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

2020 ◽  
Vol 123 (2) ◽  
pp. 181-190
Author(s):  
R.M.H. Smith

Abstract The late Permian (Lopingian) Cistecephalus Assemblage Zone (CiAZ) of the Karoo Supergroup in South Africa has recently been radiometrically-dated to range from 256 to 255 My. It encompasses approximately one million years of the late Wuchiapingian epoch, at a time when the ancient intra-continental lowlands of southern Gondwana had fully recovered from the end-Guadalupian mass extinction. The diverse Cistecephalus Assemblage Zone fauna is dominated by the small herbivorous dicynodonts Diictodon, Pristerodon and the molelike Cistecephalus, along with a range of larger dicynodont herbivores including Oudenodon, Aulacephalodon, Rhachiocephalus, Dinanomodon and rare Endothiodon. The attendant large carnivores include the gorgonopsians Aelurognathus, Rubidgea, and Smilesaurus, while smaller carnivores are represented by eutherocephalians (e.g., Ictidosuchoides, Ictidosuchops) and small gorgonopsians (e.g., Aloposaurus, Scylacocephalus). Of the parareptiles, the large-bodied taxon Pareiasaurus is most common, with the diminutive pareiasaurs Anthodon, Nanoparia, and Pumiliopareia making their first appearance. Lithostratigraphically, the biozone for the most part coincides with the arenaceous Oukloof and lower Steenkampsvlakte members in the western sub-basin and the equivalent Oudeberg and lower Daggaboersnek members in the east, where it reaches its maximum thickness of 300 m. The Cistecephalus Assemblage Zone thins westwards to 120 m at Teekloof Pass, and eastwards to approximately 100 m near the town of Fort Beaufort.

2020 ◽  
Vol 123 (2) ◽  
pp. 165-180 ◽  
Author(s):  
M.O. Day ◽  
R.M.H. Smith

Abstract The Endothiodon Assemblage Zone is the third oldest tetrapod biozone of the Beaufort Group (Adelaide Subgroup, Karoo Supergroup). It is situated between the underlying Tapinocephalus and overlying Cistecephalus assemblage zones and in the southwestern part of the basin corresponds to the majority of the Poortjie and Hoedemaker members of the Teekloof Formation. It is characterised by the dicynodont genus Endothiodon, especially in the lower part of assemblage zone, and records early ecosystem recovery from the Capitanian mass extinction. It also contains the lowest occurrence in the Karoo Basin of cynodont therapsids, eutherocephalians, bidentalian dicynodonts, and diapsids. The biozone reaches a maximum thickness of around 250 m in the southwestern part of the basin. We propose a two-fold subdivision into a lower Lycosuchus - Eunotosaurus Subzone (equivalent to the upper two-thirds of the former Pristerognathus Assemblage Zone) and an upper Tropidostoma - Gorgonops Subzone (equivalent to the former Tropidostoma Assemblage Zone), with the contact defined by the first appearance of Tropidostoma dubium. The Endothiodon Assemblage Zone is terminated by the first appearance of Aulacephalodon bainii.


2020 ◽  
Vol 123 (2) ◽  
pp. 149-164 ◽  
Author(s):  
M.O. Day ◽  
B.S. Rubidge

Abstract The Tapinocephalus Assemblage Zone is the second oldest tetrapod biozone of the Beaufort Group (Adelaide Subgroup, Karoo Supergroup), biostratigraphically positioned between the underlying Eodicynodon and overlying Endothiodon assemblage zones. It is characterised by a rich fossil tetrapod assemblage comprising basal members of most therapsid clades, but particularly dinocephalians such as Moschops capensis and basal pareiasaurs such as Bradysaurus in co-occurrence with the pylaecephalid dicynodonts Robertia and Eosimops. It corresponds to the upper two thirds of the Abrahamskraal Formation, is Capitanian (Guadalupian) in age, and reaches a maximum thickness of around 1500 m. The biozone is here separated into two subzones: a lower Eosimops – Glanosuchus Subzone and an upper Diictodon – Styracocephalus Subzone. The contact between the two subzones is defined by the first appearance of Diictodon feliceps, which closely corresponds to the base of the Moordenaars Member. The uppermost part of the biozone contains the Capitanian mass extinction and the low diversity fauna in its immediate aftermath. The zone is terminated by the first appearance of Endothiodon bathystoma.


2021 ◽  
Vol 9 ◽  
Author(s):  
Christian A. Sidor ◽  
Neil J. Tabor ◽  
Roger M. H. Smith

A new burnetiamorph therapsid, Isengops luangwensis, gen. et sp. nov., is described on the basis of a partial skull from the upper Madumabisa Mudstone Formation of the Luangwa Basin of northeastern Zambia. Isengops is diagnosed by reduced palatal dentition, a ridge-like palatine-pterygoid boss, a palatal exposure of the jugal that extends far anteriorly, a tall trigonal pyramid-shaped supraorbital boss, and a recess along the dorsal margin of the lateral temporal fenestra. The upper Madumabisa Mudstone Formation was deposited in a rift basin with lithofacies characterized by unchannelized flow, periods of subaerial desiccation and non-deposition, and pedogenesis, and can be biostratigraphically tied to the upper Cistecephalus Assemblage Zone of South Africa, suggesting a Wuchiapingian age. Isengops is the second burnetiamorph recognized from Zambia and is part of a tetrapod assemblage remarkably similar to others across southern Pangea during the Wuchiapingian. A revised cladistic analysis of Biarmosuchia yielded over 500 most parsimonious trees that generally reaffirm the results of previous analyses for burnetiamorphs: Lemurosaurus is basal, Lobalopex and Isengops are proximate burnetiid outgroups, and Bullacephalus, Burnetia, Mobaceras, Niuksenitia, and Pachydectes are burnetiines. Furthermore, Russian biarmosuchians are scattered throughout the tree and do not form sister taxon relationships with each other. Burnetiamorphs display a wide disparity of cranial adornments and are relatively speciose (13 species), especially when compared to the number of specimens discovered to date (∼16 specimens). As has been suggested in some other tetrapod clades (e.g., ceratopsian dinosaurs), the burnetiamorph fossil record supports an inferred macroevolutionary relationship between cranial adornment and increased speciation rate.


2020 ◽  
Vol 123 (2) ◽  
pp. 239-248 ◽  
Author(s):  
P.A. Viglietti ◽  
B.W. McPhee ◽  
E.M. Bordy ◽  
L. Sciscio ◽  
P.M. Barrett ◽  
...  

Abstract The Scalenodontoides Assemblage Zone (SAZ) is the oldest fossil tetrapod biozone of the Stormberg Group (Karoo Supergroup) and preserves the oldest dinosaur bearing deposits in the Karoo Basin. The SAZ represents a revision of the ‘Euskelosaurus’ Range Zone, whose taxonomic basis has been undermined because ‘Euskelosaurus’ is well demonstrated to be a nomen dubium. Recent qualitative and quantitative investigations into the biostratigraphy of the Elliot and Clarens formations have resulted in the first biostratigraphic review of all lower Elliot Formation (lEF) taxa in nearly 40 years. Thus, we replace the ‘Euskelosaurus’ Range Zone with a new biostratigraphic assemblage zone, the Scalenodontoides Assemblage Zone (SAZ). Named after the traversodontid cynodont Scalenodontoides macrodontes, which co-occurs with the sauropodomorphs Blikanasaurus cromptoni and Melanorosaurus readi. The SAZ is currently accepted to range in age between the upper Norian and Rhaetian. Our new biozone, which reaches a maximum thickness of ~200 m, is wholly contained within the lower Elliot Formation (Stormberg Group, Karoo Supergroup).


2015 ◽  
Vol 89 (4) ◽  
pp. 645-664 ◽  
Author(s):  
Adam K. Huttenlocker ◽  
Fernando Abdala

AbstractHistorically, the whaitsiid therocephalianTheriognathusOwen was one of the earliest described nonmammalian therapsids, its morphology helping to link phylogenetically the Paleozoic synapsids of North America and southern Africa to their mammalian successors. However, decades of taxonomic over-splitting and superficial descriptions obscured the morphologic diversity of the genus, hindering its utility as a study system for the evolution of synapsid cranial function as well as its biostratigraphic significance in the Late Permian of southern Africa. Here, we revise the status and provenance of all the known specimens ofTheriognathusfrom South Africa, Tanzania, and Zambia. We present both qualitative and quantitative support for the presence of a single morphospecies as proposed by some authors. Proportional differences in skulls that were previously ascribed to different morphotypes (‘Aneugomphius,’ ‘Notosollasia,’ ‘Moschorhynchus,’ and ‘Whaitsia’) are largely size-related and allometric trends are considered here in the context of jaw function and prey prehension. Our results suggest that the single species,Theriognathus microps, represented one of the most abundant Late Permian therocephalians in southern Africa and is consequently a potentially useful biostratigraphic marker for the upperCistecephalus-lowerDicynodonAssemblage Zone transition (i.e., late Wuchiapingian). The wide range of preserved sizes in conjunction with recent paleohistological evidence supports that individuals spent much of their lives in an actively-growing, subadult phase. LaterDicynodonAssemblage Zone records (e.g., upper Balfour Formation) are unconfirmed as the genus was likely replaced by other theriodont predators (e.g.,Moschorhinus) leading up to the Permo-Triassic boundary in the Karoo Basin of South Africa.


2020 ◽  
Vol 123 (2) ◽  
pp. 249-262 ◽  
Author(s):  
P.A. Viglietti ◽  
B.W. McPhee ◽  
E.M. Bordy ◽  
L. Sciscio ◽  
P.M. Barrett ◽  
...  

Abstract The Massospondylus Assemblage Zone is the youngest tetrapod biozone in the Karoo Basin (upper Stormberg Group, Karoo Supergroup) and records one of the oldest dinosaur dominated ecosystems in southern Gondwana. Recent qualitative and quantitative investigations into the biostratigraphy of the lower and upper Elliot formations (lEF, uEF) and Clarens Formation in the main Karoo Basin resulted in the first biostratigraphic review of this stratigraphic interval in nearly four decades, allowing us to introduce a new biostratigraphic scheme, the Massospondylus Assemblage Zone (MAZ). The MAZ expands upon the Massospondylus Range Zone by including the crocodylomorph Protosuchus haughtoni and the ornithischian Lesothosaurus diagnosticus as two co-occurring index taxa alongside the main index taxon, the sauropodomorph Massospondylus carinatus. With a maximum thickness of ~320 m in the southeastern portion of the basin, our new biozone is contained within the uEF and Clarens formations (upper Stormberg Group), however, based on vertebrate ichnofossils evidence, it may potentially extend into the sedimentary units of the lowermost Drakensberg Group. We do not propose any further subdivisions, and do not consider the Tritylodon Acme Zone (TAZ) as a temporal biostratigraphic marker within the MAZ. The MAZ is currently accepted to range in age between the Hettangian and Pliensbachian, however a faunal turnover, which observes an increase in the diversity of dinosaur clades, crocodylomorph, and mammaliaform taxa in the lower uEF, could reflect effects of the end-Triassic extinction event (ETE).


PeerJ ◽  
2016 ◽  
Vol 4 ◽  
pp. e1608 ◽  
Author(s):  
Christian F. Kammerer

The subfamily Rubidgeinae, containing the largest known African gorgonopsians, is thoroughly revised. Rubidgeinae is diagnosed by the absence of a blade-like parasphenoid rostrum and reduction or absence of the preparietal. Seven rubidgeine species from the Karoo Basin of South Africa are recognized as valid:Aelurognathus tigriceps,Clelandina rubidgei,Dinogorgon rubidgei,Leontosaurus vanderhorsti,Rubidgea atrox,Smilesaurus ferox, andSycosaurus laticeps. Rubidgeines are also present in other African basins:A. tigricepsandS. laticepsoccur in the Upper Madumabisa Mudstone Formation of Zambia, andD. rubidgei,R. atrox, and the endemic speciesRuhuhucerberus haughtonicomb. nov. andSycosaurus nowakicomb. nov. occur in the Usili Formation of Tanzania.Aelurognathus nyasaensisfrom the Chiweta Beds of Malawi also represents a rubidgeine, but of uncertain generic referral pending further preparation. No rubidgeine material is known outside of Africa: the purported Russian rubidgeineLeogorgon klimovensisis not clearly referable to this group and may not be diagnosable. Phylogenetic analysis of rubidgeines reveals strong support for a clade (Rubidgeini) of advanced rubidgeines includingClelandina,Dinogorgon,Leontosaurus, andRubidgea. Support forSmilesaurusas a rubidgeine is weak; it may, as previous authors have suggested, represent an independent evolution of large body size from anArctops-like ancestor. Temporally, rubidgeines are restricted to the Late Permian, first appearing in theTropidostomaAssemblage Zone and reaching highest diversity in theCistecephalusandDaptocephalusassemblage zones of the Beaufort Group.


2020 ◽  
Vol 90 (6) ◽  
pp. 609-628 ◽  
Author(s):  
Robert A. Gastaldo ◽  
Kaci Kus ◽  
Neil Tabor ◽  
Johann Neveling

ABSTRACT The fully continental succession of the Beaufort Group, Karoo Basin, South Africa, has been used in the development of environmental models proposed for the interval that spans the contact between the Daptocephalus to Lystrosaurus Assemblage Zones, associated by some workers with the end-Permian extinction event. An aridification trend is widely accepted, yet geochemical data indicate that the majority of in situ paleosols encountered in this interval developed in waterlogged environments. To date, the presence of calcic paleosols in the latest Permian can be inferred only from the presence of calcite-cemented pedogenic nodules concentrated in fluvial channel-lag deposits. Here, we report on the first empirical evidence of in situ calcic Vertisols found in the upper Daptocephalus Assemblage Zone near Old Wapadsberg Pass, one of eight classic localities in which the vertebrate turnover is reported in the Karoo Basin. Seven discrete intervals of calcic Vertisols, exposed over a very limited lateral extent, occur in an ∼ 25 m stratigraphic interval. Estimates of mean annual temperature and mean annual precipitation are calculated from geochemical measurements of one paleosol, and these estimates indicate that the prevailing climate at the time of pedogenesis was seasonally cold and humid. Correlation with adjacent stratigraphic sections indicates that the late Permian landscape experienced poorly drained and better-drained phases, interpreted to reflect a climate that varied between episodically dry and episodically wet. In contrast to a paleoenvironmental reconstruction of unidirectional aridification from strata in the Wapadsberg Pass region, this study provides new evidence for a wetting trend towards the Daptocephalus–Lystrosaurus Assemblage-Zone boundary.


2020 ◽  
Vol 123 (2) ◽  
pp. 141-148
Author(s):  
B.S. Rubidge ◽  
M.O. Day

Abstract The middle Permian Eodicynodon Assemblage Zone is the lowermost biozone of the Beaufort Group (Adelaide Subgroup, Karoo Supergroup) and occurs in the southwestern part of the main Karoo Basin. It is host to a diverse assemblage of basal therapsid genera of which Eodicynodon is the most abundant. The biozone reaches a maximum thickness of 1 100 m in the Prince Albert Road area and thins to the east and west. The biozone corresponds to the Combrinkskraal and Grootfontein members of the Abrahamskraal Formation, directly overlies the Waterford Formation of the Ecca Group, and records the earliest middle Permian terrestrial environments of Gondwana. Rocks of the biozone were deposited along the southern shoreline of the Karoo Basin in a subaerial delta plain environment as part of large-scale fan systems draining to the north and northeast within a second-order highstand systems tract.


PeerJ ◽  
2017 ◽  
Vol 5 ◽  
pp. e2913 ◽  
Author(s):  
Christian F. Kammerer ◽  
Roger M.H. Smith

Based on specimens previously identified asTropidostoma, a new taxon of dicynodont (Bulbasaurus phylloxyrongen. et sp. nov.) from the Karoo Basin of South Africa is described.Bulbasaurusis a medium-sized dicynodont (maximum dorsal skull length 16.0 cm) restricted to theTropidostomaAssemblage Zone (early Lopingian) of the Beaufort Group.Bulbasauruscan be distinguished fromTropidostomaby an array of characters including the presence of a tall, sharp premaxillary ridge, large, rugose, nearly-confluent nasal bosses, a nasofrontal ridge, massive tusks, robust pterygoids, prominently twisted subtemporal bar, and absence of a distinct postfrontal. Inclusion ofBulbasaurusin a phylogenetic analysis of anomodont therapsids recovers it as a member of Geikiidae, a clade of otherwise later Permian dicynodonts such asAulacephalodonandPelanomodon.Bulbasaurusexhibits many of the characters typical of adultAulacephalodon, but at substantially smaller skull size (these characters are absent in comparably-sizedAulacephalodonjuveniles), suggesting that the evolution of typical geikiid morphology preceded gigantism in the clade.Bulbasaurusis the earliest known geikiid and the only member of the group known from theTropidostomaAssemblage Zone; discovery of this taxon shortens a perplexing ghost lineage and indicates that abundant clades from the later Permian of South Africa (e.g., Geikiidae, Dicynodontoidea) may have originated as rare components of earlier Karoo assemblage zones.


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