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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.

Genus Species Material Notes Images

Acosmia

  • A. maotiania
  • 9 nearly complete specimens

A small ecdysozoan worm that was originally thought to have been a priapulid worm.

Fossil of Acosmia

Laojieella

  • L. thecata
  • One complete and one incomplete specimen

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.

Genus Species Material Notes Images

Cricocosmia

  • C. jinningensis

A genus of palaeoscolecid worm. Fossils have been found with smaller worms seemingly attached to them.

Reconstruction of Cricocosmia with parasitic Inquicus

Mafangscolex

  • M. sinensis
  • M. yunnanensis

Maotianshania

  • M. cylindrica

An extinct worm-like creature that might be an early member of the nematomorpha.

Fossil of Maotianshania

Tabelliscolex

  • T. chengjiangensis
  • T. hexagonus
  • T. maanshanensis

A genus of palaeoscolecid worm

Tylotites

  • T. petiolaris
Genus Species Material Notes Images

Corynetis

  • C. brevis
  • C. fortis

An archaeopriapulid worm known from the biota.

Reconstruction of Corynetis

Eximipriapulus

  • E. globocaudatus

A carnivorous priapulid-like worm that has been found living in the discarded shells of hyoliths.

Reconstruction of Eximipriapulus

Paraselkirkia

  • P. sinica

An archaeopriapulid worm that resembles another worm, Selkirkia, from the Burgess shale

Reconstruction of Paraselkirkia

Paratubiluchus

  • P. bicaudatus
Reconstruction of Paratubiluchus

Palaeopriapulites

  • P. parvus
Reconstruction of Palaeopriapulites

Selkirkia

  • S. elongata

A burrowing archaeopriapulid worm also known from the burgess shale.

Fossil of the relatedSelkirkia columbia

Sicyophorus

  • S. rarus

A genus of archaeopriapulid worm.

Reconstruction of Sicyophorus

Xiaoheiqingella

  • X. peculiaris

A priapulid worm thought to belong to the crown group.

Reconstruction of Xiaoheiqingella
Genus Species Material Notes Images

Antennacanthopodia

  • A. gracilis
  • Known from two specimens, making the species very rare in the Chengjiang.

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.

Reconstruction of Antennacanthopodia

Cardiodictyon

  • C. catenulum

A highly elongate armoured lobopodian, with saddle-shaped sclerites and a lightly sclerotized head.

Reconstruction of Cardiodictyon

Diania

  • D. cactiformis

A heavily armoured lobopodian.

Reconstruction of Diania

Lenisambulatrix

  • L. humboldti
  • A single specimen.

A simple, unarmoured lobopodian.

Reconstruction of Lenisambulatrix

Onychodictyon

  • O. ferox
  • O. gracilis
  • Several complete specimens.

An enigmatic genus of lobopodian, potentially related to either the Luolishaniidae or to Tardigrada. Not always monophyletic.

Reconstruction of Onychodictyon ferox

Paucipodia

  • P. inermis
  • Known from well-preserved body fossils.

A large unarmoured lobopodian, commonly found in association with mass death-beds of legless worms and Eldonia. Thought to have been a scavenger.

Reconstruction of Paucipodia
Genus Species Material Notes Images

Microdictyon

  • M. sinicum

Microdictyon is known from slcerite microfossils around the globe, while fossils from the Chengjiang are the only in the world to preserve the soft anatomy.

Fossil of Microdictyon sinicum
Genus Species Material Notes Images

Luolishania

  • L. longicruris

Type genus of Luolishaniidae. The species Miraluolishania haikouensis is considered a juvenile synonym of Luolishania.

Reconstruction of Luolishania

Facivermis

  • F. yunnanicus

Derived Luolishaniid lobopodian with five setiferous sieving appednages, and lacking any walking limbs. Sometimes discovered living in constructed tubes, buried into the sediment.

Reconstruction of Facivermis

Hallucigenia

  • H. fortis

A Luolishaniid lobopodian with a bulbous sclerotized head and two pairs of small feeding appendages, borne on a short neck.

Reconstruction of Hallucigenia fortis
Genus Species Material Notes Images

Megadictyon

  • M. haikouensis
  • Posterior-most section of the trunk is unknown.

A poorly known siberiid.

Reconstruction of Megadictyon cf. haikouensis

Jianshanopodia

  • J. decorata
  • Known from multiple large individuals, all from a singular exposure of the Chengjiang.

A giant siberiid lobopodian related to Megadictyon.

Reconstruction of Jianshanopodia
Genus Species Material Notes Images

Omnidens

  • O. amplus
  • Numerous well preserved oral cones.

A possible giant gilled lobopodian, closely related to Pambdelurion. Also present in the Xiaoshiba Biota.

Diagrammatic reconstruction of Omnidens

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.

Genus Species Material Notes Images

Cambroraster

  • C. sp.
  • Head sclerites.

Houcaris

  • H. saron
  • Isolated appendages.

Previously described as "Anomalocaris" saron.

Appendage of Houcaris saron
Innovatiocaris
  • I. maotianshanensis
  • I. multispiniformis
  • Complete fossil and isolated appendages
Previously assigned to the Anomalocaris genus, and one of the most completely known radiodonts.
Restoration of I. maotianshanensis

Laminacaris

  • L. chimera
  • Giant isolated appendages, up to nearly 30 centimetres long.

A giant radiodont combining features from different clades.

Appendage of Laminacaris

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.

Genus Species Material Notes Images

Amplectobelua

  • A. symbrachiata
  • Dozens of isolated appendages, and at least one complete body of a juvenile.

A large Amplectobeluid radiodont with long tail furcae. The frontal appendages formed a distinctive claw-like shape.

Reconstruction of Amplectobelua

Lyrarapax

  • L. trilobus
  • L. unguispinus
  • Known from appendages and complete body fossils, including neural anatomy.

Small Amplectobeluid radiodont. Some specimens of this genus represent some of the smallest known radiodonts.

Reconstruction of Lyrarapax unguispinus (left) and Lyrarapax trilobus (right)

Ramskoeldia

  • R. platyacantha
  • R. consimilis
  • Known from appendages, often associated with GLS plates, sclerites, and swimming flaps.

Large Amplectobeluid radiodonts, with crushing gnathobases.

Appendages of Ramskoeldia

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.

Genus Species Material Notes Images

Anomalocaris

  • A. cf. canadensis
  • Isolated appendages.
Appendage of A. cf. canadensis

Lenisicaris

  • L. lupata
  • Isolated appendages.

A second species is included in the genus, Lenisicaris pennsylvanica, from the Kinzers Formation.

Appendage of Lenisicaris

Shucaris

  • S. ankylosskelos
  • Isolated appendages.
Genus Species Material Notes Images
Chuandianella
  • C. ovata
  • Several body fossils and carapaces
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.
Reconstruction of Chuandianella

Combinivalvula

  • C. chengjiangensis

An enigmatic bivalved arthropod.

Cucumericrus

  • C. decorata

Multiple Specimens.

An enigmatic arthropod that has been tentatively classified as a possible radiodont.

Reconstructed trunk appendage of Cucumericrus

Cyathocephalus

  • C. bispinosus

Dianchia

  • D. mirabilis

Diplopyge

  • D. forcipatus
  • D. minutus

Dongshanocaris

  • D. foliiformis

A small arthropod that is currently "too poorly preserved to verify their identity as a valid taxa".

Erratus

  • E. sperare
  • Several carapaces
  • Eyes and other body parts

A bivalved arthropod that shows the origins of lungs and legs in early arthropods.

Restoration of E. sperare

Ercaia

  • E. miniscula

An arthropod that has been speculated to be an early crustacean.

Forfexicaris

  • F. valida

A small bivalved arthropod that somewhat resembles ostracods, but also has large, upward facing appendages with spines.

Reconstruction of Forfexicaris

Glossocaris

  • G. occulatus

Isoxys

  • I. auritus
  • I. curvirostratus
  • I. paradoxus

A basal nektonic arthropod that possessed a bivalved carapace, and large circular eyes.

Reconstruction of Isoxys curvirostratus

Jianshania

  • J. furcatus

A small, somewhat enigmatic arthropod known from the biota

Fossil of Jianshania

Kunmingocaris

  • K. bispinosus

Kylinxia

  • K. zhangi
  • Almost complete fossil
  • Isolated frontal appendages

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.

Reconstruction of Kylinxia

Mafangia

  • M. subscalaria

Mafangocaris

  • M. multinodus

Occacaris

  • O. oviformis

A predatory arthropod that had spiked frontal appendages similar to those of the megacheirans, and was previously thought to have belonged to that grouping

Reconstruction of Occacaris

Ovalicephalis

  • O. mirabilis

Parapaleomerus

  • P. sinensis

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.

Reconstructed carapace of Parapaleomerus

Pisinnocaris

  • P. subconigera

Pseudoiulia

  • P. cambriensis

A poorly known arthropod that, in 2013, was suggested to be a member of the family Kootenichelidae, alongside Kootenichela and Worthenella.

Pterotum

  • P. triacanthus

Rhombicalvaria

  • R. acanthi

Sunella

  • S. grandis

Syrrhaptis

  • S. intes

Tanglangia

  • T. caudata

Tauricornicaris

  • T. latizonae
  • T. oxygonae

A giant arthropod once misidentified as radiodont.

Trigoides

  • T. aclis

Urokodia

  • U. aequalis

An arthropod that had segmentation which resembled that of a millipede, as well as head and tail shields with thorny spikes. Recently, it was grouped in the mollisoniid order alongside Mollisonia, Thelxiope, and Corcorania, making it a stem-chelicerate.

Diagrammatic reconstruction of Urokodia

Wutingella

  • W. bindosa

Yiliangocaris

  • Y. ellipticus

Yunnanocaris

  • Y. megista

Zhenghecaris

  • Z. shankouensis
  • Sclerites

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.

Fossil of Zhenghecaris
Genus Species Material Notes Images

Branchiocaris

  • B. yunnanensis
  • Several isolated carapaces

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.

Reconstruction of the related Branchiocaris pretiosa

Canadaspis

  • C. laevigata

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.

Reconstruction of Canadaspis laevigata

Clypecaris

  • C. pteroidea
  • C. serrata

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.

Reconstruction of Clypecaris serrata

Ercaicunia

  • E. multinodosa

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.

Reconstruction of Ercaicunia

Erjiecaris

  • E. minusculo

A small euarthropod, and unlike other hymenocarines, it possibly had eyes directly on top of its carapace. It was most likely a nektobenthic animal.

Reconstruction of Erjiecaris

Pectocaris

  • P. spatiosa
  • P. eurypetala
  • P. inopinata

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

Reconstruction of Pectocaris spatiosa

Synophalos

  • S. xynos

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.

Reconstruction of a Synophalos chain
Tuzoia
  • T. sinensis
  • several carapaces and soft body parts
A large hymenocarine that possessed a semicircular bivalved carapace, and was one of the largest arthropods from the Cambrian
Life restoration of Tuzoia sp.
Jugatacaris A. agillis
  • several carapaces and soft body parts
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.
Reconstruction of Jugatacaris
Xiazhuangocaris X. chenggongensis
  • a single specimen (NIGP 172765), which only preserves the carapace and trunk regions
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.
Reconstruction of Xiazhuangocaris
Genus Species Material Notes Images

Chengjiangocaris

  • C. longiformis
  • C. kunmingensis

A Fuxianhuiid arthropod with numerous appendages. It is one of the most complete arthropods from the lower Cambrian.

Reconstruction of Chengjiangocaris

Fuxianhuia

  • F. protensa

A large arthropod named after Fuxian Lake. Some adult specimens are found closely associated with numerous juveniles, indicating a level of parental care.

Reconstruction of Fuxianhuia with associated juveniles
Genus Species Material Notes Images

Comptaluta

  • C. inflata
  • C. kailiensis

Indiana

  • I. sp.
CT-scan of a fossil of Indiana sp.

Jiucunella

  • J. paulula

Kunmingella

  • K. angustacostata
  • K. douvillei
  • K. guanshanensis
Reconstruction of Kunmingella douvillei

Kunyangella

  • K. cheni

Liangshanella

  • L. liangshanensis

Malongella

  • M. bituerculata

Parakunmingella

  • P. malongensis

Spinokunmingella

  • S. typica

Tsunyiella

  • T. daindongensis
Genus Species Material Notes Images

Haikoucaris

  • H. ercaiensis


Reconstruction of Haikoucaris
Genus Species Material Notes Images

Alalcomenaeus

  • A. cambricus

A small arthropod related to Leanchoilia, also known from the Burgess Shale.

Reconstruction of Alalcomenaeus

Leanchoilia

  • D. asiatica
  • L. illecebrosa

One of the several species of Leanchoilia, it has known juvenile specimens and the labrum morphology known.

Fossil of Leanchoilia illecebrosa
Genus Species Material Notes Images

Fortiforceps

  • F. foliosa
Reconstruction of Fortiforceps
Sklerolibyon *S. maomima
Reconstruction of Sklerolibyon

Jianfengia

  • J. multisegmentalis
Reconstruction of Jianfengia

Parapeytoia

  • P. yunnanensis

A jianfengiid once misidentified as a radiodont

Reconstruction of known elements from Parapeytoia
Genus Species Material Notes Images

Primicaris

  • P. larvaformis
Reconstruction of Primicaris
Genus Species Material Notes Images

Acanthomeridion

  • A. serratum

Multiple Specimens.

Reconstruction of the carapace of Acanthomeridion

Almenia

  • A. spinosa

Cindarella

  • C. eucalla
Reconstruction of Cindarella

Kuamaia

  • K. lata
  • K. muricata
Reconstruction of Kuamaia lata

Panlongia

  • P. spinosa
  • P. tetranudosa
Fossil of Panlognia

Pygmaclypeatus

  • P. daiensis
CT-scan of a specimen of Pygmaclypeatus

Retifacies

  • R. abnormalis
Reconstruction of Retifacies

Saperion

  • S. glumaceum

Sidneyia

  • S. minor
Reconstruction of Sidneyia minor

Sinoburius

  • S. lunaris
Reconstruction of Sinoburius lunaris

Skioldia

  • S. aldna

Squamacula

  • S. clypeata
Reconstruction of Squamacula

Xandarella

  • X. spectaculum
Reconstruction of Xandarella
Genus Species Material Notes Images

Misszhouia

  • M. longicaudata
Reconstruction of Misszhouia

Naraoia

  • N. spinosa
Reconstruction of Naraoia spinosa
Genus Species Material Notes Images

Eoredlichia

  • E. intermedia
Reconstruction of Eoredlichia

Kuanyangia

  • K. pustulosa

Palaeolenus

  • E. miniscula
Fossil pair of Palaeolenus

Tsunyidiscus

  • T. aclis
Line drawing of Tsunyidiscus niutitangensis

Wutingapsis

  • W. tingi

Yunnanocephalus

  • Y. yunnanensis
Fossil of Yunnanocephalus
Genus Species Material Notes Images

Amiskwia

  • A. sinica

Inquicus

  • I. fellatus
Reconstruction of Inquicus
Genus Species Material Notes Images

Ankalodous

  • A. ercainella

Eognathacantha

  • E. ercainella
Reconstruction of Eognathacantha

Protosagitta

  • P. spinosa
Reconstruction of Protosagitta

Phylum Annelida

[edit]
Genus Species Material Notes Images

Iotuba

  • I. chengjiangensis
Fossil of Iotuba
Genus Species Material Notes Images

Orthrozanclus

  • O. elongata
Reconstruction of Orthrozanclus elongata

Wiwaxia

  • W. papillo
Reconstruction of a juvenile Wiwaxia corrugata, which resembles W. papillo in its apparent lack of dorsal sclerite spines.
Genus Species Material Notes Images

Ambrolinevitus

  • A. maximus
  • A. platypluteus
  • A. ventricosus

Burithes

  • B. yunnanensis

Glossolithes

  • G. magnus

Linevitus

  • L. billingsi
  • L. flabellaris
  • L. opimus
Genus Species Material Notes Images

Eophonoris

  • E. chengjiangensis
Genus Species Material Notes Images

Cotyledion

  • C. tylodes
Reconstruction of Cotyledion
Genus Species Material Notes Images

Cambrocornulitus

  • C. rarus
Genus Species Material Notes Images

Diandongia

  • D. pista

Heliomedusa

  • H. orienta
Fossil of Heliomedusa orienta

Lingulella

  • L. orienta
Fossil of the related Lingulella caelata

Lingulellotreta

  • L. malongensis

Longtancunella

  • L. chengjiangensis
Genus Species Material Notes Images

Yunnanozoon

  • Y. lividium
Fossil of Yunnanozoon
Genus Species Material Notes Images

Cathaymyrus

  • C. diadexus
  • C. haikouensis
Reconstruction of C. diadexus and Vetulocystis

Haikouichthys

  • H. ercaicunensis
Reconstruction of Haikouichthys

Myllokunmingia

  • M. fengjiaoa
Reconstruction of Myllokunmingia

Zhongjianichthys

  • Z. rostratus
Reconstruction of Zhongjianichthys

Zhongxiniscus

  • Z. intermedius
Reconstruction of Zhongxiniscus
Genus Species Material Notes Images

Beidazoon

  • B. venustum
Reconstruction of Beidazoon

Didazoon

  • D. haoae
Reconstruction of Beidazoon

Pomatrum

  • P. ventralis
Reconstruction of Beidazoon

Vetulicola

  • V. cuneata
  • V. gantoucunensis
  • V. rectangulata
Reconstruction of Vetulicola rectangulata

Yuyuanozoon

  • Y. magnificissimi
Reconstruction of Yuyuanozoon
Genus Species Material Notes Images

Phlogites

  • P. brevis
  • P. longus

Senior synonym of Cheungkongella ancestralis.

Fossil of Phlogites

Rotadiscus

  • R. grandis
Reconstruction of Rotadiscus

Stellostomites

  • S. eumorphus
Senior synonym of Eldonia eumorpha.
Diagram of Stellostomites
Genus Species Material Notes Images

Dianchicystis

  • D. jianshanensis

Vetulocystis

  • V. catenata
Reconstruction of Vetulocystis (bottom)
Genus Species Material Notes Images

Cambrobranchus

  • C. pelagobenthos

Galeaplumosus

  • G. abilus

Shankouclava

  • S. anningense
  • S. shankouense
Reconstruction of Shankouclava anningense
  • Y. simplex
Genus Species Material Notes Images

Maotianoascus

  • M. octanarius
Reconstruction of Maotianoascus octanarius

Sinoascus

  • S. paillatus
Reconstruction of Sinoascus papillatus
Stromatoveris
  • S. psygmoglena
Reconstruction of Stromatoveris psygmoglena

Phylum Porifera

[edit]
Genus Species Material Notes Images

Allantospongia

  • A. mica

Choia

  • C. xiaolantianensis

Choiaella

  • C. radiata
Hazelia
  • H.

Leptomitella

  • L. confusa
  • L. conica
  • L. metta

Leptomitus

  • L. teretiusculus
Paraleptomitella
  • Paraleptomitella dictyodroma
  • Paraleptomitella globula

Quadrolaminiella

  • Quadrolaminiella crassa
  • Quadrolaminiella diagonalis

Saetaspongia

  • S. densa
Sinfoflabrum
  • S. antiquum
Fossil of Sinfoflabrum
Triticispongia
  • T. diagonata

Phylum Cnidaria

[edit]
Genus Species Material Notes Images

Archisaccophyllia

  • A. kunmingensis
Fossil of Archisaccophyllia

Nailiana

  • N. elegans

Priscapennamarina

  • P. angusta

Xianguangia

  • X. sinica
Fossil of Xianguangia

Enigmatic

[edit]
Genus Species Material Notes Images

Allonnia

  • A. phrixothrix

Anthrotum

  • A. robustus

Batofasciculus

  • B. ramificans

Cambrotentacus

  • C. sanwuia
Reconstruction of a pair of Cambrotentacus

Conicula

  • C. straita

Daihua

  • D. sanqiong
Fossil of Daihua

Dinomischus

  • D. venustus
Fossil of Dinomischus venustus

Discoides

  • D. abnormalis

Hippotrum

  • H. spinatus

Jiucunia

  • J. petalina

Maanshania

  • M. crusticeps

Macrocephalus

  • M. elongates

Petalilium

  • P. latus

Possible synonym of Nectocaris pteryx

Fossil of Petalilium

Parvulonoda

  • P. dubia

Phacatrum

  • P. tubifer

Phasangula

  • P. striata

Pristitoites

  • P. bifarius

Rhipitrus

  • R. calvifer


Kingdom Protista (algae)

[edit]
Genus Species Material Notes Images

Fuxianospira

  • F. gyrata

Megaspirellus

  • M. houi

Sinocylindra

  • S. yunnanensis

References

[edit]
  1. ^ "Chengjiang Maotianshan Shales". www.fossilmuseum.net. Archived from the original on 9 March 2021. Retrieved 10 September 2015.
  2. ^ Lipps, J. H.; Signor, P. W (1992). Origin and early evolution of the Metazoa. ISBN 978-0-306-44067-0.

Maotianshan Category:Maotianshan shales fossils