Early Sauropodomorph Research Articles

An early trend towards gigantism in Triassic sauropodomorph dinosaurs.

Dinosaurs dominated the terrestrial ecosystems for more than 140 Myr during the Mesozoic era, and among them were sauropodomorphs, the largest land animals recorded in the history of life. Early sauropodomorphs were small bipeds, and it was long believed that acquisition of giant body size in this clade (over 10 tonnes) occurred during the Jurassic and was linked to numerous skeletal modifications present in Eusauropoda. Although the origin of gigantism in sauropodomorphs was a pivotal stage in the history of dinosaurs, an incomplete fossil record obscures details of this crucial evolutionary change. Here, we describe a new sauropodomorph from the Late Triassic of Argentina nested within a clade of other non-eusauropods from southwest Pangaea. Members of this clade attained large body size while maintaining a plesiomorphic cyclical growth pattern, displaying many features of the body plan of basal sauropodomorphs and lacking most anatomical traits previously regarded as adaptations to gigantism. This novel strategy highlights a highly accelerated growth rate, an improved avian-style respiratory system, and modifications of the vertebral epaxial musculature and hindlimbs as critical to the evolution of gigantism. This reveals that the first pulse towards gigantism in dinosaurs occurred over 30 Myr before the appearance of the first eusauropods.

Forelimb muscle and joint actions in Archosauria: insights from Crocodylus johnstoni (Pseudosuchia) and Mussaurus patagonicus (Sauropodomorpha).

Many of the major locomotor transitions during the evolution of Archosauria, the lineage including crocodiles and birds as well as extinct Dinosauria, were shifts from quadrupedalism to bipedalism (and vice versa). Those occurred within a continuum between more sprawling and erect modes of locomotion and involved drastic changes of limb anatomy and function in several lineages, including sauropodomorph dinosaurs. We present biomechanical computer models of two locomotor extremes within Archosauria in an analysis of joint ranges of motion and the moment arms of the major forelimb muscles in order to quantify biomechanical differences between more sprawling, pseudosuchian (represented the crocodile Crocodylus johnstoni) and more erect, dinosaurian (represented by the sauropodomorph Mussaurus patagonicus) modes of forelimb function. We compare these two locomotor extremes in terms of the reconstructed musculoskeletal anatomy, ranges of motion of the forelimb joints and the moment arm patterns of muscles across those ranges of joint motion. We reconstructed the three-dimensional paths of 30 muscles acting around the shoulder, elbow and wrist joints. We explicitly evaluate how forelimb joint mobility and muscle actions may have changed with postural and anatomical alterations from basal archosaurs to early sauropodomorphs. We thus evaluate in which ways forelimb posture was correlated with muscle leverage, and how such differences fit into a broader evolutionary context (i.e. transition from sprawling quadrupedalism to erect bipedalism and then shifting to graviportal quadrupedalism). Our analysis reveals major differences of muscle actions between the more sprawling and erect models at the shoulder joint. These differences are related not only to the articular surfaces but also to the orientation of the scapula, in which extension/flexion movements in Crocodylus (e.g. protraction of the humerus) correspond to elevation/depression in Mussaurus. Muscle action is highly influenced by limb posture, more so than morphology. Habitual quadrupedalism in Mussaurus is not supported by our analysis of joint range of motion, which indicates that glenohumeral protraction was severely restricted. Additionally, some active pronation of the manus may have been possible in Mussaurus, allowing semi-pronation by a rearranging of the whole antebrachium (not the radius against the ulna, as previously thought) via long-axis rotation at the elbow joint. However, the muscles acting around this joint to actively pronate it may have been too weak to drive or maintain such orientations as opposed to a neutral position in between pronation and supination. Regardless, the origin of quadrupedalism in Sauropoda is not only linked to manus pronation but also to multiple shifts of forelimb morphology, allowing greater flexion movements of the glenohumeral joint and a more columnar forelimb posture.

Endocast of the Late Triassic (Carnian) dinosaur Saturnalia tupiniquim: implications for the evolution of brain tissue in Sauropodomorpha

The evolutionary history of dinosaurs might date back to the first stages of the Triassic (c. 250–240 Ma), but the oldest unequivocal records of the group come from Late Triassic (Carnian – c. 230 Ma) rocks of South America. Here, we present the first braincase endocast of a Carnian dinosaur, the sauropodomorph Saturnalia tupiniquim, and provide new data regarding the evolution of the floccular and parafloccular lobe of the cerebellum (FFL), which has been extensively discussed in the field of palaeoneurology. Previous studies proposed that the development of a permanent quadrupedal stance was one of the factors leading to the volume reduction of the FFL of sauropods. However, based on the new data for S. tupiniquim we identified a first moment of FFL volume reduction in non-sauropodan Sauropodomorpha, preceding the acquisition of a fully quadrupedal stance. Analysing variations in FFL volume alongside other morphological changes in the group, we suggest that this reduction is potentially related to the adoption of a more restricted herbivore diet. In this context, the FFL of sauropods might represent a vestigial trait, retained in a reduced version from the bipedal and predatory early sauropodomorphs.

Biostratigraphic significance of a new early sauropodomorph specimen from the Upper Triassic of southern Brazil

The Wachholz site (Caturrita Formation, Late Triassic), in Agudo, Rio Grande do Sul (RS), southern Brazil, has yielded several sauropodomorphs. This includes CAPPA/UFSM 0002, described here based on associated elements from the basalmost portion of the site. The specimen possesses a set of traits shared with typical ‘prosauropods’: a concave caudal margin of the trunk neural spines and a broadly convex proximal end of metacarpal V. However, it also retains some plesiomorphic features, for instance, the slender pedal digit I. Some bones closely resemble those of Unaysaurus tolentinoi, the other definitive sauropodomorph from the Caturrita Formation, an affinity corroborated by a new phylogenetic analysis. An updated biostratigraphic framework correlates the Wachholz, Água Negra (São Martinho da Serra/RS) and Botucaraí Hill (Candelária/RS) sites based on their sample of sauropodomorphs. In addition, the record of Jachaleria in the Botucaraí Hill site, a dicynodont also known from early Norian deposits of Argentina, indicates an equivalent age to deposits bearing U. tolentioni. Accordingly, a more constrained age is proposed for the Água Negra site. This is important as the early Norian marks the transition from an epoch of low sauropodomorphs representativeness to a period of extreme abundance of the group in Early-to-Middle Mesozoic ecosystems.

A new sauropod dinosaur from the Middle Jurassic of the United Kingdom.

A new record of a sauropodomorph dinosaur is here described from the Middle Jurassic (Aalenian) Saltwick Formation of Whitby (Yorkshire), UK. A single caudal vertebra represents an early sauropodomorph and signifies the earliest recognised eusauropod dinosaur from the United Kingdom. The absence of pleurocoels and a narrow, dorsoventrally deep, but craniocaudally short centrum, suggests a primitive sauropodomorph. Distinct spinopostzygopophyseal laminae rise from the lateral margins of the postzygapophyses and pass caudally along what remains of the neural spine, a character unique to a subgroup of sauropods that includes Barapasaurus, Omeisaurus and other neosauropods and eusauropods. The lack of phylogenetically robust characters in sauropod caudal vertebrae usually makes it difficult to establish affinities, but the absence of mild procoely excludes this specimen from both Diplodocoidea and Lithostrotia. The vertebra cannot be further distinguished from those of a wide range of basal sauropods, cetiosaurids and basal macronarians. However, this plesiomorphic vertebra still signifies the earliest stratigraphic occurrence for a British sauropod dinosaur.

The femoral anatomy of Pampadromaeus barberenai based on a new specimen from the Upper Triassic of Brazil

Femora of the holotype and only previously known specimen of the early sauropodomorph Pampadromaeus barberenai are incomplete and fractured. Here, we describe a new complete femur assigned to Pampadromaeus barberenai, quarried from the same outcrop of original type specimen: ‘Janner’ site, (Carnian of the Paraná Basin). It shares with the holotype of Pampadromaeus barberenai a unique combination of features: anteromedial tuber slightly posteromedially displaced; lesser trochanter proximodistally oriented with the proximal tip gently connected to the femoral shaft; fourth trochanter located on the medial margin along the mediolateral axis of the femur and with an assymetrical outline. In addition, we propose a femoral autapomorphy for Pampadromaeus barberenai: lateral condyle of the distal end of the femur extremely reduced lateromedially. Two distinct phylogenetic analyses place the new specimen as the sister-group of the Pampadromaeus barberenai type-specimen and, in one of them, they are nested within Sauropodomorpha, corroborating the original description and taxonomic assignation.

Clavicles, interclavicles, gastralia, and sternal ribs in sauropod dinosaurs: new reports from diplodocidae and their morphological, functional and evolutionary implications.

Ossified gastralia, clavicles and sternal ribs are known in a variety of reptilians, including dinosaurs. In sauropods, however, the identity of these bones is controversial. The peculiar shapes of these bones complicate their identification, which led to various differing interpretations in the past. Here we describe different elements from the chest region of diplodocids, found near Shell, Wyoming, USA. Five morphotypes are easily distinguishable: (A) elongated, relatively stout, curved elements with a spatulate and a bifurcate end resemble much the previously reported sauropod clavicles, but might actually represent interclavicles; (B) short, L-shaped elements, mostly preserved as a symmetrical pair, probably are the real clavicles, as indicated by new findings in diplodocids; (C) slender, rod-like bones with rugose ends are highly similar to elements identified as sauropod sternal ribs; (D) curved bones with wide, probably medial ends constitute the fourth morphotype, herein interpreted as gastralia; and (E) irregularly shaped elements, often with extended rugosities, are included into the fifth morphotype, tentatively identified as sternal ribs and/or intercostal elements. To our knowledge, the bones previously interpreted as sauropod clavicles were always found as single bones, which sheds doubt on the validity of their identification. Various lines of evidence presented herein suggest they might actually be interclavicles - which are single elements. This would be the first definitive evidence of interclavicles in dinosauromorphs. Previously supposed interclavicles in the early sauropodomorph Massospondylus or the theropods Oviraptor and Velociraptor were later reinterpreted as clavicles or furculae. Independent from their identification, the existence of the reported bones has both phylogenetic and functional significance. Their presence in non-neosauropod Eusauropoda and Flagellicaudata and probable absence in rebbachisaurs and Titanosauriformes shows a clear character polarity. This implicates that the ossification of these bones can be considered plesiomorphic for Sauropoda. The proposed presence of interclavicles in sauropods may give further support to a recent study, which finds a homology of the avian furcula with the interclavicle to be equally parsimonious to the traditional theory that furcula were formed by the fusion of the clavicles. Functional implications are the stabilizing of the chest region, which coincides with the development of elongated cervical and caudal vertebral columns or the use of the tail as defensive weapon. The loss of ossified chest bones coincides with more widely spaced limbs, and the evolution of a wide-gauge locomotor style.

A new species of Azendohsaurus (Diapsida: Archosauromorpha) from the Triassic Isalo Group of southwestern Madagascar: cranium and mandible

Abstract: Here, we describe a new species of Azendohsaurus from the Middle–Late Triassic of Madagascar, extending the geographical range of a taxon known otherwise only by a single species from Morocco. Although Azendohsaurus has consistently been regarded as an early dinosaur (based on various advanced dental and gnathic features resembling those characterizing certain dinosaur subgroups), the relatively complete skeletal material, now available from Madagascar, argues strongly against its dinosaurian affinities. Rather, the retention of numerous primitive cranial and postcranial features indicates a surprisingly early divergence of Azendohsaurus within Archosauromorpha and an unusual mosaic of characters in this taxon. Features considered diagnostic of Sauropodomorpha thus are inferred to occur homoplastically in at least one clade of nondinosaurian archosauromorphs, indicating a complex evolution and distribution of features traditionally thought to be derived within archosaurs. Azendohsaurus has teeth resembling those of both early sauropodomorph and ornithischian dinosaurs, yet also possesses numerous inarguable basal archosauromorph cranial and postcranial attributes. This highlights the risk of uncritically referring isolated, Middle–Late Triassic (or even later), ‘leaf-shaped’ teeth with denticles to the Dinosauria. Similarly, the occurrence of such teeth in an early diverging archosauromorph indicates that specializations for herbivory originated more frequently within this clade than conventionally assumed. For example, Azendohsaurus and numerous basal sauropodomorph dinosaur taxa share an array of convergently acquired features associated with herbivory, including tooth denticles, expanded tooth crowns, a downturned dentary and the articular located at the ventral margin of the mandible. Some of these features (denticles, expanded crowns and the ventrally deflected articular) are even more widespread among archosauromorphs, including aetosaurs, silesaurs and ornithischian dinosaurs. A downturned dentary also occurs in Trilophosaurus, a taxon further marked by unique specializations for herbivory, including transversely lophate, tricuspid teeth. An array of features associated with herbivory also occurs in rhynchosaurs and certain crocodilians (e.g. Simosuchus). This distribution suggests that craniodental features associated with herbivory were much more pervasive across the archosauromorph clade than previously recognized, possibly evolving at least six to eight times independently.

A primitive sauropodomorph from the upper Elliot Formation of Lesotho

AbstractA well-preserved, articulated dinosaur skeleton from southern Africa is described. The specimen comes from the upper Elliot Formation (?Hettangian) of Ha Ralekoala (Lesotho) and represents a new species:Ignavusaurus rachelisgenus et species nova. A cladistic analysis suggests thatIgnavusaurusis more derived thanThecodontosaurus–Pantydraco, but more primitive thanEfraasia.Ignavusaurusindeed shares a number of unambiguous synapomorphies with the taxa more derived thanThecodontosaurus–Pantydraco, such as a fully open acetabulum, but it is more plesiomorphic thanEfraasiaand more derived sauropodomorphs as shown by the evidence of, for instance, the distal extremity of its tibia that is is longer (cranio-caudally) than wide (latero-medially). The discovery ofIgnavusaurusincreases the known diversity of the early sauropodomorph fauna of the upper Elliot Formation, which stands as one of the richest horizons in the world in this respect.

P. M. Barrett & D. J. Batten (eds) 2007. Evolution and Palaeobiology of Early Sauropodomorph Dinosaurs. Special Papers in Palaeontology no. 77. 289 pp. London: The Palaeontological Association. Price £75.00 (paperback). ISBN 9781 4051 6933 2.

Despite the fact that these include some of the earliest recognized of all dinosaurs, the basal (= early) sauropodomorph dinosaurs have proved rather reluctant to reveal an agreed pattern of relationships in a strictly cladistic sense: do they constitute the monophyletic Prosauropoda (by …

The skull of Yunnanosaurus huangi Young, 1942 (Dinosauria: Prosauropoda) from the Lower Lufeng Formation (Lower Jurassic) of Yunnan, China

The Lower Lufeng Formation (Lower Jurassic: ?Hettangian-?Sinemurian) of Yunnan Province, China, has yielded an important and diverse fauna of terrestrial vertebrates that is dominated by early sauropodomorph dinosaurs (prosauropods and basal sauropods). Nevertheless, few of these animals have been studied in detail, undermining their potential significance in understanding sauropodomorph phylogeny, palaeobiology, and palaeoecology. Here, we present a detailed re-description of the cranial osteology of Yunnanosaurus huangi Young, 1942 and propose an emended diagnosis for this taxon on the basis of numerous autapomorphic characters (including an expanded internarial bar, unusual midline cranial bosses, and the possession of elongate maxillary tooth crowns lacking marginal serrations). Incorporation of these novel anatomical data into existing phylogenetic analyses of sauropodomorph interrelationships substantially affects the resolution, length, and topologies of the trees recovered. Although the phylogenetic position of Yunnanosaurus remains labile, these new analyses undermine previous suggestions that the former was the sister taxon of the southern African prosauropod Massospondylus. Several features of the skull of Yunnanosaurus (small external nares, cranial bosses, tooth crown morphology, and the lack of maxillary foramina) indicate that the palaeobiology of Yunnanosaurus may have been rather different from that of other prosauropods and basal sauropods, although more detailed functional studies and better material are needed to confirm this suggestion. (c) 2007 Natural History Museum, London. Journal compilation (c) 2007 The Linnean Society of London, Zoological Journal of the Linnean Society, 2007, 150, 319-341.