Archaeocyatha

Archaeocyatha (or archaeocyathids “ancient cups” /ˈɑːrkioʊsaɪəθə/) is a taxon of extinct, sessile, reef-building[1] marine organisms of warm tropical and subtropical waters that lived during the early (lower) Cambrian Period. It is believed that the centre of the Archaeocyatha origin are now located in East Siberia, where they are first known from the beginning of the Tommotian Age of the Cambrian, 525 million years ago (mya).[2] In other regions of the world, they appeared much later, during the Atdabanian, and quickly diversified into over a hundred families. They became the planet's very first reef-building animals and are an index fossil[3] for the Lower Cambrian worldwide.

Archaeocyatha
Temporal range: Tommotian - End Cambrian
Archaeocyatha
Scientific classification
Kingdom: Animalia
Phylum: Porifera (?)
(unranked): Archaeocyatha
Vologdin, 1937
Synonyms
  • Cyathospongia Okulitch, 1935
  • Pleospongia Okulitch, 1935

Preservation

The remains of Archaeocyatha are mostly preserved as carbonate structures in a limestone matrix. This means that the fossils cannot be chemically or mechanically isolated, save for some specimens that have already eroded out of their matrices, and their morphology has to be determined from thin cuts of the stone in which they were preserved.

Geological history

Today, the archaeocyathan families are recognizable by small but consistent differences in their fossilized structures: Some archaeocyathans were built like nested bowls, while others were as long as 300mm. Some archaeocyaths were solitary organisms, while others formed colonies. In the beginning of the Toyonian Age around 516 mya, the archaeocyaths went into a sharp decline. Almost all species became extinct by the Middle Cambrian, with the final-known species, Antarcticocyathus webberi, disappearing just prior to the end of the Cambrian period.[4] Their rapid decline and disappearance coincided with a rapid diversification of the Demosponges.

The archaeocyathids were important reef-builders in the early to middle Cambrian, with reefs (and indeed any accumulation of carbonates) becoming very rare after the group's extinction until the diversification of new taxa of coral reef-builders in the Ordovician.[5]

Archaeocyatha
*1 – Gap (intervallum) *2 – Central cavity *3 – Internal wall *4 – Pore (all the walls and septa have pores, not all are represented) *5 – Septum *6 – External wall *7 – Rizoid

Morphology

The typical archaeocyathid resembled a hollow horn coral. Each had a conical or vase-shaped porous skeleton of calcite similar to that of a sponge. The structure appeared like a pair of perforated, nested ice cream cones. Their skeletons consisted of either a single porous wall (Monocyathida), or more commonly as two concentric porous walls, an inner and outer wall separated by a space. Inside the inner wall was a cavity (like the inside of an empty ice cream cone). At the base, these pleosponges were held to the substrate by a holdfast. The body presumably occupied the space between the inner and outer shells (the intervallum).

Ecology

Flow tank experiments suggest that archaeocyathan morphology allowed them to exploit flow gradients, either by passively pumping water through the skeleton, or, as in present-day, extant sponges, by drawing water through the pores, removing nutrients, and expelling spent water and wastes through the pores into the central space.

Distribution

The archaeocyathans inhabited coastal areas of shallow seas. Their widespread distribution over almost the entire Cambrian world, as well as the taxonomic diversity of the species, might be explained by surmising that, like true sponges, they had a planktonic larval stage that enabled their wide spread.

Branching archaeocyath
Branching form archaeocyath from Rowland's Reef in Nevada
Archeocyathids
Archeocyathids from the Poleta formation, eastern California

Taxonomy

Their phylogenetic affiliation has been subject to changing interpretations, yet the consensus is growing that the archaeocyath was indeed a kind of sponge,[6] thus sometimes called a pleosponge. But some invertebrate paleontologists have placed them in an extinct, separate phylum, known appropriately as the Archaeocyatha.[7] However, one cladistic analysis[8] suggests that Archaeocyatha is a clade nested within the phylum Porifera (better known as the true sponges).

The clade Archaeocyatha have traditionally been divided into Regulares and Irregulares (Rowland, 2001):

  • Hetairacyathida (incertae sedis)
  • Regulares
    • Monocyathida
    • Capsulocyathida
    • Ajacicyathida
  • Irregulares
    • Thalassocyathida
    • Archaeocyathida
    • Kazakhstanicyathida

However, Okulitch (1955), who at the time regarded the archaeocyathans as outside of Porifera, divided the phylum in three classes:

  • Phylum Archaeocyatha Vologdin, 1937
    • Class Monocyathea Okulitch, 1943
    • Class Archaeocyathea Okulitch, 1943
    • Class Anthocyathea Okulitch, 1943

Notes

  1. ^ Archaeocyathid reef structures ("bioherms"), although not as massive as later coral reefs, might have been as deep as ten meters (Emiliani 1992:451).
  2. ^ Maloof, A.C. (2010). "Constraints on early Cambrian carbon cycling from the duration of the Nemakit-Daldynian–Tommotian boundary $$\delta$$13C shift, Morocco". Geology. 38 (7): 623–626. doi:10.1130/G30726.1.
  3. ^ Anderson, Dr. John R. "Paleozoic Life". Georgia Perimeter College. Archived from the original on 20 July 2011. Retrieved 6 July 2010.
  4. ^ The last-recorded archaeocyathan is a single species from the late (upper) Cambrian of Antarctica.
  5. ^ Munnecke, A.; Calner, M.; Harper, D. A. T.; Servais, T. (2010). "Ordovician and Silurian sea-water chemistry, sea level, and climate: A synopsis". Palaeogeography, Palaeoclimatology, Palaeoecology. 296 (3–4): 389–413. doi:10.1016/j.palaeo.2010.08.001.
  6. ^ Scuba divers have discovered living calcareous sponges, including one species that -- like the archaeocyathans -- is without spicules, thus morphologically similar to the archaeocyaths. Rowland, S.M. (2001). "Archaeocyatha: A history of phylogenetic interpretation". Journal of Paleontology. 75 (6): 1065–1078. doi:10.1666/0022-3360(2001)075<1065:AAHOPI>2.0.CO;2. Retrieved 2008-11-05.
  7. ^ Debrenne, F. and J. Vacelet. 1984. "Archaeocyatha: Is the sponge model consistent with their structural organization?" in Palaeontographica Americana, 54:pp358-369.
  8. ^ J. Reitner. 1990. "Polyphyletic origin of the 'Sphinctozoans'", in Rutzler, K. (ed.), New Perspectives in Sponge Biology: Proceedings of the Third International Conference on the Biology of Sponges (Woods Hole) pp. 33-42. Smithsonian Institution Press, Washington, DC.

References

  • Emiliani, Cesare. (1992). Planet Earth : Cosmology, Geology, & the Evolution of Life & the Environment. Cambridge University Press. (Paperback Edition ISBN 0-521-40949-7), p 451
  • Okulitch, V. J., 1955: Part E – Archaeocyatha and Porifera. Archaeocyatha, E1-E20 in Moore, R. C., (ed.) 1955: Treatise on Invertebrate Paleontology. Geological Society of America & University of Kansas Press, Lawrence, Kansas, 1955, xviii-E122.

External links

Cambrian Series 2

Cambrian Series 2 is the unnamed 2nd series of the Cambrian. It lies above the Terreneuvian series and below the Miaolingian. Series 2 has not been formally defined by the International Commission on Stratigraphy, lacking a precise lower boundary and subdivision into stages. The proposed lower boundary is the first appearance of trilobites which is estimated to be around 521 million years ago.

Cambrian Stage 10

Stage 10 of the Cambrian is the still unnamed third and final stage of the Furongian series. It follows the Jiangshanian and precedes the Ordovician Tremadocian stage. The proposed lower boundary is the first appearance of the trilobite Lotagnostus americanus around 489.5 million years ago, but other fossils are also being discussed (see below). The upper boundary is defined as the appearance of the conodont Iapetognathus fluctivagus which marks the beginning of the Tremadocian and is radiometrically dated as 485.4 million years ago.

Cambrian Stage 2

Stage 2 of the Cambrian is the unnamed upper stage of the Terreneuvian series. It lies atop the Fortunian and below Stage 3 of the Cambrian. It is commonly referred to as the Tommotian, after the Cambrian stratigraphy of Siberia. Neither the upper nor lower boundary has yet been defined by the International Commission on Stratigraphy.

The preferred definitions for the lower boundary are the first appearance of the molluscs Watsonella crosbyi or Aldanella attleborensis around 529 million years ago. The proposed upper boundary might be the first appearance of trilobites around 521 million years ago.Possible candidates for a GSSP include the first appearance of Watsonella crosbyi in the Zhujiaqing Formation in Yunnan, China or the Pestrotsvet Formation near the Aldan River on the Siberian Platform.

Cambrian Stage 3

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 approximately to the first appearance of trilobites, about 521 million years ago, though the globally asynchronous appearance of trilobites warrants the use of a separate, globally synchronous marker to define the base. The upper boundary and beginning of Cambrian Stage 4 is informally defined as the first appearance of the trilobite genera Olenellus or Redlichia around 514 million years ago.

Cambrian Stage 4

Cambrian Stage 4 is the still unnamed fourth stage of the Cambrian and the upper stage of Cambrian Series 2. It follows Cambrian Stage 3 and lies below the Wuliuan. The lower boundary has not been formally defined by the International Commission on Stratigraphy. One proposal is the first appearance of two trilobite genera, Olenellus or Redlichia. Another proposal is the first appearance of the trilobite species Arthricocephalus chauveaui. Both proposals will set the lower boundary close to 514 million years ago. The upper boundary corresponds to the beginning of the Wuliuan.

Canglangpuian

The Canglangpuian age is a Chinese regional subdivision of the Cambrian, corresponding approximately to Cambrian Stage 4.

Dorothy Hill

Dorothy Hill, AC, CBE, FAA, FRS (10 September 1907 – 23 April 1997) was an Australian geologist and palaeontologist, the first female professor at an Australian university, and the first female president of the Australian Academy of Science.

Dresbachian

The Dresbachian is a Maentwrogian regional stage of North America, lasting from 501 to 497 million years ago. It is part of the Upper Cambrian and is defined by four trilobite zones. It overlaps with the ICS-stages Guzhangian, Paibian and the lowest Jiangshanian.The Dresbachian overlies the Middle Cambrian Albertan series, and is the lowest stage of the Upper Cambrian Croixian series, followed by the Franconian stage. The Dresbachian extinction event, about 502 million years ago, was followed by the Cambrian–Ordovician extinction event about 485.4 million years ago.

Drumian

The Drumian is a stage of the Miaolingian Series of the Cambrian. It succeeds the Wuliuan and precedes the Guzhangian. The base is defined as the first appearance of the trilobite Ptychagnostus atavus around 504.5 million years ago. The top is defined as the first appearance of another trilobite Lejopyge laevigata around 500.5 million years ago.

The GSSP is defined in the Drumian section (39.5117°N 112.9915°W / 39.5117; -112.9915) in the Drum Mountains, Millard County, Utah, United States. The stage was also named after the Drum Mountains. The section is an outcrop of the Wheeler Formation, a succession of calcareous shales. The precise base of the Drumian is a laminated limestone 62 m (203 ft) above the base of the Wheeler Formation.

Fortunian

The Fortunian stage marks the beginning of the Phanerozoic eon, the Paleozoic era, and the Cambrian period. It is the first of the two stages of the Terreneuvian series. Its base is defined as the first appearance of the trace fossil Treptichnus pedum 541 million years ago. The top of the Fortunian which is the base of the Stage 2 of the Cambrian has not been formally defined yet, but will correspond to the appearance of an Archeocyatha species or "Small shelly fossils" approximately 529 million years ago.The name Fortunian is derived from a part of the Burin Peninsula, the town of Fortune near the GSSP and Fortune Bay.

Furongian

The Furongian is the fourth and final series of the Cambrian. It lasted from 497 to 485.4 million years ago. It succeeds the Miaolingian series of the Cambrian and precedes the Lower Ordovician Tremadocian stage. It is subdivided into three stages: the Paibian, Jiangshanian and the unnamed 10th stage of the Cambrian.

Guzhangian

The Guzhangian is an uppermost stage of the Miaolingian Series of the Cambrian. It follows the Drumian Stage and precedes the Paibian Stage of the Furongian Series. The base is defined as the first appearance of the trilobite Lejopyge laevigata around 500.5 million years ago. The Guzhangian-Paibian boundary is marked by the first appearance of the trilobite Glyptagnostus reticulatus around 497 million years ago.The name Guzhangian is derived from Guzhang County in Hunan Province of China.

The GSSP is defined in the Huaqiao Formation in Hunan, China. The precise base of the Guzhangian is a limestone layer 121.3 m above the base Huaqiao Formation at the Louyixi section (28.7200°N 109.9647°E / 28.7200; 109.9647), where Lejopyge laevigata has its first appearance.

Miaolingian

The Miaolingian is the third Series of the Cambrian period, and was formally named in 2018. It lasted from about 509 to 497 million years ago and is divided into 3 stages: the Wuliuan, the Drumian, and the Guzhangian. The Miaolingian is preceded by the unnamed Cambrian Series 2 and succeeded by the Furongian series.

Paibian

The Paibian is the lowest stage of Furongian series of the Cambrian. It follows the Guzhangian (3rd series of the Cambrian) and is succeeded by the Jiangshanian stage. The base is defined as the first appearance of the trilobite Glyptagnostus reticulatus around 497 million years ago. The top, or the base of the Jiangshanian is defined as the first appearance of the trilobite Agnostotes orientalis around 494 million years ago.

The name is derived from Paibi, a village in Hunan, China. The GSSP is defined in the "Paibi section" (Wuling Mountains, Huayuan County), an outcrop of the Huaqiao Formation. The base is the first occurrence of Glyptagnostus reticulatus which is 396 m above the base of the Huaqiao Formation at the type locality (28.3895°N 109.5257°E / 28.3895; 109.5257).

Reef

A reef is a bar of rock, sand, coral or similar material, lying beneath the surface of water.

Many reefs result from natural, abiotic processes—deposition of sand, wave erosion planing down rock outcrops, etc.—but the best known reefs are the coral reefs of tropical waters developed through biotic processes dominated by corals and coralline algae.

Artificial reefs (e.g. shipwrecks) sometimes have a role in enhancing the physical complexity of featureless sand bottoms, in order to attract a diverse assemblage of organisms, especially algae and fish.

Earth's largest reef system is the Great Barrier Reef in Australia, at a length of over 2,300 kilometres (1,400 miles).

Terreneuvian

The Terreneuvian is the lowermost and oldest series of the Cambrian geological system. Its base is defined by the first appearance datum of the trace fossil Treptichnus pedum around 541 million years ago. Its top is defined as the first appearance of trilobites in the stratigraphic record around 521 million years ago. This series was formally ratified by the International Commission on Stratigraphy in 2012.The Fortunian stage and presently unnamed Cambrian Stage 2 are the stages within this series. The Terreneuvian corresponds to the pre-trilobitic Cambrian.The name Terreneuvian is derived from Terre Neuve, a French name for the island of Newfoundland, Canada, where many rocks of this age are found, including the type section.

Treatise on Invertebrate Paleontology

The Treatise on Invertebrate Paleontology (or TIP) published by the Geological Society of America and the University of Kansas Press, is a definitive multi-authored work of some 50 volumes, written by more than 300 paleontologists, and covering every phylum, class, order, family, and genus of fossil and extant (still living) invertebrate animals. The prehistoric invertebrates are described as to their taxonomy, morphology, paleoecology, stratigraphic and paleogeographic range. However, taxa with no fossil record whatsoever have just a very brief listing.

Publication of the decades-long Treatise on Invertebrate Paleontology is a work-in-progress; and therefore it is not yet complete: For example, there is no volume yet published regarding the post-Paleozoic era caenogastropods (a molluscan group including the whelk and periwinkle). Furthermore, every so often, previously published volumes of the Treatise are revised.

Wuliuan

The Wuliuan stage is the fifth stage of the Cambrian, and the first stage of the Miaolingian Series of the Cambrian. It was formally defined by the ICS in 2018.

Its base is defined by the first appearance of the trilobite species Oryctocephalus indicus; it ends with the beginning of the Drumian stage, marked by the first appearance of the trilobite Ptychagnostus atavus around 504.5 million years ago.The 'golden spike' that formally defines the base of the period is driven into the Wuliu-Zengjiayan section of the Kaili formation, near Balang Village in the Miaoling Mountains, Guizhou, China.

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