Temnospondyls from the Beaufort Group (Karoo Basin) of South Africa and Their Biostratigraphy

doi:10.1016/S1342-937X(05)70315-4

Gondwana Research

Volume 7, Issue 1, January 2004, Pages 165-173

https://doi.org/10.1016/S1342-937X(05)70315-4 Get rights and content

Abstract

The Permo-Triassic Beaufort Group (Karoo Basin) of South Africa is biostratigraphically subdivided into eight, temporally successive assemblage zones based on therapsids (‘mammal-like reptiles’). The Temnospondyli, fossil tetrapods usually regarded as extinct amphibians, are second only to therapsids in terms of diversity and abundance in these strata, with nine higher-level taxa (‘families’) known. Temnospondyls are also playing an increasingly important role in biostratigraphy and correlation of the Beaufort strata. The lower Beaufort Group (Late Permian) contains six of the eight biozones, but only one temnospondyl ‘family’, the Rhinesuchidae, whose record in the Karoo is the richest in the world. However, rhinesuchid taxonomy remains in flux and the group is thus of limited biostratigraphic utility. The Early Triassic Lystrosaurus Assemblage Zone (middle Beaufort Group) contains the Rhinesuchidae, Amphibamidae, Lydekkerinidae, Tupilakosauridae, Rhytidosteidae, Mastodonsauridae and Trematosauridae, although the biostratigraphy of temnospondyls within this biozone is poorly constrained. The uppermost reaches of the Lystrosaurus biozone contain a paucity of fossils but includes ‘Kestrosaurus’ (Mastodonsauridae) and ?Trematosuchus (Trematosauridae), taxa previously thought to pertain to the lower part of the overlying Cynognathus biozone. The late Early to Middle Triassic Cynognathus Assemblage Zone (upper Beaufort Group) hosts the Mastodonsauridae, Trematosauridae, Brachyopidae, Laidleriidae and, possibly, the Rhytidosteidae. Based largely on the spatial and temporal distribution of mastodonsaurids, this biozone has been biostratigraphically subdivided into a lower A, middle B and upper C subzones, characterised by differing ages and faunas.

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Stepping out across the Karoo retro-foreland basin: Improved constraints on the Ecca-Beaufort shoreline along the northern margin
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Citation Excerpt :
Of the few tetrapod fossils recovered near to the Ecca-Beaufort contact between Brandfort and Kroonstad in the central portion of the study area, the Uranocentrodon specimen BP/1/8195 is stratigraphically most useful. This is because the stratigraphic range of Uranocentrodon is restricted to the Daptocephalus AZ (Latimer et al., 2002; Damiani, 2004; Tarailo, 2018). However, most of the known Uranocentrodon specimens were collected from a single locality near the town Senekal in the Free State Province where the Permo-Triassic boundary is considered to be absent and a hiatus of unknown duration exists between the Daptocephalus AZ and overlying Lystrosaurus declivis AZ (Hancox et al., 2002; Latimer et al., 2002).
The Karoo Supergroup of South Africa, internationally renowned for its almost continuous Carboniferous to Jurassic record of deposition in a foreland basin setting, hosts an unparalleled record of fossil tetrapods that provides unique insight into terrestrial biodiversity change over this extended period. Understanding of the litho-, bio- and chronostratigraphy of the Karoo Basin has greatly improved in recent years but most work has focused on the thicker and better-exposed rocks in the basinal foredeep. A core issue that has remained ambiguous is the stratigraphic placement of the diachronous Ecca-Beaufort contact, for which different criteria have been used during mapping in different parts of the main Karoo Basin. Comparison of biostratigraphy and contrasting lithological facies changes with those in the better-studied foredeep enabled us to revise the lithostratigraphy of the Ecca and Beaufort groups in the distal part of the main Karoo Basin. The presence of proximal marine facies associations in the distal sector conform with the definition of the Waterford Formation of the Ecca Group, so far recognised only in the southern foredeep. The upper contact of the Waterford Formation represents a change from a subaqueous to a subaerial delta plain depositional environment and marks the boundary between the Ecca and Beaufort groups, as in the south. This has consequences for Beaufort Group stratigraphic subdivision in the distal sector of the Karoo foreland basin.
Early Triassic terrestrial tetrapod fauna: a review
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Citation Excerpt :
The fauna, considering the broader taxonomic category for the abundance matrix, is dominated by Temnospondyli (59%), followed by Archosauromorpha (29.2%), Procolophonomorpha (7.9%), Chroniosuchia (2.4%), Therapsida (0.8%), non-temnospondyl Amphibia (0.4), and Lepidosauromorpha (0.3%). Early Triassic terrestrial tetrapods from South Africa are known from a huge collection of body fossils and subordinate ichnological material (Huxley, 1859; Owen, 1860, 1884; Seeley, 1878, 1894, 1905; Broom, 1903a, 1903b, 1903c, 1905, 1915, 1930, 1936, 1937, 1941, 1946; Broili, 1908; Watson, 1913, 1962; Haughton, 1915, 1924, 1925, 1965; Van Hoepen, 1915, 1916; Broili and Schröder, 1934, 1937; Broom and Robinson, 1948; Parrington, 1948; Crompton, 1955; Brink, 1958, 1959, 1965; Estes, 1961; Sigogneau, 1963; Hoffman, 1965; Carroll, 1976, 1982; Kitching, 1968, 1977; Cruickshank, 1972, 1975; Mendrez, 1972; Cluver, 1974; Colbert and Kitching, 1977; Cosgriff and Zawiskie, 1979; Carroll and Lindsay, 1985; Groenewald, 1991; Shishkin et al., 1996, 2004; Warren, 1998; Welman, 1998; Gow, 1999; Neveling et al., 1999; Damiani et al., 2000, 2001, 2003; Hancox, 2000; Shishkin and Rubidge, 2000; Damiani and Welman, 2001; Groenewald et al., 2001; Modesto et al., 2001, 2002, 2003, 2010; Damiani and Jeannot, 2002; Reisz and Scott, 2002; Bender and Hancox, 2004; Damiani, 2004; Modesto and Sues, 2004; Sidor and Smith, 2004; Schoch and Rubidge, 2005; Abdala et al., 2006, 2014; Abdala, 2007; Botha et al., 2007; Fourie and Rubidge, 2007; Cisneros, 2008; Modesto and Botha-Brink, 2008; Botha-Brink and Modesto, 2011; Botha-Brink and Smith, 2011; Gaetano et al., 2012; Novikov, 2012a; Sigurdsen et al., 2012; Huttenlocker and Botha-Brink, 2013; Gower et al., 2014; Ezcurra and Butler, 2015; Jasinoski and Abdala, 2017; Butler et al., 2019). The South African Karoo Basin represents one of the few stratigraphic sequences in the world that preserves a complete record of the terrestrial PTME (Smith and Botha-Brink, 2014; MacLeod et al., 2017; Viglietti et al., 2018) and subsequent recovery in the Early Triassic (Rubidge et al., 2016).
The Permian-Triassic mass extinction (PTME, ca. 252 Mya) was one of the most severe biotic crises of the Phanerozoic, eliminating >90% of marine and terrestrial species. This was followed by a long period of recovery in the Early and Middle Triassic which revolutionised the structure of both marine and terrestrial ecosystems, triggering the new ecosystem structure of the Mesozoic and Cenozoic. Entire new clades emerged after the mass extinction, including decapods and marine reptiles in the oceans and new tetrapods on land. In both marine and terrestrial ecosystems, the recovery is interpreted as stepwise and slow, from a combination of continuing environmental perturbations and complex multilevel interaction between species in the new environments as ecosystems reconstructed themselves. Here, we present a review of Early Triassic terrestrial tetrapod faunas, geological formations and outcrops around the world, and provide a semi-quantitative analysis of a data set of Early Triassic terrestrial tetrapods. We identify a marked regionalisation of Early Triassic terrestrial tetrapods, with faunas varying in both taxonomic composition and relative abundance according to palaeolatitudinal belt. We reject the alleged uniformity of faunas around Pangaea suggested in the literature as a result of the hot-house climate. In addition, we can restrict the “tetrapod gap” of terrestrial life in the Early Triassic to palaeolatitudes between 15°N and about 31°S, in contrast to the earlier suggestion of total absence of tetrapod taxa between 30°N and 40°S. There was fairly strong provincialism following the PTME, according to cluster analysis of a taxon presence matrix, entirely consistent with Early Triassic palaeobiogeography. Unexpectedly, the overall pattern for Early Triassic terrestrial tetrapod faunas largely reflects that of the Late Permian, suggesting that the recovery faunas in the Early Triassic retained some kind of imprint of tetrapod distributions according to palaeogeography and palaeoclimate, despite the near-total extinction of life through the PTME.
An Early Triassic polar predator ecosystem revealed by vertebrate coprolites from the Bulgo Sandstone (Sydney Basin) of southeastern Australia
2016, Palaeogeography, Palaeoclimatology, Palaeoecology
Citation Excerpt :
Indeed, our coprolite data compliments recent discoveries of new skeletal material, which evidences a range of mid-large sized temnospondyl taxa, together with novel archosauromorphs and actinopterygian fish (Greco et al., 2014). Collectively, therefore, the Bulgo Sandstone fauna parallels those from other Australian Early Triassic deposits, especially the Arcadia and Knocklofty formations, which likewise produce a predominance of aquatic tetrapods (e.g. temnospondyls and proterosuchians: Camp and Banks, 1978; Thulborn, 1979, 1986b; Warren, 1980; Warren and Black, 1985; Damiani and Warren, 1996; Warren et al., 2006; Damiani, 1999; Yates, 1999, 2000) and non-marine bony fish (e.g. saurichthyids and dipnoans: Dziewa, 1980; Northwood, 1999; Kemp, 1996; Bender, 2008) coherent with fluviatile assemblages found elsewhere in Gondwana (e.g. Warren et al., 2001; Bender and Hancox, 2004; Damiani, 2004). In fact, the Bulgo Sandstone palaeoenvironment can be very precisely reconstructed via the REE compositions of its preserved coprolite material (Fig. 7A).
Vertebrate trace fossils often provide a measure of cryptic biodiversity, and are especially pertinent when skeletal remnants are exceptionally rare. The Lower Triassic (lower Olenekian) Bulgo Sandstone at Long Reef in the Sydney Basin of southeastern Australia constitutes just such a deposit, having yielded isolated bones of giant capitosaurian temnospondyls and proterosuchid archosauriforms, together with abundant coprolites that are geochemically rich in elemental phosphate and carbon denoting vertebrate predators. Microstructural analysis of these preserved droppings reveals occasional bone fragments, fish scales, insect cuticles, plant material and bacterial traces (pseudomorph voids), as well as silicate mineral particles. REE concentrations indicate that burial and early diagenesis occurred explicitly within fluvial sediments. Furthermore, external morphological characterization permits attribution of spiral coprolites to chondrichthyan or osteichthyan fishes, polygonal, ovoid–spherical and typically flattened feces to temnospondyls, and conspicuously large cylindrical droppings to archosauriforms or other amniote apex predators. Collectively, the Bulgo Sandstone coprolite assemblage thus offers new insights into ecosystem structure and palaeoenvironment in what was an earliest Triassic near polar setting. Such data compliments the documented skeletal record, but indicates a greater range of aquatic and possibly terrestrial carnivores — the latter being enigmatically sparse in the Australian Triassic and yet detected here via the hitherto underexplored trace fossil evidence of their ecological presence.
The Daptocephalus Assemblage Zone (Lopingian), South Africa: A proposed biostratigraphy based on a new compilation of stratigraphic ranges
2016, Journal of African Earth Sciences
Citation Excerpt :
Two species of temnospondyl amphibians (Rhinesuchus muchos and Laccocephalus insperatus) were known from the DaAZ at the time of the revision of the Karoo biostratigraphy by Rubidge et al. (1995). Subsequent taxonomic revisions (Damiani and Rubidge, 2003; Latimer et al., 2002; Marsicano et al., 2015) have revealed an updated list of DaAZ temnospondyls (including Laccocephalus watsoni, Rhinesuchus capensis, Rhinesuchus waitsi, and Uranocentrodon senekalensis) but the collections database has not yet been updated to reflect these changes. Therefore, the higher taxon Rhinesuchidae was used to represent all late Permian temnospondyls in this study (Fig. 2, Table 1).
The Dicynodon Assemblage Zone (DiAZ) of South Africa's Karoo Basin is one of the eight biostratigraphic zones of the Beaufort Group. It spans the uppermost Permian strata (Balfour, Teekloof, and Normandien formations) and traditionally has been considered to terminate with the disappearance of Dicynodon lacerticeps at the Permo-Triassic Boundary. We demonstrate that the three index fossils currently used to define the Dicynodon Assemblage Zone (Dicynodon lacerticeps, Theriognathus microps, and Procynosuchus delaharpeae) have first appearance datums (FADs) below its traditionally recognized lower boundary and have ranges mostly restricted to the lower portion of the biozone, well below the Permo-Triassic Boundary. We propose re-establishing Daptocephalus leoniceps as an index fossil for this stratigraphic interval, and reinstating the name Daptocephalus Assemblage Zone (DaAZ) for this unit. Furthermore, the FAD of Lystrosaurus maccaigi in the uppermost reaches of the biozone calls for the establishment of a two-fold subdivision of the current Dicynodon Assemblage Zone. The biostratigraphic utility of Da. leoniceps and other South African dicynodontoids outside of the Karoo Basin is limited due to basinal endemism at the species level and varying temporal ranges of dicynodontoids globally. Therefore, we recommend their use only for correlation within the Karoo Basin at this time. Revision of the stratigraphic ranges of all late Permian tetrapods does not reveal a significant change in faunal diversity between the lower and upper DaAZ. However, the last appearance datums of the abundant taxa Di. lacerticeps, T. microps, P. delaharpeae, and Diictodon feliceps occur below the three extinction phases associated with the end-Permian mass extinction event. Due to northward attenuation of the strata, however, the stratigraphic position of the extinction phases may need to be reconsidered.
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Temnospondyls from the Beaufort Group (Karoo Basin) of South Africa and Their Biostratigraphy

Abstract

Zool. J. Linn. Soc.

Palaeogeogr., Palaeoclimatol., Palaeoecol.

J. Afri. Earth Sci.

Palaeogeogr., Palaeoclim., Palaeoecol.

Zool. J. Linn. Soc.

Zool. J. Linn. Soc.

Beobachtungen an Wirbeltieren der Karrooformation

XXVII. Über einen Capitosauriden aus der Cynognathus-Zone. Sitz.-Ber. bayer. Akad. Wiss. München

On a new stegocephalian (Batrachosuchus browni) from the Karroo beds of Aliwal North South Africa

Geol. Mag.

On the Permian and Triassic faunas of South Africa

Geol. Mag.

On a new labyrinthodont Rhinesuchus whaitsi, from the Permian beds of South Africa

Ann. S. Afr. Mus.

Notice of some new South African fossil amphibians and reptiles

Ann. S. Afr. Mus.

On the Triassic stegocephalians Brachyops, Bothriceps, and Lydekkerina gen. nov.

Proc. Zool. Soc. Lond.

Notes on some labyrinthodonts in the Transvaal Museum

Ann. Transvaal Mus.

Reciprocal flexural behaviour and contrasting stratigraphies: a new basin development model for the Karoo retroarc foreland system, South Africa

Basin Res.

Three capitosaurs from the Triassic of South Africa: Parotosuchus africanus (Broom 1909); Kestrosaurus dreyeri Haughton 1925, and Parotosuchus dirus sp. nov.

Palaeont. afr.

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J. Vert. Paleontol.

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Palaeont. Afri.

Parotosuchus (Amphibia, Temnospondyli) in Gondwana: biostratigraphic and palaeobiogeographic implications.

S. Afri. J. Sci.

Parotosuchus (Amphibia, Temnospondyli) from the Cynognathus Assemblage Zone (Early Triassic) of South Africa: cranial morphology and relationships.

Alcheringa

New mastodonsaurid temnospondyls from the Cynognathus Assemblage Zone (Upper Beaufort Group; Karoo Basin) of South Africa

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First record of a mastodonsaurid (Temnospondyli, Stereospondyli) from the Early Triassic Lystrosaurus Assemblage Zone (Karoo Basin) of South Africa

N. Jb. Geol. Paläont. Abh.

First trematosaurid temnospondyl from the Lystrosaurus Assemblage Zone of South Africa and its biostratigraphic implications.

Geol. Mag.

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Paleoecology and phylogeny of the Trematosauridae

Geophys. Monogr. Ser.

The continental Triassic of South Africa

Zbl. Geol. Paläont. Teil I

The role of fossils in interpreting the development of the Karoo Basin

Palaeont. Afri.

First occurrence of Paracyclotosaurus (Temnospondyli, Capitosauridae) in the Karoo Basin of South Africa and its biostratigraphic implications

S. Afri. J. Sci.