Maotianshan shales

In the Maotianshan shales is a lower Cambrian "Konservat Lagerstätte" named for Maotianshan Hill (zh-stp|t=linktext|帽|天|山|s=linktext|帽|天|山|p=Màotiānshān) in Chengjiang County, Yunnan Province, China. More commonly, the extremely diverse assemblage of organisms are referred to as the Chengjiang biota for the multiple scattered fossil sites in Chengjiang. The Chengjiang fauna occur in outcrops of the Qiongzhusi Formation that date to between 525 and Ma|520 - a period situated in the middle of the early Cambrian epoch and at least some 10 million years older than the Burgess Shale. The Maotianshan shale is 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.

History and scientific significance

Although fossils from the region have been known from the early part of the twentieth century, Chengjiang was first recognized for its exquisite states of preservation with the 1984 discovery of the naraoiid "Misszhouia", a soft-bodied relative of trilobites. Since then, the locality has been intensively studied by scientists from throughout the world, yielding a constant flow of new discoveries and an extensive literature expressing scientific debate surrounding the interpretation of discoveries. Over this time, various taxa have been revised or re-assigned to different groups. Interpretations have led to many refinements of the phylogeny of various groups and even the erection of the new phylum Vetulicolia of primitive deuterostomes. The Chengjiang biota already has all the animal groups found in the Burgess Shale; however, since it is ten million years older, it more strongly supports the deduction that metazoans diversified earlier or faster in the early Cambrian than does the Burgess Shale fauna alone. The preservation of an extremely diverse faunal assemblage renders the Maotianshan shale the world’s most important locality for understanding the evolution of early multi-cellular life, and particularly the members of phylum Chordata, which includes all vertebrates. The Chengjiang fossils comprise the oldest diverse metazoan assemblage above the Proterozoic-Phanerozoic transition, and thus the fossil record’s best data source for understanding the apparently rapid diversification of life known as the Cambrian Explosion.

Preservation and Taphonomy

Fossils occur in a section of mudstone fifty meters thick in the Yuanshan Member of the Qiongzhusi Formation. The Yuanshan Member is extensive, covering tens of thousands of square kilometers of eastern Yunnan Province, where there are numerous, scattered outcrops yielding fossils. Studies of the strata are consistent with a tropical environment with sea level changes and tectonic activity. The region is believed to have been a shallow sea with a muddy bottom. The preserved fauna is primarily benthic and was likely buried by periodic turbidity currents, since most fossils do not show evidence of post mortem transport. Like the younger Burgess Shale fossils, the paleo-environment enabled preservation of non-mineralized, soft body parts. Fossils are found in thin layers less than an inch thick. The soft parts are preserved as aluminosilicate films, often with high oxidized iron content, and often exhibiting exquisite details.

The Chengjiang beds are very deeply weathered, as evidenced by their low specific gravity (i.e. they are very lightweight).cite journal
journal=Geology
year=2008]

Fauna

The Chengjiang biota comprises an extremely diverse faunal assembly, with some 185 species described in the literature as of June 2006. Of these, nearly half are arthropods, few of which had the hard, mineral-reinforced exoskeletons characteristic of all later arthropoda; only about 3% of the organisms known from Chengjiang have hard shells, and most of those are the trilobites, of which there are five species, all of which have been found with traces of legs, antennae, and other soft body parts, an exceedingly rare occurrence in the fossil record. Phylum Porifera (sponges; 15 species) and Priapulida (16 species) are also well represented. Other phyla represented are Brachiopoda, Chaetognatha, Cnidaria, Ctenophora, Echinodermata, Hyolitha, Nematomorpha, Phoronida, Protista, and Chordata. About one in eight animals are problematic forms of uncertain affinity, some of which may have been evolutionary experiments that survived for only a brief period as benthic environments rapidly changed in the Cambrian. Chengjiang is the richest source of the lobopodia, often considered a distinct phylum, with six genera represented: "Luolishania", "Paucipodia", "Cardiodictyon"," Hallucigenia" (also known from the Burgess Shale), "Microdictyon", and "Onychodictyon".

Perhaps the most important fossils from Chengjiang are eight possible members of phylum Chordata, the phylum to which all vertebrates belong. The most famous is "Myllokunmingia", possibly a very primitive agnathid (i.e., jawless fish). Similar to "Myllokunmingia" is "Haikouichthys ercaicunensis", another primitive fish-like animal.

The enigmatic "Yunnanozoon lividum" is considered to be the earliest hemichordate, possessing many of the characteristic chordate features and providing an anatomical link between invertebrates and chordates. "Haikouella lanceolata" is described to be the earliest craniate-like chordate. This fish-like animal has many similarities to "Y. lividum", but also differs in several aspects: it has a discernible heart, dorsal and ventral aorta, gill filaments, and a notochord (neural chord).

At present, there is no agreement as to the systematic placement of the Vetulicola, represented by seven species from Chengjiang: originally described as crustacean arthropods, the Vetulicola were later erected as a new phylum of primitive deuterostomes by D.G. Shu et al. (Shu 2001). Another researcher places them with the urochordates, based on putative affinity with the Phylum Chordata. They are thought to have been swimmers that either were filter feeders or detritivores.

Some two dozen animals from the Chengjiang biota are problematic regarding phylogenetic assignment. Among these, "Anomalocaris saron", the alleged predatory terror of the early Cambrian, is the most famous. Shu (2006) recently described "Stromatoveris psygmoglena" as a possible bilateran missing link between Ediacaran fronds and Cambrian ctenophores.

List of Chengjiang Biota species by phylum

=Kingdom Animalia=

Phylum Arthropoda - 77 species
*"Acanthomeridion serratum"
*"Alalcomenaeus"
*"Almenia spinosa"
*"Anomalocaris saron"
*"Anomalocaris symbrachiata"
*"Apiocephalus elegans"
*"Branchiocaris yunnanensis"
*"Canadaspis laevigata"
*"Chengjiangocaris longiformis"
*"Chuandianella ovata"
*"Cindarella eucalla"
*"Clypecaris pteroidea"
*"Combinivalvula chengjiangensis"
*"Comptaluta inflata"
*"Comptulata leshanensis"
*"Cyathocepalus bispinosus"
*"Dianchia mirabilis"
*"Diplopyge forcipatus"
*"Diplopyge minutus"
*"Dongshanocaris foliiformis"
*"Ercaia minuscula"
*"Ercaicunia multinodosa"
*"Forfexicaris valida"
*"Forticeps foliosa"
*"Fuxianhuia protensa"
*"Glossocaris occulatus"
*"Haikoucaris ercaiensis"
*"Isoxys auritus"
*"Isoxys curvirostratus"
*"Isoxys paradoxus"
*"Jianfengia multisegmentalis"
*"Jianshania furcatus"
*"Jiucunella paulula"
*"Kuamaia lata"
*"Kuamaia muricata"
*"Kuanyangia pustulosa"
*"Kunmingella angustacostata"
*"Kunmingella douvillei"
*"Kunmingella guanshanensis"
*"Kunmingocaris bispinosus"
*"Kunyangella cheni"
*"Leanchoilia asiatica"
*"Leanchoilia illecebrosa"
*"Liangshanella liangshenensis"
*"Mafangia subscalaria"
*"Mafangocaris multinodus"
*"Malongella bituerculata"
*"Occacaris oviformis"
*"Odaraia eurypetala"
*"Ovalicephalis mirabilis"
*"Panlongia spinosa"
*"Panlongia tetranudosa"
*"Parakunmingella malongensis"
*"Parapaleomerus sinensis"
*"Pectocaris spatiosa"
*"Petalilium latus"
*"Pisinnocaris subconigera"
*"Primicaris larvaformis"
*"Pseudoiulia cambriensis"
*"Pterotum triacanthus"
*"Pygmaclypeatus daiensis"
*"Rectifacies abnormalis"
*"Rhombicalvaria acanthi"
*"Saperion glumaceum"
*"Sidneyia sinica"
*"Sinoburius lunaris"
*"Skioldia aldna"
*"Spinokunmingella typical"
*"Sunella grandis"
*"Squamacula clypeata"
*"Syrrhaptis intestinalis"
*"Urokodia aequalis"
*"Tanglangia caudata"
*"Trigoides aclis"
*"Tsunydiscus aclis"
*"Tsunyiella daindongensis"
*"Tuzoia sinensis" - a Phyllocarid
*"Waptia ovata"
*"Wutingella binodosa"
*"Xandarella spectaculum"
*"Yiliangocaris ellipticus"
*"Yunnanocaris megista"

Phylum Arthropoda - Arachnomorpha - 2 species
*"Misszhouia longicaudata"
*"Naraoia spinosa"

Phylum Arthropoda - Class Trilobita - 5 species
*"Eoredlichia intermedia"; Family Redlichiidae
*"Kuanyangia pustulosa"; Family Redlichiidae
*"Wutingaspis tingi"; Family Redlichiidae
*"Yunnanocephalus yunnanensis"; Family Yunnanocephalidae
*"Palaeolenus lantenoisi"; Family Palaeolenidae

Phylum Brachiopoda - 5 species
*"Diandongia pista"
*"Heliomedusa orienta"
*"Lingulella chengjiangensis"
*"Lingulellotreta malongensis"
*"Longtancunella chengjiangensis"

Phylum Chaetognatha - 1 species
*"Eognathacantha ercainella"

Phylum Cnidaria - 2 species
*"Priscapennamarina angusta"
*"Xianguangia sinica"

Phylum Chordata - 8 species
*"Cathaymyrus diadexus"
*"Cathaymyrus haikouensis"
*"Haikouella jianshanensis"
*"Haikouella lanceolata"
*"Haikouichthys ercaicunensis"
*"Myllokunmingia fengjiaoa"
*"Shancouclava anningense" - a Tunicate
*"Zhongxiniscus intermedius"

Phylum Ctenophora - 3 species
*"Maotianoascus octonarius"
*"Sinoascus paillatus"
*"Stromatoveris psygmoglena"

Phylum Echinodermata - 1 species
*"Cotyledion tylodes"

Phylum Hyolitha - 8 species
*"Ambrolinevitus maximus"
*"Ambrolinevitus platypluteus"
*"Ambrolinevitus ventricosus"
*"Burithes yunnanensis"
*"Glossolithes magnus"
*"Linevitus billingsi"
*"Linevitus flabellaris"
*"Linevitus opimus"

Phylum Lobopodia - 6 species
*"Cardiodictyon catenulum"
*"Hallucigenia fortis"
*"Luolishania longicruris"
*"Microdictyon sinicum"
*"Onychodictyon ferox"
*"Paucipodia inermis"

Phylum Nematomorpha - 3 species
*"Cricocosmia jinningensis"
*"Maotianshania cylindrica"
*"Palaeoscolex sinensis"

Phylum Phoronida - 1 species
*"Iotuba chengjiangensis"

Phylum Porifera - 15 species
*"Allantospongia mica"
*"Choia xiaolantianensis"
*"Choiaella radiata"
*"Hazelia" sp.
*"Leptomitella confusa"
*"Leptomitella conica"
*"Leptomitella metta"
*"Leptomitus teretiusculus"
*"Paraleptomitella dictyodroma"
*"Paraleptomitella globula"
*"Quadrolaminiella crassa"
*"Quadrolaminiella diagonalis"
*"Saetaspongia densa"
*"Sinfoflabrum antiquum"
*"Triticispongia diagonata"

Phylum Priapulida - 16 species
*"Acosmia maotiania"
*"Archotuba conoidalis"
*"Corynetis brevis"
*"Gantoucunia aspera"
*"Lagenula triolata"
*"Oligonodus specialis"
*"Paraselkirkia jinningensis"
*"Palaeopriapulites parvus"
*"Protopriapulites haikouensis"
*"Sandaokania latinodosa"
*"Selkirkia elongata"
*"Selkirkia sinica"
*"Sicyophorus rarus"
*"Xiaoheiqingella peculiaris"
*"Xishania longgiusula"
*"Yunnanopriapulus halteroformis"

Phylum Vetulicolia - 7 species
*"Banffia confusa"
*"Didazoon hoae"
*"Vetulicola cuneata"
*"Vetulicola gantoucunensis"
*"Vetulicola rectangulata"
*"Xidazoon stephanus"
*"Yunnanozoon lividum" (disputed, see Enigmatic below)

Phylum Uncertain - 4 species
*"Amplectobelua symbrachiata"
*"Anomalocaris saron"
*"Cucumericrus decoratus"
*"Parapeytoia yunnanensis "

Enigmatic - 24 species
*"Allonnia phrixothrix"
*"Amiskwia sinica"
*"Anthrotum robustus"
*"Batofasciculus ramificans"
*"Cambrotentacus sanwuia"
*"Conicula straita"
*"Dinomischus venustus"
*"Discoides abnormalis"
*"Eldonia eumorpha"
*"Facivermis yunnanicus"
*"Hippotrum spinatus"
*"Jiucunia petalina"
*"Maanshania crusticeps"
*"Macrocephalus elongates"
*"Parvulonoda dubia"
*"Phacatrum tubifer"
*"Phasangula striata"
*"Phlogites brevis"
*"Phlogites longus"
*"Priscapennamarina angusta"
*"Pristitoites bifarius"
*"Rhipitrus calvifer"
*"Rotadiscus grandis"
*"Yunnanozoon lividum" - earliest known hemichordate

Kingdom Protista (algae)

*"Fuxianospira gyrata"
*"Megaspirellus houi"
*"Sinocylindra yunnanensis"
*"Yuknessia" sp.

References

*Conway-Morris, S. (2003). The Cambrian "explosion" of metazoans and molecular biology: would Darwin be satisfied? Int J Dev Biol, 47(7-8), 505-515.
* [http://www.fossilmuseum.net/Fossil_Sites/Chengjiang.htm Fossils of the Chengjiang Maotianshan Shale] - URL retrieved September 20, 2006
*Hou, Xian-Guang; Aldridge, Richard J., Bengstrom, Jan; Siveter, David J. ;Feng, Xiang-Hong 2004; The Cambrian Fossils of Chengjang, China, Blackwell Science Ltd, 233 pp.
* [http://palaeo.gly.bris.ac.uk/Palaeofiles/Lagerstatten/chngjang/ Preservation, Taphonomy and Palaeoecology of the Chengjiang Biota] - URL retrieved Septermber 20 2006
*Shu, D-G; Luo, H-L; Conway Morris, S.; Zhang X-L; Hu, S-X; Chen, L. ;Han, J.; Zhu, M.;Li, Y; Chen, L-Z 1999, Lower Cambrian Vertebrates from South China, Nature 402 pp 42-46.
*Shu, D. G., Morris, S. C., Han, J., Chen, L., Zhang, X. L., Zhang, Z. F., et al. (2001). Primitive deuterostomes from the Chengjiang Lagerstatte (Lower Cambrian, China). Nature, 414(6862), 419-424.
*Shu, D., Morris, S. C., Zhang, Z. F., Liu, J. N., Han, J., Chen, L., et al. (2003). A new species of yunnanozoan with implications for deuterostome evolution. Science, 299(5611), 1380-1384.
*Shu, D. G., Morris, S. C., Han, J., Zhang, Z. F., & Liu, J. N. (2004). Ancestral echinoderms from the Chengjiang deposits of China. Nature, 430(6998), 422-428.
*Shu, D. G., Morris, S. C., Han, J., Li, Y., Zhang, X. L., Hua, H., et al. (2006). Lower Cambrian vendobionts from China and early diploblast evolution. Science, 312(5774), 731-734.
*Waloszek, D., & Maas, A. (2005). The evolutionary history of crustacean segmentation: a fossil-based perspective. Evol Dev, 7(6), 515-527.
*Xian-guang, H., Aldridge, R. J., Siveter, D. J., & Xiang-hong, F. (2002). New evidence on the anatomy and phylogeny of the earliest vertebrates. Proc Biol Sci, 269(1503), 1865-1869.
*Zhang, X.; Han J.; Zhang Z.; Liu H.; and Shu D. (2003) Reconsideration of the supposed naraoiid larva from the Early Cambrian Chngjiang Lagerstätte, South China. Palaeontology 46(3):447-66.
*Zhang, X. G., & Hou, X. G. (2004). Evidence for a single median fin-fold and tail in the Lower Cambrian vertebrate, Haikouichthys ercaicunensis. J Evol Biol, 17(5), 1162-1166.

See also

* Geography of China
* Stephen Jay Gould, "Wonderful Life"

External links

* cite web
url= http://www.uni-wuerzburg.de/palaeontologie/Stuff/casu30.htm
title=Chengjiang | format=Web page | accessmonthday=November 22 | accessyear=2005
* http://www.fossilmuseum.net/Fossil_Sites/Chengjiang/Chengjiang-Biota.htm