Great Ordovician Biodiversification Event

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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]

Contents

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 Late Cambrian and 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 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 Steptoean positive carbon isotope excursion 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]

In the Middle to Late Ordovician, after GOBE, 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

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. Ordovician meteor world.jpg
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.

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. [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. ZygospiraAttached.jpg
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 also 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

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The Ordovician is a geologic period and system, the second of six periods of the Paleozoic Era. The Ordovician spans 41.6 million years from the end of the Cambrian Period 485.4 Ma to the start of the Silurian Period 443.8 Ma.

The PaleozoicEra is the first of three geological eras of the Phanerozoic Eon. Beginning 538.8 million years ago (Ma), it succeeds the Neoproterozoic and ends 251.9 Ma at the start of the Mesozoic Era. The Paleozoic is subdivided into six geologic periods :

<span class="mw-page-title-main">Silurian</span> Third period of the Paleozoic Era, 443–419 million years ago

The Silurian 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. The Silurian is the shortest period of the Paleozoic Era. 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.

<span class="mw-page-title-main">Late Ordovician mass extinction</span> Extinction event around 444 million years ago

The Late Ordovician mass extinction (LOME), sometimes known as the end-Ordovician mass extinction or the Ordovician-Silurian extinction, is the first of the "big five" major mass extinction events in Earth's history, occurring roughly 445 million years ago (Ma). It is often considered to be the second-largest known extinction event just behind the end-Permian mass extinction, in terms of the percentage of genera that became extinct. Extinction was global during this interval, eliminating 49–60% of marine genera and nearly 85% of marine species. Under most tabulations, only the Permian-Triassic mass extinction exceeds the Late Ordovician mass extinction in biodiversity loss. The extinction event abruptly affected all major taxonomic groups and caused the disappearance of one third of all brachiopod and bryozoan families, as well as numerous groups of conodonts, trilobites, echinoderms, corals, bivalves, and graptolites. Despite its taxonomic severity, the Late Ordovician mass extinction did not produce major changes to ecosystem structures compared to other mass extinctions, nor did it lead to any particular morphological innovations. Diversity gradually recovered to pre-extinction levels over the first 5 million years of the Silurian period.

<span class="mw-page-title-main">Early Ordovician</span>

The Early Ordovician is the first epoch of the Ordovician period, corresponding to the Lower Ordovician series of the Ordovician system. It began after the Age 10 of the Furongian epoch of the Cambrian and lasted from 485.4 ± 1.9 to 470 ± 1.4 million years ago, until the Dapingian age of the Middle Ordovician. It includes Tremadocian and Floian ages.

<span class="mw-page-title-main">Tremadocian</span> Lowest stage of Ordovician

The Tremadocian is the lowest stage of Ordovician. Together with the later Floian Stage it forms the Lower Ordovician Epoch. The Tremadocian lasted from 485.4 to 477.7 million years ago. The base of the Tremadocian is defined as the first appearance of the conodont species Iapetognathus fluctivagus at the Global Boundary Stratotype Section and Point (GSSP) section on Newfoundland.

<span class="mw-page-title-main">Cambrian–Ordovician extinction event</span> Mass extinction event about 488 million years ago

The Cambrian–Ordovician extinction event, also known as the Cambrian-Ordovician boundary event, was an extinction event that occurred approximately 485 million years ago (mya) in the Paleozoic era of the early Phanerozoic eon. It was preceded by the less-documented End-Botomian mass extinction around 517 million years ago, and the Dresbachian extinction event about 502 million years ago.

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<span class="mw-page-title-main">Serpukhovian</span> Third stage of the Carboniferous

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<span class="mw-page-title-main">Fezouata Formation</span> Geological formation in Morocco

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<i>Aegirocassis</i> Extinct genus of radiodonts

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<span class="mw-page-title-main">Aegirocassisinae</span> Subfamily of hurdiid radiodonts

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The Steptoean positive carbon isotope excursion (SPICE) is a global chemostratigraphic event which occurred during the upper Cambrian period between 497 and 494 million years ago. This event corresponds with the ICS Guzhangian-Paibian Stage boundary and the Marjuman-Steptoean stage boundary in North America. The general signature of the SPICE event is a positive δ13C excursion, characterized by a 4 to 6 ‰ shift in δ13C values within carbonate successions around the world. SPICE was first described in 1993, and then named later in 1998. In both these studies, the SPICE excursion was identified and trends were observed within Cambrian formations of the Great Basin of the western United States.

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  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. S2CID   135413176 . Retrieved 15 July 2023.
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  27. ↑ Zhang, Junpeng; Li, Chao; Fang, Xiang; Li, Wenjie; Deng, Yiying; Tu, Chenyi; Algeo, Thomas J.; Lyons, Timothy W.; Zhang, Yuandong (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. Bibcode:2022E&PSL.59817858Z. doi: 10.1016/j.epsl.2022.117858 . ISSN   0012-821X.
  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". Nature Communications . 8: 14066. Bibcode:2017NatCo...814066L. doi:10.1038/ncomms14066. ISSN   2041-1723. PMC   5286199 . 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
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  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. Bibcode:2021NatCo..12.6430R. doi: 10.1038/s41467-021-26396-4 . PMC   8571325 . 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. Bibcode:2010Geo....38..547L. doi:10.1130/G30870.1.
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  34. ↑ Stigall, A.L; et al. (December 2016). "Biotic immigration events, speciation, and the accumulation of biodiversity in the fossil record". Global and Planetary Change . 148: 242–257. Bibcode:2017GPC...148..242S. doi:10.1016/j.gloplacha.2016.12.008.
  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. Bibcode:2023PPP...61511448S. 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 . 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 . 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 (1): 111–136. Bibcode:2023GSLSP.532..168S. doi: 10.1144/SP532-2022-168 . S2CID   254297330.
  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. Bibcode:2006PPP...232..148H. doi:10.1016/j.palaeo.2005.07.010 . Retrieved 15 July 2023.
  40. 1 2 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. Bibcode:2009PLoSO...4.7262K. doi: 10.1371/journal.pone.0007262 . PMC   2749442 . PMID   19789709.
  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. Bibcode:2023PPP...62511691E. 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. Bibcode:2023PPP...62611692M. doi:10.1016/j.palaeo.2023.111692. S2CID   259492773 . 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 . hdl: 20.500.12210/34267 .
  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. Bibcode:2023PPP...62311592S. 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. hdl: 20.500.12210/34395 . S2CID   134062318 . Retrieved 4 July 2023.
  46. ↑ Deng, Yiying; Fan, Junxuan; Yang, Shengchao; Shi, Yukun; Lu, Zhengbo; Xu, Huiqing; Sun, Zongyuan; Zhao, Fangqi; Hou, Zhangshuai (15 May 2023). "No Furongian Biodiversity Gap: Evidence from South China". Palaeogeography, Palaeoclimatology, Palaeoecology . 618. Bibcode:2023PPP...61811492D. doi: 10.1016/j.palaeo.2023.111492 .
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