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. 2023 Feb;98(1):284-315.
doi: 10.1111/brv.12907. Epub 2022 Oct 3.

The early diversification of ray-finned fishes (Actinopterygii): hypotheses, challenges and future prospects

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The early diversification of ray-finned fishes (Actinopterygii): hypotheses, challenges and future prospects

Struan Henderson et al. Biol Rev Camb Philos Soc. 2023 Feb.

Abstract

Actinopterygii makes up half of living vertebrate diversity, and study of fossil members during their Palaeozoic rise to dominance has a long history of descriptive work. Although research interest into Palaeozoic actinopterygians has increased in recent years, broader patterns of diversity and diversity dynamics remain critically understudied. Past studies have investigated macroevolutionary trends in Palaeozoic actinopterygians in a piecemeal fashion, variably using existing compendia of vertebrates or literature-based searches. Here, we present a comprehensive occurrence-based dataset of actinopterygians spanning the whole of the Palaeozoic. We use this to produce the first through-Palaeozoic trends in genus and species counts for Actinopterygii. Diversity through time generally tracks metrics for sampling, while major taxonomic problems pervading the Palaeozoic actinopterygian record obscure diversity trends. Many described species are concentrated in several particularly problematic 'waste-basket' genera, hiding considerable morphological and taxonomic diversity. This taxonomic confusion also feeds into a limited understanding of phylogenetic relationships. A heavy sampling bias towards Europe and North America exists in both occurrence databases and available phylogenetic matrices, with other regions underrepresented despite yielding important data. Scrutiny of the extent to which spatial biases influence the actinopterygian record is lacking, as is research on other forms of bias. Low richness in some time periods may be linked to geological biases, while the effects of taphonomic biases on Palaeozoic actinopterygians have not yet been investigated. Efforts are already underway both to redescribe poorly defined taxa and to describe taxa from underrepresented regions, helping to address taxonomic issues and accuracy of occurrence data. New methods of sampling standardisation utilising up-to-date occurrence databases will be critical in teasing apart biological changes in diversity and those resulting from bias. Lastly, continued phylogenetic work will enable the use of phylogenetic comparative methods to elucidate the origins of actinopterygian biogeography and subsequent patterns of radiation throughout their rise to dominate aquatic faunas.

Keywords: Actinopterygii; Palaeozoic; diversity; fossils; ichthyology; sampling biases.

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Figures

Fig. 1
Fig. 1
Diversity of Palaeozoic fishes through time presented in previous studies. (A) Family‐level diversity curves of actinopterygians and non‐actinopterygian osteichthyans (Patterson, ; using data from Benton, 1993). (B) Family‐level diversity curves of actinopterygians and non‐actinopterygian fishes (Blieck, ; using data from Benton, 1993). (C) Genus‐level diversity of marine osteichthyans and non‐osteichthyan fishes, excluding conodonts (Friedman & Sallan, ; using data from Sepkoski, 2002). (D) Genus‐level diversity of British osteichthyans and non‐osteichthyan fishes (Lloyd & Friedman, 2013). (E) Genus‐level diversity of actinopterygians and non‐actinopterygian fishes (Thomson, ; using data from Romer, 1966). (F) Genus‐level diversity of actinopterygians and non‐actinopterygian fishes (Sallan & Coates, ; Romano et al., 2016).
Fig. 2
Fig. 2
Raw counts of Palaeozoic actinopterygian genera (black, solid line), collections (brown, short‐dashed line), formations (red, dotted line) and equal‐area grid cells (orange, long‐dashed line) entered in the Paleobiology Database (PBDB).
Fig. 3
Fig. 3
Geographic spread of actinopterygian occurrences entered in the Paleobiology Database (PBDB) for the (A) Devonian; (B) Carboniferous; and (C) Permian.
Fig. 4
Fig. 4
Raw counts of Palaeozoic actinopterygian (A) genera and (B) species (black, solid line) in roughly equal‐length intervals (see Table S2). Collections (brown, short‐dashed line), formations (red, dotted line) and equal‐area grid cells (orange, long‐dashed line) are also plotted.
Fig. 5
Fig. 5
Raw counts of Palaeozoic actinopterygian (A) genera and (B) species (black, solid line) in standard International Commission on Stratigraphy (ICS) stages. Collections (brown, short‐dashed line), formations (red, dotted line) and equal‐area grid cells (orange, long‐dashed line) are also plotted.
Fig. 6
Fig. 6
Geographic spread of actinopterygian occurrences in the (A) Devonian; (B) Carboniferous; and (C) Permian.
Fig. 7
Fig. 7
Distribution of the most speciose and widespread actinopterygian genera, with occurrences coloured according to the ICS colours for the period in which they occur (Devonian – brown; Carboniferous – green; Permian – red): (A) Acrolepis; (B) Amblypterus; (C) ‘Elonichthys’; (D) Moythomasia; (E) Palaeoniscum; (F) Platysomus; and (G) Rhadinichthys.
Fig. 8
Fig. 8
Collector's curve of the global Palaeozoic actinopterygian fossil record, divided by present‐day geographic region.
Fig. 9
Fig. 9
Network depicting the flow of characters to and from phylogenetic analyses of Palaeozoic actinopterygians. Nodes are coloured according to the geographic region in which authors' listed institutions (in the primary article) are located.
Fig. 10
Fig. 10
Phylogenetic analyses of Palaeozoic actinopterygian relationships showing the geographic distribution of sampled taxa: (A) Dietze (2000); (B) Mickle et al. (2009); (C) Figueroa et al. (2019); and (D) Elliott (2016). The actinopterygian crown node is indicated in analyses that include extant taxa. Asterisks (*) indicate taxa not included in the other depicted analyses to show the lack of overlap between datasets.

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