276:
rings split or only encircle part of the trunk. Each annulus is essentially identical to its neighbours; the only trunk differentiation is at the anterior and posterior. The anterior is radially symmetrical, typically comprising an introvert, whereas the trunk is bilaterally symmetrical. The posterior hosts the terminal anus and sometimes one or two hooks. There is no one character that unites the palaeoscolecids as a clade (indeed they are likely paraphyletic), and few individual specimens contain all characteristic palaeosolecid traits.
139:
113:
312:, although as more characters are described a position closer to the priapulids becomes most probable. This said, their pharynx has the sixfold symmetry that likely characterised the ancestral ecdysozoan, rather than the fivefold symmetry of priapulans. A nematomorph affinity appears to be an artefact that results from under-sampling of the priapulid stem group. Their relationship with
293:
The group contains a wide and continuous spectrum of morphological variety, making further division of the group difficult; moreover, non-palaeoscolecid taxa likely evolved from palaeoscolecid-like ancestors, and it is thus difficult to demarcate a single clade that corresponds to the palaeoscolecid concept.
292:
Palaeoscolecids are somewhat challenging to define, and probably represent a paraphyletic grouping. Their most current systematic diagnosis references their annulated worm-like body form, the presence of rows (usually) of phosphatic plates, and a straight gut, with the anus at the end of the animal.
275:
Palaeoscolecids bear an annulated trunk ornamented with circular patterns of phosphatic tesselating plates; a layered cuticle; and an armoured proboscis. They are long and narrow, and can reach tens of centimetres in length. Their cuticle is annulated, typically in complete rings, but sometimes the
1228:
has four transverse rings of plates per annulus, arranged as two 'bands' of double rows of plates separated by a central naked zone. Occasionally, single row bifurcates into two rows (for up to 6 rows per annulus). The plates are circular, and unornamented. Platelets and microplates are absent.
339:
As palaeoscolecids may represent a grade rather than a clade, drawing up a formal taxonomy proves problematic. What is more, two parallel taxonomies exist: a form taxonomy for sclerites, and a true taxonomy for articulated fossils. The most recent holistic study of priapulids by Harvey
2351:
323:
Martin R. Smith and Alavya
Dhungana suggested in a 2022 publication that palaeoscolecids are a grade including sister taxa to Panarthropoda, highlighting similarities between the dorsal plates of taxa such as cricocosmiids and those of lobopodians such as
2536:
Whitaker, Anna F.; Jamison, Paul G.; Schiffbauer, James D.; Kimmig, Julien K. (2020). "Re‑description of the Spence Shale palaeoscolecids in light of new morphological features with comments on palaeoscolecid taxonomy and taphonomy".
2352:
https://www.researchgate.net/profile/Xi_Ping_Dong/publication/264275844_Furongian_%28_Late_Cambrian_%29_Palaeoscolecid_Cuticles_from_Hunan_Province__South_China_the_Growth_Impact_on_Worm_Cuticle/links/53d894d70cf2631430c3250c.pdf
1603:
Zhu, M.; Babcock, L.; Steiner, M. (2 May 2005). "Fossilization modes in the
Chengjiang Lagerstätte (Cambrian of China): testing the roles of organic preservation and diagenetic alteration in exceptional preservation".
666:
Other long and narrow
Palaeozoic worms that exhibit an invariant body width are commonly referred to the palaeoscolecids, even though they lack the cuticular structure that defines the group; this 'Palaeoscolecid
2349:
Duan B, Dong X. 2013. Furongian (Late
Cambrian) palaeoscolecid cuticles from Hunan Province, South China: the growth impact on the worm cuticle. Acta Scientiarum Naturalium Universitatis Pekinensis 49: 591–602.
1974:"Discussion on 'Tabelliscolex (Cricocosmiidae: Palaeoscolecidomorpha) from the early Cambrian Chengjiang Biota and the evolution of seriation in Ecdysozoa' by Shi et al. 2021 (JGS, jgs2021-060)"
1214:(unless this simplicity is taphonomic). Its introvert has a six-fold symmetry, whereas its proboscis has quincuncially arranged teeth that resemble those of other Cambrian ecdysozoan worms.
2070:
Yuning, Yang; Xingliang, Zhang (2016). "Distinctive
Scleritome with Marginal Tubercles of a New Palaeoscolecid Worm from the Shipai Fauna (Cambrian Epoch 2) at Three Gorges, South China".
2363:
Xian, Xiao-Feng; Eriksson, Mats E.; Zhang, Hua-Qiao (2023). "Growth patterns of palaeoscolecid sclerites from the
Furongian (Upper Cambrian) Wangcun section, western Hunan, South China".
2273:
Xian, Xiao-Feng; Eriksson, Mats E.; Zhang, Hua-Qiao (2023). "Growth patterns of palaeoscolecid sclerites from the
Furongian (Upper Cambrian) Wangcun section, western Hunan, South China".
1675:
Xian, Xiao-Feng; Eriksson, Mats E.; Zhang, Hua-Qiao (2023). "Growth patterns of palaeoscolecid sclerites from the
Furongian (Upper Cambrian) Wangcun section, western Hunan, South China".
1404:
Ma, X.; Aldridge, R. J.; Siveter, D. J.; Siveter, D. J.; Hou, X.; Edgecombe, G. D. (March 2014). "A New
Exceptionally Preserved Cambrian Priapulid from the Chengjiang Lagerstätte".
2238:
Topper, T. P.; Brock, G. A.; Skovsted, C. B.; Paterson, J. R. (2009). "Palaeoscolecid scleritome fragments with
Hadimopanella plates from the early Cambrian of South Australia".
2612:
Huang, D.; Chen, J.; Zhu, M.; Zhao, F. (2014). "The burrow dwelling behavior and locomotion of palaeoscolecidian worms: New fossil evidence from the Cambrian Chengjiang fauna".
806:
2501:
GarcĂa-Bellido, D. C.; Paterson, J. R.; Edgecombe, G. D. (2013). "Cambrian palaeoscolecids (Cycloneuralia) from Gondwana and reappraisal of species assigned to Palaeoscolex".
2583:
Muir, Lucy A.; Ng, Tin-Wai; Li, Xiang-Feng; Zhang, Yuan-Dong; Lin, Jih-Pai (2014). "Palaeoscolecidan worms and a possible nematode from the Early Ordovician of South China".
799:
1017:
1007:
847:
823:
344:(2010) defines a core of palaeoscolecids characterized by a cuticle that is made up of interlocking plates of multiple sizes, and a looser assemblage (palaeoscolecids
2681:
Goñi, Iban; Skovsted, Christian; Li, Luoyang; Li, Guoxiang; Betts, Marissa; Dorjnamjaa, Dorj; Altanshagai, Gundsambuu; Enkhbaatar, Batkhuyag; Topper, Timothy (2023).
792:
1821:
Peel, J. S. (March 2010). "A Corset-Like Fossil from the Cambrian Sirius Passet Lagerstätte of North Greenland and Its Implications for Cycloneuralian Evolution".
1109:
935:
966:
1573:
859:
841:
599:
2474:
1034:
778:
527:
1919:"Re-description of the Spence Shale palaeoscolecids in light of new morphological features with comments on palaeoscolecid taxonomy and taphonomy"
2857:
1044:
622:
607:
591:
575:
543:
1071:
766:
640:
583:
567:
519:
496:
440:
431:
1174:
1100:
1081:
929:
865:
480:
2023:"Tabelliscolex (Cricocosmiidae: Palaeoscolecidomorpha) from the early Cambrian Chengjiang Biota and the evolution of seriation in Ecdysozoa"
1298:"Palaeoscolecids from the Ludlow Series of Leintwardine, Herefordshire (UK): the latest occurrence of palaeoscolecids in the fossil record"
1090:
1053:
956:
897:
705:" by Harvey et al. (2010), but it has been argued that members of this family do exhibit the requisite cuticular structure, if discreetly.
551:
535:
423:
415:
2159:
Streng, Michael; Ebbestad, Jan Ove R.; Berg-Madsen, Vivianne (2016). "Cambrian palaeoscolecids (Cycloneuralia) of southern Scandinavia".
832:
691:
629:
331:
This proposal was made in response to a 2021 paper that found in a phylogenetic analysis that paleoscolecids were stem-group priapulids.
1528:
2450:"Trunk ornament on the palaeoscolecid worms Cricocosmia and Tabelliscolex from the Early Cambrian Chengjiang deposits of China"
1587:
2751:"The cuticular structure of the 495-Myr-old type species of the fossil worm ~Palaeoscolex~, ~P. piscatorum~ (?Priapulida)"
2903:
774:
Defined by the presence of an unarmoured neck between the proboscis and the trunk, and a single pair of posterior hooks.
316:
is unclear, and either group may be paraphyletic to the other. Some authors choose to include paleoscolecids within
1577:
1210:
This genus has simple sclerites with a single (small but prominent) node in the middle, so can be separated from
17:
1261:
284:
Palaeoscolecids can grow by the continuous addition of plates, or by the continued growth of individual plates.
2793:
2908:
1656:
138:
1296:
Howard, Richard J.; Parry, Luke A.; Clatworthy, Innes; D'Souza, Leila; Edgecombe, Gregory D. (May 2024).
2848:
244:); they are mainly found as disarticulated sclerites, but are also preserved in many of the Cambrian
1778:
1762:"New palaeoscolecidan worms from the Lower Cambrian: Sirius Passet, Latham Shale, and Kinzers Shale"
2021:
Shi, Xiaomei; Howard, Richard J.; Edgecombe, Gregory D.; Hou, Xianguang; Ma, Xiaoya (March 2022).
2898:
2304:
Zhang, X.-G.; Pratt, B. R. (1996). "Early Cambrian palaeoscolecid cuticles from Skaanxi, China".
1536:
2115:"First palaeoscolecid from the Cambrian (Miaolingian, Drumian) Marjum Formation of western Utah"
2113:
Leibach, Wade; Lerosey-Aubril, Rudy; Whitaker, Anna; Schiffbauer, James; Kimmig, Julien (2021).
2810:
2449:
1773:
1358:
Harvey, T. H.; Dong, X.; Donoghue, P. C. (2010). "Are palaeoscolecids ancestral ecdysozoans?".
755:
2875:
2395:
1640:
2870:
2785:
Taphonomy and palaeoecology of the early Cambrian Chengjiang Biota from eastern Yunnan, China
2468:
1809:
989:
980:
748:
2654:"A new priapulid assemblage from the early Cambrian Guanshan fossil Lagerstätte of SW China"
2652:
Hu, S.; Steiner, M.; Zhu, M.; Luo, H.; Forchielli, A.; Keupp, H.; Zhao, F.; Liu, Q. (2012).
1917:
Whitaker, Anna F.; Jamison, Paul G.; Schiffbauer, James D.; Kimmig, Julien (December 2020).
2862:
2621:
2546:
2510:
2411:
2313:
2216:
2079:
2034:
1985:
1930:
1883:
1830:
1724:
1613:
1460:
1413:
1248:
Wills, M. A. (1 April 1998). "Cambrian and Recent Disparity: the Picture from Priapulids".
8:
2722:
Repetski, John E. (1981). "An Ordovician Occurrence of Utahphospha MĂĽller & Miller".
2625:
2550:
2514:
2415:
2396:"New palaeoscolecid worms from the Furongian (upper Cambrian) of Hunan, South China: Is
2317:
2083:
2038:
1989:
1934:
1887:
1834:
1728:
1617:
1545:
1464:
1417:
1297:
2767:
2750:
2731:
2704:
2562:
2427:
2376:
2329:
2286:
2255:
2176:
2136:
2095:
2003:
1954:
1899:
1846:
1791:
1742:
1688:
1571:
1529:"Articulated palaeoscolecid sclerite arrays from the Lower Cambrian of eastern Siberia"
1478:
1429:
1383:
1273:
1265:
701:
and, by extension, the family Maotianshaniidae, was excluded from the "Palaeoscolecids
133:
1918:
2789:
2783:
2708:
2566:
2423:
2380:
2333:
2290:
2140:
2099:
2052:
2007:
1958:
1946:
1903:
1692:
1583:
1375:
1371:
1319:
1135:
734:
449:
313:
2431:
2259:
2180:
1850:
1795:
1746:
1482:
1433:
1387:
1277:
2762:
2694:
2661:
2629:
2594:
2554:
2518:
2447:
2419:
2368:
2321:
2278:
2247:
2168:
2126:
2087:
2042:
1993:
1938:
1891:
1838:
1783:
1732:
1680:
1621:
1509:
1501:
1468:
1421:
1367:
1309:
1257:
264:
1761:
2633:
2598:
2372:
2282:
1684:
1625:
1498:
Data from: A palaeoscolecid worm from the Burgess Shale. Dryad Digital Repository
300:, although their position within this group is unresolved; they may lie with the
742:
Defined by the presence of smooth, folded and sclerite-bearing cuticular regions
267:. Their taxonomic affinities within Ecdysozoa have been the subject of debate.
2833:
2558:
1942:
261:
2325:
2251:
1895:
245:
2892:
2666:
2653:
2522:
2056:
1950:
1579:
The Cambrian fossils of Chengjiang, China: the flowering of early animal life
1513:
1323:
784:
697:
685:
488:
462:
457:
297:
173:
121:
70:
2091:
1638:
2699:
2682:
2131:
2114:
1379:
881:
406:
397:
375:
366:
326:
305:
250:
2683:"New Palaeoscolecid plates from the Cambrian Stage 3 of northern Mongolia"
1787:
2842:
2585:
1868:
917:
907:
815:
679:
470:
256:
45:
2112:
2047:
2022:
1998:
1973:
1505:
2788:. Vol. 7. Berliner Paläobiologische Abhandlungen. pp. 1–197.
2735:
1737:
1712:
1473:
1448:
1269:
708:
317:
90:
55:
2172:
1842:
1639:
Jian Han; Jianni Liu; Zhifei Zhang; Xinglian Zhang; Degan Shu (2007).
1314:
112:
1182:
sclerites that have 3–10 nodes in a single circle, perhaps including
1062:
715:
673:
559:
309:
301:
233:
229:
150:
95:
2804:
1867:
Yang, Jie; Smith, Martin R.; Zhang, Xi-Guang; Yang, Xiao-yu (2020).
2827:
1425:
1138:
for palaeoscolecid macrofossils. The most recent proposal is that
241:
237:
85:
80:
65:
60:
50:
1869:"Introvert and pharynx of Mafangscolex, a Cambrian palaeoscolecid"
100:
75:
2535:
1916:
1295:
2500:
1526:
160:
2496:
2494:
2492:
2490:
2488:
2486:
2484:
2647:
2645:
2643:
348:) including other unconfirmed and palaeoscolecid-like forms:
2448:
Han, J., Liu, J., Zhang, Z., Zhang, X., and Shu, D. (2007).
2217:"Palaeoscolecid worms from the Middle Cambrian of Australia"
2481:
2237:
1262:
10.1666/0094-8373(1998)024[0177:CARDTP]2.3.CO;2
509:(from Orsten-type deposits, preserved in three dimensions)
2640:
1810:
http://gf.tmsoc.org/Documents/Mikro2009/GFSP15.pdf#page=82
891:-type plates, i.e. rectangular with parallel rows of nodes
2158:
308:. They have also been described as a sister-group to the
1403:
2214:
2020:
1649:
from the Early Cambrian Chengjiang deposits of China"
1399:
1397:
1224:
Originally described from the Spence Shale of Utah,
709:
Status impossible to determine from current material
2394:Duan, B.; Dong, X. -P.; Donoghue, P. C. J. (2012).
2345:
2343:
1866:
1357:
1178:should now be considered to include all taxa with
512:
2614:Palaeogeography, Palaeoclimatology, Palaeoecology
2362:
2272:
1759:
1674:
1606:Palaeogeography, Palaeoclimatology, Palaeoecology
1602:
1394:
1196:Though these genera have sclerites that resemble
2890:
2680:
2611:
2393:
2340:
1572:Xianguang Hou; Richard Aldridge; Jan Bergström;
351:
248:. They take their name from the typifying genus
2651:
2069:
1527:Andrey Y. Ivantsov & Ryszard Wrona (2004).
500:(2 species, Ordovician-Silurian, Great Britain)
2210:
1713:"A palaeoscolecid worm from the Burgess Shale"
1449:"A palaeoscolecid worm from the Burgess Shale"
2582:
2578:
2576:
2215:MĂĽller, K. J.; Hinz-Schallreuter, I. (1993).
2208:
2206:
2204:
2202:
2200:
2198:
2196:
2194:
2192:
2190:
1972:Smith, Martin Ross; Dhungana, Alavya (2022).
1971:
1862:
1860:
1706:
1704:
1702:
1353:
1189:
658:
296:They are considered by some to belong to the
2473:: CS1 maint: multiple names: authors list (
1641:"Trunk ornament on the palaeoscolecid worms
1351:
1349:
1347:
1345:
1343:
1341:
1339:
1337:
1335:
1333:
1291:
1289:
1287:
1243:
1241:
504:
411:(Drumian, Cambrian; Marjum Formation, Utah)
359:
2573:
2443:
2441:
2303:
2187:
1857:
1699:
111:
2781:
2766:
2755:Zoological Journal of the Linnean Society
2698:
2665:
2154:
2152:
2150:
2130:
2046:
1997:
1777:
1736:
1567:
1565:
1496:Smith, Martin R. (2015). "Lagerstatten".
1472:
1330:
1313:
1284:
2721:
2387:
1814:
1238:
1146:-type sclerites, as in the type species
719:represents an embryonic Palaeoscolecid.
492:? sp. (late early Cambrian, Scandinavia)
484:? johanssoni (mid-Cambrian, Scandinavia)
2438:
2072:Acta Geologica Sinica - English Edition
1760:Conway Morris, S.; Peel, J. S. (2010).
1753:
1632:
1119:
474:(Sirius Passet; fine structure unclear)
14:
2891:
2748:
2147:
1562:
2809:
2808:
1710:
1495:
1446:
1247:
287:
1820:
950:Other Palaeoscolecidae or unassigned
1440:
1172:Originally described from Siberia,
722:
24:
2768:10.1111/j.1096-3642.1997.tb00136.x
875:sclerites with three to ten nodes.
191:Conway Morris & Robinson, 1986
25:
2920:
2027:Journal of the Geological Society
1978:Journal of the Geological Society
2424:10.1111/j.1475-4983.2012.01148.x
1582:. Wiley-Blackwell. p. 233.
1372:10.1111/j.1525-142X.2010.00403.x
236:. They are known from the Lower
137:
43:
2775:
2742:
2715:
2674:
2605:
2529:
2356:
2297:
2266:
2231:
2106:
2063:
2014:
1965:
1910:
1802:
1668:
1203:
1124:
513:Palaeoscolecidae From Australia
41:Upper Early Cambrian–Ludfordian
1596:
1520:
1489:
1165:
1142:should only include taxa with
435:(early Ordovician, Czech Rep.)
427:(early Ordovician, Czech Rep.)
240:to the lower Ludfordian (late
13:
1:
2687:Acta Palaeontologica Polonica
2457:Acta Palaeontologica Polonica
2400:an embryonic palaeoscolecid?"
2119:Acta Palaeontologica Polonica
1766:Acta Palaeontologica Polonica
1657:Acta Palaeontologica Polonica
1232:
1217:
807:Tabelliscolex chengjiangensis
738:Conway Morris & Peel 2010
615:
402:(Burgess Shale, mid-Cambrian)
270:
215:
197:Families, genera and species
2634:10.1016/j.palaeo.2013.11.004
2599:10.1016/j.palwor.2013.06.003
2373:10.1016/j.palwor.2023.03.005
2283:10.1016/j.palwor.2023.03.005
1685:10.1016/j.palwor.2023.03.005
1626:10.1016/j.palaeo.2003.03.001
1200:, they remain valid genera.
636:(Order and Family uncertain)
7:
1360:Evolution & Development
1198:Hadimopanella knappologicum
1095:Ivantsov & Wrona, 2004
800:Tabelliscolex maanshanensis
419:(early Cambrian of Siberia)
334:
128:. Image from Smith (2015)
10:
2925:
2904:Cambrian first appearances
2782:Shixue Hu (October 2005).
2749:Conway, Morris S. (1997).
2559:10.1007/s12542-020-00516-9
1943:10.1007/s12542-020-00516-9
1191:Guanduscolex, Wudingscolex
1158:should not be included in
1018:Wudingscolex sapushanensis
232:worms resembling armoured
2817:
2326:10.1017/s0022336000023350
2252:10.1017/S0016756809990082
1896:10.1017/S0016756820000308
1711:Smith, Martin R. (2015).
1544:(1): 1–22. Archived from
1447:Smith, Martin R. (2015).
1008:Paramaotianshania zijunia
1000:Parapalaeoscolex sinensis
465:; fine structure unclear)
391:Sanxiascolex papillogyrus
279:
213:
196:
134:Scientific classification
132:
119:
110:
34:
2667:10.3140/bull.geosci.1238
2523:10.1016/j.gr.2012.12.002
1576:; Derek Siveter (2004).
1076:Kraft & Mergl, 1989
961:Kraft & Mergl, 1989
848:Maotianshania cylindrica
824:Cricocosmia jinningensis
505:Articulated microfossils
360:Articulated macrofossils
2724:Journal of Paleontology
2658:Bulletin of Geosciences
2306:Journal of Paleontology
2161:Papers in Palaeontology
2092:10.1111/1755-6724.12724
1823:Journal of Paleontology
1537:Acta Geologica Polonica
1406:Journal of Paleontology
1302:Papers in Palaeontology
887:Defined by presence of
871:Defined by presence of
793:Tabelliscolex hexagonus
539:Muller & Hinz 1993*
445:(early Cambrian, China)
393:(early Cambrian, China)
386:(early Cambrian, China)
254:. Other genera include
228:are a group of extinct
2700:10.4202/app.01030.2022
2132:10.4202/app.00875.2021
996:Mafangscolex siniensis
756:Xystoscolex boreogyrus
611:Muller & Hinz 1993
603:Muller & Hinz 1993
595:Muller & Hinz 1993
587:Muller & Hinz 1993
579:Muller & Hinz 1993
571:Muller & Hinz 1993
555:Muller & Hinz 1993
531:Muller & Hinz 1993
523:Muller & Hinz 1993
27:Extinct class of worms
2871:Paleobiology Database
1788:10.4202/app.2009.0058
1134:has been abused as a
994:Hou & Sun 1988 (=
990:Palaeoscolex sinensis
981:Louisella pedunculata
749:Chalazoscolex pharkus
2909:Silurian extinctions
1120:Genus level taxonomy
1110:Yunnanoscolex magnus
1028:Microfossil material
936:Tylotites petiolaris
902:Whitaker et al. 2020
713:It is possible that
2626:2014PPP...398..154H
2551:2020PalZ...94..661W
2515:2013GondR..24..780G
2416:2012Palgy..55..613D
2318:1996JPal...70..275Z
2240:Geological Magazine
2084:2016AcGlS..90..807Y
2048:10.1144/jgs2021-060
2039:2022JGSoc.179...60S
1999:10.1144/jgs2021-111
1990:2022JGSoc.179..111S
1935:2020PalZ...94..661W
1888:2020GeoM..157.2044Y
1876:Geological Magazine
1835:2010JPal...84..332P
1729:2015Palgy..58..973S
1618:2005PPP...220...31Z
1506:10.5061/dryad.cf493
1465:2015Palgy..58..973S
1418:2014JPal...88..371M
2033:(2): jgs2021–060.
1738:10.1111/pala.12210
1474:10.1111/pala.12210
967:Guanduscolex minor
371:(early Ordovician)
288:Taxonomic position
126:Scathascolex minor
2886:
2885:
2811:Taxon identifiers
2503:Gondwana Research
2173:10.1002/spp2.1067
1882:(12): 2044–2050.
1843:10.1666/09-102R.1
1589:978-1-4051-0673-3
1514:10255/dryad.92916
1315:10.1002/spp2.1558
1136:wastebasket taxon
1115:
1105:
1096:
1086:
1077:
1067:
1058:
1049:
1040:
1039:Goñi et al. 2023
1023:
1013:
1003:
985:
976:
962:
941:
923:
913:
903:
853:
852:Sun and Huo, 1987
771:
739:
735:Chalazoscolecidae
653:
646:
635:
634:Zhang & Pratt
450:Chalazoscolecidae
314:Archaeopriapulida
222:
221:
209:
192:
16:(Redirected from
2916:
2879:
2878:
2866:
2865:
2853:
2852:
2851:
2838:
2837:
2836:
2806:
2805:
2800:
2799:
2779:
2773:
2772:
2770:
2746:
2740:
2739:
2719:
2713:
2712:
2702:
2678:
2672:
2671:
2669:
2649:
2638:
2637:
2609:
2603:
2602:
2580:
2571:
2570:
2533:
2527:
2526:
2498:
2479:
2478:
2472:
2464:
2454:
2445:
2436:
2435:
2391:
2385:
2384:
2360:
2354:
2347:
2338:
2337:
2301:
2295:
2294:
2270:
2264:
2263:
2235:
2229:
2228:
2212:
2185:
2184:
2156:
2145:
2144:
2134:
2110:
2104:
2103:
2067:
2061:
2060:
2050:
2018:
2012:
2011:
2001:
1969:
1963:
1962:
1914:
1908:
1907:
1873:
1864:
1855:
1854:
1818:
1812:
1806:
1800:
1799:
1781:
1757:
1751:
1750:
1740:
1708:
1697:
1696:
1672:
1666:
1665:
1653:
1636:
1630:
1629:
1600:
1594:
1593:
1569:
1560:
1559:
1557:
1556:
1550:
1533:
1524:
1518:
1517:
1493:
1487:
1486:
1476:
1444:
1438:
1437:
1401:
1392:
1391:
1355:
1328:
1327:
1317:
1293:
1282:
1281:
1245:
1113:
1104:
1094:
1085:
1075:
1066:
1057:
1048:
1038:
1021:
1011:
993:
984:
970:
960:
940:Luo and Hu, 1999
939:
921:
911:
901:
860:Palaeoscolecidae
851:
842:Maotianshaniidae
769:
737:
723:Linnean taxonomy
659:Palaeoscolecids
651:
644:
633:
352:Palaeoscolecids
265:Chengjiang biota
199:
190:
185:
142:
141:
115:
105:
42:
38:Temporal range:
32:
31:
21:
2924:
2923:
2919:
2918:
2917:
2915:
2914:
2913:
2889:
2888:
2887:
2882:
2874:
2869:
2861:
2856:
2849:Palaeoscolecida
2847:
2846:
2841:
2832:
2831:
2826:
2819:Palaeoscolecida
2813:
2803:
2796:
2780:
2776:
2747:
2743:
2720:
2716:
2679:
2675:
2650:
2641:
2610:
2606:
2581:
2574:
2534:
2530:
2499:
2482:
2466:
2465:
2452:
2446:
2439:
2392:
2388:
2361:
2357:
2348:
2341:
2302:
2298:
2271:
2267:
2236:
2232:
2213:
2188:
2157:
2148:
2111:
2107:
2068:
2064:
2019:
2015:
1970:
1966:
1915:
1911:
1871:
1865:
1858:
1819:
1815:
1807:
1803:
1779:10.1.1.620.4930
1758:
1754:
1709:
1700:
1673:
1669:
1651:
1637:
1633:
1601:
1597:
1590:
1570:
1563:
1554:
1552:
1548:
1531:
1525:
1521:
1494:
1490:
1445:
1441:
1402:
1395:
1356:
1331:
1294:
1285:
1246:
1239:
1235:
1222:
1208:
1194:
1170:
1156:P. huainanensis
1129:
1122:
922:Muir et al 2014
912:Muir et al 2014
770:Han et al. 2007
728:Order Uncertain
725:
711:
671:group includes
664:
652:Duan & Dong
645:Duan & Dong
618:
600:Shergoldiscolex
515:
507:
362:
357:
337:
290:
282:
273:
226:palaeoscolecids
198:
189:
187:Palaeoscolecida
183:
136:
124:palaeoscolecid
106:
104:
103:
98:
93:
88:
83:
78:
73:
68:
63:
58:
53:
48:
40:
39:
36:
35:Palaeoscolecida
28:
23:
22:
18:Palaeoscolecida
15:
12:
11:
5:
2922:
2912:
2911:
2906:
2901:
2899:Paleoscolecids
2884:
2883:
2881:
2880:
2867:
2854:
2839:
2823:
2821:
2815:
2814:
2802:
2801:
2794:
2774:
2741:
2730:(2): 395–400.
2714:
2673:
2639:
2604:
2572:
2545:(4): 661–674.
2528:
2509:(2): 780–795.
2480:
2437:
2410:(3): 613–622.
2386:
2355:
2339:
2312:(2): 275–279.
2296:
2265:
2230:
2186:
2146:
2105:
2062:
2013:
1964:
1929:(4): 661–674.
1909:
1856:
1829:(2): 332–340.
1813:
1801:
1772:(1): 141–156.
1752:
1723:(6): 973–979.
1698:
1667:
1631:
1612:(1–2): 31–37.
1595:
1588:
1561:
1519:
1488:
1459:(6): 973–979.
1439:
1426:10.1666/13-082
1412:(2): 371–384.
1393:
1366:(2): 177–200.
1329:
1283:
1256:(2): 155–286.
1236:
1234:
1231:
1221:
1216:
1207:
1202:
1193:
1188:
1169:
1164:
1128:
1123:
1121:
1118:
1117:
1116:
1114:Hu et al 2012
1106:
1097:
1087:
1078:
1068:
1059:
1050:
1041:
1035:Floraconformis
1030:
1029:
1025:
1024:
1022:Hu et al 2012
1014:
1012:Hu et al 2012
1004:
986:
977:
963:
952:
951:
947:
946:
945:
944:
943:
942:
926:
925:
924:
914:
904:
894:
893:
892:
878:
877:
876:
856:
855:
854:
838:
837:
836:
829:
828:
827:
812:
811:
810:
803:
796:
779:Cricocosmiidae
772:
762:
761:
760:
759:
752:
744:
743:
730:
729:
724:
721:
710:
707:
663:
657:
656:
655:
637:
626:
617:
614:
613:
612:
604:
596:
588:
580:
572:
564:
556:
548:
540:
532:
528:Corallioscolex
524:
514:
511:
506:
503:
502:
501:
493:
485:
477:
476:
475:
466:
446:
436:
428:
420:
412:
403:
394:
388:
372:
361:
358:
356:
350:
336:
333:
289:
286:
281:
278:
272:
269:
262:Lower Cambrian
220:
219:
211:
210:
194:
193:
181:
177:
176:
171:
164:
163:
158:
154:
153:
148:
144:
143:
130:
129:
117:
116:
108:
107:
99:
94:
89:
84:
79:
74:
69:
64:
59:
54:
49:
44:
37:
26:
9:
6:
4:
3:
2:
2921:
2910:
2907:
2905:
2902:
2900:
2897:
2896:
2894:
2877:
2872:
2868:
2864:
2859:
2855:
2850:
2844:
2840:
2835:
2829:
2825:
2824:
2822:
2820:
2816:
2812:
2807:
2797:
2791:
2787:
2786:
2778:
2769:
2764:
2760:
2756:
2752:
2745:
2737:
2733:
2729:
2725:
2718:
2710:
2706:
2701:
2696:
2692:
2688:
2684:
2677:
2668:
2663:
2659:
2655:
2648:
2646:
2644:
2635:
2631:
2627:
2623:
2619:
2615:
2608:
2600:
2596:
2592:
2588:
2587:
2579:
2577:
2568:
2564:
2560:
2556:
2552:
2548:
2544:
2540:
2532:
2524:
2520:
2516:
2512:
2508:
2504:
2497:
2495:
2493:
2491:
2489:
2487:
2485:
2476:
2470:
2463:(2): 423–431.
2462:
2458:
2451:
2444:
2442:
2433:
2429:
2425:
2421:
2417:
2413:
2409:
2405:
2404:Palaeontology
2401:
2399:
2390:
2382:
2378:
2374:
2370:
2366:
2359:
2353:
2346:
2344:
2335:
2331:
2327:
2323:
2319:
2315:
2311:
2307:
2300:
2292:
2288:
2284:
2280:
2276:
2269:
2261:
2257:
2253:
2249:
2245:
2241:
2234:
2226:
2222:
2221:Palaeontology
2218:
2211:
2209:
2207:
2205:
2203:
2201:
2199:
2197:
2195:
2193:
2191:
2182:
2178:
2174:
2170:
2166:
2162:
2155:
2153:
2151:
2142:
2138:
2133:
2128:
2124:
2120:
2116:
2109:
2101:
2097:
2093:
2089:
2085:
2081:
2077:
2073:
2066:
2058:
2054:
2049:
2044:
2040:
2036:
2032:
2028:
2024:
2017:
2009:
2005:
2000:
1995:
1991:
1987:
1983:
1979:
1975:
1968:
1960:
1956:
1952:
1948:
1944:
1940:
1936:
1932:
1928:
1924:
1920:
1913:
1905:
1901:
1897:
1893:
1889:
1885:
1881:
1877:
1870:
1863:
1861:
1852:
1848:
1844:
1840:
1836:
1832:
1828:
1824:
1817:
1811:
1805:
1797:
1793:
1789:
1785:
1780:
1775:
1771:
1767:
1763:
1756:
1748:
1744:
1739:
1734:
1730:
1726:
1722:
1718:
1717:Palaeontology
1714:
1707:
1705:
1703:
1694:
1690:
1686:
1682:
1678:
1671:
1664:(2): 423–431.
1663:
1659:
1658:
1650:
1648:
1647:Tabelliscolex
1644:
1635:
1627:
1623:
1619:
1615:
1611:
1607:
1599:
1591:
1585:
1581:
1580:
1575:
1574:David Siveter
1568:
1566:
1551:on 2011-07-16
1547:
1543:
1539:
1538:
1530:
1523:
1515:
1511:
1507:
1503:
1499:
1492:
1484:
1480:
1475:
1470:
1466:
1462:
1458:
1454:
1453:Palaeontology
1450:
1443:
1435:
1431:
1427:
1423:
1419:
1415:
1411:
1407:
1400:
1398:
1389:
1385:
1381:
1377:
1373:
1369:
1365:
1361:
1354:
1352:
1350:
1348:
1346:
1344:
1342:
1340:
1338:
1336:
1334:
1325:
1321:
1316:
1311:
1307:
1303:
1299:
1292:
1290:
1288:
1279:
1275:
1271:
1267:
1263:
1259:
1255:
1251:
1244:
1242:
1237:
1230:
1227:
1220:
1215:
1213:
1206:
1201:
1199:
1192:
1187:
1185:
1184:Yunnanoscolex
1181:
1180:Hadimopanella
1177:
1176:
1168:
1163:
1161:
1157:
1153:
1152:P. ratcliffei
1149:
1148:P. piscatorum
1145:
1141:
1137:
1133:
1127:
1112:
1111:
1107:
1103:
1102:
1098:
1093:
1092:
1088:
1084:
1083:
1079:
1074:
1073:
1069:
1065:
1064:
1060:
1056:
1055:
1051:
1047:
1046:
1045:Hadimopanella
1042:
1037:
1036:
1032:
1031:
1027:
1026:
1020:
1019:
1015:
1010:
1009:
1005:
1001:
997:
992:
991:
987:
983:
982:
978:
974:
969:
968:
964:
959:
958:
954:
953:
949:
948:
938:
937:
933:
932:
931:
927:
920:
919:
915:
910:
909:
905:
900:
899:
895:
890:
886:
885:
884:
883:
879:
874:
873:Hadimopanella
870:
869:
868:
867:
863:
862:
861:
857:
850:
849:
845:
844:
843:
839:
835:
834:
830:
826:
825:
821:
820:
819:
817:
813:
809:
808:
804:
802:
801:
797:
795:
794:
790:
789:
788:
786:
785:Tabelliscolex
782:
781:
780:
776:
775:
773:
768:
764:
763:
758:
757:
753:
751:
750:
746:
745:
741:
740:
736:
732:
731:
727:
726:
720:
718:
717:
706:
704:
703:sensu stricto
700:
699:
698:Maotianshania
694:
693:
688:
687:
686:Tabelliscolex
682:
681:
676:
675:
670:
662:
650:
643:
642:
638:
632:
631:
627:
625:
624:
623:Dispinoscolex
620:
619:
610:
609:
608:Thoracoscolex
605:
602:
601:
597:
594:
593:
592:Schistoscolex
589:
586:
585:
581:
578:
577:
576:Pantoioscolex
573:
570:
569:
565:
562:
561:
557:
554:
553:
549:
546:
545:
544:Hadimopanella
541:
538:
537:
533:
530:
529:
525:
522:
521:
517:
516:
510:
499:
498:
494:
491:
490:
489:Maotianshania
486:
483:
482:
478:
473:
472:
467:
464:
463:Sirius Passet
460:
459:
458:Chalazoscolex
454:
453:
451:
447:
444:
442:
437:
434:
433:
429:
426:
425:
421:
418:
417:
413:
410:
408:
404:
401:
399:
395:
392:
389:
387:
383:
379:
377:
373:
370:
368:
364:
363:
355:
354:sensu stricto
349:
347:
343:
332:
330:
328:
321:
319:
315:
311:
307:
303:
299:
298:Cycloneuralia
294:
285:
277:
268:
266:
263:
259:
258:
253:
252:
247:
243:
239:
235:
231:
227:
218:
217:
212:
207:
203:
195:
188:
182:
179:
178:
175:
174:Cycloneuralia
172:
169:
166:
165:
162:
159:
156:
155:
152:
149:
146:
145:
140:
135:
131:
127:
123:
122:Burgess Shale
118:
114:
109:
102:
97:
92:
87:
82:
77:
72:
67:
62:
57:
52:
47:
33:
30:
19:
2818:
2784:
2777:
2758:
2754:
2744:
2727:
2723:
2717:
2690:
2686:
2676:
2657:
2617:
2613:
2607:
2590:
2584:
2542:
2538:
2531:
2506:
2502:
2469:cite journal
2460:
2456:
2407:
2403:
2397:
2389:
2364:
2358:
2309:
2305:
2299:
2274:
2268:
2243:
2239:
2233:
2224:
2220:
2164:
2160:
2122:
2118:
2108:
2075:
2071:
2065:
2030:
2026:
2016:
1981:
1977:
1967:
1926:
1922:
1912:
1879:
1875:
1826:
1822:
1816:
1808:conference,
1804:
1769:
1765:
1755:
1720:
1716:
1676:
1670:
1661:
1655:
1646:
1642:
1634:
1609:
1605:
1598:
1578:
1553:. Retrieved
1546:the original
1541:
1535:
1522:
1497:
1491:
1456:
1452:
1442:
1409:
1405:
1363:
1359:
1305:
1301:
1253:
1250:Paleobiology
1249:
1225:
1223:
1218:
1212:Palaeoscolex
1211:
1209:
1205:Mafangscolex
1204:
1197:
1195:
1190:
1183:
1179:
1173:
1171:
1166:
1160:Palaeoscolex
1159:
1155:
1151:
1150:. As such,
1147:
1143:
1140:Palaeoscolex
1139:
1132:Palaeoscolex
1131:
1130:
1126:Palaeoscolex
1125:
1108:
1099:
1089:
1080:
1072:Plasmuscolex
1070:
1061:
1052:
1043:
1033:
1016:
1006:
999:
995:
988:
979:
972:
965:
955:
934:
916:
906:
896:
888:
882:Palaeoscolex
880:
872:
864:
846:
831:
822:
814:
805:
798:
791:
783:
767:Cricocosmida
754:
747:
714:
712:
702:
696:
695:and others.
690:
684:
678:
672:
668:
665:
660:
649:Ornatoscolex
648:
641:Hunanoscolex
639:
628:
621:
606:
598:
590:
584:Rhomboscolex
582:
574:
568:Murrayscolex
566:
558:
550:
542:
534:
526:
520:Austroscolex
518:
508:
497:Radnorscolex
495:
487:
479:
469:
456:
441:Guanduscolex
439:
432:Plasmuscolex
430:
422:
414:
407:Arrakiscolex
405:
398:Scathascolex
396:
390:
385:
381:
380:Mafangscolex
376:Palaeoscolex
374:
367:Palaeoscolex
365:
353:
345:
341:
338:
327:Microdictyon
325:
322:
306:Nematomorpha
295:
291:
283:
274:
255:
251:Palaeoscolex
249:
246:lagerstätten
225:
223:
214:
205:
201:
186:
167:
125:
29:
2843:Wikispecies
2620:: 154–164.
2586:Palaeoworld
2365:Palaeoworld
2275:Palaeoworld
1677:Palaeoworld
1643:Cricocosmia
1175:Wronascolex
1167:Wronascolex
1101:Utahphospha
1082:Protoscolex
930:Tylotitidae
918:Sanduscolex
908:Ashetscolex
866:Wronascolex
816:Cricocosmia
680:Cricocosmia
669:sensu lato'
563:Muller 1973
481:Wronascolex
471:Xystoscolex
257:Cricocosmia
2893:Categories
2795:3980853179
2660:: 93–106.
2227:: 549–592.
2078:(3): 807.
1555:2009-06-03
1233:References
1226:Utahscolex
1219:Utahscolex
1091:Sahascolex
1054:Kaimenella
957:Gamascolex
898:Utahscolex
661:sensu lato
616:From China
552:Kaloskolex
547:Gedik 1977
536:Euryscolex
424:Gamascolex
416:Sahascolex
369:piscatorum
346:sensu lato
318:Priapulida
302:priapulids
271:Morphology
234:priapulids
230:ecdysozoan
2761:: 69–82.
2709:256711586
2593:: 15–24.
2567:211479504
2398:Markuelia
2381:257601719
2334:132183034
2291:257601719
2246:: 86–97.
2167:: 21–48.
2141:239731912
2100:132852259
2057:0016-7649
2008:244850491
1959:211479504
1951:0031-0220
1904:219092881
1774:CiteSeerX
1693:257601719
1500:. Dryad.
1324:2056-2799
1144:Milaculum
1063:Milaculum
889:Milaculum
833:Houscolex
716:Markuelia
692:Tylotites
674:Louisella
630:Houscolex
560:Milaculum
448:? Family
310:Ecdysozoa
260:from the
157:Kingdom:
151:Eukaryota
2834:Q7126500
2828:Wikidata
2432:53485143
2260:54916222
2181:89406537
1851:86256781
1796:54037571
1747:86747103
1483:86747103
1434:85627132
1388:16872271
1380:20433458
1278:88647544
975:., 2008
384:sinensis
335:Taxonomy
242:Silurian
238:Cambrian
216:See text
161:Animalia
147:Domain:
2736:1304225
2622:Bibcode
2547:Bibcode
2511:Bibcode
2412:Bibcode
2314:Bibcode
2080:Bibcode
2035:Bibcode
1986:Bibcode
1931:Bibcode
1884:Bibcode
1831:Bibcode
1725:Bibcode
1614:Bibcode
1461:Bibcode
1414:Bibcode
1270:2401237
928:Family
858:Family
840:Family
777:Family
733:Family
204:Harvey
180:Class:
2792:
2734:
2707:
2565:
2430:
2379:
2332:
2289:
2258:
2179:
2139:
2098:
2055:
2006:
1957:
1949:
1902:
1849:
1794:
1776:
1745:
1691:
1586:
1481:
1432:
1386:
1378:
1322:
1276:
1268:
765:Order
342:et al.
280:Growth
206:et al.
2858:IRMNG
2732:JSTOR
2705:S2CID
2563:S2CID
2453:(PDF)
2428:S2CID
2377:S2CID
2330:S2CID
2287:S2CID
2256:S2CID
2177:S2CID
2137:S2CID
2096:S2CID
2004:S2CID
1984:(3).
1955:S2CID
1900:S2CID
1872:(PDF)
1847:S2CID
1792:S2CID
1743:S2CID
1689:S2CID
1652:(PDF)
1549:(PDF)
1532:(PDF)
1479:S2CID
1430:S2CID
1384:S2CID
1308:(3).
1274:S2CID
1266:JSTOR
973:et al
443:minor
409:aasei
400:minor
208:2010)
202:sensu
168:Clade
2876:7273
2863:1223
2790:ISBN
2539:PalZ
2475:link
2053:ISSN
1947:ISSN
1923:PalZ
1645:and
1584:ISBN
1376:PMID
1320:ISSN
1154:and
224:The
120:The
46:Preęž’
2763:doi
2759:119
2695:doi
2662:doi
2630:doi
2618:398
2595:doi
2555:doi
2519:doi
2420:doi
2369:doi
2322:doi
2279:doi
2248:doi
2244:147
2169:doi
2127:doi
2088:doi
2043:doi
2031:179
1994:doi
1982:179
1939:doi
1892:doi
1880:157
1839:doi
1784:doi
1733:doi
1681:doi
1622:doi
1610:220
1510:hdl
1502:doi
1469:doi
1422:doi
1368:doi
1310:doi
1258:doi
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