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Silurian

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Not to be confused with Silurian (Doctor Who).
For other uses, see Silurian (disambiguation).
Silurian
443.8 ± 1.5 – 419.2 ± 3.2 Ma
Map of Earth as it appeared 430 million years ago during the Silurian period, Wenlock epoch[citation needed]
Chronology
Etymology
Name formalityFormal
Synonym(s)Gotlandian
Usage information
Celestial bodyEarth
Regional usageGlobal (ICS)
Time scale(s) usedICS Time Scale
Definition
Chronological unitPeriod
Stratigraphic unitSystem
First proposed byRoderick Murchison, 1835
Time span formalityFormal
Lower boundary definitionFAD of the Graptolite Akidograptus ascensus
Lower boundary GSSPDob's Linn, Moffat, United Kingdom
55°26′24″N 3°16′12″W / 55.4400°N 3.2700°W / 55.4400; -3.2700
Lower GSSP ratified1984[4][5]
Upper boundary definitionFAD of the Graptolite Monograptus uniformis
Upper boundary GSSPKlonk, Czech Republic
49°51′18″N 13°47′31″E / 49.8550°N 13.7920°E / 49.8550; 13.7920
Upper GSSP ratified1972[6]
Atmospheric and climatic data
Sea level above present dayAround 180 m, with short-term negative excursions[7]

The Silurian (/sɪˈljʊəri.ən, saɪ-/ sih-LURE-ee-ən, sy-)[8][9][10] is a geologic period and system spanning 24.6 million years from the end of the Ordovician Period, at 443.8 million years ago (Mya), to the beginning of the Devonian Period, 419.2 Mya.[11] The Silurian is the third and shortest period of the Paleozoic Era, and the third of twelve periods of the Phanerozoic Eon. As with other geologic periods, the rock beds that define the period's start and end are well identified, but the exact dates are uncertain by a few million years. The base of the Silurian is set at a series of major Ordovician–Silurian extinction events when up to 60% of marine genera were wiped out.

One important event in this period was the initial establishment of terrestrial life in what is known as the Silurian-Devonian Terrestrial Revolution: vascular plants emerged from more primitive land plants,[12][13] dikaryan fungi started expanding and diversifying along with glomeromycotan fungi,[14] and three groups of arthropods (myriapods, arachnids and hexapods) became fully terrestrialized.[15]

Another significant evolutionary milestone during the Silurian was the diversification of jawed fish, which include placoderms, acanthodians (which gave rise to cartilaginous fish) and osteichthyan (bony fish, further divided into lobe-finned and ray-finned fishes)，[16] although this corresponded to sharp decline of jawless fish such as conodonts and ostracoderms.

History of study

[edit]

The Silurian system was first identified by the Scottish geologist Roderick Murchison, who was examining fossil-bearing sedimentary rock strata in south Wales in the early 1830s. He named the sequences for a Celtic tribe of Wales, the Silures, inspired by his friend Adam Sedgwick, who had named the period of his study the Cambrian, from a Latin name for Wales.[17] Whilst the British rocks now identified as belonging to the Silurian System and the lands now thought to have been inhabited in antiquity by the Silures show little correlation (cf. Geologic map of Wales, Map of pre-Roman tribes of Wales), Murchison conjectured that their territory included Caer Caradoc and Wenlock Edge exposures - and that if it did not there were plenty of Silurian rocks elsewhere 'to sanction the name proposed'.[18] In 1835 the two men presented a joint paper, under the title On the Silurian and Cambrian Systems, Exhibiting the Order in which the Older Sedimentary Strata Succeed each other in England and Wales, which was the germ of the modern geological time scale.[19] As it was first identified, the "Silurian" series when traced farther afield quickly came to overlap Sedgwick's "Cambrian" sequence, however, provoking furious disagreements that ended the friendship.

The English geologist Charles Lapworth resolved the conflict by defining a new Ordovician system including the contested beds.[20] An alternative name for the Silurian was "Gotlandian" after the strata of the Baltic island of Gotland.[21]

The French geologist Joachim Barrande, building on Murchison's work, used the term Silurian in a more comprehensive sense than was justified by subsequent knowledge. He divided the Silurian rocks of Bohemia into eight stages.[22] His interpretation was questioned in 1854 by Edward Forbes,[23] and the later stages of Barrande; F, G and H have since been shown to be Devonian. Despite these modifications in the original groupings of the strata, it is recognized that Barrande established Bohemia as a classic ground for the study of the earliest Silurian fossils.

Subdivisions

[edit]
Subdivisions of the Silurian period
Epoch Age Start
(mya)		 Etymology of
Epochs and Stages		 Notes
Llandovery Rhuddanian 443.8 Cefn-Rhuddan Farm, Llandovery in Carmarthenshire, Wales
Aeronian 440.8 Cwm Coed-Aeron Farm, Wales Trefawr Track near the farm is the site of the GSSP
Telychian 438.5 Pen-lan-Telych Farm, Llandovery, Wales
Wenlock Sheinwoodian 433.4 Sheinwood village, Much Wenlock and Wenlock Edge, Shropshire, England During the Wenlock, the oldest-known tracheophytes of the genus Cooksonia, appear. The complexity of slightly later Gondwana plants like Baragwanathia, which resembled a modern clubmoss, indicates a much longer history for vascular plants, extending into the early Silurian or even Ordovician.[citation needed] The first terrestrial animals also appear in the Wenlock, represented by air-breathing millipedes from Scotland.[24]
Homerian 430.5 Homer, Shropshire, England
Ludlow Gorstian 427.4 Gorsty village near Ludlow, Shropshire, England
Ludfordian 425.6 Ludford, Shropshire, England
Přídolí — 423.0 Named after a locality at the Homolka a Přídolí nature reserve near the Prague suburb of Slivenec, Czech Republic. Přídolí is the old name of a cadastral field area.[25]

Paleogeography

[edit]
Ordovician-Silurian boundary on Hovedøya, Norway, showing brownish late Ordovician mudstone and later dark deep-water Silurian shale. The layers have been overturned by the Caledonian orogeny.

With the supercontinent Gondwana covering the equator and much of the southern hemisphere, a large ocean occupied most of the northern half of the globe.[26] The high sea levels of the Silurian and the relatively flat land (with few significant mountain belts) resulted in a number of island chains, and thus a rich diversity of environmental settings.[26]

During the Silurian, Gondwana continued a slow southward drift to high southern latitudes, but there is evidence that the Silurian icecaps were less extensive than those of the late-Ordovician glaciation. The southern continents remained united during this period. The melting of icecaps and glaciers contributed to a rise in sea level, recognizable from the fact that Silurian sediments overlie eroded Ordovician sediments, forming an unconformity. The continents of Avalonia, Baltica, and Laurentia drifted together near the equator, starting the formation of a second supercontinent known as Euramerica.

When the proto-Europe collided with North America, the collision folded coastal sediments that had been accumulating since the Cambrian off the east coast of North America and the west coast of Europe. This event is the Caledonian orogeny, a spate of mountain building that stretched from New York State through conjoined Europe and Greenland to Norway. At the end of the Silurian, sea levels dropped again, leaving telltale basins of evaporites extending from Michigan to West Virginia, and the new mountain ranges were rapidly eroded. The Teays River, flowing into the shallow mid-continental sea, eroded Ordovician Period strata, forming deposits of Silurian strata in northern Ohio and Indiana.

The vast ocean of Panthalassa covered most of the northern hemisphere. Other minor oceans include two phases of the Tethys, the Proto-Tethys and Paleo-Tethys, the Rheic Ocean, the Iapetus Ocean (a narrow seaway between Avalonia and Laurentia), and the newly formed Ural Ocean.

Fossils of the late Silurian sea bed, England

Climate and sea level

[edit]

The Silurian period was once believed to have enjoyed relatively stable and warm temperatures, in contrast with the extreme glaciations of the Ordovician before it and the extreme heat of the ensuing Devonian; however, it is now known that the global climate underwent many drastic fluctuations throughout the Silurian,[27][28] evidenced by numerous major carbon and oxygen isotope excursions during this geologic period.[29][30][31] Sea levels rose from their Hirnantian low throughout the first half of the Silurian; they subsequently fell throughout the rest of the period, although smaller scale patterns are superimposed on this general trend; fifteen high-stands (periods when sea levels were above the edge of the continental shelf) can be identified, and the highest Silurian sea level was probably around 140 metres (459 ft) higher than the lowest level reached.[26]

During this period, the Earth entered a warm greenhouse phase, supported by high CO2 levels of 4500 ppm, and warm shallow seas covered much of the equatorial land masses.[32] Early in the Silurian, glaciers retreated back into the South Pole until they almost disappeared in the middle of Silurian.[28] Layers of broken shells (called coquina) provide strong evidence of a climate dominated by violent storms generated then as now by warm sea surfaces.[33]

Perturbations

[edit]

The climate and carbon cycle appear to be rather unsettled during the Silurian, which had a higher frequency of isotopic excursions (indicative of climate fluctuations) than any other period.[26] The Ireviken event, Mulde event, and Lau event each represent isotopic excursions following a minor mass extinction[34] and associated with rapid sea-level change. Each one leaves a similar signature in the geological record, both geochemically and biologically; pelagic (free-swimming) organisms were particularly hard hit, as were brachiopods, corals, and trilobites, and extinctions rarely occur in a rapid series of fast bursts.[26][31] The climate fluctuations are best explained by a sequence of glaciations, but the lack of tillites in the middle to late Silurian make this explanation problematic.[35]

Flora and fauna

[edit]
Main article: Silurian-Devonian Terrestrial Revolution

The Silurian period has been viewed by some palaeontologists as an extended recovery interval following the Late Ordovician mass extinction (LOME), which interrupted the cascading increase in biodiversity that had continuously gone on throughout the Cambrian and most of the Ordovician.[36]

The Silurian was the first period to see megafossils of extensive terrestrial biota in the form of moss-like miniature forests along lakes and streams and networks of large, mycorrhizal nematophytes, heralding the beginning of the Silurian-Devonian Terrestrial Revolution.[12][13][37] However, the land fauna did not have a major impact on the Earth until it diversified in the Devonian.[26]

Diorama of a Silurian seafloor

The first fossil records of vascular plants, that is, land plants with tissues that carry water and food, appeared in the second half of the Silurian Period.[38] The earliest-known representatives of this group are Cooksonia. Most of the sediments containing Cooksonia are marine in nature. Preferred habitats were likely along rivers and streams. Baragwanathia appears to be almost as old, dating to the early Ludlow (420 million years)[needs update?] and has branching stems and needle-like leaves of 10–20 centimetres (3.9–7.9 in). The plant shows a high degree of development in relation to the age of its fossil remains. Fossils of this plant have been recorded in Australia,[39][40] Canada,[41] and China.[42] Eohostimella heathana is an early, probably terrestrial, "plant" known from compression fossils[43] of Early Silurian (Llandovery) age.[44] The chemistry of its fossils is similar to that of fossilised vascular plants, rather than algae.[43]

Fossils that are considered as terrestrial animals are also known from the Silurian. The definitive oldest record of millipede ever known is Kampecaris obanensis and Archidesmus sp. from the late Silurian (425 million years ago) of Kerrera.[45] There are also other millipedes, centipedes, and trigonotarbid arachnoids known from Ludlow (420 million years ago).[45][46][47] Predatory invertebrates would indicate that simple food webs were in place that included non-predatory prey animals. Extrapolating back from Early Devonian biota, Andrew Jeram et al. in 1990[48] suggested a food web based on as-yet-undiscovered detritivores and grazers on micro-organisms.[49] Millipedes from Cowie Formation such as Cowiedesmus and Pneumodesmus were considered as the oldest millipede from the middle Silurian at 428-430 million years ago,[24][50][51] although the age of this formation is later reinterpreted to be from the early Devonian instead by some researchers.[52][53] Regardless, Pneumodesmus is still an important fossil as the oldest definitive evidence of spiracles to breath in the air.[45]

Life restoration of sarcopterygian Sparalepis tingi and other fauna from the Silurian of Yunnan

The first bony fish, the Osteichthyes, appeared, represented by the Acanthodians covered with bony scales. Fish reached considerable diversity and developed movable jaws, adapted from the supports of the front two or three gill arches. A diverse fauna of eurypterids (sea scorpions)—some of them several meters in length—prowled the shallow Silurian seas and lakes of North America; many of their fossils have been found in New York state. Brachiopods were abundant and diverse, with the taxonomic composition, ecology, and biodiversity of Silurian brachiopods mirroring Ordovician ones.[54] Brachiopods that survived the LOME developed novel adaptations for environmental stress,[55] and they tended to be endemic to a single palaeoplate in the mass extinction's aftermath, but expanded their range afterwards.[56] The most abundant brachiopods were atrypids and pentamerides;[57] atrypids were the first to recover and rediversify in the Rhuddanian after LOME,[58] while pentameride recovery was delayed until the Aeronian.[57] Bryozoans exhibited significant degrees of endemism to a particular shelf.[59] They also developed symbiotic relationships with cnidarians[60] and stromatolites.[61] Many bivalve fossils have also been found in Silurian deposits,[62] and the first deep-boring bivalves are known from this period.[63] Chitons saw a peak in diversity during the middle of the Silurian.[64] Hederelloids enjoyed significant success in the Silurian, with some developing symbioses with the colonial rugose coral Entelophyllum.[65] The Silurian was a heyday for tentaculitoids,[66] which experienced an evolutionary radiation focused mainly in Baltoscandia,[67] along with an expansion of their geographic range in the Llandovery and Wenlock.[68] Trilobites started to recover in the Rhuddanian,[69] and they continued to enjoy success in the Silurian as they had in the Ordovician despite their reduction in clade diversity as a result of LOME.[70] The Early Silurian was a chaotic time of turnover for crinoids as they rediversified after LOME.[71] Members of Flexibilia, which were minimally impacted by LOME, took on an increasing ecological prominence in Silurian seas.[72] Monobathrid camerates, like flexibles, diversified in the Llandovery, whereas cyathocrinids and dendrocrinids diversified later in the Silurian.[73] Scyphocrinoid loboliths suddenly appeared in the terminal Silurian, shortly before the Silurian-Devonian boundary, and disappeared as abruptly as they appeared very shortly after their first appearance.[74] Endobiotic symbionts were common in the corals and stromatoporoids.[75][76] Rugose corals especially were colonised and encrusted by a diverse range of epibionts,[77] including certain hederelloids as aforementioned.[65] Photosymbiotic scleractinians made their first appearance during the Middle Silurian.[78] Reef abundance was patchy; sometimes, fossils are frequent, but at other points, are virtually absent from the rock record.[26]

See also

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Notes

[edit]
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  2. ^ Munnecke, A.; Samtleben, C.; Bickert, T. (2003). "The Ireviken Event in the lower Silurian of Gotland, Sweden-relation to similar Palaeozoic and Proterozoic events". Palaeogeography, Palaeoclimatology, Palaeoecology. 195 (1): 99–124. doi:10.1016/S0031-0182(03)00304-3.
  3. ^ "Chart/Time Scale". www.stratigraphy.org. International Commission on Stratigraphy.
  4. ^ Lucas, Sepncer (6 November 2018). "The GSSP Method of Chronostratigraphy: A Critical Review". Frontiers in Earth Science. 6: 191. Bibcode:2018FrEaS...6..191L. doi:10.3389/feart.2018.00191.
  5. ^ Holland, C. (June 1985). "Series and Stages of the Silurian System" (PDF). Episodes. 8 (2): 101–103. doi:10.18814/epiiugs/1985/v8i2/005. Retrieved 11 December 2020.
  6. ^ Chlupáč, Ivo; Hladil, Jindrich (January 2000). "The global stratotype section and point of the Silurian-Devonian boundary". CFS Courier Forschungsinstitut Senckenberg. Retrieved 7 December 2020.
  7. ^ Haq, B. U.; Schutter, SR (2008). "A Chronology of Paleozoic Sea-Level Changes". Science. 322 (5898): 64–68. Bibcode:2008Sci...322...64H. doi:10.1126/science.1161648. PMID 18832639. S2CID 206514545.
  8. ^ Wells, John (3 April 2008). Longman Pronunciation Dictionary (3rd ed.). Pearson Longman. ISBN 978-1-4058-8118-0.
  9. ^ "Silurian". Dictionary.com Unabridged (Online). n.d.
  10. ^ "Silurian". CollinsDictionary.com. HarperCollins.
  11. ^ "International Chronostratigraphic Chart v.2015/01" (PDF). International Commission on Stratigraphy. January 2015.
  12. ^ a b Capel, Elliot; Cleal, Christopher J.; Xue, Jinzhuang; Monnet, Claude; Servais, Thomas; Cascales-Miñana, Borja (August 2022). "The Silurian–Devonian terrestrial revolution: Diversity patterns and sampling bias of the vascular plant macrofossil record". Earth-Science Reviews. 231: 104085. Bibcode:2022ESRv..23104085C. doi:10.1016/j.earscirev.2022.104085. hdl:20.500.12210/76731. S2CID 249616013.
  13. ^ a b Xue, Jinzhuang; Huang, Pu; Wang, Deming; Xiong, Conghui; Liu, Le; Basinger, James F. (May 2018). "Silurian-Devonian terrestrial revolution in South China: Taxonomy, diversity, and character evolution of vascular plants in a paleogeographically isolated, low-latitude region". Earth-Science Reviews. 180: 92–125. Bibcode:2018ESRv..180...92X. doi:10.1016/j.earscirev.2018.03.004. Retrieved 8 December 2022.
  14. ^ Lutzoni, François; Nowak, Michael D.; Alfaro, Michael E.; Reeb, Valérie; Miadlikowska, Jolanta; Krug, Michael; Arnold, A. Elizabeth; Lewis, Louise A.; Swofford, David L.; Hibbett, David; Hilu, Khidir; James, Timothy Y.; Quandt, Dietmar; Magallón, Susana (21 December 2018). "Contemporaneous radiations of fungi and plants linked to symbiosis". Nature Communications. 9 (1): 5451. Bibcode:2018NatCo...9.5451L. doi:10.1038/s41467-018-07849-9. PMC 6303338. PMID 30575731. S2CID 56645104.
  15. ^ Garwood, Russell J.; Edgecombe, Gregory D. (September 2011). "Early Terrestrial Animals, Evolution, and Uncertainty". Evolution: Education and Outreach. 4 (3): 489–501. doi:10.1007/s12052-011-0357-y.
  16. ^ Brazeau, M. D.; Friedman, M. (2015). "The origin and early phylogenetic history of jawed vertebrates". Nature. 520 (7548): 490–497. Bibcode:2015Natur.520..490B. doi:10.1038/nature14438. PMC 4648279. PMID 25903631.
  17. ^ See:
  18. ^ Murchison, Roderick (1835). "On the Silurian System of Rocks". The London and Edinburgh Philosophical Magazine and Journal of Science. Third Series, Vol. 7: 46–52 – via Biodiversity Heritage Library.
  19. ^ Sedgwick; Murchison, R.I. (1835). "On the Silurian and Cambrian systems, exhibiting the order in which the older sedimentary strata succeed each other in England and Wales". Report of the Fifth Meeting of the British Association for the Advancement of Science. § Notices and Abstracts of Miscellaneous Communications to the Sections. 5: 59–61.
  20. ^ Lapworth, Charles (1879). "On the tripartite classification of the Lower Palaeozoic rocks". Geological Magazine. 2nd series. 6 (1): 1–15. Bibcode:1879GeoM....6....1L. doi:10.1017/s0016756800156560. S2CID 129165105. From pp. 13–14: "North Wales itself – at all events the whole of the great Bala district where Sedgwick first worked out the physical succession among the rocks of the intermediate or so-called Upper Cambrian or Lower Silurian system; and in all probability much of the Shelve and the Caradoc area, whence Murchison first published its distinctive fossils – lay within the territory of the Ordovices; … Here, then, have we the hint for the appropriate title for the central system of the Lower Palaeozoics. It should be called the Ordovician System, after this old British tribe."
  21. ^ The Gotlandian system was proposed in 1893 by the French geologist Albert Auguste Cochon de Lapparent (1839–1908): Lapparent, A. de (1893). Traité de Géologie (in French). Vol. 2 (3rd ed.). Paris, France: F. Savy. p. 748. From p. 748: "D'accord avec ces divisions, on distingue communément dans le silurien trois étages: l'étage inférieur ou cambrien (1); l'étage moyen ou ordovicien (2); l'étage supérieur ou gothlandien (3)." (In agreement with these divisions, one generally distinguishes, within the Silurian, three stages: the lower stage or Cambrian [1]; the middle stage or Ordovician [2]; the upper stage or Gotlandian [3].)
  22. ^ Barrande, Joachim (1852). Systême silurien du centre de la Bohême (in French). Paris, France and Prague, (Czech Republic): (Self-published). pp. ix–x.
  23. ^ Forbes, Edward (1854). "Anniversary Address of the President". Quarterly Journal of the Geological Society of London. 10: xxii–lxxxi. See p. xxxiv.
  24. ^ a b Paul Selden & Helen Read (2008). "The oldest land animals: Silurian millipedes from Scotland" (PDF). Bulletin of the British Myriapod & Isopod Group. 23: 36–37.
  25. ^ Manda, Štěpán; Frýda, Jiří (2010). "Silurian-Devonian boundary events and their influence on cephalopod evolution: evolutionary significance of cephalopod egg size during mass extinctions". Bulletin of Geosciences. 85 (3): 513–40. doi:10.3140/bull.geosci.1174.
  26. ^ a b c d e f g Munnecke, Axel; Calner, Mikael; Harper, David A.T.; Servais, Thomas (2010). "Ordovician and Silurian sea–water chemistry, sea level, and climate: A synopsis". Palaeogeography, Palaeoclimatology, Palaeoecology. 296 (3–4): 389–413. Bibcode:2010PPP...296..389M. doi:10.1016/j.palaeo.2010.08.001.
  27. ^ Yan, Guanzhou; Lehnert, Oliver; Männik, Peep; Calner, Mikael; Luan, Xiaocong; Gong, Fanyi; Li, Lixia; Wei, Xin; Wang, Guangxu; Zhan, Renbin; Wu, Rongchang (15 November 2022). "The record of early Silurian climate changes from South China and Baltica based on integrated conodont biostratigraphy and isotope chemostratigraphy". Palaeogeography, Palaeoclimatology, Palaeoecology. 606: 111245. Bibcode:2022PPP...60611245Y. doi:10.1016/j.palaeo.2022.111245. S2CID 252504361. Retrieved 8 December 2022.
  28. ^ a b Gambacorta, G.; Menichetti, E.; Trincianti, E.; Torricelli, S. (March 2019). "The Silurian climatic transition recorded in the epicontinental Baltica Sea". Palaeogeography, Palaeoclimatology, Palaeoecology. 517: 16–29. Bibcode:2019PPP...517...16G. doi:10.1016/j.palaeo.2018.12.016. S2CID 135118794.
  29. ^ Young, Set A.; Benayoun, Emily; Kozik, Nevin P.; Hints, Olle; Martma, Tõnu; Bergström, Stig M.; Owens, Jeremy D. (15 September 2020). "Marine redox variability from Baltica during extinction events in the latest Ordovician–early Silurian". Palaeogeography, Palaeoclimatology, Palaeoecology. 554: 109792. Bibcode:2020PPP...55409792Y. doi:10.1016/j.palaeo.2020.109792. S2CID 218930512.
  30. ^ Sproson, Adam D. (15 February 2020). "Pacing of the latest Ordovician and Silurian carbon cycle by a ~4.5 Myr orbital cycle". Palaeogeography, Palaeoclimatology, Palaeoecology. 540: 109543. Bibcode:2020PPP...54009543S. doi:10.1016/j.palaeo.2019.109543. S2CID 213445668. Retrieved 8 December 2022.
  31. ^ a b Trotter, Julie A.; Williams, Ian S.; Barnes, Christopher R.; Männik, Peep; Simpson, Andrew (February 2016). "New conodont δ18O records of Silurian climate change: Implications for environmental and biological events". Palaeogeography, Palaeoclimatology, Palaeoecology. 443: 34–48. Bibcode:2016PPP...443...34T. doi:10.1016/j.palaeo.2015.11.011.
  32. ^ Chaloner, William G. (2003). "The role of carbon dioxide in plant evolution". Evolution on Planet Earth: 65–83. doi:10.1016/B978-012598655-7/50032-X. ISBN 9780125986557.
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  34. ^ Samtleben, C.; Munnecke, A.; Bickert, T. (2000). "Development of facies and C/O-isotopes in transects through the Ludlow of Gotland: Evidence for global and local influences on a shallow-marine environment". Facies. 43 (1): 1–38. Bibcode:2000Faci...43....1S. doi:10.1007/BF02536983. S2CID 130640332.
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References

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  • Emiliani, Cesare. (1992). Planet Earth : Cosmology, Geology, & the Evolution of Life & the Environment. Cambridge University Press. (Paperback Edition ISBN 0-521-40949-7)
  • Mikulic, DG, DEG Briggs, and J Kluessendorf. 1985. A new exceptionally preserved biota from the Lower Silurian of Wisconsin, USA. Philosophical Transactions of the Royal Society of London, 311B:75-86.
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