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A NEUROVASCULAR CAVITY WITHIN THE SNOUT OF THE PREDATORY DINOSAUR SPINOSAURUS
Abstract
In 2005, Dal Sasso et al. described an almost complete snout of the unusual theropod Spinosaurus, from the Late Cretaceous of Morocco. Siemens CAT scan analysis, performed recently on the same specimen at the Ospedale Maggiore di Milano, reveals that the numerous foramina located on the outer wall of the rostrum communicate with a common internal cavity, deeply encased medially within the premaxillae, which is unique among theropod dinosaurs. The extreme retraction of the external nares in Spinosaurus permits to exclude any respiratory/olfactory function of this cavity, which in turn has a neurovascular nature. The cavity flows into two paired neurovascular passages, going through the entire snout and meeting caudally at level of the fifth maxillary tooth. This suggests that a rostral extension of the trigeminal nerve innervated the cavity.
Soares (2002) demonstrated that the foramina on the facial bones of living and extinct semi-aquatic crocodilians house dome pressure receptors, innervated by the trigeminal nerve, that are useful to hunt even in darkness, detecting prey-made pressure waves associated with disruptions to the air-water interface. According to Taquet (1984) and Holtz (2003), spinosaurids might have hunted in riparian habitats in a manner similar to herons. As for Spinosaurus, we postulate the presence of croc-like pressure receptors, that might have given its mouth, when positioned on the air-water interface, an unexpected tactile function, useful to catch swimming preys without relying on sight.
... Modern crocodilians possess trigeminal-innervated integumentary sensory organs for detecting prey in low-visibility environments (Munger and Ide, 1988;Leitch and Catania, 2012), while extant aquatic mammals possess vibrissae-related mechanoreceptors (pinnipeds, odontocetes: Ahl, 1986) and even electroreceptive organs (monotremes, odontocetes: Pettigrew and Wilkens, 2003;Czech-Damal et al., 2012). Similar adaptations have also been hypothesized for pliosaurid plesiosaurs (Foffa et al., 2014), mosasaurs ( Alvarez-Herrera et al., 2020) and even for semiaquatic theropods (Dal Sasso et al., 2009;Cau et al., 2017). However, neurovascular anatomy of the rostrum in these animals is extremely complex, with higher vascularization and denser branching than the ichthyosaurian condition (Lomax et al., 2019). ...
In 2016 a new Cretaceous ichthyosaur rostrum fragment (IG 251372) was found near Gombola, Modena Province, northern Italy. Two rostral fragments and a partial humerus were previously reported from the same locality. Despite their fragmentary nature, all these finds are relatively well preserved, with just slightly deformed shape and teeth still in place. Here we describe IG 251372, relying on CT-scan technology to allow visualization of the fossil internal structure. The new specimen has been ascribed to the sub-family Platypterygiinae based on the sub-rectangular profile of the tooth roots. Tooth crowns are robust and show a similar density of enamel ridges to many other Cretaceous ichthyosaurs historically referred to the genus Platypterygius. A detailed taphonomical analysis was performed on the material from Gombola. The ichthyosaurs previously ascribed to Platypterygius are now identified only to the subfamily level. CT-scans revealed an anterior neurovascular network within both the premaxilla and the dentary, here interpreted as mandibular and maxillary/ophthalmic divisions of the trigeminal nerve. Axial sections and videos show how these intraosseous channels emerge within the dental and premaxillary fossae. Cross section imaging of tooth roots showed an alternating pattern of tooth resorption and eruption, providing additional information on dental replacement in ichthyosaurs. IG 251372 has been attributed to the Albian-Cenomanian by means of nannofossils contained within the matrix, making it the first ichthyosaur from the Northern Apennines to be directly dated.
... Full-size DOI: 10.7717/peerj.6112/ fig-5 crocodilians and lepidosaurs, as well as those reported in theropod dinosaurs (Dal Sasso, Maganuco & Cioffi, 2009), pliosaurs (Ketchum & Benson, 2011;Foffa et al., 2014b) and plesiosaurs (Ketchum & Smith, 2010) and something similar in ichthyosaurs (Lomax & Massare, 2015). In extant taxa, these canals carry neurovascular bundles consisting of the maxillary artery and maxillary branch of the trigeminal nerve (CN V 2 ) in the upper jaw, and the inferior alveolar artery and mandibular branch of the trigeminal nerve (CN V 3 ) (Witmer, 1997). ...
Ichthyosaur fossils are abundant in Lower Jurassic sediments with nine genera found in the UK. In this paper, we describe the partial skeleton of a large ichthyosaur from the Lower Jurassic (lower Sinemurian) of Warwickshire, England, which was conserved and rearticulated to form the centrepiece of a new permanent gallery at the Thinktank, Birmingham Science Museum in 2015. The unusual three-dimensional preservation of the specimen permitted computed tomography (CT) scanning of individual braincase elements as well as the entire reassembled skull. This represents one of the first times that medical imaging and three-dimensional reconstruction methods have been applied to a large skull of a marine reptile. Data from these scans provide new anatomical information, such as the presence of branching vascular canals within the premaxilla and dentary, and an undescribed dorsal (quadrate) wing of the pterygoid hidden within matrix. Scanning also revealed areas of the skull that had been modelled in wood, clay and other materials after the specimen’s initial discovery, highlighting the utility of applying advanced imaging techniques to historical specimens. Additionally, the CT data served as the basis for a new three-dimensional reconstruction of the skull, in which minor damage was repaired and the preserved bones digitally rearticulated. Thus, for the first time a digital reconstruction of the skull and mandible of a large marine reptile skull is available. Museum records show the specimen was originally identified as an example of Ichthyosaurus communis but we identify this specimen as Protoichthyosaurus prostaxalis . The specimen features a skull nearly twice as long as any previously described specimen of P. prostaxalis , representing an individual with an estimated total body length between 3.2 and 4 m.
... Unlike in pike congers, olfaction was probably not involved in prey detection in spinosaurids. On the basis of the presence of snout neurovascular foramina (Dal Sasso et al. 2005), it has been suggested that Spinosaurus had a sensory integumentary system similar to that of other groups of long-snouted predators such as crocodilians (Dal Sasso et al. 2009Ibrahim et al. 2014) and pliosaurs (Foffa et al. 2014b). Pressure receptors and the enhanced tactile sensitivity would have helped in localizing and biting aquatic prey items under low-light conditions (i.e., in turbid waters or during crepuscular/nocturnal feeding activity), similarly to mechanoreceptive neuromasts present in the pored canals of the cephalic lateral line system of pike congers. ...
Spinosaurs represent a group of peculiar theropod dinosaurs that have often been described as “crocodile-mimic”,
predominantly fish-eating predators, and recently claimed to have been semi-aquatic animals. Here we report a suite of craniodental characters unexpectedly shared by spinosaurs and pike conger eels. Pike conger eels are predatory, mainly piscivorous bottom-dwelling anguilliform fishes that inhabit marine and brackish environments. These two groups of dinosaurs and fishes show a mediolaterally compressed, elongated rostrum, a terminal “rosette” bearing enlarged teeth in both upper and lower jaws, and a notch posterior to the premaxillary “rosette” characterized by the presence of reduced teeth. The morphological convergence observed in the jaws of these two distantly related groups of vertebrates may result from similar feeding behaviours. This typical jaw morphology likely represents an effective biomechanical adaptation for biting and grabbing elusive prey items in low-light aquatic environments. Associated with this specialized snout morphology, numerous integumentary mechanoreceptors involved in prey detection are present in both spinosaurs and pike congers. Our new observations provide an additional convincing argument regarding the decades-long and widely debated lifestyle of spinosaurs.
... Communicated by: Robert Reisz D. Foffa (*) : J. Sassoon : A. R. Cuff : M. J. Benton School of Earth Sciences, University of Bristol, Wills Memorial Building, Queens Road, Bristol BS8 1RJ, UK e-mail: df1818@my.bristol.ac.uk Spinosaurus (Dal Sasso et al. 2005 Sasso et al. , 2009), and it is sometimes possible to infer the distribution of soft tissues by making comparisons with extant taxa. However, for pliosaurids, there are no directly analogous living taxa for comparison. ...
Pliosaurs were a long-lived, ubiquitous group of Me-sozoic marine predators attaining large body sizes (up to 12 m). Despite much being known about their ecology and behaviour, the mechanisms they adopted for prey detection have been poorly investigated and represent a mystery to date. Complex neurovascular systems in many vertebrate rostra have evolved for prey detection. However, information on the occurrence of such systems in fossil taxa is extremely limited because of poor preservation potential. The neurovascular complex from the snout of an exceptionally well-preserved pliosaur from the Kimmeridgian (Late Jurassic, c. 170 Myr ago) of Weymouth Bay (Dorset, UK) is described here for the first time. Using computed tomography (CT) scans, the extensive bifurcating neurovascular channels could be traced through the rostrum to both the teeth and the foramina on the dorsal and lateral surface of the snout. The structures on the surface of the skull and the high concentrations of peripheral rami suggest that this could be a sensory system, perhaps similar to crocodile pressure receptors or shark electroreceptors.
... Due to their highly specialized skull and dentition, adapted for piscivory (e.g., Taquet 1984;Charig & Milner 1997;Sereno et al. 1998;Dal Sasso et al. 2005;Hendrickx & Buffetaut 2008;Dal Sasso et al. 2009), several features of spinosaurid teeth have already been used as synapomorphies by many authors (e.g., Sereno et al. 1998;Holtz et al. 2004;Benson 2010;Mateus et al. 2011;Carrano et al. 2012). Their teeth are indeed highly diagnostic (in fact the most diagnostic among theropods; see Table 2) as all spinosaurids possess subcircular mesialmost and lateral crowns displaying flutes (i.e., subparallel longitudinal grooves separated by acute ridges) on the lingual and/ or labial margin, minute denticles or no serrations at all on both mesial and distal carinae, and the enamel texture of Spinosaurus and baryonychines is deeply veined (or 'sculptured' sensu Hasegawa et al. 2010) and curves basally close to the carinae. ...
Theropod dinosaurs form a highly diversified clade, and their teeth are some of the most common components of the Mesozoic dinosaur fossil record. This is the case in the Lourinhã Formation (Late Jurassic, Kimmeridgian-Tithonian) of Portugal, where theropod teeth are particularly abundant and diverse. Four isolated theropod teeth are here described and identified based on morphometric and anatomical data. They are included in a cladistic analysis performed on a data matrix of 141 dentition-based characters coded in 60 taxa, as well as a supermatrix combining our dataset with six recent datamatrices based on the whole theropod skeleton. The consensus tree resulting from the dentition-based data matrix reveals that theropod teeth provide reliable data for identification at approximately family level. Therefore, phylogenetic methods will help identifying theropod teeth with more confidence in the future. Although dental characters do not reliably indicate relationships among higher clades of theropods, they demonstrate interesting patterns of homoplasy suggesting dietary convergence in (1) alvarezsauroids, therizinosaurs and troodontids; (2) coelophysoids and spinosaurids; (3) compsognathids and dromaeosaurids; and (4) ceratosaurids, allosauroids and megalosaurids. Based on morphometric and cladistic analyses, the biggest tooth from Lourinhã is referred to a mesial crown of the megalosaurid Torvosaurus tanneri, due to the elliptical cross section of the crown base, the large size and elongation of the crown, medially positioned mesial and distal carinae, and the coarse denticles. The smallest tooth is identified as Richardoestesia, and as a close relative of R. gilmorei based on the weak constriction between crown and root, the "eight-shaped" outline of the base crown and, on the distal carina, the average of ten symmetrically rounded denticles per mm, as well as a subequal number of denticles basally and at mid-crown. Finally, the two medium-sized teeth belong to the same taxon and exhibit pronounced interdenticular sulci between distal denticles, hooked distal denticles for one of them, an irregular enamel texture, and a straight distal margin, a combination of features only observed in abelisaurids. They provide the first record of Abelisauridae in the Jurassic of Laurasia and one of the oldest records of this clade in the world, suggesting a possible radiation of Abelisauridae in Europe well before the Upper Cretaceous.
At NAVEP1, Dal Sasso et al. (2009) showed the preliminary results of a computed tomography (CT) performed on a complete snout of the theropod dinosaur Spinosaurus aegyptiacus from the Late Cretaceous of Morocco (Dal Sasso et al., 2005). Recently the specimen was re-examined and the former CT data were analysed with new highly performant software. The sediment still filling even the small cavities was digitally removed, providing more detailed 3D images and allowing re-interpretation of the internal cranial anatomy of Spinosaurus.
The numerous foramina emerging on the outer wall of the rostrum do not communicate with a “common internal cavity, deeply encased medially within the premaxillae” (Dal Sasso et al., 2009), but rather with a complex neurovascular network, that does not involve neither that cavity, nor the alveoli of the teeth. The purported medial cavity is actually a combination of large deep alveoli, and of an artifact of preservation, i.e. a collapsed portion of the inter-premaxillary walls. Moreover, that cavity does not communicate with the tubular paired passages previously identified (Dal Sasso et al., 2009). In facts, rostrally these passages become fusiform and taper into pointed ends, surrounded by bone. We also found that at one-third of their length these passages connect to the subnarial foramina, and that caudally they converge into a single medial tube, which enters the aparturae nasi osseae. Consequently, we exclude the presence of rostral extensions of the ophthalmic ramus of the trigeminal nerve, and regard the fusiform passages as diverticula of the nasal sinuses. Equally reduced promaxillary sinuses show to originate from the anterior medial wall of the antorbital fenestra.
On the other hand, the neurovascular network that terminates in the numerous large foramina concentrated on the tip of the rostrum, confirms the presence of well-developed maxillary branches of the trigeminal nerve. By comparison with living crocodilians (Leitch & Catania, 2012), the rostral neurovascular network of Spinosaurus appears as an equally hypertrophied sensory organ, that likely had an important function in prey detection and manipulation.
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.
A gavialoid crocodylian from the Maastrichtian of the Oulad Abdoun phosphatic Basin (Morocco) is described, representing the oldest known crocodylian from Africa. The specimen consists of a skull that exhibits several features not found in other gavialoids, and a new genus and species is erected, Ocepesuchus eoafricanus. A phylogenetic analysis has been conducted including 201 characters and 71 taxa, where Ocepesuchus eoafricanus appears as the most basal African gavialoid, and the South American gavialoids are paraphyletic. This paraphyly has strong biogeographic implications, and the previous hypothesis of South American and Asian assemblages derived from African gavialoids should be reviewed. The historical biogeography of Gavialoidea is probably more complex than previously supposed. The phosphatic deposits of Morocco provide a unique opportunity to study the vertebrate faunal turnover across the Creta-ceous-Tertiary (KT) boundary. The crocodyliforms are very scarce in the Maastrichtian marine basins of Africa which are dominated by mosasaurid squamates. The latter became extinct by the KT boundary, while crocodyliforms survived and diversified in the Paleocene. Mosasaurids and crocodyliforms both lived in probably comparable marine environments during the Maastrichtian. The selectivity of the KT boundary extinctions remains to be explained; since freshwater environments are known for having been less affected by the KT crisis than marine ones, a freshwater lifestyle of the juveniles, like in extant marine crocodiles and unlike the fully marine mosasaurs, could explain this difference with regard to survivorship.
A large azhdarchid pterosaur is described from the Late Maastrichtian phosphatic deposits of the Oulad Abdoun Basin, near Khouribga (central Morocco). The material consists of five closely associated cervical vertebrae of a single individual. The mid-series neck vertebrae closely resemble those of azhdarchids Quetzalcoatlus and Azhdarcho in that they are elongate, with vestigial neural spines, prezygapophysial tubercles, a pair of ventral sulci near the prezygapophyses, and without pneumatic foramina on the lateral surfaces of the centra. The Moroccan pterosaur is referred to a new genus and species of Azhdarchidae: Phosphatodraco mauritanicus gen. et sp.nov. It is mainly characterized by a very long cervical vertebra eight, bearing a prominent neural spine located very posteriorly. Based on comparisons with azhdarchid vertebrae, the estimated wing span of Phosphatodraco is close to 5 m. This discovery provides the first occurrence of Late Cretaceous azhdarchids in northern Africa. Phosphatodraco is one of the few azhdarchids known from a relatively complete neck and one of the latest-known pterosaurs, approximately contemporaneous with Quetzalcoatlus.
Vertebrate microremains from the Late Devonian–Early Carboniferous of the Carnic Alps are predominantly chondrichthyan, with minor placoderm and actinopterygian remains. The faunas are sparse and, with very few exceptions, occur only in conodont-rich pelagic limestones (Pramosio Limestone) representative of the palmatolepid-bispathodid conodont biofacies. Phoebodont and jalodont chondrichthyans, also reflecting open-ocean environments, predominated during the Famennian, and eventually symmoriids seem to predominate during the Early Carboniferous. The presence of Siamodus in this assemblage gives a new locality for this genus known from few regions in the world and allows confirming its stratigraphical range (limpidus Zone) and its relation to deep-water environments. The Late Devonian vertebrate faunas are tropical and cosmopolitan, having much in common with coeval taxa from the North-Gondwanan margins and Asian terranes. Composition of the vertebrate faunas is consistent with the Carnic Alps terrane having occupied a position intermediate between Gondwana and Laurussia, as hypothesized by various authors, but because of sparsity of the taxa represented and the pronounced cosmopolitan nature of both the conodont and vertebrate faunas, the data are not compelling.
GINTER, M., HAIRAPETIAN, V. & KLUG, C. 2002. Famennian chondrichthyans from the shelves of North Gondwana. Acta Geologica Polonica, 52 (2), 169-215. Warszawa. Ichthyoliths, mainly shark teeth, from the Famennian of Iran and Northwest Africa are described. Evolution of shal-low-water chondrichthyan assemblages on the shelves of Central Iran and the Tafilalt Platform, Morocco, related to time and environmental changes, is discussed. Four new taxa, viz. Deihim mansureae gen. et sp. nov., Protacrodus serra sp. nov., Phoebodus depressus sp. nov., and Ph. gothicus transitans subsp. nov. are erected and provisional recon-structions of heterodonty in dentitions of several Famennian sharks are proposed.
Biological and physical factors govern the distribution of fossils, but it is not always clear which is more important. The preservation of late Eocene vertebrates at the UNESCO World Heritage site of Wadi Al-Hitan, Western Desert of Egypt, is controlled primarily by the physical processes responsible for sequence stratigraphic architecture on a siliciclastic shelf. Three types of stratigraphic surface, each characterized by a taxonomically and taphonomically distinct fossil assemblage, yield most of the known vertebrate fossils. Complete, partially articulated whale skeletons, primarily Basilosaurus isis, are abundant in offshore marine flooding surfaces (MFS) in the late transgressive systems tract (TST) of the first Priabonian sequence (TA4.1), where low net sedimentation rates and environmental averaging in offshore environments promoted the accumulation of carcasses on traceable stratigraphic surfaces. Complete, well-articulated whales, primarily Dorudon atrox, are more widely scattered on minor erosion surfaces in rapidly accumulating shoreface sediments of the overlying failing stage systems tract. Fragmented and abraded vertebrate remains are abundant and diverse in a discontinuous conglomerate that marks the first sequence boundary above the base of the Priabonian (Pr-2), which has not been previously recognized in Egypt, but which formed incised valleys with at least 45 in of total relief. Fossils in this variably thick lag conglomerate include skeletal elements reworked by rivers from underlying marine deposits and bones of terrestrial animals living in the fluvial environment. Marginal marine vertebrates, primarily dugongs, occur on shelly marine ravinement surfaces above Pr-2, in the early TST of the second Priabonian sequence. Most vertebrate remains in Wadi Al-Hitan occur in condensed stratigraphic intervals and taxonomic composition changes with sequence position, both important considerations in interpretation of paleobiological patterns.
We describe the first American stegosaur track of the ichnospecies Deltapodus brodricki, collected in the Upper Jurassic Morrison Formation of San Juan County, southeastern Utah, United States. The track is preserved as a natural cast on the underside of a slab of fluvial sandstone and consists of a well-preserved pes track and the eroded remains of a manus track. Previously, Deltapodus was known only from the Middle Jurassic Yorkshire coast of England and the Upper Jurassic of Portugal and Spain. The new discovery thus substantially extends the geographic record of this ichnospecies and highlights the similarities between the Late Jurassic dinosaur faunas of North America and those of Western Europe.
Known since the beginning of the century, the Iouaridène ichnosite has never been studied in detail, even if it was frequently cited for the presence of the Breviparopus taghbaloutensis r. The trampled layers belong to the lower member of the probably Upper Jurassic Iouaridène Formation. The sedimentological and stratigraphical work allowed to reconstruct the paleoenvironment as a fluvial flood basin, cyclically flooded. This work identifies 21 trampled layers, bearing probably more than 1000 tracks: a complete ichnocoenosis, with 12 different morphotypes, bearing traces of theropods, sauropods, ornithischians, thyreophora and other vertebrates and invertebrates animals. Such a large number of footprints were used to test and refine statistical and morphometrical methods applied to ichnology: PCA and Landmark analysis. The results, even if preliminary, are very promising and open new perspective for a more quantitave and objective ichnology and ichnotaxonomy.
Four trackways consisting mainly of sauropod manus prints were discovered from the Middle Jurassic of the central High Atlas Mountains, Morocco. This supports R.T.Bird's theory of sauropod swimming ability. -Author
An amiid fish of very large size (ca 3 m SL) from phosphates of probable early Eocene age in the Tilemsi Valley, Republic of Mali, is described as Maliamia gigas n. gen., n. sp. Maliamia is known only from isolated jaw bones (premaxilla, vomer, maxilla, dentary, dermopalatine, coronoid, prearticular), and is the first amiid to be described from Africa. It differs from other amiid genera in the vomerine and prearticular dentition, and so far as it and its relatives are known, is most closely related to Enneles from the early Cretaceous of Brazil.
Sauropod dinosaur remains have been discovered recently in the Late Cretaceous (Maastrichtian) phosphatic deposits of the Oulad Abdoun Basin, near Khouribga (central Morocco). The material consists of right hindlimb bones (femur, tibia and fibula) from a small-sized individual. The marine associated fauna, mainly selachians, actinopterygians, turtles, mosasaurids and plesiosaurs, suggests a marine depositional environment, so that the dinosaur remains may be a remnant of a floating carcass. The femur exhibits a prominent lateral bulge on the proximal one-third, a diagnostic feature of Titanosauriformes. The Moroccan sauropod lacks synapomorphies of Titanosauria and less inclusive clades (i.e., distal tibia expanded transversely to twice mid-shaft breadth; femoral distal condyles angled dorsomedially relative to the shaft); therefore, it is here assigned to a basal titanosauriform as Titanosauriformes indet. This is the first sauropod reported from the Maastrichtian of Morocco and one of the few dinosaur records from the uppermost Cretaceous formations of northern Africa. This discovery confirms the wide geographical distribution of Titanosauriformes during the Late Cretaceous and supports their survival into the Late Maastrichtian of Africa.
A proximal part of humerus from the basal Ypresian (lowermost Eocene) of the Ouled Abdoun Basin, Morocco, is described as a new genus and species tentatively assigned to the Phaethontidae (tropicbirds). This fossil possibly represents the oldest record of the Phaethontidae and markedly differs from Lithoptila, a contemporaneous Prophaethontidae from the same locality. This new taxon lived in a tropical climate and was probably an efficient flier with pelagic habits, like extant tropicbirds of the genus Phaethon.
A new parameter, the Trackway Ratio (TR), is proposed to supplement the previously used trackway gauge to describe and quantify the relative width of trackways in dinosaur quadrupedal gaits. It is expressed as the ratio of the width of the tracks relative to the total width of the trackway (both measured perpendicular to the long axis of the trackway). The ratio may be used with either pes (PTR) or manus (MTR) tracks. The PTR range of values for wide-, medium- and narrow-gauge trackways of previous authors are provisionally suggested to be ≤35%, 36–49% and ≥50%, respectively. The application of such a ratio would permit a more consistent ichnotaxonomy to be adopted where both track morphology and trackway parameters are used to define ichnotaxa.
Determination of the TR, as well as other parameters, will be affected by track preservation quality. Recent experiments on track simulation in the laboratory have shed further light on observations made in the field concerning the value of track measurements (in particular track length and width) recorded from below the surface on which the maker was moving. Experimental track simulations in the laboratory have shown that the dimensions of transmitted tracks preserved below the surface on which the foot was impressed may vary from 65% to 135% of the true dimensions of the indenter. Two case studies are presented that quantify the errors that may be made on calculating TR and the size, gait and speed of the maker, from trackways if the preservation of the tracks are not fully understood.
It is shown that in individual trackways the PTR may vary along the length of the trackway; so that part of the trackway may be classified as wide-gauge and other parts medium-gauge. There is a relationship between variation in PTR and that of pace angulation along the length of a single trackway. An analysis of 42 trackways, principally sauropod, shows a temporal distribution that does not agree closely with previous suggestions relating to narrow- and wide-gauge trackways.
New pterosaur remains consisting of jaw fragments of toothless taxa and isolated teeth are described from the red beds of the Kern Kern region of southern Morocco. The stratigraphic position of those red beds is discussed and it is concluded that they are in all likelihood early Cenomanian in age. At least four taxa of pterodactyloid pterosaurs are present. The toothless jaw fragments are referred to the families ?Pteranodontidae, ?Azhdarchidae and Tapejaridae. Four different morphotypes can be distinguished among the isolated teeth. They are tentatively referred to the Ornithocheiridae. This assemblage reveals a high diversity of pterosaurs in Africa during the early Upper Cretaceous. The possible occurrence of tapejarids and anhanguerids indicates relationships with the somewhat older pterosaur assemblage from the Santana Formation (Aptian/Albian) of Brazil. If confirmed, the presence of azhdarchids and pteranodontids in the early Cenomanian suggests an early differentiation of these essentially late Late Cretaceous groups of large pterosaurs.
A pterosaur is reported for the first time from a Late Cretaceous marine Lagerstätte in Lebanon, which is famous mainly for its rich ichthyofauna. The specimen, from the early Cenomanian HâqelLagerstätte , is a partial, articulated forelimb that belongs to a relatively small, adult individual of the clade Pteranodontoidea sensu Kellner. It is the most complete skeletal remnant of a pterosaur described to date from the African-Arabian Plate and the most complete of the very sparse Cenomanian pterosaur record.
Thèse--Strasbourg. Without thesis statement. t. 1. Le précambrien saharien au sud de l'Adrar des Iforas.--
Famennian chondrichthyans from the shelves of North Gondwana
- M Ginter
- V Hairapetian
- C Klug
Ginter, M., Hairapetian, V., Klug, C., 2002. Famennian chondrichthyans from the shelves of North Gondwana. Acta
Geologica Polonica, 52(2), 169-215.
Muséum national d'histoire naturelle
- Patrick De Wever
DE WEVER Patrick
Muséum national d'histoire naturelle, Géologie CP 48, 43
rue Buffon, 75005 Paris (pdewever@mnhn.fr)
18
Corresponding author: Intern. Tel. 33 320 336062; Fax
- France Villeneuve D'ascq Cedex
Villeneuve d'Ascq cedex, France. Corresponding author:
Intern. Tel. 33 320 336062; Fax. 33 320 434910;
Université Mohamed 1 er , Faculté des Sciences
- El Hammouti Kamal
EL HAMMOUTI Kamal
Université Mohamed 1 er, Faculté des Sciences, Laboratoire
Géosciences appliquées, Oujda, Maroc.
Département de Géologie
- El Kati Imad
EL KATI Imad
(LPGERN), Département de Géologie, Université Ibn Tofail,
Faculté des Sciences, BP. 133, Kenitra, Morocco.
(imadelkati@yahoo.fr)
35, 46, 47
34 GROESSENS Eric Institut Royal des Sciences Naturelles de France, DO Terre et Histoire de la Vie
- Goussard Florent
GOUSSARD Florent
CR2P, CNRS-MNHN-UPMC, Paris, France
34
GROESSENS Eric
Institut Royal des Sciences Naturelles de France, DO Terre et
Histoire de la Vie, Service Géologique de France, 13, rue
Jenner 1000 Bruxelles, France ;
(eric.groessens@sciencesnaturelles.be)
20
Les vertébrés fossiles des gisements de phosphates (Maroc-Algérie-Tunisie)
- C Arambourg
ARAMBOURG C. 1952. Les vertébrés fossiles des gisements de phosphates (Maroc-Algérie-Tunisie). Notes et
Mémoires du Service Géologique du Maroc, 92: 1-372.
A survey of pterosaurs from Africa with the description of a new specimen from Morocco
- W A Kellner A
- M S Mello A
- Ford T
KELLNER A.W.A., MELLO A.M.S. & FORD T. 2007. A survey of pterosaurs from Africa with the description of a
new specimen from Morocco. in CARVALHO I.S. et al. (eds.). Paleontologia: Cenários da Vida, Vol. 1.
Interciência, p. 257-267 (ISBN 978-85-7193-184-8).
Comments on the pterosaur fauna from Tendaguru, Upper Jurassic of Africa, with the indentification of a possible azhdarchid
- J M Sayão
- W A Kellner A
SAYÃO J.M. & KELLNER A.W.A.. 2001. Comments on the pterosaur fauna from Tendaguru, Upper Jurassic of
Africa, with the indentification of a possible azhdarchid. Boletim de resumos, 17. Congresso Brasileiro de
Paleontologia, p. 145, UFAC, Rio Branco, Acre.
Sedimentology, taphonomy, and ichnology of Late Jurassic dinosaur tracks from the Jura carbonate platform (Chevenez-Combe Ronde tracksite, NW Switzerland): insights into the tidal-flat environment and dinosaur diversity, locomotion, and palaeoecology
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carbonate platform (Chevenez-Combe Ronde tracksite, NW Switzerland): insights into the tidal-flat
environment and dinosaur diversity, locomotion, and palaeoecology. PhD Thesis University of Fribourg,
GeoFocus, 21, 278 pp.
Découverte de Dipneustes du genre Protopterus dans le Tertiaire ancient de Tamaguilelt (Soudan français)
- Lavocat R
LAVOCAT R. 1955. Découverte de Dipneustes du genre Protopterus dans le Tertiaire ancient de Tamaguilelt
(Soudan français). Compte Rendu Hebdomaire des Séances de l'Académie des Sciences. Paris, 240: 1915-1917.
Nouveaux lepidosirenides (Dipnoi) du Tertiaire africain
MARTIN M. 1995. Nouveaux lepidosirenides (Dipnoi) du Tertiaire africain. Geobios, Mémoire Special, 19: 275-280.
First evidence of Deltapodus (Dinosauria, Thyreophora) in northern Africa (Iouaridène Fm
- Belvedere M
- Mietto P. Submitted
BELVEDERE M. & MIETTO P. submitted. First evidence of Deltapodus (Dinosauria, Thyreophora) in northern
Africa (Iouaridène Fm., Upper Jurassic, Morocco). Palaeontology.
Museo del Jurásico de Asturias (MUJA), Colunga, 15-18 de octubre de
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(MUJA), Colunga, 15-18 de octubre de 2008. Museo del Jurásico de Asturias XVII, Colunga, Spain, 280 pp.
Chenini à Tataouine) connu surtout par des dents, épines et plaques de sélaciens (Protolamna, Cretodus) de Dipneustes (Neoceratodus africanus, cf. Hybodus) et de reptiles. Les vertébrés sont par contre rares dans le Crétacé supérieur
- Sauropodes
Sauropodes (Rebbachisaurus). -Albien inférieur (fm. Chenini à Tataouine) connu surtout par des
dents, épines et plaques de sélaciens (Protolamna, Cretodus) de Dipneustes (Neoceratodus
africanus, cf. Hybodus) et de reptiles. Les vertébrés sont par contre rares dans le Crétacé
supérieur. Seul un poisson Diplomystus solignaci est décrit du Sénonien d'El Hama (Gabès).
A middle-late Eocene vertebrate fauna (marine fish and mammals) from southwestern Morocco; preliminary report: age and palaeobiogeographical implications
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Adnet, S., Cappetta, H., Tabuce, R., 2010. A middle-late Eocene vertebrate fauna (marine fish and mammals) from
southwestern Morocco; preliminary report: age and palaeobiogeographical implications. Geol. Mag. 147, 860-870.
marocain): conservation et valorisation par la création d'un géoparc géo-écotouristique. Exceptional paleontological heritage of Zagora (South-eastern Morocco): conservation and valorization by the creation of a geo-ecotouristic geopark
- Le
- De Zagora
Le patrimoine paléontologique exceptionnel de Zagora (sud-est marocain):
conservation et valorisation par la création d'un géoparc géo-écotouristique.
Exceptional paleontological heritage of Zagora (South-eastern Morocco):
conservation and valorization by the creation of a geo-ecotouristic geopark.
Central Park West at 79 th Street
- H Landman Neil
- Amnh
LANDMAN Neil H.
AMNH, Central Park West at 79 th Street, New York, NY
10024, USA. (landman@amnh.org)
33
Université Montpellier 2, CC64, Place E. Bataillon, 34095-Montpellier Cedex 05, France. (laurent.marivaux@univ-montp2.fr)
- Laurent Marivaux
MARIVAUX Laurent
(ISE-M), UMR-CNRS 5554, Université Montpellier 2, CC64,
Place E. Bataillon, 34095-Montpellier Cedex 05, France.
(laurent.marivaux@univ-montp2.fr)
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