919:
637:
789:
4497:
654:
1489:
994:
684:
122:
971:
1088:
941:
1736:
539:
148:
551:
1024:
1047:
1622:
4510:
1068:
680:(2007) however, interpreted all flaps have posterior spacing where the gill blades attached. Budd and Daley (2011) reject the reconstruction by Zhang & Briggs, showing the flaps have complete posterior edges as in previous reconstructions. They mostly follow the reconstruction by Budd (1996) with modifications on some details (e.g. the first flap pair also have gills; the attachment point of gill blades located more posteriorly than previously thought).
866:
770:
899:
595:. The head bore five stalked eyes: two near the front and fairly close to the middle of the head, pointing upwards and forwards; two larger eyes with longer stalks near the rear and outer edges of the head, pointing upwards and sideways; and a single eye between the larger pair of stalked eyes, pointing upwards. It has been assumed that the eyes were all
676:(1996) thought the gill blades attached along the front edges on the dorsal side of all except the first flaps. He also found marks inside the flaps' front edges that he interpreted as internal channels connecting the gills to the interior of the body, much as Whittington interpreted the mark along the proboscis as an internal channel. Zhang and
587:'s hose and flexible, and it ended with a claw-like structure whose terminal edges bore 5 spines that projected inwards and forwards. The bilateral symmetry and lateral (instead of vertical as reconstructed by Whittington 1975) arrangement of the claw suggest it represent a pair of fused frontal appendages, comparable to those of
2106:
636:
747:
s lobe+gill arrangement as an early form of the arthropod limbs before it split into a biramous structure. However, this similar chemical composition is not only associated with the digestive tract; Budd and Daley (2011) suggest that it represents mineralization forming within fluid-filled cavities
614:
The main part of the body was typically about 5 millimetres (0.20 in) wide and had 15 segments, on each of which there were pairs of flaps (lobes) pointing downwards and outwards. The flaps overlapped so that the front of each was covered by the rear edge of the one ahead of it. The body ended
667:
Whittington (1975) interpreted the gills as paired extensions attached dorsally to the bases of all but the first flaps on each side, and thought that these gills were flat underneath, had overlapping layers on top. Bergström (1986) revealed the "overlapping layers" were rows of individual blades,
721:
views showed they were attached separately from and lower than the lobes, and extended below the body. He later found specimens that appeared to preserve the legs' exterior cuticle. He therefore interpreted the "triangles" as short, fleshy, conical legs (lobopods). He also found small mineralized
1404:
was very closely related to arthropods. In fact they presented a family tree very similar to Budd's except that theirs did not mention tardigrades. Regardless of the different morphological interpretations, all major restudies since 1980s similarly concluded that the resemblance between
3036:
1617:
to include fossil panarthropods that are thought to be close relatives of onychophorans, tardigrades and arthropods but lack jointed limbs. This group was later widely accepted as a paraphyletic grade that led to the origin of extant panarthropod phyla.
740:. Zhang and Briggs (2007) analyzed the chemical composition of the "triangles", and concluded that they had the same composition as the gut, and therefore agreed with Whittington that they were part of the digestive system. Instead they regarded
1711:
is a group of closely related living animals plus their last common ancestor plus all its descendants. A stem group contains offshoots from members of the lineage earlier than the last common ancestor of the crown group; it is a
2433:
Miroshnikov, L. D. and
Krawzov, A. G. (1960). Rare paleontological remains and traces of life in late Cambrian deposits of the northwestern Siberian platform. Palaeontology and biostratigraphy of the Soviet Arctic, 3, pp.
3140:
812:
probably used its proboscis to search the sediment for food particles and pass them to its mouth. Since there is no sign of anything that might function as jaws, its food was presumably small and soft. The paired gut
1511:
made it clear how little was known about soft-bodied animals, which do not usually leave fossils. When
Whittington described it in the mid-1970s, there was already a vigorous debate about the early evolution of
2586:
Xianguang, Hou; Bergström, Jan (2006). "Dinocaridids – anomalous arthropods or arthropod-like worms?". In Jiayu, Rong; Zongjie, Fang; Zhanghe, Zhou; Renbin, Zhan; Xiangdong, Wang; Xunlai, Yuan (eds.).
4104:
Jacobs, D. K.; Wray, C. G.; Wedeen, C. J.; Kostriken, R.; DeSalle, R.; Staton, J. L.; Gates, R. D.; Lindberg, D. R. (2000). "Molluscan engrailed
Expression, Serial Organization, and Shell Evolution".
722:
patches at the tips of some, and interpreted these as claws. Under this reconstruction, the gill-bearing flap and lobopod were homologized to the outer gill branch and inner leg branch of arthropod
3976:
Neoproterozoic-Cambrian
Biological Revolutions: Presented as a Paleontological Society Short Course at the Annual Meeting of the Geological Society of America, Denver, Colorado, November 6, 2004
1237:
was not arthropod in 1975, as he found no evidence for arthropodan jointed limbs, and that nothing like the flexible, probably fluid-filled, proboscis was known in arthropods. Although he left
1720:
are living animals that form a crown group in their own right, but Budd (1996) regarded them also as being a stem group relative to the arthropods. Viewing strange-looking organisms like
1245:
level open, the annulated but not articulated body and the unusual lateral flaps with gills persuaded him that it may have been a representative of the ancestral stock from the origin of
1789:"Extraordinary fossils reveal the nature of Cambrian life: a commentary on Whittington (1975) 'The enigmatic animal Opabinia regalis, Middle Cambrian, Burgess Shale, British Columbia'"
630:
like those of other arthropods, their bodies were flattened as they were buried and fossilized, and smaller or internal features appear as markings within the outlines of the fossils.
1438:
is widely accepted, as consistently shown by multiple phylogenetic analyses, as well as new discoveries such as the presence of arthropod-like gut glands and the intermediate taxon
3828:
312:
was a soft-bodied animal, measuring up to 7 cm in body length, and its segmented trunk had flaps along the sides and a fan-shaped tail. The head shows unusual features: five
1343:
shared by the ancestors of arthropods. After examining several sets of features shared by these and similar lobopodians he drew up a "broad-scale reconstruction of the arthropod
702:
Whittington (1975) found evidence of near-triangular features along the body, and concluded that they were internal structures, most likely sideways extensions of the gut (
1290:
lateral flaps, gill blades, stalked eyes, and specialized frontal appendages). He classified them as primitive arthropods, although he considered that arthropods are not
603:' lateral eyes, but this reconstruction, which is not backed up by any evidence, is "somewhat fanciful". The mouth was under the head, behind the proboscis, and pointed
615:
with what looked like a single conical segment bearing three pairs of overlapping tail fan blades that pointed up and out, forming a tail like a V-shaped double fan.
821:
had no legs, thought that it crawled on its lobes and that it could also have swum slowly by flapping the lobes, especially if it timed the movements to create a
571:
from head (excluding proboscis) to tail end ranged between 4 centimetres (1.6 in) and 7 centimetres (2.8 in). One of the most distinctive characters of
1532:, which views evolution as long intervals of near-stasis "punctuated" by short periods of rapid change. On the other hand, around the same time Wyatt Durham and
653:
825:
with the metachronal movement of its lobes. On the other hand, he thought the body was not flexible enough to allow fish-like undulations of the whole body.
2447:; Nedin, C. (January 1997). "The Taphonomy and Affinities of the Problematic Fossil Myoscolex from the Lower Cambrian Emu Bay Shale of South Australia".
752:
are series of circular gut glands individualized from the "triangles". While they agreed on the absence of terminal claws, the presence of lobopods in
324:
probably lived on the seafloor, using the proboscis to seek out small, soft food. Fewer than twenty good specimens have been described; 3 specimens of
2393:
918:
4237:
1933:
1891:
1724:
in this way makes it possible to see that, while the
Cambrian explosion was unusual, it can be understood in terms of normal evolutionary processes.
347:
ancestors. However, later studies since late 1990s consistently support its affinity as a member of basal arthropods, alongside the closely related
4549:
748:
within the body, which is consistent with hollow lobopods as seen in unequivocal lobopodian fossils. They also clarify that the gut diverticula of
808:
animals were buried, by a mudslide or a sediment-laden current that acted as a sandstorm, suggests they lived on the surface of the seafloor.
4150:
Craske, A. J.; Jefferies, R. P. S. (1989). "A new mitrate from the Upper
Ordovician of Norway, and a new approach to subdividing a plesion".
1325:
and over 10M years older than the
Burgess Shale, convinced him that this specimen had similar legs. He considered the legs of these two
472:, but these fossils were poorly preserved, and Whittington did not feel they provided enough information to be classified as members of the
2703:
Chen, J-Y.; Ramsköld, L.; Gui-qing Zhou, G-Q. (May 1994). "Evidence for
Monophyly and Arthropod Affinity of Cambrian Giant Predators".
2245:"Making sense of 'lower' and 'upper' stem-group Euarthropoda, with comments on the strict use of the name Arthropoda von Siebold, 1848"
710:(1994) interpreted them as contained within the lobes along the sides. Budd (1996) thought the "triangles" were too wide to fit within
1351:
of arthropods and what he considered to be their evolutionary basal members. One striking feature of this family tree is that modern
611:
formed a U-bend on its way towards the rear of the animal. The proboscis appeared sufficiently long and flexible to reach the mouth.
3657:
Anderson, Evan P.; Schiffbauer, James D.; Jacquet, Sarah M.; Lamsdell, James C.; Kluessendorf, Joanne; Mikulic, Donald G. (2021).
4496:
4554:
4294:
2107:"The morphology and phylogenetic significance of Kerygmachela kierkegaardi Budd (Buen Formation, Lower Cambrian, N Greenland)"
1413:
stalked eyes, dorsal segmentation, posterior mouth, fused appendages, gill-like limb branches) are taxonomically significant.
4051:
3037:"The "evolution" of Anomalocaris and its classification in the arthropod class Dinocarida (nov.) and order Radiodonta (nov.)"
2418:
2033:
2841:
Simonetta AM. 1970 Studies on non trilobite arthropods of the
Burgess Shale (Middle Cambrian). Palaeontogr. Ital. 66, 35–45.
1570:, considered that Early Cambrian life was much more disparate and "experimental" than any later set of animals and that the
1968:
Caron, Jean-Bernard; Jackson, Donald A. (October 2006). "Taphonomy of the
Greater Phyllopod Bed community, Burgess Shale".
4559:
4039:
452:
of some specimens and photographs of these specimens lit from a variety of angles. Whittington's analysis did not cover
134:
1466:. The stem-arthropod was actually first described in 2008, but at the time it was originally considered a specimen of
3950:
579:, whose total length was about one-third that of the body, and projected down from under the head. The proboscis was
3978:. Paleontological Society Papers. Vol. 10. New Haven, CT: Yale University Reprographics & Imaging Service;
1275:
647:
A: Whittington (1975), B: Bergström (1986), C: Budd (1996), D: Zhang & Briggs (2007), E: Budd & Daley (2011)
3827:
Briggs, Derek E. G.; Lieberman, Bruce S.; Hendricks, Jonathan R.; Halgedahl, Susan L.; Jarrard, Richard D. (2008).
3778:
Pates, Stephen; Wolfe, Joanna M.; Lerosey-Aubril, Rudy; Daley, Allison C.; Ortega-Hernández, Javier (2022-02-09).
1761:
788:
780:
515:
that was much better preserved than previous specimens, leading them to conclude that it was a close relative of
388:
was regarded as strong evidence for the "explosive" hypothesis. Later the discovery of a whole series of similar
4544:
2977:
Budd, G. E. (1997). "Stem Group Arthropods from the Lower Cambrian Sirius Passet Fauna of North Greenland". In
1448:
567:
looked so strange that the audience at the first presentation of Whittington's analysis laughed. The length of
3933:
Eldredge, N.; Gould, S. J. (1989). "APPENDIX: Punctuated Equilibria: An Alternative to Phyletic Gradualism".
1560:, cannot be accommodated in modern groups. This was one of the primary reasons why Gould in his book on the
4539:
4249:
1159:
differ, it is not surprising that the animal's classification was highly debated during the 20th century.
817:
may increase the efficiency of food digestion and intake of nutrition. Whittington (1975) believing that
1488:
4287:
3918:
Cloud, P. E. (1968). "Pre-metazoan evolution and the origins of the Metazoa.". In Drake, E. T. (ed.).
993:
1501:
1160:
409:
268:
225:
147:
3882:
Cloud, P. E. (1948). "Some problems and patterns of evolution exemplified by fossil invertebrates".
2893:
2596:
1451:, Paleontologists described a similar looking animal which was discovered in Cambrian-aged rocks of
448:
found another good specimen, and in 1975 he published a detailed description based on very thorough
718:
2757:
Vannier, Jean; Liu, Jianni; Lerosey-Aubril, Rudy; Vinther, Jakob; Daley, Allison C. (2014-05-02).
2297:"Origin and evolution of the panarthropod head – A palaeobiological and developmental perspective"
3979:
1589:
However, other discoveries and analyses soon followed, revealing similar-looking animals such as
550:
456:; Walcott's specimens of this species could not be identified in his collection. In 1960 Russian
3542:
Siveter, Derek J.; Briggs, Derek E. G.; Siveter, David J.; Sutton, Mark D.; Legg, David (2018).
2998:"Anomalocaris and other large animals in the lower Cambrian Chengjiang fauna of southwest China"
2853:"Palaeontological and Molecular Evidence Linking Arthropods, Onychophorans, and other Ecdysozoa"
538:
2591:
1529:
3544:"A three-dimensionally preserved lobopodian from the Herefordshire (Silurian) Lagerstätte, UK"
1613:, which are regarded as close relatives of arthropods. Paleontologists defined a group called
4534:
4513:
4280:
3658:
1566:
1258:
683:
249:
4023:
Glaessner, M. F. (1972). "Precambrian palaeozoology". In Jones, J. B.; McGowran, B. (eds.).
3195:
2672:"Phylogeny of the Myriapoda-Crustacea-Insecta: a new attempt using photoreceptor structure*"
2588:
Originations, Radiations and Biodiversity Changes – evidences from the Chinese fossil record
3840:
3729:
3674:
3607:
3433:
3370:
3307:
3250:
3152:
3048:
2943:
2905:
2770:
2712:
2456:
1977:
1942:
1900:
1376:
1230:
1183:
1175:
445:
3090:
Legg, David A.; Sutton, Mark D.; Edgecombe, Gregory D.; Caron, Jean-Bernard (2012-12-07).
121:
8:
4480:
1741:
392:
animals, some with closer resemblances to arthropods, and the development of the idea of
3844:
3733:
3678:
3611:
3437:
3374:
3359:"Anomalocaridid trunk limb homology revealed by a giant filter-feeder with paired flaps"
3311:
3254:
3156:
3052:
2947:
2909:
2774:
2716:
2460:
1981:
1946:
1904:
1435:
592:
424:, and published a description of all of these in 1912. The generic name is derived from
358:
4222:
4193:
4129:
4086:
3899:
3864:
3804:
3779:
3755:
3698:
3639:
3568:
3543:
3519:
3484:
3465:
3402:
3339:
3176:
3116:
3091:
3072:
2959:
2804:
2736:
2647:
2612:
2569:
2519:
2480:
2472:
2272:
2134:
2091:
2001:
1993:
1916:
1813:
1788:
1571:
1483:
519:—although this interpretation was later questioned by Dzik, who instead concluded that
425:
400:, but one that could be understood without assuming any unique evolutionary processes.
381:
142:
3239:"Arthropod fossil data increase congruence of morphological and molecular phylogenies"
2688:
2671:
2163:
Zhang, X.; Briggs, D. E. G. (2007). "The nature and significance of the appendages of
4201:
Briggs, D. E. G.; Whittington, H. B. (1987). "The affinities of the Cambrian animals
4121:
4117:
4047:
3991:
3983:
3946:
3907:
3856:
3809:
3759:
3747:
3718:"Exceptional multifunctionality in the feeding apparatus of a mid-Cambrian radiodont"
3702:
3690:
3643:
3631:
3623:
3573:
3524:
3506:
3469:
3457:
3449:
3406:
3394:
3386:
3343:
3331:
3323:
3276:
3268:
3219:
3168:
3121:
3076:
3064:
3017:
2874:
2808:
2796:
2788:
2728:
2652:
2634:
2515:
2484:
2414:
2406:
2370:
2326:
2318:
2264:
2226:
2180:
2126:
2029:
1818:
1525:
970:
623:
4133:
3868:
3595:
3180:
2523:
2138:
2005:
1087:
672:) over the upper surface of the body, with blades attached underneath each of them.
4263:
4218:
4189:
4113:
3938:
3891:
3848:
3799:
3791:
3737:
3682:
3615:
3563:
3555:
3514:
3496:
3441:
3378:
3315:
3258:
3211:
3160:
3111:
3103:
3056:
3009:
2963:
2951:
2913:
2864:
2778:
2740:
2720:
2683:
2642:
2624:
2565:
2511:
2464:
2360:
2308:
2276:
2256:
2222:
2176:
2118:
2087:
1985:
1950:
1908:
1808:
1800:
1533:
1242:
687:
496:
339:
in 1975 revealed its unusual features, it was thought to be unrelated to any known
305:
4103:
3485:"Cambrian suspension-feeding lobopodians and the early radiation of panarthropods"
3141:"Multi-Segmented Arthropods from the Middle Cambrian of British Columbia (Canada)"
2724:
1540:
history that was hidden by the lack of fossils. Whittington (1975) concluded that
4401:
3967:
2898:
Philosophical Transactions of the Royal Society of London. B, Biological Sciences
1582:
as so important to understanding this phenomenon that he wanted to call his book
1520:
argued in 1948 and 1968 that the process was "explosive", and in the early 1970s
1286:, suggested that the two animals were related, as they shared numerous features (
1250:
608:
503:
295:
30:
4015:
Durham, J. W. (1971). "The fossil record and the origin of the Deuterostomata".
1214:
2978:
2822:
1989:
1521:
940:
584:
457:
437:
429:
3942:
3780:"New opabiniid diversifies the weirdest wonders of the euarthropod stem group"
3619:
3501:
3421:
3358:
3295:
3060:
3013:
2869:
2852:
2629:
2468:
2313:
2296:
2122:
2111:
Earth and Environmental Science Transactions of the Royal Society of Edinburgh
1752: – Set of morphological features common to members of a phylum of animals
1420:
and tardigrades/cycloneuralians is no longer supported, while the affinity of
1315:
1164:
301:
4528:
4361:
4327:
3987:
3971:
3860:
3751:
3694:
3627:
3510:
3453:
3390:
3327:
3272:
3223:
3172:
3068:
3021:
2878:
2792:
2638:
2322:
2130:
1602:
1561:
1517:
1463:
1372:
1312:
1257:, respectively) which were still thought to be close relatives (united under
1218:
880:
854:
805:
506:
492:
413:
377:
329:
298:
72:
3995:
3594:
Zeng, Han; Zhao, Fangchen; Niu, Kecheng; Zhu, Maoyan; Huang, Diying (2020).
2997:
2894:"The largest Cambrian animal, Anomalocaris, Burgess Shale, British-Columbia"
1023:
580:
380:
animals appeared suddenly during the Early Cambrian, in an event called the
4424:
4410:
4383:
4347:
4303:
4125:
3911:
3813:
3795:
3635:
3577:
3528:
3461:
3398:
3335:
3280:
3125:
3107:
2917:
2800:
2732:
2656:
2444:
2374:
2365:
2348:
2330:
2268:
1954:
1912:
1822:
1804:
1597:
1591:
1430:
1340:
1307:
1291:
1270:
1265:
1083:
1046:
988:
965:
677:
596:
433:
369:
363:
353:
192:
4458:
4435:
4369:
2702:
1708:
1610:
1537:
1336:
1298:
1209:
894:
814:
703:
695:
673:
627:
205:
47:
3742:
3717:
3596:"An early Cambrian euarthropod with radiodont-like raptorial appendages"
3559:
3445:
3382:
3319:
2892:
Whittington, Harry Blackmore; Briggs, Derek Ernest Gilmor (1985-05-14).
1997:
1274:, also from the Burgess Shale. Soon after that, Swedish palaeontologist
1067:
4469:
4336:
4333:
4090:
3903:
3686:
3263:
3238:
2783:
2758:
2498:
Dzik, J. (2004). "Anatomy and relationships of the Early Cambrian worm
2476:
2295:
Ortega-Hernández, Javier; Janssen, Ralf; Budd, Graham E. (2017-05-01).
2244:
1920:
1755:
1717:
1704:
1626:
1470:
This discovery could suggest there were other animals that looked like
1425:
1352:
1344:
1254:
1226:
1171:
1146:
1063:
914:
861:
737:
588:
449:
393:
389:
348:
288:
284:
92:
57:
3422:"Hallucigenia's head and the pharyngeal armature of early ecdysozoans"
3215:
2260:
1793:
Philosophical Transactions of the Royal Society B: Biological Sciences
1621:
1382:
Although Zhang and Briggs (2007) disagreed with Budd's diagnosis that
4321:
3852:
3663:, a problematic arthropod from the Llandoverian Waukesha Lagerstätte"
3237:
Legg, David A.; Sutton, Mark D.; Edgecombe, Gregory D. (2013-09-30).
3164:
2955:
2394:
Middle Cambrian Branchiopoda, Malacostraca, Trilobita and Merostomata
1749:
1614:
1575:
1360:
1320:
1191:
1168:
935:
842:
733:
600:
576:
511:
397:
317:
291:
179:
159:
97:
41:
4017:
Proceedings of the North American Paleontological Convention, Part H
3895:
865:
4451:
4376:
3296:"Hallucigenia's onychophoran-like claws and the case for Tactopoda"
2613:"An embryological perspective on the early arthropod fossil record"
1550:
1457:
1440:
1397:
1331:
1041:
728:
723:
130:
87:
82:
67:
62:
52:
4200:
4025:
Stratigraphic Problems of the Later Precambrian and Early Cambrian
3656:
396:, suggested that the Early Cambrian was a time of relatively fast
4417:
3826:
1246:
1198:
669:
524:
465:
461:
344:
102:
77:
34:
4149:
2934:
Budd, G. E. (1993). "A Cambrian gilled lobopod from Greenland".
16:
Extinct stem-arthropod species found in Cambrian fossil deposits
4315:
4272:
3357:
Van Roy, Peter; Daley, Allison C.; Briggs, Derek E. G. (2015).
2756:
1556:
1513:
1301:
found what he considered evidence of short, un-jointed legs in
1203:
441:
340:
169:
3092:"Cambrian bivalved arthropod reveals origin of arthrodization"
1707:
was introduced to cover evolutionary "aunts" and "cousins". A
1339:(velvet worms), which are regarded as the bearers of numerous
1190:
in 1970 very different to those of Hutchinson's, with lots of
898:
769:
384:, or had arisen earlier but without leaving fossils. At first
4390:
3777:
2676:
Journal of Zoological Systematics & Evolutionary Research
2209:
Budd, G. E.; Daley, A. C. (2011). "The lobes and lobopods of
1326:
473:
4077:(Onychophora), with Description of a New Cambrian Species".
2996:
Xian-Guang, Hou; Bergström, Jan; Ahlberg, Per (1995-09-01).
316:, a mouth under the head and facing backwards, and a clawed
2970:
2294:
1545:
1452:
822:
642:
Various interpretations on the flap and gill structures of
3541:
2995:
1416:
Since the 2010s, the suggested close relationship between
1389:
s "triangles" were legs, the resemblance they saw between
3922:. New Haven, Conn.: Yale University Press. pp. 1–72.
2443:
2162:
332:, where they constitute less than 0.1% of the community.
313:
3932:
3089:
2349:"The Cambrian Fossil Record and the Origin of the Phyla"
376:
In the 1970s, there was an ongoing debate about whether
3784:
Proceedings of the Royal Society B: Biological Sciences
3096:
Proceedings of the Royal Society B: Biological Sciences
1574:
was a truly dramatic event, possibly driven by unusual
4145:
4143:
4027:. Vol. 1. University of Adelaide. pp. 43–52.
1885:
Whittington, H. B. (June 1975). "The enigmatic animal
1703:
was going on in the late 20th century, the concept of
1329:
very similar to those of the Burgess Shale lobopodian
3293:
2759:"Sophisticated digestive systems in early arthropods"
2396:. Smithsonian Miscellaneous Collections, 57: 145-228.
2078:
and the reconstruction of the arthropod stem-group".
1931:, Middle Cambrian, Burgess Shale, British Columbia".
1695:
While this discussion about specific fossils such as
2696:
2413:. London: Hutchinson Radius. p. 77 and p. 189.
1889:, Middle Cambrian Burgess Shale, British Columbia".
1731:
1155:
Considering how paleontologists' reconstructions of
4140:
4044:
The First Three Billion Years of Evolution on Earth
3935:
Time Frames: The Evolution of Punctuated Equilibria
3294:Smith, Martin R.; Ortega-Hernández, Javier (2014).
3236:
1268:and Whittington published a major redescription of
668:
interpreted the flaps as part of dorsal coverings (
3356:
3200:and its implications for the origin of arthropods"
2983:Arthropod Relationships – Special Volume Series 55
2585:
1884:
1880:
1878:
1876:
1874:
1872:
1870:
1868:
1866:
1864:
1862:
1860:
1858:
1856:
1854:
1852:
2891:
2158:
2156:
2154:
2152:
2150:
2148:
1934:Philosophical Transactions of the Royal Society B
1927:Whittington, H. B. (1975). "The Enigmatic Animal
1892:Philosophical Transactions of the Royal Society B
1850:
1848:
1846:
1844:
1842:
1840:
1838:
1836:
1834:
1832:
4526:
4248:. Virtual Museum of Canada. 2011. Archived from
4022:
3483:Caron, Jean-Bernard; Aria, Cédric (2017-01-31).
2832:(2854): 1–24. doi:10.5479/si.00963801.78-2854.1.
2827:Proceedings of the United States National Museum
2543:
2541:
2539:
2537:
2535:
2533:
2242:
2437:
1536:both argued that the animal kingdom had a long
1229:and similar taxa). After his thorough analysis
1178:in 1930, providing the first reconstruction of
622:fossils differ. Since the animals did not have
4073:Robison, R. A. (January 1985). "Affinities of
4072:
3593:
3420:Smith, Martin R.; Caron, Jean-Bernard (2015).
3196:"The affinities of the cosmopolitan arthropod
2145:
1829:
4288:
4171:
4014:
3716:Moysiuk, Joseph; Caron, Jean-Bernard (2020).
3659:"Stranger than a scorpion: a reassessment of
2547:
2530:
3959:
3917:
3881:
3715:
2405:
2023:
1967:
1498:Walcott Cambrian Geology and Paleontology II
1201:and jointed limbs) which are reminiscent of
698:individualized from the triangular extension
662:cross-section based on Budd and Daley (2011)
4046:. Princeton University Press. p. 192.
3193:
2933:
2929:
2927:
2346:
2342:
2340:
2073:
2069:
2067:
2065:
1926:
1477:
1305:. His examination of the gilled lobopodian
1249:and arthropods, two distinct animal phyla (
4295:
4281:
3419:
2663:
2204:
2202:
2200:
2198:
2196:
2194:
2192:
2190:
2063:
2061:
2059:
2057:
2055:
2053:
2051:
2049:
2047:
2045:
2028:. Hutchinson Radius. pp. 124–136 ff.
1474:and its family may have been more diverse.
1225:belonged to the so-called "trilobitoids" (
120:
3803:
3741:
3567:
3518:
3500:
3482:
3262:
3115:
2868:
2850:
2782:
2687:
2646:
2628:
2595:
2364:
2312:
2213:from the middle Cambrian Burgess Shale".
2208:
2167:from the Middle Cambrian Burgess Shale".
1812:
1758: – Extinct order of basal arthropods
343:, or perhaps a relative of arthropod and
3965:
2924:
2497:
2337:
1961:
1620:
1609:, was considered very similar to modern
1487:
1396:s lobe+gill arrangement and arthropods'
682:
502:In 1997, Briggs and Nedin reported from
320:that probably passed food to the mouth.
4550:Taxa named by Charles Doolittle Walcott
3034:
2610:
2187:
2074:Budd, G. E. (1996). "The morphology of
2042:
1182:as an anostracan swimming upside down.
403:
335:When the first thorough examination of
4527:
3829:"Middle Cambrian arthropods from Utah"
3194:Legg, David A.; Vannier, Jean (2013).
2669:
1786:
756:remain as a plausible interpretation.
491:specimens found so far come from the "
460:described specimens they found in the
4276:
4037:
3773:
3771:
3769:
3589:
3587:
2823:Restudy of some Burgess shale fossils
2752:
2750:
2581:
2579:
2388:
2386:
2384:
2301:Arthropod Structure & Development
2290:
2288:
2286:
618:Interpretations of other features of
4268:, with photo of Burgess Shale fossil
4172:Bergström, J. (1987). "The Cambrian
3937:. By Eldredge, N. pp. 193–224.
3138:
2976:
2238:
2236:
2104:
2019:
2017:
2015:
1782:
1780:
1778:
1776:
1424:as a stem-group arthropod alongside
2353:Integrative and Comparative Biology
1278:, noting in 1986 the similarity of
420:and a few of what he classified as
13:
4223:10.1111/j.1502-3931.1987.tb02036.x
4194:10.1111/j.1502-3931.1987.tb02037.x
4165:
3766:
3584:
2821:Hutchinson, George Evelyn (1930).
2747:
2576:
2570:10.1111/j.1502-3931.1986.tb00738.x
2381:
2283:
2092:10.1111/j.1502-3931.1996.tb01831.x
1787:Briggs, Derek E. G. (2015-04-19).
1363:relatives. On the other hand, Hou
1174:in 1912. The idea was followed by
14:
4571:
4230:
2857:Evolution: Education and Outreach
2689:10.1046/j.1439-0469.2000.383152.x
2243:Ortega-Hernández, Javier (2016).
2233:
2012:
1773:
1186:provided a new reconstruction of
828:
779:on the seafloor along with other
4509:
4508:
4495:
4302:
4118:10.1046/j.1525-142x.2000.00077.x
3974:; Waggoner, Benjamin M. (eds.).
2611:Chipman, Ariel D. (2015-12-18).
2516:10.1111/j.1463-6409.2004.00136.x
2227:10.1111/j.1502-3931.2011.00264.x
2181:10.1111/j.1502-3931.2007.00013.x
1734:
1605:. Another Burgess Shale animal,
1400:limbs led them to conclude that
1086:
1066:
1045:
1022:
992:
969:
939:
917:
897:
864:
794:Suggested proboscis mobility of
787:
768:
652:
635:
549:
537:
416:nine almost complete fossils of
146:
45:
4097:
4066:
4031:
4008:
3926:
3875:
3820:
3709:
3650:
3535:
3476:
3413:
3350:
3287:
3230:
3187:
3132:
3083:
3028:
2989:
2885:
2844:
2835:
2815:
2604:
2556:, unique Cambrian arthropods".
2491:
2427:
2399:
1762:Paleobiota of the Burgess Shale
495:" of the Burgess Shale, in the
2851:Edgecombe, Gregory D. (2009).
2098:
1:
4555:Fossil taxa described in 1912
2725:10.1126/science.264.5163.1304
1767:
1492:Top left: retouched image of
1141:Summarized phylogeny between
775:Ecological reconstruction of
626:armor nor even tough organic
530:
482:
4246:Burgess Shale Fossil Gallery
1217:, following earlier work by
781:fauna from the Burgess Shale
759:
7:
1727:
1595:from the Burgess Shale and
1241:s classification above the
328:are known from the Greater
304:(505 million years ago) of
10:
4576:
4560:Cambrian genus extinctions
3548:Royal Society Open Science
2985:. Systematics Association.
1990:10.2110/palo.2003.P05-070R
1528:developed their theory of
1481:
4504:
4493:
4468:
4434:
4400:
4356:
4345:
4310:
4106:Evolution and Development
3966:Bengtson, Stefan (2004).
3943:10.1515/9781400860296.193
3920:Evolution and Environment
3620:10.1038/s41586-020-2883-7
3502:10.1186/s12862-016-0858-y
3061:10.1017/S0022336000023362
3035:Collins, Desmond (1996).
3014:10.1080/11035899509546213
2870:10.1007/s12052-009-0118-3
2630:10.1186/s12862-015-0566-z
2469:10.1017/S0022336000038919
2314:10.1016/j.asd.2016.10.011
2123:10.1017/S0263593300002418
1502:Charles Doolittle Walcott
1462:and was found within the
1167:, considered it to be an
1161:Charles Doolittle Walcott
1081:
1061:
1038:
1015:
1007:
985:
962:
954:
933:
912:
892:
885:
859:
847:
717:s slender body, and that
410:Charles Doolittle Walcott
255:
248:
143:Scientific classification
141:
128:
119:
23:
3968:"Early Skeletal Fossils"
3489:BMC Evolutionary Biology
2981:; Thomas, R. H. (eds.).
2617:BMC Evolutionary Biology
2105:Budd, Graham E. (1998).
1578:mechanisms. He regarded
1478:Theoretical significance
4079:Journal of Paleontology
3980:Paleontological Society
3833:Journal of Paleontology
3145:Journal of Paleontology
3041:Journal of Paleontology
2548:Bergström, J. (1986). "
2449:Journal of Paleontology
1455:. The fossil was named
1434:and its relatives) and
1428:(a clade that includes
1371:is a member of unusual
1367:(1995, 2006) suggested
1323: million years ago
3796:10.1098/rspb.2021.2093
3108:10.1098/rspb.2012.1958
2918:10.1098/rstb.1985.0096
2670:Paulus, H. F. (2000).
1955:10.1098/rstb.1975.0033
1913:10.1098/rstb.1975.0033
1805:10.1098/rstb.2014.0313
1692:
1530:punctuated equilibrium
1505:
699:
690:(yellow highlight) of
4545:Burgess Shale fossils
4038:Knoll, A. H. (2004).
3243:Nature Communications
2763:Nature Communications
2392:WALCOTT, C. D. 1912.
2024:Gould, S. J. (1989).
1716:concept, for example
1624:
1491:
1355:(water bears) may be
804:The way in which the
686:
499:of British Columbia.
454:Opabinia ? media
422:Opabinia ? media
4264:Smithsonian page on
3661:Parioscorpio venator
3139:Legg, David (2013).
2590:. pp. 139–158.
2366:10.1093/icb/43.1.157
2347:Budd, G. E. (2003).
1379:arthropod features.
1231:Harry B. Whittington
1176:G. Evelyn Hutchinson
446:Harry B. Whittington
440:, British Columbia,
404:History of discovery
137:in Washington, D.C.
133:) on display at the
4540:Cambrian arthropods
3845:2008JPal...82..238B
3743:10.1017/pab.2021.19
3734:2021Pbio...47..704M
3679:2021Palgy..64..429A
3612:2020Natur.588..101Z
3560:10.1098/rsos.172101
3446:10.1038/nature14573
3438:2015Natur.523...75S
3383:10.1038/nature14256
3375:2015Natur.522...77V
3320:10.1038/nature13576
3312:2014Natur.514..363S
3255:2013NatCo...4.2485L
3157:2013JPal...87..493L
3102:(1748): 4699–4704.
3053:1996JPal...70..280C
2948:1993Natur.364..709B
2910:1985RSPTB.309..569W
2775:2014NatCo...5.3641V
2717:1994Sci...264.1304C
2711:(5163): 1304–1308.
2461:1997JPal...71...22B
1982:2006Palai..21..451C
1947:1975RSPTB.271....1W
1905:1975RSPTB.271....1W
1742:Paleontology portal
1678: = Crown group
1670: = Total group
694:, showing pairs of
487:All the recognized
357:and relatives) and
3790:(1968): 20212093.
3687:10.1111/pala.12534
3264:10.1038/ncomms3485
2784:10.1038/ncomms4641
2249:Biological Reviews
1799:(1666): 20140313.
1693:
1686: = Stem group
1662: = Crown node
1654: = Basal node
1645:= Lines of descent
1584:Homage to Opabinia
1572:Cambrian explosion
1506:
1484:Cambrian explosion
1436:gilled lobopodians
1199:dorsal exoskeleton
700:
593:gilled lobopodians
509:a new specimen of
382:Cambrian explosion
359:gilled lobopodians
4522:
4521:
4491:
4490:
4053:978-0-691-12029-4
3606:(7836): 101–105.
3306:(7522): 363–366.
3216:10.1111/let.12032
2942:(6439): 709–711.
2904:(1141): 569–609.
2504:Zoologica Scripta
2420:978-0-09-174271-3
2261:10.1111/brv.12168
2035:978-0-09-174271-3
1925:Free abstract at
1899:(910): 1–43 271.
1526:Stephen Jay Gould
1359:s closest living
1184:Alberto Simonetta
1153:
1152:
1135:
1134:
1126:
1125:
1117:
1116:
1108:
1107:
1099:
1098:
883:
857:
845:
470:Opabinia norilica
276:
275:
229:
129:Fossil specimen (
4567:
4512:
4511:
4499:
4354:
4353:
4297:
4290:
4283:
4274:
4273:
4260:
4258:
4257:
4240:Opabinia regalis
4226:
4197:
4160:
4159:
4147:
4138:
4137:
4101:
4095:
4094:
4070:
4064:
4063:
4061:
4060:
4040:"Cambrian Redux"
4035:
4029:
4028:
4020:
4012:
4006:
4005:
4003:
4002:
3963:
3957:
3956:
3930:
3924:
3923:
3915:
3879:
3873:
3872:
3853:10.1666/06-086.1
3824:
3818:
3817:
3807:
3775:
3764:
3763:
3745:
3713:
3707:
3706:
3654:
3648:
3647:
3591:
3582:
3581:
3571:
3539:
3533:
3532:
3522:
3504:
3480:
3474:
3473:
3417:
3411:
3410:
3354:
3348:
3347:
3291:
3285:
3284:
3266:
3234:
3228:
3227:
3191:
3185:
3184:
3165:10.1666/12-112.1
3136:
3130:
3129:
3119:
3087:
3081:
3080:
3032:
3026:
3025:
2993:
2987:
2986:
2974:
2968:
2967:
2956:10.1038/364709a0
2931:
2922:
2921:
2889:
2883:
2882:
2872:
2848:
2842:
2839:
2833:
2819:
2813:
2812:
2786:
2754:
2745:
2744:
2700:
2694:
2693:
2691:
2667:
2661:
2660:
2650:
2632:
2608:
2602:
2601:
2599:
2583:
2574:
2573:
2545:
2528:
2527:
2495:
2489:
2488:
2445:Briggs, D. E. G.
2441:
2435:
2431:
2425:
2424:
2403:
2397:
2390:
2379:
2378:
2368:
2344:
2335:
2334:
2316:
2292:
2281:
2280:
2240:
2231:
2230:
2211:Opabinia regalis
2206:
2185:
2184:
2160:
2143:
2142:
2102:
2096:
2095:
2076:Opabinia regalis
2071:
2040:
2039:
2021:
2010:
2009:
1965:
1959:
1958:
1929:Opabinia regalis
1924:
1887:Opabinia regalis
1882:
1827:
1826:
1816:
1784:
1744:
1739:
1738:
1737:
1687:
1685:
1679:
1677:
1671:
1669:
1663:
1661:
1655:
1653:
1647:
1646:
1642:
1641:
1640:
1534:Martin Glaessner
1409:and arthropods (
1395:
1388:
1341:ancestral traits
1324:
1261:) at that time.
1090:
1070:
1049:
1026:
1010:
1009:
996:
973:
957:
956:
943:
921:
901:
888:
887:
879:
868:
853:
850:
849:
841:
838:
837:
833:
832:
791:
772:
746:
716:
688:Digestive system
656:
644:Opabinia regalis
639:
569:Opabinia regalis
553:
541:
497:Canadian Rockies
444:. In 1966–1967,
418:Opabinia regalis
306:British Columbia
280:Opabinia regalis
264:
262:Opabinia regalis
260:
224:
217:
204:
191:
151:
150:
124:
114:
44:
29:Temporal range:
21:
20:
4575:
4574:
4570:
4569:
4568:
4566:
4565:
4564:
4525:
4524:
4523:
4518:
4500:
4487:
4464:
4430:
4402:Kerygmachelidae
4396:
4350:
4341:
4306:
4301:
4255:
4253:
4236:
4233:
4168:
4166:Further reading
4163:
4148:
4141:
4102:
4098:
4071:
4067:
4058:
4056:
4054:
4036:
4032:
4013:
4009:
4000:
3998:
3964:
3960:
3953:
3931:
3927:
3896:10.2307/2405523
3880:
3876:
3825:
3821:
3776:
3767:
3714:
3710:
3655:
3651:
3592:
3585:
3540:
3536:
3481:
3477:
3432:(7558): 75–78.
3418:
3414:
3369:(7554): 77–80.
3355:
3351:
3292:
3288:
3235:
3231:
3192:
3188:
3137:
3133:
3088:
3084:
3033:
3029:
2994:
2990:
2975:
2971:
2932:
2925:
2890:
2886:
2849:
2845:
2840:
2836:
2820:
2816:
2755:
2748:
2701:
2697:
2668:
2664:
2609:
2605:
2597:10.1.1.693.5869
2584:
2577:
2546:
2531:
2496:
2492:
2442:
2438:
2432:
2428:
2421:
2404:
2400:
2391:
2382:
2345:
2338:
2293:
2284:
2241:
2234:
2207:
2188:
2161:
2146:
2103:
2099:
2072:
2043:
2036:
2022:
2013:
1966:
1962:
1883:
1830:
1785:
1774:
1770:
1740:
1735:
1733:
1730:
1691:
1690:
1683:
1682:
1675:
1674:
1667:
1666:
1659:
1658:
1651:
1650:
1644:
1636:
1635:
1634:
1633:
1486:
1480:
1458:Utaurora comosa
1393:
1386:
1335:and the modern
1319:
1292:a single phylum
1251:Lophotrochozoan
1233:concluded that
1221:, thought that
1163:, the original
1136:
1127:
1118:
1109:
1100:
831:
802:
801:
800:
799:
798:
792:
784:
783:
773:
762:
744:
714:
663:
657:
648:
646:
640:
609:digestive tract
561:
557:
556:Size estimation
554:
545:
542:
533:
504:South Australia
485:
458:paleontologists
436:, southeast of
406:
296:Middle Cambrian
272:
266:
258:
257:
244:
241:O. regalis
223:
215:
202:
189:
145:
115:
113:
112:
111:
110:
105:
100:
95:
90:
85:
80:
75:
70:
65:
60:
55:
50:
39:
38:
31:Middle Cambrian
27:
17:
12:
11:
5:
4573:
4563:
4562:
4557:
4552:
4547:
4542:
4537:
4520:
4519:
4517:
4516:
4505:
4502:
4501:
4494:
4492:
4489:
4488:
4486:
4485:
4484:
4483:
4474:
4472:
4466:
4465:
4463:
4462:
4455:
4448:
4440:
4438:
4432:
4431:
4429:
4428:
4421:
4414:
4406:
4404:
4398:
4397:
4395:
4394:
4387:
4380:
4373:
4365:
4357:
4351:
4346:
4343:
4342:
4340:
4339:
4330:
4324:
4318:
4311:
4308:
4307:
4300:
4299:
4292:
4285:
4277:
4271:
4270:
4261:
4232:
4231:External links
4229:
4228:
4227:
4217:(2): 185–186.
4198:
4188:(2): 187–188.
4167:
4164:
4162:
4161:
4139:
4112:(6): 340–347.
4096:
4085:(1): 226–235.
4065:
4052:
4030:
4007:
3972:Lipps, Jere H.
3958:
3951:
3925:
3890:(4): 322–350.
3874:
3839:(2): 238–254.
3819:
3765:
3728:(4): 704–724.
3708:
3673:(3): 429–474.
3649:
3583:
3534:
3475:
3412:
3349:
3286:
3229:
3210:(4): 540–550.
3186:
3151:(3): 493–501.
3131:
3082:
3047:(2): 280–293.
3027:
3008:(3): 163–183.
2988:
2969:
2923:
2884:
2863:(2): 178–190.
2843:
2834:
2814:
2746:
2695:
2682:(3): 189–208.
2662:
2603:
2575:
2564:(3): 241–246.
2529:
2490:
2436:
2426:
2419:
2411:Wonderful Life
2398:
2380:
2359:(1): 157–165.
2336:
2307:(3): 354–379.
2282:
2255:(1): 255–273.
2232:
2186:
2175:(2): 161–173.
2144:
2117:(4): 249–290.
2097:
2041:
2034:
2026:Wonderful Life
2011:
1960:
1828:
1771:
1769:
1766:
1765:
1764:
1759:
1753:
1746:
1745:
1729:
1726:
1689:
1688:
1680:
1672:
1664:
1656:
1648:
1630:
1629:
1567:Wonderful Life
1522:Niles Eldredge
1482:Main article:
1479:
1476:
1373:cycloneuralian
1151:
1150:
1138:
1137:
1133:
1132:
1129:
1128:
1124:
1123:
1120:
1119:
1115:
1114:
1111:
1110:
1106:
1105:
1102:
1101:
1097:
1096:
1093:
1092:
1080:
1077:
1076:
1073:
1072:
1060:
1056:
1055:
1052:
1051:
1037:
1033:
1032:
1029:
1028:
1014:
1008:
1006:
1003:
1002:
999:
998:
984:
980:
979:
976:
975:
961:
955:
953:
950:
949:
946:
945:
932:
928:
927:
924:
923:
911:
908:
907:
904:
903:
891:
886:
884:
876:
875:
872:
871:
869:and relatives
858:
848:
846:
836:
830:
829:Classification
827:
793:
786:
785:
774:
767:
766:
765:
764:
763:
761:
758:
726:limbs seen in
665:
664:
658:
651:
649:
641:
634:
607:, so that the
585:vacuum cleaner
575:is the hollow
559:
558:
555:
548:
546:
543:
536:
532:
529:
484:
481:
430:Mount Hungabee
405:
402:
274:
273:
267:
253:
252:
246:
245:
237:
235:
231:
230:
213:
209:
208:
200:
196:
195:
187:
183:
182:
177:
173:
172:
167:
163:
162:
157:
153:
152:
139:
138:
126:
125:
117:
116:
108:
107:
106:
101:
96:
91:
86:
81:
76:
71:
66:
61:
56:
51:
46:
28:
15:
9:
6:
4:
3:
2:
4572:
4561:
4558:
4556:
4553:
4551:
4548:
4546:
4543:
4541:
4538:
4536:
4533:
4532:
4530:
4515:
4507:
4506:
4503:
4498:
4482:
4478:
4477:
4476:
4475:
4473:
4471:
4467:
4461:
4460:
4456:
4454:
4453:
4449:
4447:
4446:
4442:
4441:
4439:
4437:
4433:
4427:
4426:
4422:
4420:
4419:
4415:
4413:
4412:
4408:
4407:
4405:
4403:
4399:
4393:
4392:
4388:
4386:
4385:
4381:
4379:
4378:
4374:
4372:
4371:
4366:
4364:
4363:
4362:Caryosyntrips
4359:
4358:
4355:
4352:
4349:
4344:
4338:
4335:
4331:
4329:
4328:Panarthropoda
4325:
4323:
4320:Superphylum:
4319:
4317:
4313:
4312:
4309:
4305:
4298:
4293:
4291:
4286:
4284:
4279:
4278:
4275:
4269:
4267:
4262:
4252:on 2020-11-12
4251:
4247:
4243:
4241:
4235:
4234:
4224:
4220:
4216:
4212:
4208:
4204:
4199:
4195:
4191:
4187:
4183:
4179:
4175:
4170:
4169:
4157:
4153:
4152:Palaeontology
4146:
4144:
4135:
4131:
4127:
4123:
4119:
4115:
4111:
4107:
4100:
4092:
4088:
4084:
4080:
4076:
4069:
4055:
4049:
4045:
4041:
4034:
4026:
4018:
4011:
3997:
3993:
3989:
3985:
3981:
3977:
3973:
3969:
3962:
3954:
3952:9781400860296
3948:
3944:
3940:
3936:
3929:
3921:
3913:
3909:
3905:
3901:
3897:
3893:
3889:
3885:
3878:
3870:
3866:
3862:
3858:
3854:
3850:
3846:
3842:
3838:
3834:
3830:
3823:
3815:
3811:
3806:
3801:
3797:
3793:
3789:
3785:
3781:
3774:
3772:
3770:
3761:
3757:
3753:
3749:
3744:
3739:
3735:
3731:
3727:
3723:
3719:
3712:
3704:
3700:
3696:
3692:
3688:
3684:
3680:
3676:
3672:
3668:
3667:Palaeontology
3664:
3662:
3653:
3645:
3641:
3637:
3633:
3629:
3625:
3621:
3617:
3613:
3609:
3605:
3601:
3597:
3590:
3588:
3579:
3575:
3570:
3565:
3561:
3557:
3554:(8): 172101.
3553:
3549:
3545:
3538:
3530:
3526:
3521:
3516:
3512:
3508:
3503:
3498:
3494:
3490:
3486:
3479:
3471:
3467:
3463:
3459:
3455:
3451:
3447:
3443:
3439:
3435:
3431:
3427:
3423:
3416:
3408:
3404:
3400:
3396:
3392:
3388:
3384:
3380:
3376:
3372:
3368:
3364:
3360:
3353:
3345:
3341:
3337:
3333:
3329:
3325:
3321:
3317:
3313:
3309:
3305:
3301:
3297:
3290:
3282:
3278:
3274:
3270:
3265:
3260:
3256:
3252:
3248:
3244:
3240:
3233:
3225:
3221:
3217:
3213:
3209:
3205:
3201:
3199:
3190:
3182:
3178:
3174:
3170:
3166:
3162:
3158:
3154:
3150:
3146:
3142:
3135:
3127:
3123:
3118:
3113:
3109:
3105:
3101:
3097:
3093:
3086:
3078:
3074:
3070:
3066:
3062:
3058:
3054:
3050:
3046:
3042:
3038:
3031:
3023:
3019:
3015:
3011:
3007:
3003:
2999:
2992:
2984:
2980:
2979:Fortey, R. A.
2973:
2965:
2961:
2957:
2953:
2949:
2945:
2941:
2937:
2930:
2928:
2919:
2915:
2911:
2907:
2903:
2899:
2895:
2888:
2880:
2876:
2871:
2866:
2862:
2858:
2854:
2847:
2838:
2831:
2828:
2824:
2818:
2810:
2806:
2802:
2798:
2794:
2790:
2785:
2780:
2776:
2772:
2768:
2764:
2760:
2753:
2751:
2742:
2738:
2734:
2730:
2726:
2722:
2718:
2714:
2710:
2706:
2699:
2690:
2685:
2681:
2677:
2673:
2666:
2658:
2654:
2649:
2644:
2640:
2636:
2631:
2626:
2622:
2618:
2614:
2607:
2598:
2593:
2589:
2582:
2580:
2571:
2567:
2563:
2559:
2555:
2551:
2544:
2542:
2540:
2538:
2536:
2534:
2525:
2521:
2517:
2513:
2509:
2505:
2501:
2494:
2486:
2482:
2478:
2474:
2470:
2466:
2462:
2458:
2454:
2450:
2446:
2440:
2430:
2422:
2416:
2412:
2408:
2402:
2395:
2389:
2387:
2385:
2376:
2372:
2367:
2362:
2358:
2354:
2350:
2343:
2341:
2332:
2328:
2324:
2320:
2315:
2310:
2306:
2302:
2298:
2291:
2289:
2287:
2278:
2274:
2270:
2266:
2262:
2258:
2254:
2250:
2246:
2239:
2237:
2228:
2224:
2220:
2216:
2212:
2205:
2203:
2201:
2199:
2197:
2195:
2193:
2191:
2182:
2178:
2174:
2170:
2166:
2159:
2157:
2155:
2153:
2151:
2149:
2140:
2136:
2132:
2128:
2124:
2120:
2116:
2112:
2108:
2101:
2093:
2089:
2085:
2081:
2077:
2070:
2068:
2066:
2064:
2062:
2060:
2058:
2056:
2054:
2052:
2050:
2048:
2046:
2037:
2031:
2027:
2020:
2018:
2016:
2007:
2003:
1999:
1995:
1991:
1987:
1983:
1979:
1976:(5): 451–65.
1975:
1971:
1964:
1956:
1952:
1948:
1944:
1941:(910): 1–43.
1940:
1936:
1935:
1930:
1922:
1918:
1914:
1910:
1906:
1902:
1898:
1894:
1893:
1888:
1881:
1879:
1877:
1875:
1873:
1871:
1869:
1867:
1865:
1863:
1861:
1859:
1857:
1855:
1853:
1851:
1849:
1847:
1845:
1843:
1841:
1839:
1837:
1835:
1833:
1824:
1820:
1815:
1810:
1806:
1802:
1798:
1794:
1790:
1783:
1781:
1779:
1777:
1772:
1763:
1760:
1757:
1754:
1751:
1748:
1747:
1743:
1732:
1725:
1723:
1719:
1715:
1710:
1706:
1702:
1698:
1681:
1673:
1665:
1657:
1649:
1639:
1632:
1631:
1628:
1623:
1619:
1616:
1612:
1608:
1604:
1603:Sirius Passet
1600:
1599:
1594:
1593:
1587:
1585:
1581:
1577:
1573:
1569:
1568:
1563:
1562:Burgess Shale
1559:
1558:
1553:
1552:
1547:
1543:
1539:
1535:
1531:
1527:
1523:
1519:
1518:Preston Cloud
1515:
1510:
1503:
1499:
1495:
1490:
1485:
1475:
1473:
1469:
1468:Anomalocaris.
1465:
1464:Wheeler Shale
1461:
1459:
1454:
1450:
1445:
1443:
1442:
1437:
1433:
1432:
1427:
1423:
1419:
1414:
1412:
1408:
1403:
1399:
1392:
1385:
1380:
1378:
1374:
1370:
1366:
1362:
1358:
1354:
1350:
1346:
1342:
1338:
1337:onychophorans
1334:
1333:
1328:
1322:
1317:
1314:
1313:Sirius Passet
1310:
1309:
1304:
1300:
1295:
1293:
1289:
1285:
1281:
1277:
1276:Jan Bergström
1273:
1272:
1267:
1262:
1260:
1256:
1252:
1248:
1244:
1240:
1236:
1232:
1228:
1224:
1220:
1219:Percy Raymond
1216:
1212:
1211:
1206:
1205:
1200:
1197:
1193:
1189:
1185:
1181:
1177:
1173:
1170:
1166:
1162:
1158:
1148:
1144:
1140:
1139:
1131:
1130:
1122:
1121:
1113:
1112:
1104:
1103:
1095:
1094:
1091:
1089:
1085:
1079:
1078:
1075:
1074:
1071:
1069:
1065:
1058:
1057:
1054:
1053:
1050:
1048:
1044:
1043:
1035:
1034:
1031:
1030:
1027:
1025:
1021:
1020:
1012:
1011:
1005:
1004:
1001:
1000:
997:
995:
991:
990:
982:
981:
978:
977:
974:
972:
968:
967:
959:
958:
952:
951:
948:
947:
944:
942:
937:
930:
929:
926:
925:
922:
920:
916:
910:
909:
906:
905:
902:
900:
896:
890:
889:
882:
881:Panarthropoda
878:
877:
874:
873:
870:
867:
863:
856:
855:Cycloneuralia
852:
851:
844:
840:
839:
835:
834:
826:
824:
820:
816:
811:
807:
806:Burgess Shale
797:
790:
782:
778:
771:
757:
755:
751:
743:
739:
735:
731:
730:
725:
720:
719:cross-section
713:
709:
705:
697:
693:
689:
685:
681:
679:
675:
671:
661:
655:
650:
645:
638:
633:
632:
631:
629:
625:
621:
616:
612:
610:
606:
602:
599:, like other
598:
594:
590:
586:
582:
578:
574:
570:
566:
562:
552:
547:
540:
535:
534:
528:
526:
522:
518:
514:
513:
508:
507:Emu Bay Shale
505:
500:
498:
494:
493:Phyllopod bed
490:
480:
478:
475:
471:
468:and labelled
467:
463:
459:
455:
451:
447:
443:
439:
435:
431:
427:
423:
419:
415:
414:Burgess Shale
412:found in the
411:
401:
399:
395:
391:
387:
383:
379:
374:
372:
371:
366:
365:
360:
356:
355:
350:
346:
342:
338:
333:
331:
330:Phyllopod bed
327:
323:
319:
315:
311:
307:
303:
300:
299:Burgess Shale
297:
294:found in the
293:
290:
286:
282:
281:
270:
265:
263:
254:
251:
250:Binomial name
247:
243:
242:
236:
233:
232:
227:
222:
221:
214:
211:
210:
207:
201:
198:
197:
194:
188:
185:
184:
181:
178:
175:
174:
171:
168:
165:
164:
161:
158:
155:
154:
149:
144:
140:
136:
132:
127:
123:
118:
104:
99:
94:
89:
84:
79:
74:
69:
64:
59:
54:
49:
43:
36:
32:
26:
22:
19:
4535:Dinocaridida
4457:
4450:
4444:
4443:
4425:Mobulavermis
4423:
4416:
4411:Kerygmachela
4409:
4389:
4384:Pambdelurion
4382:
4375:
4368:
4360:
4348:Dinocaridida
4304:Dinocaridida
4265:
4254:. Retrieved
4250:the original
4245:
4239:
4214:
4210:
4206:
4203:Anomalocaris
4202:
4185:
4181:
4178:Anomalocaris
4177:
4173:
4155:
4151:
4109:
4105:
4099:
4082:
4078:
4074:
4068:
4057:. Retrieved
4043:
4033:
4024:
4019:: 1104–1132.
4016:
4010:
3999:. Retrieved
3975:
3961:
3934:
3928:
3919:
3887:
3883:
3877:
3836:
3832:
3822:
3787:
3783:
3725:
3722:Paleobiology
3721:
3711:
3670:
3666:
3660:
3652:
3603:
3599:
3551:
3547:
3537:
3492:
3488:
3478:
3429:
3425:
3415:
3366:
3362:
3352:
3303:
3299:
3289:
3246:
3242:
3232:
3207:
3203:
3197:
3189:
3148:
3144:
3134:
3099:
3095:
3085:
3044:
3040:
3030:
3005:
3001:
2991:
2982:
2972:
2939:
2935:
2901:
2897:
2887:
2860:
2856:
2846:
2837:
2829:
2826:
2817:
2766:
2762:
2708:
2704:
2698:
2679:
2675:
2665:
2620:
2616:
2606:
2587:
2561:
2557:
2554:Anomalocaris
2553:
2549:
2510:(1): 57–69.
2507:
2503:
2499:
2493:
2455:(1): 22–32.
2452:
2448:
2439:
2429:
2410:
2407:Gould, S. J.
2401:
2356:
2352:
2304:
2300:
2252:
2248:
2218:
2214:
2210:
2172:
2168:
2164:
2114:
2110:
2100:
2083:
2079:
2075:
2025:
1973:
1969:
1963:
1938:
1932:
1928:
1896:
1890:
1886:
1796:
1792:
1721:
1713:
1701:Anomalocaris
1700:
1696:
1694:
1637:
1606:
1598:Kerygmachela
1596:
1592:Anomalocaris
1590:
1588:
1583:
1579:
1576:evolutionary
1565:
1555:
1549:
1544:, and other
1541:
1508:
1507:
1497:
1493:
1471:
1467:
1456:
1446:
1439:
1431:Anomalocaris
1429:
1421:
1417:
1415:
1410:
1406:
1401:
1390:
1383:
1381:
1368:
1364:
1361:evolutionary
1356:
1348:
1330:
1308:Kerygmachela
1306:
1302:
1296:
1287:
1283:
1280:Anomalocaris
1279:
1271:Anomalocaris
1269:
1266:Derek Briggs
1263:
1238:
1234:
1222:
1215:Leif Størmer
1208:
1202:
1195:
1187:
1179:
1156:
1154:
1142:
1084:Euarthropoda
1082:
1062:
1040:
1039:
1018:
1017:
1016:
989:Kerygmachela
987:
986:
966:Pambdelurion
964:
963:
934:
913:
893:
860:
818:
809:
803:
795:
776:
753:
749:
741:
727:
711:
707:
701:
691:
666:
659:
643:
628:exoskeletons
619:
617:
613:
604:
572:
568:
564:
563:
560:
520:
516:
510:
501:
488:
486:
476:
469:
453:
434:Mount Biddle
421:
417:
407:
385:
378:multi-celled
375:
370:Pambdelurion
368:
364:Kerygmachela
362:
354:Anomalocaris
352:
336:
334:
325:
321:
309:
279:
278:
277:
261:
256:
240:
239:
219:
218:
193:Dinocaridida
24:
18:
4459:Mieridduryn
4436:Opabiniidae
4370:Parvibellus
3249:(1): 2485.
2769:(1): 3641.
2086:(1): 1–14.
1718:tardigrades
1709:crown group
1705:stem groups
1627:stem groups
1625:Concept of
1615:lobopodians
1611:Onychophora
1538:Proterozoic
1375:worms with
1353:tardigrades
1316:lagerstätte
1299:Graham Budd
1210:Leanchoilia
895:Onychophora
815:diverticula
738:crustaceans
704:diverticula
696:diverticula
624:mineralized
544:Restoration
438:Lake O'Hara
426:Opabin pass
394:stem groups
302:Lagerstätte
206:Opabiniidae
135:Smithsonian
4529:Categories
4481:Radiodonta
4470:Radiodonta
4337:Arthropoda
4334:Stem-group
4256:2023-01-21
4059:2009-04-22
4001:2015-02-06
2623:(1): 285.
1768:References
1756:Radiodonta
1426:Radiodonta
1377:convergent
1345:stem-group
1259:Articulata
1255:Ecdysozoan
1227:trilobites
1194:features (
1172:crustacean
1169:anostracan
1147:Ecdysozoan
1145:and other
1064:Radiodonta
936:Lobopodian
915:Tardigrada
862:Priapulida
734:trilobites
601:arthropods
589:radiodonts
531:Morphology
483:Occurrence
464:region of
450:dissection
390:lobopodian
349:radiodonts
289:stem group
180:Arthropoda
4322:Ecdysozoa
4314:Kingdom:
3988:1089-3326
3884:Evolution
3861:0022-3360
3760:236552819
3752:0094-8373
3703:234812878
3695:1475-4983
3644:226248177
3628:1476-4687
3511:1471-2148
3495:(1): 29.
3470:205244325
3454:1476-4687
3407:205242881
3391:1476-4687
3344:205239797
3328:1476-4687
3273:2041-1723
3224:1502-3931
3173:0022-3360
3077:131622496
3069:0022-3360
3022:1103-5897
2879:1936-6434
2809:205324774
2793:2041-1723
2639:1471-2148
2592:CiteSeerX
2500:Myoscolex
2485:131851540
2323:1467-8039
2221:: 83–95.
2131:1473-7116
1750:Body plan
1472:Opabinia,
1357:Opabinia'
1311:from the
1297:In 1996,
1264:In 1985,
1239:Opabinia'
1192:arthropod
1165:describer
843:Ecdysozoa
760:Lifestyle
605:backwards
577:proboscis
521:Myoscolex
512:Myoscolex
408:In 1911,
398:evolution
318:proboscis
292:arthropod
234:Species:
166:Kingdom:
160:Eukaryota
40:505
4514:Category
4452:Utaurora
4445:Opabinia
4377:Omnidens
4332:Phylum:
4316:Animalia
4266:Opabinia
4207:Opabinia
4174:Opabinia
4158:: 69–99.
4134:25274057
4126:11256378
4075:Aysheaia
3996:57481790
3912:18122310
3869:31568651
3814:35135344
3636:33149303
3578:30224988
3529:28137244
3462:26106857
3399:25762145
3336:25132546
3281:24077329
3181:86725173
3126:23055069
2801:24785191
2733:17780848
2657:26678148
2550:Opabinia
2524:85216629
2409:(1990).
2375:21680420
2331:27989966
2269:25528950
2165:Opabinia
2139:85645934
2006:53646959
1998:20173022
1823:25750235
1728:See also
1722:Opabinia
1714:relative
1697:Opabinia
1607:Aysheaia
1580:Opabinia
1551:Marrella
1548:such as
1542:Opabinia
1509:Opabinia
1494:Opabinia
1441:Kylinxia
1422:Opabinia
1418:Opabinia
1407:Opabinia
1402:Opabinia
1398:biramous
1391:Opabinia
1384:Opabinia
1369:Opabinia
1332:Aysheaia
1318:, about
1303:Opabinia
1284:Opabinia
1247:annelids
1235:Opabinia
1223:Opabinia
1188:Opabinia
1180:Opabinia
1157:Opabinia
1143:Opabinia
1042:Utaurora
1019:Opabinia
819:Opabinia
810:Opabinia
796:Opabinia
777:Opabinia
754:Opabinia
750:Opabinia
742:Opabinia
729:Marrella
724:biramous
712:Opabinia
706:). Chen
692:Opabinia
660:Opabinia
620:Opabinia
597:compound
581:striated
573:Opabinia
565:Opabinia
517:Opabinia
489:Opabinia
477:Opabinia
428:between
386:Opabinia
337:Opabinia
326:Opabinia
322:Opabinia
310:Opabinia
220:Opabinia
199:Family:
176:Phylum:
170:Animalia
156:Domain:
131:holotype
25:Opabinia
4418:Utahnax
4326:Clade:
4211:Lethaia
4182:Lethaia
4091:1304837
3904:2405523
3841:Bibcode
3805:8826304
3730:Bibcode
3675:Bibcode
3608:Bibcode
3569:6124121
3520:5282736
3434:Bibcode
3371:Bibcode
3308:Bibcode
3251:Bibcode
3204:Lethaia
3153:Bibcode
3117:3497099
3049:Bibcode
2964:4341971
2944:Bibcode
2906:Bibcode
2771:Bibcode
2741:1913482
2713:Bibcode
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2648:4683962
2558:Lethaia
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2457:Bibcode
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2215:Lethaia
2169:Lethaia
2080:Lethaia
1978:Bibcode
1970:PALAIOS
1943:Bibcode
1921:2417412
1901:Bibcode
1814:4360120
1514:animals
670:tergite
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525:annelid
523:was an
466:Siberia
462:Norilsk
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285:extinct
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