Great Ordovician Biodiversification Event

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

It followed a series of extinction events at the Cambrian-Ordovician boundary, 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][7] 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]

Causes

Possible causes include changes in palaeogeography or tectonic activity, as well as a modified nutrient supply.^[8] 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, favouring diversification.^[2] 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] On the other hand, global cooling has also been offered as a cause of the radiation, and another alternative is that the breakup of an asteroid led to the Earth being consistently pummelled by meteorites.^[3]

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, tropic layering, or by providing a new habitat.^{[clarification needed][8]} 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] 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.^[9]

Effects

Spiriferid 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,^[10] the GOBE can be considered as the 'filling out' of these phyla with the modern (and many extinct) classes and lower-level taxa.^[3]

Taxonomic diversity increased manifold; the total number of marine orders doubled, and families tripled.^[4] In addition to a diversification, the event also marked an increase in the complexity of both organisms and food webs.^[1] Taxa begun to have localized ranges, with different faunas at different parts of the globe.^[1] Communities in reefs and deeper water begun to take on a character of their own, becoming more clearly distinct from other marine ecosystems.^[1] And as ecosystems became more diverse, with more species being squeezed in to the food web, a more complex tangle of ecological interactions resulted, promoting strategies such as ecological tiering.^[1] 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]

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.^[2] The warm waters and high sea level (this 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]

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.^[11]

See also

Cambrian explosion

References

1. ^ ^{a b c d e f g h} 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.  edit
2. ^ ^{a b c d} 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: 99. doi:10.1111/j.1502-3931.2008.00115.x.  edit
3. ^ ^{a b c d e} 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. doi:10.1016/j.palaeo.2010.05.031.  edit
4. ^ ^{a b c} Droser, M. L.; Finnegan, S. (2003). "The Ordovician Radiation: A Follow-up to the Cambrian Explosion?". Integrative and Comparative Biology 43: 178. doi:10.1093/icb/43.1.178.  edit
5. ^ Marshall, C. R. (2006). "Explaining the Cambrian "explosion" of Animals". Annual Review of Earth and Planetary Sciences 34: 355–384. Bibcode 2006AREPS..34..355M. doi:10.1146/annurev.earth.33.031504.103001.  edit
6. ^ 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. doi:10.1666/06013.1.  edit
7. ^ Bambach, R. K.; Bush, A. M.; Erwin, D. H. (2007). "Autecology and the Filling of Ecospace: Key Metazoan Radiations". Palaeontology 50: 1. doi:10.1111/j.1475-4983.2006.00611.x.  edit
8. ^ ^{a b} Botting, Muir; Muir, Lucy A. (2008). "Unravelling Causal Components of the Ordovician Radiation: the Builth Inlier (Central Wales) As a Case Study". Lethaia 41: 111. doi:10.1111/j.1502-3931.2008.00118.x. 
9. ^ Solved: Mystery of Earth's First Breathable Atmosphere
10. ^ 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. doi:10.1130/G30870.1.  edit
11. ^ 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. PMC 2749442. PMID 19789709. http://www.plosone.org/article/info%3Adoi%2F10.1371%2Fjournal.pone.0007262. Retrieved 2009-10-30.  edit