Middle Miocene disruption

Short description: Mass extinction event

The Middle Miocene Climatic Transition (MMCT) was a relatively steady period of climatic cooling that occurred around the middle of the Miocene, roughly 14 million years ago (Ma), during the Langhian stage,[1] and resulted in the growth of ice sheet volumes globally, and the reestablishment of the ice of the East Antarctic Ice Sheet (EAIS).[2] The term Middle Miocene disruption, alternatively the Middle Miocene extinction or Middle Miocene extinction peak, refers to a wave of extinctions of terrestrial and aquatic life forms that occurred during this climatic interval. This period was preceded by the Middle Miocene Climatic Optimum (MMCO), a period of relative warmth from 18 to 14 Ma.[3] Cooling that led to the Middle Miocene disruption is primarily attributed CO2 being pulled out of the Earth's atmosphere by organic material before becoming caught in different locations like the Monterey Formation.[4] These may have been amplified by changes in oceanic and atmospheric circulation due to continental drift.[1] Additionally, orbitally paced factors may also have played a role.[5]

## Contents

* 1 Effects
* 2 Suggested causes
* 3 Extinction event
* 4 References
* 5 Further reading
* 6 External links

## Effects

One of the primary effects of the climatic cooling that took place during this time period was the growth of the EAIS,[6][4] termed the East Antarctic Ice Sheet Expansion (EAIE).[7] A thermal shift from wet to cold-based glaciers is recorded from the Transantarctic Mountains about 13.94 Ma, reflecting a mean annual temperature drop of 25–30 °C.[8] Significant sections of ice on the Antarctic continent are believed to have started growth at the beginning of the Middle Miocene disruption and continued to expand until about 10 Ma.[9] This growth has been attributed primarily to orbitally paced changes in oceanic and atmospheric currents, with possible amplification by a significant drop in atmospheric carbon dioxide (ppm): atmospheric CO2 fell temporarily from about 300 to 140 ppm as estimated by the relationship between atmospheric levels of CO2 and pH levels in the ocean determined by boron isotopic levels in calcium carbonate.[2] One of the primary indicators for the significant global ice sheet growth is the higher concentration of 18O found in benthic foraminifera from oceanic sediment cores during this time period. During periods of ice sheet growth, the lighter 16O isotopes found in ocean water are drawn out as precipitation and consolidate in ice sheets while a higher concentration of 18O is left behind for foraminifera to utilize.[10] The >180° phase reversal in the 41,000-year obliquity cycle around 14.0 to 13.8 Ma has also been interpreted as a signal of the EAIE.[7]

During the MMCT, the latitudinal precipitation gradient declined in Europe, though it increased during short term warming periods superimposed on the broader cooling trend,[11] whereas the seasonality of mean temperature increased.[12] The seasonality of precipitation appears to have broken down, however, with European precipitation patterns being irregular on a multiannual to decadal timescale.[13] In Anatolia, arboreal and non-arboreal vegetation fluctuated in abundance, indicating an increase in complex, vegetationally heterogeneous habitats.[14] Global cooling during the MMCT caused aridification in North Africa and South Asia.[15] In the Qaidam Basin, silicate weathering sharply decreased around 12.6 Ma, indicating a major aridification event.[16] Savanna biomes spread across Central and East Asia in response to the increased aridity.[17] In the Columbia River Basalt Group (CRBG), the cessation of kaolin-producing pedogenic processes occurred at the start of the MMCT and has been used as a proxy marker for the end of the MMCO.[18] Southwestern Australia exhibited the most arid conditions it had witnessed over any interval of the Miocene, while northwestern Australia was also hyperarid.[19]

Significant drop-off in both temperature and deep-sea ocean temperature as measured by delta 18O after the Middle Miocene Climatic Optimum

## Suggested causes

The primary cause of the cooling that came out of the MMCO was changing atmospheric CO2 levels.[1] Falling CO2 concentrations in the atmosphere has been linked to drawdown of the gas into organic material deposited along continental margins like the Monterey Formation of coastal California, an explanation known as the Monterey Hypothesis.[20] These sites of CO2 drawdown are thought to have been extensive enough to drop atmospheric concentrations in CO2 from about 300 to 140 ppm and lead to processes of global cooling that helped in the expansion of the EAIS.[2]

Organic carbon burial on land, evidenced by widespread formation of lignite deposits at this time, also contributed heavily to the reduction in pCO2.[21]

Another hypothesis is that increased silicate weathering of the uplifting Himalayas caused the MMCT, but this is contradicted by geological evidence from the Indus River system.[22]

As well significant changes in greenhouse gas concentrations, alterations to ocean circulation brought about major climatic and biotic changes. Oceanic circulation changes that took place during the MMCT are defined by increases in Antarctic Bottom Water (AABW) production, the halting of saline water delivery to the Southern Ocean from the Indian Ocean, and additional North Atlantic Deep Water production.[10] The reduction in water transport from the warm Indian Ocean to the cool Southern Ocean is believed to be responsible for the increase in AABW production.[23] The Tethys Seaway is believed to have closed around this time, exacerbating the disruptions of ocean circulation patterns that caused the MMCT.[1] The northward movement of Australia hindered equatorial circulation between the Pacific and Indian Oceans, helping to cool tropical surface waters.[3] The cooling of the Southern Ocean was coupled to the growth of the EAIS.[24]

Another suggested cause is an encounter with a dense part of the gaseous component of the Radcliffe wave. It has been suggested that the Radcliffe wavefront crossing the Solar System coincides with the period of the Middle Miocene disruption.[25][26]

An additional suggested cause for the Middle Miocene disruption has been attributed to a shift from a solar insolation cycle that is obliquity dominated to one that is dominated by eccentricity (see Milankovitch cycles). This change would have been significant enough for conditions near the Antarctic continent to allow for glaciation.[5]

Vegetational changes are known to have exacerbated the cooling of the MMCT. The changes to the Earth's surface albedo caused by the southward retreat of needleleaf trees in the high latitudes of the Northern Hemisphere amplified the drop in temperatures.[27]

## Extinction event

The Middle Miocene disruption is considered a significant extinction event and has been analyzed in terms of the importance of there being a possible periodicity between extinction events. A study from David Raup and Jack Sepkoski found that there is a statistically significant mean periodicity (where P is less than .01) of about 26 million years for 12 major extinction events. There is debate whether this potential periodicity is caused by some set of recurrent cycles or biologic factors.[28]

A sharp drop in carbonate production, known as the Miocene Carbonate Crash (MCC), occurred during the early Tortonian, shortly after the cooling event; this event is generally regarded to have been induced by the changes in thermohaline circulation resulting from the Middle Miocene disruption.[29] Changes in the intensity and seasonality of the Indian monsoon have been suggested to have brought about this change in ocean circulation.[30] Another hypothesis for the crash involves the shrinkage and shoaling of the Central American Seaway, limiting water mass exchange between the Atlantic and Pacific Oceans.[31] Evidence for this event is known from the Indian Ocean, Pacific Ocean, Atlantic Ocean, Caribbean Sea, and Mediterranean Sea, suggesting the decline of carbonate-producing marine organisms was a global phenomenon.[29]

One of the other primary effects of the climatic cooling during the Middle Miocene was the biotic impact on terrestrial and oceanic lifeforms. A primary example of these extinctions is indicated by the observed occurrence of Varanidae, chameleons, Cordylidae, Tomistominae, Alligatoridae, and giant turtles through the Miocene Climatic Optimum (18 to 16 Ma) in Central Europe (45–42 °N palaeolatitude). This was then followed by a major and permanent cooling step marked by the Mid-Miocene disruption between 14.8 and 14.1 Ma. Two crocodilians of the genera Gavialosuchus and Diplocynodon were noted to have been extant in these northern latitudes prior to the permanent cooling step, but then became extinct between 14 and 13.5 Ma.[32] Another indicator that would lead to extinctions is the conservative estimate that temperatures in the Antarctic region may have cooled by at least 8 °C in the summer months 14 Ma. This Antarctic cooling, along with significant changes in temperature gradients in Central Europe as indicated by Madelaine Böhme's study on ectothermic vertebrates, provide evidence that plant and animal life needed to migrate or adapt in order to survive.[33] In South America, the MMCT is believed to have begun the decline of hathliacynids that led to their eventual extinction.[34]

## References

1. ↑ 1.0 1.1 1.2 1.3 Hamon, N.; Sepulchre, P.; Lefebvre, V.; Ramstein, G. (28 November 2013). "The role of eastern Tethys seaway closure in the Middle Miocene Climatic Transition (ca. 14 Ma)" (in en). Climate of the Past 9 (6): 2687–2702. doi:10.5194/cp-9-2687-2013. ISSN 1814-9332. Bibcode: 2013CliPa...9.2687H. https://cp.copernicus.org/articles/9/2687/2013/. Retrieved 31 December 2023.
2. ↑ 2.0 2.1 2.2 Pearson, Paul N.; Palmer, Martin R. (2000). "Atmospheric carbon dioxide concentrations over the past 60 million years". Nature 406 (6797): 695–699. doi:10.1038/35021000. PMID 10963587. Bibcode: 2000Natur.406..695P.
3. ↑ 3.0 3.1 Scotese, Christopher Robert; Song, Haijun; Mills, Benjamin J.W.; van der Meer, Douwe G. (1 April 2021). "Phanerozoic paleotemperatures: The earth's changing climate during the last 540 million years" (in en). Earth-Science Reviews 215. doi:10.1016/j.earscirev.2021.103503. Bibcode: 2021ESRv..21503503S. https://linkinghub.elsevier.com/retrieve/pii/S0012825221000027. Retrieved 24 December 2023.
4. ↑ 4.0 4.1 Shevenell, Amelia E.; Kennett, James P.; Lea, David W. (17 September 2004). "Middle Miocene Southern Ocean Cooling and Antarctic Cryosphere Expansion" (in en). Science 305 (5691): 1766–1770. doi:10.1126/science.1100061. ISSN 0036-8075. PMID 15375266. Bibcode: 2004Sci...305.1766S.
5. ↑ 5.0 5.1 Holbourn, Ann; Kuhnt, Wolfgang; Schulz, Michael; Erlenkeuser, Helmut (24 November 2005). "Impacts of orbital forcing and atmospheric carbon dioxide on Miocene ice-sheet expansion". Nature 438 (7067): 483–487. doi:10.1038/nature04123. PMID 16306989. Bibcode: 2005Natur.438..483H. https://www.nature.com/articles/nature04123. Retrieved 19 April 2023.
6. ↑ Langebroek, P. M.; Paul, A.; Schulz, M. (22 October 2009). "Antarctic ice-sheet response to atmospheric CO<sub>2</sub> and insolation in the Middle Miocene" (in en). Climate of the Past 5 (4): 633–646. doi:10.5194/cp-5-633-2009. ISSN 1814-9332. Bibcode: 2009CliPa...5..633L. https://cp.copernicus.org/articles/5/633/2009/. Retrieved 4 July 2024.
7. ↑ 7.0 7.1 Tian, Jun; Yang, Mei; Lyle, Mitchell W.; Wilkens, Roy; Shackford, Julia K. (11 March 2013). "Obliquity and long eccentricity pacing of the Middle Miocene climate transition" (in en). Geochemistry, Geophysics, Geosystems 14 (6): 1740–1755. doi:10.1002/ggge.20108. ISSN 1525-2027. Bibcode: 2013GGG....14.1740T. https://agupubs.onlinelibrary.wiley.com/doi/10.1002/ggge.20108. Retrieved 31 December 2023.
8. ↑ Lewis, A.R.; Marchant, D.R.; Ashworth, A.C.; Hemming, S.R.; Machlus, M.L. (1 November 2007). "Major middle Miocene global climate change: Evidence from East Antarctica and the Transantarctic Mountains" (in en). Geological Society of America Bulletin 119 (11–12): 1449–1461. doi:10.1130/0016-7606(2007)119[1449:MMMGCC2.0.CO;2]. ISSN 0016-7606. https://pubs.geoscienceworld.org/gsabulletin/article/119/11-12/1449-1461/125372. Retrieved 31 December 2023.
9. ↑ Zachos, James C.; Pagani, Mark; Sloan, Lisa; Thomas, Ellen; Billups, Katharina (27 April 2001). "Trends, Rhythms, and Aberrations in Global Climate 65 Ma to Present" (in en). Science 292 (5517): 686–693. doi:10.1126/science.1059412. ISSN 0036-8075. PMID 11326091. Bibcode: 2001Sci...292..686Z. http://doc.rero.ch/record/13508/files/PAL_E304.pdf. Retrieved 19 April 2023.
10. ↑ 10.0 10.1 Flower, B. P.; Kennett, J. P. (December 1993). "Middle Miocene ocean-climate transition: High-resolution oxygen and carbon isotopic records from Deep Sea Drilling Project Site 588A, southwest Pacific". Paleoceanography and Paleoclimatology 8 (6): 811–843. doi:10.1029/93pa02196. Bibcode: 1993PalOc...8..811F.
11. ↑ Böhme, Madelaine; Winklhofer, Michael; Ilg, August (1 May 2011). "Miocene precipitation in Europe: Temporal trends and spatial gradients". Palaeogeography, Palaeoclimatology, Palaeoecology. The Neogene of Eurasia: Spatial gradients and temporal trends - The second synthesis of NECLIME 304 (3): 212–218. doi:10.1016/j.palaeo.2010.09.028. ISSN 0031-0182. Bibcode: 2011PPP...304..212B. https://www.sciencedirect.com/science/article/pii/S0031018210005997. Retrieved 11 January 2024.
12. ↑ Bruch, Angela A.; Utescher, Torsten; Mosbrugger, Volker (1 May 2011). "Precipitation patterns in the Miocene of Central Europe and the development of continentality". Palaeogeography, Palaeoclimatology, Palaeoecology. The Neogene of Eurasia: Spatial gradients and temporal trends - The second synthesis of NECLIME 304 (3): 202–211. doi:10.1016/j.palaeo.2010.10.002. ISSN 0031-0182. Bibcode: 2011PPP...304..202B. https://www.sciencedirect.com/science/article/pii/S0031018210006073. Retrieved 11 January 2024.
13. ↑ Harzhauser, Mathias; Piller, Werner E.; Müllegger, Stefan; Grunert, Patrick; Micheels, Arne (March 2011). "Changing seasonality patterns in Central Europe from Miocene Climate Optimum to Miocene Climate Transition deduced from the Crassostrea isotope archive" (in en). Global and Planetary Change 76 (1–2): 77–84. doi:10.1016/j.gloplacha.2010.12.003. Bibcode: 2011GPC....76...77H. https://www.sciencedirect.com/science/article/pii/S0921818110002596. Retrieved 21 September 2025.
14. ↑ Bouchal, Johannes Martin; Güner, Tuncay Hüseyin; Denk, Thomas (17 October 2018). "Middle Miocene climate of southwestern Anatolia from multiple botanical proxies" (in en). Climate of the Past 14 (10): 1427–1440. doi:10.5194/cp-14-1427-2018. ISSN 1814-9332. Bibcode: 2018CliPa..14.1427B. https://cp.copernicus.org/articles/14/1427/2018/. Retrieved 21 September 2025.
15. ↑ Zhang, Jian; Hu, Yongyun; Zhu, Chenguang; Flögel, Sascha; Fang, Xiaomin; Sun, Jimin (1 June 2023). "Modeling the effects of global cooling and the Tethyan Seaway closure on North African and South Asian climates during the Middle Miocene Climate Transition" (in en). Palaeogeography, Palaeoclimatology, Palaeoecology 619. doi:10.1016/j.palaeo.2023.111541. Bibcode: 2023PPP...61911541Z. https://linkinghub.elsevier.com/retrieve/pii/S0031018223001591. Retrieved 4 July 2024.
16. ↑ Bao, Jing; Song, Chunhui; Yang, Yibo; Fang, Xiaomin; Meng, Qingquan; Feng, Ying; He, Pengju (1 February 2019). "Reduced chemical weathering intensity in the Qaidam Basin (NE Tibetan Plateau) during the Late Cenozoic". Journal of Asian Earth Sciences 170: 155–165. doi:10.1016/j.jseaes.2018.10.018. ISSN 1367-9120. Bibcode: 2019JAESc.170..155B. https://www.sciencedirect.com/science/article/pii/S1367912018304425. Retrieved 31 December 2023.
17. ↑ Li, Chunxiao; Wang, Shi-Qi; Wang, Yang; Deng, Tao; Ma, Jiao; Wang, Bian; Jiangzuo, Qigao; Sun, Danhui (15 July 2025). "Savanna ecosystems and mammalian adaptations in Mid-Miocene Northern China" (in en). Scientific Reports 15 (1): 25586. doi:10.1038/s41598-025-10718-3. ISSN 2045-2322. PMID 40664786. Bibcode: 2025NatSR..1525586L.
18. ↑ Hobbs, Kevin Michael; Parrish, Judith Totman (1 September 2016). "Miocene global change recorded in Columbia River basalt–hosted paleosols" (in en). Geological Society of America Bulletin 128 (9–10): 1543–1554. doi:10.1130/B31437.1. ISSN 0016-7606. Bibcode: 2016GSAB..128.1543H. https://pubs.geoscienceworld.org/gsabulletin/article/128/9-10/1543-1554/185433. Retrieved 11 January 2024.
19. ↑ Groeneveld, Jeroen; Henderiks, Jorijntje; Renema, Willem; McHugh, Cecilia M.; De Vleeschouwer, David; Christensen, Beth A.; Fulthorpe, Craig S.; Reuning, Lars et al. (5 May 2017). "Australian shelf sediments reveal shifts in Miocene Southern Hemisphere westerlies" (in en). Science Advances 3 (5). doi:10.1126/sciadv.1602567. ISSN 2375-2548. PMID 28508066. Bibcode: 2017SciA....3E2567G.
20. ↑ Kashiwagi, Hirohiko; Shikazono, Naotatsu (25 October 2003). "Climate change during Cenozoic inferred from global carbon cycle model including igneous and hydrothermal activities". Palaeogeography, Palaeoclimatology, Palaeoecology 199 (3): 167–185. doi:10.1016/S0031-0182(03)00506-6. ISSN 0031-0182. Bibcode: 2003PPP...199..167K. https://www.sciencedirect.com/science/article/pii/S0031018203005066. Retrieved 11 January 2024.
21. ↑ Diester-Haass, Liselotte; Billups, Katharina; Gröcke, Darren R.; François, Louis; Lefebvre, Vincent; Emeis, Kay C. (18 February 2009). "Mid-Miocene paleoproductivity in the Atlantic Ocean and implications for the global carbon cycle" (in en). Paleoceanography and Paleoclimatology 24 (1). doi:10.1029/2008PA001605. ISSN 0883-8305. Bibcode: 2009PalOc..24.1209D. https://agupubs.onlinelibrary.wiley.com/doi/10.1029/2008PA001605. Retrieved 11 January 2024.
22. ↑ Clift, Peter D.; Jonell, Tara N. (28 April 2021). "Himalayan-Tibetan Erosion Is Not the Cause of Neogene Global Cooling" (in en). Geophysical Research Letters 48 (8). doi:10.1029/2020GL087742. ISSN 0094-8276. Bibcode: 2021GeoRL..4887742C. https://agupubs.onlinelibrary.wiley.com/doi/10.1029/2020GL087742. Retrieved 11 January 2024.
23. ↑ Flower, Benjamin P.; Kennett, James P. (1 April 1994). "The middle Miocene climatic transition: East Antarctic ice sheet development, deep ocean circulation and global carbon cycling". Palaeogeography, Palaeoclimatology, Palaeoecology 108 (3): 537–555. doi:10.1016/0031-0182(94)90251-8. ISSN 0031-0182. Bibcode: 1994PPP...108..537F.
24. ↑ Leutert, Thomas J.; Auderset, Alexandra; Martínez-García, Alfredo; Modestou, Sevasti; Meckler, A. Nele (31 August 2020). "Coupled Southern Ocean cooling and Antarctic ice sheet expansion during the middle Miocene" (in en). Nature Geoscience 13 (9): 634–639. doi:10.1038/s41561-020-0623-0. ISSN 1752-0908. Bibcode: 2020NatGe..13..634L. https://www.nature.com/articles/s41561-020-0623-0. Retrieved 31 December 2023.
25. ↑ Maconi, E.; Alves, J.; Swiggum, C.; Ratzenböck, S.; Großschedl, J.; Köhler, P.; Miret-Roig, N.; Meingast, S. et al. (11 February 2025). "The Solar System's passage through the Radcliffe wave during the middle Miocene". Astronomy & Astrophysics 694: A167. doi:10.1051/0004-6361/202452061. ISSN 0004-6361. Bibcode: 2025A&A...694A.167M. https://www.aanda.org/articles/aa/full_html/2025/02/aa52061-24/aa52061-24.html. Retrieved 21 September 2025.
26. ↑ Gough, Evan (12 March 2025). "Study Traces Our Solar System's Journey Through a Massive Galactic Wave" (in en-US). https://www.sciencealert.com/study-traces-our-solar-systems-journey-through-a-massive-galactic-wave.
27. ↑ Zhang, Ran; Guo, Jiaqi; Bradshaw, Catherine D.; Xu, Xiyan; Shen, Tiantian; Li, Shufeng; Nie, Junsheng; Zhang, Chunxia et al. (2 May 2025). "Vegetation feedbacks accelerated the late Miocene climate transition" (in en). Science Advances 11 (18). doi:10.1126/sciadv.ads4268. ISSN 2375-2548. PMID 40315310. Bibcode: 2025SciA...11S4268Z.
28. ↑ Raup, D. M.; Sepkoski, J. J. (1 February 1984). "Periodicity of extinctions in the geologic past" (in en). Proceedings of the National Academy of Sciences of the United States of America 81 (3): 801–805. doi:10.1073/pnas.81.3.801. ISSN 0027-8424. PMID 6583680. Bibcode: 1984PNAS...81..801R.
29. ↑ 29.0 29.1 Torfstein, Adi; Steinberg, Josh (14 August 2020). "The Oligo–Miocene closure of the Tethys Ocean and evolution of the proto-Mediterranean Sea". Scientific Reports 10 (1): 13817. doi:10.1038/s41598-020-70652-4. PMID 32796882.
30. ↑ Lübbers, Julia; Kuhnt, Wolfgang; Holbourn, Ann E.; Bolton, Clara T.; Gray, Emmeline; Usui, Yoichi; Kochhann, Karlos G. D.; Beil, Sebastian et al. (16 April 2019). "The Middle to Late Miocene "Carbonate Crash" in the Equatorial Indian Ocean". Paleoceanography and Paleoclimatology 34 (5): 813–832. doi:10.1029/2018PA003482. Bibcode: 2019PaPa...34..813L. https://agupubs.onlinelibrary.wiley.com/doi/full/10.1029/2018PA003482. Retrieved 19 April 2023.
31. ↑ Nisancioglu, Kerim Hestnes; Raymo, Maureen; Stone, Peter H. (11 February 2003). "Reorganization of Miocene deep water circulation in response to the shoaling of the Central American Seaway". Paleoceanography and Paleoclimatology 18 (1): 6-1-6-12. doi:10.1029/2002PA000767. Bibcode: 2003PalOc..18.1006N. https://www.researchgate.net/publication/228992752. Retrieved 21 April 2023.
32. ↑ Böhme, Madelaine (November 2001). "The Miocene Climatic Optimum: evidence from ectothermic vertebrates of Central Europe". Palaeogeography, Palaeoclimatology, Palaeoecology 195 (3–4): 389–401. doi:10.1016/S0031-0182(03)00367-5. http://www.wahre-staerke.com/~madelaine/Miocene_Climatic_Optimum.pdf. Retrieved 19 April 2023.
33. ↑ Lewis, Adam R.; Marchant, David R.; Ashworth, Allan C.; Hedenäs, Lars; Hemming, Sidney R.; Johnson, Jesse V.; Leng, Melanie J.; Machlus, Malka L. et al. (5 August 2008). "Mid-Miocene cooling and the extinction of tundra in continental Antarctica" (in en). Proceedings of the National Academy of Sciences of the United States of America 105 (31): 10676–10680. doi:10.1073/pnas.0802501105. ISSN 0027-8424. PMID 18678903. PMC 2495011. Bibcode: 2008PNAS..10510676L. https://www.researchgate.net/publication/23148177. Retrieved 19 April 2023.
34. ↑ Zimicz, Natalia (1 March 2014). "Avoiding Competition: the Ecological History of Late Cenozoic Metatherian Carnivores in South America" (in en). Journal of Mammalian Evolution 21 (4): 383–393. doi:10.1007/s10914-014-9255-8. ISSN 1064-7554. https://link.springer.com/article/10.1007/s10914-014-9255-8. Retrieved 21 September 2025.

## Further reading

* Allmon, Warren D.; Bottjer, David J. (2001). Evolutionary Paleoecology: The Ecological Context of Macroevolutionary Change. New York: Columbia University Press. ISBN 978-0-231-10994-9.

## External links

* Middle Miocene global map

Extinction events v * d * e

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