Earth:Great Ordovician Biodiversification Event

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Short description: Evolutionary radiation of animal life throughout the Ordovician period

The Great Ordovician Biodiversification Event (GOBE), was an evolutionary radiation of animal life throughout[1] the Ordovician period, 40 million years after the Cambrian explosion,[2] whereby the distinctive Cambrian fauna fizzled out to be replaced with a Paleozoic fauna rich in suspension feeder and pelagic animals.[3]

It followed a series of Cambrian–Ordovician extinction events, and the resulting fauna went on to dominate the Palaeozoic relatively unchanged.[4] Marine diversity increased to levels typical of the Palaeozoic,[5] and morphological disparity was similar to today's.[6] The diversity increase was neither global nor instantaneous; it happened at different times in different places.[4] Consequently, there is unlikely to be a simple or straightforward explanation for the event; the interplay of many geological and ecological factors likely produced the diversification.[1]

Duration

According to a comprehensive study of biodiversity throughout the Palaeozoic, GOBE began 497.05 Ma and ended 467.33 Ma, lasting for 29.72 Myr.[7] GOBE did not constitute one single event, as different clades diversified during different time intervals of the Early and Middle Ordovician.[8] During the Late Ordovician, diversification slowed down thanks to increased endemism and interbasinal dispersal, bringing an end to GOBE.[9]

Causes

Possible line of meteors (on the modern globe) associated with the Middle Ordovician meteor event 467.5±0.28 million years ago. Although this is suggestive of a single large meteorite shower, the exact alignment of continental plates 470 million years ago is unknown and the exact timing of meteors is also unknown.

Possible causes include an increase in marine oxygen content,[10] changes in palaeogeography or tectonic activity,[11] a modified nutrient supply,[12] or global cooling.[11]

Tectonic activity

The dispersed positions of the continents, high level of tectonic/volcanic activity, warm climate, and high CO2 levels would have created a large, nutrient-rich ecospace, favoring diversification.[2] There seems to be an association between orogeny and the evolutionary radiation,[13] with the Taconic orogeny in particular being singled out as a driver of the GOBE by enabling greater erosion of nutrients such as iron and phosphorus and their delivery to the oceans around Laurentia.[11] In addition, the changing geography led to a more diverse landscape, with more different and isolated environments; this no doubt facilitated the emergence of bioprovinciality, and speciation by isolation of populations.[1] The widespread reef development on the Baltican shelf in particular is attributable to the landmass's northward drift into more oligotrophic waters, enabling diversification of its reef biota.[14] Widespread volcanism and its delivery of biologically important trace metals has similarly been proposed as a GOBE trigger, albeit controversially.[15]

Global cooling

On the other hand, global cooling has also been offered as a cause of the radiation,[11][16][17] with long-term biodiversity trends showing a positive correlation between cooling and biodiversity during GOBE.[18][7] An uptick in fossil diversity correlates with the increasing abundance of cool-water carbonates over the course of this time interval.[19] A transient high magnitude shift towards more positive carbon isotope ratios during the Floian may reflect the initiation of a cooling through organic carbon burial that has been proposed to have kickstarted GOBE.[20] In the longer term as well, increasing carbon isotope ratios track biodiversity increase, further pointing to a link between cooling and GOBE.[21][22] The cooling during the Middle and early Late Ordovician in particular is known for its associated burst of biodiversification.[23] The volcanic activity that created the Flat Landing Brook Formation in New Brunswick, Canada may have caused rapid climatic cooling and biodiversification.[24]

Oxygenation

Thallium isotope shifts show an expansion of oxic waters throughout deep water and shallow shelf environments during the latest Cambrian and earliest Ordovician coeval with increasing burrowing depth and complexity observed among ichnofossils and increasing morphological complexity among body fossils. Thus, heightened oxygen availability may have been a key trigger for GOBE.[10] Furthermore, Ordovician biodiversification pulses were closely linked to terminations of positive carbon isotope excursions, which are characteristic of anoxia, suggesting that diversification occurred in concert with increasing oxygen content.[25] After the SPICE event about 500 million years ago, the extinction in the ocean would have opened up new niches for photosynthetic plankton, who would absorb CO2 from the atmosphere and release large amount of oxygen. More oxygen and a more diversified photosynthetic plankton as the bottom of the food chain, would have affected the diversity of higher marine organisms and their ecosystems.[26]

Following GOBE, in the Middle to Late Ordovician, an expansion of anoxic waters occurred in sync with a ~50% decline in benthic invertebrates in various epicontinental seas, providing further indirect support for a coupling of seawater oxygenation with Ordovician biodiversity.[27]

Extraterrestrial impacts

Another alternative is that the breakup of an asteroid led to the Earth being consistently pummelled by meteorites,[3] although the proposed Ordovician meteor event happened at 467.5±0.28 million years ago.[28][29] Another effect of a collision between two asteroids, possibly beyond the orbit of Mars, is a reduction in sunlight reaching the Earth's surface due to the vast dust clouds created. Evidence for this geological event comes from the relative abundance of the isotope helium-3, found in ocean sediments laid down at the time of the biodiversification event. The most likely cause of the production of high levels of helium-3 is the bombardment of lithium by cosmic rays, something which could only have happened to material which travelled through space.[30]

However, rather than sparking evolutionary diversification, other lines of evidence point to the Ordovician meteor event instead postdating the Darriwilian biodiversity burst by about 600 kyr and the start of glaciation by 800 kyr. Instead of facilitating the radiation, the meteor event may have antagonistically acted to temporarily retard and halt biological diversification according to this thesis.[31]

Positive feedbacks

The above triggers would have been amplified by ecological escalation, whereby any new species would co-evolve with others, creating new niches through niche partitioning, trophic layering, or by providing a new habitat.[clarification needed][12] As with the Cambrian Explosion, it is likely that environmental changes drove the diversification of plankton, which permitted an increase in diversity and abundance of plankton-feeding lifeforms, including suspension feeders on the sea floor, and nektonic organisms in the water column.[3]

Effects

Atrypid brachiopods (Zygospira modesta) preserved in their original positions on a trepostome bryozoan; Cincinnatian (Upper Ordovician) of southeastern Indiana.

If the Cambrian Explosion is thought of as producing the modern phyla,[32] the GOBE can be considered as the "filling out" of these phyla with the modern (and many extinct) classes and lower-level taxa.[3] The GOBE is considered to be one of the most potent speciation events of the Phanerozoic era, increasing global diversity severalfold and leading to the establishment of the Palaeozoic evolutionary fauna.[33] Notable taxonomic diversity explosions during this period include that of articulated brachiopods, gastropods, and bivalves.[34] The acritarch record (the majority of acritarchs were probably marine algae)[3] displays the Ordovician radiation beautifully; both diversity and disparity peaked in the middle Ordovician. The warm waters and high sea level (which had been rising steadily since the early Cambrian) permitted large numbers of phytoplankton to prosper; the accompanying diversification of the phytoplankton may have caused an accompanying radiation of zooplankton and suspension feeders.[2]

Taxonomic diversity increased manifold; the total number of marine orders doubled, and families tripled.[4] Marine biodiversity reached levels comparable to those of the present day.[5] Beta diversity was the most important component of biodiversity increase from the Furongian to the Tremadocian. From the Floian onward, alpha diversity dethroned beta diversity as the greater contributor to regional diversity patterns.[35] In addition to a diversification, the event also marked an increase in the complexity of both organisms and food webs.[3] The number of different life modes among hard-bodied organisms doubled.[6] Taxa began to exhibit greater provincialism and have more localized ranges, with different faunas at different parts of the globe.[36][37][38] Communities in reefs and deeper water began to take on a character of their own, becoming more clearly distinct from other marine ecosystems.[1] Benthic environments drastically increase in the amount and variety of bioturbation.[39] The planktonic realm was invaded as never before, with several invertebrate lineages colonising the open waters and initiating new food chains at the end of the Cambrian into the early Ordovician.[40] Among the newcomers colonising the planktonic realm were trilobites[41] and cephalopods.[40] Estuarine environments too experienced increased colonisation by living organisms.[42] And as ecosystems became more diverse, with more species being squeezed into the food web, a more complex tangle of ecological interactions resulted, promoting strategies such as ecological tiering. The global fauna that emerged during the GOBE went on to be remarkably stable until the catastrophic end-Permian extinction and the ensuing Mesozoic Marine Revolution.[1]

Relationship to the Cambrian Explosion

Recent work has suggested that the Cambrian Explosion and GOBE, rather than being two distinct events, represented one continual evolutionary radiation of marine life occurring over the entire Early Palaeozoic.[43] An analysis of the Paleobiology Database (PBDB) and Geobiodiversity Database (GBDB) found no statistical basis for separating the two radiations into discrete events.[44]

A proposed biodiversity gap known as the Furongian Gap is thought by some researchers to have existed between the Cambrian Explosion and GOBE existed during the Furongian epoch, the final epoch of the Cambrian. However, whether this gap is real or an artefact of an incomplete fossil record is controversial.[45] Analysis of the Guole Konservat-Lagerstätte and other sites in South China suggests the Furongian Gap did not exist, instead portraying this interval as one of rapid biotic turnovers.[46]

See also

References

  1. ↑ 1.0 1.1 1.2 1.3 1.4 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. Bibcode: 2010PPP...296..389M. 
  2. ↑ 2.0 2.1 2.2 Servais, T.; Lehnert, O.; Li, J.; Mullins, G. L.; Munnecke, A.; Nützel, A.; Vecoli, M. (2008). "The Ordovician Biodiversification: revolution in the oceanic trophic chain". Lethaia 41 (2): 99–109. doi:10.1111/j.1502-3931.2008.00115.x. 
  3. ↑ 3.0 3.1 3.2 3.3 3.4 3.5 Servais, T.; Owen, A. W.; Harper, D. A. T.; Kröger, B. R.; Munnecke, A. (2010). "The Great Ordovician Biodiversification Event (GOBE): the palaeoecological dimension". Palaeogeography, Palaeoclimatology, Palaeoecology 294 (3–4): 99–119. doi:10.1016/j.palaeo.2010.05.031. 
  4. ↑ 4.0 4.1 4.2 Droser, M. L.; Finnegan, S. (2003). "The Ordovician Radiation: A Follow-up to the Cambrian Explosion?". Integrative and Comparative Biology 43 (1): 178–184. doi:10.1093/icb/43.1.178. PMID 21680422. 
  5. ↑ 5.0 5.1 Marshall, C. R. (2006). "Explaining the Cambrian "explosion" of Animals". Annual Review of Earth and Planetary Sciences 34: 355–384. doi:10.1146/annurev.earth.33.031504.103001. Bibcode: 2006AREPS..34..355M. 
  6. ↑ 6.0 6.1 Bambach, R. K.; Bush, A. M.; Erwin, D. H. (2007). "Autecology and the Filling of Ecospace: Key Metazoan Radiations". Palaeontology 50: 1–22. doi:10.1111/j.1475-4983.2006.00611.x. 
  7. ↑ 7.0 7.1 Fan, Jun-xuan; Shen, Shu-zhong; Erwin, Douglas H.; Sadler, Peter M.; MacLeod, Norman; Cheng, Qiu-ming; Ho, Xu-dong; Yang, Jiao et al. (17 January 2020). "A high-resolution summary of Cambrian to Early Triassic marine invertebrate biodiversity". Science 367 (6475): 272–277. doi:10.1126/science.aax4953. PMID 31949075. 
  8. ↑ Servais, Thomas; Cascales-Miñana, Borja; Harper, David A. T. (1 October 2021). "The Great Ordovician Biodiversification Event (GOBE) is Not a Single Event". Paleontological Research 25 (4): 315–328. doi:10.2517/2021PR001. https://bioone.org/journals/paleontological-research/volume-25/issue-4/2021PR001/The-Great-Ordovician-Biodiversification-Event-GOBE-is-Not-a-Single/10.2517/2021PR001.short. Retrieved 22 July 2023. 
  9. ↑ Stigall, Alycia L.; Freeman, Rebecca L.; Edwards, Cole T.; Rasmussen, Christian M. Ø. (1 April 2020). "A multidisciplinary perspective on the Great Ordovician Biodiversification Event and the development of the early Paleozoic world". Palaeogeography, Palaeoclimatology, Palaeoecology 543. doi:10.1016/j.palaeo.2019.109521. https://www.sciencedirect.com/science/article/abs/pii/S0031018219310867. Retrieved 14 August 2023. 
  10. ↑ 10.0 10.1 Kozik, Nevin P.; Young, Seth A.; Lindskog, Anders; Ahlberg, Per; Owens, Jeremy D. (26 January 2023). "Protracted oxygenation across the Cambrian–Ordovician transition: A key initiator of the Great Ordovician Biodiversification Event?". Geobiology 21 (3): 323–340. doi:10.1111/gbi.12545. PMID 36703593. https://onlinelibrary.wiley.com/doi/abs/10.1111/gbi.12545. Retrieved 21 April 2023. 
  11. ↑ 11.0 11.1 11.2 11.3 Kozik, Nevin P.; Young, Seth A.; Bowman, Chelsea N.; Saltzmann, Matthew R.; Them II, Theodore R. (15 April 2019). "Middle–Upper Ordovician (Darriwilian–Sandbian) paired carbon and sulfur isotope stratigraphy from the Appalachian Basin, USA: Implications for dynamic redox conditions spanning the peak of the Great Ordovician Biodiversification Event". Palaeogeography, Palaeoclimatology, Palaeoecology 520: 188–202. doi:10.1016/j.palaeo.2019.01.032. 
  12. ↑ 12.0 12.1 Botting, Joseph P.; Muir, Lucy A. (28 June 2008). "Unravelling Causal Components of the Ordovician Radiation: the Builth Inlier (Central Wales) As a Case Study". Lethaia 41 (2): 111–125. doi:10.1111/j.1502-3931.2008.00118.x. https://onlinelibrary.wiley.com/doi/abs/10.1111/j.1502-3931.2008.00118.x. Retrieved 4 July 2023. 
  13. ↑ Miller, Arnold I.; Mao, Shuguang (1995-04-01). "Association of orogenic activity with the Ordovician radiation of marine life". Geology 23 (4): 305–308. doi:10.1130/0091-7613(1995)023<0305:AOOAWT>2.3.CO;2. ISSN 0091-7613. PMID 11539503. https://doi.org/10.1130/0091-7613(1995)0232.3.CO;2. 
  14. ↑ Pohl, Alexandre; Harper, David A. T.; Donnadieu, Yannick; Le Hir, Guillaume; Nardin, Elise; Servais, Thomas (12 October 2017). "Possible patterns of marine primary productivity during the Great Ordovician Biodiversification Event". Lethaia 51 (2): 187–197. doi:10.1111/let.12247. https://www.idunn.no/doi/full/10.1111/let.12247. Retrieved 15 July 2023. 
  15. ↑ Thompson, Cara K.; Kah, Linda C.; Astini, Ricardo; Bowring, Samuel A.; Buchwaldt, Robert (1 March 2012). "Bentonite geochronology, marine geochemistry, and the Great Ordovician Biodiversification Event (GOBE)". Palaeogeography, Palaeoclimatology, Palaeoecology 321-322: 88–101. doi:10.1016/j.palaeo.2012.01.022. https://www.sciencedirect.com/science/article/abs/pii/S0031018212000363. Retrieved 22 July 2023. 
  16. ↑ Cocks, L. Robin M.; Torsvik, Trond H. (December 2021). "Ordovician palaeogeography and climate change". Gondwana Research 100: 53–72. doi:10.1016/j.gr.2020.09.008. https://www.sciencedirect.com/science/article/pii/S1342937X20302756. Retrieved 26 May 2023. 
  17. ↑ Trotter, JA; Williams, IS; Barnes, CR; Lécuyer, C; Nicoll, RS (2008). "Did cooling oceans trigger Ordovician biodiversification? Evidence from conodont thermometry". Science 321 (5888): 550–4. doi:10.1126/science.1155814. PMID 18653889. Bibcode: 2008Sci...321..550T. 
  18. ↑ Goldberg, Samuel L.; Present, Theodore M.; Finnegan, Seth; Bergmann, Kristin D. (2021-02-09). "A high-resolution record of early Paleozoic climate" (in en). Proceedings of the National Academy of Sciences of the United States of America 118 (6): e2013083118. doi:10.1073/pnas.2013083118. ISSN 0027-8424. PMID 33526667. 
  19. ↑ Dronov, Andrei (1 November 2013). "Late Ordovician cooling event: Evidence from the Siberian Craton". Palaeogeography, Palaeoclimatology, Palaeoecology 389: 87–95. doi:10.1016/j.palaeo.2013.05.032. https://www.sciencedirect.com/science/article/abs/pii/S0031018213002745. Retrieved 20 October 2022. 
  20. ↑ Zhang, Tonggang; Shen, Yanan; Algeo, Thomas J. (1 April 2010). "High-resolution carbon isotopic records from the Ordovician of South China: Links to climatic cooling and the Great Ordovician Biodiversification Event (GOBE)". Palaeogeography, Palaeoclimatology, Palaeoecology 289 (1–4): 102–112. doi:10.1016/j.palaeo.2010.02.020. https://www.sciencedirect.com/science/article/abs/pii/S0031018210000921. Retrieved 15 July 2023. 
  21. ↑ Hu, Dongping; Zhang, Xiaolin; Li, Menghan; Xu, Yilun; Shen, Yanan (August 2021). "Carbon isotope (δ13Ccarb) stratigraphy of the Lower-Upper Ordovician of the Yangtze Platform, South China: Implications for global correlation and the Great Ordovician Biodiversification Event (GOBE)". Global and Planetary Change 203. doi:10.1016/j.gloplacha.2021.103546. https://www.sciencedirect.com/science/article/abs/pii/S0921818121001314. Retrieved 15 July 2023. 
  22. ↑ Edwards, Cole T.; Saltzman, Matthew R. (15 September 2016). "Paired carbon isotopic analysis of Ordovician bulk carbonate (δ13Ccarb) and organic matter (δ13Corg) spanning the Great Ordovician Biodiversification Event". Palaeogeography, Palaeoclimatology, Palaeoecology 458: 102–117. doi:10.1016/j.palaeo.2015.08.005. https://www.sciencedirect.com/science/article/abs/pii/S0031018215004307. Retrieved 15 July 2023. 
  23. ↑ Algeo, Thomas J.; Marenco, Pedro J.; Saltzman, Matthew R. (15 September 2016). "Co-evolution of oceans, climate, and the biosphere during the 'Ordovician Revolution': A review". Palaeogeography, Palaeoclimatology, Palaeoecology 458: 1–11. doi:10.1016/j.palaeo.2016.05.015. https://www.sciencedirect.com/science/article/abs/pii/S0031018216301559?fr=RR-2&ref=pdf_download&rr=7f6e23542b527bf7. Retrieved 14 August 2023. 
  24. ↑ "A mid-Darriwilian super volcano in northern New Brunswick, rapid climate change and the start of the great Ordovician biodiversification event". Mineralogical Association of Canada. 2012. p. 119. http://www.mineralogicalassociation.ca/doc/StJohns2012_GAC-MAC_Abstracts.pdf. 
  25. ↑ Edwards, Cole T. (March–June 2019). "Links between early Paleozoic oxygenation and the Great Ordovician Biodiversification Event (GOBE): A review". Palaeoworld 28 (1–2): 37–50. doi:10.1016/j.palwor.2018.08.006. https://www.sciencedirect.com/science/article/abs/pii/S1871174X1830043X. Retrieved 15 July 2023. 
  26. ↑ Solved: Mystery of Earth's First Breathable Atmosphere
  27. ↑ Zhang, Junpeng; Li, Chao; Fang, Xiang; Li, Wenjie; Deng, Yiying; Tu, Chenyi; Algeo, Thomas J.; Lyons, Timothy W. et al. (15 November 2022). "Progressive expansion of seafloor anoxia in the Middle to Late Ordovician Yangtze Sea: Implications for concurrent decline of invertebrate diversity". Earth and Planetary Science Letters 598: 117858. doi:10.1016/j.epsl.2022.117858. ISSN 0012-821X. https://www.sciencedirect.com/science/article/pii/S0012821X22004940. Retrieved 15 December 2023. 
  28. ↑ An extraterrestrial trigger for the mid-Ordovician ice age: Dust from the breakup of the L-chondrite parent body, Birger Schmitz et al, AAAS Science Advances, 18 Sep 2019: Vol. 5, no. 9, eaax4184; DOI: 10.1126/sciadv.aax4184, accessed 2019-10-09
  29. ↑ Lindskog, A.; Costa, M. M.; Rasmussen, C.M.Ø.; Connelly, J. N.; Eriksson, M. E. (24 January 2017). "Refined Ordovician timescale reveals no link between asteroid breakup and biodiversification" (in En). Nature Communications 8: 14066. doi:10.1038/ncomms14066. ISSN 2041-1723. PMID 28117834. "It has been suggested that the Middle Ordovician meteorite bombardment played a crucial role in the Great Ordovician Biodiversification Event, but this study shows that the two phenomena were unrelated". 
  30. ↑ McKie, Robin (12 October 2019). "New evidence shows how asteroid dust cloud may have sparked new life on Earth 470m years ago" (in en-GB). The Observer. ISSN 0029-7712. https://www.theguardian.com/science/2019/oct/12/asteroid-kicked-off-new-life-on-earth. 
  31. ↑ Rasmussen, Jan Audun; Thibault, Nicolas; Rasmussen, Christian Mac Ørum (5 November 2021). "Middle Ordovician astrochronology decouples asteroid breakup from glacially-induced biotic radiations". Nature Communications 12 (1): 6430. doi:10.1038/s41467-021-26396-4. PMID 34741034. 
  32. ↑ All mineralized phyla were present by the end of the Cambrian; see Landing, E.; English, A.; Keppie, J. D. (2010). "Cambrian origin of all skeletalized metazoan phyla--Discovery of Earth's oldest bryozoans (Upper Cambrian, southern Mexico)". Geology 38 (6): 547–550. doi:10.1130/G30870.1. Bibcode: 2010Geo....38..547L. 
  33. ↑ Bush, A. M.; Bambach, R. K.; Daley, G. M. (2007). "Changes in theoretical ecospace utilization in marine fossil assemblages between the mid-Paleozoic and late Cenozoic". Paleobiology 33: 76–97. doi:10.1666/06013.1. 
  34. ↑ Stigall, A.L (December 2016). "Biotic immigration events, speciation, and the accumulation of biodiversity in the fossil record". Global and Planetary Change 148: 242–257. doi:10.1016/j.gloplacha.2016.12.008. Bibcode: 2017GPC...148..242S. 
  35. ↑ Serra, Fernanda; Balseiro, Diego; Waisfeld, Beatriz G. (1 April 2023). "Morphospace trends underlying a global turnover: Ecological dynamics of trilobite assemblages at the onset of the Ordovician Radiation". Palaeogeography, Palaeoclimatology, Palaeoecology 615. doi:10.1016/j.palaeo.2023.111448. 
  36. ↑ Harper, David A. T.; Zhan, Ren-Bin; Jin, Jisuo (March–June 2015). "The Great Ordovician Biodiversification Event: Reviewing two decades of research on diversity's big bang illustrated by mainly brachiopod data". Palaeoworld 24 (1–2): 75–85. doi:10.1016/j.palwor.2015.03.003. https://www.sciencedirect.com/science/article/abs/pii/S1871174X15000153. Retrieved 7 July 2023. 
  37. ↑ Zhan, Renbin; Luan, Xiaocong; Huang, Bing; Liang, Yan; Wang, Guangxu; Wang, Yi (December 2014). "Darriwilian Saucrorthis Fauna: implications for the Great Ordovician Biodiversification Event (GOBE)". Estonian Journal of Earth Sciences 63 (4): 323–328. doi:10.3176/earth.2014.38. https://www.researchgate.net/publication/275956279. Retrieved 7 July 2023. 
  38. ↑ Servais, Thomas; Harper, David A. T.; Kröger, Björn; Scotese, Christopher Robert; Stigall, Alycia L.; Zhen, Yong-Yi (10 March 2023). "Changing palaeobiogeography during the Ordovician Period". Geological Society, London, Special Publications 532: 111–136. doi:10.1144/SP532-2022-168. https://www.lyellcollection.org/doi/abs/10.1144/SP532-2022-168. Retrieved 7 July 2023. 
  39. ↑ Harper, David A. T. (22 March 2006). "The Ordovician biodiversification: Setting an agenda for marine life". Palaeogeography, Palaeoclimatology, Palaeoecology 232 (2–4): 148–166. doi:10.1016/j.palaeo.2005.07.010. https://www.sciencedirect.com/science/article/abs/pii/S0031018205004141. Retrieved 15 July 2023. 
  40. ↑ 40.0 40.1 Kröger, B. R.; Servais, T.; Zhang, Y.; Kosnik, M. (2009). Kosnik, Matthew. ed. "The Origin and Initial Rise of Pelagic Cephalopods in the Ordovician". PLOS ONE 4 (9): e7262. doi:10.1371/journal.pone.0007262. PMID 19789709. Bibcode: 2009PLoSO...4.7262K. 
  41. ↑ Esteve, Jorge; López-Pachón, Matheo (1 September 2023). "Swimming and feeding in the Ordovician trilobite Microparia speciosa shed light on the early history of nektonic life habits". Palaeogeography, Palaeoclimatology, Palaeoecology 625. doi:10.1016/j.palaeo.2023.111691. 
  42. ↑ Mángano, M. Gabriela; Waisfeld, Beatriz G.; Buatois, Luis A.; Vaccari, N. Emilio; Muñoz, Diego F. (15 September 2023). "Evolutionary and ecologic controls on benthos distribution from an upper Cambrian incised estuarine valley: Implications for the early colonization of marginal-marine settings". Palaeogeography, Palaeoclimatology, Palaeoecology 626. doi:10.1016/j.palaeo.2023.111692. https://www.sciencedirect.com/science/article/abs/pii/S0031018223003103. Retrieved 8 July 2023. 
  43. ↑ Harper, David A. T.; Cascales-Miñana, Borja; Servais, Thomas (3 December 2019). "Early Palaeozoic diversifications and extinctions in the marine biosphere: A continuum of change". Geological Magazine 157 (1): 5–21. doi:10.1017/S0016756819001298. 
  44. ↑ Servais, Thomas; Cascales-Miñana, Borja; Harper, David A. T.; Lefebvre, Bertrand; Munnecke, Axel; Wang, Wenhui; Zhang, Yuandong (1 August 2023). "No (Cambrian) explosion and no (Ordovician) event: A single long-term radiation in the early Palaeozoic". Palaeogeography, Palaeoclimatology, Palaeoecology 623. doi:10.1016/j.palaeo.2023.111592. 
  45. ↑ Harper, David A. T.; Topper, Timothy P.; Cascales-Miñana, Borja; Servais, Thomas; Zhang, Yuan-Dong; Ahlberg, Per (March–June 2019). "The Furongian (late Cambrian) Biodiversity Gap: Real or apparent?". Palaeoworld 28 (1–2): 4–12. doi:10.1016/j.palwor.2019.01.007. https://www.sciencedirect.com/science/article/abs/pii/S1871174X18301628. Retrieved 4 July 2023. 
  46. ↑ Deng, Yiying; Fan, Junxuan; Yang, Shengchao; Shi, Yukun; Lu, Zhengbo; Xu, Huiqing; Sun, Zongyuan; Zhao, Fangqi et al. (15 May 2023). "No Furongian Biodiversity Gap: Evidence from South China". Palaeogeography, Palaeoclimatology, Palaeoecology 618. doi:10.1016/j.palaeo.2023.111492. 



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