Soft-body fossils

@misc{muller1983crustacea3,
    author = "Muller, K. J",
    title = "Crustacea with preserved soft parts from the Upper Cambrian of Sweden",
    year = "1983",
    howpublished = "Lethaia, v. 16, p. 93-109",
    note = "talkorigins\_source = {true}; raw\_reference = {Muller, K. J., 1983, Crustacea with preserved soft parts from the Upper Cambrian of Sweden: Lethaia, v. 16, p. 93-109.}"
}

A new biota including lightly sclerotized and soft-bodied organisms occurs in finely laminated argillaceous dolomites of late Llandoverian age in Waukesha County, Wisconsin. This discovery fills a gap between well known Cambrian and Devonian Konservat Lagerstatten. The biota is dominated by arthropods. A dalmanitid is the most numerous of 13 genera of trilobites; the crustaceans include phyllocarids and ostracods; the chelicerates are represented by the earliest well preserved xiphosure and the fauna includes a possible marine uniramian. The earliest representative of the enigmatic class Thylacocephala, and at least three arthropods of uncertain affinity are also present. There are at least four worm taxa including a possible leech and a papillate annelid. The locality has also yielded a conodont animal, Panderodus. Graptolites and conulariids are common, but echinoderms, brachiopods, bryozoans, corals and molluscs are extremely rare or absent. The unusual composition and exceptional preservation of this assemblage indicates that the biota lived and died in environments rarely represented in the Silurian fossil record.

@article{mikulic1985a,
    author = "Mikulic, D. G. and Briggs, Derek Ernest Gilmor and Kluessendorf, Joanne",
    title = "A new exceptionally preserved biota from the Lower Silurian of Wisconsin, U. S. A",
    year = "1985",
    journal = "Philosophical Transactions of the Royal Society of London. B, Biological Sciences",
    abstract = "A new biota including lightly sclerotized and soft-bodied organisms occurs in finely laminated argillaceous dolomites of late Llandoverian age in Waukesha County, Wisconsin. This discovery fills a gap between well known Cambrian and Devonian Konservat Lagerstatten. The biota is dominated by arthropods. A dalmanitid is the most numerous of 13 genera of trilobites; the crustaceans include phyllocarids and ostracods; the chelicerates are represented by the earliest well preserved xiphosure and the fauna includes a possible marine uniramian. The earliest representative of the enigmatic class Thylacocephala, and at least three arthropods of uncertain affinity are also present. There are at least four worm taxa including a possible leech and a papillate annelid. The locality has also yielded a conodont animal, Panderodus. Graptolites and conulariids are common, but echinoderms, brachiopods, bryozoans, corals and molluscs are extremely rare or absent. The unusual composition and exceptional preservation of this assemblage indicates that the biota lived and died in environments rarely represented in the Silurian fossil record.",
    url = "https://doi.org/10.1098/rstb.1985.0140",
    doi = "10.1098/rstb.1985.0140",
    number = "1148",
    pages = "75-85",
    volume = "311"
}

@article{mikulic1985a1,
    author = "Mikulic, D. G. and Briggs, D. E. G. and Kluessendorf, J",
    title = "A new exceptionally preserved biota from the Lower Silurian of Wisconsin, USA",
    year = "1985",
    journal = "Philosophical Transactions of the Royal Society, London B, v. 311, p. 75- 85",
    note = "talkorigins\_source = {true}; raw\_reference = {Mikulic, D. G., Briggs, D. E. G., and Kluessendorf, J., 1985, A new exceptionally preserved biota from the Lower Silurian of Wisconsin, USA: Philosophical Transactions of the Royal Society, London B, v. 311, p. 75- 85.}"
}

@article{mikulic1985a2,
    author = "Mikulic, D. G. and Briggs, D. E. G. and Kluessendorf, J",
    title = "A new exceptionally preserved biota from the Lower Silurian of Wisconsin, USA",
    year = "1985",
    journal = "Philosophical Transactions of the Royal Society, London B, v. 311, p. 75-85",
    note = "talkorigins\_source = {true}; raw\_reference = {Mikulic, D. G., Briggs, D. E. G., and Kluessendorf, J., 1985, A new exceptionally preserved biota from the Lower Silurian of Wisconsin, USA: Philosophical Transactions of the Royal Society, London B, v. 311, p. 75-85.}"
}

@article{whittington1985tegopelte4,
    author = "Whittington, H. B",
    title = "Tegopelte gigas, a second soft-bodied trilobite from the Burgess Shale, Middle Cambrian, British Columbia",
    year = "1985",
    journal = "Journal of Paleontology, v. 59, p. 1251-1274",
    note = "talkorigins\_source = {true}; raw\_reference = {Whittington, H. B., 1985, Tegopelte gigas, a second soft-bodied trilobite from the Burgess Shale, Middle Cambrian, British Columbia: Journal of Paleontology, v. 59, p. 1251-1274.}"
}

The Lower Cambrian Chengjiang fauna is reviewed and shown to be closely comparable with the younger Burgess Shale fauna. but with various differences in detail. A diverse group of more or less annulated lobopod animals including ‘armoured lobopods’ are regarded as representatives of the phylum Onychophora. ‘Trilobitomorphs’ include several new types. Probable protaspides of the trilobitomorph Naraoia are described. No molluses or deuterostomes have been identified. The preservational orientations of the various taxa are reviewed and compared with orientations of the Burgess Shale taxa. Orientation in the sediment is found to be closely correlated to the original shape of individuals. Several new genera and species are described: the segmented. worm‐shaped Yunnanozoon lividum gen. et sp.n., the ‘armoured lobopods’ Onychodictyon ferox gen. et sp.n. and Cardiodictyon catenulum gen. et sp.n. and the arthropods Saperion glumaceum gen. et sp.n., Sinoburius Iunaris gen. et sp.n., and Xandarella spectaculum gen. et sp.n.

@article{xianguang1991composition,
    author = "XIANGUANG, HOU and RAMSKÖLD, LARS and BERGSTRÖM, JAN",
    title = "Composition and preservation of the Chengjiang fauna –a Lower Cambrian soft‐bodied biota",
    year = "1991",
    journal = "Zoologica Scripta",
    abstract = "The Lower Cambrian Chengjiang fauna is reviewed and shown to be closely comparable with the younger Burgess Shale fauna. but with various differences in detail. A diverse group of more or less annulated lobopod animals including ‘armoured lobopods’ are regarded as representatives of the phylum Onychophora. ‘Trilobitomorphs’ include several new types. Probable protaspides of the trilobitomorph Naraoia are described. No molluses or deuterostomes have been identified. The preservational orientations of the various taxa are reviewed and compared with orientations of the Burgess Shale taxa. Orientation in the sediment is found to be closely correlated to the original shape of individuals. Several new genera and species are described: the segmented. worm‐shaped Yunnanozoon lividum gen. et sp.n., the ‘armoured lobopods’ Onychodictyon ferox gen. et sp.n. and Cardiodictyon catenulum gen. et sp.n. and the arthropods Saperion glumaceum gen. et sp.n., Sinoburius Iunaris gen. et sp.n., and Xandarella spectaculum gen. et sp.n.",
    url = "https://doi.org/10.1111/j.1463-6409.1991.tb00303.x",
    doi = "10.1111/j.1463-6409.1991.tb00303.x",
    number = "4",
    pages = "395-411",
    volume = "20"
}

@article{dong2005the,
    author = "DONG, Xiping",
    title = "The fossils of Orsten-type preservation from Middle and Upper Cambrian in Hunan, China ——Three-dimensionally preserved soft-bodied fossils (Arthropods)",
    year = "2005",
    journal = "Chinese Science Bulletin",
    url = "https://doi.org/10.1360/982005-795",
    doi = "10.1360/982005-795",
    number = "13",
    pages = "1352",
    volume = "50"
}

@article{hou2005cambrian,
    author = "Hou, Xian‐Guang and Stanley Jr., George D. and Zhao, Jie and Ma, Xiao‐Ya",
    title = "Cambrian anemones with preserved soft tissue from the Chengjiang biota, China",
    year = "2005",
    journal = "Lethaia",
    url = "https://doi.org/10.1080/00241160510013295",
    doi = "10.1080/00241160510013295",
    number = "3",
    pages = "193-203",
    volume = "38"
}

@article{caron2006a,
    author = "Caron, Jean-Bernard and Scheltema, Amélie and Schander, Christoffer and Rudkin, David",
    title = "A soft-bodied mollusc with radula from the Middle Cambrian Burgess Shale",
    year = "2006",
    journal = "Nature",
    url = "https://doi.org/10.1038/nature04894",
    doi = "10.1038/nature04894",
    number = "7099",
    pages = "159-163",
    volume = "442"
}

@article{johnston2009a,
    author = "Johnston, Kimberley J. and Johnston, Paul A. and Powell, Wayne G.",
    title = "A new, Middle Cambrian, Burgess Shale-type biota, Bolaspidella Zone, Chancellor Basin, southeastern British Columbia",
    year = "2009",
    journal = "Palaeogeography, Palaeoclimatology, Palaeoecology",
    url = "https://doi.org/10.1016/j.palaeo.2009.02.015",
    doi = "10.1016/j.palaeo.2009.02.015",
    number = "1-2",
    pages = "106-126",
    volume = "277"
}

A 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, overlain by shallower water dolostones of the Avalanche Formation, which is indicative of an overall progradational sequence. The Rockslide Formation is of similar age to the Wheeler and Marjum formations of Utah, belonging to the Bolaspidella Biozone. Only two 1 m thick units of greenish mudstone exhibit soft-bodied preservation, with most specimens coming from the lower interval. However, the biota is common but not as diverse as that of other Lagerstätten such as the Burgess Shale in its type area. The shelly fauna is dominated by the hyolith Haplophrentis carinatus Matthew, 1899 along with sparse linguliformean brachiopods, agnostoid arthropods, and ptychoparioid trilobites. The nonmineralized biota includes the macrophytic alga Margaretia dorus Walcott, 1911, priapulid worms, and the carapaces of a number of arthropods. The arthropods belong to Isoxys mackenziensis n. sp., Tuzoia cf. T. guntheri Robison and Richards, 1981; Branchiocaris? sp., Perspicaris? dilatus Robison and Richards, 1981; and bradoriids, along with fragments of arthropods of indeterminate affinities. The style of preservation indicates that most soft parts underwent complete biodegradation, leaving just the more resistant materials such as chitinous arthropod cuticles. The range of preservation and similarity to the coeval biotas preserved in Utah suggests that the composition of this Lagerstätte is probably representative of the community living on the relatively deep-water ramp or slope during middle Cambrian time in Laurentia. This would argue that the extraordinary diversity of the Burgess Shale at Mount Field is anomalous.

@article{kimmig2015softbodied,
    author = "Kimmig, Julien and Pratt, Brian R.",
    title = "Soft-bodied biota from the middle Cambrian (Drumian) Rockslide Formation, Mackenzie Mountains, northwestern Canada",
    year = "2015",
    journal = "Journal of Paleontology",
    abstract = "A 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, overlain by shallower water dolostones of the Avalanche Formation, which is indicative of an overall progradational sequence. The Rockslide Formation is of similar age to the Wheeler and Marjum formations of Utah, belonging to the Bolaspidella Biozone. Only two 1 m thick units of greenish mudstone exhibit soft-bodied preservation, with most specimens coming from the lower interval. However, the biota is common but not as diverse as that of other Lagerstätten such as the Burgess Shale in its type area. The shelly fauna is dominated by the hyolith Haplophrentis carinatus Matthew, 1899 along with sparse linguliformean brachiopods, agnostoid arthropods, and ptychoparioid trilobites. The nonmineralized biota includes the macrophytic alga Margaretia dorus Walcott, 1911, priapulid worms, and the carapaces of a number of arthropods. The arthropods belong to Isoxys mackenziensis n. sp., Tuzoia cf. T. guntheri Robison and Richards, 1981; Branchiocaris? sp., Perspicaris? dilatus Robison and Richards, 1981; and bradoriids, along with fragments of arthropods of indeterminate affinities. The style of preservation indicates that most soft parts underwent complete biodegradation, leaving just the more resistant materials such as chitinous arthropod cuticles. The range of preservation and similarity to the coeval biotas preserved in Utah suggests that the composition of this Lagerstätte is probably representative of the community living on the relatively deep-water ramp or slope during middle Cambrian time in Laurentia. This would argue that the extraordinary diversity of the Burgess Shale at Mount Field is anomalous.",
    url = "https://doi.org/10.1017/jpa.2014.5",
    doi = "10.1017/jpa.2014.5",
    number = "1",
    pages = "51-71",
    volume = "89"
}

We describe an extraordinarily preserved non-trilobite artiopod Bailongia longicaudata gen. et sp. nov. from the Cambrian Stage 4 Guanshan Biota in Yiliang, Kunming of the Yunnan Province in China. Its exoskeleton consists of a large semi-elliptical cephalon with paired lateral posterior eyes, nine tapering homonomous tergites and a long slender tailspine. Appendages include paired small antennae, at least three pairs of post-antennal cephalic limbs, and trunk biramous limbs consisting of an endopod and an exopod with lamellae. B. longicaudata does not conform to any taxon within Artiopoda, although the eyes invite comparisons with Xandarella spectaculum, Sinoburius lunaris and Phytophilaspis. Parsimony analyses indicate Bailongia is a member of Artiopoda and cannot be readily accommodated within any of the major artiopod clades.

@article{jiao2022a,
    author = "Jiao, De-Guang and Du, Kun-Sheng and Zhang, Xi-Guang and Yang, Jie and Eggink, Daniel",
    title = "A new small soft-bodied non-trilobite artiopod from the Cambrian Stage 4 Guanshan Biota",
    year = "2022",
    journal = "Geological Magazine",
    abstract = "We describe an extraordinarily preserved non-trilobite artiopod Bailongia longicaudata gen. et sp. nov. from the Cambrian Stage 4 Guanshan Biota in Yiliang, Kunming of the Yunnan Province in China. Its exoskeleton consists of a large semi-elliptical cephalon with paired lateral posterior eyes, nine tapering homonomous tergites and a long slender tailspine. Appendages include paired small antennae, at least three pairs of post-antennal cephalic limbs, and trunk biramous limbs consisting of an endopod and an exopod with lamellae. B. longicaudata does not conform to any taxon within Artiopoda, although the eyes invite comparisons with Xandarella spectaculum, Sinoburius lunaris and Phytophilaspis. Parsimony analyses indicate Bailongia is a member of Artiopoda and cannot be readily accommodated within any of the major artiopod clades.",
    url = "https://doi.org/10.1017/s0016756821001254",
    doi = "10.1017/s0016756821001254",
    number = "5",
    pages = "730-734",
    volume = "159"
}

Before Cambrian Stage 3, unambiguous body fossils of segmented bilaterians were rare, severely hampering our understanding of the early history of such important animals. Here we report a variety of microfossils with quintessential features such as paired appendages, dorsoventral and anteroposterior differentiations from the basal Cambrian Fortunian of South China, representing the earliest known three-dimensional body fossils of segmented bilaterians. These fossils were all microbial pseudomorphs built up by secondarily phosphatized bacteria aggregations, testifying microbial pseudomorph could serve as a novel and important pathway to preserve tiny, fragile bilaterian progenitors. This finding unveils a diversified segmented bilaterian world at the very beginning of Cambrian and would arouse a more comprehensive perspective on the early evolution of bilaterian body plans.

@misc{yang2024basal,
    author = "Yang, Xiaoguang and Wang, Deng and Zhang, Zhiliang and Wang, Xing and Sun, Jie and Hao, Wenjing and Liu, Yiqun and Uesugi, Kentaro and Komiya, Tsuyoshi and Han, Jian",
    title = "Basal Cambrian soft-bodied segmented bilaterians preserved as microbial pseudomorphs",
    year = "2024",
    abstract = "Before Cambrian Stage 3, unambiguous body fossils of segmented bilaterians were rare, severely hampering our understanding of the early history of such important animals. Here we report a variety of microfossils with quintessential features such as paired appendages, dorsoventral and anteroposterior differentiations from the basal Cambrian Fortunian of South China, representing the earliest known three-dimensional body fossils of segmented bilaterians. These fossils were all microbial pseudomorphs built up by secondarily phosphatized bacteria aggregations, testifying microbial pseudomorph could serve as a novel and important pathway to preserve tiny, fragile bilaterian progenitors. This finding unveils a diversified segmented bilaterian world at the very beginning of Cambrian and would arouse a more comprehensive perspective on the early evolution of bilaterian body plans.",
    url = "https://doi.org/10.1101/2024.07.03.601876",
    doi = "10.1101/2024.07.03.601876"
}

@article{zeng2026a,
    author = "Zeng, Han and Liu, Qi and Zhao, Fangchen and Luo, Cui and Wang, Dezhi and Zhu, Yuyan and Liu, Yao and Chen, Kai and Sun, Zhixin and Hong, Yanjie and Miao, Lanyun and Hu, Chunlin and Sun, Haijing and Pan, Bing and Zhao, Jialin and Yin, Zongjun and Li, Guoxiang and Yang, Xinglian and Yang, Aihua and Hu, Shixue and Zhu, Maoyan",
    title = "A Cambrian soft-bodied biota after the first Phanerozoic mass extinction",
    year = "2026",
    journal = "Nature",
    url = "https://doi.org/10.1038/s41586-025-10030-0",
    doi = "10.1038/s41586-025-10030-0",
    number = "8106",
    pages = "679-687",
    volume = "651"
}