Florisbad Quaternary Research

Florisbad Quaternary Research

Quaternary Fossils

The Florisbad Quaternary Research Department was established with the acquisition of the Florisbad fossil site in 1980, with the initial aim of excavating the Florisbad site and developing on-site research infrastructure. In March 1984, with the departure of Dr Ron Clarke, who was the scientist initially appointed, the research aim of the Department was redefined to be that of an interdisciplinary Quaternary research unit. However, in time comparative mammal osteology, as applied to the study of Quaternary mammal faunas, has become the main research focus of the Department.


Oppenheimer De Beers Group Research Conference attended

Oppenheimer De Beers
Dr Daryl Codron receiving the Best Poster Award from Mr Nicky Oppenheimer of E. Oppenheimer & Son.

11 January 2017
Drs Nico Avenant (Mammalogy Department), Daryl Codron (Quarternary Research) and their post-Doc student Jacqui Codron attended the 7th annual Oppenheimer De Beers Group Research Conference in October in Johannesburg.  They were involved in three separate contributions at the conference.  Daryl won the Best Poster Award.

Conference Paying Tribute to Prof Louis Scott

26 September 2014
From 07 to 11 July, the Florisbad Department, together with the Archaeology and Karoo Palaeontology Departments, and the University of the Free State (UFS), hosted an international palaeoclimate conference - “From Past to Present: Changing Climates, Ecosystems and Environments of southern Africa – A Tribute to Louis Scott”. The conference was organized as a tribute to Professor Louis Scott, who recently retired from the UFS’s Department of Plant Sciences. Louis is globally recognized as one of the leading scholars in Quaternary Palaeobotany and Palaeoecology, having made an outstanding contribution to teaching and research in South Africa and on the international front. His pioneering research in the fields of palynology, long-term continental environmental change, and interpretation of palaeoenvironmental records associated with archaeological sites, has contributed insights into the origin of our current environment by identifying long-term patterns of climate change. Results of his studies have been applied in numerical models of vegetation change in Africa and globally, being relevant across the fields of botany, geology, climatology, archaeology, anthropology and palaeontology.

The conference was attended by leading South African and international scholars, and covered topics as diverse as ecosystems ecology, palaeoenvironmental change, and archaeology. Proceedings are to be published in the journals Palaeoecology of Africa and the Transactions of the Royal Society of Africa. For more information, visit www.lscott-tribute.co.za.

Postdoctoral Research Fellow

Michael Toffolo

27 March 2014
Michael Toffolois an archaeologist, especially interested in geoarchaeology (i.e. the study of the formation and post-depositional processes of a site) and microarchaeology (i.e. the study of the microscopic archaeological record – whatever requires instruments in order to be seen). He received his BA (2006) and MA (2009) in Archaeology from the University Of Padua, Italy, where he worked at several Roman and Medieval archaeological sites. Michael subsequently started a PhD in Archaeology at Tel Aviv University, Israel, which was completed in 2014. During this study he carried out fieldwork at major Iron Age sites such as Megiddo, Ashkelon, Tell es-Safi/Gath, Tel Dor (all located in Israel), Lefkandi, Kalapodi, Corinth (Greece) and Idalion (Cyprus). He also worked at Upper Paleolithic sites (Manot and BokerTachtit – Israel). All his laboratory work was carried out as a Visiting PhD student at the Kimmel Center for Archaeological Science, Weizmann Institute (Israel). Michael’s dissertation is focused on the radiocarbon chronology of the southern Levant and Aegean during the Iron Age, with special attention to the characterization of archaeological contexts using a microarchaeological approach, which involves the application of several analytical techniques: archaeological soil micromorphology, Fourier-Transform Infrared spectrometry and micro-spectrometry (FTIR), Raman microscopy, quantification of soil free phosphates, phytolith analysis, Scanning Electron Microscopy coupled with Energy Dispersive Spectrometry (SEM-EDS) and Thermo-Gravimetric Analysis (TGA). He routinely uses this approach in the study of Eastern Mediterranean Late Bronze and Iron Age chronology; household activities in Mediterranean Late Bronze and Iron Age tell sites; pyrogenic aragonite and its importance in ancient pyrotechnology and in radiocarbon dating of archaeological ash and plaster.

This postdoctoral fellowship forms part of a National Research Foundation (NRF) African Origins Platform (AOP) research grant to James Brink for a project titled “Early and Middle Pleistocene evolution of large mammal faunas and modern landscapes in southern Africa”. Michael is in charge of studying the formation and post-depositional processes of the Middle to Late Pleistocene sedimentary deposits of the Florisbad site, and the sedimentary sequence of the end-Early Pleistocene site of Cornelia-Uitzoek.

Complete anterior part of an extinct equid


23 February 2009
The final preparation of the excavation of a complete anterior part of an extinct equid from the Modder River site of Erfkroon. The specimen was discovered by Britt Bousman, Dept. of Anthropology, Texas State University, and excavated during October 2008. Here Mr Isaac Thapo is preparing the removal of the left forelimb.


Senior Specialist Museum Scientist and Head of Department

James S. Brink DPhil arjbrink@nasmus.co.za

James-BrinkIn 1978 James Brink received a BA degree (Archaeology, Anthropology & Latin) and in 1980 BA Honours (Archaeozoology) from the University of Stellenbosch. He was appointed in 1983 as a junior Researcher in Archaeozoology at the National Museum. Based at the newly established Florisbad Quaternary Research Station, he studied the fossil animal remains from the new excavations at Florisbad, as well as the old fossil collections from this site. He completed this study in 1987, for which he received the degree MA (Archaeozoology) from the University of Stellenbosch and the Frank Schweitzer postgraduate student prize (Universities of Cape Town and Stellenbosch). He also received study grants from the Human Sciences Research Council (now incorporated in the National Research Foundation, NRF) and from the German Foreign Exchange program (DAAD) to study at a German University. In 1987 and 1988 he completed a two-semester course at the Institut für Palaeoanatomie, Faculty of Veterinary Science, University of Munich, which included courses on the comparative osteology of Old World mammals, the history of domestication in the Old World, the comparative osteology of Nile fishes and an introductory course in Latin and Greek as used in scientific nomenclature. In 1992 he received the degree BA Honours in Latin from the University of the Free State, while preparing for a comparative osteological study of alcelaphine bovids and the evolution of wildebeest. Through funding received from the Centre for Science Development (CSD, now part of the NRF) he invited Prof. R. Grün (Canberra) to do a comprehensive Electron Spin Resonance / Optically Stimulated Luminescence (ESR/OSL) study of the Florisbad deposits and fossils, including the human skull. In the process a pioneering technique of non-destructive ESR dating was developed in order to date the molar associated with the human skull. This work, published in 1996, provided the temporal framework needed for studying the alcelaphine fossil record in southern Africa. In 2001, as part of this study, he worked for two months in the Palaeontology Laboratory of the National Museum of Natural History, Paris, on North African alcelaphine fossils. In 2005 he received the degree DPhil from the University of Stellenbosch for his study on the evolution of the black wildebeest (Connochaetes gnou) and modern large mammal faunas in southern Africa. James is currently expanding his interests to the comparison of the large mammal record of East Africa with that of southern Africa. He is a NRF rated scientist, with a B3 rating.

Since March 1984 James has been in charge of the research activities at the Florisbad Quaternary Research Station. In this time he oversaw the founding of an extensive osteological reference collection and the expansion of the Quaternary fossil vertebrate collections through field work at key sites, such as Florisbad, Cornelia-Uitzoek and Erfkroon.

Principal Museum Scientist

Daryl Codron PhD d.codron@nasmus.co.za

Daryl CodronDaryl is an animal ecologist, with specialist expertise in stable isotope analysis and applications of this technique to ecology and palaeoecology. Based on research relying largely on stable isotope approaches, he was awarded a PhD degree from the Department of Archaeology, University of Cape Town (UCT) in 2006, for his studies on the trophic dynamics of savanna ungulates and the implications for evolutionary diversification of this group. He had earlier received an MSc degree in Quaternary Science, from UCT in 2003, for similar research on baboons as extant analogues for interpreting stable isotope patterns of early hominins. Though much of Daryl's postgraduate research was within the broader discipline of palaeoscience, his initial training was in biology. In 1998 he earned a BSc degree in Zoology and Botany from the Univeristy of Johannesburg (formerly the Rand Afrikaans University), and in 2001 earned a BSc Honours degree in Biodiversity and Conservation from the same institution. This diversity in training resulted in him being awarded several postdoctoral fellowships over the years. In 2006/07 he was appointed as postdoctoral fellow of the Florisbad Quaternary Research Department of the National Museum (the first such appointment within the National Museum), where he conducted research on the evolution of landscapes and mammalian faunas of the South African central interior through the Late Quaternary. Thereafter, he was appointed as full-time lecturer in animal ecology at the School of Biological and Conservation Sciences, University of KwaZulu-Natal, until 2009. Following that, Daryl was awarded a Marie Curie fellowship from the European Union, to conduct experimental stable isotope research hosted at the University of Zürich (UZ), Switzerland for two years. Following successful completion of the Marie Curie fellowship, he was awarded a further two-year postdoctoral position at UZ, funded by the Deutsche Forschungsgemeinschaft Research Unit 533 “The Biology of Sauropod Dinosaurs”.

Daryl's research interests span a variety of ecological and evolutionary sub-disciplines, including behavioural and evolutionary ecology, community ecology, and macroecology, all with implications for advancing our understanding of large mammal evolution particular within African savanna landscapes. Stable isotope analysis remains at the core of most of this research, and many of his studies focus on key issues regarding the most fundamental principles of this technique. The combination of ecological and palaeontological skills within his repertoire has made it possible for Daryl to explore novel questions about the processes driving macroevolutionary patterns observed in the South African mammal fossil record. The strengths of this skill set were well-received by his peers, and accordingly Daryl was awarded a C1 NRF rating as of January 2014.

Current research projects:

A conceptual model for interpreting variability in isotope niche breadths of fossil populations
Does C4 grass limit the diversity of intermediate-feeding ruminants in Africa?
Factors influencing stable isotope distributions and abundances in African savanna and grassland ecosystems
Life histories, ecology, and community dynamics of terrestrial vertebrates across the Permian-Triassic extinction
Daryl is a member of the Bloemfontein Palaeosystems Centre (www.palaeosystems.co.za), an inter-departmental partnership within the Museum, and including members from the University of the Free State.

Chief Research Assistant

Sharon-HoltSharon Holt MSc sholt@nasmus.co.za

Sharon Holt obtained her BA degree at the University of Pretoria in 1991 and her BA Honours (Archaeology) in 1995. She obtained her MSc degree in Archaeology at the University of the Witwatersrand in 2009. Thesis title: The faunal remains from the Makgabeng Plateau, Limpopo Province. Her career started as a Research Assistant in the Archaeology Department of the National Museum in 1992. In 2006 she was transferred to the Florisbad Quaternary Research Department, where she is currently curating the modern mammal osteological collection housed at Florisbad.

General Assistants

Abel Dichakane

Isaac Thapo

Jacob Maine
Jacob Dintine Maine


The research focus of the Department is the evolution of large mammal faunas and the appearance of modern environments in central southern Africa. For this purpose we use the fossil mammal collections and the modern osteology collections, both housed at the Florisbad Quaternary Research Station. Florisbad is the type locality of the Florisian Land Mammal Age and the site is the base from where we conduct fieldwork in the central interior. We excavate at various sites, including Cornelia-Uitzoek (the type locality of the Cornelian Land Mammal Age) and at Erfkroon, a Florisian locality on the Modder River.

The Department is a participating member of the Bloemfontein Palaeosystems Centre (BPC).

Publications - Scientific Articles

Van Zyl, W., S. Badenhorst & J.S. Brink. 2016. Pleistocene Bovidae from X Cave on Bolt’s Farm in the Cradle of Humankind in South Africa. Annals of the Ditsong National Museum of Natural History 6: 39–73.

Stone, O.M.L. A.I.R. Herries, J.S. Brink, & S.W. Laffan.2016. The chacma baboon (Papio ursinus) through time: A model of potential core habitat regions during a glacial-interglacial cycle. Evolutionary Ecology30:755–782.

Brink, J.S. 2016. Faunal evidence for mid- and late Quaternary environmental change in southern Africa. In: Knight, J. and Grab, S.W. (eds) Quaternary environmental change in southern Africa: physical and human dimensions. Cambridge University Press, pp. 286-307 .

Brink, J.S., S. Holt & L. Kolska Horwitz. 2016. The Oldowan and early Acheulean Mammalian Fauna of Wonderwerk Cave (Northern Cape Province, South Africa). African Archaeological Review.33: 223–250

Leichliter, J.N., Sponheimer, M., Avenant, N.L., Sandberg, P.A., Paine, O.C.C., Codron, D., Codron, J., Passey, B.H. 2016. Small mammal insectivore stable carbon isotope compositions as habitat proxies in a South African savanna ecosystem. J Archaeol Sci Reports 8:335-345

Botha-Brink, J., Codron, D., Huttenlocker, A., Angielczyk, K.D., Ruta, M. 2016. Breeding young  as a survival strategy during Earth’s greatest mass extinction. Sci Reports. 6:24053

Codron, D., Codron, J., Sponheimer, M., Clauss, M. 2016. Within-population isotopic niche variability in savanna mammals: disparity between carnivores and herbivores. Frontiers Ecol Evol. 4: doi:10.3389/fevo.2016.00015

Clauss, M., Fritz, J., Tschuor, A., Braun, U., Hummel, J., Codron, D. 2016. Dry matter and digesta particle size gradients along the goat digestive tract on grass and browse diets. J Anim Physiol Anim Nutr. doi:10.1111/jpn.12505

Brink, J.S., S. Holt & L. Kolska Horwitz. 2015. Preliminary Findings on Macro-Faunal Taxonomy, Taphonomy,Biochronology and Palaeoecology from the Basal Layers of Wonderwerk Cave, South Africa. In I. Thiaw & H. Bocoum (Eds.),Dakar: Memoires de lIFAN - CA. DIOP (93, 137-147).Preserving African Cultural Heritage.(Proceedings of the 13th Congress of the Panafrican Archaeological Association for Prehistory and Related Studies – PAA and of the 20th Meeting of the Society of Africanist Archaeologists – Safa).

Gowlett, J.A.J., J S. Brink, Andy A.I.R. Herries, S. Hoare, I. Onjala & S.M. Rucina. 2015. At the heart of the African Acheulean: the physical, social and cognitive landscapes of Kilombe. In. Coward, F., Hosfield, R., Pope, M. & Wenban-Smith, F.  (eds.), Settlement, society and cognition in human evolution: Landscapes in mind. Cambridge University Press, pp. 75-93.

Smith, P., J. S. Brink, J. W. Hoffman, L.C. Bam, R. Nshimirimana & F.C. de Beer. 2015. The late Middle Pleistocene upper third molar from Florisbad: metrics and morphology. Transactions of the Royal Society of South Africa, 70: 233 -244

Toffolo, M.B., J.S. Brink, F. Berna. 2015. Bone diagenesis at the Florisbad spring site, Free State Province (South Africa): implications for the taphonomy of the Middle and Late Pleistocene faunal assemblages. Journal of Archaeological Science: Reports, 4:152–163.

Brink, J.S., C.B. Bousman & R. Grün. 2015.  A reconstruction of the skull of Megalotragus priscus(Broom, 1909), based on a find from Erfkroon, Modder River, South Africa, with notes on the chronology and biogeography of the species. Palaeoecology of Africa, 33:71-94.

Van Aardt, A., C.B. Bousman, J.S. Brink, G.A. Brook, Z. Jacobs, P.J. du Preez,L. Rossouw and L. Scott. 2015. Baden-Baden: A fossil spring site in the western Free State, South Africa Palaeoecology of Africa, 33: 117-152.

Codron, J., Duffy, K.J., Avenant, N.L., Sponheimer, M., Leichliter, J., Paine O, Sandberg, P., Codron, D. 2015. Stable isotope evidence for trophic niche partitioning in a South African savanna rodent community. Curr Zool. 61: 397-411

Müller, J., Clauss,  M., Codron, D., Schulz, E., Hummel, J., Kircher, P., Hatt, J.-M. 2015. Tooth length and incisal wear and growth in guinea pigs (Cavia porcellus) fed diets of different abrasiveness.J Anim Physiol Anim Nutr. 99:591-604

Backwell, L.R., T. S. McCarthy, L.Wadley, Z. Henderson, C. M. Steininger, B. deKlerk,M. Barré, M.Lamothe, B. M. Chase,S. Woodborne, G.J. Susino, M.K. Bamford, C.Sievers, J.S. Brink, L.Rossouw, L.Pollarolo, G.Trower, L. Scott, F. d’Errico. 2014. Multiproxy record of late Quaternary climate change and Middle Stone Age human occupation at Wonderkrater, South Africa.Quaternary Science Reviews. 99: 42–59.

Carbone, C,, Codron, D., Scofield, C., Clauss, M., Bielby, J. 2014. Geometric factors influencing the diet of vertebrate predators in marine and terrestrial environments. Ecol Lett. 17: 1553-1559

Harris, J.A., M. Bar-Matthews, J.S. Brink, K. Braun, E. Fisher, Z. Jacobs, P. Karkanas & C. W. Marean. 2014. Herolds Bay Cave: New Evidence for Middle Stone Age Coastal Adaptations in Cape Province, South Africa "Paleoanthropology Society Meeting Abstracts, Calgary, Canada, 8-9 April 2014" PaleoAnthropology 2014:A10-A11.

Müller, J., M. Clauss, D. Codron, E. Schulz, J. Hummel, M. Fortelius, P. Kircher & J.-M. Hatt. 2014.Growth and wear of incisor and cheek teeth in domestic rabbits (Oryctolagus cuniculus) fed diets of different abrasiveness. J Exp Zool. 321A: 283-298

Clauss, M., M.T. Dittmann, D.W.H. Müller, P. Zerbe & D. Codron. 2014. Low scaling of a life history variable: analysing eutherian gestation periods with and without phylogeny-informed statistics. Mamm Biol. 79: 9-16

Balsiger, A., S. Keller, I. Espie, E.P. Lane, A. Kotze, M. Clauss & D. Codron. 2014. Reproductive seasonality and newborn mortality at the National Zoological Gardens of South Africa. Proceedings of the International Conference on Diseases of Zoo and Wild Animals, p 102

Gattiker, C., I. Espie, E.P. Lane, A. Kotze, D. Codron & M. Clauss. 2014. Correlations of diet composition and diet-related disorders in a collection of captive wild ruminants at the National Zoological Gardens of South Africa. Proceedings of the International Conference on Diseases of Zoo and Wild Animals, p 89

Germitsch, N., P.-R. Vybiral, D. Codron, M. Clauss, A. Kotze & E.P. Lane. 2014. Adrenal mass as a retrospective indicator of health problems in captive cheetahs (Acinonyx jubatus) at the Ann van Dyk Cheetah Centre, South Africa. Proceedings of the International Conference on Diseases of Zoo and Wild Animals, p 138

Hagen, K., I. Espie, E.P. Lane, A. Kotze, F. Wyss, M. Clauss & D. Codron. 2014. Mortality and diseases in greater flamingo (Phoenicopterus roseus) and Carribean flamingo (Phoenicopterus ruber) kept at the National Zoological Garderns of South Africa. Proceedings of the International Conference on Diseases of Zoo and Wild Animals, p 57-64

Keller, S., A. Balsiger, I. Espie, E.P. Lane, A. Kotze, M. Clauss & D. Codron. 2014. Density-dependent effect on mortality in 42 zoo animal species at the National Zoological Gardens of South Africa. Proceedings of the International Conference on Diseases of Zoo and Wild Animals, p 56

Vybiral, P.-R., N. Germitsch, D. Codron, M. Clauss, A. Kotze & E.P. Lane. 2014. Neonatal mortality in captive cheetahs (Acinonyx jubatus) at the Ann van Dyk Cheetah Centre, South Africa. Proceedings of the International Conference on Diseases of Zoo and Wild Animals, p 215

Taylor, L.A., D.W.H. Müller, C. Schwitzer, T.M. Kaiser, D. Codron, E. Schulz & M. Clauss. 2014.Tooth wear in captive rhinos differs from that of free-ranging conspecifics. Contributions to Zoology 83: 107-117

Taylor, L.A., T.M. Kaiser, C. Schwitzer, D.W.H. Müller, D. Codron, M. Clauss & E. Schulz. 2013.Detecting inter-cusp and inter-tooth wear patterns in rhinocerotids. Plos One. 8:e80921

Clauss, M., P. Steuer, D.W.H. Müller, D. Codron & J. Hummel. 2013. Herbivory and body size: allometries of diet quality and gastrointestinal physiology, and implications for herbivore ecology and dinosaur gigantism. PloS One. 8:e68714

Brink, J.S., F. De Beer, J. Hoffman & L. Bam. 2013. The evolutionary meaning of Raphicerus-like morphology in the dentitions and postcrania of Antidorcas bondi (Antilopini). Zitteliana vol. 31 (Series B): 21.

Müller, D.W.H., D. Codron, C. Meloro, A. Munn, A. Schwarm, J. Hummel & M. Clauss. 2013.Assessing the Jarman–Bell Principle: scaling of intake, digestibility, retention time and gut fill with body mass in mammalian herbivores. Comp Biochem Physiol A. 164:129-140

Codron, J., J.A. Lee-Thorp, M. Sponheimer, & D. Codron. 2013. Plant stable isotope composition across habitat gradients in a semi-arid savanna: implications for environmental reconstruction. J Quat Sci. 28:301-310

Codron, D., Carbone, C., Müller, D.W.H., Clauss, M. 2013. Ecological modelling, size distributions and taphonomic size bias in dinosaur faunas: reply to Brown et al.Biol Lett. 9:20120922. doi:10.1098/rsbl.2012.0922

Codron, D., C. Carbone & M. Clauss. 2013. Ecological interactions in dinosaur communities: Influences of small offspring and complex ontogenetic life histories. Plos One. 8:e77110

Djagoun, C.A.M.S., D. Codron, J. Sealy, G.A. Mensah & B. Sinsin. 2013. Stable carbon isotope analysis of the diets of West African bovids in Pendjari Biosphere Reserve, Northern Benin. S Afr J Wildl Res. 43:33-43

Codron, J., K. Kirkman, K.J. Duffy, M. Sponheimer, J.A. Lee-Thorp, A. Ganswindt, M. Clauss & D. Codron. 2013. Stable isotope turnover and variability in tail hairs of captive and free-ranging African elephants (Loxodonta africana) reveal dietary niche differences within populations. Can J Zool. 91:124-134

Clauss, M., M.T. Dittmann, D.W.H.Müller, C. Meloro & D. Codron. 2013. Bergmann’s rule in mammals: a cross-species interspecific pattern. Oikos 122:1465-1472

Lauper, M., I. Lechner, P. Barboza, W. Collins, J. Hummel, D. Codron & M. Clauss. 2013. Rumination of different-sized particles in muskoxen (Ovibos moschatus) and moose (Alces alces) on grass and browse diets, and implications for rumination in different ruminant feeding types. Mamm Biol. 78:142-152

Berna, F., P. Goldberg, L.K. Horwitz, J.S. Brink, S. Holt, M. Bamford & M. Chazan. 2012.Microstratigraphic evidence of in situ fire in the Acheulean strata of Wonderwerk Cave, Northern Cape province, South Africa. Proc. Natl. Acad. Sci. U.S.A. 109:E1215-E1220

Brink, J.S., A.I.R. Herries, J. Moggi-Cecchi, J.A.J. Gowlett, C.B. Bousman, P.J. Hancox, R. Grun, V. Eisenmann, J.W. Adams & L. Rossouw. 2012. First hominine remains from a ~1.0 million year old bone bed at Cornelia-Uitzoek, Free State Province, South Africa. J Hum Evol.E 63:527-535

Brink, J.S. 2012. The post-1.0 Ma evolution of large mammal endemism in southern Africa in relation to East Africa and subsequent biogeographic isolation of the Cape coastal region. Quaternary International vol. 279–280: 69.

Chazan, M.,  D.M. Avery, M. Bamford,F. Berna, J.S. Brink, S. Holt, Y. Fernandez-Jalvo,  P. Goldberg, A. Matmon, N. Porat, H. Ron, L. Rossouw, L. Scott and L. Kolska Horwitz. 2012. The Oldowan horizon in Wonderwerk Cave (South Africa): archaeological, geological, paleontological and paleoclimatic evidenceJournal of Human Evolution63: 859-866.

Brink, J.S. 2012. The fauna from Lithakong. Appendix in: Kaplan, J. & Mitchell, P. The archaeology of the Lesotho Highlands Water Project Phases IA and IB. Southern African Humanities24: 30 -32.

Meier, H., Bousman, C.B., Brink, J.S., Bateman, M.D., Trower, G., Grün, R., Codron, D., Rossouw, L., Ramsey, C. and Scott, L. 2012. The Middle and Late Pleistocene alluvial terraces with Middle and Later Stone Age occupations in the Modder River valley, South Africa. Geological Society of America Abstracts with Programs.Vol. 44, No. 7, p.569.

Codron, D., Carbone, C., Müller D.W.H. & Clauss, M. 2012. Ontogenetic niche shifts in dinosaurs influenced size, diversity and extinction in terrestrial vertebrates. Biol Lett. 8:620-623

Codron, D., M. Sponheimer, J. Codron, I. Newton, J.L. Lanham & M. Clauss.  2012. The confounding effects of source isotopic heterogeneity on consumer-diet and tissue-tissue stable isotope relationships. Oecologia 169:939-953

Zerbe, P. M. Clauss, D. Codron, L.B. Lackey, E. Rensch, W.J. Streich, J.-M. Hatt & D.W.H. Müller. 2012. Reproductive seasonality in captive wild ruminants: implications for biogeographical adaptation, photoperiodic control, and life history. Biol Rev. 87:965-990

Codron, D. J. Hull, J.S. Brink, J. Codron, D. Ward & M. Clauss. 2011. Influence of competition on nichedynamics of syntopic grazing ungulates: contrasting the predictions of habitat selection models using stable isotope analysis. Evol Ecol Res. 13:217-235

Herries, A.I.R., S. Davies, J.S. Brink, D. Curnoe, G. Warr, M. Hill, S. Rucina, I. Onjala, J. A.J. Gowlett. 2011. New Explorations and Preliminary Magnetobiostratigraphical Analysis of the Kilombe Acheulian Locality, Central Rift, Kenya. "Abstracts of the Paleoanthropology Society 2011 Meeting." PaleoAnthropology 2011: A16

Manegold, A. & J.S. Brink. 2011. Descriptions and palaeoecological implications of bird remains from the Middle Pleistocene of Florisbad, South Africa. Palaeontologische Zeitschrift  85:19–32.

Manegold, A. & J.S. Brink. 2011. A chat (Saxicolinae, Muscicapidae) from the mid-Holocene of Florisbad, South AfricaOstrich  82(1): 57–63.

Lewis, P., J.S. Brink, A. Kennedy & T. Campbell. 2011. Examination of the Florisbad microvertebrates. South African Journal of Science 107 (7/8): 64-67. 

Grobler, J.P., I. Rushworth, J. S. Brink, P. Bloomer,, A. Kotze, B. Reilly, S. Vrahimis. 2011.Management of hybridization in an endemic species: decision making in the face of imperfect information in the case of the black wildebeest — Connochaetes gnou.  European Journal of Wildlife Research57 (5): 997-1006.

Bousman, B., J.S. Brink, S. Tooth, L. Rossouw, L. Scott, R. Grün, S. Barnett, M Espino & G. Trower. 2010. Middle and Later Stone Age Occupations Dating to Marine Isotope Stage 3 and 4 at Erfkroon, South Africa. Abstracts of the PaleoAnthropology Society 2010 Meeting." PaleoAnthropology:A4 & A5.

Ackermann, R.R., J.S. Brink, S. Vrahimis & B. de Klerk. 2010. Hybrid wildebeest (Artiodactyla: Bovidae) provide further evidence for shared signatures of admixture in mammalian crania. S. Afr. J. Sci. 106: 1-5.

Fernandez-Jalvo, Y., L. Scott, J.S. Carrion, G. Gil-Romera, J.S Brink, F. Neumann & L. Rossouw. 2010. Pollen taphonomy of hyaena coprolites: an experimental approach. Zona Arqueologica 13: 148-156.

Curnoe, D. & J.S. Brink. 2010. Palaeopathology of the Florisbad Cranium. Journal of Human Evolution59: 504-513

Codron, D., J. Codron J. Lee-Thorp, M. Sponheimer, C.C. Grant & J.S. Brink. 2009. Stable isotope evidence for nutritional stress, competition and loss of functional habitat as factors in limiting recovery of rare antelope in southern Africa. Journal of Arid Environments 73: 449-457.

Herries, A.I.R., J.S. Brink, B. Bousman, V. Eisenmann, J. Gowlett, R. Grün, J. Hancox, J. Moggi-Cecchi, L. Rossouw. 2009. A palaeomagnetic age of about 990,000 years for the Cornelia-Uitzoek fossil vertebrate, hominin and Acheulean site. South Africa. American Journal of Physical Anthropology, 138 (S48): 149.

Herries, A.I.R., Adams, J., Braun, D., Brink, J., Brown, K., Curnoe, D., Fisher, E, Marean, C., Roberts, D. & Warr, G. 2009. Palaeomagnetism through space and time in Africa: dating landmarks in hominin and mammalian evolution. Geological Society of Australia Abstracts 93: 36.

Brink, J.S. & D.D. Stynder. 2009. Morphological and trophic distinction in the dentitions of two early alcelaphine bovids from Langebaanweg (genus Damalacra). Palaeontologia Africana 44: 148-150.

Brink,J.S.,B. Bousman & R. Grün. 2009. The Interior of Southern Africa as a Unique habitat and Evolutionary Source Area for Periodic large mammal and human Dispersals During the middle and late Pleistocene. Abstracts of the PaleoAnthropology Society 2009 Meetings..PaleoAnthropology2009: A6

Bousman, B., J.S. Brink, G. Trower, S. Tooth, S. Woodbourne, R. Grün, L. Rossouw, L. Scott & H. Meier. 2009. Notes From the Interior: Recent Archaeological and Paleontological Investigations at Erfkroon, South AfricaPaleoAnthropology 2009: A5.

Brophy, J., D. de Ruiter & J.S. Brink. 2009. Mimosa: a new Middle Stone Age fossil locality in the Free State, South Africa. American Journal of Physical Anthropology, 138 (S48): 98.

Brink, J.S. 2008. Vertebrate remains from Sandy Bay Valley, St Helena Island, South Atlantic. in: Colin A. Lewis. The Late Glacial and Holocene avifauna of the island of St Helena, South Atlantic Ocean. Transactions of the Royal Society of South Africa 63(2): 143 - 144.

Codron, D. J. S. Brink, L. Rossouw, M. Clauss, J. Codron, J.A. Lee-Thorp & M. Sponheimer. 2008. Functional differentiation of African grazing ruminants: an example of specialized adaptations to very small changes in diet. Biological Journal of the Linnean Society 94: 755–764.

Codron, D., J.S. Brink, L. Rossouw  & M. Clauss. 2008. The evolution of ecological specialization in southern African ungulates: competition- or physical environmental turnover? Oikos 117: 344-353.

Codron, D., J. Lee-Thorp, M. Sponheimer, J. Codron, D. de Ruiter & J.S. Brink. 2007. Significance of diet type and diet quality for ecological diversity of African ungulates. Journal of Animal  Ecology 76: 526–537.

Herries, A., D. Curnoe, J.S. Brink, Z.L. Henderson, D. Morris, K. Van Ryneveld & E. Hodge. 2007.Landscape evolution, palaeoclimate and Later Stone Age occupation of the Ghaap Plateau escarpment, Northern Cape Province, South Africa. Antiquity Vol 81 (313).

Codron, D. & J.S. Brink. 2007. Trophic ecology of two savanna grazers, blue wildebeest Connochaetes taurinus and black wildebeest Connochaetes gnouEuropean Journal of Wildlife Research 53: 90-99.

Codron, D., J. Codron, J.A. Lee-Thorp, M. Sponheimer, D. de Ruiter & J.S. Brink. 2007. Stable isotope characterization of mammalian predator–prey relationships in a South African savanna. Eur J Wildl Res. 53: 161-170.

Backwell, L.R., C.M. Steininger, J.S. Brink, J. Neveling, L. Rossouw & L. Pereira. 2007. Large mammal mass death accumulation in the Holocene of South Africa. Palaeontologia Africana. 42: 118.

Brink, J.S. 2007. Vicariance in coastal large ungulate populations during the Middle and Late Pleistocene in southern Africa. Palaeontologia Africana. 42: 120.

Curnoe, D., A. Herries, J.S. Brink, P. Hopley, K. van Reyneveld, Z. Henderson & D. Morris. 2007.Landscape evolution, palaeoclimate and Later Stone Age occupation of the Ghaap Plateau escarpment, Northern Cape Province, South Africa. Antiquity 81(3): 313.

Bousman, B., Brink, J. S., Tooth, S., Oksanen, E., Meier, H. 2007. Middle and LAter Stone Age occupations in a Late Pleistocene Terrace at Erfkroon, South Africa. Abstracts of the Society for American Archaeology 72nd Annual Meeting, April 25-29, Austin: 76.

Brink, J.S. R. Grün, L. Rossouw, D. Codron. 2007. Mid-Quaternary large mammal succession and extinction in southern Africa. Quaternary International 167-168: 46-47

Webley, L. & J.S. Brink. 2006/7. Ethnographic observations on the butchering of domestic stock amongst the descendants of Nama-speaking pastoralists in Namaqualand, northern Cape. Southern African Field Archaeology 15 & 16: 12 – 25.

Codron, D., Codron, J., Lee-Thorp, J.A., Sponheimer, M., de Ruiter, D, & Brink, J.S. 2006. Dietary variation in impala Aepyceros melampus recorded by carbon isotope composition of feces. Acta Zoologica Sinica 52(6): 1015-1025.

Curnoe, D., A. Herries, J.S. Brink, P. Hopley, K. van Reyneveld, Z. Henderson & D. Morris. 2006.Discovery of Middle Pleistocene fossils and stone tool-bearing deposits at Groot Kloof, Ghaap Escarpment, Northern Cape Province. South African Journal of Science 102: 180-184.

Brink, J.S. 2004. The taphonomy of an Early/Middle Pleistocene hyaena burrow at Cornelia-Uitzoek, South Africa. Revue de Paléobiologie 23(2): 731-740.

Brink J.S. 2004. The evolution of the black wildebeest (Connochaetes gnou) as an indication of the appearance and spread of open grasslands in southern Africa. In L.C. Maul & R.-D. Kahlke (eds.) Late Neogene and Quaternary biodiversity and evolution: Regional developments and interregional correlations. Terra Nostra, Schriften der Alfred Wegener Stiftung 2: 81. (Abstracts of the proceedings of the VI International Palaeontological colloquium in Weimar, 25 – 30 April 2004).

Thackeray, J.F. & J. S. Brink. 2004. Damaliscus niro horns from Wonderwerk Cave and other Pleistocene sites: Morphological and chronological considerations. Palaeontologia Africana 40: 89-93.

Lacruz, R. P.Ungar, P.J. Hancox, J.S. Brink & L.R. Berger. 2003. Gladysvale: Fossils, strata and GIS analysis. South African Journal of Science 99: 283-285.

Scott, L., Y. Fernandes-Jalvo, J. Carrion & J.S. Brink. 2003. Taphonomy of pollen coprolites from Spain and Southern Africa. Palaeontologia Africana 39: 83-91.

Coetzee, L. & J.S. Brink. 2003. Fossil oribatid mites (Acari, Oribatida) from the Florisbad Quaternary deposits, South Africa. Quaternary Research 59: 246-254

Lacruz, R., J.S. Brink, P.J. Hancox, A.R. Skinner, A. Herries, P. Schmid & L.R. Berger. 2002.Palaeontology and geological context of a Middle Pleistocene faunal assemblage from the Gladysvale Cave, South Africa. Palaeontologia Africana 38: 99-114

Roberts, D. & J.S. Brink. 2002. Dating and correlation of Neogene coastal deposits in the Western Cape (South Africa): Implications for Neotectonism. South African Journal of Geology 105: 337-352

Brink, J.S. & Z.L. Henderson. 2001. A high-resolution Last Interglacial MSA horizon at Florisbad in the context of other open-air occurrences in the central interior of southern Africa: an interim statement. In: N. Conard (Ed.) Middle Palaeolithic and Middle Stone Age settlement systems. Kerns Verlag, Tuebingen.pp. 1-20

Carrion, J.S., J.S. Brink, L. Scott, & J.N.F. Binneman. 2000. Palynology of  Pleistocene hyaena coprolites from Oyster Bay, southeastern Cape coast, South Africa: the palaeo-environment of an open-air Howieson’s Poort occurrence. South African Journal of Science 96: 449-453

Brink, J.S. & L. Rossouw. 2000. New trial excavations at the Cornelia-Uitzoek type locality. Navorsinge van die Nasionale Museum, Bloemfontein 16:141-156

Eisenmann, V. & J.S. Brink. 2000. Koffiefontein 'quaggas' and true Cape quaggas: the importance of basic skull morphology. South African Journal of Science 96: 529-533

Churchill, S.E., J.S. Brink, L.R. Berger, R. A. Hutchison, L. Rossouw, D. Stynder, P. J. Hancox, D. Brandt, S. Woodborne, J. C. Loock, L. Scott, and P. Ungar. 2000. Erfkroon: A new Florisian fossil locality from fluvial contexts in the western Free State. South African Journal of Science 96: 161-163

Brink, J.S. 1999. Preliminary report on a caprine from the Cape mountains, South Africa. Archaeozologia 10: 11-26

Brink, J.S., L.R. Berger & S.E. Churchill. 1999. Mammalian fossils from erosional gullies (dongas) in the Doring River drainage. Central Free State Province, South Africa. In: C. Becker, H. Manhart, J. Peters & J. Schibler (eds.), Historium animalium ex ossibusBeiträge zur Paläoanatomie, Archäologie, Ägyptologie, Ethnologie und Geschichte der Tiermedizin: Festschrift für Angela von den Driesch. Rahden/Westf : Verlag Marie Leidorf GmbH, 79-90

Brink, J.S. & L. Webley. 1996. Faunal evidence for pastoralist settlement at Jakkalsberg, Richtersveld, Northern Cape Province. Southern African Field Archaeology 5(2): 70-78

Grün, R., J.S. Brink, N.A. Spooner, L. Taylor, C.B.Stringer, R.B. Franciscus, & A. Murray. 1996.Direct dating of the Florisbad hominid. Nature 382: 500-501

Berger, L.R. & J.S. Brink. 1996. Late Quaternary fossils, including a human patella, from the Riet River Gravels, Free State, South Africa. South African Journal of Science 92: 277-278

Thackeray, J.F., J.S. Brink & I. Plug. 1996. Temporal variability in horncore dimensions of Damaliscus niro from Olduvai, Sterkfontein, Cornelia and Florisbad. In: Stewart, K. & Seymour, K (eds.). Palaeoecology and Palaeoenvironments of late Cenozoic Mammals: Tributes to the career of C.S. (Rufus) Churcher. University of Torornto Press, Canada. pp. 630-636.

Brink, J.S.,  H. de Bruiyn, L.B. Rademeyer and W.A. van der Westhuizen. 1995. A new find of Megalotragus priscus (Alcelaphini, Bovidae) from the central Karoo, South Africa. Palaeontologia Africana 32: 17-22.

Brink, J.S. 1994. An ass, Equus (Asinus) sp., from late Quaternary mammalian assemblages of Florisbad and Vlakkraal, central Southern Africa. South African Journal of Science 90: 497-500.

Peters, J., A. Gautier, J.S. Brink & W. Haenen. 1994. Late Quaternary extinction of ungulates in Sub-Saharan Africa: a reductionist's approach. Journal of Archaeological Science 21: 17-28.

Webley, L, F. Archer & J.S. Brink. 1993. Die Toon: a Late Holocene site in the Richtersveld National Park, northern Cape. Koedoe 36(2): 1-9.

 Brink, J.S. 1993. Postcranial evidence for the evolution of the black wildebeest, Connochaetes gnou: an exploratory study. Palaeontologia Africana 30: 61-69.

Scott, L. & J.S. Brink. 1992. Quaternary palynology, palaeontology and palaeoenvironments in central South Africa. South African Geographer 19: 22-34.

Loubser, J.N.H. & J.S. Brink. 1992. Unusual paintings of wildebeest and a zebra-like animal from north-western Lesotho. Southern African Field Archaeology 1: 103-107.

Brink, J.S. & S. Holt. 1992. A small goat, Capra hircus, from a Late Iron Age site in the eastern Orange Free State. Southern African Field Archaeology 1: 88-91.

Bender, P.A. & J.S. Brink. 1992. A preliminary report on new large mammal fossil finds from the Cornelia-Uitzoek site. South African Journal of Science 88: 512-515.

Peters, J. & J.S. Brink. 1992. Comparative postcranial osteomorphology and osteometry of springbok, Antidorcas marsupialis (Zimmerman, 1780) and Grey rhebok, Pelea capreolus (Forster, 1790) (Mammalia, Bovidae). Navorsinge van die Nasionale Museum, Bloemfontein 8: 162-207.

Brink, J.S., J.J.B. Dreyer & J. Loubser. 1992. Rescue excavations at Pramberg, Jacobsdal, south-western Orange Free State. Southern African Field Archaeology 1: 54-60.

Brink, J.S. & J.A. Lee-Thorp. 1992. The feeding niche of an extinct springbok, Antidorcas bondi(Antilopini, Bovidae), and its palaeoenvironmental meaning. South African Journal of Science 88: 227-229.

Partridge, T.C., D.M. Avery, G.A. Botha, J.S. Brink, J. Deacon, R.S. Herbert, R.R. Maud, L.Scott, A.S. Talma and J.C. Vogel. 1990. Late Pleistocene and Holocene climatic change in Southern Africa. South African Journal of Science 86: 302-306.

Brink, J.S. 1990. Middle Neolithic animal remains from Bajuwarenkaserne, district Regensburg. In J. Schibler, J. Sedlmeier & H. Spycher (eds.) Festschrift für Hans Stampfli: Beiträge zur Archäologie, Anthropologie, Geologie und Paläontologie. Helbing & Lichtenhahn. pp. 31-37.

Bousman, C.B., T.C. Partridge, L. Scott, M. Seaman, S.E. Metcalfe, J.C. Vogel & J.S. Brink. 1988.Palaeoenvironmental implications of late Pleistocene and Holocene valley fills in Blydefontein Basin, Noupoort, C.P., South Africa. Palaeoecology of Africa 19:43-67.

Brink, J.S. 1988. The taphonomy and palaeoecology of the Florisbad spring fauna. Palaeoecology of Africa 19: 169-179.

Brink, J.S. 1987. The archaeozoology of Florisbad, Orange Free State. Memoirs van die Nasionale Museum. 24. 151 pp.

Rubidge, B.S. & J.S. Brink. 1986. Life through the Ages: A guide to the Palaeontology Exhibits of the National Museum. National Museum, Bloemfontein. 65 pp.

Rubidge, B.S. & J.S. Brink. 1985. Preliminary survey of the extent and nature of the Pleistocene sedimentary deposits at Florisbad. Navorsinge van die Nasionale Museum, Bloemfontein 5:69-76.

Deacon, H.J., J. Deacon, A. Scholtz, J.F. Thackeray & J.S.Brink. 1984. Correlation of palaeoenvironmental data from the Late Pleistocene and Holocene deposits at Booplaas cave, southern Cape. In Vogel, J.C. (ed.) Late Cainozoic Palaeoclimates of the Southern Hemisphere. A.A. Balkema. pp. 339-351.

Brink, J.S. & H.J. Deacon. 1982. A study of a Last Interglacial shell midden and bone accumulation at Herolds Bay, Cape Province, South Africa. Palaeoecology of Africa 15: 31-39.

Publications - Popular Articles

Brink, J.S. & N.A. Avenant. 2007. The mountain reedbuck and grey rhebok. ToGoTo  19: 42–43.

Brink, J.S. & S. Vrahimis. 2006. The Highveld Grasslands – Habitat for the Black Wildebeest. ToGoTo 16: 8-9.

Brink, J.S. 2002. Discovery of the first human fossil material from Cornelia-Uitzoek, c. 800 000 years old. Culna 57: 4-7.

Brink, J.S. 1997. Direct dating of the Florisbad fossil hominid. Culna 52: 7-9.

Brink, J.S. 1995. 'n Reuse uitgestorwe wildebees vanuit die sentraal-Karoo. Culna 49: 28-29.

Brink, J.S. 1995. Die herkoms van die mens is van universele belang. Vuka SA 1(1): 81.

Brink, J.S. 1995. New archaeological discoveries (An ass-like equid, Equus (Asinus) sp, from Late Quaternary sites in southern Africa. Paper read at the 7th congress of the International Council for Archaeozoology, Konstanz.) Centre for Science Development Bulletin.

Brink, J.S. 1994. Florisbad: 'n nuwe dateringsmetode. Culna 47:14-15.

Brink, J.S. 1994. Florisbad: esels vir die eerste keer in suidelike Afrika uitgeken. Culna 46: 26-27.

Brink, J.S. & M. Naude. 1993. Florisbad, a research station in the Orange Free State. Conserva 8(3): 24.

Brink, J.S. 1991. Modern people from Africa. Culna 41: 7-8.

Brink, J.S. & F. Prins. 1990. Vroeë tekens van honde in die Transkei. Culna 39: 20.

Brink, J.S. 1986. Die Florisbadman: Was Prof. Dreyer reg in sy interpretasie van die skedel? Nasionale Museum Nuus 31: 12.

Brink, J.S. 1986. Florisbad: 'n kampeerterrein van meer as 40 000 jaar oud. Nasionale Museum Nuus30: 22-23.

Rubidge, B.S. & J.S. Brink. 1985. Geological research at Florisbad. Nasionale Museum Nuus 29: 6-7.

Brink, J.S. 1983. Florisbad: 'n Beeld van die oerjagvelde van die Vrystaat. Nasionale Museum Nuus25: 19.


The Florisbad comparative osteological collection consists largely of ungulates, i.e. suids, bovids and equids that are adapted to the open grasslands of the Free State region and adjacent areas. The carnivores include zoo specimens and re-introduced wild-living specimens.

The original collection, which forms the core of the existing, expanded collection, was assembled by E.C.N. van Hoepen in early 20th century. Van Hoepen collected widely, but the plains zebra collection (Equus quagga) from the Kruger National Park and the Zululand area, which dates to the 1930s and 1940s, is one of the highlights.

Since the mid-1980s we have expanded the collection substantially by adding specimens mainly from the Free State Provincial nature reserves and occasionally from National Parks. The Kalahari springbok collection is a unique collection of 16 individuals from a single herd in the Nossob area. The suids are mainly the common warthog, Phacochoerus africanus, but we do have a few desert warthog, Phacochoerus aethiopicus. Virtually all southern African bovid tribes are represented, with emphasis on Alcelaphini and Antilopini. Black wildebeest, Connochaetes gnou, and blesbok, Damaliscus pygargus, are abundantly represented.







































































Florisbad Quaternary Research Station

Florisbad is a fossil-bearing spring mound situated 45 km northwest of Bloemfontein in the central Free State Province. It is widely known for producing an archaic modern human skull, now dated to c. 260 000 years old. The mammalian remains from the Florisbad spring, dated to between 400 000 and 100 000 years ago, is the type assemblage of the Florisian Land mammal Age and it is a key site for understanding the appearance of modern environments as well as modern human origins in southern Africa. Three osteological collections are housed at the site. It also has a small educational centre, aimed at primary school visits.

Until the late 1970s Florisbad was a privately owned holiday resort and mineral spa. In March 1980 the property was acquired by the National Government and made available to the National Museum to establish a Quaternary Research Station. The buildings on the site now house the offices, laboratories, and collections of the Florisbad Quaternary Research Department. The site was declared a National Monument in 1995, but due to a change in legislation, it is now a Provincial Heritage Site, pending declaration as a National Heritage Site.


Researchers wishing to work on the collections should contact Dr James Brink jbrink@nasmus.co.za.

School groups or groups wishing to go on a conducted educational tour should contact the Education Department Tel: + 27 51 447 9609.
Entrance fee (per appointment only)
Adults: R15.00; Children: R5.00

Groups (more than 10) Adults: R10.00; Children: R2.00


Jaco Smith
Site Manager
Tel: +27 51 831 1132

Tria Oersen
General Assistant

Johannes Motshabi
Johannes Motshabi
General Assistant

Services Offered
  • Collections are made available for study.
  • Training in Quaternary Palaeontology is provided on an ad hoc basis.
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