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Home , Bothriospondylus, Dinosaur Beds, Eucnemesaurus, Paralititan

Review and analysis of African sauropodomorph diversity

P.D. Mannion Department of Earth Sciences, University College London, WC1E 6BT, U.K. E-mail: [email protected]

Introduction Sauropodomorphs were a major Mesozoic terrestrial radiation of gigantic, herbivorous . Their remains have been discovered on all continents and by the close of the they had achieved a global distribution (Galton & Upchurch 2004). African sauropodomorphs are known from the (; Raath 1996), right through to the Maastrichtian (Late ; Curry Rogers & Forster 2001), thus spanning the entire known temporal range of dinosaurs. In addition, their remains have been recovered from 16 different African countries, ranging across the entire continent (Fig. 1). Here, the sauropodomorph diversity from each geological Epoch will be summarized, with a concluding analysis of African sauropodomorph diversity through time. Figure 1. Map of African sauropodomorph localities. Late Triassic The Late Triassic of Africa is almost entirely dominated of (M. thabanensis) based on an by remains from . These include numerous isolated femur from Lesotho; however, Yates (2007b) basal sauropodomorphs ( and Plateosauravus, recently expressed doubt as to the validity of this species as well as undiagnostic remains; Van Hoepen 1920; and it is here considered a . Another basal Haughton 1924; Yates 2003, 2007a; Galton et al. 2005), sauropod, , is known from the Toarcian of several indeterminate prosauropods (including remains Morocco (Allain et al. 2004). from Zimbabwe; Raath 1996), as well as Azendohsaurus from Morocco (originally described as an ornithischian Middle and then reinterpreted as a prosauropod; Dutuit 1972; By the Middle Jurassic, African sauropodomorphs were Gauffre 1993a), which is now considered a non-dinosaurian composed entirely of sauropods, with prosauropods archosauriform (Irmis et al. 2007). Additionally, two South seemingly having become extinct. The first Madagascan African taxa originally considered as prosauropods remains are known from this time period, with the basal (Melanorosaurus and ; Haughton 1924; eusauropod Archaeodontosaurus (Buffetaut 2005) and the Galton & van Heerden 1985) have more recently been titanosauriform (Bonaparte 1986; resolved as basal sauropods (Upchurch et al. 2004, 2007; Upchurch et al. 2004) roughly contemporaneous alongside Yates 2007b). A third South African Triassic sauropod material described by Lydekker (1895) as ‘ () was named by Yates & Kitching (2003). madagascariensis’. This latter material represents a derived non-neosauropod eusauropod which is distinct from the Early Jurassic other named Madagascan taxa (Mannion, in press). The The Early Jurassic is dominated by the South African and basal macronarian is known from Morocco Zimbabwean prosauropod (Owen 1854), (Monbaron et al. 1999), as are also remains originally which is known from over 80 skeletons (Galton & described as ‘ mogrebiensis’ (Lapparent 1955), Upchurch 2004). Barrett (2004) noted the presence of which are in need of revision and are currently considered another diagnostic Early Jurassic South African sauro- undiagnostic. Most recently, Mahammed et al. (2005) podomorph, though this is considered a new species of described a basal eusauropod, Chebsaurus, from Algeria. Massospondylus (Barrett, in press) and is not included as a distinct taxon in the diversity analysis. As well as various indeterminate sauropodomorph remains, there are also Currently known Late Jurassic African sauropods are recently excavated skeletons from South Africa that have restricted to Tanzania and Zimbabwe. Most remains were been suggested to represent three new sauropodomorph collected during the German Tendaguru expeditions of taxa, including a basal sauropod (Yates et al. 2007). The 1909–1913. Diplodocoids (Australodocus, Zimbabwean taxon was originally identified and ), , the putative titanosaur as a prosauropod (Raath 1972) but has subsequently been and a taxon of unknown affinities (Tendaguria) demonstrated to be a basal sauropod (Cruickshank 1975; are all known from Tanzania (Fraas 1908; Janensch 1914; Upchurch et al. 2004). Gauffre (1993b) named a second Wild 1991; Bonaparte et al. 2000; Remes 2007). Material

108 ISSN 0078-8554 Palaeont. afr. (December 2009) 44: 108–111 Figure 2. African sauropodomorph diversity through time. Solid black line represents African sauropodomorph taxic diversity. Dashed grey line represents global sauropodomorph taxic diversity. Solid grey line represents a phylogenetic diversity estimate of global sauropod diversity. These last two diversity curves are based on Barrett & Upchurch (2005) and Upchurch & Barrett (2005). Geological timescale based on Gradstein et al. (2005). from Zimbabwe has also been referred to several of these Discussion and conclusions genera (Raath & McIntosh 1987). Thirty-one African sauropodomorphs are considered generically distinct, spanning the full temporal interval that dinosaurs existed and including representatives of all The first sauropod material known from is the main sauropodomorph clades. An African sauropodo- preserved in Early Cretaceous deposits. This is repre- morph taxic diversity curve (Fig. 2) shows peaks in the sented by the titanosaurs and Karongasaurus (Late Triassic), Hettangian-Sinemurian (Early Juras- (Haughton 1928; Jacobs et al. 1993; Gomani 2005). sic), Bathonian (Middle Jurassic) and - is known from Morocco (Lavocat 1954) and Tithonian (Late Jurassic), as well as a shallower peak in another rebbachisaurid () has been described the mid-Cretaceous. The curve also suggests that the from Niger (Sereno et al. 1999). Also described from Niger Pliensbachian-Bajocian (Early-Mid Jurassic), Oxfordian is the eusauropod (or basal macronarian; Upchurch et al. (Late Jurassic), and much of the Cretaceous were periods 2004) (Sereno et al. 1999). Material described from of apparent lower diversity. Niger, Tunisia and Algeria as ‘Rebbachisaurus tamesnensis’ In general, this matches quite closely with diversity (Lapparent 1960) may be a mixture of Nigersaurus and curves based on global taxic and phylogenetic diversity Jobaria. A brachiosaurid (‘Brachiosaurus nougaredi’) was also estimates (i.e. Barrett & Upchurch 2005; Upchurch & named by Lapparent (1960) based on Algerian remains, but Barrett 2005; Fig. 2), although the African diversity curve is not here considered valid. Lastly, indeterminate unsurprisingly shows lower diversity levels based on only sauropod material is known from South Africa, Cameroon sampling from one continent. The main difference is and Kenya (Weishampel et al. 2004, and references the lack of any known African taxa for much of the Late therein; Sertich et al. 2005; De Klerk 2008). Cretaceous, in comparison to global values. Upchurch & Barrett (2005) used dinosaur-bearing formations (DBFs) to help tease apart genuine global diversity signals from preservational biases. During the Cretaceous, DBFs were Valid,named Late Cretaceous taxa are composed entirely at their highest; however,the number of DBFs for the Afri- of titanosaurs, with Aegyptosaurus and both can Late Cretaceous (based on Weishampel et al. 2004) is described from the of Egypt (Stromer 1932; extremely low,thus this low diversity may be an artefact of Smith et al. 2001), and known from the late a poor African Late Cretaceous rock record. The overall Maastrichtian of Madagascar (Curry Rogers & Forster closeness in fit suggests that African sauropodomorphs 2001). Indeterminate remains (predominantly of titano- were at least as diverse as in other areas of the world, and saurs, but also including some possible diplodocoids) are followed similar fluctuations temporally. also known from Angola, Morocco, Niger, Sudan, Swazi- land and Tanzania (Weishampel et al. 2004, and references Thanks to P.Upchurch and an anonymous reviewer for reading earlier versions of this paper and to the Palaeontological Society of South Africa for allowing me to therein; Pereda Suberbiola et al. 2004; Jacobs et al. 2006; present my research at the 2008 meeting. This work was supported by a University O’Connor et al. 2006). College London NERC studentship (NER/S/A/2006/14347).

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