| [1] | Raup, D.M., and Sepkoski, J.J.J. (1982). Mass extinctions in the marine fossil record. Science 215: 1501-1503. https://doi.org/10.1126/science.215.4539.1501. |
| [2] | Song, H., Kemp, D.B., Tian, L., et al. (2021). Thresholds of temperature change for mass extinctions. Nat. Commun. 12(1): 4694. https://doi.org/10.1038/s41467-021-25019-2. |
| [3] | Knoll, A.H., Bambach, R.K., Payne, J.L., et al. (2007). Paleophysiology and end-Permian mass extinction. Earth Planet Sci. Lett. 256(3–4): 295-313. https://doi.org/10.1016/j.epsl.2007.02.018. |
| [4] | Dal Corso, J., Song, H., Callegaro, S., et al. (2022). Environmental crises at the Permian–Triassic mass extinction. Nat. Rev. Earth Environ. 3(3): 197-214. https://doi.org/10.1038/s43017-021-00259-4. |
| [5] | Jablonski, D. (1986). Background and mass extinctions: the alternation of macroevolutionary regimes. Science 231: 129-133. https://doi.org/10.1126/science.231.4734.129. |
| [6] | Payne, J.L., Bush, A.M., Chang, E.T., et al. (2016). Extinction intensity, selectivity and their combined macroevolutionary influence in the fossil record. Biol. Lett. 12(10): 20160202. https://doi.org/10.1098/rsbl.2016.0202. |
| [7] | Stanley, S.M. (2016). Estimates of the magnitudes of major marine mass extinctions in earth history. Proc. Natl. Acad. Sci. USA 113(42): E6325-E6334. https://doi.org/10.1073/pnas.1613094113. |
| [8] | Song, H., Wignall, P.B., Tong, J., et al. (2013). Two pulses of extinction during the Permian-Triassic crisis. Nat. Geosci. 6(1): 52-56. https://doi.org/10.1038/ngeo1649. |
| [9] | Sepkoski, J.J., Jr. (1984). A kinetic model of Phanerozoic taxonomic diversity, Ⅲ. Post-Paleozoic families and mass extinctions. Paleobiology 10(2): 246-267. https://doi.org/10.1017/S0094837300008186. |
| [10] | Dai, X., Davies, J.H.F.L., Yuan, Z., et al. (2023). A Mesozoic fossil lagerstätte from 250.8 million years ago shows a modern-type marine ecosystem. Science 379(6632): 567-572. https://doi.org/10.1126/science.adf1622. |
| [11] | Muscente, A.D., Prabhu, A., Zhong, H., et al. (2018). Quantifying ecological impacts of mass extinctions with network analysis of fossil communities. Proc. Natl. Acad. Sci. USA 115(20): 5217-5222. https://doi.org/10.1073/pnas.1719976115. |
| [12] | Rojas, A., Calatayud, J., Kowalewski, M., et al. (2021). A multiscale view of the Phanerozoic fossil record reveals the three major biotic transitions. Commun. Biol. 4(1): 309. https://doi.org/10.1038/s42003-021-01805-y. |
| [13] | Clapham, M.E., and Payne, J.L. (2011). Acidification, anoxia, and extinction: A multiple logistic regression analysis of extinction selectivity during the Middle and Late Permian. Geology 39(11): 1059-1062. https://doi.org/10.1130/g32230.1. |
| [14] | Kiessling, W., and Simpson, C. (2011). On the potential for ocean acidification to be a general cause of ancient reef crises. Glob. Chang. Biol. 17(1): 56-67. https://doi.org/10.1111/j.1365-2486.2010.02204.x. |
| [15] | Foster, W.J., Ayzel, G., Münchmeyer, J., et al. (2022). Machine learning identifies ecological selectivity patterns across the end-Permian mass extinction. Paleobiology 48(3): 357-371. https://doi.org/10.1017/pab.2022.1. |
| [16] | Brayard, A., Meier, M., Escarguel, G., et al. (2015). Early Triassic Gulliver gastropods: Spatio-temporal distribution and significance for biotic recovery after the end-Permian mass extinction. Earth Sci. Rev. 146: 31-64. https://doi.org/10.1016/j.earscirev.2015.03.005. |
| [17] | Romano, C., Koot, M.B., Kogan, I., et al. (2016). Permian–Triassic Osteichthyes (bony fishes): diversity dynamics and body size evolution. Biol. Rev. 91(1): 106-147. https://doi.org/10.1111/brv.12161. |
| [18] | Schaal, E.K., Clapham, M.E., Rego, B.L., et al. (2016). Comparative size evolution of marine clades from the Late Permian through Middle Triassic. Paleobiology 42(1): 127-142. https://doi.org/10.1017/pab.2015.36. |
| [19] | He, W.H., Shi, G., Twitchett, R., et al. (2015). Late Permian marine ecosystem collapse began in deeper waters: evidence from brachiopod diversity and body size changes. Geobiology 13(2): 123-138. https://doi.org/10.1111/gbi.12119. |
| [20] | Jablonski, D., and Raup, D.M. (1995). Selectivity of end-Cretaceous marine bivalve extinctions. Science 268(5209): 389-391. https://doi.org/10.1126/science.11536722. |
| [21] | Payne, J.L., and Finnegan, S. (2007). The effect of geographic range on extinction risk during background and mass extinction. Proc. Natl. Acad. Sci. USA 104(25): 10506-10511. https://doi.org/10.1073/pnas.0701257104. |
| [22] | Suzuki, T., and Imai, K. (1998). Evolution of myoglobin. Cell. Mol. Life Sci. 54(9): 979-1004. https://doi.org/10.1007/s000180050227. |
| [23] | Terwilliger, N.B. (1998). Functional adaptations of oxygen-transport proteins. J. Exp. Biol. 201(8): 1085-1098. https://doi.org/10.1242/jeb.201.8.1085. |
| [24] | Song, H., Wignall, P.B., and Dunhill, A.M. (2018). Decoupled taxonomic and ecological recoveries from the Permo-Triassic extinction. Sci. Adv. 4(10): eaat5091. https://doi.org/10.1126/sciadv.aat5091. |
| [25] | Payne, J.L., Bush, A.M., Heim, N.A., et al. (2016). Ecological selectivity of the emerging mass extinction in the oceans. Science 353(6305): 1284-1286. https://doi.org/10.1126/science.aaf2416. |
| [26] | Song, S., Starunov, V., Bailly, X., et al. (2020). Globins in the marine annelid Platynereis dumerilii shed new light on hemoglobin evolution in bilaterians. BMC Evol. Biol. 20(1): 165. https://doi.org/10.1186/s12862-020-01714-4. |
| [27] | Mangum, C.P. (2011). Invertebrate blood oxygen carriers. Compr. Physiol. : 1097-1135. https://doi.org/10.1002/cphy.cp130215. |
| [28] | Heim, N.A., Bakshi, S.H., Buu, L., et al. (2020). Respiratory medium and circulatory anatomy constrain size evolution in marine macrofauna. Paleobiology 46(3): 288-303. https://doi.org/10.1017/pab.2020.16. |
| [29] | Lau, K.V., Maher, K., Altiner, D., et al. (2016). Marine anoxia and delayed Earth system recovery after the end-Permian extinction. Proc. Natl. Acad. Sci. USA 113(9): 2360-2365. https://doi.org/10.1073/pnas.1515080113. |
| [30] | Zhang, F., Romaniello, S.J., Algeo, T.J., et al. (2018). Multiple episodes of extensive marine anoxia linked to global warming and continental weathering following the latest Permian mass extinction. Sci. Adv. 4(4): e1602921. https://doi.org/10.1126/sciadv.1602921. |
| [31] | Pimentel-Galvan, M., Lau, K.V., Maher, K., et al. (2022). Duration and intensity of end-Permian marine anoxia. Geochem. Geophys. Geosyst. 23(1): e2021GC010130. https://doi.org/10.1029/2021GC010130. |
| [32] | Deutsch, C., Ferrel, A., Seibel, B., et al. (2015). Climate change tightens a metabolic constraint on marine habitats. Science 348(6239): 1132-1135. https://doi.org/10.1126/science.aaa1605. |
| [33] | Penn, J.L., Deutsch, C., Payne, J.L., et al. (2018). Temperature-dependent hypoxia explains biogeography and severity of end-Permian marine mass extinction. Science 362(6419): eaat1327. https://doi.org/10.1126/science.aat1327. |
| [34] | Hülse, D., Lau, K.V., van de Velde, S.J., et al. (2021). End-Permian marine extinction due to temperature-driven nutrient recycling and euxinia. Nat. Geosci. 14(11): 862-867. https://doi.org/10.1038/s41561-021-00829-7. |
| [35] | Song, H., Wignall, P.B., Chu, D., et al. (2014). Anoxia/high temperature double whammy during the Permian-Triassic marine crisis and its aftermath. Sci. Rep. 4: 4132. https://doi.org/10.1038/srep04132. |
| [36] | Dabruzzi, T.F., and Bennett, W.A. (2014). Hypoxia effects on gill surface area and blood oxygen-carrying capacity of the Atlantic stingray, Dasyatis sabina. Fish Physiol. Biochem. 40(4): 1011-1020. https://doi.org/10.1007/s10695-013-9901-8. |
| [37] | Rubalcaba, J.G., Verberk, W.C.E.P., Hendriks, J., et al. (2020). Oxygen limitation may affect the temperature and size dependence of metabolism in aquatic ectotherms. Proc. Natl. Acad. Sci. USA 117(50): 31963-31968. https://doi.org/10.1073/pnas.2003292117. |
| [38] | Feng, Y., Song, H., and Bond, D.P.G. (2020). Size variations in foraminifers from the early Permian to the Late Triassic: implications for the Guadalupian–Lopingian and the Permian–Triassic mass extinctions. Paleobiology 46(4): 511-532. https://doi.org/10.1017/pab.2020.37. |
| [39] | Wu, Y., Chu, D., Tong, J., et al. (2021). Six-fold increase of atmospheric pCO2 during the Permian–Triassic mass extinction. Nat. Commun. 12(1): 2137. https://doi.org/10.1038/s41467-021-22298-7. |
| [40] | Wu, Y., Cui, Y., Chu, D., et al. (2023). Volcanic CO2 degassing postdates thermogenic carbon emission during the end-Permian mass extinction. Sci. Adv. 9(7): eabq4082. https://doi.org/10.1126/sciadv.abq4082. |
| [41] | Orr, J.C., Fabry, V.J., Aumont, O., et al. (2005). Anthropogenic ocean acidification over the twenty-first century and its impact on calcifying organisms. Nature 437(7059): 681-686. https://doi.org/10.1038/nature04095. |
| [42] | Watson, S. -A., Morley, S.A., and Peck, L.S. (2017). Latitudinal trends in shell production cost from the tropics to the poles. Sci. Adv. 3(9): e1701362. https://doi.org/10.1126/sciadv.1701362. |
| [43] | Jurikova, H., Gutjahr, M., Wallmann, K., et al. (2020). Permian–Triassic mass extinction pulses driven by major marine carbon cycle perturbations. Nat. Geosci. 13(11): 745-750. https://doi.org/10.1038/s41561-020-00646-4. |
| [44] | Pörtner, H.O., Langenbuch, M., and Michaelidis, B. (2005). Synergistic effects of temperature extremes, hypoxia, and increases in CO2 on marine animals: From Earth history to global change. J. Geophys. Res. Oceans 110: C09S10. https://doi.org/10.1029/2004JC002561. |
| [45] | Stumpp, M., Hu, M.Y., Melzner, F., et al. (2012). Acidified seawater impacts sea urchin larvae pH regulatory systems relevant for calcification. Proc. Natl. Acad. Sci. USA 109(44): 18192-18197. https://doi.org/10.1073/pnas.1209174109. |
| [46] | Pan, T. -C.F., Applebaum, S.L., and Manahan, D.T. (2015). Experimental ocean acidification alters the allocation of metabolic energy. Proc. Natl. Acad. Sci. USA 112(15): 4696-4701. https://doi.org/10.1073/pnas.1416967112. |
| [47] | Manwell, C. (1960). Oxygen equilibrium of brachiopod Lingula hemerythrin. Science 132(3426): 550-551. https://doi.org/10.1126/science.132.3426.550. |
| [48] | Isozaki, Y. (1997). Permo-Triassic boundary superanoxia and stratified superocean: records from lost deep sea. Science 276: 235-238. https://doi.org/10.1126/science.276.5310.235. |
| [49] | Racki, G. (1999). Silica-secreting biota and mass extinctions: survival patterns and processes. Palaeogeogr. Palaeoclimatol. Palaeoecol. 154(1–2): 107-132. https://doi.org/10.1016/S0031-0182(99)00089-9. |
| [50] | Bagarinao, T. (1992). Sulfide as an environmental factor and toxicant: tolerance and adaptations in aquatic organisms. Aquat. Toxicol. 24(1–2): 21-62. https://doi.org/10.1016/0166-445X(92)90015-F. |
| [51] | Montes-Rodríguez, I.M., Rivera, L.E., López-Garriga, J., et al. (2016). Characterization and expression of the Lucina pectinata oxygen and sulfide binding hemoglobin genes. PLoS One 11(1): e0147977. https://doi.org/10.1371/journal.pone.0147977. |
| [52] | Martín-Durán, J.M., de Mendoza, A., Sebé-Pedrós, A., et al. (2013). A broad genomic survey reveals multiple origins and frequent losses in the evolution of respiratory hemerythrins and hemocyanins. Genome Biol. Evol. 5(7): 1435-1442. https://doi.org/10.1093/gbe/evt102. |
| [53] | Pushie, M.J., Pratt, B.R., Macdonald, T.C., et al. (2014). Evidence for biogenic copper (hemocyanin) in the middle Cambrian arthropod Marrella from the Burgess Shale. Palaios 29(10): 512-524. https://doi.org/10.2110/palo.2014.073. |
| [54] | Sun, Y., Joachimski, M.M., Wignall, P.B., et al. (2012). Lethally hot temperatures during the Early Triassic greenhouse. Science 338(6105): 366-370. https://doi.org/10.1126/science.1224126. |
| [55] | Wignall, P.B., and Twitchett, R.J. (1996). Oceanic anoxia and the end-Permian mass extinction. Science 272(5265): 1155-1158. https://doi.org/10.1126/science.272.5265.1155. |
| [56] | Yin, H., and Song, H. (2013). Mass extinction and Pangea integration during the Paleozoic-Mesozoic transition. Sci. China Earth Sci. 56(11): 1791-1803. https://doi.org/10.1007/s11430-013-4624-3. |
| [57] | Bond, D.P., and Grasby, S.E. (2017). On the causes of mass extinctions. Palaeogeogr. Palaeoclimatol. Palaeoecol. 478: 3-29. https://doi.org/10.1016/j.palaeo.2016.11.005. |
| [58] | Zhang, H., Zhang, F., Chen, J. -b., et al. (2021). Felsic volcanism as a factor driving the end-Permian mass extinction. Sci. Adv. 7(47): eabh1390. https://doi.org/10.1126/sciadv.abh1390. |
| [59] | Benton, M.J. (2018). Hyperthermal-driven mass extinctions: killing models during the Permian–Triassic mass extinction. Phil. Trans. R. Soc. A. 376(2130): 20170076. https://doi.org/10.1098/rsta.2017.0076. |
| [60] | Song, H., and Scotese, C.R. (2023). The end-Paleozoic great warming. Sci. Bull. 68(21): 2523-2526. https://doi.org/10.1016/j.scib.2023.09.009. |
| [61] | McGhee, G. (2018). Carboniferous Giants and Mass Extinction. In Carboniferous Giants and Mass Extinction, (Columbia University Press). |
| [62] | Hönisch, B., Ridgwell, A., Schmidt, D.N., et al. (2012). The geological record of ocean acidification. Science 335(6072): 1058-1063. https://doi.org/10.1126/science.1208277. |
| [63] | Alroy, J. (2010). The shifting balance of diversity among major marine animal groups. Science 329(5996): 1191-1194. https://doi.org/10.1126/science.1189910. |
| [64] | Ceballos, G., Ehrlich, P.R., Barnosky, A.D., et al. (2015). Accelerated modern human–induced species losses: Entering the sixth mass extinction. Sci. Adv. 1(5): e1400253. https://doi.org/10.1126/sciadv.1400253. |
| [65] | Wang, F., Harindintwali, J.D., Wei, K., et al. (2023). Climate change: Strategies for mitigation and adaptation. The Innovation Geoscience 1(1): 100015. https://doi.org/10.59717/j.xinn-geo.2023.100015. |
| [66] | Yin, Z., Zhou, B., Duan, M., et al. (2023). Climate extremes become increasingly fierce in China. Innovation 4(2): 100406. https://doi.org/10.1016/j.xinn.2023.100406. |
| [67] | Penn, J.L., and Deutsch, C. (2022). Avoiding ocean mass extinction from climate warming. Science 376(6592): 524-526. |
| [68] | Kwiatkowski, L., Torres, O., Bopp, L., et al. (2020). Twenty-first century ocean warming, acidification, deoxygenation, and upper-ocean nutrient and primary production decline from CMIP6 model projections. Biogeosciences 17(13): 3439-3470. https://doi.org/10.5194/bg-17-3439-2020. |
| Song H., Wu Y., Dai X., et al., (2024). Respiratory protein-driven selectivity during the Permian-Triassic mass extinction. The Innovation 5(3), 100618. https://doi.org/10.1016/j.xinn.2024.100618 |
To request copyright permission to republish or share portions of our works, please visit Copyright Clearance Center's (CCC) Marketplace website at marketplace.copyright.com.
Extinction and body-size reduction of marine animals at the genus level during the Permian-Triassic mass extinction
Extinction and size reduction in different groups
Logistic regression shows the selectivity of extinction during the Permian-Triassic crisis
Ocean O2, H2S, and pH changes during the Permian-Triassic extinction using the cGENIE model and geological records