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Correction for Song Et Al., Flat Latitudinal Diversity Gradient Caused Correction EARTH, ATMOSPHERIC, AND PLANETARY SCIENCES, EVOLUTION Correction for “Flat latitudinal diversity gradient caused by the Permian–Triassic mass extinction,” by Haijun Song, Shan Huang, Enhao Jia, Xu Dai, Paul B. Wignall, and Alexander M. Dunhill, which was first published July 6, 2020; 10.1073/pnas.1918953117 (Proc. Natl. Acad. Sci. U.S.A. 117, 17578–17583). The editors wish to note that the following competing interest statement regarding personal associations with the authors should have been disclosed: Editor M.J.B. shares a research grant with P.B.W. and has coauthored papers with A.M.D. in 2019 and 2020, and with H.S. in 2017 and 2019. The editors apologize for the oversight in failing to disclose this information to readers. The competing interest statement in the article has been corrected. Published under the PNAS license. First published August 10, 2020. www.pnas.org/cgi/doi/10.1073/pnas.2015344117 20334 | PNAS | August 18, 2020 | vol. 117 | no. 33 www.pnas.org Downloaded by guest on September 28, 2021 Flat latitudinal diversity gradient caused by the Permian–Triassic mass extinction Haijun Song (宋海军)a,1, Shan Huangb, Enhao Jia (贾恩豪)a, Xu Dai (代旭)a, Paul B. Wignallc, and Alexander M. Dunhillc aState Key Laboratory of Biogeology and Environmental Geology, School of Earth Sciences, China University of Geosciences, 430074 Wuhan, China; bSenckenberg Biodiversity and Climate Research Center, 60325 Frankfurt am Main, Germany; and cSchool of Earth and Environment, University of Leeds, LS2 9JT Leeds, United Kingdom Edited by Michael J. Benton, University of Bristol, Bristol, UK, and accepted by Editorial Board Member Neil H. Shubin May 21, 2020 (received for review October 29, 2019) The latitudinal diversity gradient (LDG) is recognized as one of the of marine species (16) and caused profound temporary and most pervasive, global patterns of present-day biodiversity. How- permanent ecological restructuring of marine ecosystems (23), ever, the controlling mechanisms have proved difficult to identify which ultimately catalyzed the transformation of marine com- because many potential drivers covary in space. The geological munities dominated by Paleozoic faunas to those dominated by record presents a unique opportunity for understanding the mecha- the Modern fauna (15). The effect of the largest mass extinction nisms which drive the LDG by providing a direct window to deep-time in Earth’s history on global marine biogeography is largely un- biogeographic dynamics. Here we used a comprehensive database known, but the rich marine fossil record from this time can be a containing 52,318 occurrences of marine fossils to show that the shape powerful tool for illuminating the fundamental principles that of the LDG changed greatly during the Permian–Triassic mass extinc- shape global biodiversity. tion from showing a significant tropical peak to a flattened LDG. The LDG dynamics across the P-Tr mass extinction has received flat LDG lasted for the entire Early Triassic (∼5 My) before reverting to little attention except for a few case studies that have all sug- a modern-like shape in the Middle Triassic. The environmental ex- gested a strong impact of environmental changes on global di- tremes that prevailed globally, especially the dramatic warming, likely versity patterns. For example, the early-middle Permian diversity induced selective extinction in low latitudes and accumulation of di- of terrestrial tetrapods reportedly peaked in temperate regions versity in high latitudes through origination and poleward migration, as a result of profound climate-induced biome shift (24). By the EARTH, ATMOSPHERIC, AND PLANETARY SCIENCES which combined together account for the flat LDG of the Early Triassic. Early Triassic, the distribution of terrestrial tetrapods had moved 10 to 15° poleward (25, 26), and the group was apparently absent biogeography | end-Permian mass extinction | global warming | ocean from 40 °S to 30 °N as a result of tropical overheating (18). anoxia | biodiversity Phylogenetic network analysis also found a marked increase in biogeographic connectedness, resulting in a more homogeneous he increase in species richness from the poles to the tropics, composition of diversity across latitudes in tetrapods during this long known as the latitudinal diversity gradient (LDG), is one time (27). Similar poleward migrations were also found in ma- T EVOLUTION of the most pervasive first-order biological patterns on Earth rine invertebrates in the Northern Hemisphere during the Early today (1, 2), both on land (3) and in the oceans (4–6). Yet, the Triassic (28), and a variable LDG during this time was reported relative importance of the diverse ecological and evolutionary mechanisms (e.g., reviews in refs. 7–9) for generating this pattern Significance remains unclear. Paleontological data provide a unique per- spective in the search for the dominant driver(s) of LDGs, The deep-time dynamics of the latitudinal diversity gradient allowing diversity and distribution dynamics to be tracked (LDG), especially through dramatic events like mass extinctions, through the long history of environmental changes (10, 11). In can provide invaluable insights into the biotic responses to particular, climate (e.g., temperature or precipitation) is regar- global changes, yet they remain largely underexplored. Our ded as a key driver of the LDG, and its large-scale changes have study shows that the shape of marine LDGs changed sub- been postulated to have altered the general shape of the LDG stantially and rapidly during the Permian–Triassic mass ex- through time (12, 13). Steep, normal LDGs (i.e., with a signifi- tinction from a modern-like steep LDG to a flat LDG. The flat cant tropical peak like today) have been found primarily during LDG lasted for ∼5 My and was likely a consequence of the icehouse times, whereas bimodal or even reverse LDGs with extreme global environment, including extreme warming and diversity peaks at mid to high latitudes occurred during green- ocean anoxia, which ensured harsh conditions prevailing from house climates (13, 14). These findings call for assessments of the tropics to the poles. Our findings highlight the fundamental the relative importance of climate per se and environmental role of environmental variations in concert with severe bio- stability, especially through comparing the dynamics across sev- diversity loss in shaping the first-order biogeographic patterns. eral time intervals with different environmental templates. The dramatic changes in environmental conditions and the Author contributions: H.S., P.B.W., and A.M.D. designed research; H.S., E.J., and X.D. performed research; H.S., S.H., and X.D. analyzed data; and H.S., S.H., P.B.W., and severe mass extinction at the end of the Permian provide an A.M.D. wrote the paper. excellent opportunity for investigating LDGs and their control- The authors declare no competing interest. – ling mechanisms. The Permian Triassic (P-Tr) mass extinction, This article is a PNAS Direct Submission. M.J.B. is a guest editor invited by the which occurred ca. 252 My ago, was the largest extinction event Editorial Board. of the Phanerozoic (15, 16). This biological crisis was linked to Published under the PNAS license. extreme and prolonged environmental changes, many of which Data deposition: All data used to conduct analyses and plot figures are available for are probably the most serious of the past 500 My. The contem- download at Dryad, https://datadryad.org/stash/dataset/doi:10.5061/dryad.41ns1rn9z. poraneous eruption of the Siberian Traps large igneous province 1To whom correspondence may be addressed. Email: [email protected]. (17) drove ∼10 °C global warming within ca. 30,000 to 60,000 y This article contains supporting information online at https://www.pnas.org/lookup/suppl/ through greenhouse gas emissions (18, 19) and widespread doi:10.1073/pnas.1918953117/-/DCSupplemental. oceanic anoxia (20–22). These disastrous events killed over 90% www.pnas.org/cgi/doi/10.1073/pnas.1918953117 PNAS Latest Articles | 1of6 in ammonoids (29). These findings suggest a great change in 201.3 Rh biogeographic distribution during the P-Tr crisis, which would (Ma) have had a profound impact on LDG. In this study, we in- Late No vestigate LDG dynamics through the P-Tr mass extinction event, Triassic and the later recovery of biodiversity, using the most compre- hensive fossil database thus far for this time period. Ca 237 Global Changes in Biogeography La Middle In order to assess the effect of the P-Tr extinction and associated Triassic environmental extremes on latitudinal diversity patterns, we An 247.2 analyzed biogeographic distributions using a database consisting Sp Sm of occurrences of marine genera (including 20 major clades; see Early Methods for details) from the late Permian (Changhsingian, Triassic Di 254.1 Ma) to the Late Triassic (Rhaetian, 201.3 Ma). This da- Gr tabase is an update of an earlier P-Tr marine fossil database (23) 251.90 Late and includes 52,318 generic occurrences at a substage- or stage- Permian Ch level resolution from 1,768 literature sources (Dataset S1). 254.14 Among these, 7,752, 12,676, 13,456, and 18,634 occurrences are 0-15º 15-30º 30-45º 45-90º in the late Permian and Early, Middle, and Late Triassic, re- spectively. Based on reconstructed paleolatitudes, the collections were binned into four paleolatitudinal zones for each hemi- 46.4 65.5 84.6 103.7 122.8 sphere: 0 to 15°, 15 to 30°, 30 to 45°, and 45 to 90°. The larger size Subsampled diversity of the 45 to 90° bin was chosen to accommodate the relatively Fig. 2. Rarefied genus-level diversity trends related to latitude from the low sample density at higher latitudes in most time intervals. At late Permian to the end of the Triassic. Data are standardized by repeatedly the epoch level, data from the paleolatitudinal zones of the subsampling from a randomly generated set until reaching a quota of 136 Northern and Southern Hemispheres are analyzed separately occurrences in each time bin at each latitudinal interval (SI Appendix, Table (Fig.
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