Blackwell Publishing LtdOxford, UKZOJZoological Journal of the Linnean Society0024-40822007 The Linnean Society of London? 2007 1493 477492 Original Article THE OLDEST CYNODONTJ. BOTHA ET AL. Zoological Journal of the Linnean Society, 2007, 149, 477–492. With 6 figures The oldest cynodont: new clues on the origin and early diversification of the Cynodontia J. BOTHA1*, F. ABDALA2 and R. SMITH3 1Karoo Palaeontology, National Museum, PO Box 266, Bloemfontein, 9300, South Africa 2Bernard Price Institute for Palaeontological Research, University of the Witwatersrand, Johannesburg, Wits, 2050, South Africa 3Iziko: South African Museum of Cape Town, PO Box 61, Cape Town, 8000, South Africa Received August 2005; accepted for publication June 2006 Early mammaliaforms and their extinct relatives, nonmammaliaform cynodonts, have long been the focus of intense research in attempting to unravel how and when major changes toward mammalness occurred. The earliest well- known representatives of cynodonts are latest Late Permian in age. Here, we describe Charassognathus gracilis gen. et sp. nov., from the early Late Permian of South Africa, representing the oldest cynodont yet found. This spec- imen displays a notch on the dentary in the same location as the base of the masseteric fossa in the basal cynodonts Procynosuchus and Dvinia, and represents the first indication in theriodonts of an invasion of occlusal musculature onto the dentary. A phylogenetic analysis of seven therocephalians and ten non-mammaliaform cynodonts and equally weighted characters resulted in nine most parsimonious trees, the strict consensus of which shows a basal polytomy in cynodonts, including Charassognathus, Dvinia, Procynosuchus and a clade including the remaining cyn- odonts. The basal polytomy in the majority rule consensus tree is reduced, as Procynosuchus and Dvinia form a clade. One most parsimonious tree, from an analysis using implied weights, positions Charassognathus as the most basal cynodont. This result implies that the Cynodontia initially diversified in Permian Gondwana, in what is now south- ern Africa. © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society, 2007, 149, 477–492. ADDITIONAL KEYWORDS: Karoo Basin – non-mammaliaform cynodont – Permian – South Africa. INTRODUCTION 2000). Procynosuchus is traditionally recognized as the earliest known cynodont (Rubidge & Sidor, 2001; The Cynodontia has long been recognized as a crucial Sidor & Smith, 2004) ranging from the base to the top therapsid group because it includes mammals as its of the Dicynodon Assemblage Zone [AZ] (Wuchiapin- extant subclade (Rubidge & Sidor, 2001). Extinct gian–Changhsingian) in South Africa (Rubidge, 1995; members of the group form a paraphyletic assemblage Sidor & Smith, 2004) and in biostratigraphically known as non-mammaliaform cynodonts and repre- equivalent levels in East Africa, western Germany sent the stage at which many mammalian character- (von Huene, 1950; Kemp, 1979; Sues & Boy, 1988) and istics first appeared in therapsids. The earliest Russia (= Cyrbasiodon; Tatarinov, 2004). A new cyn- cynodont records are known from the latest Late Per- odont from the older Tropidostoma AZ (early Late Per- mian and include the genera Procynosuchus from mian) described here indicates a South African origin South Africa, Tanzania, Germany and Russia (Kemp, for cynodonts. 1979; Sues & Boy, 1988; Rubidge, 1995; Tatarinov, 2004), Cynosaurus and Nanictosaurus from South Africa (Rubidge, 1995; Van Heerden & Rubidge, 1990), MATERIAL AND METHODS and Dvinia, Uralocynodon and Nanocynodon from The specimen described herein, SAM-PK-K10369 (in Russia (Tatarinov, 1968, 1987; Battail & Surkov, the collections of the Iziko: South African Museum, Cape Town, South Africa), consists of a complete, lat- *Corresponding author. E-mail: [email protected] erally compressed skull, a partially preserved axis and © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society, 2007, 149, 477–492 477 478 J. BOTHA ET AL. third cervical vertebra, and articulated femur, tibia and Dvinia by the absence of maxillary precanine and fibula. Institutional abbreviations and specimens teeth. The anterior postcanines have a single high of the therocephalians and basal cynodonts studied for cusp in labial view, whereas the posterior postcanines comparative purposes are presented in Appendices 1 exhibit a main cusp directed slightly backward, and and 2, respectively. tiny anterior and posterior accessory cusps, with the A cladistic analysis using the program TNT anterior accessory cusps positioned slightly higher (Goloboff, Farris & Nixon, 2003) was performed to than the posterior accessory cusps. determine the phylogenetic position of Charas- Comments: The combined presence of the following sognathus. A data matrix was constructed for 59 characters indicates that C. gracilis is a cynodont: cranio-dental characters and 18 theriodont taxa. multicuspidated postcanines; groove in the squamosal Cyonosaurus, ‘one of the rare gorgonopsian skulls to for the quadratojugal; two occipital condyles; elon- have been thoroughly studied’ (Sigogneau-Russell, gated ascending process of the epipterygoid; frontal 1989: 83), was used to root the cladograms. In addition excluded from the orbital margin; prominent angle of to ten cynodont taxa, seven therocephalian taxa were the dentary; reflected laminar of the angular with included in the analysis, as Therocephalia is a sister small lateral crests and femoral head set off dorsally group of the Cynodontia. A heuristic search was per- from the shaft (see Description for more details). formed with all characters having equal weights. This search consisted of ten random addition sequences Holotype: SAM-PK-K10369 consists of a complete (ten Wagner trees randomizing the order of the termi- skull with lower jaw in occlusion, the axis, third cer- nals) and tree-bisection-reconnection swapping, stor- vical vertebra, ?cervical ribs and a few indeterminate ing ten trees per replication. The run was performed bones. Associated material includes a right articulated with collapsing rule one (Coddington & Scharff, 1994), femur, tibia and fibula. which collapses branches with ambiguous support. Etymology: Greek, charasso, prefix meaning ‘notch’ Increasing the number of replicates did not change the and gnathus, ‘jaw’, referring to the characteristic result obtained. A second analysis was performed with notch on the dentary. The specific epithet, gracilis, similar settings, but using implied weights (Goloboff, Latin, meaning ‘slender’, refers to the delicate nature 1993). The weighting was made by means of a con- of the specimen. stant of concavity K. A possible outcome is the decrease in the number of most parsimonious trees by Locality and horizon: The specimen was found in a reducing the influence of homoplastic characters. roadside cutting of highway R353 in Teekloof Pass, Characters showing many extra steps in the most par- between the towns Leeu Gamka and Fraserburg in the simonious trees are thus down-weighted in relation to Beaufort West District, Western Cape Province, South the characters that better fit those trees. Analyses Africa. The locality lies on the farm Willowdene, a por- were performed with the constant of concavity set at tion of Beato 238 (Fig. 1); exact coordinates are given intermediate and low values. in the Iziko South African Museum. It is stratigraph- Material examined and literature consulted for each ically positioned in the upper half of a thick mudrock- taxon included in the cladistic analysis is presented in dominated succession of fluvio-lacustrine strata, Appendix 2, the list of characters and data matrix in known locally as the Hoedemaker Member of the Appendix 3, and unambiguous synapomorphies for Teekloof Formation (Rubidge, 1995; Figs 1A, 2A), nodes of the most parsimonious tree obtained with which is considered to be early Late Permian (Wuchi- implied weights are presented in Appendix 4. apingian) in age (Catuneanu et al., 2005). The few lat- erally extensive sandstone bodies in the Hoedemaker Member (Fig. 2B) are interpreted as having been SYSTEMATIC PALAEONTOLOGY deposited as amalgamated point bars within high-sin- uosity Mississippi-sized meandering rivers (Smith, THERAPSIDA BROOM, 1905 1987). Most of the vertebrate fossils are found in the CYNODONTIA OWEN, 1861 thick greenish-grey massively bedded siltstone with CHARASSOGNATHUS GRACILIS GEN. ET SP. NOV. minor mudstone intercalations that occur between the Diagnosis: This new cynodont is distinguished by the main channel sandstones. These sediments occur in 5– presence of a small notch in the base of the coronoid 10-m-thick coarsening-upward sequences interpreted process, located in a similar position to the base of the as prograding crevasse splay sequences. They were masseteric fossa in Dvinia and Procynosuchus. The laid down by repeated overbank flood events emanat- angle of the dentary is prominent, appearing more ing from the channel banks and ponding in the low- developed than in Procynosuchus and Dvinia, but less land flood basins. The new cynodont specimen was so than in Nanictosaurus. The dental formula is ?4·1· recovered from a 0.25-m-thick lens of fissile, dark pur- 8/3·1·?, and it is distinguished from Procynosuchus ple, mudstone on a parting between two 1-m-thick © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society, 2007, 149, 477–492 THE OLDEST CYNODONT 479 A Hoedemaker Member Africa Beaufort Group N Johannesburg South Africa 30° Fraserburg