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New fossils of Abelisauridae (Dinosauria: Theropoda) from the upper Maastrichtian of Morocco, North Africa
Abstract
The end of the Cretaceous saw the evolution of endemic dinosaur faunas on different landmasses, driven by continental fragmentation. Understanding the evolution of these biogeographic patterns is important for understanding the evolution of Mesozoic ecosystems. However, the faunas of the southern land masses remain understudied relative to the intensively sampled dinosaur faunas of western North America and Asia. In particular, the latest Cretaceous of Africa remains largely unknown, with only a handful of taxa reported so far, including titanosaurian sauropods, the lambeosaurine Ajnabia odysseus, and the large abelisaurid theropod Chenanisaurus barbaricus. We report two new abelisaurid fossils from the upper Maastrichtian phosphates of the Ouled Abdoun Basin, in northern Morocco. The first is the tibia of a medium-sized abelisaurid from Sidi Chennane, with an estimated length of ~5 m. The tibia has a strongly hooked cnemial crest resembling that of the South American Quilmesaurus and Aucasaurus. The highly rugose bone texture suggest the animal was mature, rather than a juvenile of the larger Chenanisaurus. The second is a small right second metatarsal from Sidi Daoui,. The metatarsal measures 190 mm in length, suggesting a small animal, ~2.6 m in length. The metatarsal shows strong mediolateral compression, a feature present in noasaurids and some early abelisaurids, but absent in most Late Cretaceous abelisaurids. It is distinct from other abelisauroids in the strong constriction and bowing of the shaft in lateral view, and the medial curvature of the bone in anterior view. Bone texture suggests it comes from a mature individual. The small size, gracile proportions and unusual shape of the metatarsal suggest it is not closely related to other latest Cretaceous abelisaurids. The new fossils suggest as many as three abelisaurid taxa coexisted in the late Maastrichtian of Morocco, showing dinosaurs were highly diverse in North Africa prior to the end-Cretaceous mass extinction.
... Such a history predicts a classic vicariance scenario, whereby phylogenies should reflect the recency of connectedness of the geographic units inhabited [177]. There is evidence for vicariance, including endemism among Late Cretaceous faunas [178], statistically supported area relationships in phylogenies that conform to predictions derived from palaeogeography [106,132,161], maximum likelihood ancestral area estimations that support 'best-fit' models that incorporate vicariance [104,115,154,155] and phylogenetic network biogeographic analyses that demonstrate greater faunal similarities between geographic areas that were more recently in contact [133,179] ( §6b). However, other studies have expressed scepticism about the importance of continent-scale vicariance, pointing to the supposedly cosmopolitan nature of Cretaceous dinosaurian faunas and anomalous phylogenetic relationships that are incongruent with palaeogeography [108][109][110]144,[180][181][182] ( §4c). ...
... There has been a considerable influx of new data on dinosaurs during the past few decades (e.g. [235,236]), partly driven by fieldwork that has targeted biogeographically important or under-sampled regions such as Antarctica [204,215], Alaska [140,213], Australia [111,148,237,238] and Africa [115,154,155,167,178,181,195]. These new data, combined with ongoing database construction [231,239] and phylogenetic work, are filling some key gaps in our understanding of dinosaur biogeography, as illustrated by progress regarding Australian faunas ( §4c). ...
Dinosaurs potentially originated in the mid-palaeolatitudes of Gondwana 245–235 million years ago (Ma) and may have been restricted to cooler, humid areas by low-latitude arid zones until climatic amelioration made northern dispersals feasible ca 215 Ma. However, this scenario is challenged by new Carnian Laurasian fossils and evidence that even the earliest dinosaurs had adaptations for arid conditions. After becoming globally distributed in the Early–Middle Jurassic (200–160 Ma), dinosaurs experienced vicariance driven by Pangaean fragmentation. Regional extinctions and trans-oceanic dispersals also played a role, and the formation of ephemeral land connections meant that older vicariance patterns were repeatedly overprinted by younger ones, creating a reticulate biogeographic history. Palaeoclimates shaped dispersal barriers and corridors, including filters that had differential effects on different types of dinosaurs. Dinosaurian biogeographic research faces many challenges, not the least of which is the patchiness of the fossil record. However, new fossils, extensive databasing and improved analytical methods help distinguish signal from noise and generate fresh perspectives. In the future, developing techniques for quantifying and ameliorating sampling biases and modelling the dispersal capacities of dinosaurs are likely to be two of the key components in our modern research programme.
... The presence of abelisaurid remains in a marine sedimentary environment is uncommon but not exceptional. Previous records of abelisaurids in marine formations include the holotype of Genusaurus sisteronis, from Albian deposits in south-eastern France [25] and various remains from the Maastrichtian phosphates of Morocco [26][27][28] and Egypt [29,30]. However, only very few dinosaur remains, belonging to ornithopods and ankylosaurs, have hitherto been reported from the Chalk of the Anglo-Paris basin [31]. ...
... As for the European majungasaurines, a connection via Africa seems to be the only possible option, although furileusaurians have not yet been recorded from Africa. It should be noted, however, that an incomplete tibia from the upper Maastrichtian phosphates at Sidi Chennane (Morocco) shows a markedly downturned end of the cnemial crest [28]. As noted above, this is considered as a distinctive character of Furileusauria by Filippi et al. [16]. ...
An articulated group of skeletal elements comprising a sacrum, both ilia and a first caudal vertebra, plus an isolated tooth found in immediate proximity to the bones, from the lower Cenomanian Chalk at Saint-Jouin-Bruneval (Seine-Maritime, Normandy, France) is described and attributed to a new genus and species of abelisaurid theropod, Caletodraco cottardi, on the basis of several characters of the sacrum and pelvis. The peculiar shape of the transverse process of the first caudal vertebra shows that Caletodraco cottardi differs from majungasaurine abelisaurids previously described from Europe, such as Arcovenator escotae, and belongs to the Furileusauria, a group of derived abelisaurids hitherto recognized only from South America. The presence of a furileusaurian abelisaurid in the Cenomanian of Normandy suggests that the biogeographical history of the Abelisauridae in Europe was more complex than hitherto admitted. Several previously described European abelisaurids, such as the Albian Genusaurus sisteronis, may in fact belong to the Furileusauria.
... Other marine reptiles include pachyvaranid squamates [42], elasmosaurid plesiosaurs [43,44], chelonioid turtles [45,46], and rare crocodylomorphs [47]. A diverse pterosaur fauna is also known [48,49]; dinosaurs are rare but diverse [50][51][52][53]. Bony fish [54] and selachians [54,55] are highly diverse and abundant. ...
Late Cretaceous marine ecosystems saw a major adaptive radiation of mosasaurids, which evolved highly disparate jaw and tooth morphologies to feed on different prey. A striking pattern seen in mosasaurids was a high diversity of durophagous forms. Durophagy likely evolved several times independently in the mosasaurine genera Globidens, Prognathodon, and Carinodens. Carinodens is unusual in having low, rectangular, laterally compressed teeth. The genus is known from around the world, with the species Carinodens minalmamar and C. belgicus previously reported from the Maastrichtian Phosphates of Morocco. Here, we report a new species of Carinodens, C. acrodon, from the same Maastrichtian Phosphates. It is characterized by teeth with tall crowns, triangular apices, and broad bases. Many of the diagnostic features of this species appear to be plesiomorphies, suggesting the persistence of a primitive species of Carinodens into the latest Maastrichtian alongside the more derived C. minalmamar and C. belgicus. The new species contributes to an emerging pattern of mosasaurid hyperdiversity in the late Maastrichtian of Morocco. A revised faunal list, including stratigraphic range extensions of Khinjaria and Stelladens into the latest Maastrichtian of upper Couche III of the Phosphates of Morocco, suggests at least 16 species of mosasauroid coexisted here.
... Wilson et al., 2003), northern Africa (e.g. Longrich et al., 2023), and Madagascar (e.g. Krause et al., 2007;Farke and Sertich, 2013). ...
Gondwanan dinosaur faunae during the 20 Myr preceding the Cretaceous-Palaeogene (K/Pg) extinction included several line-ages that were absent or poorly represented in Laurasian landmasses. Among these, the South American fossil record contains diverse abelisaurids, arguably the most successful groups of carnivorous dinosaurs from Gondwana in the Cretaceous, reaching their highest diversity towards the end of this period. Here we describe Koleken inakayali gen. et sp. n., a new abelisaurid from the La Colonia Formation (Maastrichtian, Upper Cretaceous) of Patagonia. Koleken inakayali is known from several skull bones, an almost complete dorsal series, complete sacrum, several caudal vertebrae, pelvic girdle and almost complete hind limbs. The new abelisaurid shows a unique set of features in the skull and several anatomical differences from Carnotaurus sas-trei (the only other abelisaurid known from the La Colonia Formation). Koleken inakayali is retrieved as a brachyrostran abeli-saurid, clustered with other South American abelisaurids from the latest Cretaceous (Campanian-Maastrichtian), such as Aucasaurus, Niebla and Carnotaurus. Leveraging our phylogeny estimates, we explore rates of morphological evolution across ceratosaurian lineages, finding them to be particularly high for elaphrosaurine noasaurids and around the base of Abelisauridae, before the Early Cretaceous radiation of the latter clade. The Noasauridae and their sister clade show contrasting patterns of morphological evolution, with noasaurids undergoing an early phase of accelerated evolution of the axial and hind limb skeleton in the Jurassic, and the abelisaurids exhibiting sustained high rates of cranial evolution during the Early Cretaceous. These results provide much needed context for the evolutionary dynamics of ceratosaurian theropods, contributing to broader understanding of macroevolutionary patterns across dinosaurs.
... new ichnosp., has an uncommon morphology. The fact that its pedal ray III has the main supporting function in walking or running gait, would then probably depend on laterally compressed metatarsals II and IV, such as those of V. paranaensis (or also, for example, the II one of the Sidi Daoui small-sized, gracile abelisaurid of Morocco, of Longrich et al., 2023) so that the central element (metatarsus III) would have borne most of the body load during locomotion. Besides, similar conditions (Langer et al., 2019) are present in other Noasaurinae. ...
The Botucatu Formation (Paraná Basin, Brazil and the southern neighboring countries) comprises one of the richest tetrapod ichnological deposits of the Lower Cretaceous in South America. The ichnofossils are found in reddish sandstones lithofacies -interpreted to be dune and interdune deposits. The sandstones of Botucatu Formation originally covered a surface estimated in at least 1,300,000 km2, the largest known fossil desert in the Earth’s history. The distribution area of the Botucatu paleodesert presents one of the world’s largest megatracksites. The tetrapod ichnofauna from the Botucatu Formation comprises minor
bipedal dinosaur tracks (almost all attributable to theropods, with one exception, a doubt ornithopod trackway) along with many thousands of footprints of early mammals, and a single trackway of a lacertiform reptile. Among the bipedal dinosaur footprints, the most common and typical are considered theropod tracks, with long strides and high step angle and always with an acuminate termination. These trackways are straight and very narrow, with long strides and step angles showing high values. The footprints have a relatively large and very wide III digit and small, short, pointed, bladelike outer digits. The most special characteristic
is that the II digit is longer and more important than the IV digit. Because of this morphology, the general outline of the footprint often reminds that of a waterdrop and, although it is structurally tridactyl, it usually looks like functionally monodactylous. These tracks are herein assigned to small theropods adapted to desert life with a prevalently cursorial gait, probably ancestors of clades such as noasaurs and velocisaurs. Due its unique morphological aspects it is defined a new ichnogenus and ichnospecies.
In the Late Cretaceous, northern and southern hemispheres evolved distinct dinosaurian faunas. Titanosaurians and abelisaurids dominated the Gondwanan continents; hadrosaurids, ceratopsians and tyrannosaurs dominated North America and Asia. Recently, a lambeosaurine hadrosaurid, Ajnabia odysseus, was reported from the late Maastrichtian phosphates of the Oulad Abdoun Basin Morocco, suggesting dispersal between Laurasia and Gondwana. Here we report new fossils from the phosphates of Morocco showing lambeosaurines achieved high diversity in the late Maastrichtian of North Africa. A skull represents a new dwarf lambeosaurine, Minqaria bata. Minqaria resembles Ajnabia odysseus in size, but differs in the ventrally positioned jugal facet and sinusoidal toothrow. The animal is small, ~ 3.5 m long, but the fused braincase shows it was mature. A humerus and a femur belong to larger hadrosaurids, ~ 6 m long, implying at least three species coexisted. The diversity of hadrosaurids in Europe and Africa suggests a dispersal-driven radiation, with lambeosaurines diversifying to take advantage of low ornithischian diversity. African lambeosaurines are small compared to North American and Asia hadrosaurids however, perhaps due to competition with titanosaurians. Hadrosaurids are unknown from eastern Africa, suggesting Moroccan hadrosaurids may be part of a distinct insular fauna, and represent an island radiation.
Tyrannosaurs are among the most intensively studied and best-known dinosaurs. Despite this, their relationships and systematics are highly controversial. An ongoing debate concerns the validity of Nanotyrannus lancensis, interpreted either as a distinct genus of small-bodied tyrannosaur or a juvenile of Tyrannosaurus rex. We examine multiple lines of evidence and show that the evidence strongly supports recognition of Nanotyrannus as a distinct species for the following reasons: 1. High diversity of tyrannosaurs and predatory dinosaurs supports the idea that multiple tyrannosaurids inhabited the late Maastrichtian of Laramidia; 2. Nanotyrannus lacks characters supporting referral to Tyrannosaurus or Tyrannosaurinae but differs from T. rex in >150 morphological characters, while intermediate forms combining the features of Nanotyrannus and T. rex are unknown; 3. Histology shows specimens of Nanotyrannus showing (i) skeletal fusions, (ii) mature skull bone textures, (iii) slow growth rates relative to T. rex, (iv) decelerating growth in their final years of life, and (v) growth curves predicting adult masses of~1500 kg or less, showing these animals are subadults and young adults, not juvenile Tyrannosaurus; 4. growth series of other tyrannosaurids, including Tarbosaurus and Gorgosaurus, do not show morphological changes proposed for a Nanotyrannus-Tyrannosaurus growth series, and deriving Tyrannosaurus from Nanotyrannus requires several changes inconsistent with known patterns of dinosaur development; 5. Juvenile T. rex exist, showing diagnostic features of Tyrannosaurus; 6. Phylogenetic analysis suggests that Nanotyrannus may lie outside Tyrannosauridae. Tyrannosaur diversity before the K-Pg extinction is higher than previously appreciated. The challenges inherent in diagnosing species based on fossils mean paleontologists may be systematically underestimating the diversity of ancient ecosystems.
In the dusk of the Mesozoic, advanced duck-billed dinosaurs (Hadrosauridae) were so successful that they likely outcompeted other herbivores, contributing to declines in dinosaur diversity. From Laurasia, hadrosaurids dispersed widely, colonizing Africa, South America, and, allegedly, Antarctica. Here, we present the first species of a duck-billed dinosaur from a subantarctic region, Gonkoken nanoi, of early Maastrichtian age in Magallanes, Chile. Unlike duckbills further north in Patagonia, Gonkoken descends from North American forms diverging shortly before the origin of Hadrosauridae. However, at the time, non-hadrosaurids in North America had become replaced by hadrosaurids. We propose that the ancestors of Gonkoken arrived earlier in South America and reached further south, into regions where hadrosaurids never arrived: All alleged subantarctic and Antarctic remains of hadrosaurids could belong to non-hadrosaurid duckbills like Gonkoken. Dinosaur faunas of the world underwent qualitatively different changes before the Cretaceous-Paleogene asteroid impact, which should be considered when discussing their possible vulnerability.
Mosasaurids, a clade of specialized marine squamates, saw a major adaptive radiation in the Late Cretaceous, evolving a wide range of body sizes, shapes, and specialized tooth morphologies. The most diverse known mosasaurid faunas come from the late Maastrichtian phosphates of Morocco. Here, we report an unusual new mosasaurid, Stelladens mysteriosus, based on a partial jaw and associated tooth crowns from lower Couche III phosphatic deposits at Sidi Chennane, Oulad Abdoun Basin, Morocco. Stelladens is characterized by short, triangular tooth crowns with a series of strong, elaborate, and serrated ridges on the lingual surface of the tooth, functioning as accessory carinae. Morphology of the teeth and associated jaw fragment suggest affinities with Mosasaurinae. No close analogues to the unique tooth morphology of Stelladens are known, either extant or extinct. It may have had an unusual and highly specialized diet, a specialized prey-capture strategy, or both. The diversity of mosasaurid teeth is much higher than that of plesiosaurs, ichthyosaurs, or extant marine mammals, and likely reflects both the ecological diversity of mosasaurids and complex developmental mechanisms responsible for tooth formation in mosasaurines. Mosasaurid diversity continued to increase up to the Cretaceous–Paleogene boundary.
Abelisaurids are medium–large-sized theropod dinosaurs that were predominant in the carnivorous fauna during the Late Cretaceous of Gondwana. These predators are abundant in the Cretaceous fossil strata of Patagonia, which yield the best record for this group. In the Late Cretaceous, abelisaurids appear in almost all regions of Gondwana and in all stages, except for the Coniacian, in which they are globally unknown. Here we describe a new abelisaurid, Elemgasem nubilus gen. et sp. nov., from the Portezuelo Formation (Turonian–Coniacian), Patagonia, Argentina. The palaeohistology of the appendicular bones of Elemgasem shows that the holotype was a subadult individual, but had achieved sexual maturity. This taxon is based on several axial and appendicular elements, and is diagnosed by the presence of a marked pattern of rugosity on the lateral surface of the fibula and a dorsoventrally deep lateral wall of the calcaneum. Moreover, the posterior caudal vertebrae have a morphology slightly different from any other abelisaurid. Elemgasem nubilus is recovered as an unstable taxon within Brachyrostra, given that it was recovered as sister taxon of Furileusauria or in several positions within this clade. Despite the problematic phylogenetic relationships of Elemgasem nubilus, it is important because it is the first abelisaurid from the Turonian–Coniacian interval and it increases the diversity of this theropod family at a time of marked turnover in the tetrapod fauna of South America, global climate change, and mass extinction events recorded worldwide in the marine realm.
The early evolution of thyreophoran dinosaurs is thought to have occurred primarily in northern continents since most evidence comes from the Lower and Middle Jurassic of Europe and North America. The diversification into stegosaurs and ankylosaurs is obscured by a patchy fossil record comprising only a handful of fragmentary fossils, most with uncertain phylogenetic affinities. Here we report the discovery of a new armoured dinosaur from the early Late Cretaceous of Argentina, recovered phylogenetically using various datasets either as a basal thyreophoran or a stem ankylosaur, closely related to Scelidosaurus. It bears unusual anatomical features showing that several traits traditionally associated with the heavy Cretaceous thyreophorans did not occur universally. Jakapil kaniukura gen. et sp. nov. is the first definitive thyreophoran species from the Argentinian Patagonia. Unlike most thyreophorans, it seems to show a bipedal stance, as in Scutellosaurus. Jakapil also shows that early thyreophorans had a much broader geographic distribution than previously thought. It is a member of an ancient basal thyreophoran lineage that survived until the Late Cretaceous in South America.
The oxygen isotope compositions of carbonate and phosphatic fossils hold the key to understanding Earth-system evolution during the last 500 million years. Unfortunately, the validity and interpretation of this record remain unsettled. Our comprehensive compilation of Phanerozoic δ¹⁸O data for carbonate and phosphate fossils and microfossils (totaling 22,332 and 4615 analyses, respectively) shows rapid shifts best explained by temperature change. In calculating paleotemperatures, we apply a constant hydrosphere δ¹⁸O, correct seawater δ¹⁸O for ice volume and paleolatitude, and correct belemnite δ¹⁸O values for ¹⁸O enrichment. Similar paleotemperature trends for carbonates and phosphates confirm retention of original isotopic signatures. Average low-latitude (30° S–30° N) paleotemperatures for shallow environments decline from 42.0 ± 3.1 °C in the Early-to-Middle Ordovician to 35.6 ± 2.4 °C for the Late Ordovician through the Devonian, then fluctuate around 25.1 ± 3.5 °C from the Mississippian to today. The Early Triassic and Middle Cretaceous stand out as hothouse intervals. Correlations between atmospheric CO2 forcing and paleotemperature support CO2’s role as a climate driver in the Paleozoic.
Megaraptora is a theropod clade known from former Gondwana landmasses and Asia. Most members of the clade are known from the Early to Late Cretaceous (Barremian–Santonian), with Maastrichtian megaraptorans known only from isolated and poorly informative remains. The aim of the present contribution is to describe a partial skeleton of a megaraptorid from Maastrichtian beds in Santa Cruz Province, Argentina. This new specimen is the most informative megaraptoran known from Maastrichtian age, and is herein described as a new taxon. Phylogenetic analysis nested the new taxon together with other South American megaraptorans in a monophyletic clade, whereas Australian and Asian members constitute successive stem groups. South American forms differ from more basal megaraptorans in several anatomical features and in being much larger and more robustly built.
Our understanding of sauropodomorph evolution is continually improved with the recognition of new species and their inclusion in phylogenetic data sets, along with the incorporation of novel characters. Whereas some subclades remain stable in terms of their original taxonomic content, the definitions of others have proven to be less easily applicable as increased taxon and character sampling has changed sauropodomorph interrelationships. Relatively minor differences in the position of clade specifiers between alternative phylogenies result in substantial differences in clade composition in several instances. Titanosauria is the most successful and diverse clade of sauropodomorph dinosaurs, but its interrelationships are poorly understood. With an ever-growing number of recognized species and increased taxon sampling in phylogenies, resulting in the recovery of newly identified, diverse subclades, the need to re-evaluate titanosaurian supra-specific taxonomy becomes increasingly important. Using fifteen phylogenetic data sets, we conducted a series of analyses to detect the most stable clades and taxa amongst Titanosauria. Based on this, we propose several changes to titanosaur systematics, including a newly defined node-stem triplet composed of Eutitanosauria and its constituent lineages, Colossosauria and Saltasauroidea, developing a new framework to improve stability and facilitate communication of phylogenetic results.
Here we describe a new hadrosaurid from the Upper Cretaceous (Campanian-Maastrichtian) Allen Formation of R ıo Negro Province, northwest Patagonia. The new taxon is based on cranial and postcranial elements from subadult and adult specimens. The new taxon may have reached 8-9 m in total body length, and it is diagnosed by a unique combination of characters, including a very low maxilla with respect to the dentary, and a dentary with a prominent and elongate anterior process with a deep groove on its anterior end, among other features. The new taxon is known from well-preserved elements and constitutes one of the most complete hadrosaurids known from South America. Features of the teeth, cranial and postcranial bones of the new taxon overlap with those of other hadrosaurid specimens previously recorded from this continent. This new evidence allows us to recognize that Secernosaurus koerneri, Bonapartesaurus rionegrensis and 'Kritosaurus' australis are valid taxa. The latter can be distinguished from the other South American taxa, as well as from Kritosaurus from North America. Thus, a new generic name is proposed for the species 'Kritosaurus' australis. Phylogenetic analysis recovers a new clade of South American hadrosaurids composed of the new taxon, 'Kritosaurus' australis, Bonapartesaurus and Secernosaurus. The new South American clade is tentatively nested among Kritosaurini and is supported by several traits, including an ilium with a subhorizontal ridge separating the preacetabular notch from the pubic pedicle, a longitudinal ridge on the dorsal surface of the postacetabular process, and a twisted distal end of the postacetabular process. The recovery of a monophyletic clade of South American hadrosaurids indicates that the history of the clade on Gondwanan landmasses is far from well understood, and new discoveries may change the current picture of the taxonomy and phylogenetic relationships of southern duck-billed dinosaurs. http://zoobank.org/urn:lsid:zoobank.org:pub:6C6F0056-9D3A-4097-A10E-2E33C9DB76B9
Armoured dinosaurs are well known for their evolution of specialized tail weapons—paired tail spikes in stegosaurs and heavy tail clubs in advanced ankylosaurs1. Armoured dinosaurs from southern Gondwana are rare and enigmatic, but probably include the earliest branches of Ankylosauria2–4. Here we describe a mostly complete, semi-articulated skeleton of a small (approximately 2 m) armoured dinosaur from the late Cretaceous period of Magallanes in southernmost Chile, a region that is biogeographically related to West Antarctica5. Stegouros elengassen gen. et sp. nov. evolved a large tail weapon unlike any dinosaur: a flat, frond-like structure formed by seven pairs of laterally projecting osteoderms encasing the distal half of the tail. Stegouros shows ankylosaurian cranial characters, but a largely ancestral postcranial skeleton, with some stegosaur-like characters. Phylogenetic analyses placed Stegouros in Ankylosauria; specifically, it is related to Kunbarrasaurus from Australia6 and Antarctopelta from Antarctica7, forming a clade of Gondwanan ankylosaurs that split earliest from all other ankylosaurs. The large osteoderms and specialized tail vertebrae in Antarctopelta suggest that it had a tail weapon similar to Stegouros. We propose a new clade, the Parankylosauria, to include the first ancestor of Stegouros—but not Ankylosaurus—and all descendants of that ancestor. Stegouros elengassen, an ankylosaur from the late Cretaceous of Chile, has a large tail weapon, named a macuahuitl after the Aztec club, with a frond-like structure formed by seven pairs of laterally projecting osteoderms encasing the distal half of the tail.
The question why non-avian dinosaurs went extinct 66 million years ago (Ma) remains unresolved because of the coarseness of the fossil record. A sudden extinction caused by an asteroid is the most accepted hypothesis but it is debated whether dinosaurs were in decline or not before the impact. We analyse the speciation-extinction dynamics for six key dinosaur families, and find a decline across dinosaurs, where diversification shifted to a declining-diversity pattern ~76 Ma. We investigate the influence of ecological and physical factors, and find that the decline of dinosaurs was likely driven by global climate cooling and herbivorous diversity drop. The latter is likely due to hadrosaurs outcompeting other herbivores. We also estimate that extinction risk is related to species age during the decline, suggesting a lack of evolutionary novelty or adaptation to changing environments. These results support an environmentally driven decline of non-avian dinosaurs well before the asteroid impact.
Mosasaurids (Mosasauridae) were specialized marine lizards that evolved and radiated in the Late Cretaceous. Their diversity peaked in the Maastrichtian, with the most diverse faunas known from Morocco. Here we describe a new species of mosasaurid from this fauna. Pluridens serpentis sp. nov. is described based on two complete skulls and referred jaws. It is referred to Pluridens based on the elongate and robust jaws, small teeth, and specialized tooth implantation. Pluridens is referred to Halisaurinae based on the posteriorly expanded premaxilla, long premaxilla-maxilla suture, broad premaxillary facet on the maxilla, closed otic notch, and small, striated, hooked teeth. The orbits are reduced relative to other halisaurines while the snout is robust and flat with a broad, rounded tip. The jaws bear numerous small, hooked, snake-like teeth. Skulls imply lengths of 5-6 meters; referred material suggests lengths of ≥10 meters. Pluridens’ specialized morphology – especially the contrasting large size and small teeth - suggests a distinct feeding strategy. Small orbits imply that P. serpentis relied on nonvisual cues including touch and chemoreception during foraging, as in modern marine snakes. Numerous neurovascular foramina on the premaxillae are consistent with this idea. The small teeth suggest proportionately small prey. The dentary becomes massive and robust in the largest individuals, suggesting sexual selection and perhaps sexual dimorphism, with the mandibles possibly functioning for combat as in modern beaked whales and lizards. The new mosasaur emphasizes how Maastrichtian mosasaurids were characterized by high species richness, functional diversity of niches occupied, and a certain degree of endemism, i.e. geographic specialization. and continued diversifying until the end of the Cretaceous, just prior to the K-Pg extinction.
We report theropod caudal vertebrae found at Phu Wiang Mountain, Thailand. They resemble the Portuguese Baryonyx and pertain to the Spinosauridae based on the presence of striations on the surface of the transverse process of the caudals, well developed double keels and a deep ventral groove on the centra, two laminae, delimiting three fossae below the transverse process, and posterior caudals having curved, rod-like neural spines with small process at the base. This supports the presence of spinosaurids in the Sao Khua Formation of Thailand. Furthermore, the putative basal ceratosaur Camarillasaurus Sánchez- Hernández and Benton, 2014 from Spain is found here to be a spinosaurid based on the resemblance of the type materials to the Thai spinosaurid described in the present work, as well as the phylogenetic analysis and the similarity to other spinosaurids. In Europe, spinosaurids have been reported from England, Portugal, and Spain. The reassessment of Camarillasaurus adds to the number of this group in this region. The presence of more than one spinosaurid taxon in the same region is common and can be found in the Kem Kem Beds of Morocco, the Araripe Basin of Brazil, the Iberian Peninsula, and the Khorat Plateau of Southeast Asia.
The mosasaurids (Mosasauridae) were a group of lizards that became highly specialized for marine life in the mid-Cretaceous. By the end of the Cretaceous, they had undergone an adaptive radiation, and showed a wide range of body sizes, locomotor styles, and diets. Their ranks included piscivores, apex predators, and durophages. Here, we report a new taxon, Xenodens calminechari gen. et sp. nov., from the upper Maastrichtian phosphates of Morocco, with dental specializations unlike those of any known reptile. Teeth form a unique dental battery in which short, laterally compressed and hooked teeth formed a saw-like blade. Unique features of tooth structure and implantation suggest affinities with the durophagous Carinodens. The tooth arrangement seen in Xenodens not only expands known disparity of mosasaurids, but is unique among Squamata, or even Tetrapoda. The specialized dentition implies a previously unknown feeding strategy, likely involving a cutting motion used to carve pieces out of large prey, or in scavenging. This novel dental specialization adds to the already considerable disparity and functional diversity of the late Maastrichtian mosasaurids and marine reptiles. This provides further evidence for a diverse marine fauna just prior to the K-Pg extinction.
We describe two partial postcranial skeletons belonging to the enigmatic theropod dinosaur clade Megaraptoridae from the Upper Cretaceous (lower Cenomanian-upper Turonian) Bajo Barreal Formation of southern Chubut Province, central Patagonia, Argentina. The specimens are assigned to Megaraptoridae due to their possession of multiple anatomical features that are considered synapomorphies of that predatory dinosaur group, such as a greatly enlarged, laterally compressed ungual of manual digit I that possesses asymmetrical lateral and medial vascular grooves. Overlapping elements of the two skeletons are nearly identical in morphology, suggesting that they probably represent the same taxon, a large-bodied theropod that was previously unknown from the early Late Cretaceous of southern South America. The Bajo Barreal specimens constitute the most ancient unquestionable records of Megaraptoridae from that continent, and exhibit particularly strong osteological resemblances to penecontemporaneous megaraptorids from the Winton Formation of Australia. Phylogenetic analysis recovers the unnamed Bajo Barreal taxon as the earliest-diverging South American megaraptorid and the oldest-known representative of this clade that likely attained a body length of at least seven meters and a mass of at least one metric ton. Overall, the balance of the evidence suggests that megaraptorids originated in eastern Gondwana (Australia) during the Early Cretaceous, then subsequently dispersed to western Gondwana (South America) during the mid-Cretaceous, where they attained substantially larger body sizes, ultimately coming to occupy the apex predator niches in their respective habitats.
The Late Cretaceous saw distinctly endemic dinosaur faunas evolve in the northern and southern hemispheres. The Laurasian continents of North America and Asia were dominated by hadrosaurid and ceratopsian ornithischians, with tyrannosaurs as apex predators. In Gondwanan communities, including Africa, South America, India and Madagascar, titanosaurian sauropods dominated as herbivores and abelisaurids as predators. These patterns are thought to be driven by the breakup of Pangaea and formation of seaways limiting dispersal. Here, we report a new lambeosaurine hadrosaurid, Ajnabia odysseus gen. et sp. nov., from the upper Maastrichtian of Morocco, North Africa, the first Gondwanan representative of a clade formerly thought to be restricted to Laurasia. The new animal shows features unique to Hadrosauridae and specifically Lambeosaurinae. Phylogenetic analysis recovers it within Arenysaurini, a clade of lambeosaurines previously known only in Europe. Biogeographic modelling shows that lambeosaurines dispersed from Asia to Europe, then to Africa. Given the existence of large, persistent seaways isolating Africa and Europe from other continents, and the absence of the extensive, bidirectional interchange characterizing land bridges, these patterns suggest dispersals across marine barriers, similar to those seen in Cenozoic mammals, reptiles, and amphibians. Dispersal across marine barriers also occurs in other hadrosaurid lineages and titanosaurian sauropods, suggesting oceanic dispersal played a key role in structuring Mesozoic terrestrial dinosaur faunas.
Abelisaurid theropods dominated the predator role across Gondwana during the Late Cretaceous. They are characterized by highly reduced forelimbs and one of the most specialized cranial morphologies among carnivorous dinosaurs, exemplified by a broad skull, short rostrum, high occipital region, and highly kinetic intramandibular joint, suggestive of a specialized feeding strategy. Late Cretaceous abelisaurids are known from some remarkably complete taxa with well-preserved skulls. However, little is known about the pattern of character transformation that led to their highly modified condition because there are no well-preserved abelisaurids before the Late Cretaceous. Here we report a basal abelisaurid from the Early Cretaceous of Brazil that preserves a complete skull and reveals an early stage in the cranial evolution of the group. It lacks the specialized temporal and mandibular features characteristic of derived abelisaurids, including the kinetic intramandibular joint and knob-like dorsal projection of the parietals.
Noasaurines form an enigmatic group of small-bodied predatory theropod dinosaurs known from the Late Cretaceous of Gondwana. They are relatively rare, with notable records in Argentina and Madagascar, and possible remains reported for Brazil, India, and continental Africa. In south-central Brazil, the deposits of the Bauru Basin have yielded a rich tetrapod fauna, which is concentrated in the Bauru Group. The mainly aeolian deposits of the Caiuá Group, on the contrary, bear a scarce fossil record composed only of lizards, turtles, and pterosaurs. Here, we describe the first dinosaur of the Caiuá Group, which also represents the best-preserved theropod of the entire Bauru Basin known to date. The recovered skeletal parts (vertebrae, girdles, limbs, and scarce cranial elements) show that the new taxon was just over 1 m long, with a unique anatomy among theropods. The shafts of its metatarsals II and IV are very lateromedially compressed, as are the blade-like ungual phalanges of the respective digits. This implies that the new taxon could have been functionally monodactyl, with a main central weight-bearing digit, flanked by neighbouring elements positioned very close to digit III or even held free of the ground. Such anatomical adaptation is formerly unrecorded among archosaurs, but has been previously inferred from footprints of the same stratigraphic unit that yielded the new dinosaur. A phylogenetic analysis nests the new taxon within the Noasaurinae clade, which is unresolved because of the multiple alternative positions that Noasaurus leali can acquire in the optimal trees. The exclusion of the latter form results in positioning the new dinosaur as the sister-taxon of the Argentinean Velocisaurus unicus.
In the lead-up to the Cretaceous/Paleogene mass extinction, dinosaur diversity is argued to have been either in long-term decline, or thriving until their sudden demise. The latest Cretaceous (Campanian–Maastrichtian [83–66 Ma]) of North America provides the best record to address this debate, but even here diversity reconstructions are biased by uneven sampling. Here we combine fossil occurrences with climatic and environmental modelling to quantify latest Cretaceous North American dinosaur habitat. Ecological niche modelling shows a Campanian-to-Maastrichtian habitability decrease in areas with present-day rock-outcrop. However, a continent-wide projection demonstrates habitat stability, or even a Campanian-to-Maastrichtian increase, that is not preserved. This reduction of the spatial sampling window resulted from formation of the proto-Rocky Mountains and sea-level regression. We suggest that Maastrichtian North American dinosaur diversity is therefore likely to be underestimated, with the apparent decline a product of sampling bias, and not due to a climatically-driven decrease in habitability as previously hypothesised.
Pterosaurs were the first vertebrates to evolve powered flight and the largest animals to ever take wing. The pterosaurs persisted for over 150 million years before disappearing at the end of the Cretaceous, but the patterns of and processes driving their extinction remain unclear. Only a single family, Azhdarchidae, is definitively known from the late Maastrichtian, suggesting a gradual decline in diversity in the Late Cretaceous, with the Cretaceous–Paleogene (K-Pg) extinction eliminating a few late-surviving species. However, this apparent pattern may simply reflect poor sampling of fossils. Here, we describe a diverse pterosaur assemblage from the late Maastrichtian of Morocco that includes not only Azhdarchidae but the youngest known Pteranodontidae and Nyctosauridae. With 3 families and at least 7 species present, the assemblage represents the most diverse known Late Cretaceous pterosaur assemblage and dramatically increases the diversity of Maastrichtian pterosaurs. At least 3 families—Pteranodontidae, Nyctosauridae, and Azhdarchidae—persisted into the late Maastrichtian. Late Maastrichtian pterosaurs show increased niche occupation relative to earlier, Santonian-Campanian faunas and successfully outcompeted birds at large sizes. These patterns suggest an abrupt mass extinction of pterosaurs at the K-Pg boundary.
Prominent hypotheses advanced over the past two decades have sought to characterize the Late Cretaceous continental vertebrate palaeobiogeography of Gondwanan landmasses, but have proved difficult to test because terrestrial vertebrates from the final ~30 million years of the Mesozoic are extremely rare and fragmentary on continental Africa (including the then-conjoined Arabian Peninsula but excluding the island of Madagascar). Here we describe a new titanosaurian sauropod dinosaur, Mansourasaurus shahinae gen. et sp. nov., from the Upper Cretaceous (Campanian) Quseir Formation of the Dakhla Oasis of the Egyptian Western Desert. Represented by an associated partial skeleton that includes cranial elements, Mansourasaurus is the most completely preserved land-living vertebrate from the post-Cenomanian Cretaceous (~94-66 million years ago) of the African continent. Phylogenetic analyses demonstrate that Mansourasaurus is nested within a clade of penecontemporaneous titanosaurians from southern Europe and eastern Asia, thereby providing the first unambiguous evidence for a post-Cenomanian Cretaceous continental vertebrate clade that inhabited both Africa and Europe. The close relationship of Mansourasaurus to coeval Eurasian titanosaurians indicates that terrestrial vertebrate dispersal occurred between Eurasia and northern Africa after the tectonic separation of the latter from South America ~100 million years ago. These findings counter hypotheses that dinosaur faunas of the African mainland were completely isolated during the post-Cenomanian Cretaceous.
The phosphates of Morocco range over a period of about 24 million years (m.y.), from the end of the Cretaceous (Maastrichtian) up to the base of the Middle Eocene (Lutetian), which is the longest interval of all phosphate deposits from the Mediterranean and Atlantic “Phosphogenic Provinces”. These marine sediments have recorded the evolution of life in the epicontinental seas outside of the western African craton and its continental hinterland; they are characterised by their remarkable richness in fossil vertebrate remains and constitute an almost unique palaeobiodiversity “hot-spot” at the Cretaceous-Tertiary turnover, in close relations with the rich organic inqux by the deep upwelling currents along the West African oceanic margin.
These outcrops are characterised by a singular combination of factors that highlights their value: 1) the fossil faunas of the Oulad Abdoun and Ganntour basins range along the whole phosphatic series and include all main vertebrate groups; 2) fossils are exceptional in terms of abundance, diversity and preservation; 3) they document a 24 m.y. key-period of vertebrate evolution that can be followed in situ, and record two major biological crises (Cretaceous/ Tertiary (K/Pg) and Palaeocene/Eocene (P/E) boundaries); 4) they originate from still poorlyknown palaeobiogeographical provinces (South-Tethys and Atlantic margins and Arabo-Africanprovince); 5) the scientiWc exploitation of these outcrops is possible in the long term; 6) the collected fossils not only have a great scientiWc signiWcance (several ongoing studies), but also patrimonial (worldwide reference collection) and museological (long term discoveries; creation of a museum) values.
Since the pioneer works of the French palaeontologist Camille Arambourg between 1935 and 1952, this exceptional palaeontological heritage has been scientiWcally neglected except for the wellstudied selachians. In 1996, the unexpected discovery of continental mammals in these phosphate marine deposits revived the interest of palaeonlogists and lead to the signature in 1997 of a French- Moroccan research Agreement between the Centre National de la Recherche ScientiWque (CNRS), the Muséum National d’Histoire Naturelle (MNHN), the OfWce ChériWen des Phosphates Group (OCP), and the Ministère de l’Energie, des Mines, de l’Eau et de l’Environnement (MEMEE).
In 2005, this Agreement was extended to the Universities Cadi Ayyad of Marrakech (UCAM) and Chouaîb Doukkali of El Jadida (UCDJ). Our multidisciplinary studies in the framework of this Agreement called “Phosphapal” not only permitted to improve our knowledge of the fossil vertebrates from the Phosphates of Morocco, their palaeoenvironments and palaeobiogeography, but also constitute a large palaeontological reference collection that motivates the creation by the OCP Group of a Museum of Paleontology in the city of Khouribga.
Fossil remains from the phosphates of Morocco cover all vertebrate groups, except amphibians: cartilaginous (selachians) and bony (actinopterygians) Wshes, reptiles (including birds) and mammals, which, in the current state of knowledge, correspond to 332 species, belonging to 192 genera and 86 families. Most taxa are marine but some very rare ones are terrestrial. Some of these groups, such as non-avian dinosaurs, birds, pterosaurs and mammals were totally unknown at the time of Arambourg’s works.
Selachians are, by far, the most abundant and diversiWed fossil group with 250 species of sharksand rays. Beyond their strictly palaeontological (taxonomical) interest, they constitute a standard biostratigraphical tool to date and correlate phosphate deposits along the whole South-Tethyan margin. Actinopterygian Wshes are mainly represented by teleosts and are abundant in all levels. They remain mostly undescribed apart from some punctual works. Reptiles (including birds) are also very abundant and diversiWed (at least 55 species) and are often represented by complete and spectacular specimens. Except some terrestrial taxa (non-avian dinosaurs and pterosaurs), they mainly include marine forms belonging to chelonians, squamates, plesiosaurians, crocodyliformes and birds. This palaeoherpetofauna of the phosphates of Morocco represents a worldwide reference for Late Cretaceous - Early Palaeogene marine reptile faunas of the South-Tethyan margin. Continental mammals, the most unexpected group in these marine series, are represented by, at least, ten species belonging to the hyaenodontidan, proboscidean, hyracoid and “condylarth” orders. They are among the oldest placentals of Africa and the oldest modern ungulate fossils.
During the latest Cretaceous, distinct dinosaur faunas were found in Laurasia and Gondwana. Tyrannosaurids, hadrosaurids, and ceratopsians dominated in North America and Asia, while abelisaurids and titanosaurids dominated in South America, India, and Madagascar. Little is known about dinosaur faunas from the latest Cretaceous of Africa, however. Here, a new abelisaurid theropod, Chenanisaurus barbaricus, is described from the upper Maastrichtian phosphates of the Ouled Abdoun Basin in Morocco, North Africa on the basis of a partial dentary and isolated teeth. Chenanisaurus is both one of the largest abelisaurids, and one of the youngest known African dinosaurs. Along with previously reported titanosaurid remains, Chenanisaurus documents the persistence of a classic Gondwanan abelisaurid-titanosaurid fauna in mainland Africa until just prior to the end-Cretaceous mass extinction. The animal is unusual both in terms of its large size and the unusually short and robust jaw. Although it resembles South American carnotaurines in having a deep, bowed mandible, phylogenetic analysis suggests that Chenanisaurus may represent a lineage of abelisaurids that is distinct from those previously described from the latest Cretaceous of South America, Indo-Madagascar, and Europe, consistent with the hypothesis that the fragmentation of Gondwana led to the evolution of endemic dinosaur faunas during the Late Cretaceous.
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.
We describe the partially preserved femur of a large-bodied theropod dinosaur from the Cenomanian “Kem Kem Compound Assemblage” (KKCA) of Morocco. The fossil is housed in the Museo Geologico e Paleontologico “Gaetano Giorgio Gemmellaro” in Palermo (Italy). The specimen is compared with the theropod fossil record from the KKCA and coeval assemblages from North Africa. The combination of a distally reclined head, a not prominent trochanteric shelf, distally placed lesser trochanter of stout, alariform shape, a stocky shaft with the fourth trochanter placed proximally, and rugose muscular insertion areas in the specimen distinguishes it from Carcharodontosaurus, Deltadromeus and Spinosaurus and supports referral to an abelisaurid. The estimated body size for the individual from which this femur was derived is comparable to Carnotaurus and Ekrixinatosaurus (up to 9 meters in length and 2 tons in body mass). This find confirms that abelisaurids had reached their largest body size in the “middle Cretaceous,” and that large abelisaurids coexisted with other giant theropods in Africa. We review the taxonomic status of the theropods from the Cenomanian of North Africa, and provisionally restrict the Linnean binomina Carcharodontosaurus iguidensis and Spinosaurus aegyptiacus to the type specimens. Based on comparisons among the theropod records from the Aptian-Cenomanian of South America and Africa, a partial explanation for the so-called “Stromer’s riddle” (namely, the coexistence of many large predatory dinosaurs in the “middle Cretaceous” record from North Africa) is offered in term of taphonomic artifacts among lineage records that were ecologically and environmentally non-overlapping. Although morphofunctional and stratigraphic evidence supports an ecological segregation between spinosaurids and the other lineages, the co-occurrence of abelisaurids and carcharodontosaurids, two groups showing several craniodental convergences that suggest direct resource competition, remains to be explained.
Resumen
The Late Cretaceous (Maastrichtian) Lameta Formation of central India has yielded dissociated elements of a variety of predatory dinosaurs, most of them coming from a quarry named the "Carnosaur bed." The materials were described by Huene and Matley nearly 70 years ago. They recognized nine theropod species, which they sorted out into the theropod subgroups "Carnosauria" and "Coelurosauria". Huene and Matley also described a considerable amount of theropod hindlimb bones (e.g., femora, tibiae, metatarsals, and pedal phalanges) that they could not refer to any of these species, but vaguely interpreted as corresponding to "allosaurid" or "coelurosaurid" theropods. We reviewed the available collection of Cretaceous theropods from Bara Simla housed at the Geological Survey of India, Calcutta, arriving to the following conclusions: 1) Indosuchus and Indosaurus are abelisaurids, as recognized by previous authors, but available information is not enough to judge whether they are synonyms; 2) Laevisuchus indicus is a small abelisauroid, related to Noasaurus and Masiakasaurus on the basis of their peculiar cervical vertebrae; 3) the controversial taxa " Compsosuchus", " Dryptosauroides", " Ornithomimoides", and " Jubbulpuria" are represented by isolated vertebrae corresponding to different portions of the neck and tail, and also exhibit abelisauroid features; 4) hindlimb bones originally referred to as "allosaurid" and "coelurosaurian" also exhibit abelisauroid characters, and bones of large size are tentatively referred to as corresponding to Indosuchus or Indosaurus, whereas some pedal bones of smaller size may belong to Laevisuchus; 5) two kinds of abelisaurid feet are apparent: one in which the phalanges of digit III and IV are robust, and another type in which the phalanges of digit IV are transversely narrow and dorsoventrally deep. This review demonstrates that all of the theropod elements discovered at the "Carnosaur bed" belong to a single theropod clade, the Abelisauroidea
Tyrannosaurs and hadrosaurs from the Late Cretaceous of eastern North America (Appalachia) are distinct from those found in western North America (Laramidia), suggesting that eastern North America was isolated during the Late Cretaceous. However, the Late Cretaceous fauna of Appalachia remains poorly known. Here, a partial maxilla from the Campanian Tar Heel Formation (Black Creek Group) of North Carolina is shown to represent the first ceratopsian from the Late Cretaceous of eastern North America. The specimen has short alveolar slots, a ventrally projected toothrow, a long dentigerous process overlapped by the ectopterygoid, and a toothrow that curves laterally, a combination of characters unique to the Leptoceratopsidae. The maxilla has a uniquely long, slender and downcurved posterior dentigerous process, suggesting a specialized feeding strategy. The presence of a highly specialized ceratopsian in eastern North America supports the hypothesis that Appalachia underwent an extended period of isolation during the Late Cretaceous, leading the evolution of a distinct dinosaur fauna dominated by basal tyrannosauroids, basal hadrosaurs, ornithimimosaurs, nodosaurs, and leptoceratopsids. Appalachian vertebrate communities are most similar to those of Laramidia. However some taxa-including leptoceratopsids-are also shared with western Europe, raising the possibility of a Late Cretaceous dispersal route connecting Appalachia and Europe.
Skorpiovenator bustingorryi is a derived abelisaurid theropod represented by a fairly complete skeleton from the Late Cretaceous sedimentary beds of north-western Patagonia. Although some features were described in the original paper, mainly related to the skull, the appendicular anatomy remains undescribed. The aim of the present contribution is to provide a detailed description and analysis of the available appendicular bones, including comparisons with other ceratosaurian theropods close to Skorpiovenator. In this way, new autapomorphies emerged to further distinguish Skorpiovenator from its relatives. Furthermore, a comprehensive phylogenetic analysis was performed and several characteristics of the hind limb, in particular some of the autopodium, resulted in the identification of new apomorphic traits for Ceratosauria and Abelisauridae. These features might prove to be useful for future phylogenetic analyses and may help to resolve the still confusing and debated internal relationships of abelisaurid theropods.
A B S T R A C T
The Cretaceous-Paleogene (K-Pg) transition saw mass extinctions in terrestrial and marine ecosystems. Terrestrial vertebrate diversity patterns across the K-Pg boundary have seen extensive study, but less is known about marine vertebrates. We describe a new mosasaurid from the latest Maastrichtian phosphatic beds of Morocco, showing how mosasaurids evolved to become apex predators in the latest Cretaceous. Thalassotitan atrox n. gen. et sp., from the Oulad Abdoun Basin of Khouribga Province, Morocco is characterized by large size, a broad skull, massive jaws, and reduced cranial kinesis, suggesting it was highly adapted for carnivory. Teeth resemble those of killer whales in their robust, conical shape, and show heavy wear and damage. Phylogenetic analysis recovers Thalassotitan as a close relative of Prognathodon currii and P. saturator within the Prognathodontini. Among the associated fauna, three genera of mosasaurids, elasmosaurid plesiosaur, chelonioid turtle, and enchodontid fish show acid damage, and could be prey ingested by mosasaurids, likely Thalassotitan. Thalassotitan shows mosasaurids evolved to fill the marine apex predator niche, a niche occupied by orcas and white sharks today. Mosasaurs continued to diversify and fill new niches until their extinction at the end of the Cretaceous.
Although osteohistology has proven to be useful to discern several palaeobiological traits and systematics in non-avian dinosaurs, the bone microstructure of several groups (e.g., abelisaurid theropods) is still poorly documented. Here, we provide a microstructural description of different bones (a cervical epipophysis, three dorsal ribs, three grastralia, and the left femur) of the type specimen of the abelisaurid Aucasaurus garridoi, its ontogenetic stage and compare its growth with other theropods, especially abelisauroid taxa. The cortical bone is mostly composed by fibrolamellar bone tissue composed of a woven-fibred matrix. Secondary remodelling varies among the different elements. The compacta of the rib shows eleven growth marks (with an additional seven in the external fundamental system), implying that the specimen was at least eleven years old and had reached somatic maturity when it died. The presence of lines of arrested growth/annuli supports a cyclical growth strategy, as in Niebla, Quilmesaurus, and Abelisauridae indet. MMCh-PV 69. All these abelisaurids indicate that the maturity based on cyclical growth marks spacing is not reliable, due to considerable variability of this parameter within a single element. This study represents the first intensive osteohistological investigation for an Abelisauridae member, providing new information on palaeobiological traits of this clade.
The fossil record of ornithischians in South America is sparse, and they are clearly underrepresented when compared with sauropod dinosaurs. However, recent discoveries indicate that ornithischians were more diversified than thought. The aim of the present contribution is to describe isolated remains belonging to ankylosaurs, and ornithopods, including basal euiguanodontians and hadrosaurs coming from the Chorrillo Formation (upper Campanian–lower Maastrichtian), Santa Cruz province, southern Argentina. The fossil remains of ankylosaurs reported here are the southernmost recorded for the continent. They show a unique combination of plesiomorphic features, indicating that they may belong to a basal ankylosaur. Ankylosaurs and hadrosaurids are thought to have arrived in South America during the latest Cretaceous through Central America. However, a detailed overview of the fossil record of Gondwana indicates that both clades were present and probably diversified along southern continents. This indicates that their presence in South America may be alternatively interpreted as the result of migration from other landmasses, including Africa and Europe, or may even be the result of Jurassic–Early Cretaceous vicariance from their northern counterparts.
An epicontinental sea bisected West Africa periodically from the Late Cretaceous to the early Eocene, in dramatic contrast to the current Sahara Desert that dominates the same region today. Known as the Trans-Saharan Seaway, this warm and shallow ocean was a manifestation of globally elevated sea level associated with the rapid break-up of the supercontinent Gondwana in the late Mesozoic. Although it varied in size through time, the Trans-Saharan Seaway is estimated to have covered as much as 3000 km² of the African continent and was approximately 50 m deep. The edges of the sea were defined in part by the high topography of the Precambrian cratons and mobile belts of West Africa. Over its approximately 50 million year episodic existence, through six major periods of transgression and regression, the Trans-Saharan Seaway left behind extensive nearshore marine sedimentary strata with abundant fossils. The waters that yielded these deposits supported and preserved the remains of numerous vertebrate, invertebrate, plant, and microbial species that are now extinct. These species document a regional picture of ancient tropical life that spanned two major Earth events: the Cretaceous-Paleogene (K-Pg) boundary and the Paleocene-Eocene Thermal Maximum (PETM). Whereas extensive epeiric seas flooded the interior portions of most continents during these intervals, the emerging multicontinental narrative has often overlooked the Trans-Saharan Seaway, in part because fundamental research, including the naming of geological formations and the primary description and analysis of fossil species, remained to be done. We provide such synthesis here based on two decades of fieldwork and analyses of sedimentary deposits in the Republic of Mali. Northern parts of the Republic of Mali today include some of the farthest inland reaches of the ancient sea.
We bring together and expand on our prior geological and paleontological publications and provide new information on ancient sedimentary rocks and fossils that document paleoequatorial life of the past. Ours is the first formal description of and nomenclature for the Upper Cretaceous and lower Paleogene geological formations of this region and we tie these names to regional correlations over multiple modern territorial boundaries. The ancient seaway left intriguing and previously unclassified phosphate deposits that, quite possibly, represent the most extensive vertebrate macrofossil bone beds known from anywhere on Earth. These bone beds, and the paper shales and carbonates associated with them, have preserved a diverse assemblage of fossils, including a variety of new species of invertebrates and vertebrates, rare mammals, and trace fossils. The shallow marine waters included a wide range of paleoenvironments from delta systems, to hypersaline embayments, protected lagoons, and carbonate shoals.
Our overarching goal has been to collect vertebrate fossils tied to a K-Pg stratigraphic section in Africa. We provide such a section and, consistent with prior ideas, indicate that there is a gap in sedimentation in Malian rocks in the earliest Paleocene, an unconformity also proposed elsewhere in West Africa. Our phylogenetic analyses of several vertebrate clades across the K-Pg boundary have clarified clade-by-clade species-level survivorship and range extensions for multiple taxa. Few macrofossil species from the Trans-Saharan Seaway show conspicuous change at either the K-Pg boundary or the PETM based on current evidence, although results are very preliminary. Building on our earlier report of the first record of rock-boring bivalves from the Paleocene of West Africa, we further describe here a Cretaceous and Paleogene mollusk fauna dominated by taxa characteristic of the modern tropics. “This abstract has been truncated. To view the complete abstract please download the PDF”
The Gondwanan hadrosaurid dinosaur record is relatively poorly known and very complicated. A new hadrosaurid, Bonapartesaurus rionegrensis, gen. et sp. nov., from the late Campanian–early Maastrichtian of the Salitral Moreno site, Argentina, is characterized by the following combination of characters: the ratio between the height of the neural spines and that of the centrum of the sacral vertebrae greater than 3.5; neural spines of the anterior section of the caudal vertebrae extremely long and evenly expanding distally; a preacetabular process of the ilium slightly deflected ventrally, with the angle greater than 150°; the ratio between the maximum dorsoventral depth of the posterior end of the preacetabular process and the dorsoventral distance from the pubic peduncle to the dorsal margin of the ilium less than 0.50; the ratio between the dorsoventral height and anteroposterior length of the iliac blade 0.8 or greater; asymmetrical lateral profile of the supraacetabular process; the posterior portion of the postacetabular process markedly thicker mediolaterally as a result of the dorsomedial twist of the postacetabular process; an anteriorly expanded cnemial crest restricted to the proximal end of the tibia; and articular surface of the astragalus for the internal malleolus of the tibia moderately expanded medially, articulating with only part of the ventral surface of the tibial internal malleolus. Our phylogenetic analysis indicates that Bonapartesaurus is located within the Tribe Saurolophini. Paleobiogeographically, its presence suggests at least two saurolophine dispersal events from North America, one toward South America and another toward Asia, no later than the late Campanian.
http://zoobank.org/urn:lsid:zoobank.org:pub:1ADEDB71-1FD6-4081-915D-6C73D8389408
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Citation for this article: Cruzado-Caballero, P., and J. Powell. 2017. Bonapartesaurus rionegrensis, a new hadrosaurine dinosaur from South America: implications for phylogenetic and biogeographic relations with North America. Journal of Vertebrate Paleontology. DOI: 10.1080/02724634.2017.1289381.
The upper Campanian-lower Maastrichtian site of Lo Hueco (Cuenca, Spain) has provided a set of well-preserved partial skeletons in anatomical connection or with a low dispersion of their skeletal elements. One partial skeleton is herein described and a new titanosaurian sauropod is established, Lohuecotitan pandafilandi. This titanosaur is diagnosed by eight autapomorphic features: dorsally and ventrally widened or bifurcated posterior centrodiapophyseal lamina in anterior and middle dorsal vertebrae; short postspinal lamina with a transversely expanded distal end represented by smooth scars in the dorsal vertebrae; anteriormost caudals with the medial spinoprezygapophyseal and medial spinopostzygapophyseal laminae ventrally connected with the prespinal and postspinal laminae, respectively; anterior caudal neural spines with a dorsal projection of the prespinal and postspinal laminae; anterior caudal neural spines bears a “greek-cross”-like cross-section; middle caudal centra having two round and rough structures in the dorsal edge of the posterior articulation, which extends to the dorsal surface of the centrum; the articular ends of the rami of the haemal arches are divided in two articular surfaces; and tuberosity between the anterior and the lateral trochanter of the fibula. The herein performed phylogenetic analysis considered L. pandafilandi as a member of Lithostrotia more derived than Malawisaurus. The known palaeodiversity of the Late Cretaceous Ibero-Armorican titanosaurs is increasing, and further analyses focused on this group will be necessary to better understand the evolutionary history of European titanosaurs and to clarify their relationships within Titanosauria.
Theropod dinosaurs from the Late Jurassic of Gondwana are still poorly known, with Elaphrosaurus bambergi Janensch, 1920, from the late Kimmeridgian of Tendaguru, Tanzania, being the only taxon represented by more than isolated remains from Africa. Having long been considered a coelurosaurian, more specifically an ornithomimosaur, Elaphrosaurus is currently regarded as a basal ceratosaur. Here, we revise the osteology and phylogenetic position of this important taxon. Elaphrosaurus shows many unusual osteological characters, including extremely elongated and constricted cervical vertebrae, an expansive shoulder girdle with strongly modified forelimbs, a relatively small ilium, and elongate hindlimbs with a very small ascending process of the astragalus that is fused to the tibia. We found this taxon to share many derived characters with noasaurids, such as: strongly elongate cervical and dorsal vertebrae; low, rectangular neural spines in the mid-caudal vertebrae; presence of only an anterior centrodiapophyseal lamina in anterior caudal vertebrae; presence of a wide, U-shaped notch between the glenoid and the anteroventral hook in the coracoid; a laterally flared postacetabular blade of the ilium; a flat anterior side of the distal tibia; and a reduced shaft of metatarsal II. Our analysis placed Elaphrosaurus within a dichotomous Noasauridae as part of a Jurassic subclade, here termed Elaphrosaurinae, that otherwise includes taxa from eastern Asia. These results underscore the long and complex evolutionary history of abelisauroids, which is still only beginning to be understood.
Tyrannosauroids are among the most distinctive and best known of Mesozoic theropods. Diagnostic skeletal material for Tyrannosauridae is at present limited to the last part of the Late Cretaceous of eastern and central Asia (China and Mongolia) and North America; the more inclusive Tyrannosauroidea includes taxa from the Late Jurassic of Europe and North America and the Early Cretaceous of Europe and Asia. Tyrannosaurids include some of the largest known theropod taxa (up to 13 m long and weighing six or more tonnes). The five most completely known species (Tyrannosaurus rex, Tarbosaurus bataar, Daspletosaurus torosus, Albertosaurus sarcophagus, and Gorgosaurus libratus) are all represented by individuals with femora more than 100 cm in length, reaching 138 cm in the largest Tyrannosaurus rex.
