
2157:
2153:
2149:
2145:
2141:
2137:
2133:
2129:
2125:
2118:
2103:
2102:
2097:
2090:
2082:
2078:
2074:
2070:
2066:
2062:
2058:
2054:
2050:
2046:
2042:
2035:
2027:
2023:
2019:
2015:
2011:
2007:
2003:
1999:
1992:
1988:
1980:
1978:
1967:
1966:
1958:
1957:
1949:
1948:
1940:
1939:
1936:
1934:
1929:
1928:
1921:
1920:
1917:
1916:
1913:
1911:
1907:
1901:
1900:
1897:
1896:
1893:
1891:
1887:
1881:
1880:
1874:
1873:
1867:
1864:
1863:
1860:
1859:
1851:533 mya
1850:
1849:
1841:509 mya
1840:
1839:
1831:
1830:
1822:
1821:
1813:
1812:
1804:
1803:
1800:
1799:
1793:
1792:
1789:
1788:
1785:
1783:
1779:
1778:Ptychoderidae
1773:
1772:
1766:
1765:
1762:
1761:
1758:
1757:
1751:
1750:
1744:
1743:
1740:
1739:
1736:
1734:
1730:
1729:Harrimaniidae
1724:
1723:
1717:
1716:Enteropneusta
1714:
1713:
1710:
1709:
1706:
1704:
1700:
1699:
1692:
1691:
1688:
1687:
1679:
1678:
1675:
1673:
1669:
1668:
1661:
1660:
1657:
1656:
1653:
1651:
1647:
1641:
1640:
1634:
1633:Pterobranchia
1631:
1630:
1624:
1621:
1620:
1617:
1616:
1608:
1607:
1599:
1598:
1590:
1589:
1586:
1584:
1580:
1579:Holothuroidea
1574:
1573:
1570:
1569:
1566:
1564:
1560:
1554:
1553:
1547:
1544:
1543:
1540:488 mya
1539:
1538:
1530:
1529:
1526:
1524:
1520:
1514:
1513:
1510:
1509:
1506:
1504:
1500:
1494:
1493:
1487:
1484:
1483:
1477:
1476:
1473:
1472:
1469:
1467:
1463:
1457:
1456:
1450:
1449:Echinodermata
1447:
1446:
1440:
1437:
1436:
1433:
1432:
1424:
1423:
1415:518 mya
1414:
1413:
1405:440 mya
1404:
1403:
1395:425 mya
1394:
1393:
1385:
1384:
1381:
1379:
1375:
1374:Sarcopterygii
1369:
1368:
1365:
1364:
1361:
1359:
1355:
1349:
1348:
1342:
1339:
1338:
1335:
1334:
1331:
1329:
1325:
1319:
1318:
1312:
1311:Gnathostomata
1309:
1308:
1305:
1304:
1301:
1299:
1295:
1289:
1288:
1282:
1279:
1278:
1275:
1274:
1271:
1269:
1265:
1259:
1258:
1252:
1249:
1248:
1245:
1244:
1241:
1239:
1235:
1229:
1228:
1222:
1219:
1218:
1212:
1211:Deuterostomia
1209:
1208:
1202:
1199:
1198:
1194:
1191:
1189:
1188:Ptychoderidae
1185:
1181:
1171:
1169:
1165:
1161:
1157:
1153:
1148:
1146:
1145:
1140:
1136:
1135:
1130:
1126:
1125:Burgess Shale
1122:
1121:
1116:
1115:
1110:
1109:
1108:Haikouichthys
1104:
1103:
1096:
1091:
1087:
1083:
1079:
1074:
1072:
1068:
1067:
1062:
1061:
1056:
1052:
1048:
1044:
1043:
1038:
1037:
1032:
1028:
1021:Fossil record
1018:
1016:
1011:
1009:
1005:
1001:
997:
991:
986:
982:
981:
975:
970:
966:
959:
950:
948:
942:
937:
927:
923:
921:
917:
913:
909:
905:
904:hemichordates
900:
898:
893:
891:
887:
883:
879:
874:
872:
868:
864:
860:
855:
853:
849:
844:
842:
841:lophophorates
838:
834:
830:
820:
811:
809:
805:
789:
788:Holothuriodea
786:
784:(sea urchins)
783:
780:
779:
778:
774:
771:
767:
763:
761:
758:
754:
749:
745:
742:
738:
737:
736:
732:
731:
730:
729:Echinodermata
726:
722:
721:Pterobranchia
718:
716:(acorn worms)
715:
714:Enteropneusta
711:
709:
705:
704:
703:
699:
698:
697:
694:
691:
680:
676:
672:
669:
665:
664:
663:(jawed fish)
662:
661:Gnathostomata
658:
655:
651:
650:
649:
645:
642:
638:
637:
636:
633:
630:
627:
623:
619:
618:
617:
613:
612:
610:
609:
608:
606:
600:
590:
588:
583:
579:
575:
574:lophophorates
571:
567:
563:
559:
544:
542:
538:
534:
531:
527:
524:
519:
517:
516:
511:
507:
503:
499:
495:
491:
487:
483:
479:
475:
474:panarthropods
471:
467:
463:
458:
456:
452:
448:
444:
440:
436:
431:
429:
426:
422:
418:
414:
410:
409:Echinodermata
406:
402:
398:
394:
390:
384:
333:
332:Deuterostomia
330:
326:
323:
303:
299:
298:Deuterostomes
287:
284:
281:
279:
278:Echinodermata
276:
274:
271:
270:
269:
266:
263:
260:
258:
255:
252:
250:
247:
244:
242:
239:
238:
236:
231:
226:
221:
220:Deuterostomia
218:
216:Superphylum:
215:
214:
211:
208:
205:
202:
201:
198:
195:
192:
189:
188:
185:
182:
179:
176:
175:
172:
169:
166:
165:
162:
159:
156:
155:
150:
145:
141:
136:
114:
109:
102:
97:
92:
87:
82:
77:
72:
67:
62:
57:
52:
47:
41:
37:
31:Deuterostomes
29:
26:
22:
5378:
5351:Potentially
5346:
5299:
5275:
5269:Gymnolaemata
5264:Stenolaemata
5253:Ectoproctans
5251:
5222:
5193:
5169:
5163:Pterobranchs
5150:
5126:
5102:
5078:
5054:
5048:within phyla
5046:Major groups
5007:
4916:
4903:Bryozoa s.l.
4894:Lophophorata
4882:
4858:
4836:
4828:
4791:
4780:Orthonectida
4755:
4702:
4695:
4684:
4603:
4596:
4525:
4483:Scalidophora
4432:
4420:Ambulacraria
4407:
4320:(see belowâ)
4300:
4292:
4256:
4207:Opisthokonta
4071:
4066:
4019:
4015:
3956:
3952:
3942:
3904:(2): 89â93.
3901:
3897:
3887:
3849:(2): 94â97.
3846:
3842:
3832:
3807:
3803:
3793:
3742:
3738:
3728:
3677:
3673:
3663:
3610:
3606:
3595:
3554:
3550:
3540:
3497:
3493:
3435:
3431:
3421:
3404:
3400:
3396:
3390:
3357:
3353:
3349:
3343:
3310:
3306:
3301:Shu, D.-G.;
3296:
3255:
3251:
3247:
3242:Shu, D.-G.;
3237:
3226:. Retrieved
3219:the original
3206:
3202:
3193:
3185:
3134:
3130:
3119:
3068:
3064:
3054:
3046:
3041:
3029:. Retrieved
3024:
3014:
3003:
2968:
2964:
2954:
2921:
2917:
2911:
2877:
2873:
2867:
2842:
2838:
2788:
2784:
2774:
2754:
2747:
2710:
2706:
2696:
2653:
2649:
2639:
2580:
2576:
2569:
2520:
2516:
2509:
2498:
2463:
2459:
2449:
2406:
2402:
2391:
2350:
2346:
2340:
2325:
2287:
2283:
2273:
2248:
2244:
2190:
2186:
2176:
2127:
2123:
2117:
2105:. Retrieved
2099:
2089:
2048:
2044:
2034:
2001:
1997:
1991:
1973:
1930:(â 555 mya)
1925:
1924:
1904:
1884:
1796:
1776:
1754:
1727:
1696:
1695:
1667:Rhabdopleura
1665:
1664:
1644:
1623:Hemichordata
1577:
1557:
1517:
1497:
1460:
1439:Ambulacraria
1372:
1352:
1341:Osteichthyes
1322:
1292:
1262:
1232:
1210:
1192:
1177:
1168:Catellocaula
1167:
1159:
1149:
1143:
1132:
1118:
1112:
1106:
1102:Helicoplacus
1100:
1075:
1064:
1058:
1040:
1034:
1024:
1012:
1006:in the late
995:
984:
978:
963:
943:
939:
924:
901:
894:
875:
863:evaginations
856:
845:
826:
801:
702:Hemichordata
696:Ambulacraria
692:
675:Osteichthyes
659:Infraphylum
631:
602:
566:Chaetognatha
555:
530:infrakingdom
520:
513:
459:
446:
432:
428:Cambroernida
413:Hemichordata
331:
315:mouth second
297:
296:
283:Cambroernida
273:Hemichordata
268:Ambulacraria
219:
203:
190:
177:
25:
5508:Bilaterians
5403:Wikispecies
5327:Tusk shells
5312:Cephalopods
5283:Polychaetes
5230:Turbellaria
5211:Crustaceans
5187:Secernentea
5177:Chromadorea
5158:Acorn worms
5128:Echinoderms
5120:Vertebrates
5062:Demosponges
4996:is disputed
4458:Protostomia
4346:is disputed
4273:ParaHoxozoa
3680:(1): 1847.
3545:Han, Jian;
3397:Rhabdotubus
3137:(1): 1286.
3071:(1): 4121.
2839:Development
2785:BMC Biology
2193:(1): 0005.
2039:Han, Jian;
2004:(1): 1â11.
1866:Protostomia
1756:Spengelidae
1519:Ophiuroidea
1178:Below is a
1152:bone tissue
1139:pterobranch
1086:sea urchins
1078:echinoderms
1066:Yanjiahella
1027:Phanerozoic
1017:diversity.
969:protostomes
920:echinoderms
886:echinoderms
871:enterocoely
770:sea lillies
748:Ophiuroidea
673:Superclass
652:Superclass
643:(tunicates)
582:arrow worms
558:Brachiopoda
547:Systematics
541:germ layers
510:arrow worms
486:mud dragons
482:penis worms
470:ecdysozoans
462:Protostomia
455:enterocoely
443:protostomes
419:known from
417:Vetulicolia
329:superphylum
246:Vetulicolia
184:ParaHoxozoa
21:Deuterosome
5518:Superphyla
5502:Categories
5307:Gastropods
5288:Clitellata
5195:Arthropods
5134:Sea lilies
5080:Cnidarians
4998:; see also
4928:Brachiozoa
4691:Syndermata
4650:Gnathifera
4348:; see also
4293:Trichoplax
4291:Placozoa (
4278:Planulozoa
3228:2015-09-01
3027:(19): 1â13
2874:Kimberella
2791:(79): 79.
2107:31 January
1983:References
1927:Kimberella
1559:Echinoidea
1499:Asteroidea
1294:Cyclostomi
1281:Vertebrata
1251:Olfactores
1164:Ordovician
1051:cnidarians
985:Kimberella
980:Kimberella
878:gill slits
782:Echinoidea
775:Subphylum
764:Subphylum
755:Subphylum
743:(starfish)
741:Asteroidea
733:Subphylum
648:Vertebrata
646:Subphylum
639:Subphylum
635:Olfactores
620:Subphylum
597:See also:
539:and three
515:Kimberella
490:spiralians
488:etc.) and
435:blastopore
322:bilaterian
106:(Possible
5293:Sipuncula
5240:Monogenea
5235:Trematoda
5206:Myriapods
5171:Nematodes
5115:Tunicates
5110:Lancelets
5104:Chordates
5096:Myxozoans
5003:Nephrozoa
4992:phylogeny
4792:Salinella
4747:Rouphozoa
4588:Tactopoda
4517:Nematoida
4469:Ecdysozoa
4390:Bilateria
4377:Bilateria
4358:Benthozoa
4353:Eumetazoa
4342:phylogeny
4312:Bilateria
4202:Amoebozoa
4180:Alveolata
4168:Cryptista
4143:Eukaryota
3934:205247296
3767:0028-0836
3702:2045-2322
3647:0960-9822
3209:: 67â78.
3159:2041-1723
3093:2045-2322
2918:BioEssays
2904:156823511
2713:(1): 24.
2680:0960-9822
2631:155104811
2615:0960-9822
2545:0028-0836
2265:1063-5157
2152:0028-0836
2026:128415270
1886:Ecdysozoa
1546:Echinozoa
1486:Asterozoa
1462:Crinoidea
1201:Nephrozoa
1174:Phylogeny
1082:sea stars
1071:Fortunian
1047:Ediacaran
1045:from the
1000:Ediacaran
965:Bilateria
908:chordates
902:Both the
897:tunicates
880:into the
861:forms as
777:Echinozoa
757:Blastozoa
735:Asterozoa
679:tetrapods
626:lancelets
605:phylogeny
570:Phoronida
533:Bilateria
502:flatworms
478:nematoids
210:Nephrozoa
197:Bilateria
167:Kingdom:
161:Eukaryota
108:Ediacaran
5430:BioLib:
5388:Wikidata
5317:Bivalves
5301:Molluscs
5277:Annelids
4636:Spiralia
4243:Animalia
4197:Amorphea
4175:Rhizaria
4163:Hacrobia
4153:Excavata
4138:Bacteria
4046:18192178
3993:33310849
3926:26842059
3871:26842060
3824:32169826
3785:21307940
3720:28500313
3655:31104936
3579:28135722
3532:33741592
3472:27701429
3432:PLOS ONE
3280:12556891
3177:32152310
3111:33602958
3031:13 March
2995:37167976
2946:29130876
2938:17120226
2859:12070079
2817:23031503
2739:23020903
2688:31211978
2623:31104936
2553:17051155
2490:30639106
2441:18339954
2433:16890510
2383:12196991
2306:21708765
2215:28812551
2160:28135722
2073:28135722
1906:Spiralia
1264:Tunicata
1221:Chordata
1129:Cambrian
1090:crinoids
1036:Burykhia
1031:tunicate
906:and the
890:mitrates
859:mesoderm
833:blastula
766:Crinozoa
641:Tunicata
616:Chordata
593:Taxonomy
523:outgroup
506:rotifers
498:annelids
494:mollusks
421:Cambrian
405:Chordata
241:Chordata
171:Animalia
157:Domain:
36:Cambrian
5446:8814528
5394:Q150866
5322:Chitons
5245:Cestoda
5182:Enoplea
5056:Sponges
4829:Symbion
4772:Mesozoa
4212:Animals
4133:Archaea
4109:Extant
4037:2615822
3984:7732190
3961:Bibcode
3906:Bibcode
3879:3870574
3851:Bibcode
3776:4025995
3747:Bibcode
3711:5431833
3682:Bibcode
3615:Bibcode
3559:Bibcode
3523:7978419
3502:Bibcode
3463:5049775
3440:Bibcode
3401:Lethaia
3382:4234408
3362:Bibcode
3335:4402854
3315:Bibcode
3288:4401274
3260:Bibcode
3168:7063041
3139:Bibcode
3102:7893023
3073:Bibcode
2973:Bibcode
2808:3462677
2730:3564725
2707:Evodevo
2658:Bibcode
2585:Bibcode
2561:4366885
2525:Bibcode
2468:Bibcode
2411:Bibcode
2375:7886451
2355:Bibcode
2347:Science
2195:Bibcode
2132:Bibcode
2053:Bibcode
2006:Bibcode
1156:Permian
1131:fossil
1060:Arkarua
1055:sponges
1015:innexin
882:pharynx
810:group.
727:Phylum
700:Phylum
654:Agnatha
614:Phylum
562:Bryozoa
395:during
327:of the
325:animals
309:
233:Clades
225:Grobben
40:Present
5459:914156
5347:bolded
4190:Plants
4127:Domain
4111:animal
4044:
4034:
3991:
3981:
3932:
3924:
3898:Nature
3877:
3869:
3843:Nature
3822:
3783:
3773:
3765:
3739:Nature
3718:
3708:
3700:
3653:
3645:
3587:353780
3585:
3577:
3551:Nature
3530:
3520:
3470:
3460:
3380:
3354:Nature
3350:Pikaia
3333:
3307:Nature
3286:
3278:
3252:Nature
3175:
3165:
3157:
3109:
3099:
3091:
2993:
2944:
2936:
2902:
2892:
2857:
2815:
2805:
2762:
2737:
2727:
2686:
2678:
2629:
2621:
2613:
2559:
2551:
2543:
2517:Nature
2488:
2439:
2431:
2381:
2373:
2304:
2263:
2221:
2213:
2168:353780
2166:
2158:
2150:
2124:Nature
2081:353780
2079:
2071:
2045:Nature
2024:
1158:-aged
1120:Pikaia
947:cloaca
867:coelom
829:zygote
746:Class
739:Class
719:Class
712:Class
706:Class
666:Class
580:. The
568:, and
451:coelom
439:cloaca
320:) are
300:(from
227:, 1908
5485:67145
5472:33511
5433:14961
4509:N+L+P
4217:Fungi
4114:phyla
3930:S2CID
3875:S2CID
3820:S2CID
3583:S2CID
3378:S2CID
3331:S2CID
3284:S2CID
3222:(PDF)
3199:(PDF)
2942:S2CID
2627:S2CID
2557:S2CID
2437:S2CID
2379:S2CID
2223:90795
2219:S2CID
2164:S2CID
2077:S2CID
2022:S2CID
1042:Ausia
990:538.8
916:heart
912:aorta
693:Clade
632:Clade
425:clade
401:phyla
393:mouth
302:Greek
204:Clade
191:Clade
178:Clade
5467:NCBI
5454:ITIS
4990:The
4340:The
4062:UCMP
4042:PMID
3989:PMID
3922:PMID
3867:PMID
3781:PMID
3763:ISSN
3716:PMID
3698:ISSN
3651:PMID
3643:ISSN
3575:PMID
3528:PMID
3468:PMID
3276:PMID
3173:PMID
3155:ISSN
3107:PMID
3089:ISSN
3033:2024
2991:PMID
2934:PMID
2900:OCLC
2890:ISBN
2855:PMID
2813:PMID
2760:ISBN
2735:PMID
2684:PMID
2676:ISSN
2619:PMID
2611:ISSN
2549:PMID
2541:ISSN
2486:PMID
2429:PMID
2371:PMID
2302:PMID
2261:ISSN
2211:PMID
2156:PMID
2148:ISSN
2109:2017
2069:PMID
1111:and
1088:and
1039:and
996:i.e.
892:).
389:anus
306:lit.
46:Preê
5441:EoL
5418:ADW
4847:M+K
4739:R+M
4676:M+S
4545:L+P
4070:at
4032:PMC
4024:doi
4020:363
3979:PMC
3969:doi
3914:doi
3902:530
3859:doi
3847:530
3812:doi
3771:PMC
3755:doi
3743:470
3706:PMC
3690:doi
3633:hdl
3623:doi
3567:doi
3555:542
3518:PMC
3510:doi
3458:PMC
3448:doi
3409:doi
3370:doi
3358:384
3323:doi
3311:402
3268:doi
3256:421
3250:".
3211:doi
3163:PMC
3147:doi
3097:PMC
3081:doi
2981:doi
2926:doi
2882:doi
2847:doi
2843:129
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