Karoo Fossils

Of all the transitional series amongst the major groups of vertebrates, one of the most extraordinary and best documented is the transition from primitive amniotes to mammals. Amniotes are vertebrates whose embryos are totally enclosed in a fluid-filled sac, called the amnion and include living reptiles, birds and mammals. About 350 million years ago, the amniotes gave rise to two different lineages, namely the reptiles and the mammals. The group of animals that include mammals and all their ancestors is called the Synapsida. The early synapsids that lived between 260 and 180 million years ago are known as therapsids, a particularly significant group in the study of mammalian evolution as they are the direct ancestors of living mammals.

The South African Karoo Basin contains the best therapsid fossil record in the whole world giving us the exciting opportunity of being able to study the intricate details of the transition into mammals. These animals, that span a period of more than 80 million years from the Middle Permian to the Middle Jurassic, show the gradual acquisition of mammal-like characteristics until it is almost impossible to distinguish the latest most mammal-like therapsids from the earliest true mammals.

The research conducted in the Karoo Palaeontology Department focuses on the biology, ecology and biostratigraphy of these very important animals. Regular field excursions to the Karoo have allowed us to expand our current vertebrate collection and thus conserve these fossils, which form an important part of South Africa’s natural heritage.


A new giant dinosaur from the South African Free State Province

Ledumahadi reconstruction

2 October 2018
A new species of a giant dinosaur has been found in South Africa’s Free State Province. The plant-eating dinosaur, named Ledumahadi mafube, weighed 12 tonnes and stood about four metres high at the hips.Ledumahadi mafube was the largest land animal alive on Earth when it lived, nearly 200 million years ago. It was roughly double the size of a large African elephant. A team of international scientists, led by University of the Witwatersrand (Wits) palaeontologist Professor Jonah Choiniere, described the new species in the journal Current Biology. Ledumahadi mafube is one of the closest relatives of sauropod dinosaurs. Sauropods, weighing up to 60 tonnes, include well-known species like Brontosaurus. All sauropods ate plants and stood on four legs, with a posture like modern elephants. Ledumahadi evolved its giant size independently from sauropods, and although it stood on four legs, its forelimbs would have been more crouched. This caused the scientific team to consider Ledumahadi an evolutionary “experiment” with giant body size. Dr Botha-Brink investigated the life history of this animal by studying its bone microstructure. The animal grew rapidly to adulthood and was practically fully grown when it died. Its bone tissues display aspects of both basal sauropodomorphs and the more derived sauropods, showing that Ledumahadi represents a transitional stage between these two major groups of dinosaurs.

The palaeobiology of Brazilian cynodonts, the ancestors of mammals


2 October 2018
Dr Jennifer Botha-Brink from the Karoo Palaeontology Department at the National Museum has just published research in the journal PeerJ entitled “Osteohistology of Late Triassic prozostrodontian cynodonts from Brazil” in collaboration with Brazilian and Argentinean colleagues Drs Marina Bento Soares and Agustín Martinelli from the Universidade Federal do Rio Grande do Sul, Brazil. This contribution sheds new light on the palaeobiology of the ancestors of mammals. Known as non-mammaliaform cynodonts, these animals gave rise to mammals during the Early Jurassic, around 200 million years ago. Dr Botha-Brink and colleagues studied the bone microstructure of several members of the Prozostrodontia, the cynodont group most closely related to mammals, in order to decipher the growth patterns of these animals and compare them with those of living mammals. Some of the earliest prozostrodontians were about the size of an opposum whereas the later brasilodontids were tiny, similar in size to that of a modern shrew. All prozostrodontian cynodonts grew rapidly to maturity, but the later, tiny brasilodontid prozostrodontians grew relatively more slowly than their ancestors and more like that of the mammaliaform Morganucodon, an immediate precursor of the modern mammal lineage (monotremes and therians). This may be due to similar body sizes or the close family relationship between brasilodontids and mammaliaforms. Interestingly, when compared with similar-sized living mammals, they appear to have grown more slowly to adult size. Although the Prozostrodontia acquired increasingly mammalian features during their evolution, including rapid juvenile growth, the small brasilodontid prozostrodontians still exhibit an extended growth period compared to similar-sized living mammals.

Visit by previous head of department

Visit by previous head of department

12 January 2017
Prof. Steve Fourie (right) was the head of the Karoo Palaeontology Department in the early 1970s. During this time he and John Nyaphuli worked closely together. John joined the Museum in 1973 and soon showed talent for preparing fossils. During his time here he developed into the best fossil preparator in the country and one of the best in the world. After his retirement John remains involved at the Museum where he assists with training and also prepares delicate fossils. Prof. Fourie later left the Museum to accept a post at the University of the Free State, where he retired in 1998. He visited the Karoo Palaeontology Department where he and John, both 83 years old, caught up on days gone by.

Postdoctoral Fellow


10 January 2017
Dr Lucas Legendre from Paris joined the Karoo Palaeontology Department in July as a Postdoctoral Fellow for two years. He is working with Dr Jennifer Botha-Brink on the bone microstructure of burrowing animals. The ecology of an animal plays an important role in influencing the way bone tissue grows. Dr Legendre’s research aims to identify how a burrowing lifestyle affects bone tissue patterns. Results will be used to infer burrowing or digging behaviours from the fossil bone microstructure of extinct animals. This kind of evidence can shed light on the ecology of an extinct animal when other avenues (e.g. skeletal anatomy) remain ambiguous.


Specialist Museum Scientist and Head of Department

Jennifer Botha PhD, Pr Sci Nat

JenniferMy research interests are in palaeobiology and palaeoecology, focusing on the life history responses of extinct vertebrates to environmental change. Using the Permo-Triassic Mass Extinction (PTME) as a model, I use a variety of techniques, including biostratigraphy, morphology and osteohistology, to test theories regarding differential species survival during mass extinctions. I have a special interest in osteohistology, the study of fossil bone microstructure, which provides novel information about the life history of the vertebrates associated with the PTME.

My PhD research (University of Cape Town, 2002) entailed the development and application of a variety of techniques used to assess the biology of extinct vertebrates, including osteohistology and stable isotope analysis, focusing on the palaeohistology of cynodont therapsids (the ancient ancestors of mammals). Thereafter, I took up a postdoctoral fellowship at the Iziko Museums of South Africa, during which time I focused largely on the biostratigraphy of the Permo-Triassic Karoo Basin. Since 2005, I have been head of the Karoo Palaeontology Department of the National Museum, Bloemfontein and Affiliated Researcher with the Department of Zoology and Entomology at the University of the Free State, where I have established a long-term research programme studying macro- and micro-skeletal structure, as well as spatial and temporal distributions of Permo-Triassic fossil vertebrates to gain a deeper understanding of the biotic crisis associated with the PTME.

I have presented or co-authored more than 30 conference presentations at both national conferences and abroad, published more than 35 articles in national and international scientific journals and am a B-rated scientist (National Research Foundation). Current research interests include the biology and ecology of Permo-Triassic vertebrates, the end-Permian mass extinction event and subsequent recovery, and the bone histology of fossil vertebrates.

Collections Manager and Principal Research Assistant

Elize Butler MSc

Elize ButlerElize Butler has been working at the National Museum since 1993. She obtained a BSc degree from the University of the Free State, majoring in Zoology and Botany and an Honours degree in Zoology. She recently completed an MSc thesis (cum laude) entitiled "The post-cranial skeleton of the Early Triassic non-mammalian cynodont Galesaurus planiceps: implications for biology and lifestyle". As Collections Manager she is responsible for the Geology and Palaeontology Collections at the Museum. This includes curating the collection, managing and maintaining the database and handling enquiries by scientists and members of the public.

Osteohistology Technican

Sekhomotso Gubuza

Sekhomotso joined the National Museum in 2017 to manage the Osteohistology Laboratory in the Karoo Palaeontology Department. Her tasks include producing bone histology thin sections of extant and extinct vertebrates, storing the thin sections and maintaining the digital databases. She also digitally renders each section so that we have an image database of each slide. Sekho is responsible for running the laboratory, training students on the equipment and trouble shooting.



Senior Fossil Preparator

Nthaopa Ntheri

Nthaopa worked at the Evolutionary Studies Institute (ESI) at the University of the Witwatersrand for five years before he joined the Palaeontology Department of the Museum in 2006. He prepares both small and larger fossil specimens, and assists with casting. He is also an experienced fossil finder in the field and was responsible for finding an almost complete, fully articulated Erythrosuchus africanus skeleton in 2008.

Sharon-LedibaneSenior Fossil Preparator

Sharon Ledibane

Sharon has been a member of the Palaeontology Department of the Museum since 2006. She prepares small and large fossils and occasionally assists with field work.

Sina ChakaFossil Preparator

Sina Chaka

Sina began working for the Museum in 2011. She mostly prepares small intricate fossils.

William MoleheFossil Preparator

William Molehe

William became a member of the Palaeontology Department in 2011. He prepares larger fossils, does most of the casting for the department and assists with field work.

Thabang NtsalaFossil Preparator

Thabang Ntsala

Thabang began working for the Museum in 2013. He prepares many large skeletons and assists with casting and field work.


Organismal Response to Severe Environmental Perturbation

Permo-Triassic Boundary

Permo-Triassic boundary site in the South African Karoo Basin

This research programme focuses on the response of extinct vertebrates - from the microscopic to macroscopic level - during severe environmental shifts that occur during mass extinctions. One of the greatest challenges in understanding the nature of past ecosystems is being able to accurately assess the biology and ecology of the extinct taxa living in these systems. This is particularly the case when attempting to understand how the life histories of extinct vertebrates varied through time, particularly during periods of extreme environmental fluctuation, such as those demonstrated by mass extinctions. Mass extinctions present an excellent opportunity for testing theories about differential survival in the face of dramatic environmental perturbation. The most catastrophic mass extinction event in Phanerozoic history is the Permo-Triassic Mass Extinction (PTME), which occurred approximately 252 million years ago. The South African Karoo Basin preserves the most complete terrestrial record of the Permo-Triassic sequence facilitating detailed studies on the effect of the extinction on vertebrate biology and ecology. Using the PTME as a model, a variety of techniques, including biostratigraphy, geochemistry, morphology and osteohistology, are used to test hypotheses regarding differential species survival.

Osteohistology of Fossil Vertebrates

Cynognathus histology

Life histories can be assessed based on the bone microanatomy and histology of vertebrates because these are known to reflect growth rates and patterns, ontogenetic stages, reproductive maturity, biomechanical adaptations, lifestyles and potentially the effects of significant environmental perturbation on growth. The osteohistology (bone microstructure) of fossil animals is thus one of the most powerful avenues for investigating the biology and ecology of extinct vertebrates. This research programme involves the study of extinct vertebrate life histories and aims to assess life history variation through time, partiularly during times of environmental stress, such as those caused by mass extinctions, and to identify osteohistological characteristics that may aid in survival.

Osteohistology of Living Vertebrates

aardvark histology

The aim of this programme is to develop a comparative modern vertebrate osteohistology database that can be used to study the osteohistology of extinct vertebrates. Various South African vertebrates have been collected for the purpose of developing this database. Research on this collection is currently focused on fossorial (burrowing) and semi-fossorial mammals. Numerous studies have shown that lifestyle preference affects the bone microstructure of an animal. Although much research has been done on the osteohistology of terrestrial and aquatic vertebrates, relatively little has been done on fossorial and semi-fossorial animals.

Students and Opportunities

A variety of projects in vertebrate palaeontology (including palaeohistology) at the postgraduate (BScHons, MSc and PhD) and postdoctoral levels are available.  Enquiries should be made to Dr Jennifer Botha (Head of the Karoo Palaeontology Department) at


Dr Lucas Legendre (2016 - 2018)

Bone microstructure of fossorial modern mammals.

This study aimed to investigate the fossorial activity of modern mammals, using histological thin sections of limb bones to evaluate biomechanical and structural constraints associated with burrowing. This study focused on the South African aardvark, Orycteropus afer to assess the influence of fossorial behaviour on the osteohistology of this animal.


Brandon Stuart MSc candidate (2020-)

The postcranial skeleton of Therocephalia (Synapsida: Therapsida) from the Karoo Basin of South Africa.

This study aims to describe the postcranial skeletons of various South Africa therocephalian species from the Karoo Basin of South Africa. It is an expansion of his honours thesis, which focused on the postcranial description of the therocephalian Moschorhinus kitchingi.

Bailey Weiss MSc candidate (2019-)

Bone microanatomy of Anomodontia Owen, 1859 (Synapsida: Therapsida) from the Karoo Basin of South Africa.

This study aims to study the bone microanatomy, i.e. the internal structure and organization of bone, of various dicynodont therapsid taxa from South Africa in order to assess what biomechanical and ecological signals can be obtained from bone microanatomy.


Kimberley Chapelle PhD (2018-2019)

The ontogeny and taxonomy of the basal sauropodomorph dinosaur Massospondylus carinatus (Owen, 1854).

This study examined the taxonomy and osteohistology of the Early Jurassic sauropodomorph dinosaur Massospondylus carinatus from the upper Elliot Formation, South Africa. She revised the cranial description and assessed an ontogenetic series of various postcranial elements to examine whether there was a postural shift during the growth of this taxon.

Luke Norton PhD (2013-2020)

Tooth replacement patterns in the Eutheriodontia from South Africa.

This study assessed the tooth replacement of selected therocephalian and cynodont therapsids using micro-CT scanning in order to determine tooth replacement patterns and rates within derived members of the Therapsida.

Alexander Botha MSc (2015-2017)

Comparative bone histology of Stigmochelys pardalis (Leopard tortoise), with specific reference to ontogeny and biomechanics.

This study examined the limb bone histology and bone microanatomy of the Leopard tortoise Stigmochelys pardalis from South Africa. He studied an ontogenetic series of various limb elements allowing him to describe inter-elemental variation as well as the changes in growth pattern through ontogeny.

Adam Huttenlocker PhD (2009-2013)

Osteohistology and skeletal growth in Permo-Triassic therocephalian therapsids (Amniota: Synapsida).

This study analysed the growth patterns in therocephalian and cynodont therapsids using osteohistological and skeletochronological methods, and re-assessed the phylogenetic relationships amongst Therocephalia. Changes in body size and growth rates were examined in order to shed light on therocephalian evolution with special emphasis on taxa associated with the Permo-Triassic mass extinction.

Publications - Scientific Articles

Current Researchers

Botha, J. and A. K. Huttenlocker. 2021. Non-mammalian synapsids; pp. 550-563 in V. de Buffrénil, A. de Ricqlès, L. Zylberberg and K. Padian (eds.), Comparative skeletal histology and paleohistology, CRC Press, Taylor and Francis, Boca Raton, 824 pp. PDF 2.8MB

Chapelle, K. E. J., J. Botha and J. N. Choiniere. 2021. Extreme growth plasticity in the early branching sauropodomorph Massospondylus carinatus. Biology Letters 17: 20200843. DOI: PDF 416 MB

Viglietti, P. A., R. B. J. Benson, R. M. H. Smith, J. Botha, C. Kammerer, Z. Skosan, N. Mtalana, S. Mtungata, D. Wolvaardt, M. Strong, G. Skinner, P. October, K. D. Angielczyk. 2021. Evidence from South Africa for a protracted end-Permian extinction on land. Proceedings of the National Academy of Science 118(17): e2017045118. DOI: view link. PDF on request.

Norton, L. A., F. Abdala, B. S. Rubidge and J. Botha. 2020. Tooth replacement patterns in the Early Triassic epicynodont Galesaurus planiceps (Therapsida, Cynodontia). PLOS One 15(12) e0243985. DOI: PDF 6.2 MB.

Botha, J. 2020. The paleobiology and paleoecology of South African Lystrosaurus. PeerJ 8:e10408. DOI: PDF 7.9 MB.

Botha, J. and R. M. H. Smith. 2020. Biostratigraphy of the Lystrosaurus declivis Assemblage Zone in R. M. H. Smith (ed.), Karoo biostratigraphy. Special Issue. South African Journal of Geology 123(2): 207-216. PDF 3.7 MB.

Smith, R. M. H., Rubidge, B. S., Day, M. O. and J. Botha. 2020. Introduction to the tetrapod biozonation of the Karoo Supergroup in R. M. H. Smith (ed.), Karoo biostratigraphy. Special Issue. South African Journal of Geology 123(2): 131-140. PDF 4.1 MB.

Botha, J., A. K. Huttenlocker, R. M. H. Smith, R. Prevec, P. Viglietti and S. P. Modesto. 2020. New geochemical and palaeontological data from the Permian-Triassic boundary in the South African Karoo Basin test the synchronicity of terrestrial and marine extinctions. Palaeogeography, Palaeoclimatology, Palaeoecology. 540: 109467. DOI: PDF 8.7 MB. | SUPP tables. | SUPP info.

Mayer, E. L., A. Hubbe, J. Botha-Brink, A. M. Ribeiro, X. S. Villagran and M. de Almeida Rizzutto. 2020. Diagenetic changes on bone histology of Quaternary mammals from a tropical cave deposit in southeastern Brazil. Palaeogeography, Palaeoclimatology, Palaeoecology 537: 109372. PDF 3.7 MB.

Botha, A. E. and J. Botha. 2019. Ontogenetic and inter-elemental osteohistological variability in the leopard tortoise Stigmochelys pardalis. PeerJ 7e:8030 DOI 10.7717/peerj.8030. PDf 2.5 MB.

Chapelle, K. E. J., P. M. Barrett, J. Botha and J. N. Choiniere. 2019. Ngwevu intloko: a new early sauropodomorph dinosaur from the Lower Jurassic Elliot Formation of South Africa and comments on cranial ontogeny in Massospondylus carinatus. PeerJ. DOI:10.7717/peerj.7240. PDF 5 MB

Barrett, P. M., K. E. J. Chapelle, C. K. Staunton, J. Botha and J. Choiniere. 2019. Postcranial osteology of the neotype specimen of Massospondylus carinatus Owen, 1854 (Dinosauria: Sauropodomorpha) from the upper Elliot Formation of South Africa. Palaeontologia Africana 53: 114-178. | PDF 8 MB.

Veiga, F. H., J. Botha-Brink, A. Maria Ribeiro, J. Ferigolo and M. Bento Soares. 2019.  Osteohistology of the Late Triassic silesaurid Sacisaurus agudoensis from southern Brazil and implications for growth in early dinosaurs. Anais da Academia Brasileira de Ciências 91(2): e20180643. DOI 10.1590/0001-3765201920180643. PDF 2.7 MB.

McPhee, B. R. Benson, J. Botha-Brink, E. Bordy and J. Choiniere. 2018. A giant dinosaur from the earliest Jurassic of South Africa and the transition to quadrupedality in early sauropodomorphs. Current Biology 28: 1-9. DOI: view link.

Legendre, L. J. and J. Botha-Brink. 2018. Digging the compromise: the functional link between bone microstructure and fossoriality in the aardvark (Orycteropus afer). PeerJ 6:e5216. DOI: PDF 4 MB

Botha-Brink, J., M. B. Soares and A. G. Martinelli. 2018. Osteohistology of Late Triassic prozostrodontian cynodonts from Brazil. PeerJ 6:e5029. DOI  PDF 2 MB.

Butler, E., F. Abdala andJ. Botha-Brink. 2018. Postcranial anatomy of the Early Triassic non-mammaliaform cynodont Galesaurus planiceps (Owen, 1859) from the Karoo Basin, South Africa. Papers in Palaeontology. DOI: PDF on request

Roopnarine, P. D., K. A. Angielczyk, S. Olroyd, S. J. Nesbitt, J. Botha-Brink, B. R. Peecook, M. O. Day and R. M. H. Smith. 2018. Comparative ecological dynamics of Permian-Triassic communities from the Karoo, Luangwa and Ruhuhu basins of southern Africa; pp. 254-272 in C. A. Sidor and S. J. Nesbitt (eds.), Vertebrate and climate evolution in the Triassic rift basin of Tanzania and Zambia. Society of Vertebrate Paleontology Memoir 17. Journal of Vertebrate Paleontology 37(6, Supplement). DOI: view link. PDF on request

Veiga, F. H., J. Botha-Brink and M. B. Soares. 2018. Osteohistology of the non-mammaliaform traversodontids Protuberum cabralense and Exaeretodon riograndensis from southern Brazil. Historial Biology DOI: view link. PDF on request

Botha-Brink, J. 2017. Burrowing in Lystrosaurus: preadaptation to a postextinction environment? Journal of Vertebrate Paleontology 37(5): e1365080. DOI: view link. PDF on request

Codron, J., J. Botha-Brink, D. Codron, A. Huttenlocker and K. Angielczyk. 2017. Predator prey interactions among Permo-Triassic terrestrial vertebrates as a deterministic factor influencing faunal collapse and turnover. Journal of Evolutionary Biology 30(1):40-54. DOI: 10.1111/jeb.12983. PDF on request

Legendre, L. J., G. Guénard, J. Botha-Brink and J. Cubo. 2016. Palaeohistological evidence for ancestral endothermy in archosaurs. Systematic Biology DOI: 10.1093/sysbio/syw033. PDF on request

Lyson, T. R., B. S. Rubidge, T. M. Scheyer, K. de Queiroz, E. R. Schachner, R. M. H. Smith and J. Botha-Brink. 2016. Fossorial origin of the turtle shell. Current Biology 26:1-8. DOI: 10.1016/j.cub.2016.05.020. PDF on request

Botha-Brink, J., D. Codron, A. K. Huttenlocker, K. D. Angielczyk and M. Ruta. 2016. Breeding young as a survival strategy during Earth's greatest mass extinction. Scientific Reports. 6: 24053. PDF 1.3 MBPDF SUPP 4.7 MB. Excel SUPP: view link.

Ecker, M., J. Botha-Brink, J. A. Lee-Thorp and L. K. Horwitz. 2015. Ostrich eggshell as a source of palaeoenvironmental information in the arid interior of South Africa; pp. 95-111 in J. Runge (ed.), Changing climates, ecosystems and environments within arid southern Africa and adjoining regions: Palaeoecology of Africa 33. CRC Press, Florence, Kentucky, 228 pp. PDF on request

Lyson, T. R., T. M. Scheyer, E. Schachner, J. Botha-Brink, G. S. Bever, K. de Queiroz,C. Farmer, B. S. Rubidge and J. A. Gauthier. 2014. Fossorial ancestry in the origin of turtles? Insights into the origin of the turtle lung ventilation mechanism and their shell. Nature Communications 5:5211 DOI: 10.1038/ncomms6211. PDF on request

Gower, D. J., R. J. Butler, A. G. Sennikov, J. Hancox and J. Botha-Brink. 2014. A new species of Garjainia ochev, 1958 (Diapsida: Archosauriformes: Erythrosuchidae) from the Early Triassic of South Africa. PLoS ONE 9: 11: e111154. PDF 1.8 MB

Smith, R. M. H. and J. Botha-Brink. 2014. Sedimentological and taphonomic evidence for drought-induced die-offs during the Permo-Triassic mass extinction in the main Karoo Basin, South Africa. Palaeogeography, Palaeoclimatology, Palaeoecology 312: 40-53.  PDF 2.9 MB SUPP 155 KB

Huttenlocker, A. K. and J Botha-Brink. 2014. Growth patterns and the evolution of bone microstructure in permo-Triassic therocephalians (Amniota, Therapsida) of South Africa. PeerJ 2:e325; DOI 10.7717/peerj.325. PDF 1.1 MB SUPP 243 KB

Botha-Brink, J., A. K. Huttenlocker, and S. P. Modesto. 2014. Vertebrate Paleontology of Nooitgedacht 68: A Lystrosaurus maccaigi-rich Permo-Triassic boundary locality in South Africa in pp. 289-304. C. F. Kammerer, K. D. Angielczyk, and J. Fröbisch (eds.), The Early Evolutionary History of the Synapsida. Springer, Dordrecht, Netherlands. PDF 2.2 MB

MacDougall, M. J., S. P. Modesto and J Botha-Brink. 2013. The postcranial skeleton of the Early Triassic parareptile Sauropareion anoplus, with a discussion of possible life history. Acta Palaeontologica Polonica 58(4).

Ruta, M., J. Botha-Brink, S. A. Mitchell and M. J. Benton. 2013. The radiation of cynodonts and the ground plan of mammalian morphological diversity. Proceedings of the Royal Society B 280. view linkPDF 1.2 MB  SUPP Methods & Results  SUPP files S1-S12

Huttenlocker, A. K. and J. Botha-Brink. 2013. Body size and growth patterns in the therocephalian Moschorhinus kitchingi (Therapsida: Eutheriodontia) before and after the end-Permian extinction in South Africa. Paleobiology 39: 253-277. PDF on request

Botha-Brink, J. and R. M. H. Smith. 2012. Palaeobiology of Triassic procolophonids inferred from bone microstructure. Comptes Rendus Palevol 11: 419-433. PDF 3.5 MB

Botha-Brink, J., F. Abdala and A. Chinsamy. 2012. Paleobiology and radiation of Permo-Jurassic non-mammaliaform cynodonts; pp. 223-246 in A. Chinsamy-Turan (ed.), The biology of non-mammalian therapsids: insights from bone microstructure. Indiana University Press, Bloomington, 360 pp.

Ray, S., J. Botha-Brink, and A. Chinsamy. 2012. Dicynodont growth dynamics and lifestyle adaptations; pp. 121-146 in A. Chinsamy-Turan (ed.), The biology of non-mammalian therapsids: insights from bone microstructure. Indiana University Press, Bloomington, 360 pp.

Smith, R. M. H. and J. Botha-Brink. 2011. Morphology and composition of bone-bearing coprolites from the Late Permian Beaufort Group, Karoo Basin, South Africa. Palaeogeography, Palaeoclimatology, Palaeoecology 312: 40-53. PDF on request

Botha-Brink, J and R. M. H. Smith. 2011. Osteohistology of the Triassic archosauromorphs Prolacerta, Proterosuchus, Erythrosuchus and Euparkeria from the Karoo Basin of South Africa. Journal of Vertebrate Paleontology 31(6): 1238-1254. PDF 1.68 MB

Botha-Brink, J. and S. P. Modesto. 2011. A new specimen of the therocephalian synapsid Olivierosuchus parringtoni from the Lower Triassic South African Karoo Basin. Palaeontology 54: 591-606.  PDF 1.3 MB

Botha-Brink, J. and K. D. Angielczyk. 2010. Do extraordinarily high growth rates in Permo-Triassic dicynodonts (Therapsida, Anomodontia) explain their success before and after the end-Permian extinction? Zoological Journal of the Linnean Society 160: 341-365 PDF on request

Botha-Brink, J. 2010. Comments on the proposed precedence of Procynosuchus Broom, 1937 (Therapsida, Cynodontia) over Cyrbasiodon Broom, 1931 and Parathrinaxodon Parrington, 1936.Bulletin of Zoological Nomenclature 67(1): 95. PDF 46 KB

Modesto, S. P., D. M. Scott, J. Botha-Brink and R. R. Reisz. 2010. A new and unusual procolophonid parareptile from the Lower Triassic Katberg Formation of South Africa. Journal of Vertebrate Paleontology 30(3): 715-723. PDF 3.3Mb

Modesto, S. P. and J. Botha-Brink. 2010. A burrow cast with Lystrosaurus skeletal remains from the Lower Triassic of South Africa. Journal of Vertebrate Paleontology. 25: 274-281.PDF 2.7Mb

Modesto, S. P. and J. Botha-Brink. 2010. Problems of correlation of South African and South American tetrapod faunas across the Permian-Triassic boundary. Journal of African Earth Sciences. 52: 242-248.PDF 619 KB

Botha-Brink, J. and S. P. Modesto. 2009. Anatomy and relationships of the Middle Permian varanopid Heleosaurus scholtzi based on a social aggregation from the Karoo Basin of South Africa. Journal of Vertebrate Paleontology. 29(2): 389-400. PDF 2.5Mb

Modesto, S. and J. Botha-Brink. 2008. A new archosauriform reptile from the Lower Triassic Lystrosaurus Assemblage Zone of South Africa. Journal of Vertebrate Paleontology 28(3): 914-917 PDF 445 KB

Botha-Brink, J. and F. Abdala. 2008. A new cynodont record from the Tropidostoma Assemblage Zone of the Beaufort Group: implications for the early diversity of cynodonts in South Africa. Palaeontologia Africana 43: 1-6.PDF 688 KB

Botha-Brink, J. and S. Modesto. 2007. A mixed-age classed ‘pelycosaur’ aggregation from South Africa: earliest evidence of parental care in amniotes? Proceedings of the Royal Society of London B274: 2829-2834.PDF 548 KB

Botha, J. and K. Angielczyk. 2007. An integrative approach to distinguishing the Late Permian dicynodont species Oudenodon bainii and Tropidostoma microtrema (Therapsida: Anomodontia). Palaeontology. 50(5): 1175-1209.PDF 1.4Mb

Botha, J. and T. Gaudin. 2007. An early Pliocene pangolin (Mammalia: Pholidota) from Langebaanweg, South Africa. Journal of Vertebrate Paleontology. 27(2): 484-491.PDF 721 KB

Botha, J., S. Modesto and R. Smith. 2007. A specimen of the procolophonoid reptile Sauropareionanoplus from the Katberg Formation of South Africa. South African Journal of Science. 103: 54-56.PDF 605 KB

Botha, J. and R. Smith. 2007Lystrosaurus species composition across the Permo-Triassic boundary of South Africa. Lethaia. 40(2): 125-137.PDF 421 KB

Botha, J., F. Abdala and R. Smith. 2007. The oldest cynodont: new clues on the origin and diversification of Cynodontia. Zoological Journal of the Linnean Society 149: 477-492.PDF 430 KB

Botha, J. and R. Smith. 2006. Rapid vertebrate recuperation in the Karoo Basin of South Africa following the end-Permian extinction. Journal of African Earth Sciences 45 (4-5): 502-514.PDF 993 KB

Smith, R. and J. Botha. 2005. The recovery of terrestrial vertebrate diversity in the South African Karoo Basin after the End-Permian extinction. Comptes Rendus Palevol 4: 555-568.PDF 635 KB

Botha, J., J. Lee-Thorp and A. Chinsamy. 2005. The palaeoecology of the non-mammalian cynodonts Diademodon  and Cynognathus from the Karoo Basin of South Africa, using stable light isotope analysis. Palaeogeography, Palaeoclimatology, Palaeoecology 223: 303-316.PDF 234 KB

Botha, J. and A. Chinsamy. 2005. Growth patterns of Thrinaxodon, a non-mammalian cynodont from the Early Triassic of South Africa. Palaeontology 48: 385-394.PDF 542 KB

Ward, P. D., J. Botha, R. Buick, M. O. de Kock, D. H. Erwin, G. Garrison, J. Kirschvink and R. Smith. 2005. Abrupt and gradual extinction among land vertebrates in South Africa: Evidence against a K/P-type impact extinction at the end of the Permian. Science 307: 709-713.PDF 371 KB

Botha, J. and A. Chinsamy. 2004. Growth and lifestyle adaptations of the Triassic non-mammalian cynodont Trirachodon. Acta Palaeontologica Polonica 49(4): 619-627.PDF 613 KB

Ray, S., J. Botha and A. Chinsamy. 2004. Bone histology and growth patterns of some non-mammalian therapsids. Journal of Vertebrate Paleontology 24(3): 634-648.PDF 849 KB

Botha, J., J. Lee-Thorp and M. Sponheimer. 2004. An examination of Triassic cynodont tooth enamel chemistry using Fourier Transform Infrared spectroscopy. Calcified Tissue International 74(2): 162-169.PDF 155 KB

Botha, J. 2003. Biological aspects of the Permian dicynodont Oudenodon (Therapsida, Dicynodontia), deduced from bone histology and cross-sectional geometry. Palaeontologia Africana 39: 37-44.PDF 3.4Mb

Botha, J. and A. Chinsamy. 2000. Growth patterns deduced from the bone histology of the cynodonts Diademodon  and Cynognathus. Journal of Vertebrate Paleontology 20 (4): 705-711.PDF 595 Kb

Publications - Popular Articles

Current Researhers

Botha, 2020. Our ancient ancestors the Therapsida. Culna

Botha, 2020. The biggest mass extinction in Earth’s history. Culna

Botha, 2020. New analysis sheds important light on an ancient mass extinction event. The Conversation.

Botha-Brink, J. 2016. How looking 250 million years into the past could save modern species. The Conversation.

Botha-Brink, J. 2011. A Diamond in the Rough. The Life Story of an Extraordinary Man. Culna 66: 32-33.

Botha-Brink, J. 2007. Survivor of the most catastrophic extinction. The Voyage of the Planet pp. 8-17.

Botha, J. 2006. A long time ago and not so far away – an ancient world in crisis? Culna 61 pp. 61-62.

Botha, J. 2004. Bizarre creatures from an ancient past. Getaway Magazine pp. 74-78.


The fossil collection predates the founding of the National Museum in 1877. The present vertebrate collection contains more than 3600 fossil specimens and includes unique material such as the tritheledontid cynodont Diarthrognathus, virtually all the material of the primitive dicynodont, Eodicynodon, the only known skull and skeleton of the early prosauropod dinosaur, Melanorosaurus, and a complete, fully articulated skeleton of the amphibian Uranocentrodon and the archosauriform Erythrosuchus. We also have more than 3000 invertebrate specimens, mainly consisting of Cretaceous ammonites, 126 specimens of fossil plants, which were collected mostly during the 1920s and 1950s, as well as more than 2000 mineral specimens, which include several meteorites. The fossil display, which is open to the public, consists mainly of animals that represent early relatives of mammals, namely the Therapsida, and are integral to understanding the origin of mammals.

Karoo vertebrates

The Karoo vertebrate collection comprises mostly therapsid and dinosaur fossils. The collection includes approximately:

  • 50 fish fossils including mostly Semionotus capensis
  • 100 amphibian fossils, including a complete, fully articulated skeleton of Uranocentrodon and several skulls and skeletons of Micropholis and Lydekkerina
  • 130 parareptile fossils, including several complete skulls and skeletons of Procolophon, Sauropareion and Saurodektes
  • 70 archosauromorph fossils including the holotype of Proterosuchus fergusi Broom 1903 and a complete fully articulated skeleton of Erythrosuchus africanus
  • 50 dinosaur fossils, including several Massospondylus carinatus specimens, one Fabrosaurus australis specimen, a partial skeleton of Euskelosaurus browni and the only known skull and skeleton of Melanorosaurus readi
  • 3285 therapsid fossils:
    • 30 dinocephalian specimens
    • 3020 dicynodont specimens
    • 60 gorgonopsian specimens
    • 30 therocephalian specimens, including a complete articulated skeleton of Theriognathus and several specimens of Olivierosuchus
    • 145 cynodont specimens, including several complete skulls and skeletons of Galesaurus and Thrinaxodon

126 plant fossils comprising mostly fossilized trees from Senekal

3100 invertebrate fossils comprising mostly ammonites, but also some trace fossils

2259 minerals, comprising mostly semi-precious stones

Type List

Phonodus dutoitorum Modesto, Scott, Botha-Brink and Reisz 2010

Proterosuchus fergusi Broom 1903 (Chasmatosaurus alexanderi Hoffman 1965)

Pedeticosaurus leviseuri van Hoepen 1915

Gryponyx africanus Haughton 1924

Melanorosaurus readi Haughton 1924 (Euskelosaurus browni Huxley 1866)

Eodicynodon oelofseni Rubidge 1990

Patranomodon nyaphulii Rubidge and Hopson 1990

Colobodectes cluveri Modesto, Rubidge and Welman 2003

Australosyodon nyaphuli Rubidge 1994

Oudenodon margaritae van Hoepen 1934

Compsodon helmoedi van Hoepen 1934

Lanthanostegus mohoii Modesto, Rubidge and Welman 2003

Langbergia modisei Abdala, Neveling and Welman 2006

Diarthrognathus broomi Crompton 1958 (holotype & paratype)

Services Offered


Our fossil preparators are fully trained in the casting of fossil vertebrates and traces. A list of available casts and models is given below:

Casts of fossils for sale at the National Museum

1GlossopterisPermian30 cm, several leaves on brown background
2DicroidiumTriassic21 cm, single leaf on grey background
3Deiradoceras ammoniteCretaceous26 cm, complete shell
4Semionotus (small)Triassic15 cm, fish in sandstone
5Semionotus (large)Triassic20 cm, fish in sandstone
6LydekkerinaTriassic12 cm, skull with palate and lower jaw
7LydekkerinaTriassic30 cm, articulated skeleton
8MesosaurusPermian28 cm, skeleton in grey-brown rock
9NanopariaPermian18 cm, skull with palate
10ProcolophonTriassic12 cm, skull and lower jaw
11PedeticosaurusTriassic31 cm, skull and skeleton
12ProterosuchusTriassic119 cm, skull and skeleton
13ProterosuchusTriassic31 cm, skull
14EuskelosuarusTriassic70 cm, redbrown femur
15EodicynodonPermian6 cm, skull and lower jaw
16EodicynodonPermian30 cm, skull and skeleton with lower jaw
17OudenodonPermian32 cm, skull and palate
18LystrosaurusTriassic12 cm, skull with palate and lower jaw
19LemurosaurusPermian15 cm, skull with lower jaw
20TheriognathusPermian19 cm, skull with palate
21MoschorhinusTriassic23 cm, skull with palate and lower jaw
22ScaloposaurusTriassic4 cm, skull
23OlivierosuchusTriassic20 cm, skull and partial skeleton
24ProcynosuchusPermian10 cm, skull with palate and lower jaw
25ProcynosuchusPermian42 cm, skull and skeleton
26ThrinaxodonTriassic6 cm, skull and lower jaw
27ThrinaxodonTriassic11 cm, skull with palate and lower jaw
28ThrinaxodonTriassic41 cm, skull and skeleton
29GalesaurusTriassic9 cm, skull with palate and lower jaw
30CynognathusTriassic26 cm, skull with palate, no teeth, lower jaw
 31DiademodonTriassic30 cm, skull with lower jaw

Models of fossils for sale at the National Museum
(approx. 25 cm)


Thin sectioning
The Department contains a fully functional bone histology laboratory for the thin sectioning and analysis of fossil bone microstructure.

Loan Policy

The acceptance of material on loan implies that the following conditions have been accepted:

  • Loan material must be handled with the utmost care and must be securely packed for return by courier.
  • National Museum specimens described as holotypes, paratypes, allotypes or some other category or type, or used to supplement the description of a new taxon or the redescription of an existing species, are to be clearly labeled as such, and must be returned.
  • All labels and numbers given to specimens/objects must be retained and under no circumstances should any of these be discarded or altered. Where applicable the catalogue numbers of specimens/objects must be given in descriptions.
  • The National Museum must be credited in any publication when referring to this material. At least two copies of any such publication must be sent to this Museum.
  • Without express permission no specimen or object may be altered, dissected, prepared, restored, cleaned or treated — either by mechanical or chemical means.
  • Casting of specimens/objects is only allowed under specific conditions. Permission must be obtained beforehand.
PSSA Conference

Palaeontological Society of Southern Africa Biennial Meeting 2018

July 4th to July 6th 2018



It is our pleasure to invite you to the 20th Palaeontological Society of Southern Africa Biennial Meeting to be hosted by the National Museum, Bloemfontein from 4 July to 6 July 2018.

Please see the second circular here (download here).

Please see the third circular here (download here).

PSSA 2018 Registration Forms are now available (download here).

Please find the PSSA programme and abstracts here.
Do not print as you will receive a printed volume at the conference.

Please see details about the pre-conference workshop (download here)

Notices are also up on the PSSA website and Facebook, and will continue to be distributed via membership lists. Please forward the circular to your networks.Any queries can be directed to the Program Chair, Jennifer Botha-Brink at

Please check this site regularly for updates.

We look forward to seeing you here!

Dr Jennifer Botha-Brink
Karoo Palaeontology Department
Program Chair

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