Volume 41 December 2005

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Preliminary report of a large theropod dinosaur trackway in Clarens Formation sandstone (Early Jurassic) in the Paul Roux district, northeastern Free State, South Africa
(BERNARD PRICE INSTITUTE FOR PALAEONTOLOGICAL RESEARCH, 2005) Raath, Michael A.; Yates, Adam M.
An isolated fallen block of Clarens Formation sandstone near the small northeastern Free State town of Paul Roux preserves part of the trackway of a bipedal dinosaur. Although well known as a local curiosity, this trackway has not previously been formally reported or described. It consists of five successive paces of what is interpreted as a medium-sized to large theropod dinosaur, and represents the largest known theropod trackway in the ‘Stormberg’ sequence in South Africa. The tracks are assigned to the ichnotaxon Grallator sp., and show similarities to North American tracks of comparable age originally described as Dilophosauripus. Until now no body fossils of a likely candidate trackmaker were known, but elsewhere in this volume a possible candidate is described by the second author.
Makapansgat suids and Metridiochoerus
(BERNARD PRICE INSTITUTE FOR PALAEONTOLOGICAL RESEARCH, 2005) Cooke, H. B. S.
Fossil suid material from Member 3 of the Makapansgat Formation was described as Potamochoeroides shawi but some authors have regarded it as an early stage of the Metridiochoerus andrewsi lineage. There is no complete cranium so a new reconstruction has been based on combining the data from a number of partial specimens. Comparison with Metridiochoerus andrewsi is difficult as there are reasons to suspect that the so-called ‘male’ and ‘female’ crania from Koobi Fora could be specifically distinct. An undescribed cranium from the Omo Shungura Formation is morphologically similar to the ‘male’ and is attributed to Metridiochoerus jacksoni. Comparison with the reconstructed Makapansgat suid indicates that the latter already shows an early stage in the distinctive architecture of the M. jacksoni cranium, as well as some resemblances in dental features. Accordingly it is suggested that the Makapansgat suid be designated as Metridiochoerus shawi. A few teeth from Member B11 of the Shungura Formation, with an age close to 2.95 Ma, are placed provisionally as M. cf. shawi; previously described molars referred to an early Notochoerus scotti are probably from a similar level. While this is suggestive, correlation is not firmly established.
New information on the palate and lower jaw of Massospondylus (Dinosauria: Sauropodomorpha)
(BERNARD PRICE INSTITUTE FOR PALAEONTOLOGICAL RESEARCH, 2005) Barrett, Paul M.; Yates, Adam M.
Additional anatomical details of the palate and lower jaw of the prosauropod dinosaur Massospondylus Owen are documented on the basis of a previously undescribed skull from the upper Elliot Formation. The palate is generally similar to that of other early sauropodomorphs, but can be shown to differ from those of Plateosaurus, Lufengosaurus and Thecodontosaurus in several respects. For example, Massospondylus lacks the well-developed palatine boss seen in Plateosaurus and the pneumatic recess that is present on the ectopterygoid of Thecodontosaurus. In addition, Massospondylus possesses an expanded medioventral premaxillary process that is much larger than that of any other basal sauropodomorph.
A new specimen of Dicynodon traquairi (Newton) (Synapsida: Anomodontia) from the Late Permian (Tartarian) of northern Scotland
(BERNARD PRICE INSTITUTE FOR PALAEONTOLOGICAL RESEARCH, 2005) Cruickshank, Arthur R. I.; Clark, Neil D. L.; Adams, Calum
A recently discovered natural mould of a complete, almost undistorted, skull and lower jaw of a dicynodont (c. 237mmoverall length), in a block of Upper Permian sandstone (= Dicynodon Assemblage Zone: Hopeman Sandstone Formation) from Clashach Quarry, Hopeman, Morayshire, is described using novel techniques, including Computed Tomography scanning (CT), Magnetic Resonance Imaging (MRI) and rapid-prototype modelling. It is assigned to the taxon Dicynodon traquairi (Newton, 1893). When compared with Dicynodon lacerticeps Owen, 1845, it is distinguished principally by having the pineal opening sunk deeply between the diverging parietals, subparallel pterygoid rami narrowly separated, with no transverse flanges, and in addition, a deeply grooved lower jaw symphysis. The southern African fauna lived on river flats in a higher (southern) palaeolatitude than the possibly desert-dwelling Scottish species. The Hopeman Sandstone Formation is of the same age as the better-known Cutties Hillock Sandstone Formation, whose fauna is briefly discussed and reviewed.
A fossil peat deposit from the Late Triassic (Carnian) of Zimbabwe with preserved cuticle of Pteridospermopsida and Ginkgoales, and its geological setting
(BERNARD PRICE INSTITUTE FOR PALAEONTOLOGICAL RESEARCH, 2005) Barale, Georges; Bamford, Marion K.; Gomez, Bernard; Broderick, Timothy J.; Raath, Michael A.; Cadman, Ann
Well-preserved cuticular material of Pteridospermopsida and Ginkgoales from the Late Triassic of Zimbabwe is described here for the first time. It is preserved within a brown peat-like lens in the Upper Karoo Angwa Sandstone Formation. The locality is on the Manyima River in the lower portion of the mid-Zambezi Valley of Zimbabwe. Using SEM and light microscopy to identify the taxa, the fragmentary cuticles are of Pteridospermopsida type and have been assigned to Lepidopteris sp. (Peltaspermales) and Dicroidium sp. A, B, (Corystospermales). Cuticles of the ginkgoalean leaf genus, Sphenobaiera, are also described. Well-preserved ovules were found in close association with the cuticles, but as the stomata are not visible they cannot be assigned to any genus. Based on their close similarity to the Dicroidium flora of the South African Upper Karoo, the plants are considered to be equivalent to the South African Molteno Formation in age (Carnian). The palynoflora supports this age bracket, as does fauna preserved nearby. The taphonomic process was one of transport, sorting and deposition in a fluvial system.
The taxonomic status of Parathrinaxodon proops (Therapsida: Cynodontia), with comments on the morphology of the palate in basal cynodonts
(BERNARD PRICE INSTITUTE FOR PALAEONTOLOGICAL RESEARCH, 2005) Abdala, Fernando; Allinson, Matthew
The holotype and only specimen of Parathrinaxodon proops, a cynodont from the Upper Permian Kawinga Formation, Tanzania, is redescribed. Upper postcanines from the middle of the tooth row are ovoid in outline, presenting a large main cusp and tiny anterior and posterior accessory cusps on the sectorial margin. Anterior and posterior lingual cusps on the crown indicate the presence of a lingual cingulum. The overall postcanine morphology is remarkably similar to that of Procynosuchus delaharpeae, a Late Permian cynodont particularly common in the lower Beaufort Group of South Africa. The presence of a complete osseous palate and a medial palatal opening between the maxillae (=vomerine fossa) in Parathrinaxodon proops remain the main differences previously reported between this species and Procynosuchus delaharpeae. Restudy of the palate of Parathrinaxodon proops indicates that there exists some degree of deformation, particularly notable in the broken and distorted vomer. The supposed presence of the complete secondary palate and of the medial palatal opening in Parathrinaxodon proops are interpreted as resulting from a slight horizontal displacement of the long, and originally free, palatal processes of the maxilla and palatine. It is concluded that Parathrinaxodon proops is synonymous with Procynosuchus delaharpeae. This synonymy is problematic because Parathrinaxodon proops Parrington 1936 would have priority over Procynosuchus delaharpeae Broom 1937, but the latter is the best known Late Permian cynodont. Consequently, we propose to conserve Procynosuchus delaharpeae as the valid name for this cynodont based on article 23, section 9 (Reversal of precedence) of the International Code of Zoological Nomenclature. An analysis of the Kawinga fauna, using genus as the taxonomic unit for comparison, indicates strong similarity (67%) with faunas from theTropidostoma, Cistecephalus and Dicynodon assemblage zones from the SouthAfrican Karoo.
A new theropod dinosaur from the Early Jurassic of South Africa and its implications for the early evolution of theropods
(BERNARD PRICE INSTITUTE FOR PALAEONTOLOGICAL RESEARCH, 2005) Yates, Adam M.
A new theropod, Dracovenator regenti, from the upper Elliot Formation is described, based upon a fragmentary skull. It can be diagnosed on the basis of a bilobed fossa on the lateral surface of the premaxilla that is connected to the alveolar margin by a narrow channel, the presence of a deep, oblique, lateral notch on the articular and hypertrophied dorsal processes on the articular. Other aspects of its morphology display a mosaic of coelophysoid and advanced theropod characteristics. A cladistic analysis of basal Theropoda, including the new taxon finds that the new taxon is closely related to Dilophosaurus wetherilli and Zupaysaurus rougieri although the clade formed by these three taxa is not robustly supported. It also finds that Coelophysoidea sensu lato is paraphyletic with respect to Ceratosauria + Tetanurae but that this topology is not a significantly better explanation of the data than an inclusive, monophyletic Coelophysoidea.
James William Kitching (1922–2003): a tribute
(BERNARD PRICE INSTITUTE FOR PALAEONTOLOGICAL RESEARCH, 2005) Raath, Michael A.; Rubidge, Bruce S.
On 24 December 2003, James William Kitching, regarded by many as one of the world’s greatest fossil finders, died at his home in Johannesburg. His passing marks the end of a pioneering era of palaeontological giants in South Africa.
Palaeontologia africana Volume 41
(Bernard Price Institute for Palaeontological Research, 2005)
Late Triassic traversodontids (Synapsida: Cynodontia) in southern Africa
(BERNARD PRICE INSTITUTE FOR PALAEONTOLOGICAL RESEARCH, 2005) Battail, Bernard
Scalenodontoides macrodontes was described in 1957 by Crompton & Ellenberger as a new genus and species of the family Traversodontidae. For many years it was known only by its type specimen, a lower jaw from the Upper Triassic of Lesotho. The specimen was redescribed in more detail by Hopson in 1984, who established the close affinities of Scalenodontoides with Exaeretodon. In 1993, Gow & Hancox described the first skull of Scalenodontoides, discovered, together with fragmentary remains, in South Africa. The skull from South Africa looked very much like a skull from Lesotho, housed in the Muséum National d’Histoire Naturelle, Paris, and initially attributed, in an unpublished work, to the chiniquodontid Belesodon (Costedoat, 1962). Further preparation of the skull from Lesotho was carried out; the specimen proved not to belong to a chiniquodontid, but to a large traversodontid, described in this paper. A revision of the traversodont remains known from the Late Triassic lower Elliot Formation of Lesotho and South Africa leads to the conclusion that they can all be attributed to the species Scalenodontoides macrodontes. Detailed comparisons between Scalenodontoides and Exaeretodon confirm Scalenodontoides as a valid genus, with only one species, Scalenodontoides macrodontes. A new diagnosis of Scalenodontoides macrodontes, based on an analysis of all available material, is given.
The ownership of the Taung skull and of other fossil hominids and the question of repatriation
(BERNARD PRICE INSTITUTE FOR PALAEONTOLOGICAL RESEARCH, 2005) Tobias, Phillip V.
The ownership of fossils, and for purposes of this paper I refer to that of hominid fossils, was long assumed to be vested in the individuals who made the discoveries. The author reviews here a series of case histories with which he has had direct or indirect personal contact, that illustrate claims for ownership. Some have been explicit, some implicit. They are drawn from South Africa, East Africa, North Africa, England, France, Germany, Italy, Russia, the Netherlands, Indonesia and China. This historical essay reviews the replacement of this practice by a policy that fossils are not seen as personal property, but as part of the heritage of the country of origin. During the colonial era, many specimens were removed from former colonies to the ‘home countries’, where they remained for decades, at least until the subject territories attained their independence from the former imperial powers. The new policy about ownership, in such cases, entails the return (repatriation) of the expatriate fossils to the source country. Examples of success stories and of tardy responses are given. A policy for the future is set forth.
A juvenile gomphodont cynodont specimen from the Cynognathus Assemblage Zone of South Africa: implications for the origin of gomphodont postcanine morphology
(BERNARD PRICE INSTITUTE FOR PALAEONTOLOGICAL RESEARCH, 2005) Hopson, James A.
The partial skull and lower jaws of a small gomphodont cynodont from the Cynognathus Assemblage Zone of South Africa has a well-preserved postcanine dentition distinctly different from that of contemporaneous adult Diademodon and Trirachodon. On the basis of its small size and great amount of tooth replacement it is interpreted to be a juvenile individual. The postcanines are compared with those of adults and juveniles of Diademodon and traversodontids and is seen to differ from them. Comparison with adults of Trirachodon shows some unique postcanine resemblances, such as well-developed anterior and posterior many-cusped cingula and three transverse cusps joined by a prominent ridge. Thus it is identified as a probable juvenile Trirachodon of uncertain species. Unlike in Trirachodon adults, tall central and internal cusps of the upper postcanines lie close together on the medial side of the crown, separated from the tall external cusp by a deep valley. In these features it shows a striking resemblance to the traversodontid Scalenodon angustifrons, but not to more primitive traversodontids. The lower postcanines superficially resemble those of traversodontids in that two cusps (central and internal) are very tall and the posterior basin is elongated, but, unlike in traversodontids, the external cusp is present, though relatively small. Evidence of tooth replacement occurs in the incisors, canines, and postcanines. At least two replacement waves of gomphodont teeth are indicated, as well as replacement of small, possibly sectorial teeth at the rear of the tooth row. Probable homology of (at least) the external and internal cusps in the three gomphodont families suggests that the common ancestor also possessed transversely-expanded crowns developed from an external sectorial position (homologous with the ancestral blade-like tooth) and a hypertrophied internal cingulum.
A new deep-bodied Late Permian actinopterygian fish from the Beaufort Group, South Africa
(BERNARD PRICE INSTITUTE FOR PALAEONTOLOGICAL RESEARCH, 2005) Bender, Patrick
Anew genus of actinopterygian (ray-finned) fish, Blourugia seeleyi is described from Late Permian (Tatarian) fluvio-lacustrine, siltstone dominated deposits within the lower Beaufort Group of South Africa. It was originally provisionally assigned to the globally known genus Atherstonia by Woodward (1893), but indications are that the genus is distinct from Atherstonia on the basis of its deep-bodied form and the associated skull characters; thus Blourugia seeleyi is placed in Gardiner&Schaeffer’s (1989) Platysomus Group. The new genus is characterized by a uniquely shaped prominent high triangular posterior blade of the maxilla, dermosphenotic triangular shaped, pointed marginal teeth, 8–10 branchiostegal rays, flank scales that exhibit a well-developed dermal ornamentation consisting of numerous transverse ganoine ridges, and the presence of a dermopterotic that contacts the nasal. Blourugia appears to be a primitive deep-bodied form, basal to lower actinopterygian deep-bodied forms such as Adroichthys, Amphicentrum, Cheirodopsis, Paramesolepis and Platysomus. As a member of Gardiner & Schaeffer’s Platysomus Group, it is therefore derived relative to stem-actinopterans such as Howqualepis, Mimia and Moythomasia, and also derived relative to earlier southern African Palaeozoic actinopterygians such as Atherstonia scutata, Mentzichthys jubbi, Namaichthys schroederi and the newly/recently described lower Beaufort Group taxa Bethesdaichthys kitchingi and Kompasia delaharpei, but basal to stem-neopterygians such as Australosomus and Saurichthys.
On the stratigraphic range of the dicynodont taxon Emydops (Therapsida: Anomodontia) in the Karoo Basin, South Africa
(BERNARD PRICE INSTITUTE FOR PALAEONTOLOGICAL RESEARCH, 2005) Angielczyk, Kenneth D.; Fröbisch, Jörg; Smith, Roger M. H.
The dicynodont specimen SAM-PK-708 has been referred to the genera Pristerodon and Emydops by various authors, and was used to argue that the first appearance of Emydops was in the Tapinocephalus Assemblage Zone in the Karoo Basin of South Africa. However, the specimen never has been described in detail, and most discussions of its taxonomic affinities were based on limited data. Here we redescribe the specimen and compare it to several small dicynodont taxa from the Tapinocephalus and Pristerognathus assemblage zones. Although the specimen is poorly preserved, it possesses a unique combination of features that allows it to be assigned confidently to Emydops. The locality data associated with SAM-PK-708 are vague, but they allow the provenance of the specimen to be narrowed down to a relatively limited area southwest of the town of Beaufort West. Strata from the upper Tapinocephalus Assemblage Zone and the Pristerognathus Assemblage Zone crop out in this area, but we cannot state with certainty from which of these biostratigraphic divisions the specimen was collected. Nevertheless, SAM-PK-708 is an important datum because it demonstrates that the stratigraphic range of Emydops must be extended below its widely-accepted first appearance in the Tropidostoma Assemblage Zone. This range extension is significant because it implies that the divergence between the emydopid and dicynodontid lineages must have occurred no later than Pristerognathus Assemblage Zone times, and that most of the major lineages of Permian dicynodonts had emerged by a relatively early point in the history of the group.
Fossil hyraxes (Hyracoidea: Mammalia) from the Late Miocene and Plio-Pleistocene of Africa, and the phylogeny of the Procaviidae
(BERNARD PRICE INSTITUTE FOR PALAEONTOLOGICAL RESEARCH, 2005) Pickford, Martin
A palate with much of the dentition from Aragai, Lukeino Formation (6 Ma) Kenya, is the most complete known specimen of a Late Miocene procaviid hyracoid. It shares several features with Dendrohyrax. The specimen is as large as the western tree hyrax, Dendrohyrax dorsalis, but it is attributed to a new species. D. dorsalis ranges through the tropical forests of Central and Western Africa, from Uganda to Gambia. As such the presence of a similar species at Lukeino provides evidence of the humid forest nature of the palaeoenvironment in the Tugen Hills during the Late Miocene. The fossil hyracoid specimens from the Early Pliocene of Langebaanweg, South Africa, are close in morphology to, but somewhat larger than, the extant bush hyrax, Heterohyrax brucei, but have some derived characters found in Procavia capensis. The cheek teeth are brachyodont, the lower premolar row is complete with a well-developed p/1 and there is a long diastema between the second incisor and the first premolar, all features recalling Heterohyrax. However, the depth of the mandible, the hypsodonty of the lower incisor, and the length of the premolar row relative to the length of the molar row are similar to the condition in Procavia and attest to the onset of molar enlargement relative to the rest of the dentition. In the overall context of the Procaviidae, Procavia is the most derived genus, and the presence of a few Procavia-like features in the Langebaanweg fossils indicate that the species concerned was probably already evident on the Procavia lineage, but the presence of several plesiomorphic characters reveals that it is a primitive member of the lineage. These also reveal that the specimens do not belong to Procavia cf. antiqua into which they were previously tentatively classified by Hendey (1976) as they are somewhat more derived. The detailed systematic status of the large extinct hyracoid Gigantohyrax maguirei, Kitching, 1965, from Pliocene cave fillings at Makapansgat, South Africa, has not previously been satisfactorily demonstrated, even though it is clear that most authors have considered it to be a procaviid closely related to Procavia. Kitching (1965) compared it only to species of Procavia. Re-study of the original sample, as well as additional fossils (three partial skulls, isolated upper premolar, fragment of mandible with a premolar) reveal that Gigantohyrax shares many features with the genus Dendrohyrax, fewer with Heterohyrax and even fewer with Procavia. It is concluded that among the Procaviidae, Gigantohyrax is most closely related to Dendrohyrax. The new discoveries of Late Miocene and Pliocene procaviids inKenya and SouthAfrica, when added to recently described associated upper and lower dental elements of Meroehyrax bateae from the base of the Middle Miocene of Uganda, permit a reappraisal of procaviid phylogeny. It is concluded that procaviids probably descended from Saghatheriidae, and that Pliohyracidae did not give rise to procaviids as previously thought by some authors.
Biostratigraphy of the lower Burgersdorp Formation (Beaufort Group; Karoo Supergroup) of South Africa – implications for the stratigraphic ranges of early Triassic tetrapods
(BERNARD PRICE INSTITUTE FOR PALAEONTOLOGICAL RESEARCH, 2005) Neveling, J.; Hancox, P. J.; Rubidge, B. S.
The Beaufort Group (Karoo Supergroup) of South Africa comprises a thick sequence of fluvio-lacustrine sedimentary rocks that accumulated in a landlocked, intracratonic foreland basin in southwestern Gondwana during the Middle Permian to Middle Triassic. To the south this basin was bounded by the Cape Fold Belt, which acted as the major source of both sediment and discharge. Rocks of the Beaufort Group are renowned for their rich fossil record and eight tetrapod-based biozones are currently recognized. The uppermost two biozones of the Beaufort Group, the Lystrosaurus and Cynognathus assemblage zones, record terrestrial biotic recovery following the Permo-Triassic mass extinction event. Stratigraphic overlap between these biozones occurs in the proximal sector, but their separation by an unconformity in the distal sector reflects the incomplete preservation of the sequence in this part of the basin. Our results afford chronostratographic control that impacts on current theories on the development of the Karoo Basin, and on the relative age of the sequence.
The ownership of the Taung skull and of other fossil hominids and the question of repatriation
(Bernard Price Institute for Palaeontological Research, 2005-12) Tobias, Phillip V.
The ownership of fossils, and for purposes of this paper I refer to that of hominid fossils, was long assumed to be vested in the individuals who made the discoveries. The author reviews here a series of case histories with which he has had direct or indirect personal contact, that illustrate claims for ownership. Some have been explicit, some implicit. They are drawn from South Africa, East Africa, North Africa, England, France, Germany, Italy, Russia, the Netherlands, Indonesia and China. This historical essay reviews the replacement of this practice by a policy that fossils are not seen as personal property, ut as part of the heritage of the country of origin. During the colonial era, many specimens were removed from former colonies to the ‘home countries’, where they remained for decades, at least until the subject territories attained their independence from the former imperial powers. The new policy about ownership, in such cases, entails the return (repatriation) of the expatriate fossils to the source country. Examples of success stories and of tardy responses are given. A policy for the future is set forth.

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