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A possible later stone age painting of a dicynodont (Synapsida) from the South African Karoo [1]

['Julien Benoit', 'Evolutionary Studies Institute', 'School Of Geosciences', 'University Of The Witwatersrand', 'Johannesburg', 'South Africa']

Date: 2024-10

Abstract The Horned Serpent panel at La Belle France (Free State Province, South Africa) was painted by the San at least two hundred years ago. It pictures, among many other elements, a tusked animal with a head that resembles that of a dicynodont, the fossils of which are abundant and conspicuous in the Karoo Basin. This picture also seemingly relates to a local San myth about large animals that once roamed southern Africa and are now extinct. This suggests the existence of a San geomyth about dicynodonts. Here, the La Belle France site has been visited, the existence of the painted tusked animal is confirmed, and the presence of tetrapod fossils in its immediate vicinity is supported. Altogether, they suggest a case of indigenous palaeontology. The painting is dated between 1821 and 1835, or older, making it at least ten years older than the formal scientific description of the first dicynodont, Dicynodon lacerticeps, in 1845. The painting of a dicynodont by the San would also suggest that they integrated (at least some) fossils into their belief system.

Citation: Benoit J (2024) A possible later stone age painting of a dicynodont (Synapsida) from the South African Karoo. PLoS ONE 19(9): e0309908. https://doi.org/10.1371/journal.pone.0309908 Editor: Dawid Surmik, University of Silesia, POLAND Received: July 23, 2024; Accepted: August 20, 2024; Published: September 18, 2024 Copyright: © 2024 Julien Benoit. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. Data Availability: All relevant data are within the manuscript and its Supporting Information files. Funding: This research was financially supported by the DSI-NRF African Origins Platform (AOP210218587003; UID: 136505). The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript. Competing interests: The authors have declared that no competing interests exist.

Introduction A crucial, yet outstanding question about the history of palaeosciences is that of palaeontological indigenous knowledge in the South African Karoo. The almost continuous Permo-Jurassic fossil record of the South African Karoo chronicles the rise, diversification, and fall of the Therapsida, as well as the evolutionary origins of mammals, turtles, dinosaurs, and lizards, providing many intermediate species each documented by dozens of specimens [1–4]. The abundance of tetrapod fossils in the Karoo and their quality of preservation both make it a konzentrat and konservat lagerstatten [1,5,6]. Given the wealth of fossils that the Main Karoo Basin and other Karoo-aged basins have delivered, and the long human occupation of this part of the African continent, the existence of a long-standing indigenous knowledge of fossils is very likely [7,8]. Yet, the study of African indigenous palaeontology is still fairly young [7,9–12], and the evidence remains accordingly sparse and debatable, especially given the scarcity of written accounts. Although patchy, a growing record of geomyths, place names, written accounts, and archaeological evidence supports that many southern African cultures knew and, in some cases, inquired about the fossils around them [7,8]. The Horned Serpent panel, in the Koesberg mountains, gorgeously depicts elements of the San culture and was partly figured and described by Stow and Bleek [13] in plates 36–39. The rock art was made at an unknown date by the “Bushmen of the East” [14], now referred to as the /Xam speaking San [15]. One of these elements is an unidentified animal that bears two enlarged tusks, making it superficially look like a walrus (Fig 1A). This walrus-like figure has been the object of much speculation regarding its identity, because no representative of the Odobenidae lives or has ever lived nearby sub-saharan African coasts [16–18]. Tusked creatures (more or less imaginary and composite) are not rare in San rock art, including tusked lion, snake, antelopes, and people; but in those cases, the tusks are always curved upwards, like they do in warthogs and bushpigs [19–21], not downwards as in the tusked animal from La Belle France (Fig 1A). PPT PowerPoint slide

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TIFF original image Download: Fig 1. The tusked animal of the Horned Serpent panel compared to the skull of a dicynodont. A, the tusked animal of the Horned Serpent panel redrawn from Stow and Bleek [13]. B, skull of a Diictodon feliceps (BP/1/8140, Jasfontein, Victoria West District) photographed by the author in situ at the moment of its discovery, before excavation, and unprepared. https://doi.org/10.1371/journal.pone.0309908.g001 The question of the identity of this animal has been dismissed because it is seemingly a rain-animal from the San “spirit-realm” [13], which is spiritual by essence and does not have to be realistic [18]; however, this does not address the question of what inspired this figure in the first place. Even the most fantastic elements of San art, such as therianthropes and rain-animals, are based on actual animals and phenomena [13,21–25]. The San spiritual pantheon is directly inspired from their real-life environment and the fantastic beings they painted are thus always an amalgamation of different existing animals [13,14,21–23,25–27]. As such, whether fantastic or not, the question of what inspired this tusked animal remains open. The /Xam speaking San, who made the Horned Serpent painting, occupied the Karoo area [15], a landscape in which the fossil-richness is mostly due to the overly abundant and often well-preserved dicynodonts, a group of tusked therapsids [1,2,5,28,29]. In many cases, their skulls are naturally exposed by erosion in spectacular ways (Fig 1B), making them easy to find and collect, and their tusks are so conspicuous that their anatomy is not difficult to interpret, even to the untrained eyes. The downturned tusks of dicynodonts resemble those of the tusked animal of the Horned Serpent Panel (Fig 1). Archaeological evidence directly supports that the San did find and transport fossils over long distances [7,8], and could interpret them in surprisingly accurate ways [11,12]. If the San could identify the fossilised skulls of dicynodonts as belonging to once alive animals, it is possible that their tusked faces could have contributed to their rock art. In this respect, it is noteworthy that, to the San of the Koesberg, the animals depicted on the Horned Serpent panel were real and used to live among them: “The Bushmen of the east […] declare that there were at one time a number of animals living in the country in the days of their forefathers, which are now extinct and nowhere to be found in Southern Africa. Some of these are described as great monstrous brutes, exceeding the elephant or hippopotamus in bulk.” (p. 131 in [14]). In the next page of the same book, the tusked animal is described as an entity distinct from the rain-animal (referred to as ‘Kou-teign-Koo-rou) and the Serpent (referred to as ‘Koo-be-eng) [14]. In addition to its tusks, the extraordinary size of the animal evokes the heavily mineralised bones and disproportionately enlarged skulls of some dicynodonts found in abundance in the Main Karoo Basin, such as Kannemeyeria, Lystrosaurus, and Daptocephalus [2,6,28,30] If the La Belle France painting pictures a dicynodont, it would have been made independently from, and perhaps before, the discovery of Dicynodon lacerticeps, the first dicynodont described in the western scientific literature [31]. This could be the oldest depiction of a dicynodont ever made. This possibility is intriguing and warrants scrutiny. In this contribution, the Horned Serpent panel was re-examined to address its accuracy and determine its age. The accuracy of Stow and Bleek’s [13] reproductions has been questioned, which may alter significantly the interpretation of the depicted scenes [32], and as such, first hand photographs of the paintings were made. The palaeontology and geology of La Belle France were also explored to assess the fossil richness of the area and its potential for supporting a geomyth.

Conclusion The ethnographic, archaeological, and palaeontological evidence are consistent with the hypothesis that the Horned Serpent panel could possibly depict a dicynodont. This is supported by i) the downward orientation of the tusks on the La Belle France tusked animal (Fig 3A and 3B), which does not match that of any modern African animals, but does match the tusks of dicynodonts (Figs 1B and 3C), ii) the abundance of dicynodont fossils in the area (Fig 5), and iii) the local San belief into a long extinct, large animal [14]. This would imply that the San may have i) discovered dicynodont fossils, ii) interpreted them as long-extinct species, iii) made a painting of one of them at La Belle France, and iv) integrated them into their worldview. If so, this would evidence that Later Stone Age people were aware of dicynodont fossils at least a decade before their formal scientific description by western scientists [31], and made the first known reconstruction of one of them. Even if one considers that the Horned Serpent panel has a purely spiritual meaning, it does not invalidate the hypothesis that the tusked animal itself may have been imagined based on a dicynodont fossil. The spiritual and palaeontological interpretation of this painting are not mutually exclusive. Helm et al. [7] were the first to propose that some of the strangest animals in the San rock art may be interpreted as depictions of Permian reptiles, and the current contribution supports that perhaps there may be more dicynodont depictions to be found among some of the paintings and petroglyphs preserved in the Karoo [7,19–21,60,61].

Acknowledgments To Mr. and Mrs. Swanepoel and Mr. and Mrs. Kruger for access to La Belle France and showing the location of the paintings. To Luyanda Gcaza, Nomatile Nombewu, the Eastern Cape Province Tourism Agency, and Eastern Cape Province Heritage Resources Agency for access to the Oviston Nature Reserve. To the two anonymous reviewers for their help towards improving this work.

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