P. John Hancox

University of the Witwatersrand

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Bruce S. Rubidge

University of the Witwatersrand

Christian Koeberl

University of Vienna

W. U. Reimold

Universidade de Brasília

D. Brandt

Medtronic (United States)

Jonah N. Choiniere

University of the Witwatersrand

Richard J. Butler

University of Birmingham

Pia A. Viglietti

University of the Witwatersrand

Johann Neveling

Council for Geoscience

Christian F. Kammerer

University of the Witwatersrand

Stephen Tooth

Aberystwyth University

Cooperative Institutions

University of the Witwatersrand

University of Cape Town

Iziko Museums of South Africa

Rice Institute

University of Pretoria

Consejo Nacional de Investigaciones Científicas y Técnicas

University of Washington

University of Johannesburg

National Research Foundation

American Museum of Natural History

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Biostratigraphy of the Cynognathus Assemblage Zone (Beaufort Group, Karoo Supergroup), South Africa

South African Journal of Geology (2020)

P. John Hancox Johann Neveling Bruce S. Rubidge

Abstract The Cynognathus Assemblage Zone is the youngest tetrapod biozone of the Beaufort Group (Tarkastad Subgroup, Karoo Supergroup). It is situated between the underlying Lystrosaurus declivis Assemblage Zone and the base of the overlying Molteno Formation (Stormberg Group) and corresponds to the entire Burgersdorp Formation. It is characterised by the presence throughout of the cynodont genus Cynognathus. The biozone reaches a maximum thickness of around 650 m in the southeast part of the basin and thins dramatically to the north, where it is only a maximum of 50 m thick. We here propose a three-fold subdivision into a lower Langbergia-Garjainia Subzone, a middle Trirachodon-Kannemeyeria Subzone and an upper Cricodon-Ufudocyclops Subzone. The basal contact is deﬁned biostratigraphically by the ﬁrst appearance of Cynognathus crateronotus and Langbergia modisei. The Cynognathus Assemblage Zone lacks a deﬁned biostratigraphic upper limit, being unconformably terminated by the base of the overlying Molteno Formation, which lacks a terrestrial vertebrate fossil record other than trackways.

Biozone

Supergroup

Assemblage (archaeology)

10.25131/sajg.123.0016

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Citations (39)

A new kannemeyeriiform dicynodont (Ufudocyclops mukanelai, gen. et sp. nov.) from Subzone C of the Cynognathus Assemblage Zone, Triassic of South Africa, with implications for biostratigraphic correlation with other African Triassic faunas (project)

MorphoBank datasets (2020)

Christian F. Kammerer Pia A. Viglietti P. John Hancox Richard J. Butler Jonah N. Choiniere

Assemblage (archaeology)

10.7934/p3602

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AngonisaurusandShansiodon, dicynodonts (Therapsida, Anomodontia) from subzone C of theCynognathusAssemblage Zone (Middle Triassic) of South Africa

Journal of Vertebrate Paleontology (2013)

P. John Hancox Kenneth D. Angielczyk Bruce S. Rubidge

ABSTRACT Dicynodonts are the most abundant therapsids from the Beaufort Group (Karoo Basin, South Africa), and they have been used as index fossils to define most of the Beaufort biozones. Although speciose during the Late Permian, dicynodont diversity was greatly reduced by the end-Permian mass extinction. The Cynognathus Assemblage Zone, the uppermost biozone of the Beaufort Group, is traditionally known to host only the dicynodonts Kannemeyeria simocephalus and Kombuisia frerensis. Recent biostratigraphic research in this biozone has revealed the presence of two more poorly known dicynodont genera, Angonisaurus and Shansiodon, which are biostratigraphically constrained to the uppermost Cynognathus Assemblage Zone (subzone C). Although these taxa have been reported from subzone C of the Cynognathus Assemblage Zone previously, we provide the first detailed description of the material in question and justifications for the taxonomic identifications. The presence of Angonisaurus in the Cynognathus subzone C allows a direct correlation between this assemblage and that of the Lifua Member of the Manda beds in Tanzania. A correlation with the Antarctic upper Fremouw Formation also may be possible, but is less certain. The presence of Shansiodon allows a direct correlation with the upper Ermaying Formation of China, but we reject the previously proposed global 'Shansiodon biochron' because Shansiodon sensu stricto only occurs in China and South Africa. Biogeographically, Angonisaurus appears to have been widespread but rare in southern Gondwana, and the unusual distribution of Shansiodon mirrors that of Diictodon feliceps in the Permian. Southern African dicynodont assemblages display increased provinciality in the Middle Triassic.

Biozone

Assemblage (archaeology)

Chronostratigraphy

10.1080/02724634.2013.723551

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Citations (39)

Permian climate change recorded in palynomorph assemblages of Mozambique (Moatize Basin, eastern Tete Province)

Acta Palaeobotanica (2017)

Annette E. Götz P. John Hancox Andrew Lloyd

Abstract Knowledge of Late Permian biodiversity patterns, following the end-Guadalupian crisis, is still in its infancy, since most recent studies have focused on the end-Permian biotic crisis. The palynological record of southern Africa, however, reveals major climatic changes during the Late Permian. Here we report new palynological data from eastern Tete Province of Mozambique, documenting the change from cool to warm temperate climates during the Lopingian. This prominent climate signal was also detected recently in other depositional environments elsewhere in southern Africa, and thus enables interregional temporal correlations.

10.1515/acpa-2017-0001

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Complex multicusped postcanine teeth from the Lower Triassic of South Africa

Journal of Vertebrate Paleontology (2012)

Leandro C. Gaetano Helke Mocke Fernando Abdala P. John Hancox

ABSTRACT Two incisors and five postcanine teeth of complex crown morphology were found in the lower levels of the Burgersdorp Formation of the Beaufort Group, corresponding to the basal Subzone A of the Cynognathus Assemblage Zone (late Olenekian). All the teeth bear a single root and the postcanines show two rows of mesiodistally aligned cusps and a central basin or groove. Among the postcanines, two general patterns of crown morphology are recognized on the basis of relative length of the cusp rows, number of cusps per row, and width of the central area. Phylogenetic affinities of the described specimens remain unclear, and thus they are regarded as Eucynodontia incertae sedis. However, comparisons with non-mammalian cynodonts with labiolingually expanded postcanines show that they are most similar to those of Aleodon, Candelariodon, and some haramiyids. Many cynodont taxa such as Aleodon, Candelariodon, Cromptodon, haramiyids, and tritylodontids, which are probably not closely related to each other, have labiolingually expanded postcanines with cusps arranged mesiodistally in rows and a central basin; thus pointing to the homoplastic nature of dental morphologies in the cynodont lineage. The teeth presented here are the oldest record of therapsid teeth with crowns having parallel rows of cusps, representing a temporal extension of approximately 10 million years for this crown pattern.

Incertae sedis

10.1080/02724634.2012.697086

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Waterford Formation in the south-eastern Karoo : implications for basin development : research letter

South African Journal of Science (2012)

Bruce S. Rubidge P. John Hancox Richard Mason

Extensive research on the rocks of the Karoo Supergroup has shown that this sequence, which contains an unsurpassed record of Permian-Jurassic tetrapods, records a largely unbroken stratigraphic succession from 300 Ma to 180 Ma. This Gondwanan succession was deposited in a changing environmental setting reflecting glacial marine through deltaic to fluvial and aeolian desert conditions. The contact between the Ecca and Beaufort Groups (at the top of the Waterford Formation of the Ecca Group) in the southern and western Karoo represents a change in depositional environment from a subaqueous to a subaerial delta plain. By contrast, the Waterford Formation has not yet been recognised in the south-eastern Karoo Basin, which might imply that a major unconformity is present between the Fort Brown Formation of the Ecca Group, deposited in a prodelta environment, and the overlying fluvially deposited Koonap Formation of the Beaufort Group. From careful documentation of lithofacies and sedimentological data, it can be demonstrated that the Waterford Formation is indeed present in the south-eastern part of the basin and that no major unconformity is present - a fact that has implications for the mapping of Karoo rocks in the Eastern Cape Province of South Africa, for understanding the depositional environment of 'reptilian' fossils from the lowermost Beaufort in this part of the Karoo basin, and for basin development models.

Beaufort scale

Subaerial

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The Earliest Post-Paleozoic Freshwater Bivalves Preserved in Coprolites from the Karoo Basin, South Africa

PLoS ONE (2012)

Adam M. Yates Frank Neumann P. John Hancox

Several clades of bivalve molluscs have invaded freshwaters at various times throughout Phanerozoic history. The most successful freshwater clade in the modern world is the Unionoida. Unionoids arose in the Triassic Period, sometime after the major extinction event at the End-Permian boundary and are now widely distributed across all continents except Antarctica. Until now, no freshwater bivalves of any kind were known to exist in the Early Triassic.Here we report on a faunule of two small freshwater bivalve species preserved in vertebrate coprolites from the Olenekian (Lower Triassic) of the Burgersdorp Formation of the Karoo Basin, South Africa. Positive identification of these bivalves is not possible due to the limited material. Nevertheless they do show similarities with Unionoida although they fall below the size range of extant unionoids. Phylogenetic analysis is not possible with such limited material and consequently the assignment remains somewhat speculative.Bivalve molluscs re-invaded freshwaters soon after the End-Permian extinction event, during the earliest part of the recovery phase during the Olenekian Stage of the Early Triassic. If the specimens do represent unionoids then these Early Triassic examples may be an example of the Lilliput effect. Since the oldest incontrovertible freshwater unionoids are also from sub-Saharan Africa, it is possible that this subcontinent hosted the initial freshwater radiation of the Unionoida. This find also demonstrates the importance of coprolites as microenvironments of exceptional preservation that contain fossils of organisms that would otherwise have left no trace.

Early Triassic

Permian–Triassic extinction event

Extinction (optical mineralogy)

Freshwater bivalve

10.1371/journal.pone.0030228

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Citations (19)

ELLIOTHERIUM KERSTENI, A NEW TRITHELEDONTID FROM THE LOWER ELLIOT FORMATION (UPPER TRIASSIC) OF SOUTH AFRICA

Journal of Paleontology (2006)

Christian A. Sidor P. John Hancox

We describe a new tritheledontid, Elliotherium kersteni n. gen. and sp., on the basis of a partial skull collected from the lower Elliot Formation (Upper Triassic; Euskelosaurus Range Zone) on the farm Beatrix, Free State Province, South Africa. Although similar to Pachygenelus, the genus differs from other South African tritheledontids in its higher maxillary postcanine tooth count, lack of labial cingula on the upper postcanine teeth, shorter secondary palate, retention of a median vomerine ridge behind the secondary palate, and smaller interpterygoid vacuity. With regard to these features, Elliotherium is most similar to the South American tritheledontid Chaliminia. Elliotherium is autapomorphic in its possession of a long parasphenoid rostrum, ventrally depressed secondary palate, medially convex margins of the interpterygoid vacuity, and a thin lamina of palatine covering the interpterygoid vacuity. Although Elliotherium is the stratigraphically lowest positioned tritheledontid from southern Africa, it is still probably later occurring than the South American taxa Chaliminia or Riograndia. Based on a revision of a previous analysis of advanced cynodont relationships, we propose a sister-group relationship between Elliotherium and Chaliminia, although support for this pairing is weak.

Rostrum

Autapomorphy

10.1666/0022-3360(2006)080[0333:ekantf]2.0.co;2

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Citations (40)

Unique trackway on Permian Karoo shoreline provides evidence of temnospondyl locomotory behaviour

PLoS ONE (2023)

David P. Groenewald Ashley Krüger Michael O. Day Cameron R. Penn-Clarke P. John Hancox

Large-bodied temnospondyl amphibians were the dominant predators in non-marine aquatic ecosystems from the Carboniferous to the Middle Triassic. In the Permian-aged lower Beaufort Group of the main Karoo Basin, South Africa, temnospondyls are represented exclusively by the family Rhinesuchidae and are well represented by body fossils, whereas trace fossils are scarce. Accordingly, most interpretations of the behaviour of this family are based on skeletal morphology and histological data. Here we document the sedimentology and palaeontology of a late Permian palaeosurface situated immediately below the palaeoshoreline of the Ecca Sea (transition from the Ecca Group to the Beaufort Group) near the town of Estcourt in KwaZulu-Natal Province. The surface preserves numerous ichnofossils, including tetrapod footprints and fish swim-trails, but most striking are seven body impressions and associated swim trails that we attribute to a medium-sized (~1.9 m long) rhinesuchid temnospondyl. These provide valuable insight into the behaviour of these animals. The sinuous shape of some of the traces suggest that the tracemaker swam with continuous sub-undulatory propulsion of the tail.

Tetrapod (structure)

10.1371/journal.pone.0282354

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THE PERMIAN-TRIASSIC BOUNDARY, KAROO BASIN, SOUTH AFRICA: GEOCHEMICAL INSIGHT INTO THIS MASS EXTINCTION EVENT. L. Coney

L. Coney W. U. Reimold P. John Hancox Dieter Mader Christian Koeberl

Permian–Triassic extinction event

Extinction (optical mineralogy)

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