Albert G. Sellés

3-D Modelling of Megaloolithid Clutches: Insights about Nest Construction and Dinosaur Behaviour

Background Megaloolithid eggs have long been associated with sauropod dinosaurs. Despite their extensive and worldwide fossil record, interpretations of egg size and shape, clutch morphology, and incubation strategy vary. The Pinyes locality in the Upper Cretaceous Tremp Formation in the southern Pyrenees, Catalonia provides new information for addressing these issues. Nine horizons containing Megaloolithus siruguei clutches are exposed near the village of Coll de Nargó. Tectonic deformation in the study area strongly influenced egg size and shape, which could potentially lead to misinterpretation of reproductive biology if 2D and 3D maps are not corrected for bed dip that results from tectonism. Methodology/Findings Detailed taphonomic study and three-dimensional modelling of fossil eggs show that intact M. siruguei clutches contained 20–28 eggs, which is substantially larger than commonly reported from Europe and India. Linear and grouped eggs occur in three superimposed levels and form an asymmetric, elongate, bowl-shaped profile in lateral view. Computed tomography data support previous interpretations that the eggs hatched within the substrate. Megaloolithid clutch sizes reported from other European and Indian localities are typically less than 15 eggs; however, these clutches often include linear or grouped eggs that resemble those of the larger Pinyes clutches and may reflect preservation of incomplete clutches. Conclusions/Significance We propose that 25 eggs represent a typical megaloolithid clutch size and smaller egg clusters that display linear or grouped egg arrangements reported at Pinyes and other localities may represent eroded remnants of larger clutches. The similarity of megaloolithid clutch morphology from localities worldwide strongly suggests common reproductive behaviour. The distinct clutch geometry at Pinyes and other localities likely resulted from the asymmetrical, inclined, and laterally compressed titanosaur pes unguals of the female, using the hind foot for scratch-digging during nest excavation.

Allodaposuchus palustris sp. nov. from the Upper Cretaceous of Fumanya (South-Eastern Pyrenees, Iberian Peninsula): Systematics, Palaeoecology and Palaeobiogeography of the Enigmatic Allodaposuchian Crocodylians

The controversial European genus Allodaposuchus is currently composed of two species (A. precedens, A. subjuniperus) and it has been traditionally considered a basal eusuchian clade of crocodylomorphs. In the present work, the new species A. palustris is erected on the base of cranial and postcranial remains from the lower Maastrichtian of the southern Pyrenees. Phylogenetic analyses here including both cranial and postcranial data support the hypothesis that Allodaposuchus is included within Crocodylia. The studied specimen suggests little change in postcranial skeleton along the evolutionary history of crocodylians, except for some bone elements such as the axis, the first caudal vertebra and the ilium. The specimen was found in an organic mudstone corresponding to a coastal wetland environment. Thus, A. palustris from Fumanya is the first Allodaposuchus reported in lacustrine-palustrine settings that expand the ecological range for this genus. The S-DIVA palaeobiogeographic reconstruction of ancestral area suggests that early members of Crocodylia rapidly widespread for the Northern Hemisphere landmasses no later than the Campanian, leading the apparition of endemic groups. In that way “Allodaposuchia” represents an endemic European clade probably originated in the Ibero-Armorican domain in the late Campanian and dispersed by the Southern European archipelago prior to the early Maastrichtian.

A re-evaluation of aff. Megaloolithidae eggshell fragments from the uppermost Cretaceous of the Pyrenees and implications for crocodylomorph eggshell structure

The Upper Cretaceous outcrops of the Pyrenees yield one of the most extensive and continuous records of paleoological remains anywhere in the world. Most of eggs and eggshells have been referred to the oofamily Megaloolithidae. In this study, we present a revision of eggshell fragments from the Blasi 2 locality, lattermost Maastrichtian in age, previously assigned to aff. Megaloolithidae. The presence of a blocky extinction pattern and basal knobs supports a crocodilian affinity of these materials. We classify them as Krokolithidae indet. Three structural layers can be recognised in the Blasi 2 eggshells, a feature that is shared with other recent eggshells (e.g. Crocodylus porosus and Crocodylus niloticus) and fossil crocodylomorph eggshells (Krokolitheswilsoni), which were previously described as single layered. The new proposed affinity of the Blasi 2 eggshells reduces the Megaloolithidae oodiversity of the last few million years of the Cretaceous in the Pyrenees to only two valid ootaxa, Megaloolithusmamillare and Megaloolithusbaghensis. The lack of more complete material precludes the erection of new ootaxa based on the Blasi 2 material.

A new species of Allodaposuchus (Eusuchia, Crocodylia) from the Maastrichtian (Late Cretaceous) of Spain: phylogenetic and paleobiological implications

Background. The Late Cretaceous is a keystone period to understand the origin and early radiation of Crocodylia, the group containing all extant lineages of crocodilians. Among the taxa described from the latest Cretaceous of Europe, the genus Allodaposuchus is one of the most common but also one of the most controversial. However, because of its fragmentary record, several issues regarding its phylogenetic emplacement and its ecology remain unsolved or unknown. The discovery of a single specimen attributed to Allodaposuchus, represented by both cranial and postcranial remains, from the Casa Fabà site (Tremp Basin, NE Spain) in the lower red unit of the Tremp Fm. (early Maastrichtian, Late Cretaceous) offers a unique opportunity to deepen in the phylogenetic relationships of the group and its ecological features. Methods. The specimen is described in detail, and CT scan of the skull is performed in order to study the endocranial morphology as well as paratympanic sinuses configuration. In addition, myological and phylogenetic analyses are also carried out on the specimen for to shed light in ecological and phylogenetic issues, respectively. Results. The specimen described herein represents a new species, Allodaposuchus hulki sp. nov., closely related to the Romanian A. precedens. The CT scan of the skull revealed an unexpected paratympanic sinuses configuration. Allosaposuchus hulki exhibits an “anterodorsal tympanic sinus” not observed in any other extant or extinct crocodilian. The caudal tympanic recesses are extremely enlarged, and the expanded quadratic sinus seems to be connected to the middle-ear channel. Phylogenetic analyses confirm the emplacement of the informal taxonomic group ‘Allodaposuchia’ at the base of Crocodylia, being considered the sister group of Borealosuchus and Planocraniidae. Discussion. Although this is a preliminary hypothesis, the unique paratympanic configuration displayed by A. hulki suggests that it could possess a high-specialized auditory system. Further, the large cranial cavities could help to reduce the weight of the cranium. Concerning the postcranial skeleton, Allodaposuchus hulki shows massive and robust vertebrae and forelimb bones, suggesting it could have a bulky body. The myological study performed on the anterior limb elements supports this interpretation. In addition, several bone and muscular features seem to point at a semi-erected position of the forelimbs during terrestrial locomotion. Taking all the above results into consideration, it seems plausible to suggest that A. hulki could conduct large incursions out of the water and have a semi-terrestrial lifestyle.

Spheroolithus europaeus, oosp. nov. (Late Maastrichtian, Catalonia), the Youngest Oological Record of Hadrosauroids in Eurasia

Hadrosauroids were the most abundant dinosaurs in the European archipelago during the Maastrichtian, being present in the Ibero-Armorican (current areas of Spain, Portugal, and France), the Adriatic-Dinaric Platform (northeast Italy), and the Tisia-Dacia Block (Romania) islands. Five valid species of hadrosauroids have been described from these areas to date: Pararhabdodon isonensis (Casanovas-Cladellas et al., 1993), Arenysaurus ardevoli (Pereda-Suberbiola et al., 2009b), Blasisaurus canudoi (Cruzado-Caballero et al., 2010a), Tethyshadros insularis (Dalla Vecchia, 2009), Telmatosaurus transsylvanicus (Nopcsa, 1899), and Canardia garonnensis (Prieto-Márquez et al., 2013). In addition, some indeterminate forms, comprising basal hadrosauroids, hadrosaurines, and lambeosaurines, have also been reported (Pereda-Suberbiola et al., 2009a; CruzadoCaballero et al., 2010b). Despite the abundance of osteological and ichnological remains of European hadrosauroids (Dalla Vecchia, 2006; Riera et al., 2009; Vila et al., 2012), oological evidence remains unreported. In the present study, we describe a new ootaxon attributed to hadrosauroid dinosaurs, Spheroolithus europaeus, oosp. nov., representing the first occurrence of this fossil egg type in the Late Cretaceous of Europe and the youngest evidence in Eurasia. The affiliation of this new oospecies with hadrosauroids is based on abundant eggshell fragments collected from the Tremp Formation (southern Pyrenees, Catalonia; Fig. 1). Institutional Abbreviation—IPS, Institut Català de Paleontologia Miquel Crusafont, Barcelona, Spain.

The youngest sauropod evidence in Europe

Titanosaur sauropods constituted one of the major dinosaur faunal components in Europe along the latest Cretaceous. Although they were abundant in the late Campanian–early Maastrichtian, titanosaurs apparently demised in terms of diversity and abundance in the late Maastrichtian Ibero-Armorican domain. Thus, any finding in this crucial period provides new clues to understand the biodiversity trend of the group prior to the K–Pg boundary. Here, we report the occurrence of a vertebral element attributed to a titanosaur in fluvial deposits dated as uppermost Maastrichtian (C29r chrone). The specimen corresponds to the right posterior part of a posterior cervical vertebra. CT-scan and histological analyses revealed high pneumatisation with laminar structures composed of compact bone, strongly remodelled with at least three generations of secondary osteons. These evidences suggest a relatively advanced age for the individual at death. Given that the partial vertebra was recovered from the uppermost portion of the late Maastrichtian, it might represent the youngest evidence for titanosaur sauropods in Europe discovered so far. Further, alongside to other remains (i.e. eggshells and footprints), the studied partial vertebra provides evidence for the persistence of this taxonomic group at the very end of the Cretaceous in southwestern Europe.

Reassessing the endemic European Upper Cretaceous dinosaur egg Cairanoolithus

The fossil record of dinosaur eggs and eggshells from the uppermost Cretaceous strata of south-western Europe is composed of both worldwide-distributed and endemic egg types. In this study, we are reviewing the enigmatic European oogenus Cairanoolithus, which after analysing abundant material from classic and new localities it is reassigned to a new oofamily (Cairanoolithidae oofam. nov.) in the light of the unique combination of structural characters. The new oofamily includes one oogenus and two oospecies. Cairanoolithid eggs share several features with other ornithopod egg types indeed. Furthermore, our phylogenetic analysis places Cairanoolithus as the sister ootaxon of the ornithopod ootaxa, being considered the most basal ornithischian egg type known so far. Although neither embryonic nor bones remains are known in association with cairanoolithid eggs so far, several taxonomic attributions have been proposed for this egg type over time. On the basis of microstructural features, phylogenetic results and anatomical constrains, we discuss in this paper previous taxonomic attributions and provide new evidence for suggesting a plausible nodosaurid affinity.

Evidence of Reproductive Stress in Titanosaurian Sauropods Triggered by an Increase in Ecological Competition

The occurrence of dinosaur pathologic eggs in the Late Cretaceous of Europe is well known, but their origin remains unclear. Here we expose the results of a detailed sampling of the conspicuous fossil record of Late Cretaceous titanosaurian eggs (oogenus Megaloolithius) from several southwestern Europe basins. After examining more than 450 samples, we observed a remarkable and statistically supported occurrence of multiple pathologic eggs in a relatively short stratigraphic range at the end of the early Maastrichtian, circa 71-70 Ma. All pathologic specimens exhibit multi-layered eggshell condition, a characteristic related to dystocia, or egg retention within the female uterus for an abnormal prolonged period of time. After exploring various scenarios, the occurrence of pathologic eggs is strongly correlated with an intense dinosaur faunal replacement that occurred during the early Maastrichtian in the Ibero-Armorican Island. Given that inter-species competiveness is proved to produce major affects in ecological communities, our results suggest that pathologies in the eggs of European titanosaurians could be a consequence of an increase in reproductive stress triggered by direct ecological competition between different dinosaurs. Thus, the present study provides a new perspective of how dinosaurs might have been affected by ecological/environmental disturbance.