Abstract
Current hypotheses of early tetrapod evolution posit close ecological and biogeographic ties to the extensive coal-producing wetlands of the Carboniferous palaeoequator with rapid replacement of archaic tetrapod groups by relatives of modern amniotes and lissamphibians in the late Carboniferous (about 307 million years ago). These hypotheses draw on a tetrapod fossil record that is almost entirely restricted to palaeoequatorial Pangea (Laurussia)1,2. Here we describe a new giant stem tetrapod, Gaiasia jennyae, from high-palaeolatitude (about 55° S) early Permian-aged (about 280 million years ago) deposits in Namibia that challenges this scenario. Gaiasia is represented by several large, semi-articulated skeletons characterized by a weakly ossified skull with a loosely articulated palate dominated by a broad diamond-shaped parasphenoid, a posteriorly projecting occiput, and enlarged, interlocking dentary and coronoid fangs. Phylogenetic analysis resolves Gaiasia within the tetrapod stem group as the sister taxon of the Carboniferous Colosteidae from Euramerica. Gaiasia is larger than all previously described digited stem tetrapods and provides evidence that continental tetrapods were well established in the cold-temperate latitudes of Gondwana during the final phases of the Carboniferous–Permian deglaciation. This points to a more global distribution of continental tetrapods during the Carboniferous–Permian transition and indicates that previous hypotheses of global tetrapod faunal turnover and dispersal at this time2,3 must be reconsidered.
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Data availability
The authors declare that all the data supporting the findings of this study are available within the paper and the Supplementary Information. All nomenclatural acts are registered with ZooBank (https://zoobank.org) under the Life Science Identifier urn:lsid:zoobank.org:pub:62D1B947-D36F-49D2-8490-EAD49C46A22B.
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Acknowledgements
We thank the National Heritage Council and the Geological Survey of Namibia for fieldwork permits and loan of specimens; R. Swart, A. Wanke, F. Abdala and P. October for logistical and field support; S. Mtungata, who found and expertly prepared the type specimen; and G. Lio for the artistic reconstruction of Gaiasia. The 2014, 2015 and 2019 expeditions were supported by National Geographic Society Research Grant 9360-13 and PAST. Research was also possible thanks to the support of the Agencia Nacional de Promoción Científica y Tecnológica (Argentina, PICT 2019-01127 to C.A.M.) and the Universidad de Buenos Aires and CONICET (to C.A.M. and L.C.G.). J.D.P. was supported by Banting Postdoctoral Fellowship 473316. This study is contribution R-346 by C.A.M. and L.C.G. to the Instituto de Estudios Andinos Don Pablo Groeber (IDEAN, UBA-CONICET).
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The project was conceived by C.A.M., R.M.H.S. and J.D.P.; phylogenetic analysis was carried out by C.A.M. and J.D.P.; R.M.H.S., C.A.M., A.C.M., L.C.G. and H.M. conducted fieldwork to collect specimens and geological context. C.A.M., J.D.P. and R.M.H.S. wrote the manuscript, with comments from all authors.
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Extended data figures and tables
Extended Data Fig. 1 Gaiasia jennyae gen. et sp. nov. (F-1528).
a,b, skull in posterior view. a, Photograph. b, Interpretative drawing. Scale bar, 50 mm. et, excavatio tympanica; exo, exoccipital; occf, occipital artery foramen; op, opisthotic; par, prearticular; psph, parasphenoid; pt, pterygoid; q, quadrate; qj, quadratojugal; sa, surangular;?socc, supraoccipital; X, XII, cranial nerves X and XII, respectively.
Extended Data Fig. 2 Gaiasia jennyae gen. et sp. nov. (F-1528).
a,b, Right lateral view of the posterior part of the skull and anterior vertebrae during preparation of the specimen. a, Photograph. b, Interpretative drawing. c,d, Posterior view of the left side of the skull. c, Photograph. d, Interpretative drawing. Scale bar, 50 mm. exo, exoccipital; Md, mandible; occf, occipital artery foramen; op, opisthotic; psph, parasphenoid; pt, pterygoid; q, quadrate; qj, quadratojugal;?socc, supraoccipital; sq, squamosal; X, XII, cranial nerves X and XII, respectively. Yellow shade, spiracular cleft; red shade, depressor mandibulae; cross-hatching, matrix.
Extended Data Fig. 3 Gaiasia jennyae gen. et sp. nov. (F-1528).
a–d, Composite reconstruction of the mandible. a, Dorsal. b, Ventral. c, Labial. d, Lingual. e, f. Photographs of the right hemimandible. e, Ventral view of the posterior half. f, Dorsal view of the symphyseal area. adsym, adsymphysial plate; an, angular; anf, angular fenestra; c1, anterior coronoid; c2, middle coronoid; chf, chordatympanic foramen; d, dentary; par, prearticular; pospl, postsplenial; sa, surangular; spl, splenial. Dotted white circles show the position of the symphysial fangs. Scale bar, 50 mm.
Extended Data Fig. 4 Gaiasia jennyae gen. et sp. nov. (F-1528).
Ventral branchial element in a, Dorsal and b, Ventral views. Anterior to the left. Scale bar, 50 mm.
Extended Data Fig. 5 Gaiasia jennyae gen. et sp. nov. (F-1528).
Articulated axial skeleton. a, Articulated column in right lateral view. Scale bar, 100 mm. b, Detail of the block just behind the star in a. Scale bar, 50 mm. c, Same block as in b in anterior view showing a pleurocentrum. Scale bar, 50 mm. Light green, pleurocentrum; yellow, intercentrum; dark green, neural arch; brown, rib.
Extended Data Fig. 6 Gaiasia jennyae gen. et sp. nov. (F-1528).
a–b, Composite reconstruction of the skull. a, Dorsal. b, Ventral. c, Artistic reconstruction of Gaiasia in lateral view; artwork by Gabriel Lio. ect, ectopterygoid; exo, exoccipital; f, frontal; it, intertemporal; j, jugal; l, lacrimal; mx, maxilla; n, nasal; op, opisthotic; p, parietal; pfr, prefrontal; pl, palatine; pmx, premaxilla; po, postorbital; pof, postfrontal; pp, postparietal; psph, parasphenoid; pt, pterygoid; qj, quadratojugal; socc?, supraoccipital; sq, squamosal; st, supratemporal; t, tabular; v, vomer. Scale bar, 50 mm.
Extended Data Fig. 7 Phylogenetic analysis.
a, Unweighted strict-consensus tree. b—f, Strict consensus of trees recovered using implying weights49. Implied weights inferred with b, k = 2, c, k = 4, d, k = 6, e, k = 8; f, k = 10.
Supplementary information
Supplementary Information
Supplementary Fig. 1 (anatomical figure of paratype specimen), Data including Figs. 2–7 (geological setting and taphonomic information, character list and phylogenetic discussion), Tables 1 and 2 and References.
Supplementary Data
The new phylogenetic dataset generated during the current study.
Supplementary Data
Clack et al. 2019 data matrix.
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Marsicano, C.A., Pardo, J.D., Smith, R.M.H. et al. Giant stem tetrapod was apex predator in Gondwanan late Palaeozoic ice age. Nature 631, 577–582 (2024). https://doi.org/10.1038/s41586-024-07572-0
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DOI: https://doi.org/10.1038/s41586-024-07572-0
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