Maotianshan Shales
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The Maotianshan Shales are a series of
Early Cambrian The Cambrian Period ( ; sometimes symbolized Ꞓ) was the first geological period of the Paleozoic Era, and of the Phanerozoic Eon. The Cambrian lasted 53.4 million years from the end of the preceding Ediacaran Period 538.8 million years ago ( ...
deposits in the Chiungchussu Formation, 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 The Burgess Shale is a fossil-bearing deposit exposed in the Canadian Rockies of British Columbia, Canada. It is famous for the exceptional preservation of the soft parts of its fossils. At old (middle Cambrian), it is one of the earliest foss ...
. They take their name from Maotianshan Hill (, Literal meaning: Hat Sky Mountain) in
Chengjiang County Chengjiang (; earlier Tchinkiang) is a city located in Yuxi, Yunnan Province, China, just north of Fuxian Lake. Administrative divisions Chengjiang City has 2 subdistricts and 4 townships. ;2 subdistricts * Fenglu () * Longjie () ;4 towns C ...
,
Yunnan Yunnan , () is a landlocked Provinces of China, province in Southwest China, the southwest of the People's Republic of China. The province spans approximately and has a population of 48.3 million (as of 2018). The capital of the province is ...
Province,
China China, officially the People's Republic of China (PRC), is a country in East Asia. It is the world's most populous country, with a population exceeding 1.4 billion, slightly ahead of India. China spans the equivalent of five time zones and ...
. The most famous assemblage of organisms are referred to as the Chengjiang biota for the multiple scattered
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
sites in Chengjiang. The age of the Chengjiang Lagerstätte is locally termed Qiongzhusian, a stage correlated to the late
Atdabanian Cambrian Stage 3 is the still unnamed third stage of the Cambrian. It succeeds Cambrian Stage 2 and precedes Cambrian Stage 4, although neither its base nor top have been formally defined. The plan is for its lower boundary to correspond approx ...
Stage in
Siberia Siberia ( ; rus, Сибирь, r=Sibir', p=sʲɪˈbʲirʲ, a=Ru-Сибирь.ogg) is an extensive geographical region, constituting all of North Asia, from the Ural Mountains in the west to the Pacific Ocean in the east. It has been a part of ...
n sequences of the middle of the
Early Cambrian The Cambrian Period ( ; sometimes symbolized Ꞓ) was the first geological period of the Paleozoic Era, and of the Phanerozoic Eon. The Cambrian lasted 53.4 million years from the end of the preceding Ediacaran Period 538.8 million years ago ( ...
. The shales date to ≤. The shales also contain the slightly younger Guanshan biota from
Malong District Malong District () is a district of the city of Qujing, Yunnan Yunnan , () is a landlocked province in the southwest of the People's Republic of China. The province spans approximately and has a population of 48.3 million (as of 2018). T ...
in Yunnan.


History and scientific significance

Although fossils from the region have been known from the early part of the 10th century, Chengjiang was first recognized for its exquisite states of preservation with the 1984 discovery of the naraoiid ''
Misszhouia ''Misszhouia'' is a genus of small to average sized (up to long) marine arthropods within the Naraoiidae family, that lived during the early Cambrian period. The only species presently known is ''Misszhouia longicaudata'' (the genus is mono ...
'', a soft-bodied relative of
trilobite Trilobites (; meaning "three lobes") are extinct marine arthropods that form the class Trilobita. Trilobites form one of the earliest-known groups of arthropods. The first appearance of trilobites in the fossil record defines the base of the At ...
s. Since then, the locality has been intensively studied by scientists throughout the world, yielding a constant flow of new discoveries and triggering an extensive scientific debate surrounding the interpretation of discoveries. Over this time, taxa have been revised or reassigned to different groups. Interpretations have led to many refinements of the
phylogeny A phylogenetic tree (also phylogeny or evolutionary tree Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, MA.) is a branching diagram or a tree showing the evolutionary relationships among various biological spec ...
of groups and even the erection of the new phylum
Vetulicolia VetulicoliaThe taxon name, Vetulocolia, is derived from the type genus, ''Vetulicola'', which is a compound Latin word composed of ''vetuli'' "old" and ''cola'' "inhabitant". is a taxon (either phylum or subphylum in rank) encompassing several ex ...
of primitive
deuterostomes Deuterostomia (; in Greek) are animals typically characterized by their anus forming before their mouth during embryonic development. The group's sister clade is Protostomia, animals whose digestive tract development is more varied. Some exampl ...
. The Chengjiang biota 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 for understanding the evolution of early multi-cellular life, particularly the members of phylum
Chordata A chordate () is an animal of the phylum Chordata (). All chordates possess, at some point during their larval or adult stages, five synapomorphies, or primary physical characteristics, that distinguish them from all the other taxa. These fiv ...
, which includes all
vertebrates Vertebrates () comprise all animal taxa within the subphylum Vertebrata () ( chordates with backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the phylum Chordata, ...
. The Chengjiang fossils comprise the oldest diverse
metazoan Animals are multicellular, eukaryotic organisms in the biological kingdom Animalia. With few exceptions, animals consume organic material, breathe oxygen, are able to move, can reproduce sexually, and go through an ontogenetic stage in ...
assemblage above the
Proterozoic The Proterozoic () is a geological eon spanning the time interval from 2500 to 538.8million years ago. It is the most recent part of the Precambrian "supereon". It is also the longest eon of the Earth's geologic time scale, and it is subdivided ...
-
Phanerozoic The Phanerozoic Eon is the current geologic eon in the geologic time scale, and the one during which abundant animal and plant life has existed. It covers 538.8 million years to the present, and it began with the Cambrian Period, when anima ...
transition and, thus, the
fossil record A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved in ...
's best data source for understanding the apparently rapid diversification of life known as the
Cambrian Explosion The Cambrian explosion, Cambrian radiation, Cambrian diversification, or the Biological Big Bang refers to an interval of time approximately in the Cambrian Period when practically all major animal phyla started appearing in the fossil recor ...
. One of the most intriguing locations of the Chengjiang biota is the Haiyan Lagerstätte where hundreds of juvenile specimens have been found. This unique location offers insights into the development of most animal groups and as such is a unique deposit in the Cambrian


IUGS geological heritage site

In respect of 'the Chengjiang fossils represent ngan uparalleled record of the fundamentally important rapid diversification of metazoan life in the early Cambrian', the International Union of Geological Sciences (IUGS) included the 'Cambrian Chengjiang fossil site and lagerstätte' in its assemblage of 100 'geological heritage sites' around the world in a listing published in October 2022. The organisation defines an 'IUGS Geological Heritage Site' as 'a key place with geological elements and/or processes of international scientific relevance, used as a reference, and/or with a substantial contribution to the development of geological sciences through history.'


Preservation and taphonomy

The fossils occur in a section of mudstone thick in the Yuanshan Member of the Qiongzhusi Formation. The Yuanshan Member is extensive, covering multiple of eastern Yunnan Province, where there are many 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 The benthic zone is the ecological region at the lowest level of a body of water such as an ocean, lake, or stream, including the sediment surface and some sub-surface layers. The name comes from ancient Greek, βένθος (bénthos), meaning "t ...
and was likely buried by periodic
turbidity current A turbidity current is most typically an underwater current of usually rapidly moving, sediment-laden water moving down a slope; although current research (2018) indicates that water-saturated sediment may be the primary actor in the process. T ...
s, 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). Trace fossils are abundant.


Chengjiang 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. Most of those are the
trilobite Trilobites (; meaning "three lobes") are extinct marine arthropods that form the class Trilobita. Trilobites form one of the earliest-known groups of arthropods. The first appearance of trilobites in the fossil record defines the base of the At ...
s (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 Priapulida (priapulid worms, from Gr. πριάπος, ''priāpos'' 'Priapus' + Lat. ''-ul-'', diminutive), sometimes referred to as penis worms, is a phylum of unsegmented marine worms. The name of the phylum relates to the Greek god of fertility ...
(16 species) are also well represented. Other phyla represented are Brachiopoda,
Chaetognatha The Chaetognatha or chaetognaths (meaning ''bristle-jaws'') are a phylum of predatory marine worms that are a major component of plankton worldwide. Commonly known as arrow worms, about 20% of the known Chaetognatha species are benthic, and ca ...
,
Cnidaria Cnidaria () is a phylum under kingdom Animalia containing over 11,000 species of aquatic animals found both in freshwater and marine environments, predominantly the latter. Their distinguishing feature is cnidocytes, specialized cells that ...
,
Ctenophora Ctenophora (; ctenophore ; ) comprise a phylum of marine invertebrates, commonly known as comb jellies, that inhabit sea waters worldwide. They are notable for the groups of cilia they use for swimming (commonly referred to as "combs"), an ...
,
Echinodermata An echinoderm () is any member of the phylum Echinodermata (). The adults are recognisable by their (usually five-point) radial symmetry, and include starfish, brittle stars, sea urchins, sand dollars, and sea cucumbers, as well as the s ...
,
Hyolitha Hyoliths are animals with small conical shells, known as fossils from the Palaeozoic era. They are at least considered as lophotrochozoan, and possibly being lophophorates, a group which includes the brachiopods, while others consider them as be ...
,
Nematomorpha Nematomorpha (sometimes called Gordiacea, and commonly known as horsehair worms, hairsnakes, or Gordian worms) are a phylum of parasitoid animals superficially similar to nematode worms in morphology, hence the name. Most species range in size f ...
, Phoronida, and
Chordata A chordate () is an animal of the phylum Chordata (). All chordates possess, at some point during their larval or adult stages, five synapomorphies, or primary physical characteristics, that distinguish them from all the other taxa. These fiv ...
. Possible molluscs include ''Wiwaxia'' and ''Nectocaris''. 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 The lobopodians, members of the informal group Lobopodia (from the Greek language, Greek, meaning "blunt feet"), or the formally erected phylum Lobopoda Cavalier-Smith (1998), are panarthropods with stubby legs called lobopods, a term which may ...
, a group including many early
panarthropod Panarthropoda is a proposed animal clade containing the extant phyla Arthropoda, Tardigrada (water bears) and Onychophora (velvet worms). Panarthropods also include extinct marine legged worms known as lobopodians ("Lobopodia"), a paraphylet ...
s, with six genera represented: ''
Luolishania ''Luolishania'' is an extinct genus of lobopodian worm and known from the Lower Cambrian Chiungchussu Formation (Maotianshan Shales) of the Chengjiang County, Yunnan Province, China. A monotypic genus, it contains one species ''Luolishania longi ...
'', '' Paucipodia'', '' Cardiodictyon'',''
Hallucigenia ''Hallucigenia'' is a genus of Cambrian animal resembling worms, known from articulated fossils in Burgess Shale-type deposits in Canada and China, and from isolated spines around the world. The generic name reflects the type species' unusual ...
'' (also known from the Burgess Shale), ''
Microdictyon ''Microdictyon'' is an extinct armoured worm-like animal coated with net-like scleritic plates, known from the Early Cambrian Maotianshan shale of Yunnan China and other parts of the world. ''Microdictyon'' is part of the ill-defined taxon &ndas ...
'', and ''
Onychodictyon ''Onychodictyon'' is a genus of extinct lobopodian known from the Lower Cambrian Chengjiang Maotianshan Shales in the Yunnan Province in China. It was characterized by a stout body covered by fleshy papillae and pairs of sclerotized plates with ...
''. Perhaps the most important fossils from Chengjiang are eight possible members of phylum
Chordata A chordate () is an animal of the phylum Chordata (). All chordates possess, at some point during their larval or adult stages, five synapomorphies, or primary physical characteristics, that distinguish them from all the other taxa. These fiv ...
, the phylum to which all vertebrates belong. The most famous is ''
Myllokunmingia ''Myllokunmingia'' is a genus of basal chordate from the Lower Cambrian Maotianshan shales of China 518 to 490 mya and is thought to be a vertebrate, although this is not conclusively proven. The species M. fengjiaoa is 28 mm long and 6&n ...
'', possibly a very primitive agnathid (i.e.,
jawless fish Agnatha (, Ancient Greek 'without jaws') is an infraphylum of jawless fish in the phylum Chordata, subphylum Vertebrata, consisting of both present ( cyclostomes) and extinct (conodonts and ostracoderms) species. Among recent animals, cyclosto ...
). Similar to ''Myllokunmingia'' is ''
Haikouichthys ''Haikouichthys'' is an extinct genus of craniate (animals with notochords and distinct heads) that lived 518 million years ago, during the Cambrian explosion of multicellular life. ''Haikouichthys'' had a defined skull and other characteristi ...
ercaicunensis'', another primitive fish-like animal. The enigmatic '' Yunnanozoon lividum'' is considered to be the earliest
hemichordate Hemichordata is a phylum which consists of triploblastic, enterocoelomate, and bilaterally symmetrical marine deuterostome animals, generally considered the sister group of the echinoderms. They appear in the Lower or Middle Cambrian and includ ...
, possessing many of the characteristic chordate features and providing an anatomical link between
invertebrates Invertebrates are a paraphyletic group of animals that neither possess nor develop a vertebral column (commonly known as a ''backbone'' or ''spine''), derived from the notochord. This is a grouping including all animals apart from the chordate ...
and chordates. ''
Haikouella lanceolata ''Haikouella'' is an agnathan chordate from the Lower Cambrian Maotianshan Shales of Chengjiang County in Yunnan Province, China. An analysis in 2015 placed ''Haikouella'' as a junior synonym of ''Yunnanozoon,'' another Maotianshan shale Cambr ...
'' is described to be the earliest
craniate A craniate is a member of the Craniata (sometimes called the Craniota), a proposed clade of chordate animals with a skull of hard bone or cartilage. Living representatives are the Myxini (hagfishes), Hyperoartia (including lampreys), and the m ...
-like chordate. This fish-like animal has many similarities to ''Y. lividum'', but 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 ''Vetulicola'' is an extinct genus of marine animal from the Cambrian of China. It is the eponymous member of the enigmatic phylum Vetulicolia, which is of uncertain affinities but may belong to the deuterostomes. Description The type species ...
, 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 ''Anomalocaris'' ("unlike other shrimp", or "abnormal shrimp") is an extinct genus of Radiodonta, radiodont, an Order (biology), order of early-diverging stem-group arthropods. The first fossils of ''Anomalocaris'' were discovered in the ''Ogygop ...
saron'', the alleged predatory terror of the early Cambrian, is the most famous. Shu (2006) recently described ''
Stromatoveris ''Stromatoveris psygmoglena'' is a genus of basal petalonam from the Chengjiang Lagerstätte, Chengjiang deposits of Yunnan that was originally aligned with the fossil ''Charnia'' (strictly, the Charniomorpha) from the Ediacara biota. However, s ...
psygmoglena'' as a possible bilateran missing link between
Ediacaran The Ediacaran Period ( ) is a geological period that spans 96 million years from the end of the Cryogenian Period 635 million years ago (Mya), to the beginning of the Cambrian Period 538.8 Mya. It marks the end of the Proterozoic Eon, and th ...
fronds and Cambrian ctenophores. '' Cambrocornulitus'' had a tubicolous shell which probably was biomineralized. It shares some affinities with cornulitids and lophophorates.


Guanshan fauna

This fauna is similar and also contains
vetulicolian VetulicoliaThe taxon name, Vetulocolia, is derived from the type genus, ''Vetulicola'', which is a compound Latin word composed of ''vetuli'' "old" and ''cola'' "inhabitant". is a taxon (either phylum or subphylum in rank) encompassing several ex ...
s.


Gallery

File:Misszhouia longicaudata 2.jpg, ''
Misszhouia ''Misszhouia'' is a genus of small to average sized (up to long) marine arthropods within the Naraoiidae family, that lived during the early Cambrian period. The only species presently known is ''Misszhouia longicaudata'' (the genus is mono ...
longicaudata'' File:Heliomedusa orienta.jpg, '' Heliomedusa orienta'' File:Leanchoilia illecebrosa.jpg, ''
Leanchoilia ''Leanchoilia'' is an megacheiran arthropod known from Cambrian deposits of the Burgess Shale in Canada and the Chengjiang biota of China. It was about long and had long, whip-like feelers mounted on frontal arm-like appendages. Its internal o ...
illecebrosa''


See also

*
Geography of China China has great physical diversity. The eastern plains and southern coasts of the country consist of fertile lowlands and foothills. They are the location of most of China's agricultural output and human population. The southern areas of th ...
*
Stephen Jay Gould Stephen Jay Gould (; September 10, 1941 – May 20, 2002) was an American paleontologist, evolutionary biologist, and historian of science. He was one of the most influential and widely read authors of popular science of his generation. Gould sp ...
, '' Wonderful Life''


References


Further reading

*
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.
Preservation, Taphonomy and Palaeoecology of the Chengjiang Biota
- URL retrieved September 20, 2006 * * * * * * * * *


External links

*

at fossilmuseum.net {{Burgess Shale type preservation Geologic formations of China Cambrian System of Asia Cambrian China Shale formations Cambrian northern paleotemperate deposits Lagerstätten Paleontology in Yunnan Geography of Yuxi First 100 IUGS Geological Heritage Sites