This is a list of fossils found at Maotianshan Shales, whose most famous assemblage of organisms are referred to as the Chengjiang biota. [1]
The Maotianshan Shales are a series of Early Cambrian sedimentary deposits in the Chiungchussu Formation, [2] famous for their Konservat Lagerstätten , deposits known for the exceptional preservation of fossilized organisms or traces. The Maotianshan Shales form one of some forty Cambrian fossil locations worldwide exhibiting exquisite preservation of rarely preserved, non-mineralized soft tissue, comparable to the fossils of the Burgess Shale of British Columbia, Canada.
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| A small ecdysozoan worm that was originally thought to have been a priapulid worm. | ||
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| An ecdysozoan worm considered close to the priapulids, it has a spiny proboscis, an elliptical theca on the posterior part of the trunk, and a caudal appendage. |
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| A genus of palaeoscolecid worm. Fossils have been found with smaller worms seemingly attached to them. | |||
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| An extinct palaeoscolecid worm closely related to Cricocosmia. | |||
| A genus of palaeoscolecid worm | |||
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| An archaeopriapulid worm known from the biota. | |||
| A single specimen. | A stem-group priapulid. | ||
| A carnivorous priapulid-like worm that has been found living in the discarded shells of hyoliths. | |||
| An archaeopriapulid worm that resembles another worm, Selkirkia , from the Burgess shale | |||
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| A burrowing archaeopriapulid worm also known from the burgess shale, alongside the Fezouata Shale. | |||
| A genus of archaeopriapulid worm. | |||
| A priapulid worm thought to belong to the crown group. |
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| An unarmoured lobopodian, apparently closely related to Onychophora. The long, paired, cirriform structures at the end of the body could have been sensory, or perhaps represent viscera not externally present in life. | ||
| A highly elongate armoured lobopodian, with saddle-shaped sclerites and a lightly sclerotized head. | |||
| A heavily armoured lobopodian. | |||
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| A simple, unarmoured lobopodian. | ||
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| An enigmatic genus of lobopodian, potentially related to either the Luolishaniidae or to Tardigrada. Not always monophyletic. | ||
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| A large unarmoured lobopodian, commonly found in association with mass death-beds of legless worms and Eldonia. Thought to have been a scavenger. |
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| Microdictyon is known from sclerite microfossils around the globe, while fossils from the Chengjiang are the only in the world to preserve the soft anatomy. |
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| Type genus of Luolishaniidae. The species Miraluolishania haikouensis is considered a juvenile synonym of Luolishania. | |||
| Derived Luolishaniid lobopodian with five setiferous sieving appendages, and lacking any walking limbs. Sometimes discovered living in constructed tubes, buried into the sediment. | |||
| A hallucigeniid lobopodian with a bulbous sclerotized head and two pairs of small feeding appendages, borne on a short neck. |
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| A poorly known siberiid. | ||
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| A giant siberiid lobopodian related to Megadictyon. |
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| A giant gilled lobopodian, closely related to Pambdelurion. Also present in the Xiaoshiba Biota. |
Radiodonta are a group of highly successful panarthropods. The Chengjiang Fauna includes a large number of Radiodont species, primarily from the clade Amplectobeluidae. While hurdiids are poorly known from the formation and surrounding region, several specimens attributable to the family have been discovered, but remain unnamed. Some radiodonts from the Chengjiang defy easy classification, or are otherwise debated in their exact affinity.
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| Previously described as "Anomalocaris" saron. | ||
Innovatiocaris |
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| Previously assigned to the Anomalocaris genus, and one of the most completely known radiodonts. | |
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| A giant radiodont combining features from different clades. |
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| A large Amplectobeluid radiodont with long tail furcae. The frontal appendages formed a distinctive claw-like shape. | ||
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| Small Amplectobeluid radiodont. Some specimens of this genus represent some of the smallest known radiodonts. | ||
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| Large Amplectobeluid radiodonts, with crushing gnathobases. The species R. consimilis may belong to Houcaris instead. [3] |
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| A second species is included in the genus, Lenisicaris pennsylvanica, from the Kinzers Formation. | ||
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| An unusual radiodont seemingly transitional between anomalocaridids and amplectobeluids, with an oral cone similar to the former but gnathobase-like structures similar to the latter. | ||
Hurdiids (peytoiids [3] ) have historically been considered absent, or at least rare, from the Chengjiang Biota, an observation that researchers have found confusing given their regularity at other sites. In recent years, a number of fragmentary hurdiid fossils have been described, including 3 partial appendages which remain unnamed, [4] an isolated sclerite assigned to Cambroraster, [5] Zhenghecaris (the affinities of which, even its identity as a radiodont, remains questionable), and isolated lateral scelrites (also questionable). [6]
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| A large nektobenthic hurdiid, first known from the Burgess Shale. Fossils of Cambroraster are also known from Mantou Formation of North China, [7] making this a very widespread genus. This genus was the first definitive peytoiid known from China. | ||
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| Large-sized arthropod carapace, originally described as bivalved arthropod close to Thylacocephala, later reinterpreted as hurdiid radiodont close to Cambroraster , its classification is still discussed. Some fossils, apparently the lateral scelrites of a radiodont, connected anteriorly by a "beak" and featuring two-pronged, wing-like processes posteriorly (similar to the lateral processes of Zhenghecaris's central head sclerite), have also been attributed to Zhenghecaris. [6] | ||
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Astutuscaris |
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| A small arthropod of uncertain affinities superficially similar to fuxianhuiids. | |
Chuandianella |
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| An enigmatc bivalved arthropod originally considered close to Waptia . It was then discovered that it lacked mandibles and therefore does not belong to Hymenocarina, the Waptiidae family, and even the larger mandibulata clade. | |
| An enigmatic bivalved arthropod. | |||
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| An enigmatic arthropod that has been tentatively classified as a possible radiodont, although a position closely related to Erratus as the most primitive deuteropod is more likely. | ||
Cyathocephalus |
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| A small arthropod that is currently "too poorly preserved to verify their identity as a valid taxa". | |||
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| A bivalved arthropod currently placed as the most basal deuteropod, that shows the origins of lungs and legs in early arthropods. | ||
| An arthropod that has been speculated to be an early crustacean. | |||
| A small bivalved arthropod that somewhat resembles ostracods, but also has large, upward facing appendages with spines. | |||
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| A basal nektonic arthropod that possessed a bivalved carapace, and large circular eyes, related to Surusicaris from the Burgess Shale. Also found in various other Cambrian aged sites. | |||
| A small, somewhat enigmatic arthropod known from the biota | |||
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| An early arthropod that may show the "missing link" between early stem-arthropods, and more advanced true arthropods. It possessed frontal appendages similar to radiodonts. | ||
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| A predatory arthropod that had spiked frontal appendages similar to those of the megacheirans, and was previously thought to have belonged to that grouping | |||
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| An arachnomorph arthropod of the group strabopida. Unlike the other members of strabopida, Parapaleomerus lacks dorsal eyes and only possesses ten trunk tergites. The telson has been described as trapezoidal in shape. | |||
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| A poorly known arthropod that, in 2013, was suggested to be a member of the family Kootenichelidae, alongside Kootenichela and Worthenella. | |||
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| A giant arthropod once misidentified as a peytoiid radiodont. | |||
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| Also known in the Burgess Shale by B. pretiosa, B. yunnanensis is only known from isolated carapaces, although it is thought that this may be because in this species the carapace covered the entire body. It also has two morphotypes. | ||
| A hymenocarine arthropod, also known from the Burgess Shale by C. perfecta, although some authors believe that both species should be separated into different genera. They were mainly benthic animals. | |||
| A hymenocarine arthropod. The species in the genus are primarily distinguished by the presence of a serrated edge on the front of the carapace of C. serrata. C. serrata is noted for the modification of an anterior pair of limbs into spined grasping appendages, indicating a predatory lifestyle. | |||
| A hymenocarine arthropod that had a bivalved carapace which covered about a third of its total body-length, and had up to six serrations on its forward edge. The head had a pair of large uniramous antennae, as well as a smaller pair of secondary antennae, and a pair of mandibles and maxillae. | |||
| A small euarthropod, and unlike other hymenocarines, it possibly had eyes directly on top of its carapace. It was most likely a nektobenthic animal. | |||
| A bivalved arthropod of the hymenocarina grouping. The species P. spatiosa is regarded as a filter feeder, using the setae on the endites of their limbs to filter out matter from the water column | |||
| A hymenocarine notable for fossils showing a bizarre chain of multiple individuals. Although the purpose for this behavior is unknown, it may have served a reproductive, migratory, or defensive purpose, with the authors of the describing paper of this genus considering migration as its most likely function. | |||
Tuzoia |
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| A large hymenocarine that possessed a semicircular bivalved carapace, and was one of the largest arthropods from the Cambrian | |
Jugatacaris | A. agillis |
| A hymenocarine arthropod. The carapace of the animal was around 28 to 37 millimetres (1.1 to 1.5 in) in length, with a pronounced ridge at the top of the carapace separating the two valves, which formed a fin-like structure raised above the carapace. The head had a pair of stalked eyes, as well as a dumbbell shaped medial eye between them. The head also bore a pair of mandibles as well as at least one and possibly two pairs of antennules. | |
Xiazhuangocaris | X. chenggongensis |
| A hymenocarine with no known head fossils. The material known show that the carapace had a pronounced pair of notches at its front, as well as a posterior notch at its rear. The body had at least 13 tergite-pleurite rings, which terminate in a pair of rounded caudal rami, which are fringed with setae. |
Genus | Species | Material | Notes | Images |
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| A Fuxianhuiid arthropod with numerous appendages. It is one of the most complete arthropods from the lower Cambrian. | |||
| A large arthropod named after Fuxian Lake. Some adult specimens are found closely associated with numerous juveniles, indicating a level of parental care. |
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| A small arthropod related to Leanchoilia, also known from the Burgess Shale. | |||
| One of the several species of Leanchoilia, it has known juvenile specimens and the labrum morphology known. |
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Sklerolibyon | *S. maomima | |||
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| A jianfengiid once misidentified as a radiodont |
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Acanthomeridion |
| Multiple Specimens. | ||
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| An arthropod that had segmentation which resembled that of a millipede, as well as head and tail shields with thorny spikes. Recently, it was recovered as the most basal artiopod. | |||
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| A species of deuterostome animal with controversial taxonomy, with some authors placing it as an chordate, stem-vertebrate, hemichordate, etc. |
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| A chordate known from the Yunnan Province with potential relations to cephalochordates like lancelets. | |||
| An early craniate (an animal with a distinct notochord and head) of the Myllokunmingiida order. Some authors consider it potentially synonymous with the contemporary Myllokunmingia. [8] | |||
| A myllokunmingiid known for its wide body plan, sail-like fin on its dorsal plane, and a non-biomineralized skeleton. | |||
| Another myllokunmingiid known for lacking a dorsal fin, and having a more slim body compared to its two contemporary relatives. | |||
| An early chordate thought to represent a transitional form between earlier chordates, and more derived craniates. |
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| Senior synonym of Cheungkongella ancestralis. | |||
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| Senior synonym of Eldonia eumorpha. |
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| Likely the earliest known tunicate, alongside Palaeoikopleuria . | |||
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Stromatoveris |
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Hazelia |
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Paraleptomitella |
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Sinfoflabrum |
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Triticispongia |
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| Possible synonym of Nectocaris pteryx | |||
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Macroalgae rank fourth in species-level diversity behind arthropods, priapulids, and sponges, and account for 71.5% of total abundance from the biota, particularly dominated by unattached (floating, planktonic, and drifting) species. [9]
Genus | Species | Material | Notes | Images |
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| Sessile, attached to the substrate. | |||
| The most common species of macroalgae present, accounting for 52% of all macroalgae fossils reported from the Chengjiang. | Unattached (planktonic, drifting, floating) | ||
| Unattached (planktonic, drifting, floating) | |||
| Sessile, attached to the substrate. | |||
| Unattached (planktonic, drifting, floating) | |||
| Sessile, attached to the substrate. | |||
| Unattached (planktonic, drifting, floating) | |||
| Sessile, attached to the substrate. | |||
| Sessile, attached to the substrate. | |||
| Unattached (planktonic, drifting, floating) | |||
| Unattached (planktonic, drifting, floating) | |||
| Unattached (planktonic, drifting, floating) | |||
Dinocaridida is a proposed fossil taxon of basal arthropods, which flourished during the Cambrian period and survived up to Early Devonian. Characterized by a pair of frontal appendages and series of body flaps, the name of Dinocaridids refers to the suggested role of some of these members as the largest marine predators of their time. Dinocaridids are occasionally referred to as the 'AOPK group' by some literatures, as the group compose of Radiodonta, Opabiniidae, and the "gilled lobopodians" Pambdelurion and Kerygmachelidae. It is most likely paraphyletic, with Kerygmachelidae and Pambdelurion more basal than the clade compose of Opabiniidae, Radiodonta and other arthropods.
Anomalocaris is an extinct genus of radiodont, an order of early-diverging stem-group arthropods.
Anomalocarididae is an extinct family of Cambrian radiodonts, a group of stem-group arthropods.
Schinderhannes bartelsi is a species of hurdiid radiodont (anomalocaridid), known from one specimen from the Lower Devonian Hunsrück Slates. Its discovery was astonishing because the latest definitive radiodonts were known only from the Early Ordovician, at least 66 million years earlier than this taxon.
Radiodonta is an extinct order of stem-group arthropods that was successful worldwide during the Cambrian period. Radiodonts are distinguished by their distinctive frontal appendages, which are morphologically diverse and used for a variety of functions. Radiodonts are among the earliest large predators, but they also included sediment sifters and filter feeders. Some of the most famous species of radiodonts are the Cambrian taxa Anomalocaris canadensis, Hurdia victoria, Peytoia nathorsti, Titanokorys gainesi, Cambroraster falcatus and Amplectobelua symbrachiata. The later surviving members include the subfamily Aegirocassisinae from the Early Ordovician of Morocco and the Early Devonian member Schinderhannes bartelsi from Germany.
Hurdia is an extinct genus of hurdiid radiodont that lived 505 million years ago during the Cambrian Period. Fossils have been found in North America, China and the Czech Republic.
Cucumericrus ("cucumber-leg") is an extinct genus of stem-arthropod. The type and only species is Cucumericrus decoratus, with fossils discovered from the Maotianshan Shales of Yunnan, China.
Amplectobeluidae is a clade of Cambrian radiodonts. It currently includes five definitive genera, Amplectobelua, Lyrarapax, Ramskoeldia, Guanshancaris and a currently unnamed genus from the lower Cambrian aged Sirius Passet site in Greenland. There is also a potential fifth genus, Houcaris, but that genus has become problematic in terms of its taxonomic placement.
Hurdiidae is an extinct cosmopolitan family of radiodonts, a group of stem-group arthropods, which lived during the Paleozoic Era. It is the most long-lived radiodont clade, lasting from the Cambrian period to the Devonian period.
Ramskoeldia is a genus of amplectobeluid radiodont described in 2018. It was the second genus of radiodont found to possess gnathobase-like structures and an atypical oral cone after Amplectobelua. The type species, Ramskoeldia platyacantha, was discovered in the Chengjiang biota of China, the home of numerous radiodontids such as Amplectobelua and Lyrarapax.
Cambroraster is an extinct monotypic genus of hurdiid radiodont, dating to the middle Cambrian, and represented by the single formally described species Cambroraster falcatus. Hundreds of specimens were found in the Burgess Shale, and described in 2019. A large animal at up to 30 centimetres (12 in), it is characterized by a significantly enlarged horseshoe-shaped dorsal carapace (H-element), and presumably fed by sifting through the sediment with its well-developed tooth plates and short frontal appendages with hooked spines. Nicknamed the "spaceship" fossil when first found, for the way its dorsal carapace resembles the fictional Millennium Falcon, the specific epithet falcatus in its scientific name is a nod to that resemblance.
Houcaris is a possibly paraphyletic radiodont genus, tentatively assigned to either Amplectobeluidae, Anomalocarididae or Tamisiocarididae, known from Cambrian Series 2 of China and the United States. The type species is Houcaris saron which was originally described as a species of the related genus Anomalocaris. Other possible species include H. magnabasis and H. consimilis. The genus Houcaris was established for the two species in 2021 and honors Hou Xianguang, who had discovered and named the type species Anomalocaris saron in 1995 along with his colleagues Jan Bergström and Per E. Ahlberg.
Titanokorys is a genus of extinct hurdiid (peytoiid) radiodont that existed during the mid Cambrian. It is the largest member of its family from the Cambrian, with a body length of 50 cm (20 in) long, making it one of the largest animals of the time. It bears a resemblance to the related genus Cambroraster. Fossils of T. gainesi were first found within the Marble Canyon locality within the Burgess Shale in 2018. The fossils were not named until 2021 because they were assumed to be giant specimens of Cambroraster.
Laminacaris is a genus of extinct stem-group arthropods (Radiodonta) that lived during the Cambrian period. It is monotypic with a single species Laminacaris chimera, the fossil of which was described from the Chengjiang biota of China in 2018. Around the same time, two specimens that were similar or of the same species were discovered at the Kinzers Formation in Pennsylvania, USA. The first specimens from China were three frontal appendages, without the other body parts.
Innovatiocaris is a genus of radiodont of uncertain family from the early Cambrian Chengjiang Lagerstätte of Yunnan Province, China. The genus contains two named species, I. maotianshanensis, known from a nearly complete young individual measuring up to 15.2 cm (6.0 in) and isolated frontal appendages, and I.? multispiniformis, known from a complete frontal appendage.
Wufengella is a genus of extinct camenellan "tommotiid" that lived during the Early Cambrian. Described in 2022, the only species Wufengella bengtsonii was discovered from the Maotianshan Shales of Chiungchussu (Qiongzhusi) Formation in Yunnan, China. The fossil indicates that the animal was an armoured worm that close to the common ancestry of the phyla Phonorida, Brachiozoa and Bryozoa, which are collectively grouped into a clade called Lophophorata.
Cordaticaris is a genus of extinct hurdiid (peytoiid) radiodont that lived in what is now northern China during the middle Cambrian period. This animal was described in 2020 based on remains found in the Zhangxia Formation, located in the Shandong Province. It is differentiated from other members of its family by its unique heart-shaped frontal sclerite, and its frontal appendages bearing nine endites and seven more elongated subequal endites. This animal was important as it was the first Miaolingian aged hurdiid known from rock layers outside of laurentia, allowing paleontologists to get a better grasp of this families geographic range in life.
Zhenghecaris shankouensis is an enigmatic arthropod from the Lower Cambrian Maotianshan Shales, tentatively classified as a hurdiid (peytoiid) radiodont, and originally as a thylacocephalan. It is known from several specimens mostly preserving the carapace and eyes measuring roughly 15 cm (5.9 in) in width, which would have marked it as one of the largest thylacocephalans, behind Ostenocaris, Dollocaris and Ainiktozoon, as well as the earliest since all other thylacocephalans are Ordovician or younger. Better preserved fossils show that it was more similar to the domed sclerites of radiodonts such as Cambroraster, with two lateral spine processes on either side of the carapace, the eyes apparently fitting into the posterior notches. Additional isolated sclerites from the Chengjiang have been described as the lateral sclerites of Zhenghecaris due to their broad similarity to the P-elements of hurdiids (peytoiids), and because of their similar construction, ornamentation, and possession of two-pronged lateral spine processes. It has also been classified conservatively as Arthropoda incertae sedis, as the fragmentary remains cannot confidently be classified further due to the lack of associated appendages of trunk elements.
Shucaris is a genus of radiodont of uncertain taxonomic placement from the Lower Cambrian Maotianshan Shales in Yunnan, South China. The type and only species is S. ankylosskelos, known from multiple specimens comprising frontal appendages, multiple endites, gnathobase‐like structures, a nearly complete body, a head carapace complex, and one body flap associated with setal blades.