Small carbonaceous fossil

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A fragment of Silurian arthropod cuticle with attached setae, extracted by delicate acid maceration Silurian arthropod cuticle with setae.jpg
A fragment of Silurian arthropod cuticle with attached setae, extracted by delicate acid maceration

Small carbonaceous fossils (SCFs) are sub-millimetric organic remains of organisms preserved in sedimentary strata.

Contents

This category of fossils has traditionally included robust or thick-walled entities such as plant spores, acritarchs and chitinozoa, but the term 'SCFs' is usually applied to more fragile remnants of animals that can only be extracted through a delicate maceration technique. [1] SCFs are relatively widespread and abundant, and can potentially preserve both mineralized and non-mineralized parts of organisms. Since SCFs can preserve the remains of non-biomineralized organisms, they have been viewed as a relatively untapped record of animal evolution, which has the potential to circumvent some of the biases of the shelly fossil record. [1]

Extraction

SCFs are typically preserved in fine-grained siliciclastic rocks, and are too small to be fruitfully examined on bedding planes. Instead, they are extracted by dissolving the rock in acid. Traditional palynological preparations involve high-energy steps such as centrifuging that destroy large and fragile fossils. In the more delicate technique pioneered by Butterfield, [2] individual microfossils are picked from sieved acid residues by hand. The sieving stage removes crystalline residue, making it easier to extract fossils, but introduces a filter: the smallest fossils (<~40 µm) pass through the sieve and are lost. [1] Once extracted, fossils can be mounted for light or scanning electron microscopy: transmitted light illuminates internal microstructures, whereas SEM picks out surface features.

Preservation

SCFs are best preserved in environments that had anoxic conditions [3] and where the sediments have not been subject to high temperatures (limited thermal maturity); [4] the presence of oxygen is particularly deleterious at high temperatures. [5]

Biota

Scanning electron micrograph of a late Silurian chitinozoan, an enigmatic Palaeozoic SCF that can be recovered by the standard palynological technique. Scale bar: 50 um Whole chitinozoan cropped.jpg
Scanning electron micrograph of a late Silurian chitinozoan, an enigmatic Palaeozoic SCF that can be recovered by the standard palynological technique. Scale bar: 50 µm

Traditional palynological methods are designed for extracting fossilized plant spores and other resistant organic microfossils such as acritarchs and chitinozoa. By using the modified SCFs extraction technique, more delicate fossil structures can also be recovered, including fragments of animals. In particular, this technique has been applied to sediments deposited during the Cambrian Period, since there is great interest in tracking how soft-bodied animals evolved during this time interval. [1] [6] [7] [4] Animal SCFs extracted from Cambrian sediments include the minute scales of priapulid worms, [7] Wiwaxia sclerites, [8] and arthropod feeding parts, [6] for example. These organisms are not represented in the conventional (shelly) fossil record, and so the SCFs record provides data on their distribution and evolution that would not otherwise be available. Lagerstätten such as the Burgess Shale provide isolated snapshots of Palaeozoic life, whereas SCFs provide a more continuous record, albeit blighted by the fragmentary (and consequently enigmatic) nature of many of its constituents. [1] As such, SCFs can help to fill in some of the details of the fossil record outside the rare Lagerstätten sites: for instance, highlighting the rapid nature of the Cambrian explosion. [6]

Related Research Articles

<span class="mw-page-title-main">Priapulida</span> Phylum of unsegmented marine worms

Priapulida, 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, because their general shape and their extensible spiny introvert (eversible) proboscis may resemble the shape of a human penis. They live in the mud and in comparatively shallow waters up to 90 metres (300 ft) deep. Some species show a remarkable tolerance for hydrogen sulfide, anoxia and low salinity. Halicryptus spinulosus appears to prefer brackish shallow waters. They can be quite abundant in some areas. In an Alaskan bay as many as 85 adult individuals of Priapulus caudatus per square meter has been recorded, while the density of its larvae can be as high as 58,000 per square meter.

<i>Hallucigenia</i> Genus of Cambrian animals

Hallucigenia is a genus of lobopodian, known from Cambrian aged 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 appearance and eccentric history of study; when it was erected as a genus, H. sparsa was reconstructed as an enigmatic animal upside down and back to front. Lobopodians are a grade of Paleozoic panarthropods from which the velvet worms, water bears, and arthropods arose.

<span class="mw-page-title-main">Acritarch</span> Microfossils

Acritarchs are organic microfossils, known from approximately 1800 million years ago to the present. The classification is a catch all term used to refer to any organic microfossils that cannot be assigned to other groups. Their diversity reflects major ecological events such as the appearance of predation and the Cambrian explosion.

<span class="mw-page-title-main">Chitinozoan</span> Group of marine microfossils

Chitinozoa are a group of flask-shaped, organic walled marine microfossils produced by an as yet unknown organism. Common from the Ordovician to Devonian periods, the millimetre-scale organisms are abundant in almost all types of marine sediment across the globe. This wide distribution, and their rapid pace of evolution, makes them valuable biostratigraphic markers.

<i>Wiwaxia</i> Genus of Cambrian animals

Wiwaxia is a genus of soft-bodied animals that were covered in carbonaceous scales and spines that protected it from predators. Wiwaxia fossils – mainly isolated scales, but sometimes complete, articulated fossils – are known from early Cambrian and middle Cambrian fossil deposits across the globe. The living animal would have measured up to 5 cm (2 inch) when fully grown, although a range of juvenile specimens are known, the smallest being 2 millimetres (0.079 in) long.

<i>Ottoia</i> Extinct genus of priapulid worms

Ottoia is a stem-group archaeopriapulid worm known from Cambrian fossils. Although priapulid-like worms from various Cambrian deposits are often referred to Ottoia on spurious grounds, the only clear Ottoia macrofossils come from the Burgess Shale of British Columbia, which was deposited 508 million years ago. Microfossils extend the record of Ottoia throughout the Western Canada Sedimentary Basin, from the mid- to late- Cambrian. A few fossil finds are also known from China.

<span class="mw-page-title-main">Microfossil</span> Fossil that requires the use of a microscope to see it

A microfossil is a fossil that is generally between 0.001 mm and 1 mm in size, the visual study of which requires the use of light or electron microscopy. A fossil which can be studied with the naked eye or low-powered magnification, such as a hand lens, is referred to as a macrofossil.

<span class="mw-page-title-main">Halkieriid</span> Family of incertae sedis

The halkieriids are a group of fossil organisms from the Lower to Middle Cambrian. Their eponymous genus is Halkieria, which has been found on almost every continent in Lower to Mid Cambrian deposits, forming a large component of the small shelly fossil assemblages. The best known species is Halkieria evangelista, from the North Greenland Sirius Passet Lagerstätte, in which complete specimens were collected on an expedition in 1989. The fossils were described by Simon Conway Morris and John Peel in a short paper in 1990 in the journal Nature. Later a more thorough description was undertaken in 1995 in the journal Philosophical Transactions of the Royal Society of London and wider evolutionary implications were posed.

<i>Odontogriphus</i> Genus of soft-bodied animals from middle Cambrian

Odontogriphus is a genus of soft-bodied animals known from middle Cambrian Lagerstätte. Reaching as much as 12.5 centimetres (4.9 in) in length, Odontogriphus is a flat, oval bilaterian which apparently had a single muscular foot and a "shell" on its back that was moderately rigid but of a material unsuited to fossilization.

The Burgess Shale of British Columbia is famous for its exceptional preservation of mid-Cambrian organisms. Around 69 other sites have been discovered of a similar age, with soft tissues preserved in a similar, though not identical, fashion. Additional sites with a similar form of preservation are known from the Ediacaran and Ordovician periods.

<span class="mw-page-title-main">Tommotiid</span> Extinct order of brachiopods

Tommotiids are an extinct group of Cambrian invertebrates thought to be early lophophorates.

The fossils of the Burgess Shale, like the Burgess Shale itself, are fossils that formed around 505 million years ago in the mid-Cambrian period. They were discovered in Canada in 1886, and Charles Doolittle Walcott collected over 65,000 specimens in a series of field trips up to the alpine site from 1909 to 1924. After a period of neglect from the 1930s to the early 1960s, new excavations and re-examinations of Walcott's collection continue to reveal new species, and statistical analysis suggests that additional discoveries will continue for the foreseeable future. Stephen Jay Gould's 1989 book Wonderful Life describes the history of discovery up to the early 1980s, although his analysis of the implications for evolution has been contested.

Large ornamented Ediacaran microfossils are microscopic acritarchs, usually over 100 μm in diameter, which are common in sediments of the Ediacaran period, 635 to 538.8 million years ago. They largely disappear from the Ediacaran period fossil record before 560 million years ago, roughly coeval with the origin of the Ediacara biota. They differ from Palaeozoic microfossils in many respects; they are larger, often have internal contents, have a differently-constructed cell wall, and differ in shape.

<span class="mw-page-title-main">Archaeopriapulida</span> Class of marine worms

Archaeopriapulida is a group of priapulid worms known from Cambrian lagerstätte. The group is closely related to, and very similar to, the modern Priapulids. It is unclear whether it is mono- or polyphyletic. Despite a remarkable morphological similarity to their modern cousins, they fall outside of the priapulid crown group, which is not unambiguously represented in the fossil record until the Carboniferous. In addition to well-preserved body fossils, remains of several archaeopriapulid taxa are known to have been preserved primarily as organic microfossils, such as isolated scalids and pharyngeal teeth. They are probably closely related or paraphyletic to the palaeoscolecids; the relationship between these basal worms is somewhat unresolved.

<i>Selkirkia</i> Extinct genus of priapulid worms

Selkirkia is a genus of predatory, tubicolous priapulid worms known from the Middle Cambrian Burgess Shale, Ogygopsis Shale and Puncoviscana Formation. 142 specimens of Selkirkia are known from the Greater Phyllopod bed, where they comprise 0.27% of the community. In the Burgess Shale, 20% of the tapering, organic-walled tubes are preserved with the worm inside them, whereas the other 80% are empty. Whilst alive, the tubes were probably vertical, whereas trilobite-occupied tubes are horizontal.

<i>Omnidens</i> Extinct genus of Cambrian animals

Omnidens amplus, meaning "large all-tooth", is an extinct species of large Cambrian animal known only from a series of large mouth apparatus, originally mistaken as the mouthparts of anomalocaridids. When first named, it was interpreted as a giant priapulid, but is now considered a panarthropod. Its mouth apparatus closely resembles that of the smaller gilled lobopodian Pambdelurion, indicating it is likely to have been a close relative of that species, with which it may be synonymous. With a maximum estimated body length of 1.5 metres (4.9 ft), Omnidens is suggested to have been the largest known free-living Cambrian organism. Omnidens fossils are found in the Maotianshan Shales.

<span class="mw-page-title-main">Soom Shale</span> Late Ordovician lagerstatte formation in South Africa

The Soom Shale is a member of the Late Ordovician (Hirnantian) Cederberg Formation in South Africa, renowned for its remarkable preservation of soft-tissue in fossil material. Deposited in still waters, the unit lacks bioturbation, perhaps indicating anoxic conditions.

<i>Paraselkirkia</i> Genus of priapulid worms

Paraselkirkia is a genus of archaeopriapulid known from the Chengjiang biota, resembling Selkirkia.

Corynetis is a genus of archaeopriapulid known from the Chengjiang biota, and a senior synonym of Anningvermis multispinosus.

<i>Cochleatina</i> Cambrian microfossil

Cochleatina is an organic-walled microfossil known from the late Ediacaran period and early Cambrian Fortunian Stage. Cochleatina comprises a complex spiral ribbon structure, with a serrated outer margin. These spirals are frequently found embedded in an organic sheet. Cochleatina is a rare example of a fossil taxon known to span the Ediacaran–Cambrian boundary.

References

  1. 1 2 3 4 5 Butterfield, N. J.; Harvey, T. H. P. (2011). "Small carbonaceous fossils (SCFs): A new measure of early Paleozoic paleobiology". Geology. 40: 71–74. doi:10.1130/G32580.1.
  2. Butterfield, N. J. (1990). "A reassessment of the enigmatic Burgess Shale fossil Wiwaxia corrugata (Matthew) and its relationship to the polychaete Canadia spinosa Walcott". Paleobiology. 16 (3): 287–303. doi:10.1017/s0094837300010009. JSTOR   2400789. S2CID   88100863.
  3. Guilbaud, Romain; Slater, Ben; Poulton, Simon; Harvey, Thomas; Brocks, Jochen; Nettersheim, Benjamin; Butterfield, Nicholas. "Oxygen minimum zones in the early Cambrian ocean". Geochemical Perspectives Letters. 6: 33–38.
  4. 1 2 Slater, Ben J.; Harvey, Thomas H. P.; Butterfield, Nicholas J. (2018). "Small carbonaceous fossils (SCFs) from the Terreneuvian (lower Cambrian) of Baltica". Palaeontology. 61 (3): 417–439. doi: 10.1111/pala.12350 . hdl: 2381/41261 . ISSN   1475-4983.
  5. Schiffbauer, J. D.; Wallace, A. F.; Hunter, J. L.; Kowalewski, M.; Bodnar, R. J.; Xiao, S. (2012). "Thermally-induced structural and chemical alteration of organic-walled microfossils: An experimental approach to understanding fossil preservation in metasediments". Geobiology. 10 (5): 402–423. doi:10.1111/j.1472-4669.2012.00332.x. PMID   22607551. S2CID   43319350.
  6. 1 2 3 Harvey, T. H. P.; Velez, M. I.; Butterfield, N. J. (2012). "Exceptionally preserved crustaceans from western Canada reveal a cryptic Cambrian radiation". Proceedings of the National Academy of Sciences. 109 (5): 1589–1594. Bibcode:2012PNAS..109.1589H. doi: 10.1073/pnas.1115244109 . PMC   3277126 . PMID   22307616.
  7. 1 2 Smith, Martin R.; Harvey, Thomas H. P.; Butterfield, Nicholas J. (2015). "The macro- and microfossil record of the Cambrian priapulid Ottoia". Palaeontology. 58 (4): 705–721. doi: 10.1111/pala.12168 . ISSN   1475-4983.
  8. Slater, Ben J.; Harvey, Thomas H. P.; Guilbaud, Romain; Butterfield, Nicholas J. (2017). "A cryptic record of Burgess Shale-type diversity from the early Cambrian of Baltica". Palaeontology. 60 (1): 117–140. doi: 10.1111/pala.12273 . hdl: 2381/38663 . ISSN   1475-4983. S2CID   55097552.