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121-467: Before early Cambrian diversification, most organisms were relatively simple, composed of individual cells or small multicellular organisms, occasionally organized into colonies . As the rate of diversification subsequently accelerated, the variety of life became much more complex and began to resemble that of today. Almost all present-day animal phyla appeared during this period, including the earliest chordates . The seemingly rapid appearance of fossils in

242-681: A "stunning" find of lagerstätten, known as the Qingjiang biota , was reported from the Danshui river in Hubei province, China . More than 20,000 fossil specimens were collected, including many soft bodied animals such as jellyfish, sea anemones and worms, as well as sponges, arthropods and algae. In some specimens the internal body structures were sufficiently preserved that soft tissues, including muscles, gills, mouths, guts and eyes, can be seen. The remains were dated to around 518 Mya and around half of

363-465: A GSSP, but it is expected to be defined in strata marking the first appearance of trilobites in Gondwana . There was a rapid diversification of metazoans during this epoch, but their restricted geographic distribution, particularly of the trilobites and archaeocyaths , have made global correlations difficult, hence ongoing efforts to establish a GSSP. The Miaolingian is the third series/epoch of

484-601: A coelom—for example flatworms , whose organs are surrounded by unspecialized tissues . Changes in the abundance and diversity of some types of fossil have been interpreted as evidence for "attacks" by animals or other organisms. Stromatolites, stubby pillars built by colonies of microorganisms , are a major constituent of the fossil record from about 2,700  million years ago , but their abundance and diversity declined steeply after about 1,250  million years ago . This decline has been attributed to disruption by grazing and burrowing animals. Precambrian marine diversity

605-461: A colony to survive ( obligate ). For example, some carpenter bees will form colonies when a dominant hierarchy is formed between two or more nest foundresses (facultative colony), while corals are animals that are physically connected by living tissue (the coenosarc ) that contains a shared gastrovascular cavity . Unicellular and multicellular unitary organisms may aggregate to form colonies. For example, Modular organisms are those in which

726-407: A few regions. Relative dating ( A was before B ) is often assumed sufficient for studying processes of evolution, but this, too, has been difficult, because of the problems involved in matching up rocks of the same age across different continents . Therefore, dates or descriptions of sequences of events should be regarded with some caution until better data become available. In 2004, the start of

847-405: A genet (or genetic individual formed from a sexually-produced zygote ) asexually reproduces to form genetically identical clones called ramets . A clonal colony is when the ramets of a genet live in close proximity or are physically connected. Ramets may have all of the functions needed to survive on their own or be interdependent on other ramets. For example, some sea anemones go through

968-487: A head and a tail, the burrowers probably had bilateral symmetry—which would in all probability make them bilaterian animals. They fed above the sediment surface, but were forced to burrow to avoid predators. Trace fossils (burrows, etc.) are a reliable indicator of what life was around, and indicate a diversification of life around the start of the Cambrian, with the freshwater realm colonized by animals almost as quickly as

1089-545: A large (about 30 cm) organism, and would have provided great potential for diversification: Specialised feeding apparatus allows a number of different approaches to feeding and development, and creates a number of different approaches to avoid being eaten. The earliest generally accepted echinoderm fossils appeared in the Late Atdabanian ; unlike modern echinoderms, these early Cambrian echinoderms were not all radially symmetrical. These provide firm data points for

1210-401: A lesser degree, they serve as armor for protection against predators and adverse conditions of environment. Some Cloudina fossils show small holes in shells. The holes possibly are evidence of boring by predators sufficiently advanced to penetrate shells. A possible " evolutionary arms race " between predators and prey is one of the hypotheses that attempt to explain the Cambrian explosion. In

1331-409: A mass extinction, or with a massive warming resulting from the release of methane ice . Such changes may reflect a cause of the Cambrian explosion, although they may also have resulted from an increased level of biological activity—a possible result of the explosion. Despite these uncertainties, the geochemical evidence helps by making scientists focus on theories that are consistent with at least one of

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1452-582: A profound change in life on Earth ; prior to the Period, the majority of living organisms were small, unicellular and poorly preserved. Complex, multicellular organisms gradually became more common during the Ediacaran, but it was not until the Cambrian that the rapid diversification of lifeforms, known as the Cambrian explosion , produced the first representatives of most modern animal phyla . The Period

1573-861: A single-celled egg to a larva or juvenile form. The innermost layer forms the digestive tract (gut); the outermost forms skin; and the middle one forms muscles and all the internal organs except the digestive system. Most types of living animal are triploblastic—the best-known exceptions are Porifera (sponges) and Cnidaria (jellyfish, sea anemones, etc.). The bilaterians are animals that have right and left sides at some point in their life histories. This implies that they have top and bottom surfaces and, importantly, distinct front and back ends. All known bilaterian animals are triploblastic, and all known triploblastic animals are bilaterian. Living echinoderms ( sea stars , sea urchins , sea cucumbers , etc.) 'look' radially symmetrical (like wheels) rather than bilaterian, but their larvae exhibit bilateral symmetry and some of

1694-517: A sliver of continental terrane rifted from Laurentia with the narrow Taconic seaway opening between them. The remains of this terrane are now found in southern Scotland, Ireland, and Newfoundland. Intra-oceanic subduction either to the southeast of this terrane in the Iapetus, or to its northwest in the Taconic seaway, resulted in the formation of an island arc . This accreted to the terrane in

1815-558: A subduction zone was closing the narrow seaway between the North West Kunlun region of Tarim and the South West Kunlun terrane. North China lay at equatorial to tropical latitudes during the early Cambrian, although its exact position is unknown. Much of the craton was covered by shallow seas, with land in the northwest and southeast. Northern North China was a passive margin until the onset of subduction and

1936-463: A successful analysis is a hierarchy of clades —groups whose members are believed to share a common ancestor. The cladistic technique is sometimes problematic, as some features, such as wings or camera eyes , evolved more than once, convergently —this must be taken into account in analyses. From the relationships, it may be possible to constrain the date that lineages first appeared. For instance, if fossils of B or C date to X million years ago and

2057-573: A virgin queen of the colony replaces the dead queen as the primary reproductive) or colony inheritance (when a worker takes over as primary reproductive). Individuals in social colonies and modular organisms receive benefit to such a lifestyle. For example, it may be easier to seek out food, defend a nesting site, or increase competitive ability against other species. Modular organisms' ability to reproduce asexually in addition to sexually allows them unique benefits that social colonies do not have. The energy required for sexual reproduction varies based on

2178-457: Is a colony of microorganisms often comprising several species, with properties and capabilities greater than the aggregate of capabilities of the individual organisms. Colony ontogeny refers to the developmental process and progression of a colony. It describes the various stages and changes that occur within a colony from its initial formation to its mature state. The exact duration and dynamics of colony ontogeny can vary greatly depending on

2299-455: Is also unique in its unusually high proportion of lagerstätte deposits, sites of exceptional preservation where "soft" parts of organisms are preserved as well as their more resistant shells. By the end of the Cambrian, myriapods , arachnids , and hexapods started adapting to the land, along with the first plants . The term Cambrian is derived from the Latin version of Cymru ,

2420-409: Is based on living organisms, it accommodates extinct organisms poorly, if at all. The concept of stem groups was introduced to cover evolutionary "aunts" and "cousins" of living groups, and have been hypothesized based on this scientific theory. A crown group is a group of closely related living animals plus their last common ancestor plus all its descendants. A stem group is a set of offshoots from

2541-441: Is characterised by complex, sediment-penetrating Phanerozoic-type trace fossils , and its upper part by small shelly fossils. The second series/epoch of the Cambrian is currently unnamed and known as Cambrian Series 2 . It lasted from c. 521 Ma to c. 509 Ma. Its two stages are also unnamed and known as Cambrian Stage 3 , c. 521 Ma to c. 514 Ma, and Cambrian Stage 4 , c. 514 Ma to c. 509 Ma. The base of Series 2 does not yet have

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2662-405: Is controversial. If it does represent a cnidarian larva, Eolarva would represent the first fossil evidence of indirect development in metazoans in the earliest Cambrian. Medusozoans developed complex life cycles with a medusa stage during the Cambrian explosion, as evidenced by the discovery of Burgessomedusa phasmiformis . The earliest trilobite fossils are about 530 million years old, but

2783-513: Is left behind when the rock is dissolved with hydrofluoric acid . The diversity of this assemblage is similar to that of modern crustacean faunas. Analysis of fragments of feeding machinery found in the formation shows that it was adapted to feed in a very precise and refined fashion. This contrasts with most other early Cambrian arthropods, which fed messily by shovelling anything they could get their feeding appendages on into their mouths. This sophisticated and specialised feeding machinery belonged to

2904-419: Is not a fundamental division of nature, such as the difference between electrons and protons . It is simply a very high-level grouping in a classification system created to describe all currently living organisms. This system is imperfect, even for modern animals: different books quote different numbers of phyla, mainly because they disagree about the classification of a huge number of worm-like species. As it

3025-634: Is partially an artefact of missing strata at the Tommotian-type section, and most of this fauna in fact began to diversify in a series of pulses through the Nemakit-Daldynian and into the Tommotian. Colony (biology) In biology , a colony is composed of two or more conspecific individuals living in close association with, or connected to, one another. This association is usually for mutual benefit such as stronger defense or

3146-552: Is that the history of life on Earth goes back 3,850 million years : Rocks of that age at Warrawoona, Australia , were claimed to contain fossil stromatolites , stubby pillars formed by colonies of microorganisms . Fossils ( Grypania ) of more complex eukaryotic cells, from which all animals, plants and fungi are built, have been found in rocks from 1,400  million years ago , in China and Montana . Rocks dating from 580 to 543 million years ago contain fossils of

3267-624: Is the first geological period of the Paleozoic Era, and the Phanerozoic Eon . The Cambrian lasted 53.4 million years from the end of the preceding Ediacaran period 538.8 Ma (million years ago) to the beginning of the Ordovician Period 485.4 Ma. Most of the continents lay in the southern hemisphere surrounded by the vast Panthalassa Ocean . The assembly of Gondwana during the Ediacaran and early Cambrian led to

3388-649: The Altai-Sayan terranes. Some models show a convergent plate margin extending from Greater Avalonia, through the Timanide margin of Baltica, forming the Kipchak island arc offshore of southeastern Siberia and curving round to become part of the Altai-Sayan convergent margin. Along the then western margin, Late Neoproterozoic to early Cambrian rifting was followed by the development of a passive margin. To

3509-587: The Arequipa-Antofalla block united with the South American sector of Gondwana in the early Cambrian. The Kuunga Orogeny between northern ( Congo Craton , Madagascar and India ) and southern Gondwana ( Kalahari Craton and East Antarctica ), which began c. 570 Ma, continued with parts of northern Gondwana over-riding southern Gondwana and was accompanied by metamorphism and the intrusion of granites . Subduction zones , active since

3630-490: The Cambrian and Silurian periods. By 1859, leading geologists including Roderick Murchison were convinced that what was then called the lowest Silurian stratum showed the origin of life on Earth, though others, including Charles Lyell , differed. In On the Origin of Species , Darwin considered this sudden appearance of a solitary group of trilobites, with no apparent antecedents, and absent other fossils, to be "undoubtedly of

3751-581: The Ediacara biota , organisms so large that they are likely multicelled, but very unlike any modern organism. In 1948, Preston Cloud argued that a period of "eruptive" evolution occurred in the Early Cambrian, but as recently as the 1970s, no sign was seen of how the 'relatively' modern-looking organisms of the Middle and Late Cambrian arose. The intense modern interest in this "Cambrian explosion"

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3872-553: The Jiangshanian c. 494 Ma to c. 489.5 Ma, which have defined GSSPs; and the unnamed Cambrian Stage 10 , c. 489.5 Ma to 485.4 ± 1.9 Ma. The GSSP for the Cambrian–Ordovician boundary is at Green Point , western Newfoundland , Canada, and is dated at 485.4 Ma. It is defined by the appearance of the conodont Iapetognathus fluctivagus . Where these conodonts are not found the appearance of planktonic graptolites or

3993-512: The Tommotian has historically been understood to mark an explosive increase of the number and variety of fossils of molluscs, hyoliths and sponges , along with a rich complex of skeletal elements of unknown animals, the first archaeocyathids , brachiopods , tommotiids and others. Also soft-bodied extant phyla such as comb jellies , scalidophorans , entoproctans , horseshoe worms and lobopodians had armored forms. This sudden increase

4114-626: The continental flood basalts of the Kalkarindji large igneous province (LIP) began to erupt. These covered an area of > 2.1 × 10 km across northern, central and Western Australia regions of Gondwana making it one of the largest, as well as the earliest, LIPs of the Phanerozoic. The timing of the eruptions suggests they played a role in the early to middle Cambrian mass extinction . The terranes of Ganderia , East and West Avalonia , Carolinia and Meguma lay in polar regions during

4235-568: The trilobite Jujuyaspis borealis can be used. The boundary also corresponds with the peak of the largest positive variation in the δ C curve during the boundary time interval and with a global marine transgression. Major meteorite impact structures include: the early Cambrian (c. 535 Ma) Neugrund crater in the Gulf of Finland , Estonia, a complex meteorite crater about 20 km in diameter, with two inner ridges of about 7 km and 6 km diameter, and an outer ridge of 8 km that formed as

4356-472: The "Primordial Strata" was noted by William Buckland in the 1840s, and in his 1859 book On the Origin of Species , Charles Darwin discussed the then-inexplicable lack of earlier fossils as one of the main difficulties for his theory of descent with slow modification through natural selection . The long-running puzzlement about the seemingly-sudden appearance of the Cambrian fauna without evident precursor(s) centers on three key points: whether there really

4477-858: The "end" of the explosion, or at least indications that the crown groups of modern phyla were represented. Around the start of the Cambrian (about 539  million years ago ), many new types of traces first appear, including well-known vertical burrows such as Diplocraterion and Skolithos , and traces normally attributed to arthropods, such as Cruziana and Rusophycus . The vertical burrows indicate that worm-like animals acquired new behaviours, and possibly new physical capabilities. Some Cambrian trace fossils indicate that their makers possessed hard exoskeletons , although they were not necessarily mineralised. Meiofaunal as well as macrofaunal bilaterians participated in this invasion of infaunal niches. Burrows provide firm evidence of complex organisms; they are also much more readily preserved than body fossils, to

4598-609: The "explosion" looks a lot less sudden than if all these organisms represent an unrelated "experiment", and were replaced by the animal kingdom fairly soon thereafter (40 million years is "soon" by evolutionary and geological standards). The traces of organisms moving on and directly underneath the microbial mats that covered the Ediacaran sea floor are preserved from the Ediacaran period, about 565  million years ago . They were probably made by organisms resembling earthworms in shape, size and how they moved. The burrow-makers have never been found preserved, but, because they would need

4719-701: The 19 km diameter Glikson crater (c. 508 Ma) in Western Australia; the 5 km diameter Mizarai crater (500±10 Ma) in Lithuania; and the 3.2 km diameter Newporte structure (c. 500 Ma or slightly younger) in North Dakota , U.S.A. Reconstructing the position of the continents during the Cambrian is based on palaeomagnetic , palaeobiogeographic , tectonic , geological and palaeoclimatic data. However, these have different levels of uncertainty and can produce contradictory locations for

4840-550: The Amazonia region of Gondwana with a narrow Iapetus Ocean that only began to open once Gondwana was fully assembled c. 520 Ma. Those not in favour of the existence of Pannotia show the Iapetus opening during the Late Neoproterozoic, with up to c. 6,500 km (c. 4038 miles) between Laurentia and West Gondwana at the beginning of the Cambrian. Of the smaller continents, Baltica lay between Laurentia and Gondwana,

4961-516: The Cambrian and Early Ordovician. Gondwana was a massive continent, three times the size of any of the other Cambrian continents. Its continental land area extended from the south pole to north of the equator. Around it were extensive shallow seas and numerous smaller land areas. The cratons that formed Gondwana came together during the Neoproterozoic to early Cambrian. A narrow ocean separated Amazonia from Gondwana until c. 530 Ma and

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5082-461: The Cambrian system, I can give no satisfactory answer. American paleontologist Charles Walcott , who studied the Burgess Shale fauna , proposed that an interval of time, the "Lipalian", was not represented in the fossil record or did not preserve fossils, and that the ancestors of the Cambrian animals evolved during this time. Earlier fossil evidence has since been found. The earliest claim

5203-501: The Cambrian was dated to 542 Ma . In 2012, it was revised to 541 Ma then in 2022 it was changed again to 538.8 Ma. Some theory suggest Cambrian explosion occurred during the last stages of Gondwanan assembly, which is formed following Rodinia splitting, overlapped with the opening of the Iapetus Ocean between Laurentia and western Gondwana. The largest Cambrian faunal province is located around Gondwana, which extended from

5324-540: The Cambrian, Laurentia lay across or close to the equator.  It drifted south and rotated c. 20° anticlockwise during the middle Cambrian, before drifting north again in the late Cambrian. After the Late Neoproterozoic (or mid-Cambrian) rifting of Laurentia from Gondwana and the subsequent opening of the Iapetus Ocean, Laurentia was largely surrounded by passive margins with much of the continent covered by shallow seas. As Laurentia separated from Gondwana,

5445-738: The Cambrian, lasting from c. 509 Ma to c. 497 Ma, and roughly identical to the middle Cambrian in older literature [1] . It is divided into three stages: the Wuliuan c. 509 Ma to 504.5 Ma; the Drumian c. 504.5 Ma to c. 500.5 Ma; and the Guzhangian c. 500.5 Ma to c. 497 Ma. The name replaces Cambrian Series 3 and was ratified by the IUGS in 2018. It is named after the Miaoling Mountains in southeastern Guizhou Province , South China, where

5566-593: The Doushantou Formation in China and from deposits of the same age in Mongolia, although the interpretation of these fossils as spicules has been challenged. In the late Ediacaran-lowest Cambrian, numerous tube dwellings of enigmatic organisms appeared. It was organic-walled tubes (e.g. Saarina ) and chitinous tubes of the sabelliditids (e.g. Sokoloviina , Sabellidites , Paleolina ) that prospered up to

5687-544: The Ediacaran Timanian Orogeny was coming to an end. In this region the early to middle Cambrian was a time of non-deposition and followed by late Cambrian rifting and sedimentation. Its southeastern margin was also a convergent boundary , with the accretion of island arcs and microcontinents to the craton, although the details are unclear. Siberia began the Cambrian close to western Gondwana and north of Baltica. It drifted northwestwards to close to

5808-584: The GSSP marking its base is found. This is defined by the first appearance of the oryctocephalid trilobite Oryctocephalus indicus . Secondary markers for the base of the Miaolingian include the appearance of many acritarchs forms, a global marine transgression , and the disappearance of the polymerid trilobites, Bathynotus or Ovatoryctocara. Unlike the Terreneuvian and Series 2, all the stages of

5929-609: The ICS ratify rock units based on a Global Boundary Stratotype Section and Point (GSSP) from a single formation (a stratotype ) identifying the lower boundary of the unit. Currently the boundaries of the Cambrian System, three series and six stages are defined by global stratotype sections and points. The lower boundary of the Cambrian was originally held to represent the first appearance of complex life, represented by trilobites . The recognition of small shelly fossils before

6050-475: The Iapetus and from Gondwana by the Ran Ocean. It was composed of two continents, Fennoscandia and Sarmatia , separated by shallow seas. The sediments deposited in these unconformably overlay Precambrian basement rocks. The lack of coarse-grained sediments indicates low lying topography across the centre of the craton. Along Baltica's northeastern margin subduction and arc magmatism associated with

6171-419: The Miaolingian are defined by GSSPs . The olenellids , eodiscids , and most redlichiids trilobites went extinct at the boundary between Series 2 and the Miaolingian. This is considered the oldest mass extinction of trilobites. The Furongian , c. 497 Ma to 485.4 ± 1.9 Ma, is the fourth and uppermost series/epoch of the Cambrian. The name was ratified by the IUGS in 2003 and replaces Cambrian Series 4 and

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6292-423: The Neoproterozoic, extended around much of Gondwana's margins, from northwest Africa southwards round South America, South Africa , East Antarctica , and the eastern edge of West Australia. Shorter subduction zones existed north of Arabia and India. The Famatinian continental arc stretched from central Peru in the north to central Argentina in the south. Subduction beneath this proto- Andean margin began by

6413-457: The Ran Ocean (an arm of the Iapetus) opening between it and Gondwana. Siberia lay close to the western margin of Gondwana and to the north of Baltica. Annamia and South China formed a single continent situated off north central Gondwana. The location of North China is unclear. It may have lain along the northeast Indian sector of Gondwana or already have been a separate continent. During

6534-552: The Sedgwick's "Upper Cambrian", claiming all fossilised strata for "his" Silurian series. Matters were complicated further when, in 1852, fieldwork carried out by Sedgwick and others revealed an unconformity within the Silurian, with a clear difference in fauna between the two. This allowed Sedgwick to now claim a large section of the Silurian for "his" Cambrian and gave the Cambrian an identifiable fossil record. The dispute between

6655-791: The Welsh name for Wales, where rocks of this age were first studied. It was named by Adam Sedgwick in 1835, who divided it into three groups; the Lower, Middle, and Upper. He defined the boundary between the Cambrian and the overlying Silurian, together with Roderick Murchison , in their joint paper " On the Silurian and Cambrian Systems, Exhibiting the Order in which the Older Sedimentary Strata Succeed each other in England and Wales ". This early agreement did not last. Due to

6776-826: The ability to attack bigger prey. Colonies can form in various shapes and ways depending on the organism involved. For instance, the bacterial colony is a cluster of identical cells (clones). These colonies often form and grow on the surface of (or within) a solid medium, usually derived from a single parent cell. Colonies, in the context of development, may be composed of two or more unitary (or solitary) organisms or be modular organisms. Unitary organisms have determinate development (set life stages) from zygote to adult form and individuals or groups of individuals (colonies) are visually distinct. Modular organisms have indeterminate growth forms (life stages not set) through repeated iteration of genetically identical modules (or individuals), and it can be difficult to distinguish between

6897-461: The alga Volvox (a coenobium ), or multicellular, as in the phylum Bryozoa . Colonial organisms may have been the first step toward multicellular organisms . Individuals within a multicellular colonial organism may be called ramets , modules , or zooids . Structural and functional variation ( polymorphism ), when present, designates ramet responsibilities such as feeding, reproduction, and defense. To that end, being physically connected allows

7018-479: The beginning of the Tommotian . The mineralized tubes of Cloudina , Namacalathus , Sinotubulites and a dozen more of the other organisms from carbonate rocks formed near the end of the Ediacaran period from 549 to 542 million years ago , as well as the triradially symmetrical mineralized tubes of anabaritids (e.g. Anabarites , Cambrotubulus ) from uppermost Ediacaran and lower Cambrian. Ediacaran mineralized tubes are often found in carbonates of

7139-444: The body fossils of animals that are thought to have been capable of making them. While exact assignment of trace fossils to their makers is generally impossible, traces may, for example, provide the earliest physical evidence of the appearance of moderately complex animals (comparable to earthworms ). Several chemical markers indicate a drastic change in the environment around the start of the Cambrian. The markers are consistent with

7260-491: The calculated "family tree" says A was an ancestor of B and C, then A must have evolved more than X million years ago. It is also possible to estimate how long ago two living clades diverged—i.e. about how long ago their last common ancestor must have lived—by assuming that DNA mutations accumulate at a constant rate. These " molecular clocks ", however, are fallible, and provide only a very approximate timing: they are not sufficiently precise and reliable for estimating when

7381-557: The class was already quite diverse and cosmopolitan , suggesting they had been around for quite some time. The fossil record of trilobites began with the appearance of trilobites with mineral exoskeletons—not from the time of their origin. Crustaceans, one of the four great modern groups of arthropods, are very rare throughout the Cambrian. Convincing crustaceans were once thought to be common in Burgess Shale-type biotas, but none of these individuals can be shown to fall into

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7502-405: The colonial organism to distribute nutrients and energy obtained by feeding zooids throughout the colony. The hydrozoan Portuguese man o' war is a classic example of a colonial organism, one of many in the taxonomic class. A microbial colony is defined as a visible cluster of microorganisms growing on the surface of or within a solid medium, presumably cultured from a single cell. Because

7623-448: The colony as a whole and the modules within. In the latter case, modules may have specific functions within the colony. In contrast, solitary organisms do not associate with colonies; they are ones in which all individuals live independently and have all of the functions needed to survive and reproduce. Some organisms are primarily independent and form facultative colonies in reply to environmental conditions while others must live in

7744-415: The colony is clonal , with all organisms in it descending from a single ancestor (assuming no contamination ), they are genetically identical, except for any mutations (which occur at low frequencies). Obtaining such genetically identical organisms (or pure strains ) can be useful; this is done by spreading organisms on a culture plate and starting a new stock from a single resulting colony. A biofilm

7865-622: The crown group of "true crustaceans". The Cambrian record of crown-group crustaceans comes from microfossils . The Swedish Orsten horizons contain later Cambrian crustaceans, but only organisms smaller than 2 mm are preserved. This restricts the data set to juveniles and miniaturised adults. A more informative data source is the organic microfossils of the Mount Cap formation , Mackenzie Mountains, Canada. This late Early Cambrian assemblage ( 510 to 515 million years ago ) consists of microscopic fragments of arthropods' cuticle, which

7986-425: The curator of Oxford Museum , in 1698. Although their evolutionary importance was not known, on the basis of their old age, William Buckland (1784–1856) realized that a dramatic step-change in the fossil record had occurred around the base of what we now call the Cambrian. Nineteenth-century geologists such as Adam Sedgwick and Roderick Murchison used the fossils for dating rock strata, specifically for establishing

8107-604: The development of new convergent plate boundaries and continental-margin arc magmatism along its margins that helped drive up global temperatures. Laurentia lay across the equator, separated from Gondwana by the opening Iapetus Ocean . The Cambrian was a time of greenhouse climate conditions, with high levels of atmospheric carbon dioxide and low levels of oxygen in the atmosphere and seas. Upwellings of anoxic deep ocean waters into shallow marine environments led to extinction events, whilst periods of raised oxygenation led to increased biodiversity . The Cambrian marked

8228-663: The development of the Bainaimiao arc in the late Cambrian. To its south was a convergent margin with a southwest dipping subduction zone, beyond which lay the North Qinling terrane (now part of the Qinling Orogenic Belt ). South China and Annamia formed a single continent. Strike-slip movement between it and Gondwana accommodated its steady drift northwards from offshore the Indian sector of Gondwana to near

8349-409: The diagnostic features that allow taxonomists to classify more recent organisms, such as similarities to living organisms, are generally absent in the ediacarans. However, there seems little doubt that Kimberella was at least a triploblastic bilaterian animal. These organisms are central to the debate about how abrupt the Cambrian explosion was. If some were early members of the animal phyla seen today,

8470-466: The division of labor becomes more pronounced. Some individuals may specialise in tasks like foraging, defense, or tending to the brood, while others may take on general tasks within the nest. These specialised tasks can change throughout the life of a worker. At a certain point in the colony ontogeny, usually after a period of growth and maturation, the colony produces reproductives, including new virgin queens (princesses) and males. These individuals have

8591-671: The earliest echinoderms may have been bilaterally symmetrical. Porifera and Cnidaria are radially symmetrical, not bilaterian, and not triploblastic (but the common Bilateria-Cnidaria ancestor's planula larva is suspected to be bilaterally symmetric). The term Coelomate means having a body cavity (coelom) containing the internal organs. Most of the phyla featured in the debate about the Cambrian explosion are coelomates: arthropods, annelid worms, molluscs, echinoderms and chordates —the noncoelomate priapulids are an important exception. All known coelomate animals are triploblastic bilaterians, but some triploblastic bilaterian animals do not have

8712-456: The early 1970s and which views evolution as long intervals of near-stasis "punctuated" by short periods of rapid change. Other analyses, some more recent and some dating back to the 1970s, argue that complex animals similar to modern types evolved well before the start of the Cambrian. Radiometric dates for much of the Cambrian, obtained by analysis of radioactive elements contained within rocks, have only recently become available, and for only

8833-540: The early Cambrian Maotianshan shale beds of Chengjiang ( Yunnan , China ) and Sirius Passet ( Greenland ), the middle Cambrian Burgess Shale ( British Columbia , Canada ) and the late Cambrian Orsten ( Sweden ) fossil beds. While lagerstätten preserve far more than the conventional fossil record, they are far from complete. Because lagerstätten are restricted to a narrow range of environments (where soft-bodied organisms can be preserved very quickly, e.g. by mudslides), most animals are probably not represented; further,

8954-461: The early Cambrian, and high-to-mid southern latitudes by the mid to late Cambrian. They are commonly shown as an island arc-transform fault system along the northwestern margin of Gondwana north of northwest Africa and Amazonia, which rifted from Gondwana during the Ordovician. However, some models show these terranes as part of a single independent microcontinent , Greater Avalonia, lying to

9075-557: The equator as the Ægir Ocean opened between it and Baltica. Much of the continent was covered by shallow seas with extensive archaeocyathan reefs . The then northern third of the continent (present day south; Siberia has rotated 180° since the Cambrian) adjacent to its convergent margin was mountainous. From the Late Neoproterozoic to the Ordovician, a series of island arcs accreted to Siberia's then northeastern margin, accompanied by extensive arc and back-arc volcanism. These now form

9196-545: The exact details can vary depending on the species, the general progression typically involves a number of well-defined stages, detailed below. In this initial stage, a single female individual or small group of female individuals, often called the foundress(es), queen(s) (and kings for termites) or primary reproductive(s), establish a new colony. The foundresses build a basic nest structure and begin to lay eggs. The foundresses can also perform non-reproductive tasks at this early stage, such as nursing these first eggs and leaving

9317-475: The exceptional conditions that create lagerstätten probably do not represent normal living conditions. In addition, the known Cambrian lagerstätten are rare and difficult to date, while Precambrian lagerstätten have yet to be studied in detail. The sparseness of the fossil record means that organisms usually exist long before they are found in the fossil record—this is known as the Signor–Lipps effect . In 2019,

9438-516: The extent that the absence of trace fossils has been used to imply the genuine absence of large, motile, bottom-dwelling organisms. They provide a further line of evidence to show that the Cambrian explosion represents a real diversification, and is not a preservational artifact. The first Ediacaran and lowest Cambrian ( Nemakit-Daldynian ) skeletal fossils represent tubes and problematic sponge spicules. The oldest sponge spicules are monaxon siliceous, aged around 580  million years ago , known from

9559-508: The first trilobites, and Ediacara biota substantially earlier, has led to calls for a more precisely defined base to the Cambrian Period. Despite the long recognition of its distinction from younger Ordovician rocks and older Precambrian rocks, it was not until 1994 that the Cambrian system/period was internationally ratified. After decades of careful consideration, a continuous sedimentary sequence at Fortune Head, Newfoundland

9680-436: The fossil record is very incomplete, increasingly so as earlier times are considered. Despite this, they are often adequate to illustrate the broader patterns of life's history. Also, biases exist in the fossil record: different environments are more favourable to the preservation of different types of organism or parts of organisms. Further, only the parts of organisms that were already mineralised are usually preserved, such as

9801-626: The frequency and length of reproductive activity, number and size of offspring, and parental care. While solitary individuals bear all of those energy costs, individuals in some social colonies share a portion of those costs. Modular organisms save energy by using asexual reproduction during their life. Energy reserved in this way allows them to put more energy towards colony growth, regenerating lost modules (due to predation or other cause of death), or response to environmental conditions. Cambrian The Cambrian ( / ˈ k æ m b r i . ə n , ˈ k eɪ m -/ KAM -bree-ən, KAYM - )

9922-517: The globe that corresponded to the base of the Cambrian. An early date of 570 Ma quickly gained favour, though the methods used to obtain this number are now considered to be unsuitable and inaccurate. A more precise analysis using modern radiometric dating yields a date of 538.8 ± 0.2 Ma. The ash horizon in Oman from which this date was recovered corresponds to a marked fall in the abundance of carbon-13 that correlates to equivalent excursions elsewhere in

10043-424: The gravest nature" among the difficulties in his theory of natural selection. He reasoned that earlier seas had swarmed with living creatures, but that their fossils had not been found because of the imperfections of the fossil record. In the sixth edition of his book, he stressed his problem further as: To the question why we do not find rich fossiliferous deposits belonging to these assumed earliest periods prior to

10164-580: The groups that feature in the Cambrian explosion first evolved, and estimates produced by different techniques vary by a factor of two. However, the clocks can give an indication of branching rate, and when combined with the constraints of the fossil record, recent clocks suggest a sustained period of diversification through the Ediacaran and Cambrian. A phylum is the highest level in the Linnaean system for classifying organisms . Phyla can be thought of as groupings of animals based on general body plan. Despite

10285-571: The last 1 billion years may indicate an increased need for defence against predation. Other groups of small organisms from the Neoproterozoic era also show signs of antipredator defenses. A consideration of taxon longevity appears to support an increase in predation pressure around this time. In general, the fossil record shows a very slow appearance of these lifeforms in the Precambrian, with many cyanobacterial species making up much of

10406-553: The late Cambrian, triggering southeast-dipping subduction beneath the terrane itself and consequent closure of the marginal seaway. The terrane collided with Laurentia in the Early Ordovician. Towards the end of the early Cambrian, rifting along Laurentia's southeastern margin led to the separation of Cuyania (now part of Argentina) from the Ouachita embayment with a new ocean established that continued to widen through

10527-613: The late Cambrian. Along the northern margin of Gondwana, between northern Africa and the Armorican Terranes of southern Europe, the continental arc of the Cadomian Orogeny continued from the Neoproterozoic in response to the oblique subduction of the Iapetus Ocean. This subduction extended west along the Gondwanan margin and by c. 530 Ma may have evolved into a major transform fault system. At c. 511 Ma

10648-474: The likely environmental changes. Cladistics is a technique for working out the "family tree" of a set of organisms. It works by the logic that, if groups B and C have more similarities to each other than either has to group A, then B and C are more closely related to each other than either is to A. Characteristics that are compared may be anatomical , such as the presence of a notochord , or molecular , by comparing sequences of DNA or protein . The result of

10769-422: The lineage at a point earlier than the last common ancestor of the crown group; it is a relative concept, for example tardigrades are living animals that form a crown group in their own right, but Budd (1996) regarded them as also being a stem group relative to the arthropods. The term Triploblastic means consisting of three layers, which are formed in the embryo , quite early in the animal's development from

10890-512: The low northern latitudes to the high southern latitudes, just short of the South Pole. By the middle and later parts of the Cambrian, continued rifting had sent the paleocontinents of Laurentia, Baltica and Siberia on their separate ways. Fossils of organisms' bodies are usually the most informative type of evidence. Fossilization is a rare event, and most fossils are destroyed by erosion or metamorphism before they can be observed. Hence,

11011-543: The lower boundary of the Cambrian at the base of the Tommotian Stage, characterized by diversification and global distribution of organisms with mineral skeletons and the appearance of the first Archaeocyath bioherms. The Terreneuvian is the lowermost series/ epoch of the Cambrian, lasting from 538.8 ± 0.2 Ma to c. 521 Ma. It is divided into two stages: the Fortunian stage, 538.8 ± 0.2 Ma to c. 529 Ma; and

11132-482: The lowest Cambrian, the stromatolites were decimated. This allowed animals to begin colonization of warm-water pools with carbonate sedimentation. At first, it was anabaritids and Protohertzina (the fossilized grasping spines of chaetognaths ) fossils. Such mineral skeletons as shells, sclerites, thorns and plates appeared in uppermost Nemakit-Daldynian ; they were the earliest species of halkierids , gastropods , hyoliths and other rare organisms. The beginning of

11253-436: The major continents. This, together with the ongoing debate around the existence of the Neoproterozoic supercontinent of Pannotia , means that while most models agree the continents lay in the southern hemisphere, with the vast Panthalassa Ocean covering most of northern hemisphere, the exact distribution and timing of the movements of the Cambrian continents varies between models. Most models show Gondwana stretching from

11374-420: The matter into the public eye and raised questions about what the explosion represented. While differing significantly in details, both Whittington and Gould proposed that all modern animal phyla had appeared almost simultaneously in a rather short span of geological period. This view led to the modernization of Darwin's tree of life and the theory of punctuated equilibrium , which Eldredge and Gould developed in

11495-403: The nest to gather resources. This is also known as the ergonomic stage. As the eggs laid by the foundresses develop, they give rise to the first generation of workers. These workers can assume various tasks, such as foraging, brood care, and nest maintenance. Initially, the worker population is relatively small, and their tasks are not as specialised. As the colony grows, more workers emerge, and

11616-641: The oceans. Fossils known as " small shelly fauna " have been found in many parts on the world, and date from just before the Cambrian to about 10 million years after the start of the Cambrian (the Nemakit-Daldynian and Tommotian ages; see timeline ). These are a very mixed collection of fossils: spines, sclerites (armor plates), tubes, archeocyathids (sponge-like animals) and small shells very like those of brachiopods and snail-like molluscs—but all tiny, mostly 1 to 2 mm long. While small, these fossils are far more common than complete fossils of

11737-419: The organisms that produced them; crucially, they cover the window from the start of the Cambrian to the first lagerstätten: a period of time otherwise lacking in fossils. Hence, they supplement the conventional fossil record and allow the fossil ranges of many groups to be extended. The first cnidarian larvae, represented by the genus Eolarva , appeared in the Cambrian, although the identity of Eolarva as such

11858-471: The potential to leave the nest and start new colonies, ensuring the transmission of the gene pool of its natal colony. Over time, colonies may go through a senescence phase where the reproductive output declines, and the colony's overall vitality diminishes. Eventually, the colony may die off or be replaced by a new generation of reproductives. After the death of the queen in a monogyne colony, possible fates other than colony death include serial polygyny (when

11979-452: The process of pedal laceration in which a genetically identical individual is asexually produced from tissue broken off from the anemone's pedal disc. In plants, clonal colonies are created through the propagation of genetically identical individuals by stolons or rhizomes . Colonial organisms are clonal colonies composed of many physically connected, interdependent individuals. The subunits of colonial organisms can be unicellular, as in

12100-470: The result of an impact of an asteroid 1 km in diameter; the 5 km diameter Gardnos crater (500±10 Ma) in Buskerud , Norway, where post-impact sediments indicate the impact occurred in a shallow marine environment with rock avalanches and debris flows occurring as the crater rim was breached not long after impact; the 24 km diameter Presqu'ile crater (500 Ma or younger) Quebec , Canada;

12221-409: The scarcity of fossils, Sedgwick used rock types to identify Cambrian strata. He was also slow in publishing further work. The clear fossil record of the Silurian, however, allowed Murchison to correlate rocks of a similar age across Europe and Russia, and on these he published extensively. As increasing numbers of fossils were identified in older rocks, he extended the base of the Silurian downwards into

12342-471: The seemingly different external appearances of organisms, they are classified into phyla based on their internal and developmental organizations. For example, despite their obvious differences, spiders and barnacles both belong to the phylum Arthropoda, but earthworms and tapeworms , although similar in shape, belong to different phyla. As chemical and genetic testing becomes more accurate, previously hypothesised phyla are often entirely reworked. A phylum

12463-563: The shells of molluscs . Since most animal species are soft-bodied, they decay before they can become fossilised. As a result, although 30-plus phyla of living animals are known, two-thirds have never been found as fossils. The Cambrian fossil record includes an unusually high number of lagerstätten , which preserve soft tissues. These allow paleontologists to examine the internal anatomy of animals, which in other sediments are only represented by shells, spines, claws, etc.—if they are preserved at all. The most significant Cambrian lagerstätten are

12584-577: The south of North China. To the south of these the Tarim microcontinent lay between Gondwana and Siberia. Its northern margin was passive for much of the Paleozoic, with thick sequences of platform carbonates and fluvial to marine sediments resting unconformably on Precambrian basement. Along its southeast margin was the Altyn Cambro–Ordovician accretionary complex, whilst to the southwest

12705-410: The south polar region to north of the equator. Early in the Cambrian, the south pole corresponded with the western South American sector and as Gondwana rotated anti-clockwise, by the middle of the Cambrian, the south pole lay in the northwest African region. Laurentia lay across the equator, separated from Gondwana by the Iapetus Ocean . Proponents of Pannotia have Laurentia and Baltica close to

12826-409: The species and environmental conditions. Factors such as resource availability, competition, and environmental cues can influence the progression and outcome of colony development. During colony ontogeny for eusocial insects such as ants and bees, a colony goes through several distinct phases, each characterised by specific behavioural patterns, division of labor, and structural modifications. While

12947-438: The species identified at the time of reporting were previously unknown. Trace fossils consist mainly of tracks and burrows, but also include coprolites (fossil feces ) and marks left by feeding. Trace fossils are particularly significant because they represent a data source that is not limited to animals with easily fossilized hard parts, and reflects organisms' behaviour. Also, many traces date from significantly earlier than

13068-416: The start of the Cambrian. Most of this "Ediacara biota" were at least a few centimeters long, significantly larger than any earlier fossils. The organisms form three distinct assemblages, increasing in size and complexity as time progressed. Many of these organisms were quite unlike anything that appeared before or since, resembling discs, mud-filled bags, or quilted mattresses—one paleontologist proposed that

13189-414: The strangest organisms should be classified as a separate kingdom , Vendozoa. At least some may have been early forms of the phyla at the heart of the "Cambrian explosion" debate, having been interpreted as early molluscs ( Kimberella ), echinoderms ( Arkarua ) and arthropods ( Spriggina , Parvancorina , Yilingia ). Still, debate exists about the classification of these specimens, mainly because

13310-448: The stromatolite reefs and thrombolites , i.e. they could live in an environment adverse to the majority of animals. Although they are as hard to classify as most other Ediacaran organisms, they are important in two other ways. First, they are the earliest known calcifying organisms (organisms that built shells from calcium carbonate ). Secondly, these tubes are a device to rise over a substrate and competitors for effective feeding and, to

13431-693: The then north, Siberia was separated from the Central Mongolian terrane by the narrow and slowly opening Mongol-Okhotsk Ocean . The Central Mongolian terrane's northern margin with the Panthalassa was convergent, whilst its southern margin facing the Mongol-Okhotsk Ocean was passive. During the Cambrian, the terranes that would form Kazakhstania later in the Paleozoic were a series of island arc and accretionary complexes that lay along an intra-oceanic convergent plate margin to

13552-577: The traditional "Upper Cambrian". The GSSP for the base of the Furongian is in the Wuling Mountains , in northwestern Hunan Province , China. It coincides with the first appearance of the agnostoid trilobite Glyptagnostus reticulatus , and is near the beginning of a large positive δ C isotopic excursion. The Furongian is divided into three stages: the Paibian , c. 497 Ma to c. 494 Ma, and

13673-664: The two geologists and their supporters, over the boundary between the Cambrian and Silurian, would extend beyond the life times of both Sedgwick and Murchison. It was not resolved until 1879, when Charles Lapworth proposed the disputed strata belong to its own system, which he named the Ordovician. The term Cambrian for the oldest period of the Paleozoic was officially agreed in 1960, at the 21st International Geological Congress . It only includes Sedgwick's "Lower Cambrian series", but its base has been extended into much older rocks. Systems , series and stages can be defined globally or regionally. For global stratigraphic correlation,

13794-514: The underlying sediment. At the start of the Ediacaran period, much of the acritarch fauna, which had remained relatively unchanged for hundreds of millions of years, became extinct, to be replaced with a range of new, larger species, which would prove far more ephemeral. This radiation, the first in the fossil record, is followed soon after by an array of unfamiliar, large fossils dubbed the Ediacara biota, which flourished for 40 million years until

13915-569: The unnamed Stage 2, c. 529 Ma to c. 521 Ma. The name Terreneuvian was ratified by the International Union of Geological Sciences (IUGS) in 2007, replacing the previous "Cambrian Series 1". The GSSP defining its base is at Fortune Head on the Burin Peninsula, eastern Newfoundland, Canada (see Ediacaran - Cambrian boundary above). The Terreneuvian is the only series in the Cambrian to contain no trilobite fossils. Its lower part

14036-415: The west of Baltica and aligned with its eastern ( Timanide ) margin, with the Iapetus to the north and the Ran Ocean to the south. During the Cambrian, Baltica rotated more than 60° anti-clockwise and began to drift northwards. This rotation was accommodated by major strike-slip movements in the Ran Ocean between it and Gondwana. Baltica lay at mid-to-high southerly latitudes, separated from Laurentia by

14157-505: The world, and to the disappearance of distinctive Ediacaran fossils ( Namacalathus , Cloudina ). Nevertheless, there are arguments that the dated horizon in Oman does not correspond to the Ediacaran-Cambrian boundary, but represents a facies change from marine to evaporite-dominated strata – which would mean that dates from other sections, ranging from 544 to 542 Ma, are more suitable. *Most Russian paleontologists define

14278-456: Was a mass diversification of complex organisms over a relatively short period during the early Cambrian, what might have caused such rapid change, and what it would imply about the origin of animal life. Interpretation is difficult, owing to a limited supply of evidence based mainly on an incomplete fossil record and chemical signatures remaining in Cambrian rocks. The first discovered Cambrian fossils were trilobites , described by Edward Lhuyd ,

14399-478: Was dominated by small fossils known as acritarchs . This term describes almost any small organic-walled fossil—from the egg cases of small metazoans to resting cysts of many different kinds of green algae . After appearing around 2,000  million years ago , acritarchs underwent a boom around 1,000  million years ago , increasing in abundance, diversity, size, complexity of shape, and especially size and number of spines. Their increasingly spiny forms in

14520-444: Was settled upon as a formal base of the Cambrian Period, which was to be correlated worldwide by the earliest appearance of Treptichnus pedum . Discovery of this fossil a few metres below the GSSP led to the refinement of this statement, and it is the T. pedum ichnofossil assemblage that is now formally used to correlate the base of the Cambrian. This formal designation allowed radiometric dates to be obtained from samples across

14641-717: Was sparked by the work of Harry B. Whittington and colleagues, who, in the 1970s, reanalysed many fossils from the Burgess Shale and concluded that several were as complex as, but different from, any living animals. The most common organism, Marrella , was clearly an arthropod , but not a member of any known arthropod class . Organisms such as the five-eyed Opabinia and spiny slug-like Wiwaxia were so different from anything else known that Whittington's team assumed they must represent different phyla, seemingly unrelated to anything known today. Stephen Jay Gould 's popular 1989 account of this work, Wonderful Life , brought

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