The Permian and Triassic were key time intervals in the history of life on Earth. Both periods are marked by a series of biotic crises including the most catastrophic of such events, the end-Permian mass extinction, which eventually led to a major turnover from typical Palaeozoic faunas and floras to those that are emblematic for the Mesozoic and Cenozoic. Here we review patterns in Permian-Triassic bony fishes, a group whose evolutionary dynamics are understudied. Based on data from primary literature, we analyse changes in their taxonomic diversity and body size (as a proxy for trophic position) and explore their response to Permian-Triassic events. Diversity and body size are investigated separately for different groups of Osteichthyes (Dipnoi, Actinistia, 'Palaeopterygii', 'Subholostei', Holostei, Teleosteomorpha), within the marine and freshwater realms and on a global scale (total diversity) as well as across palaeolatitudinal belts. Diversity is also measured for different palaeogeographical provinces. Our results suggest a general trend from low osteichthyan diversity in the Permian to higher levels in the Triassic. Diversity dynamics in the Permian are marked by a decline in freshwater taxa during the Cisuralian. An extinction event during the end-Guadalupian crisis is not evident from our data, but 'palaeopterygians' experienced a significant body size increase across the Guadalupian-Lopingian boundary and these fishes upheld their position as large, top predators from the Late Permian to the Late Triassic. Elevated turnover rates are documented at the Permian-Triassic boundary, and two distinct diversification events are noted in the wake of this biotic crisis, a first one during the Early Triassic (dipnoans, actinistians, 'palaeopterygians', 'subholosteans') and a second one during the Middle Triassic ('subholosteans', neopterygians). The origination of new, small taxa predominantly among these groups during the Middle Triassic event caused a significant reduction in osteichthyan body size. Neopterygii, the clade that encompasses the vast majority of extant fishes, underwent another diversification phase in the Late Triassic. The Triassic radiation of Osteichthyes, predominantly of Actinopterygii, which only occurred after severe extinctions among Chondrichthyes during the Middle-Late Permian, resulted in a profound change within global fish communities, from chondrichthyan-rich faunas of the Permo-Carboniferous to typical Mesozoic and Cenozoic associations dominated by actinopterygians. This turnover was not sudden but followed a stepwise pattern, with leaps during extinction events.
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Chondrichthyans are newly reported from the autochthonous Wordian Khuff Formation (middle Permian), cropping out in well-exposed, low-palaeolatitude sections in the interior Haushi-Huqf area of Oman. The shark remains comprise isolated teeth, dermal denticles and fin spines and have been recovered by processing limestone in buffered acetic acid from bulk rock samples. The fauna consists of mainly ctenacanthiform and hybodontiform taxa, identified as Glikmanius cf. myachkovensis, Glikmanius culmenis sp. nov., Omanoselache hendersoni gen. et sp. nov., Omanoselache angiolinii gen. et sp. nov., cf. Omanoselache sp., Reesodus underwoodi gen et sp. nov., Teresodus amplexus gen. et sp. nov., Gunnellodus bellistriatus, Khuffia lenis gen. et sp. nov., Khuffia prolixa gen. et sp. nov. and Euselachii sp. indet. Additional specimens include rare teeth of the lonchidiid cf. 'Palaeozoic Genus 1' sp., of the neoselachian Cooleyella cf. fordi and a further indeterminate neoselachian, of an indeterminate petalodont and of the holocephalan Deltodus aff. mercurei and Solenodus cf. crenulatus. Fin spines add a further two taxa, Nemacanthus sp. and Amelacanthus cf. sulcatus, which have neoselachian affinities and therefore an unclear relationship to the recovered teeth. The occurrence of Nemacanthus within this Wordian fauna represents the oldest record of this taxon and its only known occurrence in the Palaeozoic. Of the remaining genera, Glikmanius has previously been recorded from the Wordian, whereas for all the others, this study represents their youngest known stratigraphic occurrence and first occurrence in Guadalupian (middle Permian) strata. This adds significantly to our knowledge of the global diversity of chondrichthyans preceding the end-Guadalupian biotic crisis. Palaeogeographically, for all taxa, this study represents the first record from the western fringe of the marine Neotethyan basin, and only Cooleyella was previously known from the southern (Gondwanan) part of the Pangaean continental margin.
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