SciCombinator

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Concept: Spinosauridae

184

Countershading is common across a variety of lineages and ecological time [1-4]. A dark dorsum and lighter ventrum helps to mask the three-dimensional shape of the body by reducing self-shadowing and decreasing conspicuousness, thus helping to avoid detection by predators and prey [1, 2, 4, 5]. The optimal countershading pattern is dictated by the lighting environment, which is in turn dependent upon habitat [1, 3, 5, 6]. With the discovery of fossil melanin [7, 8], it is possible to infer original color patterns from fossils, including countershading [3, 9, 10]. Applying these principles, we describe the pattern of countershading in the diminutive theropod dinosaur Sinosauropteryx from the Early Cretaceous Jehol Biota of Liaoning, China. From reconstructions based on exceptional fossils, the color pattern is compared to predicted optimal countershading transitions based on 3D reconstructions of the animal’s abdomen, imaged in different lighting environments. Reconstructed patterns match well with those predicted for animals living in open habitats. Jehol is presumed to have been a predominantly closed forested environment [3, 11, 12], but our results indicate a more heterogeneous range of habitats. Sinosauropteryx is also shown to exhibit a “bandit mask,” a common pattern in many living vertebrates, particularly birds, that serves multiple functions including camouflage [13-18]. Sinosauropteryx therefore shows multiple color pattern features likely related to the habitat in which it lived. Our results show how reconstructing the color of extinct animals can inform on their ecologies beyond what may be obvious from skeletal remains alone. VIDEO ABSTRACT.

Concepts: Ecology, Bird, Extinction, Cretaceous, Dinosaur, Spinosauridae

108

Here we report on angiosperm-like pollen and Afropollis from the Anisian (Middle Triassic, 247.2-242.0 Ma) of a mid-latitudinal site in Northern Switzerland. Small monosulcate pollen grains with typical reticulate (semitectate) sculpture, columellate structure of the sexine and thin nexine show close similarities to early angiosperm pollen known from the Early Cretaceous. However, they differ in their extremely thin inner layer (nexine). Six different pollen types (I-VI) are differentiated based on size, reticulation pattern, and exine structure. The described pollen grains show all the essential features of angiosperm pollen. However, considering the lack of a continuous record throughout the lower part of the Mesozoic and the comparison with the oldest Cretaceous finds we suggest an affinity to an angiosperm stem group. Together with the previously published records from the Middle Triassic of the Barents Sea area the angiosperm-like pollen grains reflect a considerable diversity of the parent plants during the Middle Triassic. Sedimentological evidence and associated palynofloras also suggest a remarkable ecological range for these plants. Associated with these grains we found pollen comparable to the genus Afropollis. Representatives of this genus are commonly recorded in Lower Cretaceous sediments of low latitudes, but until now had no record from the lower part of the Mesozoic.

Concepts: Flowering plant, Pollen, Cretaceous, Triassic, Psittacosaurus, Early Cretaceous, Spinosauridae, Anisian

60

The discovery of large, complex, internal canals within the rostra of fossil reptiles has been linked with an enhanced tactile function utilised in an aquatic context, so far in pliosaurids, the Cretaceous theropod Spinosaurus, and the related spinosaurid Baryonyx. Here, we report the presence of a complex network of large, laterally situated, anastomosing channels, discovered via micro-focus computed tomography (μCT), in the premaxilla and maxilla of Neovenator, a mid-sized allosauroid theropod from the Early Cretaceous of the UK. We identify these channels as neurovascular canals, that include parts of the trigeminal nerve; many branches of this complex terminate on the external surfaces of the premaxilla and maxilla where they are associated with foramina. Neovenator is universally regarded as a ‘typical’ terrestrial, predatory theropod, and there are no indications that it was aquatic, amphibious, or unusual with respect to the ecology or behaviour predicted for allosauroids. Accordingly, we propose that enlarged neurovascular facial canals shouldn’t be used to exclusively support a model of aquatic foraging in theropods and argue instead that an enhanced degree of facial sensitivity may have been linked with any number of alternative behavioural adaptations, among them defleshing behaviour, nest selection/maintenance or social interaction.

Concepts: Cretaceous, Dinosaur, Theropoda, Early Cretaceous, Carcharodontosauridae, Spinosauridae, Spinosaurus, Neovenatoridae

57

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.

Concepts: Dinosaur, Theropoda, Tyrannosaurus, Ceratosauria, Giganotosaurus, Carcharodontosaurus, Spinosauridae, Spinosaurus

30

Six quadrate bones, of which two almost certainly come from the Kem Kem beds (Cenomanian, Upper Cretaceous) of south-eastern Morocco, are determined to be from juvenile and adult individuals of Spinosaurinae based on phylogenetic, geometric morphometric, and phylogenetic morphometric analyses. Their morphology indicates two morphotypes evidencing the presence of two spinosaurine taxa ascribed to Spinosaurus aegyptiacus and? Sigilmassasaurus brevicollis in the Cenomanian of North Africa, casting doubt on the accuracy of some recent skeletal reconstructions which may be based on elements from several distinct species. Morphofunctional analysis of the mandibular articulation of the quadrate has shown that the jaw mechanics was peculiar in Spinosauridae. In mature spinosaurids, the posterior parts of the two mandibular rami displaced laterally when the jaw was depressed due to a lateromedially oriented intercondylar sulcus of the quadrate. Such lateral movement of the mandibular ramus was possible due to a movable mandibular symphysis in spinosaurids, allowing the pharynx to be widened. Similar jaw mechanics also occur in some pterosaurs and living pelecanids which are both adapted to capture and swallow large prey items. Spinosauridae, which were engaged, at least partially, in a piscivorous lifestyle, were able to consume large fish and may have occasionally fed on other prey such as pterosaurs and juvenile dinosaurs.

Concepts: Mandible, Cretaceous, Dinosaur, Theropoda, Spinosauridae, Spinosaurus, Irritator, Sigilmassasaurus

26

A new species of pterosaur, Maaradactylus kellneri gen. nov., sp. nov. (Archosauria: Pterosauria) from the Romualdo Formation (Aptian/Albian), is herein described. The specimen (MPSC R 2357) was found at Sítio São Gonçalo, Santana do Cariri city (State of Ceará, northeast Brazil) and consists of the skull, atlas and axis, and represents one of the largest skulls of the Anhangueridae from the Araripe Basin described. The autapomorphies of the new pterosaur include the following characters: a premaxillary sagittal crest that is relatively long and high, beginning at the anterior part of the skull (rostrum) and extending to the 22nd pair of alveoli, not covering the nasoantorbital fenestra or the choanaes, and also the presence of 35 pairs of alveoli; smooth palatal ridge, which starts on the 5th pair of alveoli and ends on the 13th pair; palate is convex shaped in the anterior region; choanae not extending laterally; small and convex palatal elevation; the 5th, 6th and 7th alveoli smaller than the 4th and 8th; the alveoli decreasing in size from the 9th to the 12th and increasing from the 13th to 18th, and from the 18th to the 35th they are arranged in triplets. Furthermore, the lateral surface of the premaxillary crest shows grooves and tridimensional structures that may have housed blood vessels.

Concepts: Monotonic function, Skull, Cretaceous, Pterosaur, Pterodactyloidea, Northeast Region, Brazil, Skull and Bones, Spinosauridae

24

Fossil sites from Brazil have yielded specimens of spinosaurid theropods, among which the most informative include the cranial remains of Irritator, Angaturama, and Oxalaia. In this work some of their craniodental features are reinterpreted, providing new data for taxonomic and evolutionary issues concerning this particular clade of dinosaurs. The mesial-most tooth of the left maxilla of the holotype of Irritator is regarded as representing the third tooth position, which is also preserved in the holotype of Angaturama. Thus, both specimens cannot belong to the same individual, contrary to previous assumptions, although they could have been the same taxon. In addition, the position of the external nares of Irritator is more comparable to those of Baryonyx and Suchomimus instead of other spinosaurine spinosaurids. In fact, with regards to some craniodental features, Brazilian taxa represent intermediate conditions between Baryonychinae and Spinosaurinae. Such a scenario is corroborated by our cladistic results, which also leave open the possibility of the former subfamily being non-monophyletic. Furthermore, the differences between spinosaurids regarding the position and size of the external nares might be related to distinct feeding habits and degrees of reliance on olfaction. Other issues concerning the evolution and taxonomy of Spinosauridae require descriptions of additional material for their clarification.

Concepts: Cladistics, Theropoda, Spinosauridae, Spinosaurus, Irritator, Spinosauroidea, Baryonyx, Spinosaurids

19

A number of extant and extinct archosaurs evolved an elongate, narrow rostrum. This longirostrine condition has been associated with a diet comprising a higher proportion of fish and smaller prey items compared to taxa with broader, more robust snouts. The evolution of longirostrine morphology and a bulbous anterior rosette of premaxillary teeth also occurs in the spinosaurid theropod dinosaurs, leading to suggestions that at least some members of this clade also had a diet comprising a notable proportion of fish or other small vertebrates. Here we compare the rostral biomechanics of the spinosaurs Baryonyx walkeri and Spinosaurus c.f. S. aegyptiacus to three extant crocodilians: two longistrine taxa, the African slender-snouted crocodile Mecistops cataphractus and the Indian gharial Gavialis gangeticus; and the American alligator Alligator mississippiensis. Using computed tomography (CT) data, the second moments of area and moments of inertia at successive transverse slices along the rostrum were calculated for each of the species. Size-independent results tested the biomechanical benefits of material distribution within the rostra. The two spinosaur rostra were both digitally reconstructed from CT data and compared against all three crocodilians. Results show that African slender-snouted crocodile skulls are more resistant to bending than an equivalent sized gharial. The alligator has the highest resistances to bending and torsion of the crocodiles for its size and greater than that of the spinosaurs. The spinosaur rostra possess similar resistance to bending and torsion despite their different morphologies. When size is accounted for, B. walkeri performs mechanically differently from the gharial, contradicting previous studies whereas Spinosaurus does not. Biomechanical data support known feeding ecology for both African slender-snouted crocodile and alligator, and suggest that the spinosaurs were not obligate piscivores with diet being determined by individual animal size.

Concepts: Reptile, Crocodilia, Crocodile, Theropoda, Alligator, Spinosauridae, Gharial, Spinosaurus

16

The view of spinosaurs as dinosaurs of semi-aquatic habits and strongly associated with marginal and coastal habitats are deeply rooted in both scientific and popular knowledge, but it was never statistically tested. Inspired by a previous analysis of other dinosaur clades and major paleoenvironmental categories, here we present our own statistical evaluation of the association between coastal and terrestrial paleoenvironments and spinosaurids, along with other two theropod taxa: abelisaurids and carcharodontosaurids. We also included a taphonomic perspective and classified the occurrences in categories related to potential biases in order to better address our interpretations. Our main results can be summarized as follows: 1) the taxon with the largest amount of statistical evidence showing it positively associated to coastal paleoenvironments is Spinosauridae; 2) abelisaurids and carcharodontosaurids had more statistical evidence showing them positively associated with terrestrial paleoenvironments; 3) it is likely that spinosaurids also occupied spatially inland areas in a way somehow comparable at least to carcharodontosaurids; 4) abelisaurids may have been more common than the other two taxa in inland habitats.

Concepts: Scientific method, Phylogenetic nomenclature, Cladistics, Dinosaur, Theropoda, Spinosauridae, Spinosaurus, Spinosauroidea

15

Sigilmassasaurus brevicollis is an enigmatic theropod dinosaur from the early Late Cretaceous (Cenomanian) of Morocco, originally based on a few isolated cervical vertebrae. Ever since its original description, both its taxonomic validity and systematic affinities were contentious. Originally considered to represent its own family, Sigilmassasauridae, the genus has variously been suggested to represent a carcharodontosaurid, an ornithischian, and, more recently, a spinosaurid. Here we describe new remains referrable to this taxon and re-evaluate its taxonomic status and systematic affinities. Based on the new remains, a re-evaluation of the original materials, and comparisons with other spinosaurids, the holotype of Sigilmassasaurus brevicollis is identified as an anterior dorsal, rather than a cervical vertebra, and differences between elements referred to this taxon can be explained by different positions of the elements in question within the vertebral column. Many characters used previously to diagnose the genus and species are found to be more widespread among basal tetanurans, and specifically spinosaurids. However, the taxon shows several autapomorphies that support its validity, including the presence of a strongly rugose, ventrally offset triangular platform that is confluent with a ventral keel anteriorly in the mid-cervical vertebral centra and a strongly reduced lateral neural arch lamination, with no or an incomplete distinction between anterior and posterior centrodiapophyseal laminae in the posterior cervical and anterior dorsal vertebrae. We argue furthermore that Spinosaurus maroccanus, also described on the basis of isolated cervical vertebrae from the same stratigraphic unit and in the same paper as Sigilmassasaurus brevicollis, is a subjective synonym of the latter. Both a detailed comparison of this taxon with other theropods and a formal phylogenetic analysis support spinosaurid affintities for Sigilmassasaurus. However, we reject the recently proposed synonymy of both Spinosaurus maroccanus and Sigilmassasurus brevicollis with Spinosaurus aegyptiacus from the Cenomanian of Egypt, as there are clear differences between the vertebrae of these taxa, and they do not share any derived character that is not found in other spinosaurids. Together with a comparison with other spinosaurid vertebral material from the Kem Kem, this suggests that more than one taxon of spinosaurid was present in the Kem Kem assemblage of Morocco, so the referral of non-overlapping material from this unit to a single taxon should be regarded with caution.

Concepts: Vertebral column, Vertebra, Cervical vertebrae, Dinosaur, Theropoda, Carcharodontosaurus, Spinosauridae, Spinosaurus