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Beyond the Stromer's Riddle: the impact of lumping and splitting hypotheses on the systematics of the giant predatory dinosaurs from northern Africa
Abstract
The “mid-Cretaceous” record from northern Africa is characterized by a peculiar fauna of theropod dinosaurs, mostly referred to a “triumvirate” of clades: Abelisauroidea, Allosauroidea and Spinosauridae. The majority of this material is based on unassociated bones and has ignited a debate on the validity and inclusiveness of the named species, between “lumping” and “splitting” approaches. Although this debate has mostly focused on the alpha taxonomy, the impact of minimizing the a priori assumptions on the diversity and inclusiveness of the taxa coded in the numerical analyses (“methodological splitting”) has barely been investigated. Here, we use new theropod material from the “Kem Kem beds” (Morocco) to test the “methodological splitting” approach on theropod phylogenetics. Revision of the theropod material from the Bahariya Formation (Egypt) described by Ernst Stromer in 1934 leads us to consider the Moroccan Deltadromeus as a junior synonym of Bahariasaurus. Using a large-scale phylogenetic analysis integrating ontogenetic information, all Kem Kem material results nested in the three lineages of the “triumvirate”. The “noasaurids” are reconstructed as a paraphyletic grade of Abelisauroidea, with Bahariasaurus as the largest representative and related to non-predatory taxa showing several convergences with the ornithomimosaurs. Kryptops palaios hypodigm is confirmed an abelisauridallosauroid chimaera. Our analysis also indicates that Eocarcharia dinops hypodigm is a spinosaurid-allosauroid chimaera, and supports recent suggestions for the exclusion of Carcharodontosaurus iguidensis from the latter genus. The Egyptian carcharodontosaurid specimen described by Ernst Stromer and recently renamed Tameryraptor markgrafi is reconstructed as sister taxon of Carcharodontosaurus saharicus neotype even following the updated coding of its morphology. A couple of fused frontals shows several similarities with the holotype of the enigmatic carcharodontosaurid Sauroniops but is less robustly built despite the comparable size. The “mid-Cretaceous” northern African theropod diversity cannot be resolved following “splitting” or “lumping” aprioristic approaches. Direct overlap of diagnostic elements is the only valid criterion for lumping non-associated material. In the absence of overlap in the sample, hypodigms based on “methodological splitting” should be preferred because they prevent topological artifacts biased by the unrecognised inclusion of chimaeras in the taxon sample.
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The first partial skeleton of a carcharodontosaurid theropod was described from the Egyptian Bahariya Oasis by Ernst Stromer in 1931. Stromer referred the specimen to the species Megalosaurus saharicus, originally described on the basis of isolated teeth from slightly older rocks in Algeria, under the new genus name Carcharodontosaurus saharicus. Unfortunately, almost all of the material from the Bahariya Oasis, including the specimen of Carcharodontosaurus was destroyed during World War II. In 1996, a relatively complete carcharodontosaurid cranium was described from similar aged rocks in Morocco and designated the neotype of the species Carcharodontosaurus saharicus in 2007. However, due to the destruction of the original material, comparisons of the neotype to the Egyptian fossils have so far only been done cursorily. A detailed reexamination of the available information on the Egyptian carcharodontosaurid, including a previously undescribed photograph of the exhibited specimen, reveals that it differs from the Moroccan neotype in numerous characters, such as the development of the emargination of the antorbital fossa on the nasals, the presence of a horn-like rugosity on the nasal, the lack of a dorsoventral expansion of the lacrimal contact on the frontals, and the relative enlargement of the cerebrum. The referability of the Egyptian specimen to the Algerian M. saharicus is found to be questionable, and the neotype designation of the Moroccan material for C. saharicus is accepted here under consideration of ICZN Atricle 75, as it both compares more favorably to M. saharicus and originates from a locality closer to the type locality. A new genus and species, Tameryraptor markgrafi gen. et sp. nov, is proposed for the Egyptian taxon. The theropods of the Bahariya Oasis and the Moroccan Kem Kem Group are thus not as closely related as previously thought, and the proposed faunal similarities between these two strata need further examination.
Saurophaganax maximus is the designation of material attributed to a massive theropod dino-saur recovered from the Kenton 1 Quarry in the Kenton Member of the Upper Jurassic Morrison Formation of Oklahoma. The theropod was originally given the name Saurophagus maximus, but was later revised to Saurophaganax maximus because the former did not conform to the International Commission on Zoological Nomenclature standards. Several autapomorphies were described for S. maximus including the postorbital lacking a postorbital boss, the atlas lacking facets for a proatlas, cervical vertebrae with nearly vertical post-zygapophyses, unique accessory laminae on the holotype neural arch, pneumatized post-pectoral dorsal cen-tra, craniocaudally expanded chevrons, laterally bowed femora, tibiae without an astragalar buttress and more prominent distomedial crest, and less distally divergent fourth metatarsals. However, our re-evaluation shows that some of the elements originally used to distinguish Saurophaganax from Allosaurus are more parsimoni-ously referred to diplodocid sauropods found in the same quarry rather than an allosaurid. Additionally, the holotype neural arch cannot be confidently assigned to a theropod, making Saurophaganax maximus a nomen dubium. The presence of at least one skeletally mature theropod was confirmed through paleohistology of a fourth metatarsal. Despite the similarity of the decisively theropod material to known species of Allosaurus, some elements feature subtle autapomorphies that suggest they pertain to a distinct species, which we de-scribe as Allosaurus anax sp. nov.
Dinosaurs thrived for over 160 million years in Mesozoic ecosystems, displaying diverse ecological and evolutionary adaptations. Their ecology was shaped by large-scale climatic and biogeographic changes, calling for a ‘deep-time’ macroecological investigation. These factors include temperature fluctuations and the break up of Pangaea, influencing species richness, ecological diversity and biogeographic history. Recent improvements in the dinosaur fossil record have enabled large-scale studies of their responses to tectonic, geographic and climatic shifts. Trends in species diversity, body size and reproductive traits can now be analysed using quantitative approaches like phylogenetic comparative methods, machine learning and Bayesian inference. These patterns sometimes align with, but also deviate from, first-order macroecological rules (e.g. species–area relationship, latitudinal biodiversity gradient, Bergmann’s rule). Accurate reconstructions of palaeobiodiversity and niche partitioning require ongoing taxonomic revisions and detailed anatomical descriptions. Interdisciplinary research combining sedimentology, geochemistry and palaeoclimatology helps uncover the environmental conditions driving dinosaur adaptations. Fieldwork in under-sampled regions, particularly at latitudinal extremes, is crucial for understanding the spatial heterogeneity of dinosaur ecosystems across the planet. Open science initiatives and online databases play a key role in advancing this field, enriching our understanding of deep-time ecological processes, and offering new insights into dinosaur macroecology and its broader implications.
The end of the Cretaceous saw the Western Interior Seaway divide North America into two land masses, Laramidia in the west and Appalachia in the east. Laramidian dinosaurs inhabited a narrow strip of land extending from Mexico to Alaska. Within this geographically restricted area, dinosaurs evolved high diversity and endemism, with distinct species in the north and south. Here, we report a new tyrannosaurid from the Late Campanian-aged Cerro del Pueblo Formation of Coahuila, Mexico, which is part of a tribe of tyrannosaurs originating in southern Laramidia. Phylogenetic analysis recovers the new tyrannosaur as part of a clade including Labocania anomala from the La Bocana Roja Formation of Baja California Norte, Bistahieversor sealeyi from the Kirtland Formation of New Mexico, Teratophoneus curriei from the Kaiparowits Formation in Utah, and Dynamoterror dynastes from the Menefee Formation of New Mexico. Distinct frontal morphology and the younger age (~72.5–73 Ma versus >75.8 Ma for L. anomala) support recognition of the new tyrannosaur as a distinct species of Labocania, Labocania aguillonae. The Labocania clade dominated southern Laramidia at a time when the north was dominated by daspletosaurins and albertosaurines. The high endemism seen in tyrannosaurids is remarkable, given that modern apex predators have large geographic ranges and hints that the diversity of carnivorous dinosaurs has been underestimated.
Known since the 19 th Century, the compsognathids are among the smallest predatory dinosaurs, and include the first feathered non-avian species found. Traditionally, compsognathids have been considered small and unspecialized coelurosaurs, closer to birds than large-bodied forms like allosauroids and megalosaurids. Yet, all known compsognathids are based on skeletally-immature specimens, and this challenges the accuracy of their traditional phyletic placement. Despite the role of heterochrony in dinosaur evolution is widely recognized, the impact of ontogenetic-biased miscodings in shaping theropod phylogenetics is mostly underestimated. Herein, I show that the standard framework of theropod macroevolution is biased by a series of coding artifacts which violate semaphoront equality prescribed by phylogenetic systematics. I introduce "Ontogenetic State Partitioning" (OSP), a novel coding protocol which integrates ontogenetic and morphological variation under a total evidence approach, and apply it to a densely sampled data set focusing on Mesozoic theropods. The phylogenetic analysis dismissed "Compsognathidae" from being a natural group: its members are identified as juvenile morphs nested among several non-maniraptoriform tetanuran lineages. Conservatism in the immature body plan and greater disparity among large-sized adults differentiate the predatory communities dominated by non-coelurosaurian species (e.g., the so called "triumvirates") from the maniraptoriform-tyrannosaurid faunas (herein named "tyrannies"). This clade-specific differentiation among the communities is confirmed by an analysis of the predatory guild structures including all growth stages: triumvirates and tyrannies result as particular cases along a continuum of communities regulated mainly by alternative contributions of the small-and medium-sized classes. The oldest tyrannies (early Late Cretaceous in age) cluster among non-tyranny communities, supporting the hypothesis that tyrannosaurid-dominated faunas acquired their peculiar structure only after the extinction of the non-coelurosaurian components. The macroevolutionary trajectory that led the maniraptoriforms to realize the avian-like biology may have precluded them from occupying hypercarnivorous large-bodied niches: this bauplan constraint would have favored the tyrannosauroids in opportunistically assuming the apex predatory roles in Late Cretaceous Asiamerica but not elsewhere. The large-scale structure of the Cenozoic radiation of birds is coherent with the framework introduced herein.
Although originally described almost three decades ago, the holotype of Irritator challengeri from the Lower Cretaceous Romualdo Formation of Brazil still represents the most complete spinosaurid skull known to science. Here, we present a detailed description of the skull of Irritator based on digital reconstructions from medical and micro computed tomography (μCT) data. Segmentation reveals the near-complete palatal complex and braincase, an unusual morphology of the retroarticular process, a large, ventrally inclined surangular shelf and the tooth replacement pattern. The digitally reconstructed skull anatomy indicates a robust dentition, a field of binocular vision in front of the skull with an inclined snout orientation, a relatively weak but fast bite, as well as laterally spreading and rotating lower jaw rami during jaw opening. We modified an existing phylogenetic matrix of Tetanurae to account for new observations on the morphology of Irritator and analysed this using parsimony and Bayesian methods. Results support Spinosauridae as members of Megalosauroidea and recover a monophyletic Carnosauria (Megalosauroidea + Allosauroidea). Parsimony analysis recovers Monolophosaurus nested within Megalosauroidea as sister taxon to spinosaurids, but this is not supported by the Bayesian analysis. Bayesian time-calibration and evolutionary rate analysis indicate that spinosaurid evolution happened fast, despite a long ghost lineage of at least 35 million years. High evolutionary rates over a prolonged time can explain the highly derived skull morphology of spinosaurids. This study provides an in-depth look into the evolution of spinosaurid skull anatomy and refines our understanding of these specialized Mesozoic predators.
A predominantly fish-eating diet was envisioned for the sail-backed theropod dinosaur Spinosaurus aegyptiacus when its elongate jaws with subconical teeth were unearthed a century ago in Egypt. Recent discovery of the high-spined tail of that skeleton, however, led to a bolder conjecture that S. aegyptiacus was the first fully aquatic dinosaur. The 'aquatic hypothesis' posits that S. aegyptiacus was a slow quadruped on land but a capable pursuit predator in coastal waters, powered by an expanded tail. We test these functional claims with skeletal and flesh models of S. aegyptiacus. We assembled a CT-based skeletal reconstruction based on the fossils, to which we added internal air and muscle to create a posable flesh model. That model shows that on land S. aegyptiacus was bipedal and in deep water was an unstable, slow-surface swimmer (<1 m/s) too buoyant to dive. Living reptiles with similar spine-supported sails over trunk and tail are used for display rather than aquatic propulsion, and nearly all extant secondary swimmers have reduced limbs and fleshy tail flukes. New fossils also show that Spinosaurus ranged far inland. Two stages are clarified in the evolution of Spinosaurus, which is best understood as a semiaquatic bipedal ambush piscivore that frequented the margins of coastal and inland waterways. Editor's evaluation This article evaluates the hypothesis that Spinosaurus was a specialized aquatic dinosaur, by developing a CT-based skeletal restoration and examining its hydrodynamic properties. In this reappraisal of the "aquatic hypothesis", new results support the alternative "semi-aquatic hypothesis". This article will be of interest to vertebrate paleontologists and functional morphologists, as well as wider academic and non-academic audiences.
Tarbosaurus bataar is a sister taxon of the well-studied theropod dinosaur Tyrannosaurus rex, and numerous fossils of
this tyrannosaurid have been discovered in the Upper Cretaceous Nemegt Formation of Mongolia. Although specimens
of different sizes of Tarbosaurus bataar have been discovered since its initial description, few rigorous studies on its
growth changes have been done. Here we examine growth changes in the frontal bones of seven Tarbosaurus bataar
specimens using bivariate analyses and the Björk superimposition method to demonstrate trends in their ontogenetic
allometry. The width and depth of the frontal undergoes positive allometry during growth, whereas the length shows a
trend of negative allometry. The details of growth changes in Tarbosaurus bataar frontals are largely similar to those
of Tyrannosaurus rex. Furthermore, generic allometric trends of tyrannosaurid frontals, including those of Tarbosaurus
bataar, are shared with other large-bodied theropod clades and may represent a consequence of strengthening parts of
the braincase as an anchor for the jaw musculature.
Numerous non-avian theropod dinosaur fossils have been reported from the Upper Cretaceous (Cenomanian) Bahariya Formation, Bahariya Oasis, Western Desert of Egypt, but unambiguous materials of Abelisauridae have yet to be documented. Here we report Mansoura University Vertebrate Paleontology Center (MUVP) specimen 477, an isolated, well-preserved tenth cervical vertebra of a medium-sized abelisaurid from the Bahariya Formation. The new vertebra shows affinities with those of other Upper Cretaceous abelisaurids from Madagascar and South America, such as Majungasaurus crenatissimus , Carnotaurus sastrei , Viavenator exxoni and a generically indeterminate Patagonian specimen (Museo Padre Molina specimen 99). Phylogenetic analysis recovers the Bahariya form within Abelisauridae, either in a polytomy of all included abelisaurids (strict consensus tree) or as an early branching member of the otherwise South American clade Brachyrostra (50% majority rule consensus tree). MUVP 477, therefore, represents the first confirmed abelisaurid fossil from the Bahariya Formation and the oldest definitive record of the clade from Egypt and northeastern Africa more generally. The new vertebra demonstrates the wide geographical distribution of Abelisauridae across North Africa during the middle Cretaceous and augments the already extraordinarily diverse large-bodied theropod assemblage of the Bahariya Formation, a record that also includes representatives of Spinosauridae, Carcharodontosauridae and Bahariasauridae.
Secondary aquatic adaptations evolved independently more than 30 times from terrestrial vertebrate ancestors1,2. For decades, non-avian dinosaurs were believed to be an exception to this pattern. Only a few species have been hypothesized to be partly or predominantly aquatic3–11. However, these hypotheses remain controversial12,13, largely owing to the difficulty of identifying unambiguous anatomical adaptations for aquatic habits in extinct animals. Here we demonstrate that the relationship between bone density and aquatic ecologies across extant amniotes provides a reliable inference of aquatic habits in extinct species. We use this approach to evaluate the distribution of aquatic adaptations among non-avian dinosaurs. We find strong support for aquatic habits in spinosaurids, associated with a marked increase in bone density, which precedes the evolution of more conspicuous anatomical modifications, a pattern also observed in other aquatic reptiles and mammals14–16. Spinosaurids are revealed to be aquatic specialists with surprising ecological disparity, including subaqueous foraging behaviour in Spinosaurus and Baryonyx, and non-diving habits in Suchomimus. Adaptation to aquatic environments appeared in spinosaurids during the Early Cretaceous, following their divergence from other tetanuran theropods during the Early Jurassic17. In extinct species including non-avian dinosaurs, bone density is shown to be a reliable indicator of aquatic behavioural adaptations, which emerged in spinosaurids during the Early Cretaceous.
Spinosaurids are among the most distinctive and yet poorly-known of large-bodied theropod dinosaurs, a situation exacerbated by their mostly fragmentary fossil record and competing views regarding their palaeobiology. Here, we report two new Early Cretaceous spinosaurid specimens from the Wessex Formation (Barremian) of the Isle of Wight. Large-scale phylogenetic analyses using parsimony and Bayesian techniques recover the pair in a new clade within Baryonychinae that also includes the hypodigm of the African spinosaurid Suchomimus. Both specimens represent distinct and novel taxa, herein named Ceratosuchops inferodios gen. et sp. nov. and Riparovenator milnerae gen. et sp. nov. A palaeogeographic reconstruction suggests a European origin for Spinosauridae, with at least two dispersal events into Africa. These new finds provide welcome information on poorly sampled areas of spinosaurid anatomy, suggest that sympatry was present and potentially common in baryonychines and spinosaurids as a whole, and contribute to updated palaeobiogeographic reconstructions for the clade.
In recent decades, intensive research on non-avian dinosaurs has strongly suggested that these animals were restricted to terrestrial environments¹. Historical proposals that some groups, such as sauropods and hadrosaurs, lived in aquatic environments2,3 were abandoned decades ago4–6. It has recently been argued that at least some of the spinosaurids—an unusual group of large-bodied theropods of the Cretaceous era—were semi-aquatic7,8, but this idea has been challenged on anatomical, biomechanical and taphonomic grounds, and remains controversial9–11. Here we present unambiguous evidence for an aquatic propulsive structure in a dinosaur, the giant theropod Spinosaurus aegyptiacus7,12. This dinosaur has a tail with an unexpected and unique shape that consists of extremely tall neural spines and elongate chevrons, which forms a large, flexible fin-like organ capable of extensive lateral excursion. Using a robotic flapping apparatus to measure undulatory forces in physical models of different tail shapes, we show that the tail shape of Spinosaurus produces greater thrust and efficiency in water than the tail shapes of terrestrial dinosaurs and that these measures of performance are more comparable to those of extant aquatic vertebrates that use vertically expanded tails to generate forward propulsion while swimming. These results are consistent with the suite of adaptations for an aquatic lifestyle and piscivorous diet that have previously been documented for Spinosaurus7,13,14. Although developed to a lesser degree, aquatic adaptations are also found in other members of the spinosaurid clade15,16, which had a near-global distribution and a stratigraphic range of more than 50 million years¹⁴, pointing to a substantial invasion of aquatic environments by dinosaurs.
The geological and paleoenvironmental setting and the vertebrate taxonomy of the fossiliferous, Cenomanian-age deltaic sediments in eastern Morocco, generally referred to as the “Kem Kem beds”, are reviewed. These strata are recognized here as the Kem Kem Group, which is composed of the lower Gara Sbaa and upper Douira formations. Both formations have yielded a similar fossil vertebrate assemblage of predominantly isolated elements pertaining to cartilaginous and bony fishes, turtles, crocodyliforms, pterosaurs, and dinosaurs, as well as invertebrate, plant, and trace fossils. These fossils, now in collections around the world, are reviewed and tabulated. The Kem Kem vertebrate fauna is biased toward largebodied carnivores including at least four large-bodied non-avian theropods (an abelisaurid, Spinosaurus, Carcharodontosaurus, and Deltadromeus), several large-bodied pterosaurs, and several large crocodyliforms.
No comparable modern terrestrial ecosystem exists with similar bias toward large-bodied carnivores. The Kem Kem vertebrate assemblage, currently the best documented association just prior to the onset of the Cenomanian-Turonian marine transgression, captures the taxonomic diversity of a widespread northern African fauna better than any other contemporary assemblage from elsewhere in Africa.
Keywords
Africa, Cretaceous, dinosaur, Gara Sbaa Formation, Douira Formation, paleoenvironment, vertebrate
Allosaurus is one of the best known theropod dinosaurs from the Jurassic and a crucial taxon in phylogenetic analyses. On the basis of an in-depth, firsthand study of the bulk of Allosaurus specimens housed in North American institutions, we describe here a new theropod dinosaur from the Upper Jurassic Morrison Formation of Western North America, Allosaurus jimmadseni sp. nov., based upon a remarkably complete articulated skeleton and skull and a second specimen with an articulated skull and associated skeleton. The present study also assigns several other specimens to this new species, Allosaurus jimmadseni , which is characterized by a number of autapomorphies present on the dermal skull roof and additional characters present in the postcrania. In particular, whereas the ventral margin of the jugal of Allosaurus fragilis has pronounced sigmoidal convexity, the ventral margin is virtually straight in Allosaurus jimmadseni . The paired nasals of Allosaurus jimmadseni possess bilateral, blade-like crests along the lateral margin, forming a pronounced nasolacrimal crest that is absent in Allosaurus fragilis .
Abelisaurid theropods first appear in the fossil record in the early Jurassic and survived at least until the end of the Mesozoic. They were known to have dominated South America, India and Madagascar but were not so abundant in North America or Asia. Much less is known about their presence in Africa, although there has been several recent discoveries of abelisaurid material in Morocco. Here we add a partially preserved ilium to a growing body of evidence that suggests abelisaurs might also have dominated Africa.
Here we describe an extremely large and relatively complete (roughly 65%) skeleton of Tyrannosaurus rex (RSM P2523.8). Multiple measurements (including those of the skull, hip, and limbs) show that RSM P2523.8 was a robust individual with an estimated body mass exceeding all other known T. rex specimens and representatives of all other gigantic terrestrial theropods. Histological sampling of the fibula confirms that RSM P2523.8 is skeletally mature. The prevalence of incompletely coossified elements contradicts previous assertions that such unfused elements can be taken as indicators of somatic immaturity. As an extreme example of both ontogenetic maturity and osteological robustness, RSM P2523.8 offers support for prior hypotheses that a sampling bias occurs throughout the Dinosauria, making it likely that most taxa grew to significantly greater size than current known specimens indicate. Anat Rec, 303:656–672, 2020. © 2019 Wiley Periodicals, Inc.
The Spinosauridae is a specialised clade of theropod dinosaurs known from the Berriasian to the Cenomanian of Africa, South America, Europe and Asia. Spinosaurs were unusual among non-avian dinosaurs in exploiting a piscivorous niche within riverine and estuarine habitats, and they include the largest known theropod. Although fossils of giant spinosaurs are increasingly well-represented in the fossil record, little juvenile material has been described. Here, we describe new examples of juvenile spinosaurines from the middle Cretaceous (Cenomanian) Kem Kem beds of Morocco. The fossils include material from a range of sizes and are relatively common within the Kem Kem deposits, suggesting that juveniles exploited the same semiaquatic niche as the adults throughout ontogeny. This implies that the Cenomanian delta habitats supported an age-inclusive population of spinosaurs that was neither geographically or environmentally separated, though some ecological separation between juveniles and adults is likely based on the large variation in size. Bones or teeth of very small (<2 m) spinosaurs have not been found, however. This could represent a taphonomic bias, or potentially an ecological signal that the earliest ontogenetic stages inhabited distinct environments. Skeletal remains include individuals referable to Sigilmassasaurus brevicollis and a second spinosaurine taxon. Consistent with this, we also identify two distinct cranial morphs of Spinosauridae present within the Kem Kem, supporting previous recognition of distinct taxa in the assemblage.
We describe a nearly complete pedal ungual phalanx, discovered in the Kem Kem Beds (Cenomanian) of Tafilalt region, south-eastern Morocco. The bone is symmetric, pointed, low, elongate, and almost flat ventrally in lateral aspect. This peculiar morphology allows to refer the specimen to the smallest known individual of the genus Spinosaurus. The bone belongs to an early juvenile individual and it is proportionally identical to the ungual of the third digit of a large partial skeleton recently found, suggesting an isometric growth for this part of the pes and the retention of peculiar locomotor adaptations—such as traversing soft substrates or paddling—during the entire lifespan.
Birds are one of the most successful groups of vertebrates. The origin of birds from their reptilian ancestors is traditionally rooted near the Jurassic "Urvogel" Archaeopteryx, an approach that has contributed in defining the dichotomy between the "reptilian" (pre-Archaeopteryx) and "avian" (post-Archaeopteryx) phases of what is instead a single evolutionary continuum. A great and still ever increasing amount of evidence from the fossil record has filled the gaps between extinct dinosaurs, Mesozoic birds and modern avians, and led to the revision of the misleading dichotomy between pre-and post-Archaeopteryx stages in the evolution of bird biology. Herein, the progressive assembly of the modern avian body plan from the archosaurian ancestral condition is reviewed using a combination of phylogenetic methods. The stem lineage leading to modern birds is described using 38 internodes, which identity a series of progressively less inclusive ancestors of modern birds and their Mesozoic sister taxa. The 160-million-year long assembly of the avian bauplan is subdivided into three main stages on the basis of analyses of skeletal modularity, cladogenetic event timing, divergence rate inference and morphospace occupation. During the first phase ("Huxleyian stage": Early Triassic to Middle Jurassic), the earliest ancestors of birds acquired postcranial pneumatisation, an obligate bipedal and digitigrade posture, the tridactyl hand and feather-like integument. The second phase ("Ostromian stage": second half of Jurassic) is characterised by a higher evolutionary rate, the loss of hypercarnivory, the enlargement of the braincase, the dramatic reduction of the caudofemoral module, and the development of true pennaceous feathers. The transition to powered flight was achieved only in the third phase ("Marshian stage": Cretaceous), with the re-organisation of both forelimb and tail as flight-adapted organs and the full acquisition of the modern bauplan. Restricting the investigation of the avian evolution to some Jurassic paravians or to the lineages crown-ward from Archaeopteryx ignores the evolutionary causes of over 60% of the features that define the avian body. The majority of the key elements forming the third phase are exaptations of novelties that took place under the different ecological and functional regimes of the Huxleyian and Ostromian stages, and cannot be properly interpreted without making reference to their original historical context.
RIASSUNTO - [La costruzione del piano corporeo aviano: un processo lungo 160 milioni di anni] - Gli uccelli sono uno dei gruppi di vertebrati di maggiore successo. L'origine degli uccelli dai loro antenati rettiliani è tradizionalmente ancorata intorno allo "Urvogel" giurassico Archaeopteryx; questo approccio ha consolidato la distinzione tra una fase "rettiliana" (precedente Archaeopteryx) ed una "aviana" (successiva ad Archaeopteryx) in quello che è invece un singolo continuum evolutivo. Una crescente quantità di evidenze dal registro fossilifero ha colmato le lacune esistenti tra i dinosauri non-avialiani, gli uccelli mesozoici e quelli moderni, e ha portato alla revisione della fuorviante dicotomia tra fasi pre-e post-Archaeopteryx nell'evoluzione della biologia aviana. Il progressivo assemblaggio del moderno piano corporeo aviano è qui discusso usando una combinazione di metodi fi logenetici. La linea filetica che conduce agli uccelli moderni è descritta da 38 internodi, che identifi cano una serie progressiva di antenati condivisi tra gli uccelli attuali e i loro sister group mesozoici. I 160 milioni di anni di durata della costruzione del bauplan aviano sono suddivisi in tre fasi principali sulla base di analisi della modularità scheletrica, della cronologia degli eventi cladogenetici, dei tassi di divergenza, e delle regioni del morfospazio occupate. Durante la prima fase (detta "huxleyiana": dal Triassico Inferiore al Giurassico Medio), gli antenati degli uccelli svilupparono la pneumatizzazione postcraniale, una postura bipede obbligata e digitigrada, la mano tridattila e un tegumento simile al piumaggio. La seconda fase ("ostromiana": seconda metà del Giurassico) è caratterizzata da un più elevato tasso di evoluzione divergente, la perdita dell'ecologia ipercarnivora, l'espansione dell'endocranio, la drammatica riduzione del modulo caudofemorale, e lo sviluppo di piumaggio pennaceo. La transizione al volo battuto fu sviluppata solo nella terza fase ("marshiana": Cretacico), con la riorganizzazione dell'arto anteriore e della coda in organi adatti al volo, e la completa acquisizione del bauplan moderno. Restringere l'indagine sull'evoluzione aviana ad alcuni paraviani giurassici o alle linee successive ad Archaeopteryx significa ignorare la causa di oltre il 60% delle caratteristiche che definiscono il modello corporeo degli uccelli. La maggioranza degli elementi chiave che defi niscono la moderna fase dell'evoluzione aviana sono exaptation di novità occorse sotto diff erenti regimi ecologico-funzionali nelle fasi huxleyiana e ostromiana, e non possono essere propriamente interpretati senza fare riferimento al contesto storico della loro origine.
Understanding ontogenetic patterns is important in vertebrate paleontology because the assessed skeletal maturity of an individual often has implications for paleobiogeography, species synonymy, paleobiology, and body size evolution of major clades. Further, for many groups the only means of confidently determining ontogenetic status of an organism is through the destructive process of histological sampling. Although the ontogenetic patterns of Late Jurassic and Cretaceous dinosaurs are better understood, knowledge of the ontogeny of the earliest dinosaurs is relatively poor because most species-level growth series known from these groups are small (usually, maximum of n ~ 5) and incomplete. To investigate the morphological changes that occur during ontogeny in early dinosaurs, I used ontogenetic sequence analysis (OSA) to reconstruct developmental sequences of morphological changes in the postcranial ontogeny of the early theropods Coelophysis bauri and Megapnosaurus rhodesiensis, both of which are known from large sample sizes (n = 174 and 182, respectively). I found a large amount of sequence polymorphism (i.e. intraspecific variation in developmental patterns) in both taxa, and especially in C. bauri, which possesses this variation in every element analyzed. Megapnosaurus rhodesiensis is similar, but it possesses no variation in the sequence of development of ontogenetic characters in the tibia and tarsus. Despite the large amount of variation in development, many characters occur consistently earlier or later in ontogeny and could therefore be important morphological features for assessing the relative maturity of other early theropods. Additionally, there is a phylogenetic signal to the order in which homologous characters appear in ontogeny, with homologous characters appearing earlier or later in developmental sequences of early theropods and the close relatives of dinosaurs, silesaurids. Many of these morphological features are important characters for the reconstruction of archosaurian phylogeny (e.g. trochanteric shelf). Because these features vary in presence or appearance with ontogeny, these characters should be used with caution when undertaking phylogenetic analyses in these groups, since a specimen may possess certain character states owing to ontogenetic stage, not evolutionary relationships.
Bayesian phylogenetic methods integrating simultaneously morphological and strati-graphic information have been applied increasingly among paleontologists. Most of these studies have used Bayesian methods as an alternative to the widely-used parsimony analysis, to infer macroevolutionary patterns and relationships among species-level or higher taxa. Among recently introduced Bayesian methodologies, the Fossilized Birth-Death (FBD) model allows incorporation of hypotheses on ancestor-descendant relationships in phylogenetic analyses including fossil taxa. Here, the FBD model is used to infer the relationships among an ingroup formed exclusively by fossil individuals, i.e., dipnoan tooth plates from four localities in the Ain el Guettar Formation of Tunisia. Previous analyses of this sample compared the results of phylogenetic analysis using parsimony with stratigraphic methods, inferred a high diversity (five or more genera) in the Ain el Guettar Formation, and interpreted it as an artifact inflated by depositional factors. In the analysis performed here, the uncertainty on the chronostratigraphic relationships among the specimens was included among the prior settings. The results of the analysis confirm the referral of most of the specimens to the taxa Asiatoceratodus, Equinoxiodus, Lavocatodus and Neoceratodus, but reject those to Ceratodus and Ferganoceratodus. The resulting phylogeny constrained the evolution of the Tunisian sample exclusively in the Early Cretaceous, contrasting with the previous scenario inferred by the stratigraphically-calibrated topology resulting from parsimony analysis. The phylogenetic framework also suggests that (1) the sampled localities are laterally equivalent, (2) but three localities are restricted to the youngest part of the section; both results are in agreement with previous stratigraphic analyses of these localities. The FBD model of specimen-level units provides a novel tool for phylogenetic inference among fossils but also for independent tests of stratigraphic scenarios.
We describe the osteology of the new small theropod dinosaur Masiakasaurus knopfleri, from the Late Cretaceous Maevarano Formation of northwestern Madagascar. Approximately 40% of the skeleton is known, including parts of the jaws, axial column, forelimb, pelvic girdle, and hind limb. The jaws of Masiakasaurus are remarkably derived, bearing a heterodont, procumbent dentition that is unknown elsewhere among dinosaurs. The vertebrae are similar to those of abelisauroids in the reduction of the neural spine, lack of pleurocoelous fossae on the centrum, and extensively pneumatized neural arch. The limb skeleton is relatively gracile and bears numerous abelisauroid synapomorphies, including a rounded humeral head, peg-and-socket iliac-pubic articulation, prominent femoral medial epicondyle, expanded tibial cnemial crest, and double-grooved pedal unguals. The femora and tibiae show evidence of dimorphism. More specific features shared between Masiakasaurus, the Argentine Noasaurus, and the Indian Laevisuchus suggest that these taxa form a clade (Noasauridae) within Abelisauroidea. This is supported by a cladistic phylogenetic analysis of 158 characters and 23 theropod taxa. Additionally, Ceratosauria is rendered paraphyletic in favor of a sister-taxon relationship between Neoceratosauria and Tetanurae that is exclusive of Coelophysoidea. The unique dental and jaw specializations of Masiakasaurus suggest deviation from the typical theropod diet. Finally, the distribution of noasaurids further supports a shared biogeographic history between South America, Madagascar, and India into the Late Cretaceous.
Background
Late Cretaceous terrestrial strata of the Neuquén Basin, northern Patagonia, Argentina have yielded a rich fauna of dinosaurs and other vertebrates. The diversity of saurischian dinosaurs is particularly high, especially in the late Cenomanian-early Turonian Huincul Formation, which has yielded specimens of rebacchisaurid and titanosaurian sauropods, and abelisaurid and carcharodontosaurid theropods. Continued sampling is adding to the known vertebrate diversity of this unit.
Methodology/ Principal Findings
A new, partially articulated mid-sized theropod was found in rocks from the Huincul Formation. It exhibits a unique combination of traits that distinguish it from other known theropods justifying erection of a new taxon, Gualicho shinyae gen. et sp. nov. Gualicho possesses a didactyl manus with the third digit reduced to a metacarpal splint reminiscent of tyrannosaurids, but both phylogenetic and multivariate analyses indicate that didactyly is convergent in these groups. Derived characters of the scapula, femur, and fibula supports the new theropod as the sister taxon of the nearly coeval African theropod Deltadromeus and as a neovenatorid carcharodontosaurian. A number of these features are independently present in ceratosaurs, and Gualicho exhibits an unusual mosaic of ceratosaurian and tetanuran synapomorphies distributed throughout the skeleton.
Conclusions/ Significance
Gualicho shinyae gen. et sp. nov. increases the known theropod diversity of the Huincul Formation and also represents the first likely neovenatorid from this unit. It is the most basal tetatanuran to exhibit common patterns of digit III reduction that evolved independently in a number of other tetanuran lineages. A close relationship with Deltadromaeus from the Kem Kem beds of Niger adds to the already considerable biogeographic similarity between the Huincul Formation and coeval rock units in North Africa.
Many recent studies of theropod relationships have been focused on the phylogeny of coelurosaurs and the question of the origin of birds, but the interrelationships and evolution of basal theropods are still poorly understood. Thus, this paper presents a phylogenetic analysis of all theropods, but focuses on the basal members of this clade. The result supports the inclusion of Eoraptor and herrerasaurids in the Theropoda, but differs from other recent studies in two main aspects: (1) The taxa usually grouped as ceratosaurs form two monophyletic clades that represent successively closer outgroups to tetanurans. The more basal of these clades, the Coelophysoidea, comprise the majority of Late Triassic and Early Jurassic theropods. The other clade of basal theropods that are usually included in the Ceratosauria comprises Ceratosaurus, Elaphrosaurus, and abelisaurids. (2) Two monophyletic groups of basal tetanurans are recognized: the Spinosauroidea and the Allosauroidea. In contrast to other recent phylogenetic hypotheses, both clades are united in a monophyletic Carnosauria. The branching pattern of the present cladogram is in general accordance with the stratigraphic occurrence of theropod taxa. Despite the differences in recent analyses, there is a significant level of consensus in theropod phylogeny. At least four different radiations of non-avian theropods can be recognized. These radiations show different patterns in Laurasia and Gondwana, and there are increasing differences between the theropod faunas of the two hemispheres from the Triassic to the Cretaceous.
Six quadrate bones, of which two almost certainly come from the Kem Kem beds (Cenomanian, Upper Cretaceous) of south-eastern Morocco, are determined to be from juvenile and adult individuals of Spinosaurinae based on phylogenetic, geometric morphometric, and phylogenetic morphometric analyses. Their morphology indicates two morphotypes evidencing the presence of two spinosaurine taxa ascribed to Spinosaurus aegyptiacus and? Sigilmassasaurus brevicollis in the Cenomanian of North Africa, casting doubt on the accuracy of some recent skeletal reconstructions which may be based on elements from several distinct species. Morphofunctional analysis of the mandibular articulation of the quadrate has shown that the jaw mechanics was peculiar in Spinosauridae. In mature spinosaurids, the posterior parts of the two mandibular rami displaced laterally when the jaw was depressed due to a lateromedially oriented intercondylar sulcus of the quadrate. Such lateral movement of the mandibular ramus was possible due to a movable mandibular symphysis in spinosaurids, allowing the pharynx to be widened. Similar jaw mechanics also occur in some pterosaurs and living pelecanids which are both adapted to capture and swallow large prey items. Spinosauridae, which were engaged, at least partially, in a piscivorous lifestyle, were able to consume large fish and may have occasionally fed on other prey such as pterosaurs and juvenile dinosaurs.
Although the Late Jurassic of Portugal has provided abundant dinosaur fossils, material from the Early Cretaceous is scarce. This paper reports new cranial and postcranial material of the theropod dinosaur Baryonyx walkeri found in the Barremian (Papo Seco Formation) of Portugal. This specimen, found at Praia das Aguncheiras, Cabo Espichel, consists of a partial dentary, isolated teeth, pedal ungual, two calcanea, presacral and caudal vertebrae, fragmentary pubis, scapula, and rib fragments. It represents the most complete spinosaurid yet discovered in the Iberian Peninsula and the most complete dinosaur from the Early Cretaceous of Portugal. This specimen is confidently identified as a member of Baryonychinae due to the presence of conical teeth with flutes and denticles in a dentary rosette. The specimen ML1190 shares the following characteristics with Baryonyx walkeri: enamel surface with small (nearly vertical) wrinkles, variable denticle size along the carinae, 6-7 denticles per mm, wrinkles forming a 45 degree angle near the carinae, and tooth root longer than crown. In addition, dubious taxa based on teeth morphology such as Suchosaurus cultridens (Owen, 1840-1845), and Suchosaurus girardi (Sauvage 1897-98; Antunes & Mateus 2003) are discussed, based on comparisons with well-known material such as Baryonyx walkeri Charig & Milner, 1986. Suchosaurus cultridens and S. girardi are considered as nomina dubia due to the lack of diagnostic apomorphies, but both specimens are referred to Baryonychinae incertae sedis.
Sigilmassasaurus brevicollis is an enigmatic theropod dinosaur from the early Late Cretaceous (Cenomanian) of Morocco, originally based on a few isolated cervical vertebrae. Ever since its original description, both its taxonomic validity and systematic affinities were contentious. Originally considered to represent its own family, Sigilmassasauridae, the genus has variously been suggested to represent a carcharodontosaurid, an ornithischian, and, more recently, a spinosaurid. Here we describe new remains referrable to this taxon and re-evaluate its taxonomic status and systematic affinities. Based on the new remains, a re-evaluation of the original materials, and comparisons with other spinosaurids, the holotype of Sigilmassasaurus brevicollis is identified as an anterior dorsal, rather than a cervical vertebra, and differences between elements referred to this taxon can be explained by different positions of the elements in question within the vertebral column. Many characters used previously to diagnose the genus and species are found to be more widespread among basal tetanurans, and specifically spinosaurids. However, the taxon shows several autapomorphies that support its validity, including the presence of a strongly rugose, ventrally offset triangular platform that is confluent with a ventral keel anteriorly in the mid-cervical vertebral centra and a strongly reduced lateral neural arch lamination, with no or an incomplete distinction between anterior and posterior centrodiapophyseal laminae in the posterior cervical and anterior dorsal vertebrae. We argue furthermore that Spinosaurus maroccanus, also described on the basis of isolated cervical vertebrae from the same stratigraphic unit and in the same paper as Sigilmassasaurus brevicollis, is a subjective synonym of the latter. Both a detailed comparison of this taxon with other theropods and a formal phylogenetic analysis support spinosaurid affintities for Sigilmassasaurus. However, we reject the recently
Recent fieldwork in Upper Cretaceous terrestrial deposits in northwestern Madagascar has yielded a remarkable diversity of vertebrates, including several specimens of the abelisaurid theropod Majungasaurus crenatissimus (Depéret, 1896) Lavocat, 1955. Featured among the discoveries is an exquisite specimen (UA 8678) that preserves a virtually complete precaudal vertebral column, numerous costal elements, and portions of the skull and appendicular skeleton. This contribution represents the first description highlighting the postcranial axial skeleton of Majungasaurus. Owing to the completeness and quality of preservation, this specimen allows an examination of the serial transformation of features along the length of the axial skeleton, including a detailed analysis of postcranial pneumaticity in a nontetanuran theropod. Notable features of Majungasaurus include pneumatic cervical ribs with caudally bifurcate shafts and extensive pneumaticity of all postatlantal, precaudal vertebrae. Several postcranial features exhibited by Majungasaurus - including a well-developed cervical epipophysis, laterally expanded dorsal parapophysis, and sub-divided infradiapophyseal fossa in middle dorsal series - support previous phylogenetic studies placing it within Abelisauroidea and Abelisauridae. Majungasaurus (and abelisaurids generally) exhibit a robust cervical skeleton that features tightly interlocking cervical ribs, hyperossification of cervical rib shafts, and hypertrophied muscle attachment sites relative to other basal theropods. These features together highlight an axial core constructed to withstand high stresses, likely reflecting feeding adaptations for predation on large-bodied prey.
New specimens of the unusual theropod Spinosaurus cf. S. aegyptiacus from the Late Cretaceous (early Cenomanian) of Morocco reveal new information about the structure of the snout and the very large adult body size attained by the species. The external naris is retracted farther caudally on the snout than in other spinosaurids and is bordered exclusively by the maxilla and nasal. The fused nasals preserve a longitudinal, fluted crest. The size of the snout suggests that Spinosaurus may well have exceeded the maximum adult body size of other large Cretaceous theropods such as Tyrannosaurus and Giganotosaurus. The new material also supports the monophyly of the Spinosaurinae and the separation of Spinosaurus and Irritator.
Noasaurus leali is a small (~2 m) carnivorous theropod and the nominal genus of the clade Noasauridae, one of the two radiations of abelisauroid ceratosaurs predominantly present in the Southern Hemisphere during the Mesozoic. This eponymous theropod from the Maastrichtian Lecho Formation of Salta, Argentina, is known from an incomplete skeleton of which the strongly curved manual ungual is the most peculiar element. We here provide for the first time a comprehensive description of the holotypic specimens of Noasaurus, whose phylogenetic position was explored using three independent datamatrices on theropod relationships. This species is diagnosed by several apomorphies such as a dorsal ridge in the maxillary fossa, a strongly arched quadrate, a cervical neural arch with anterior epipophyseal prongs, and a manual ungual with a subtriangular flexor fossa delimited by a V-shaped ridge. Results of the phylogenetic analyses recovered Noasaurus closely related to Velocisaurus, Masiakasaurus, and Laevisuchus, which together form a Late Cretaceous radiation of small-bodied noasaurids restricted to the Southern Hemisphere. The peculiar morphology of the lateral dentition and manual unguals suggests that Noasaurus was an opportunistic carnivore feeding on small prey items and a possible piscivore gaffing fish with its specialized hand claws.
The Kem Kem Group is a lowermost lithostratigraphic unit from the Upper Cretaceous that extends along the border between Algeria and Morocco, in the northern region of Africa. This geological unit has yielded several tetrapod fossils, including a well-represented assemblage of theropod dinosaurs, after more than eight decades of research. Here, we report new occurrences of spinosaurid theropods from the spinosaurine clade in the Kem Kem Group by providing anatomical descriptions and taxonomic identifications of 11 new specimens derived from the Tafilalt region of Morocco. Among the findings, we describe a cervical vertebra of Sigilmassasaurus, in addition to several cranial, axial, and appendicular elements that can safely be attributed to Spinosaurinae. Moreover, based on a unique combination of characteristics, we also describe an isolated and partial ischium belonging to an indeterminate carcharodontosaurid. We also deliver a detailed redescription of one of the most complete snouts of a spinosaurine known to date. Therefore, the theropod dinosaurs of the Kem Kem Group show considerable diversity, but many questions, especially related to the diversity of spinosaurids and the general abundance of carnivorous dinosaurs in this region, remain unclear until new materials are discovered and complete descriptions are made.
A new graphical user interface (GUI) for the parsimony program TNT is presented that works under the Linux and Mac operating systems, as well as the Cygwin environment (which runs under Windows). The new interface is based on the GIMP Tool Kit, GTK (version 3). Formerly, only Windows versions of TNT had a GUI. The new interface improves upon the existing Windows GUI in several respects. These changes, together with several additions to the program since the publication of version 1.5, warrant a change in minor version, thus moving from version 1.5 to 1.6. Among the most notable improvements are the possibility to access graphical user dialogs by means of simple commands, to easily save trees in SVG format (“Scalable Vector Graphics”) directly from any tree‐diagram being displayed, and to manage analyses in parallel (using multiple processors, by means of the PVM system or “Parallel Virtual Machine”), as well as a generally more stable and consistent behaviour. As the binaries for the new version are compiled as native 64‐bit applications, this removes the limitations for accessing large amounts of memory in the previous GUI Windows interface (which is a 32‐bit application).
The ‘Kem Kem Compound Assemblage’ (KKCA) along the Algerian-Moroccan border is a series of fossiliferous localities, Cenomanian in age, particularly rich in large-bodied theropod dinosaurs. Two species of carcharodontosaurid allosauroids have been identified in these units, Carcharodontosaurus saharicus and Sauroniops pachytholus. Recently, the validity of the second species has been challenged, and all carcharodontosaurid material from the KKCA has been referred uniquely to the former species. Here, we describe a new theropod cranial material from the KKCA which is referred to Carcharodontosauridae, including one partial maxilla which shows a morphology distinct from that of C. saharicus. We review the arguments used to interpret Sauroniops holotype as an immature individual of Carcharodontosaurus and show that they were based on non-homologous comparisons and on the misinterpretation of the original description of S. pachytholus: The latter is confirmed to be a valid taxon, distinct from C. saharicus and with an inferred body size comparable to the largest carcharodontosaurids. The presence of more than one giant carcharodontosaurid species in the Cenomanian of Morocco recalls the carcharodontosaurid diversity from penecontemporary units from Argentina.
Giant carnivorous dinosaurs such as Tyrannosaurus rex and abelisaurids are characterized by highly reduced forelimbs that stand in contrast to their huge dimensions, massive skulls, and obligate bipedalism.¹,² Another group that follows this pattern, yet is still poorly known, is the Carcharodontosauridae: dominant predators that inhabited most continents during the Early Cretaceous3, 4, 5 and reached their largest sizes in Aptian-Cenomanian times.6, 7, 8, 9, 10 Despite many discoveries over the last three decades, aspects of their anatomy, especially with regard to the skull, forearm, and feet, remain poorly known. Here we report a new carcharodontosaurid, Meraxes gigas, gen. et sp. nov., based on a specimen recovered from the Upper Cretaceous Huincul Formation of northern Patagonia, Argentina. Phylogenetic analysis places Meraxes among derived Carcharodontosauridae, in a clade with other massive South American species. Meraxes preserves novel anatomical information for derived carcharodontosaurids, including an almost complete forelimb that provides evidence for convergent allometric trends in forelimb reduction among three lineages of large-bodied, megapredatory non-avian theropods, including a remarkable degree of parallelism between the latest-diverging tyrannosaurids and carcharodontosaurids. This trend, coupled with a likely lower bound on forelimb reduction, hypothesized to be about 0.4 forelimb/femur length, combined to produce this short-armed pattern in theropods. The almost complete cranium of Meraxes permits new estimates of skull length in Giganotosaurus, which is among the longest for theropods. Meraxes also provides further evidence that carchardontosaurids reached peak diversity shortly before their extinction with high rates of trait evolution in facial ornamentation possibly linked to a social signaling role.
Well-sampled dinosaur communities from the Jurassic through the early Late Cretaceous show greater taxonomic diversity among larger (>50 kg) theropod taxa than communities of the Campano-Maastrichtian, particularly to those of eastern/central Asia and Laramidia. The large carnivore guilds in Asiamerican assemblages are monopolized by tyrannosaurids, with adult medium-sized (50–500 kg) predators rare or absent. In contrast, various clades of theropods are found to occupy these body sizes in earlier faunas, including early tyrannosauroids. Assemblages with “missing middle-sized” predators are not found to have correspondingly sparser diversity of potential prey species recorded in these same faunas. The “missing middle-sized” niches in the theropod guilds of Late Cretaceous Laramidia and Asia may have been assimilated by juvenile and subadults of tyrannosaurid species, functionally distinct from their adult ecomorphologies. It is speculated that if tyrannosaurids assimilated the niches previously occupied by mid-sized theropod predators, we would expect the evolution of distinct transitions in morphology and possibly the delay of the achievement of somatic maturity in species of this taxon.
Inferring the body mass of fossil taxa, such as non‐avian dinosaurs, provides a powerful tool for interpreting physiological and ecological properties, as well as the ability to study these traits through deep time and within a macroevolutionary context. As a result, over the past 100 years a number of studies advanced methods for estimating mass in dinosaurs and other extinct taxa. These methods can be categorized into two major approaches: volumetric‐density (VD) and extant‐scaling (ES). The former receives the most attention in non‐avian dinosaurs and advanced appreciably over the last century: from initial physical scale models to three‐dimensional (3D) virtual techniques that utilize scanned data obtained from entire skeletons. The ES approach is most commonly applied to extinct members of crown clades but some equations are proposed and utilized in non‐avian dinosaurs. Because both approaches share a common goal, they are often viewed in opposition to one another. However, current palaeobiological research problems are often approach specific and, therefore, the decision to utilize a VD or ES approach is largely question dependent. In general, biomechanical and physiological studies benefit from the full‐body reconstruction provided through a VD approach, whereas large‐scale evolutionary and ecological studies require the extensive data sets afforded by an ES approach. This study summarizes both approaches to body mass estimation in stem‐group taxa, specifically non‐avian dinosaurs, and provides a comparative quantitative framework to reciprocally illuminate and corroborate VD and ES approaches. The results indicate that mass estimates are largely consistent between approaches: 73% of VD reconstructions occur within the expected 95% prediction intervals of the ES relationship. However, almost three quarters of outliers occur below the lower 95% prediction interval, indicating that VD mass estimates are, on average, lower than would be expected given their stylopodial circumferences. Inconsistencies (high residual and per cent prediction deviation values) are recovered to a varying degree among all major dinosaurian clades along with an overall tendency for larger deviations between approaches among small‐bodied taxa. Nonetheless, our results indicate a strong corroboration between recent iterations of the VD approach based on 3D specimen scans suggesting that our current understanding of size in dinosaurs, and hence its biological correlates, has improved over time. We advance that VD and ES approaches have fundamentally (metrically) different advantages and, hence, the comparative framework used and advocated here combines the accuracy afforded by ES with the precision provided by VD and permits the rapid identification of discrepancies with the potential to open new areas of discussion.
Dans les grès surtout siliceux et un peu poreux du Continental intercalaire (Crétacé inférieur) du Sahara (Laghouat, In Aménas, Timimoun, Adrar) les Kerkoubs sont des concrétions de 1 à 150 mm et plus, indurées, plus ou moins sphériques, donc à croissance plus ou moins isotrope, parfois coalescentes, dues à une cimentation carbonatée. Formés au Crétacé inférieur ou plus "récemment ? Par voie purement physico-chimique, ou avec intervention de microorganismes (bactéries chlorophylliennes ou autres ... etc.) ? A Laghouat, au Rocher des Pigeons, les Kerkoubs, loin d'être uniformément répartis, sont localisés dans certains bancs des grès, souvent en essaims (photo 1) d'une même famille de formes, essaims à limite parfois très tranchée (photo 14). Sur la surface d'un gros Kerkoub, existent quelquefois, sous forme de verrues hémisphériques identiques à ce que donnerait un bourgeonnement (photo 10), des Kerkoubs moyens (4 à 5 mm) et sur la surface de ceux-ci et aussi du gros, des Kerkoubs en tête d'épingle (1 mm). Quelquefois aussi il y a eu deux actes de concrétionnement, emboîtés (photo 13), de petits Kerkoubs sphériques très durs étant eux-mêmes englobés dans un agrégat lui aussi plus ou moins sphérique, mais moins fortement induré. Le vent érodant le grès encaissant, mal cimenté, plus vite que les Kerkoubs, ceux-ci sont mis en relief (photo 5) et finalement se détachent. En place, la partie en relief d'un Kerkoub est plus dure, mieux cimentée que la partie encore engagée dans le grès ; elle est souvent teintée de rouille. Dans la falaise de grès, au fur et à mesure qu'on s'éloigne de la surface actuelle et qu'on observe la roche à l'intérieur, la distinction entre le Kerkoub et l'encaissant devient à l'oeil nu de moins en moins nette et au bout de quelques centimètres ou décimètres, elle disparaît. Tout ceci inclinerait à penser que le concrétionnement est récent, ou même s'il a débuté au Crétacé, qu'il s'est poursuivi encore récemment.
Quelquefois des Kerkoubs sont entourés chacun d'un anneau circulaire ferruginisé (photo 6) qui évoque soit des anneaux de diffusion de Liesegang, soit un dépôt bactérien. Parfois un seul et même banc renferme par places quelques Kerkoubs et ailleurs des tubulures (photos 16 et 17) d'origine évidemment biologique.
L'apport de calcaire, ciment des Kerkoubs, exige bien entendu une certaine circulation d'eau, mais sous climat chaud et aride ou plutôt semi-aride.
Spinosauridae is a specialized group of theropod dinosaurs characterised by a long, narrow skull, robust forelimbs with a hooked thumb claw, and tall neural spines forming a dorsal sail. The ecology of these unusual dinosaurs has been debated since the original discovery of Spinosaurus aegyptiacus in 1912. Morphological similarities to crocodilians, including tooth shape and an elongated rostrum, indicate a piscivorous diet, and in the giant Spinosaurus, a long body and short limbs suggest semi-aquatic habits. However, the hypothesized aquatic habits of Spinosaurus have been called into question, and the distribution of aquatic habits within Spinosauridae remain unclear. Here, new spinosaurid specimens from the Kem Kem beds of Morocco reveal aquatic adaptations in the cranium. Elevated orbits and bending of the frontals placed the eyes atop the skull, as in semiaquatic animals such as crocodiles and hippos. Two morphologies are present, a smaller morph characterized by narrow, triangular frontals, and a larger morph characterized by broad, subrectangular frontals overlapping the prefrontals. The two morphs suggest two distinct spinosaurine taxa, and are tentatively referred to the spinosaurines Spinosaurus cf. aegyptiacus and Sigilmassasaurus brevicollis, respectively. Semiaquatic habits were widespread within the Spinosaurinae and at least two distinct aquatic spinosaurines inhabited the Cenomanian of North Africa, challenging previous assumptions that non-avian dinosaurs were solely terrestrial. The appearance of giant semiaquatic dinosaurs may have followed the disappearance of giant pholidosaurid crocodylomorphs, suggesting that the extinction of large crocodylomorphs was associated with the rise of dinosaurs as apex predators in the freshwater ecosystem in North Africa.
The alpha taxonomy of dinosaurs encounters difficulties from the inability to apply biological species criteria to fossil material, and the prevalence of small sample sizes and of incomplete specimens. These enforce a quasi-typological praxis, in which differences in form must be used to distinguish species. Ecological arguments that one or few conspecific herbivores will occupy a given area and that the larger the body size of a terrestrial herbivore the less likely the existence of a sympatric closely closely related herbivore species, are not compelling. Ontogenetic variation in large theropods involves predominantly changes in proportion of skeletal structures, such as limbs, and some changes in proportions of individual bones. Individual variation involves largely changes in proportions of individual bones and also in the number of serial elements (teeth). -from Author
Ontogenetic variation is documented within many dinosaur species, but extreme ontogenetic changes are rare among dinosaurs, particularly among theropods. Here, we analyze 19 specimens of the Jurassic ceratosaurian theropod Limusaurus inextricabilis, representing six ontogenetic stages based on body size and histological data. Among 78 ontogenetic changes we identify in these specimens, the most unexpected one is the change from fully toothed jaws in the hatchling and juvenile individuals to a completely toothless beaked jaw in the more mature individuals, representing the first fossil record of ontogenetic edentulism among the jawed vertebrates. Jaw morphological data, including those derived from Mi-CT and SR-μCT scanning of Limusaurus specimens, reveal dental alveolar vestiges and indicate that ontogenetic tooth loss in Limusaurus is a gradual, complex process. Our discovery has significant implications for understanding the evolution of the beak, an important feeding structure present in several tetrapod clades, including modern birds. This radical morphological change suggests a dietary shift, probably from omnivory for juvenile Limusaurus to herbivory for adult Limusaurus, which is also supported by additional evidence from gastroliths and stable isotopes. Incorporating new ontogenetic information from Limusaurus into phylogenetic analyses demonstrates surprisingly little effect on its placement when data from different stages are used exclusively, in contrast to previous analyses of tyrannosaurids, but produces subtle differences extending beyond the placement of Limusaurus.
Recent fieldwork in the Upper Cretaceous (Maastrichtian) Maevarano Formation, northwest Madagascar, has yielded important new skull material of the abelisaurid theropod, Majungasaurus crenatissimus. One of these specimens in particular - a virtually complete, disarticulated, and well preserved skull - greatly elucidates the craniofacial osteology of abelisaurids. Herein we describe the skull and lower jaws of this mid-sized theropod dinosaur. A number of features of the facial skeleton and cranium (as well as the postcranium) appear to result from increased levels of mineralization and ossification, which, at least in some instances, can be related directly to specific soft-tissue structures; examples include lacrimal-postorbital contact dorsal to the orbit, suborbital processes of the lacrimal and postorbital, presence of a mineralized interorbital septum, fused interdental plates, and mineralization of the overlying integument. Autapomorphic features include a highly derived nasal - greatly thickened and fused to its counterpart, with a large interior pneumatic chamber - and a median, 'dome'-like thickening of the frontals, which appear to have been variably pneumatized by a paranasal air sac. Majungasaurus also possesses a derived suite of skull morphologies, including: a rostrocaudally abbreviated, dorsoventrally deep, and transversely broad skull; an expanded occiput, likely associated with expanded cervical musculature; short-crowned dentition; and an enlarged external mandibular fenestra consistent with a moderate degree of intramandibular movement or accommodation. A number of characters, present on both the skull and postcranial skeleton, suggest a divergent mode of predation relative to other, non-abelisaurid theropods.