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Carboniferous-Earliest Permian Biodiversification Event

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The Carboniferous-Earliest Permian Biodiversification Event (CPBE) was an evolutionary radiation of marine life, predominantly of fusulinid foraminifera and brachiopods, during most of the Carboniferous period and the earliest part of the Permian period, beginning in the late Visean and ending in the late Asselian, lasting about 41.2 million years in total. The radiation increased marine species richness and most profoundly diversified fusulinids and brachiopods, although the diversity of nektonic conodonts and cephalopods declined during this time, while the diversity of corals increased for only a brief period before plateauing in the middle of the radiation.

Causes[edit]

The Hangenberg event, marking the boundary between the Devonian and Carboniferous, is widely recognised as one of the largest mass extinction events of the Phanerozoic, with a species extinction rate of 70-82%. The event most severely affected trilobites, ammonoids, foraminifera, stromatoporoids, tabulate corals, and phytoplankton, leaving an abundance of vacant ecological niches in its wake. Following a brief re-radiation during the early Tournaisian occurred an episode of ocean hypoxia spurred by increased marine productivity known as the Mid-Tournaisian Event.[1] Though much less severe than the Hangenberg event, this extinction event further reduced marine biodiversity and vacated more niches. A third drop in species richness similar in magnitude to the Mid-Tournaisian Event occurred during the middle Visean.[2]

Two distinct pulses of global cooling marked by positive δ13C and δ15N excursions, one during the middle Tournaisian and another during the early Visean, precipitated glaciations marking the start of the Late Palaeozoic Ice Age (LPIA).[3] The high-frequency, high-amplitude glacioeustasy and consequent rapid shifts in littoral and neritic zones that characterised the LPIA have been hypothesised to be a driving factor of the high rates of speciation among fusulinids.[4] Around the same time as the beginning of the LPIA, the closure of the Rheic Ocean became complete as the continents of Gondwana and Euramerica collided to form the supercontinent Pangaea, reducing ocean circulation and resulting in widespread oligotrophic conditions favourable for fusulinids. In addition, the geographic isolation of different marine ecoregions created by the simultaneous onset of glaciation and closure of the Rheic Ocean would have further spurred rapid rates of diversification among fusulinids and other marine clades.[2]

Intervals[edit]

Six distinct intervals of biotic change leading up to and during the CPBE have been identified and labelled I1-I6.[2]

I1[edit]

I1 is a stasis interval spanning from the Famennian to the middle Visean. With the exception of the Hangenberg event, the interval was marked by relative stasis in the overall species richness. The average species origination rate (23.65%/2 Myr) closely matches the species extinction rate (25.08%/2 Myr) across the interval, albeit with high turnover, with sub-intervals of high species origination and low species extinction rates being followed by sub-intervals of low species origination and high species extinction and vice versa. Species richness during this interval peaks at the end of the Tournaisian. Species origination during this interval was dominated by conodonts, with speciation of brachiopods and corals being secondary and tertiary, respectively, with regards to overall species origination.

I2[edit]

I2 marks the beginning of the CPBE proper and represents the first distinct pulse of increase in species richness, lasting from the middle Visean to the early Bashkirian. The species origination rate was around 15%/2 Myr at the beginning of the interval, but it dropped to only around 5%/2 Myr by the end of it, with the species extinction rate remaining constant at just above 5%/2 Myr throughout the interval. The Serpukhovian extinction, an unusual extinction event caused not by an increase in extinction rates but a decline in speciation rates, a decline characteristic of this interval, occurred during I2. Conodont diversity declined during this period, while brachiopods represented most of the increase in diversity, with foraminifera and corals also contributing.

I3[edit]

I3 marks an interval of rapid increase in diversity spanning from the early Bashkirian to the late Moscovian, marking the beginning of the second distinct pulse of biodiversification. Both species origination and species extinction rates increased, but the former's increase was greater than the latter's, with the average rates across the whole interval being 25.57%/2 Myr and 18.21%/2 Myr, respectively. At the very beginning of the interval, both origination and extinction rates climbed to the same levels as in I1, although the extinction rate quickly dropped and remained relatively low for the rest of the interval, whereas the origination rate remained high. During I3, foraminifera became the largest contributor to species origination, with brachiopods becoming secondary in proportional contributions to diversification for the remainder of the CPBE. Coral and conodont diversity slightly declined during I3.

I4[edit]

I4 represents an interval marked by decline in species richness, interrupting the broader trend of increase in diversity. The average species extinction rate was 26.28%/2 Myr, while the species origination rate was only 15.59%/2 Myr. I4 spans from the late Moscovian to the late Kasimovian, roughly corresponding to the terrestrial extinction brought about by the Carboniferous rainforest collapse. Foraminifera, brachiopods, conodonts, and corals all declined in diversity throughout this interval.

I5[edit]

I5 represents the interval of fastest biodiversity increase, which was driven overwhelmingly by foraminifera as brachiopod diversity increased less significantly and conodont and coral diversity increased very slightly and virtually plateaued. Species origination was at 29.21%/2 Myr and species extinction at only 17.32%/2 Myr over the course of the interval. I5 spans from the Late Kasimovian to the end of the Asselian.

I6[edit]

I6 spans the Sakmarian and represents the aftermath of the CPBE, with the average species extinction rate (23.10%/2 Myr) slightly the species exceeding origination rate (21.43%/2 Myr).

See also[edit]

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References[edit]

  1. Yao, Le; Qie, Wenkun; Luo, Genming; Liu, Jiangsi; Algeo, Thomas J.; Bai, Xiao; Yang, Bo; Wang, Xiangdong (24 April 2015). "The TICE event: Perturbation of carbon–nitrogen cycles during the mid-Tournaisian (Early Carboniferous) greenhouse–icehouse transition". Chemical Geology. 401: 1–14. doi:10.1016/j.chemgeo.2015.02.021. Retrieved 4 September 2022.
  2. 2.0 2.1 2.2 Shi, Yukun; Wang, Xiangdong; Fan, Junxuan; Huang, Hao; Xu, Huiqing; Zhao, Yingying; Shen, Shuzhong (September 2021). "Carboniferous-earliest Permian marine biodiversification event (CPBE) during the Late Paleozoic Ice Age". Earth-Science Reviews. 220. doi:10.1016/j.earscirev.2021.103699. Retrieved 4 September 2022.
  3. Liu, Jiangsi; Algeo, Thomas J.; Qie, Wenkun; Saltzman, Matthew R. (1 October 2019). "Intensified oceanic circulation during Early Carboniferous cooling events: Evidence from carbon and nitrogen isotopes". Palaeogeography, Palaeoclimatology, Palaeoecology. 531. doi:10.1016/j.palaeo.2018.10.021. Retrieved 4 September 2022.
  4. Groves, John R.; Yue, Wang (1 September 2009). "Foraminiferal diversification during the late Paleozoic ice age". Paleobiology. 35 (3): 367–392. doi:10.1666/0094-8373-35.3.367. Retrieved 4 September 2022.


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