Research Letter Biostratigraphic correiation and Page 1 of 4

Biostratigraphic correiation in tiie Karoo: The case AUTHORS: of the Middie parareptiie Eunotosaurus Mike Dayi Bruce Rubidge^ The richness of from the of has enabled biostratigraphic John Almond^ subdivision of this Permo- succession, with global applicability. Despite being the thickest of the seven Sifeiani Jirah^ recognised, attempts at further subdivision of the Middie Permian TafJinocephalus Assemblage AFFILIATIONS: Zone () have not been successful, largely because the exact stratigraphie ranges 'Bernard Price Institute of of fossil taxa are unknown. This gap in knowledge has limited stratigraphie eorrelation of the Abrahamskraal Paiaeontoiogy, University of the Formation and hindered understanding of Middle Permian Karoo basin development. Currently, the lowermost , Johannesburg, South Africa Beaufort Group is split between an eastern and a western stratigraphie scheme and, beeause of poor outerop ^Natura Viva, Cape Town, and the reiative paueity of fossiis in the east, stratigraphie eorrelation between the two areas has been South Africa uneertain. Reeent fossil diseoveries of the parareptile Eunotosaurus africafius in the and Free State provinees have extended its known geographic range in ttie east. An additional speeimen from the lower CORRESPONOENCE TO: in the Eastern Cape has, forthe first time, enabied the biostratigraphie eorrelation of this Mike Day unit with the Poortjie Member of the in the west. These finds eonfirm the diaehroneity of the boundary between the marine Eeea Group and the terrestrial Beaufort Group. EMAIL: [email protected]. acza Introduction POSTAL AOORESS: The taxonomic affinity of the enigmatic Permian parareptiie Eunotosaurus africanus, easily recognised in the BPi Palaeontology, fieid by its distinctive broad ribs (Figure 1), has iong intrigued paiaeontoiogists. The genus has been placed in Paiaecsciences Centre, Schooi the Parareptiiia as the soie member ot a sister taxon to the Miiierettidae^'^ and, most recently, as a sister taxon of Geosciences, university to modern turties.^ Aii specimens of Eunotosaurus come from the upper Tapinocephaius and Pristerognathus ot the Witwatersrand, Private Assembiage Zones of the Permian Beaufort Group of South Africa"'^ and most specimens have been recovered Bag 3, Johannesburg 2050, from the western part of the Main Karoo Basin (Tabie 1, Figure 2) where the genus has aiready been used for South Africa biostratigraphic correlation.^^

OATES: Received: 11 June 2012 Revised: 02 Oct. 2012 Accepted: 08 Oct. 2012

KEYWOROS: Eunotosaurus] ; Permian; Karoo; parareptiie; Beaufort Group

HOW TO CITE: Day M, Rubidge B, Almond J, Jirah S. Biostratigraphic correlation in the Karoo: The case of the Middie Permian parareptiie Eunotosaurus. S AfrJSci. 2013;109(3/4), Art. #0030,4 pages. http://dx.doi.org/10.1590/ sajs.2013/20120030

Srmrm: Councii for Geoscience, Pretoria

Figure 1 : The Middie Permian parareptiie Eunotosaurus africanus, specimen M777.

In this part of the basin the Tapinocephaius and Pristerognathus Assemblage Zones correspond approximateiy to the Abrahamskraal Formation and the overlying Poortjie Member of the Teekioof Formation, respectiveiy.^ © 20Í3. The Authors. Correiation of these biozones with iithostratigraphic units in the east of the basin has been difficuit as fossiis are Pubiished under a Creative iess abundant and the Tapinocephaius Assembiage Zone was oniy recently demonstrated to be present in there Commons Attribution Licence. at aii.'

South African Journai of Science Voiume 109 \ Number 3/4 http://www.sajs.eo.za March/Aprii 2013 Researeh Letter Biostratigrapbic correlation and Eunotosaurus Page 2 ot 4

Table 1 : List ot Eunotosaurus specimens included in this study.

Specimen number Collecfion Localify Disfricf Province Unif Certainfy (to within)

AÍVI5999 AiVI Buckiands108 Aibany Eastern Cape K Low

B46 CGS Sterkfontein 147 Murraysburg Westem Cape AorP 100 m

BP/1/3514 BPI Die Bad 286 Beaufort Wesf A 150 m

BP/1/3515 BPI Die Bad 286 Beaufort Wesf Western Cape A 150 m

BP/1/5677 BPI Katferskraai 536 Carnarvon A 50 m

BP/1/6218 BPi The Grant 39 Aibany Eastern Cape K 50 m

BP/1/7024 BPI Bullekraal251 Beautort West Westem Cape A 50 m

BP/1/7027 BPI Buiiekraai251 Westem Cape A 50 m

BP/1/7121 BPI Bullekraal 251 Beautort West Westem Cape A 50 m

BP/1/7198 BPi Zwargershoek151 DeAar Northern Cape ? Low

C()b3 CGS Droogvoetstontein 356 Northem Cape A 50 m

cm341 CGS Kromelboog 68 Beautort West Western Cape P 50 m

cm45 CGS Esferville57 BeauforfWesf Western Cape PorH 100 m

GM71 CGS Bord Voi Water 313 Fraserburg Northern Cape A 100 m

M275 CGS Duiker Kranse 45 Beautort West Western Cape P 50 m

M775 CGS Duiker Kranse 45 Beautort West Western Cape P 50 m

M777 CGS One Kop 53 Beautort West Westem Cape P 100 m

NM3299 NM Philippoiis Commonage Philippoiis Free State Low

NM3466 NM Katterskraai 536 Carnarvon Northem Cape A 50 m

NM3474 NM Kareepoort138 Phiiippoiis Free State 7 Low

NM348S NM Somerstontein 69 Philippoiis Free State ? Low

NM3486 NM Somersfontein 69 Phiiippoiis Free State 7 Low

NM3490 NM Somerstontein 69 Philippoiis Free State ? Low

NM3500 NM Somerstontein 69 Phiiippoiis Free State 7 Low

SAM-PK-011954 SAM De Kruis van Bioem Fontein 323 Fraserburg Northern Cape A 100 m

SAM-PK-K00207 SAM Lammers Kraai 11 Prince Albert Westem Cape A 200 m

SAM-PK-K01132 SAM BushmansRiver312 Beautort West Western Cape A 100 m

SAM-PK-K01133 SAM BushmansRiver312 Beaufort West Westem Cape A 100 m

SAM-PK-K01509 SAM Bumonfein387 Beaufort Wesf Western Cape PorH loom

SAM-PK-K01625 SAM Bulffonfein 387 Beaufort Wesf Western Cape PorH 100 m

SAM-PK-K01673 SAM Bulffonfein 387 Beaufort West Wesfem Cape PorH 100 m

SAM-PK-K06732 SAM Ryers Vaiiey 401 Fraserburg Northem Cape P 100 m

SAM-PK-K07611 SAM Bloukrans 31 Prince Alberf Western Cape A 100 m

SAM-PK-K07670 SAM Springtontein 305 Beautort West Western Cape A 50 m

SAÍVÍ-PK-K07909 SAM Kiipbani« Fontein 137 Beaufort West Western Cape AorP 100 m

SAÍVI-PK-K07910 SAM Kiipbanks Fontein 137 Beaufort West Westem Cape AorP 100 m

SAÍVÍ-PK-K07911 SAM Kiipbanks Fontein 137 Beaufort West Western Cape AorP 100 m

Af[/I, Albany Museum, Grafiamstown; CGS, Councii for Geoscience, Pretoria: BPI, Bernard Price Institute (Palaeontology), Johannesburg: NM, National l^useum, Bloemfontein: SAM, Iziko South African Museum, Cape Town.

A, Abrahamskraal Formation: p Poortjie Member: H, Hoedemaker Member; K, Koonap Formation.

Specimens in bold are at the extremes ot the range.

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Etinoiosaufus specimens 3122, Vietoria West, 1:250 000 Geoiogieai Series 1989), the ioeaiity Western Region data of CGS CM45 indicate that this specimen was eoiieeted from the Soutn.eastern Region Centrai Region iowermost Hoedemai

The stratigraphie position of eaeh Eunotosaurus specimen was then Waterfor d | Koona p : piotted on the stratigraphie seetion tor its respeetive region, spread ¥ Eunotosaurus range across the number of 50-m intervais eaeh eouid possibly oceupy. This approaeh eompensated for the ievei of uneertainty aecompanying aii J Eunotosaurus fossii fossiis, either because ot the iack ot precise iocaiity data or because ot the compiexity ot the geoiogy or iocai topography. The range uitimateiy 500 ^^k Waterford Formation caicuiated is the minimum range torthe genus based on the existing data Poorljie Member with a maximum error ot 50 m. Middleton Formation Institution abbreviations are as foiiows: SAM, iziko South African Abrahamskraal/Koonap Museum, Cape Town: BR Bernard Price Institute (Paiaeontology), • formations Johannesburg: AM, Albany Museum, Grahamstown: NM, Nationai Museum, Bioemtontein: CGS, Couneii for Geoseienee, Pretoria. Results Compilation of aii the ioeaiity data reiating to Eunotosaurus indieates A, Abrafiamsi

South African Journal of Science Voiume 109 | Number 3/4 http://www.sajs.co.za 70 March/Aprii 2013 Research Letter Biostratigraphic correlation and Eunotosaurus Page 4 ot 4

(Eastern Cape Province), occurs in the lower Middleton Formation Acknowledgements (Figure 4). We thank the Paiaeontoiogicai Scientitic Trust (PAST), the NRF and the Department of Science and Technoiogy tor the generous tunding making this research possibie. We also acknowledge two anonymous reviewers tor their usetui and heiptui comments on the manuscript. Authors' contributions M.D. wrote the manuscript, caiculated specimen stratigraphie positions and was involved in the tieldwork; B.R. was the project leader and assisted in writing the manuscript; J.A. conducted some ot the tieldwork; S.J. was involved in tieldwork and assisted in stratigraphie interpretation. References 1. Laurin M, Reisz RR. A réévaluation ot early phyiogeny Zool J Linn Soc. 1995:113(2):165-223. http://dx.doi.0rg/10.111t/j.1096-3642.1995. tb00932.x

2. Tsuii LA, Müller J. Assembling the history ot the Parareptiiia; Phyiogeny, diversification, and a new definition of the ciade. Foss Rec. 2009:12(1 ):71 -81. Figure 4: Eunotosaurus fossil impression embedded in http://dx.d0i.0rg/i O.t O02/mmng.200800011 on the tarm Wiiton, near Middleton, Eastern Cape Province (32°58'40.2"S, 25°45'58.5"E). 3. Lyson TR, Bever GS, Bhuliar B-AS, Joyce WG, Gauthier JA. Transitional tossils and the origin ot turttes. Biol Lett. 2010;6(6):830-833. http://dx.dol. In the Free State, the specimens ot Eunotosaurus are associated orQ/10.1098/rsbi.2010.0371, PMid:20534602, PiVICid:3001370 with scylacosaurid therocephalians and, as no dinocephaiians have 4. Keyser AW, Smith RHM. biozonation ot the Beautort Group been discovered, these specimens are considered to taii within the with special reterence to the Western Karoo Basin. Ann Geol Sur S Atr. Pristerognathus Assemblage Zone'; however, the lithostratigraphic 1979:12:1-36. context is uncertain here.'" 5. Rubidge BS, editor Biostratigraphy ot the Beaufort Group (). The scarcity ot tossii material in the eastern part ot the Basin is S Afr Comm Strat Biostrat Ser. 1995;1:1-46. problematic. The ditterence in stratigraphie regime, contounded by 6. Rubidge BS, Modesto SR Sidor CP Eunotosaurus africanus trom the Ecca- the absence ot the Eodicynodon Assemblage Zone and an unknown Beautort contact in Northern Cape Province, South Atrica - implications tor proportion otthe Tapinocephalus Assemblage Zone near Grahamstown,' Karoo Basin developmenL S Atr J Sei. 1999:95(11/12):553-555. makes subdivision ot the eastern tormations uncertain. Although 7. Modesto SP Rubidge BS, De Klerk WJ, Weiman J. A dinocephalian BP/1/6218 may prove to be the iowest occurrence ot Eunotosaurus, tauna on the Ecca-Beautort contact in Eastern Cape Province, South Atrica. until collecting ettorts create a iarger sample size, the stratigraphie SAtrJ Sei. 2001:97:161-163. ranges ot specimens trom the eastern Karoo Basin have limited use 8. Nicolas M. Tetrapod biodiversity through the Permo-Triassic Beautort Group unless compared to the west. (Karoo Supergroup) ot South Atrica [dissertation]. Johannesburg: University of the Witwatersrand: 2007. Age 9. Weiman J, Loock JC, Rubidge BS. i\lew evidence tor diachroneity of the Traditionally, the age ot the Tapinocephaius and Pristerognathus Ecca-Beautort contact (Karoo Supergroup, South Atrica). S Atr J Sei. assemblage zones has reiied upon comparisons to the Russian Permian, 2001:97(7/8):320-322. and more speciticaily to the Ischeevo tauna.^'" This comparison piaces 10. Jirah S. Sedimentoiogical, paiaeontoiogicai & stratigraphie analysis them in the stage, dated at 265.8-260.4 million years of the Abrahamskraal Formation (Beautort Group) in an area south ot ago.'* From existing material, the stratigraphie range ot Eunotosaurus Mervi/evilie, South Africa [dissertation]. Johannesburg: University ot the corresponds to the upper third ot the Tapinocephaius Assemblage Zone Witwatersrand; 2009. and the entirety ot the Pristerognathus Assemblage Zone. The upper end t1. Coie Dl, Wippiinger PE. Sedimentology and molybdenum potentiai of the ot the Eunotosaurus range is theretore dated to about 260 million years Beautort Group in the main Karoo basin. South Atrica. C tor Geosci Mem. ago, at the Capitanian- boundary.'^ The lower extent 2001:80:1-225. ot its range is more ditticuit to determine, but given the data avaiiabie trom biostratigraphic correlation with , the tirst appearance ot the 12. Mason R. A bio- and litho-stratigraphic study ot the Ecca-Beautort contact in the southeastern Karoo Basin (Albany district. Eastern Cape Province) genus would probably tall within the middle Capitanian. [dissertation]. Johannesburg: University otthe Witvi/atersrand; 2007. Conclusion 13. Rubidge BS, Hancox PJ, Catuneanu 0. Sequence anaiysis ot the Ecca Beautort contact in the southern Karoo ot South Africa. S Atr J Geoi. Our study demonstrates that Eunotosaurus has a restricted stratigraphie 2000:103(1):81-96. range ot about 450 m within the tluvial lower Beautort Group ot the Main Karoo Basin and can theretore be usetui tor biostratigraphic correlation. 14. Rubidge BS, Erwin DH, Ramezani J, Bowring SA, De Klerk WJ. High-precision The presence ot this genus in only the upper part ot the Koonap Formation temporal calibration of Late Permian vertebrate biostratigraphy: U-Pb zircon constraints trom the Karoo Supergroup, South Atrica. Geology. In press 2013, and iowermost Middieton Formation supports the lithostratigraphic doi:10.1130/G33622.1. http://dx.d0i.0rg/i 0.2113/103.1.81 correlation ot the Poortjie Member in the western Karoo Basin to the base ot the Middleton Formation in the East.'^ As the distribution ot 15. Wardiaw BR, Davydov V Gradstein FM. The Permian period. In: Gradstein Eunotosaurus corresponds to a tixed time period, the recognition that the FM, Ogg JG, Smith AG, editors. A geologic time scaie 2004. Cambridge: Cambridge University Press; 2004. p. 249-270. genus occurs in the upper but not the lower Tapinocephaius Assemblage Zone in the south has turther utility; the presence ot Eunotosaurus 16. Hill RS. Expianation to geological map sheet 3224 Graatt-Reinet. Pretoria: with a Tapinocephalus Assemblage Zone tauna on the Ecca-Beautort GeolSurSAtr;1993. p. 1-31. boundary in the Carnarvon district. Northern Cape,^ and its presence with a Pristerognathus Assemblage Zone tauna immediateiy above the Ecca-Beautort boundary at Phiiippoiis, southern Free State," contirm the younging ot the Karoo paiaeoshoreline towards the north.

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