Bernat Vila

High-resolution LiDAR and photogrammetric survey of the Fumanya dinosaur tracksites (Catalonia): implications for the conservation and interpretation of geological heritage sites

Increasing political and social awareness of the importance of protecting the geological heritage is compelling geoscientists to consider new methods for reconciling conservation and exploration of their research sites. Terrestrial Light Detection And Range (LiDAR) imaging is an accurate method of collecting 3D spatial data that has so far been under-utilized in the geological sciences. This aim of this paper is to assess the value of integrated LiDAR and photogrammetric imaging as a tool for synchronizing scientific exploration with conservation of geological heritage sites.   Fumanya (Catalonia) is one of the most important Cretaceous tracksites in Europe, but the nature of exposure of the track-bearing surface has hindered quantitative documentation of the ichnites. Using integrated Light Detection And Range (LiDAR) imaging and photogrammetry it has been possible to construct high-resolution Digital Outcrop Models (DOM) of the tracksites. Photo-textured DOMs are a powerful visualization tool and function as fully 3D interactive databases that preserve information about the site that would otherwise be lost to erosion. LiDAR-derived DOMs have the potential to contribute profoundly to future geoconservation projects, particularly as a tool for documenting and monitoring heritage sites and promoting education and tourism. LiDAR scanning also provides sufficient resolution to perform robust quantitative analysis of dinosaur tracks.

Comparison of water vapor conductance in a titanosaur egg from the Upper Cretaceous of Argentina and a Megaloolithus siruguei egg from Spain

Abstract We calculated water vapor conductance (a product of eggshell porosity) from the first definitively identified sauropod egg (Megaloolithus patagonicus) from the Auca Mahuevo locality in Argentina. We then compared the results with those from M. siruguei (an egg type long associated with sauropod dinosaurs) from the Pinyes locality in Spain. The 14-cm Auca Mahuevo egg has a thinner eggshell and 47 times fewer pores than the 22-cm M. siruguei specimen. The resulting water vapor conductance (GH2O) of the titanosaur and M. siruguei eggs is 341 and 3979 mg H2O day−1 Torr−1, respectively; these values are two and ten times greater than in avian eggs of comparable size, but lower than in eggs of most modern reptiles. Clutches from Auca Mahuevo typically contain 20–40 eggs; in contrast, M. siruguei clutches from the Pinyes site average nine eggs. The GH2O of M. siruguei exceeds that of the Argentine egg by an order of magnitude, supporting previous inferences of egg burial. The GH2O of the Argentine titanosaur egg closely approximates that of Troodon and some oviraptorid eggs, previously calculated as equal to or two times greater than, respectively, the GH2O of avian eggs of similar size. Higher embryonic growth rates (relative to modern reptiles), especially in some dinosaurs with large clutch mass, may have required incubation in a more open environment, where water conservation represented a more critical factor than in a buried clutch. The lower GH2O calculated for the two megaloolithid eggs is consistent with previous interpretations of nesting mode that are based on site taphonomy and nesting traces. This study indicates that at least some dinosaurs did not fully bury their eggs.

Discriminating between Medium-Sized Tridactyl Trackmakers: Tracking Ornithopod Tracks in the Base of the Cretaceous (Berriasian, Spain)

Background Recent work on the Jurassic-Cretaceous transition of the Iberian Range (Spain) has opened a new window onto the interpretation of the trackmakers of some medium-sized tridactyl tracks. The ichnotaxon Therangospodus oncalensis has been described in the Huérteles Formation (Berriasian) and is one of the classical tracks from the area assigned to medium-sized theropods. Methodology/Principal Findings A review of the type locality of Therangospodus oncalensis (Fuentesalvo tracksite) and other tracksites from the Huérteles Formation (Berriasian) has yielded new information on the morphology, gait and trackmaker identity of the aforementioned ichnospecies. The new data suggest that the trackmaker is an ornithopod rather than a theropod on the basis of the length/width ratio, the anterior triangle length-width ratio, the short steps, the round to quadrangular heel pad impression and the probable manus impressions. Conclusions/Significance T. oncalensis shows similarities with various tracks from the Berriasian of Europe assigned to Iguanodontipus. The ichnotaxonomical status of this ichnospecies is here considered as Iguanodontipus? oncalensis due to the current state of knowledge of the ichnotaxonomy of medium-sized ornithopod tracks. This reassessment of I? oncalensis also has two significant implications for the palaeoecology of the faunas during the deposition of the Huérteles Formation: 1- the high number and percentage of theropod tracks would be lower than previous papers have suggested. 2- the gregarious behaviour described in the type locality (Fuentesalvo) would be among ornithopods instead of theropods.

The Latest Succession of Dinosaur Tracksites in Europe: Hadrosaur Ichnology, Track Production and Palaeoenvironments

A comprehensive review and study of the rich dinosaur track record of the Tremp Formation in the southern Pyrenees of Spain (Southwestern Europe) shows a unique succession of footprint localities prior to the end-Cretaceous mass extinction event. A description of some 30 new tracksites and data on sedimentary environments, track occurrence and preservation, ichnology and chronostratigraphy are provided. These new track localities represent various facies types within a diverse set of fluvial environments. The footprint discoveries mostly represent hadrosaurian and, less abundantly, to sauropod dinosaurs. The hadrosaur tracks are significantly smaller in size than, but morphologically similar to, those of North America and Asia and are attributable to the ichnogenus Hadrosauropodus. The track succession, with more than 40 distinct track levels, indicates that hadrosaur footprints in the Ibero-Armorican region occur predominantly in the late Maaastrichtian (at least above the early Maastrichtian–late Maastrichtian boundary). The highest abundance is found noticeably found in the late Maastrichtian, with tracks occurring in the C29r magnetochron, within about the latest 300,000 years of the Cretaceous.

Manus-only titanosaurid trackway from Fumanya (Maastrichtian, Pyrenees): further evidence for an underprint origin

The origin of manus-only and manus dominated sauropod trackways has been a matter of intense debate since two hyphothesis exist: (a) manus-only and manus-dominated trackways result from a ‘swimming’ sauropod, and (b) they result from a selective underprint phenomenon that only leaves the manus recorded. Several new sauropod trackways are reported in the Fumanya tracksite area (Maastrichtian), in SE Pyrenees, where both tracks and undertracks are found on the same stratigraphic bedding surface. In one of the trackways, footprint morphology together with the trackway pattern displays a clear succession of manus-only impressions attributed to a sauropod dinosaur in a walking gait. The ichnological comparison between the manus-only trackway with the other complete trackway (manus-pes) display an identical distribution of the manus pattern. This fact clearly points towards an underprint phenomenon as the origin for manus-only trackways, since it is rather unlikely that the same pattern would completely match different locomotion behaviours such as walking and swimming. Therefore, we suggest an interpretation based on the differential loading between the hindfoot and the forefoot on an upper stratigraphic track-level, for the studied manus-only trackway. □ Fumanya tracksite, manus-only trackways, titanosaurs, trackway pattern, underprint, Upper Cretaceous.

Manus track preservation bias as a key factor for assessing trackmaker identity and quadrupedalism in basal ornithopods.

Background The Las Cerradicas site (Tithonian–Berriasian), Teruel, Spain, preserves at least seventeen dinosaur trackways, some of them formerly attributed to quadrupedal ornithopods, sauropods and theropods. The exposure of new track evidence allows a more detailed interpretation of the controversial tridactyl trackways as well as the modes of locomotion and taxonomic affinities of the trackmakers. Methodology/Principal Findings Detailed stratigraphic analysis reveals four different levels where footprints have been preserved in different modes. Within the tridactyl trackways, manus tracks are mainly present in a specific horizon relative to surface tracks. The presence of manus tracks is interpreted as evidence of an ornithopod trackmaker. Cross-sections produced from photogrammetric digital models show different depths of the pes and manus, suggesting covariance in loading between the forelimbs and the hindlimbs. Conclusions/Significance Several features (digital pads, length/width ratio, claw marks) of some ornithopod pes tracks from Las Cerradicas are reminiscent of theropod pedal morphology. This morphological convergence, combined with the shallow nature of the manus tracks, which reduces preservation potential, opens a new window into the interpretation of these tridactyl tracks. Thus, trackmaker assignations during the Jurassic–Cretaceous interval of purported theropod trackways may potentially represent ornithopods. Moreover, the Las Cerradicas trackways are further evidence for quadrupedalism among some basal small- to medium-sized ornithopods from this time interval.

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.

Intra-trackway morphological variations due to substrate consistency: the El Frontal dinosaur tracksite (lower Cretaceous, Spain).

An ichnological and sedimentological study of the El Frontal dinosaur tracksite (Early Cretaceous, Cameros basin, Soria, Spain) highlights the pronounced intra-trackway variation found in track morphologies of four theropod trackways. Photogrammetric 3D digital models revealed various and distinct intra-trackway morphotypes, which reflect changes in footprint parameters such as the pace length, the track length, depth, and height of displacement rims. Sedimentological analyses suggest that the original substrate was non-homogenous due to lateral changes in adjoining microfacies. Multidata analyses indicate that morphological differences in these deep and shallow tracks represent a part of a continuum of track morphologies and geometries produced by a gradient of substrate consistencies across the site. This implies that the large range of track morphologies at this site resulted from similar trackmakers crossing variable facies. The trackways at the El Frontal site present an exemplary case of how track morphology, and consequently potential ichnotaxa, can vary, even when produced by a single trackmaker.

Dynamic similarity in titanosaur sauropods: ichnological evidence from the Fumanya dinosaur tracksite (southern Pyrenees).

The study of a small sauropod trackway from the Late Cretaceous Fumanya tracksite (southern Pyrenees, Catalonia) and further comparisons with larger trackways from the same locality suggest a causative relationship between gait, gauge, and body proportions of the respective titanosaur trackmakers. This analysis, conducted in the context of scaling predictions and using geometric similarity and dynamic similarity hypotheses, reveals similar Froude numbers and relative stride lengths for both small and large trackmakers from Fumanya. Evidence for geometric similarity in these trackways suggests that titanosaurs of different sizes moved in a dynamically similar way, probably using an amble gait. The wide gauge condition reported in trackways of small and large titanosaurs implies that they possessed similar body (trunk and limbs) proportions despite large differences in body size. These results strengthen the hypothesis that titanosaurs possessed a distinctive suite of anatomical characteristics that are well reflected in their tracks and trackways.