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1

Trott, Thomas J. "Gustatory responses ofpriapulus caudatus delamarck, 1816 (priapulida, priapulidae): Feeding behavior and chemoreception by a living fossil." Marine and Freshwater Behaviour and Physiology 31, no. 4 (1998): 251–57. http://dx.doi.org/10.1080/10236249809387076.

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2

Ferraguti, Marco, and Claudio Garbelli. "The spermatozoon of a ‘living fossil’: Tubiluchus troglodytes (Priapulida)." Tissue and Cell 38, no. 1 (2006): 1–6. http://dx.doi.org/10.1016/j.tice.2005.05.001.

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3

HU, SHI-XUE, MAO-YAN ZHU, FANG-CHEN ZHAO, and MICHAEL STEINER. "A crown group priapulid from the early Cambrian Guanshan Lagerstätte." Geological Magazine 154, no. 6 (2017): 1329–33. http://dx.doi.org/10.1017/s001675681700019x.

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AbstractA well-preserved fossil priapulid worm, Xiaoheiqingella sp., is reported from the early Cambrian Guanshan Lagerstätte (Cambrian Series II, Stage 4) near Kunming City, Yunnan Province, SW China. The body of the animal consists of four sections: a swollen introvert, a constricted neck, a finely annulated trunk and a caudal appendage. The body configuration exhibits a close resemblance to that of the crown group priapulid Xiaoheiqingella peculiaris from the early Cambrian Chengjiang Lagerstätte. The new discovery provides another striking example of crown group priapulids, representing th
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4

Ma, Xiaoya, Richard J. Aldridge, David J. Siveter, Derek J. Siveter, Xianguang Hou, and Gregory D. Edgecombe. "A New Exceptionally Preserved Cambrian Priapulid from the Chengjiang Lagerstätte." Journal of Paleontology 88, no. 2 (2014): 371–84. http://dx.doi.org/10.1666/13-082.

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A fossil priapulid, Eximipriapulus globocaudatus new genus new species, is described from the Cambrian Chengjiang Lagerstätte of Yunnan, China. The exceptional preservation of the animal reveals morphological details that allow direct comparison with extant priapulids. The body is divisible into a partially eversible pharynx, a smooth collar, a scalid-bearing introvert, a neck with triangular scalids, an unsegmented trunk with annulations, and a distinctly expanded terminal region. Several specialized regions of the alimentary canal are recognized: a pharynx (lined by cuticle and bearing teeth
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5

Yang, Jie, Javier Ortega-Hernández, Nicholas J. Butterfield, et al. "Fuxianhuiid ventral nerve cord and early nervous system evolution in Panarthropoda." Proceedings of the National Academy of Sciences 113, no. 11 (2016): 2988–93. http://dx.doi.org/10.1073/pnas.1522434113.

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Panarthropods are typified by disparate grades of neurological organization reflecting a complex evolutionary history. The fossil record offers a unique opportunity to reconstruct early character evolution of the nervous system via exceptional preservation in extinct representatives. Here we describe the neurological architecture of the ventral nerve cord (VNC) in the upper-stem group euarthropodChengjiangocaris kunmingensisfrom the early Cambrian Xiaoshiba Lagerstätte (South China). The VNC ofC. kunmingensiscomprises a homonymous series of condensed ganglia that extend throughout the body, ea
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6

Kesidis, Giannis, Ben J. Slater, Sören Jensen, and Graham E. Budd. "Caught in the act: priapulid burrowers in early Cambrian substrates." Proceedings of the Royal Society B: Biological Sciences 286, no. 1894 (2019): 20182505. http://dx.doi.org/10.1098/rspb.2018.2505.

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The fossilized traces of burrowing worms have taken on a considerable importance in studies of the Cambrian explosion, partly because of their use in defining the base of the Cambrian. Foremost among these are the treptichnids, a group of relatively large open probing burrows that have sometimes been assigned to the activities of priapulid scalidophoran worms. Nevertheless, most Cambrian burrows have an uncertain progenitor. Here we report a suite of exceptionally preserved trace and body fossils from sandstones of the lower Cambrian (Stage 4) File Haidar Formation of southern Sweden that can
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7

MORRIS, S. CONWAY. "The cuticular structure of the 495-Myr-old type species of the fossil worm Palaeoscolex, P. piscatorum (?Priapulida)." Zoological Journal of the Linnean Society 119, no. 1 (1997): 69–82. http://dx.doi.org/10.1111/j.1096-3642.1997.tb00136.x.

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8

Wills, Matthew A. "Cambrian and Recent disparity: the picture from priapulids." Paleobiology 24, no. 2 (1998): 177–99. http://dx.doi.org/10.1666/0094-8373(1998)024[0177:cardtp]2.3.co;2.

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Abstract An understanding of several macroevolutionary trends has been greatly advanced in recent years by a focus on disparity (morphological variety) rather than taxic diversity. A seminal issue has been the nature of the Cambrian Radiation, and the question of whether problematical Cambrian fossils embody a range of anatomical design far exceeding that observed thereafter. Arthropods have hitherto furnished the only case study, revealing comparable levels of Cambrian and Recent disparity. The generality of this observation needs to be tested in other groups, and the priapulid worms provide
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9

Peel, John S. "A corset-like fossil from the Cambrian Sirius Passet Lagerstätte of North Greenland and its implications for cycloneuralian evolution." Journal of Paleontology 84, no. 2 (2010): 332–40. http://dx.doi.org/10.1666/09-102r.1.

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A large (maximum length 80 mm), tubular, corset-like problematic fossil from the early Cambrian (Cambrian Series 2, Stage 3) Sirius Passet Lagerstätte of North Greenland is interpreted as the lorica of an ancestral loriciferan. in addition to the double circlet of 7 plates composing the lorica, Sirilorica carlsbergi new genus, new species also preserves up to six multicuspidate cuticular denticles that are similar in shape to the pharyngeal teeth of priapulid worms, although their location is suggestive of scalids. Whilst traditionally placed as a sister group of priapulid worms within Vinctip
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10

Zhao, Fangchen, Jean-Bernard Caron, David J. Bottjer, Shixue Hu, Zongjun Yin, and Maoyan Zhu. "Diversity and species abundance patterns of the Early Cambrian (Series 2, Stage 3) Chengjiang Biota from China." Paleobiology 40, no. 1 (2014): 50–69. http://dx.doi.org/10.1666/12056.

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Lagerstättenfrom the Precambrian–Cambrian transition have traditionally been a relatively untapped resource for understanding the paleoecology of the “Cambrian explosion.” This quantitative paleoecological study is based on 10,238 fossil specimens belonging to 100 animal species, 11 phyla, and 15 ecological categories from the lower Cambrian (Series 2, Stage 3) Chengjiang biota (Mafang locality near Haikou, Yunnan Province, China). Fossils were systematically collected within a 2.5-meter-thick sequence divided into ten stratigraphic intervals. Each interval represents an induced time-averaged
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11

Rudkin, David M. "A possible archaeopriapulid trace fossil from the Middle Cambrian Stephen Formation, British Columbia." Paleontological Society Special Publications 6 (1992): 255. http://dx.doi.org/10.1017/s2475262200008157.

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It is generally accepted that the extraordinary preservation of fossils in the Burgess Shale is at least partly due to the exclusion of large metazoan scavengers through rapid burial in an inimical environment. Disruption of individual organisms appears to have been largely the result of mechanical processes and/or microbial decay. The absence of bioturbation through much of the classic section, including Walcott's Phyllopod Bed, supports these arguments. However, macroscopic traces are known from other horizons within the Burgess Shale section and from lateral equivalents in the Stephen Forma
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12

Telford, Maximilian J., Sarah J. Bourlat, Andrew Economou, Daniel Papillon, and Omar Rota-Stabelli. "The evolution of the Ecdysozoa." Philosophical Transactions of the Royal Society B: Biological Sciences 363, no. 1496 (2008): 1529–37. http://dx.doi.org/10.1098/rstb.2007.2243.

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Ecdysozoa is a clade composed of eight phyla: the arthropods, tardigrades and onychophorans that share segmentation and appendages and the nematodes, nematomorphs, priapulids, kinorhynchs and loriciferans, which are worms with an anterior proboscis or introvert. Ecdysozoa contains the vast majority of animal species and there is a great diversity of body plans among both living and fossil members. The monophyly of the clade has been called into question by some workers based on analyses of whole genome datasets. We review the evidence that now conclusively supports the unique origin of these p
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13

Wang, Deng, Jean Vannier, Isabell Schumann, et al. "Origin of ecdysis: fossil evidence from 535-million-year-old scalidophoran worms." Proceedings of the Royal Society B: Biological Sciences 286, no. 1906 (2019): 20190791. http://dx.doi.org/10.1098/rspb.2019.0791.

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With millions of extant species, ecdysozoans (Scalidophora, Nematoida and Panarthropoda) constitute a major portion of present-day biodiversity. All ecdysozoans secrete an exoskeletal cuticle which must be moulted periodically and replaced by a larger one. Although moulting (ecdysis) has been recognized in early Palaeozoic panarthropods such as trilobites and basal groups such as anomalocaridids and lobopodians, the fossil record lacks clear evidence of ecdysis in early scalidophorans, largely because of difficulties in recognizing true exuviae. Here, we describe two types of exuviae in micros
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14

Vinn, Olev, Magdy El Hedeny, Mansour I. Almansour, Saleh Alfarraj, and Ursula Toom. "The first record of a possible 'priapulid' from the lower Cambrian (Series 2) of Estonia." Carnets Geol. 25, no. 8 (2025): 169–75. https://doi.org/10.2110/carnets.2025.2508.

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The Yunnanpriapulus? sp. can be divided into three distinct parts: an anterior introvert, a slightly constricted elongated neck, and a slightly bulbous elongated posterior trunk. The fossil is preserved as a three-dimensional cast in the fine-grained sandstone and is oriented parallel to the bedding plane. Rapid burial helped protect the organism from scavengers and decay and provided the opportunity for fossilization via pyritization. The preservation of the specimen on a bedding plane resulted from the post-mortem transportation of the dead animal, likely due to a storm event. The priapulid
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15

Srivastava, Purnima. "Problematic Worms and Priapulid-like Fossils from the Nagaur Group, the Marwar Supergroup, Western Rajasthan, India." Ichnos 19, no. 3 (2012): 156–64. http://dx.doi.org/10.1080/10420940.2012.702606.

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16

Baliński, Andrzej, Yuanlin Sun, and Jerzy Dzik. "Traces of marine nematodes from 470 million years old Early Ordovician rocks in China." Nematology 15, no. 5 (2013): 567–74. http://dx.doi.org/10.1163/15685411-00002702.

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Cylindrical, mostly horizontal, burrows of 20-60 μm diam. and sinusoidal course, found in the middle part of the Early Ordovician (early Floian) Fenxiang Formation in the Hubei Province of China, represent the oldest record of activity by marine nematodes, preceding known nematode body fossils by 70 million years. The burrows are filled with secondarily oxidised pyrite framboids and clay mineral flakes, indicating low oxygen content in the mud and proving that the animals lined their burrows with organic matter, being bacteriovores and mud-eaters. The marine bottom environment enabling such a
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17

Schreiber, A., M. Eisinger, H. Rumohr, and V. Storch. "Icy heritage: ecological evolution of the postglacial Baltic Sea reflected in the allozymes of a living fossil, the priapulid Halicryptus spinulosus." Marine Biology 125, no. 4 (1996): 671–85. http://dx.doi.org/10.1007/bf00349249.

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18

Lieberman, Bruce S. "A new soft-bodied fauna: The Pioche Formation of Nevada." Journal of Paleontology 77, no. 4 (2003): 674–90. http://dx.doi.org/10.1017/s0022336000044413.

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A new Burgess Shale-type soft-bodied fauna crossing the Lower-Middle Cambrian boundary in the Comet Shale Member of the Pioche Formation in Lincoln County, Nevada, contains common remains of soft-bodied ecdysozoan taxa. These fossils provide important new information about the nature and variety of Cambrian soft-bodied organisms. Arthropod taxa include one species of Canadaspis Novozhilov in Orlov, 1960, one species of ?Perspicaris Briggs, 1977, three species of Tuzoia Walcott, 1912, and at least two species of Anomalocaris Whiteaves, 1892. A priapulid referable to Ottoia Walcott, 1911a, was a
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19

Kimmig, Julien, and Brian R. Pratt. "Soft-bodied biota from the middle Cambrian (Drumian) Rockslide Formation, Mackenzie Mountains, northwestern Canada." Journal of Paleontology 89, no. 1 (2015): 51–71. http://dx.doi.org/10.1017/jpa.2014.5.

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AbstractA new Burgess Shale-type Lagerstätte is described from the middle Cambrian (Series 3, Drumian) Rockslide Formation of the Mackenzie Mountains, Northwest Territories, Canada. The Rockslide Formation is a unit of deeper water ramp to slope, mixed carbonate, and siliciclastic facies deposited on the northwestern margin of Laurentia. At the fossil-bearing locality, the unit onlaps a fault scarp cutting lower Cambrian sandstones. There it consists of a succession of shale and thick-laminated to thin-bedded lime mudstone, calcareous sandstone, and greenish-colored calcareous mudstone, overla
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20

Peel, John S., Simon Conway Morris, and Jon R. Ineson. "The Sirius Passet Fauna, an Early Cambrian Lagerstätte from North Greenland." Paleontological Society Special Publications 6 (1992): 233. http://dx.doi.org/10.1017/s2475262200007930.

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The Sirius Passet Fauna of North Greenland is one of the oldest Cambrian lagerstätten from the North American continent. It is known from a single locality in Peary Land (83°N, 40°W), on the shores of the Arctic Ocean, where outer shelf mudstones from the lower part of the Buen Formation (Early Cambrian) yield a rich assemblage of mainly poorly skeletised organisms with preserved soft parts. The steeply-dipping fossiliferous mudstones occur in close proximity to horizontally-bedded platform carbonates of the underlying Portfjeld Formation (Early Cambrian) in a structurally complex terrane. The
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21

Howard, Richard J., Mattia Giacomelli, Jesus Lozano-Fernandez, et al. "The Ediacaran origin of Ecdysozoa: integrating fossil and phylogenomic data." Journal of the Geological Society, February 1, 2022, jgs2021–107. http://dx.doi.org/10.1144/jgs2021-107.

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Ecdysozoans (Phyla Arthropoda, Kinorhyncha, Loricifera, Nematoda, Nematomorpha, Onychophora, Priapulida, Tardigrada) are invertebrates bearing a tough, periodically moulted cuticle that predisposes them to exceptional preservation. Ecdysozoans dominate the oldest exceptionally preserved bilaterian animal biotas in the early to mid-Cambrian (c. 520–508 Ma), with possible trace fossils in the latest Ediacaran (<556 Ma). The fossil record of Ecdysozoa is among the best understood of major animal clades and is believed to document their origins and evolutionary history well. Strikingly, however
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22

Wernström, Joel Vikberg, Ben J. Slater, Martin V. Sørensen, Denise Crampton, and Andreas Altenburger. "Geometric morphometrics of macro- and meiofaunal priapulid pharyngeal teeth provides a proxy for studying Cambrian “tooth taxa”." Zoomorphology, August 12, 2023. http://dx.doi.org/10.1007/s00435-023-00617-4.

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AbstractPriapulids are marine, benthic ecdysozoan worms that feed using a distinctive toothed pharynx. While only a handful of lineages have survived to the present day, the Cambrian priapulid stem group left behind a rich record of articulated body fossils and characteristic trace fossils in the form of burrows. Recently, the fossil record of isolated priapulid cuticular elements including pharyngeal teeth has gained increased attention as a means of revealing cryptic priapulid taxa otherwise unknown among macrofossils. In this study, we focus on the ecological implications of shape variation
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23

Turk, Katherine A., Achim Wehrmann, Marc Laflamme, and Simon A. F. Darroch. "Priapulid neoichnology, ecosystem engineering, and the Ediacaran–Cambrian transition." Palaeontology 67, no. 4 (2024). http://dx.doi.org/10.1111/pala.12721.

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AbstractThe evolutionary rise of powerful new ecosystem engineering impacts is thought to have played an important role in driving waves of biospheric change across the Ediacaran–Cambrian transition (ECT; c. 574–538 Ma). Among the most heavily cited of these is bioturbation (organism‐driven sediment disturbance) as these activities have been shown to have critical downstream geobiological impacts. In this regard priapulid worms are crucial; trace fossils thought to have been left by priapulan‐grade animals are now recognized as appearing shortly before the base of the Cambrian and represent so
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24

Wang, Deng, Jean Vannier, José M. Martín-Durán, María Herranz, and Chiyang Yu. "Preservation and early evolution of scalidophoran ventral nerve cord." Science Advances 11, no. 2 (2025). https://doi.org/10.1126/sciadv.adr0896.

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Ecdysozoan worms (Nematoida + Scalidophora) are typified by disparate grades of neural organization reflecting a complex evolutionary history. The fossil record offers a unique opportunity to reconstruct the early character evolution of the nervous system via the exceptional preservation of extinct representatives. We focus on their nervous system as it appears in early and mid-Cambrian fossils. We show that some of the oldest known representatives of the group either preserved in carbonaceous compression (early and mid-Cambrian Burgess-type preservation) or secondarily phosphatized in three d
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25

Mussini, Giovanni, and Nicholas J. Butterfield. "Exotic cuticular specializations in a Cambrian scalidophoran." Proceedings of the Royal Society B: Biological Sciences 292, no. 2040 (2025). https://doi.org/10.1098/rspb.2024.2806.

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Scalidophora, the ecdysozoan group including priapulids, kinorhynchs and loriciferans, comprises some of the most abundant and ecologically important Cambrian animals. However, reconstructions of the morphology and lifestyles of fossil scalidophorans are often hampered by poor preservation of their submillimetre-scale cuticular specializations. Based on exceptionally preserved small carbonaceous fossils (SCFs), we describe a new scalidophoran-grade animal, Scalidodendron crypticum gen. et sp. nov., from the Early to Middle Cambrian Hess River Formation of northern Canada. The Hess River SCFs c
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26

Turk, Katherine A., Mikaela A. Pulsipher, Helke Mocke, Marc Laflamme, and Simon A. F. Darroch. "Himatiichnus mangano igen. et isp. nov., a scalidophoran trace fossil from the late Ediacaran of Namibia." Royal Society Open Science 11, no. 10 (2024). http://dx.doi.org/10.1098/rsos.240452.

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Himatiichnus mangano igen. et isp. nov., a new trace fossil from the late Ediacaran Huns Member of the Urusis Formation, southern Namibia, comprises intertwining tubes exhibiting dual lineation patterns and reminiscent of both modern and early Cambrian examples of priapulid worm burrows. These similarities support the interpretation of a total-group scalidophoran tracemaker for H. mangano , thus providing direct evidence for the first appearance date of Scalidophora in the late Ediacaran ca 539 Ma. This new material is thus indicative of the presence of total-group scalidophorans below the Cam
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27

Wang, Deng, Jean Vannier, Cédric Aria, Jie Sun, and Jian Han. "Tube-dwelling in early animals exemplified by Cambrian scalidophoran worms." BMC Biology 19, no. 1 (2021). http://dx.doi.org/10.1186/s12915-021-01172-4.

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Abstract Background The radiation of ecdysozoans (moulting animals) during the Cambrian gave rise to panarthropods and various groups of worms including scalidophorans, which played an important role in the elaboration of early marine ecosystems. Although most scalidophorans were infaunal burrowers travelling through soft sediment at the bottom of the sea, Selkirkia lived inside a tube. Results We explore the palaeobiology of these tubicolous worms, and more generally the origin and evolutionary significance of tube-dwelling in early animals, based on exceptionally preserved fossils from the e
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28

Hu, S., M. Steiner, M. Zhu, et al. "A new priapulid assemblage from the early Cambrian Guanshan fossil Lagerstätte of SW China." Bulletin of Geosciences, February 24, 2012, 93–106. http://dx.doi.org/10.3140/bull.geosci.1238.

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29

Turk, Katherine A., Katie M. Maloney, Marc Laflamme, and Simon A. F. Darroch. "Paleontology and ichnology of the late Ediacaran Nasep–Huns transition (Nama Group, southern Namibia)." Journal of Paleontology, May 16, 2022, 1–17. http://dx.doi.org/10.1017/jpa.2022.31.

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Abstract The Nasep and Huns members of the Urusis Formation (Nama Group), southern Namibia, preserve some of the most diverse trace-fossil assemblages known from the latest Ediacaran worldwide, including potentially the world's oldest “complex” vertical sediment-penetrating burrows. These sediments record relatively diverse communities of bilaterian metazoans existing before the base of the Cambrian and an increase in the intensity of metazoan ecosystem engineering behaviors that could eventually produce profound changes in the character of the Phanerozoic sedimentary record (the “agronomic re
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30

Pari, Giovanni, Derek E. G. Briggs, and Robert R. Gaines. "The soft-bodied biota of the Cambrian Series 2 Parker Quarry Lagerstätte of northwestern Vermont, USA." Journal of Paleontology, February 16, 2022, 1–21. http://dx.doi.org/10.1017/jpa.2021.125.

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Abstract A non-biomineralized arthropod, Protocaris marshi, was described from the lower Cambrian (Dyeran Series 2, Stage 4) of Parker's Cobble in northwestern Vermont in 1884. It represents the first fossil exhibiting Burgess Shale-type preservation to have been discovered. The locality was presumed to have been worked out and was not collected in a significant way for more than 100 years. Rediscovery of productive layers has yielded soft-bodied and lightly sclerotized taxa new to the locality, including the alga Fuxianospora, a possible priapulid, a radiodont, and a specimen tentatively assi
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31

Pari, Giovanni, Derek E. G. Briggs, and Robert R. Gaines. "The Parker Quarry Lagerstätte of Vermont—The first reported Burgess Shale–type fauna rediscovered." Geology, February 22, 2021. http://dx.doi.org/10.1130/g48422.1.

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Soft-bodied fossils of Cambrian age, now known as Burgess Shale–type biotas, were first described from the Parker Slate of the northwest Vermont (USA) slate belt in the late 19th century, 25 years before the discovery of the Burgess Shale in British Columbia, Canada. Here, we report the rediscovery of fossiliferous horizons at Parker’s Cobble, the site of the original quarry, which was thought to have been exhausted by excavation. New discoveries include a radiodont, multiple specimens of a new bivalved arthropod, a priapulid, and other undescribed forms. Pervasive soft-sediment deformation su
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32

Mussini, Giovanni, James W. Hagadorn, Anne E. Miller, et al. "Evolutionary escalation in an exceptionally preserved Cambrian biota from the Grand Canyon (Arizona, USA)." Science Advances 11, no. 30 (2025). https://doi.org/10.1126/sciadv.adv6383.

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Exceptionally preserved fossil assemblages, or Konservat-Lagerstätten, open direct windows on non-biomineralized faunas that chronicle the Cambrian radiation of animal phyla. However, these assemblages do not typically capture the well-oxygenated, resource-rich environments sustaining most metazoan diversity in modern marine systems. We describe exceptionally preserved and articulated carbonaceous mesofossils from the middle Cambrian (~507 to 502 million years) Bright Angel Formation of the Grand Canyon (Arizona, USA). This biota preserves probable algal and cyanobacterial photosynthesizers to
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33

"Icy heritage: ecological evolution of the postglacial Baltic Sea reflected in the allozymes of a living fossil, the priapulid Halicryptus spinulosus." Oceanographic Literature Review 44, no. 7 (1997): 736. https://doi.org/10.1016/s0967-0653(97)85724-9.

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34

Mussini, Giovanni, Yorick P. Veenma, and Nicholas J. Butterfield. "A peritidal Burgess‐Shale‐type fauna from the middle Cambrian of western Canada." Palaeontology 68, no. 1 (2025). https://doi.org/10.1111/pala.70001.

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AbstractBurgess‐Shale‐type (BST) faunas have proven critical for mapping the Cambrian assembly of animal‐dominated ecosystems, but have so far only been reported from fully subaqueous deposits. Here we integrate evidence from ichnofossils, sedimentary features, and small carbonaceous fossils (SCFs) from the middle Cambrian (Late Guzhangian, Series 3) Pika Formation of western Jasper National Park, Alberta (Canada) to document a unique BST fauna, occupying a peritidal habitat near the outer margin of a large epicratonic sea. Finely laminated shales with mudcracks and dumbbell‐shaped Arthraria‐t
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35

Wood, Rachel A., and Mary L. Droser. "The evolution of reproduction in Ediacaran–Cambrian metazoans." Biological Reviews, May 15, 2025. https://doi.org/10.1111/brv.70036.

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ABSTRACTThe evolution of reproductive style is a fundamental aspect of metazoan life history but has not been explored holistically through the Ediacaran–Cambrian rise of metazoans. Recent molecular clock analyses based on only unequivocal metazoan fossil calibrations suggest that Porifera were present by at least 590 million years ago (Ma), all major eumetazoan clades originated in the mid–late Ediacaran, and bilaterians were probably present by the late Ediacaran. An alternating pelagic larval (potentially for dispersal) and benthic adult life cycle appears to be an ancestral feature of meta
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36

Shan, Longlong, Thomas H. P. Harvey, Kui Yan, Jun Li, Yuandong Zhang, and Thomas Servais. "Palynological recovery of small carbonaceous fossils (SCFs) indicates that the late Cambrian acritarch Goniomorpha Yin 1986 represents the teeth of a priapulid worm." Palynology, December 14, 2022. http://dx.doi.org/10.1080/01916122.2022.2157504.

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