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Revision of Amygdalodon Patagonicus Cabrera, 1947 (Dinosauria, Sauropoda)

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Revision of Amygdalodon Patagonicus Cabrera, 1947 (Dinosauria, Sauropoda) Mitt. Mus. Nat.kd. Berl., Geowiss. Reihe 6 (2003) 173-181 10.11.2003 Revision of Amygdalodon patagonicus Cabrera, 1947 (Dinosauria, Sauropoda) Oliver W. M. Rauhut’ With 3 figures Abstract The type material of the oldest known sauropod dinosaur from South America, Amygdulodon patagonicus, from the Cerro Carnerero Formation (Toarcian-Bajocian) of Chubut province, Argentina, is reviewed. The material includes elements of at least two individuals plus a remain of another, indeterminate vertebrate, and a posterior dorsal vertebra is designated as the lectotype of Amygdalodon. All of the sauropod material originally referred to this species represents very basal, non-neo- sauropodan eusauropods. Amygdalodon is the only South American evidence for a probable Early to early Middle Jurassic global dispersal of basal eusauropodan sauropods. Key words: Dinosauria, Sauropoda, Early to Middle Jurassic, Argentina. Zusammenfassung Das Typusmaterial des altesten siidamerikanische sauropoden Dinosauriers, Amygdalodon putugonicus von der Cerro Carner- ero Formation (Toarc-Bajoc) der Provinz Chubut, Argentinien, wird revidiert. Das Material enthalt Elemente von mindestens zwei Individuen sowie eines anderen, nicht identifizierbaren Wirbeltieres. Ein hinterer Dorsalwirbel wird als Lektotypus von Amygdulodon bestimmt. Das gesamte ursprunglich zu diesem Taxon gestellte Sauropodenmaterial reprasentiert sehr basale Eusauropoden auRerhalb der Neosauropoda. Amygdalodon ist der einzige siidamerikanische Hinweis auf eine vermutlich globale Verbreitung der Eusauropoden im unteren bis friihen mittleren Jura. Schliisselworter: Dinosauria, Sauropoda, Unterer bis Mittlerer Jura, Argentinien. Introduction Despite its potential importance as South America’s oldest known sauropod, Amygdalodon In 1947, Cabrera named some fragmentary re- has received surprisingly little attention in the mains from Jurassic strata in the province of literature, possibly due to the fragmentary nature Chubut, Argentina, as a new sauropod taxon, of the material. Since Casamiquela’s redescrip- Amygdalodon patagonicus. The material came tion of the taxon, it was only mentioned in some from the Cerro Carnerero Formation at Pampa rather general overview articles of South Ameri- de Agnia and represented the first dinosaur de- can vertebrates or dinosaurs (e.g. Bonaparte scribed from the Jurassic of South America and 1978, 1981, 1996, Novas 1997) or sauropods in one of the oldest sauropod occurrences then general (e.g. McIntosh 1990a). Amygdalodon has known. Casamiquela (1963) redescribed all the generally been regarded as a member of the original material and referred several other ele- Cetiosauridae (Cabrera 1947, Casamiquela 1963, ments in the collections of the Museo de La Pla- Bonaparte 1978, McIntosh 1990a), an assem- ta, Argentina, to the same taxon. The latter ma- blage of basal sauropods that is now considered terial had been collected by the geologist A. to be paraphyletic (Upchurch 1995, 1998, Wilson Piatnitzky prior to the holotype, apparently from 2002). Although listed in his work as a valid tax- the same locality, and Casamiquela argued that it on, McIntosh (1990a) also indirectly doubted its probably represents the same individual. validity, stating that the elements “indicate cetio- ’ Institut fiir Palaontologie, Museum fur Naturkunde, InvalidenstraRe 43, 10115 Berlin, Germany, and Museo Paleontobgi- co Egidio Feruglio, Fontana 140, 9100 Trelew, Argentina. Received May, accepted July 2003 174 Rauhut, 0. W. M., Revision of Amygddodon pafagonicus saurid affinities but do not serve to clearly differ- In the following descriptions, the terminology entiate this animal from the other members of of the vertebral laminae follows that of Wilson this family” (p. 378). (1999), and that of the lateral vertebral recesses The age of the material is still uncertain. that of Makovicky (1997). Although usually cited as Bajocian (e.g. McIn- tosh 1990a, Novas 1997), it has also been re- garded as Early Jurassic by some authors (e.g. The type material Bonaparte 1981). The Cerro Carnerero Forma- tion, which yielded the material, is the basal for- Te e t h : Of the originally four complete and three mation of the Lonco Trapial Group and uncon- fragmentary teeth of Amygdulodon, only one formably overlies marine units of early Toarcian almost complete element could be located in the age (Riccardi et al. 1992). It is overlain by the collections of the La Plata Museum (Fig. lA, B). Lonco Trapial (= Caiiadh Puelman) Formation, It has an asymmetrical spatulate crown with a a unit of mainly volcanic and volcanoclastic well-developed wear-facet on the distal carina api- rocks. The age of the Lonco Trapial Group is cally, which is more strongly developed laterally generally considered to be Toarcian to Middle than lingually. No marginal denticles are present Jurassic (probably Bathonian in the area con- on the anterior carina up to where it meets the cerned; Riccardi et al. 1992, Figari & Courtade posterior wear-facet. Well-developed longitudinal 1993). Page et al. (1999: 466) noted that the vol- grooves parallel to the carina are present lingually canism in this area started in the Aalenian, so a and laterally along the anterior carina and at least late Toarcian to Aalenian age seems to be most laterally along the posterior carina. Apart from likely for the Cerro Carnerero Formation, but the marginal groove, the lingual side of the crown more data is needed to confirm this. is slightly convex mesio-distally, as in Curdiodon The purpose of the present paper is to review (Upchurch & Martin 2003). The root is considera- the original holotype and referred material of bly longer and mesio-distally narrower than the Amygdulodon patugonicus and to evaluate its crown and approximately round in cross-section. systematic position on the basis of modern ideas Cervical vertebrae: Only the body of an of sauropod phylogeny (Upchurch 1995, 1998, anterior cervical vertebra and an isolated prezy- Wilson & Sereno 1998, Wilson 2002). gapophysis of a more posterior cervical are pre- Institution a1 a b br evi at i o n : MLP, Museo sent. de La Plata, La Plata, Argentina. The preserved centrum represents an anterior or middle cervical (Fig. 1C-E). The centrum is opisthocoelous, elongate and very strongly flexed The material of Amygdalodon ventrally in its anterior part, so that the anterior articular surface is strongly offset from the pos- The material referred to Amygdulodon is very terior. The parapophyses are elongate, oval in fragmentary. Cabreras’ (1947) original material shape and placed on the anteroventral edge of (MLP 46-VIII-21-1) consisted of several teeth, a the centrum. A small ridge runs from the poster- cervical, two dorsal and two caudal vertebral ior end of the parapophysis posteriorly along the centra and some vertebral fragments, several ventral edge of the lateral side and disappears at fragmentary ribs, a partial right pubis, and a approximately half the length of the centrum. A plate-like bone that Cabrera identified as a frag- pleurocoel is absent; only a large, very shallow ment of the scapula (Figs 1, 2). Casamiquela depression is present on the sides and is deeper (1963) referred another dorsal vertebra, a sacral anteriorly than posteriorly. Likewise, no excava- vertebral centrum, and a distal end of a tibia, tions are present on the dorsal surfaces of the apparently from the same locality as the type, to parapophyses. In ventral view, the centrum is this taxon (MLP 36-XI-10-3; Fig. 3). For meas- strongly constricted behind the parapophyses urements of the elements see Casamiquela and has a well-developed, sharp ventral keel in (1963). its flattened anterior half. The posterior half of Fig. 1. Type material of Aniygdalodon parctgorlrcus Cabrera. 1947. A. B. Tooth, MLP 46-VIII-21-1/13, in lingual (A) and labial (B) view. C. D, E. Cervical vertebra, MLP 46-VIII-21-1/8. in right lateral (C, stereopair), ventral (D) and left lateral (E) view. F, G. Right cervical prezygapophysis, MLP 46-VIII-21-117. in dorsal (F) and medial (G) view. H. Anterior dorsal neural spine, MLP 46-VIII-21-116. in posterior view. I, J, K. Posterior dorsal vertebra, MLP 46-VIII-21-112, lectotype, in right lateral view Mitt. Mus. Nat.kd. Berl., Geowiss. Reihe 6 (2003) 175 (I, stereopair), right lateral view, with right wall of neural canal remived (J), and posterior (K) view. L, M. Caudal vertebra, MLP 46-VIII-21-1/3, in left lateral (L) and posterior (M) view. N. Caudal vertebra, MLP 46-VIII-21-1/4, in right lateral view. 0. Proximal fragment of a dorsal rib, MLP 46-VIII-21-1/9, in anterolateral view. Abbreviations: acdl, anterior centrodiapophy- seal lamina; cprl, centroprezygapophyseal lamina; g, groove; k, ventral keel; pcdl, posterior centrodiapophyseal lamina; pod, postzygodiapophyseal lamina; poz, postzygapophysis; pp, parapophysis; prdl, prezygodiapophyseal lamina; spol, spinopostzga- pophyseal lamina; sprl, spinoprezygapophyseal lamina; tprl, intraprezygapophyseal lamina. Scale bars 1cm (A, B) and 10 cm (C-0). 176 Rauhut, 0. W. M., Revision of Amygdalodon patagonicus the ventral side is transversely rounded and with- shape in combination with the presence of spino- out a keel. prezygapophyseal and spinopostzygapophyseal Only the bases of the neural arch with the laminae clearly indicate that it is one of the ante- neural canal is present. The latter is collapsed, so riormost dorsal neural spines (Fig. 1H). The that not much can be said about its shape, but it spine is strongly flattened anteroposteriorly and seems to have been relatively small in compari- transversely expanded dorsally, although this ex- son with the vertebral centrum. Most of the ver- pansion terminates a short way below the distal tebral laminae are broken off and only the bases end, above which the tip of the spine is broadly of some are visible, better preserved on the right rounded. A small anterior and posterior projec- than on the left side. The anterior centrodiapo- tion is present at the distalmost tip of the spine, physeal lamina is slender and short and does not probably for the attachment of an interspinal reach the anterior end of the centrum.
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