The vertebrate fauna of the Upper Permian of Niger—III, morphology and ontogeny of the hindlimb of Moradisaurus grandis (Reptilia, Captorhinidae)

We describe hindlimb elements of the large captorhinid Moradisaurus grandis (Reptilia: Captorhinidae) recently collected from the Upper Permian Moradi Formation of north-central Niger. This description is based primarily on an associated partial skeleton comprising a hemimandible, vertebral column, and partial left and nearly complete right hindlimb. Additionally, we report on a well-preserved, articulated, and essentially complete juvenile pes that provides important information on the ontogeny of the captorhinid tarsus. The hindlimb of Moradisaurus is stout and more massively built than in any other known captorhinid. The femur displays several features interpreted as adaptations to the demands of large body size, and the tibia and fibula have short, wide shafts and hypertrophied condyles and epicondyles. The astragalus is very derived, possessing two accessory ossifications and a relatively horizontal tibial articulation, indicating that the tibia was held more vertically than in other captorhinids. The calcaneum is co-ossified with distal tarsal five. The juvenile pes indicates that the captorhinid centrale arose from the fusion of two ossification centers, and that the captorhinid astragalus arose from the fusion of at least three ossification centers

The vertebrate fauna of the Upper Permian of Niger. IV. Nigerpeton ricqlesi (Temnospondyli: Cochleosauridae), and the Edopoid Colonization of Gondwana

We describe the edopoid temnospondyl Nigerpeton ricqlesi from the Upper Permian Moradi Formation of northern Niger on the basis of two partial skulls and tentatively associated postcranial material. This crocodile-like taxon displays several edopoid characters states such as a long prenarial region with enlarged premaxillae, elongated vomers, large, posteriorly tapering choanae, and a jugal that broadens anteriorly. Nigerpeton possesses a unique carnivorous dentition. It is autapomorphic in its possession of an extremely elongate snout bearing a maxillary bulge that accommodates three hypertrophied caniniform teeth, inner premaxillary tusks, and anterior paired fenestrae, which pierce the skull roof. In addition, both the maxilla and dentary tooth rows show the sporadic appearance of ‘doubled’ tooth positions. The lateral-line system is present at the adult stage, which suggests an aquatic habitat for this taxon. A phylogenetic analysis of Edopoidea and its relatives places Nigerpeton as the sister taxon to the Permo-Carboniferous genus Chenoprosopus from the U.S.A. As with other members of the Moradi tetrapod fauna, the discovery of Nigerpeton strengthens support for the hypothesis that West Africa hosted an endemic fauna at the close of the Paleozoic Era. Biogeographically, we propose that Late Carboniferous and Permian edopoids were geographically widespread and that they twice crossed the Central Pangean mountain chain (between Laurussia and Gondwana) during their evolution. This distribution was later fragmented with the onset of Late Permian climatic warming

Structural Extremes in a Cretaceous Dinosaur

Fossils of the Early Cretaceous dinosaur, Nigersaurus taqueti, document for the first time the cranial anatomy of a rebbachisaurid sauropod. Its extreme adaptations for herbivory at ground-level challenge current hypotheses regarding feeding function and feeding strategy among diplodocoids, the larger clade of sauropods that includes Nigersaurus. We used high resolution computed tomography, stereolithography, and standard molding and casting techniques to reassemble the extremely fragile skull. Computed tomography also allowed us to render the first endocast for a sauropod preserving portions of the olfactory bulbs, cerebrum and inner ear, the latter permitting us to establish habitual head posture. To elucidate evidence of tooth wear and tooth replacement rate, we used photographic-casting techniques and crown thin sections, respectively. To reconstruct its 9-meter postcranial skeleton, we combined and size-adjusted multiple partial skeletons. Finally, we used maximum parsimony algorithms on character data to obtain the best estimate of phylogenetic relationships among diplodocoid sauropods. Nigersaurus taqueti shows extreme adaptations for a dinosaurian herbivore including a skull of extremely light construction, tooth batteries located at the distal end of the jaws, tooth replacement as fast as one per month, an expanded muzzle that faces directly toward the ground, and hollow presacral vertebral centra with more air sac space than bone by volume. A cranial endocast provides the first reasonably complete view of a sauropod brain including its small olfactory bulbs and cerebrum. Skeletal and dental evidence suggests that Nigersaurus was a ground-level herbivore that gathered and sliced relatively soft vegetation, the culmination of a low-browsing feeding strategy first established among diplodocoids during the Jurassic

A New Basal Sauropod Dinosaur from the Middle Jurassic of Niger and the Early Evolution of Sauropoda

The early evolution of sauropod dinosaurs is poorly understood because of a highly incomplete fossil record. New discoveries of Early and Middle Jurassic sauropods have a great potential to lead to a better understanding of early sauropod evolution and to reevaluate the patterns of sauropod diversification.A new sauropod from the Middle Jurassic of Niger, Spinophorosaurus nigerensis n. gen. et sp., is the most complete basal sauropod currently known. The taxon shares many anatomical characters with Middle Jurassic East Asian sauropods, while it is strongly dissimilar to Lower and Middle Jurassic South American and Indian forms. A possible explanation for this pattern is a separation of Laurasian and South Gondwanan Middle Jurassic sauropod faunas by geographic barriers. Integration of phylogenetic analyses and paleogeographic data reveals congruence between early sauropod evolution and hypotheses about Jurassic paleoclimate and phytogeography.Spinophorosaurus demonstrates that many putatively derived characters of Middle Jurassic East Asian sauropods are plesiomorphic for eusauropods, while South Gondwanan eusauropods may represent a specialized line. The anatomy of Spinophorosaurus indicates that key innovations in Jurassic sauropod evolution might have taken place in North Africa, an area close to the equator with summer-wet climate at that time. Jurassic climatic zones and phytogeography possibly controlled early sauropod diversification

Evolution and Homology of the Astragalus in Early Amniotes: New Fossils, New Perspectives

ABSTRACT The reorganization of the ankle in basal amniotes has long been considered a key innovation allowing the evolution of more terrestrial and cursorial behavior. Understanding how this key innovation arose is a complex problem that largely concerns the homologizing of the amniote astragalus with the various ossifications in the anamniote tarsus. Over the last century, several hypotheses have been advanced homologizing the amniote astragalus with the many ossifications in the ankle of amphibian-grade tetrapods. There is an emerging consensus that the amniote astragalus is a complex structure emerging via the co-ossification of several originally separate elements, but the identities of these elements remain unclear. Here we present new fossil evidence bearing on this contentious question. A poorly ossified, juvenile astragalus of the large captorhinid Moradisaurus grandis shows clear evidence of four ossification centers, rather than of three centers or one center as posited in previous models of astragalus homology. Comparative material of the captorhinid Captorhinikos chozaensis is also interpretable as demonstrating four ossification centers. A new, four-center model for the homology of the amniote astragalus is advanced, and is discussed in the context of the phylogeny of the Captorhinidae in an attempt to identify the developmental transitions responsible for the observed pattern of ossification within this clade. Lastly, the broader implications for amniote phylogeny are discussed, concluding that the neomorphic pattern of astragalus ossification seen in all extant reptiles (including turtles) arose within the clade Diapsida. J. Morphol. 267:415-425, 2006

The vertebrate fauna of the upper Permian of Niger—IX. The appendicular skeleton of <i>Bunostegos akokanensis</i> (Parareptilia: Pareiasauria)

<div><p>ABSTRACT</p><p>Pareiasaurs were a group of herbivorous reptiles that lived during the middle to late Permian (˜265–252 Ma) in what is modern-day Europe, Asia, South America, and Africa. Field work in the Moradi Formation of northern Niger has produced multiple elements of the appendicular skeleton of the pareiasaur <i>Bunostegos akokanensis</i>. The considerable size disparity and morphological variation among the elements suggest that they represent ontogenetic stages ranging from relatively juvenile to adult. Here we present the first description of the scapulocoracoid, humerus, radius, ulna, pelvis, and femur of <i>Bunostegos</i> as well as some of the first ontogenetic data for postcranial osteology in pareiasaurs. As with the skull, numerous postcranial autapomorphies characterize <i>Bunostegos</i>, including laterally originating acromion process of the scapula; radius and ulna with continuous articular surface on humerus; paired crests on the olecranon process; ulna longer than humerus; pinched posterior margin of the acetabular rim; robust pelvic symphysis extending the length of the puboischiatic plate; lack of a distinct postaxial flange of the femur; and an elaborated femoral lateral condyle wrapping over the medial condyle. We incorporated data from the appendicular skeleton of <i>Bunostegos</i> into a revised phylogenetic analysis of pareiasaur relationships. The results of this analysis corroborate previous cranial analyses that place <i>Bunostegos</i> between Guadalupian taxa and the Lopingian velosaur subclade. Interestingly, several aspects of its postcranial anatomy suggest that <i>Bunostegos</i> possessed relatively upright forelimb posture, which would be unique among pareiasaurs and possibly Permian amniotes as a whole.</p><p>SUPPLEMENTAL DATA—Supplemental materials are available for this article for free at www.tandfonline.com/UJVP</p></div

Structural Extremes in a Cretaceous Dinosaur

Fossils of the Early Cretaceous dinosaur, Nigersaurus taqueti, document for the first time the cranial anatomy of a rebbachisaurid sauropod. Its extreme adaptations for herbivory at ground-level challenge current hypotheses regarding feeding function and feeding strategy among diplodocoids, the larger clade of sauropods that includes Nigersaurus. We used high resolution computed tomography, stereolithography, and standard molding and casting techniques to reassemble the extremely fragile skull. Computed tomography also allowed us to render the first endocast for a sauropod preserving portions of the olfactory bulbs, cerebrum and inner ear, the latter permitting us to establish habitual head posture. To elucidate evidence of tooth wear and tooth replacement rate, we used photographic-casting techniques and crown thin sections, respectively. To reconstruct its 9-meter postcranial skeleton, we combined and size-adjusted multiple partial skeletons. Finally, we used maximum parsimony algorithms on character data to obtain the best estimate of phylogenetic relationships among diplodocoid sauropods. Nigersaurus taqueti shows extreme adaptations for a dinosaurian herbivore including a skull of extremely light construction, tooth batteries located at the distal end of the jaws, tooth replacement as fast as one per month, an expanded muzzle that faces directly toward the ground, and hollow presacral vertebral centra with more air sac space than bone by volume. A cranial endocast provides the first reasonably complete view of a sauropod brain including its small olfactory bulbs and cerebrum. Skeletal and dental evidence suggests that Nigersaurus was a ground-level herbivore that gathered and sliced relatively soft vegetation, the culmination of a low-browsing feeding strategy first established among diplodocoids during the Jurassic.</p