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Analysis of Millerettid Parareptile Relationships in the Light of New Material of BROOMIA PERPLEXA Watson, 1914, from the Permian of South Africa

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Journal of Systematic Palaeontology 6 (4): 453–462 Issued 24 November 2008 doi:10.1017/S147720190800254X Printed in the United Kingdom C The Natural History Museum ! Analysis of millerettid parareptile relationships in the light of new material of BROOMIA PERPLEXA Watson, 1914, from the Permian of South Africa Juan Carlos Cisneros Departamento de Paleontologia† e Estratigrafia, Universidade Federal do Rio Grande do Sul, Porto Alegre, CP 15001, 91501-970, Brazil; and Bernard Price Institute for Palaeontological Research, University of the Witwatersrand, Private Bag 3, WITS 2050, Johannesburg, South Africa Bruce S. Rubidge Bernard Price Institute for Palaeontological Research, University of the Witwatersrand, Private Bag 3, WITS 2050, Johannesburg, South Africa Richard Mason Bernard Price Institute for Palaeontological Research, University of the Witwatersrand, Private Bag 3, WITS 2050, Johannesburg, South Africa Charlton Dube Bernard Price Institute for Palaeontological Research, University of the Witwatersrand, Private Bag 3, WITS 2050, Johannesburg, South Africa SYNOPSIS A second specimen of the poorly known millerettid Broomia perplexa is reported. It consists of a cranium and partially articulated postcranium. As indicated by its small size (ca.35% shorter skull than the holotype) and unfused pelvis and limbs, the new specimen is a sub-adult indi- vidual. Comparison with the holotype reveals only minor differences that are ascribed to ontogenetic or individual variation. Accordingly we consider the new specimen to represent the second record of Broomia,discovered90yearsafterthedescriptionoftheholotype.Apreliminaryphylogenetic analysis of millerettid inter-relationships identifies Broomia as a millerettid related to the genus Millerosaurus.TheenigmaticparareptileEunotosaurus africanus is nested within Millerettidae as the sister taxon of Milleretta rubidgei.WhereasmillerettidsarewellknownfromthelatestPermian Dicynodon Assemblage Zone of the Beaufort Group of South Africa, Broomia and Eunotosaurus are found in the Middle Permian Tapinocephalus Assemblage Zone. The new Broomia specimen not only complements our knowledge of this early millerettid but also shows, in addition to recent findings in the same area, that the lowermost Beaufort Group is at least 1200 m thick in this area compared with ca. 2500 m in the western part of the Karoo Basin. KEY WORDS Parareptilia, phylogeny, Gondwana, Middle Permian, Tapinocephalus Assemblage Zone Contents Introduction 454 Institutional abbreviations 454 Material and methods 454 Geological setting 454 Systematic palaeontology 455 Parareptilia Olson, 1947 sensu deBraga & Reisz, 1996 455 Millerettidae Romer, 1956 455 Broomia perplexa Watson, 1914 455 Description 457 Cranial skeleton 457 E-mail: [email protected] † 454 J. C. Cisneros et al. Postcranial skeleton 458 Comparisons with other millerettids 458 Interrelationships of the Millerettidae 459 Methods 459 Results 459 Discussion 459 Conclusions 460 Acknowledgements 460 References 461 Appendix A: Character list 461 Appendix A: Data matrix 462 Introduction Institutional abbreviations Millerettids are small lizard-like reptiles considered to be the BMNH Natural History Museum, London. = basalmost members of the Parareptilia (Laurin & Reisz 1995; BP Bernard Price Institute for Palaeontological Re- = deBraga & Reisz 1996; Modesto 1999; Berman et al.2000; search, University of the Witwatersrand, Johan- Reisz & Scott 2002). Some features of the group include a nesburg. cranial ornamentation formed by shallow tubercles and the RC Rubidge Collection, Wellwood, Camdeboo = presence of a temporal fenestra (Gow 1972). Five species are (formerly Graaff-Reinet) Municipality, Eastern known: Milleretta rubidgei Broom, 1938, Millerosaurus or- Cape, South Africa. natus Broom, 1948, Millerosaurus nuffieldi Watson, 1957, Milleropsis pricei (Watson, 1957) and Broomia perplexa Watson, 1914. All are from the Permian of the Karoo Basin in South Africa. Whereas the millerettids from the latest Material and methods Permian Dicynodon Assemblage Zone (AZ) are known from The new specimen, BP/1/6222, comprises a skull and post- several specimens, the Middle Permian Broomia perplexa is cranial elements. Most of the specimen had been exposed by the only millerettid from the Tapinocephalus AZ, and it is erosion and was badly weathered, with only the impression represented only by the holotype (Kitching 1977). of bone preserved in places. An air hammer was used as the In the past, Broomia has been considered by several main instrument to prepare the palate, but manual cleaning authors to be a taxon that is closely related, or ancestral, to with fine needles was employed for delicate areas. The pel- the Millerettidae (Broom 1921, 1941; Romer 1956; Watson vic girdle and the right hindlimb were preserved largely as 1957; Kuhn 1969), but was not mentioned in the review of impressions in the mudstone, and negative preparation (i.e. the Millerettidae by Gow (1972). The taxon has also been the remaining bone was removed) was undertaken on these regarded as a probable araeoscelid diapsid (Nopcsa 1923; areas. This procedure was not applied to the right knee be- Piveteau 1955) or procolophonoid parareptile (Romer 1966). cause in this area the bone is in good condition. The pelvis In a detailed and well-illustrated re-description of the genus, and right hindlimb areas that had been negatively prepared Thommasen & Carroll (1981) justified the classification of were filled with silicone in order to produce a positive cast this taxon as a millerettid mainly on the basis of palatal fea- for study. Numerous morphological details of the palate and tures, but they could not evaluate the presence of distinctive the manus became visible only through the application of millerettid features in other areas of the skull because the roof immersion oil. of the cranium is not preserved in the holotype. Broomia was The holotype of Broomia perplexa,BMNHR4065,con- also excluded from the phylogenetic definition of the group sists of a natural impression and a silicone cast was employed proposed by Laurin & Reisz (1995) and was not included for its study. Other comparative material included: Miller- in any other reptilian group by these authors. Scepticism in etta rubidgei,BP/1/2040,BP/1/2610,BP/1/2614,BP/1/2876, regarding Broomia as a millerettid may be due to the incom- BP/1/3818, BP/1/3821, BP/1/3822, RC 14 (holotype) and pletenessoftheholotype,whichlackstheroofofthecranium, RC 70; Millerosaurus ornatus,RC78(holotype);Millerop- where some features that diagnose the Millerettidae are loc- sis pricei, BP/1/720 (a block with at least eight specimens, ated (Gow 1972; Laurin & Reisz 1995). In addition, a con- including the holotype) and BP/1/4203. siderable stratigraphical gap exists between Broomia,known from the Tapinocephalus AZ, and the millerettids from the Dicynodon AZ. This paper reports a new parareptile speci- men from the Tapinocephalus AZ in Eastern Cape Province that can be clearly identified as a millerettid and exhibits fea- Geological setting tures that are comparable with only Broomia perplexa.The BP/1/6222 was discovered by Charlton Dube on the farm, new specimen also provides important cranial information The Grant, in Eastern Cape Province (Fig. 1). It was re- not present in the holotype. The stratigraphical implications covered from an argillaceous interval comprising dark olive- of the presence of Broomia in Eastern Cape Province are green mudstone with minor sandstones, 1705 m above the discussed. Ecca/Beaufort boundary. Stratigraphically the locality falls South African Permian millerettid parareptile relationships 455 Figure 1 Locality map showing part of the Grahamstown area of South Africa and the farm The Grant. The site where the specimen was recovered is indicated by an asterisk. within the middle part of the Koonap Formation (lower Systematic palaeontology Beaufort Group), which is considered to have been depos- ited in a subaerial delta plain setting (Johnson & le Roux PARAREPTILIA Olson, 1947 sensu deBraga & 1994). Reisz, 1996 At present the biostratigraphy of this part of the Karoo MILLERETTIDAE Romer, 1956 Basin is poorly known (Rubidge 1995). However, as a res- ult of recent collecting activities in the area, dinocephali- NEW DEFINITION. A clade that comprises all taxa more re- ans (Modesto et al. 2001), scylacosaurid therocephalians lated to Milleretta rubidgei Broom, 1938, than to Macroleter and the parareptile Eunotosaurus have been found on The poezicus Tverdokhlebova & Ivakhnenko, 1984. Grant, thus indicating that the lowermost Beaufort Tapino- cephalus AZ (Boonstra 1969; Smith & Keyser 1995) is present in the area. This is in addition to an earlier dis- Broomia perplexa Watson, 1914 (Figs 2–3) covery of Eunotosaurus 15 km to the east (Gow & de Klerk HOLOTYPE. BMNH R4065, an impression of a skull and 1997). fairly complete and articulated postcranium in ventral view. Figure 2 Broomia perplexa BP/1/6222. A, block containing the fossil, most elements are exposed in dorsal view. Pelvis and right hindlimb are shown prior to negative preparation. B, reverse side of the block, showing palate, the tibia–femur articulation and some unidentified postcranial elements. Abbreviations: cr, cranium; cl, clavicle; fe, femur; fi, fibula; h, humerus; is, ischia; ph, phalanx; m, manus; pb, pubis; r, radius; ti, tibia; ul, ulna; vt, vertebrae. 456 J. C. Cisneros et al. Figure 3 Broomia perplexa BP/1/6222. A, cranium, left lateral and dorsal surfaces. B, cranium in palatal view and attached
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  • Biarmosuchus

    Biarmosuchus

    Meet the Amniotes: The great terrestrial adaptation Pterosauria Archaic archosaurs Crocodiles Dinosauria Lepidosaurs Anapsids Synapsids Most ‘amphibians’ Most ‘fishes’ Assorted jawless fish Amniota Urochordata Tetrapoda Cephalochordata Gnathostomata Vertebrata Amniotic egg Chordata Thick skin Distinctive skulls The cleidoic egg: a private pond Eggshell: Semipermeable Calcareous or leathery Albumen: Egg cytoplasm Amnion: Protection / Gas transfer Yo l k Sac: Nutrient Pool Allantois: Waste Pool Synapsida Anapsida Lepidosauria Archosauria First amniotes Diapsida in record (!!) Eureptilia Amniotes Walking with Monsters Chapter 2 1:10-5:00 Evolution of Eggs? Some modern amphibians To deal with longer time Eggs became larger, lay eggs on land... why? periods on dry land, tougher. Large eggs can - One inner membrane tougher shells were produce larger babies, 1. escape predation selected for. Gas exchange which had a higher and waste devices evolved likelihood of survival in a for complete eggtonomy tough world. Evolution of Hair? Amniotes all have the gene for hair: alpha keratin In birds/lizards, it’s expressed in claws In mammals, it’s used in hair & nails 310 Ma Thrinaxodon Blood vessel channels on premaxillae, maxillae ~vibrassae (whiskers) (early Triassic) Castorcauda First direct fossil evidence of hair (mid-Jurassic) Meet the Amniotes No temporal fenestra Upper temporal fenestra Lower temporal fenestra Single temporal fenestra = ‘window’ fenestra The Permian 299-251 Ma The Permian 299-251 Ma Convergence of Pangaea The effects of the landscape on climate: Gondwana icecap disappeared Heat distributed more equally through fluids than solids as continent drifted north Oceans slower to warm/cool than continents Pangaea: Rapid warming/cooling ~ more intense than today Temperature extremes Our modern continents are ‘tempered’ by oceans between them.