Cranial bones and atlas of titanosaurs (Dinosauria,Sauropoda) from Late Cretaceous (Bauru Group) of Uberaba,Minas Gerais State,Brazil

Isolated left prefrontal, left squamosal and atlas of titanosaur dinosaurs are described and compared. They come from the Late Cretaceous Serra da Galga Member of the Marília Formation at the Serra do Veadinho region, Peirópolis (Uberaba County, Minas Gerais State, Brazil). Due to the sparse cranial elements of titanosaurs already known from Brazil, these specimens are noticeable to be presented. In addition, the atlas vertebra is described for the first time for Brazilian titanosaurs. The morphology of the cranial bones closely resembles lithostratian titanosaurs, such as Rapetosaurus, rather than basal titanosaurs. The atlas is similar to that of other titanosaurs, suggesting that the anatomy of this element seems to be more conservative than other vertebral elements, in which vertebral laminae play an important rule in titanosaur taxonomy.     

Pachycephalosaurs (Dinosauria: Ornithischia) from the Upper Cretaceous (upper Campanian) of New Mexico: A reassessment of Stegoceras novomexicanum

Pachycephalosaurs, a group of ornithischian dinosaurs with distinctive cranial ornamentation and skull domes, underwent dramatic changes in cranial morphology during ontogeny. This has caused debate about whether some specimens belong to juveniles or adults, which impacts studies of pachycephalosaur phylogeny and evolution. One such debate concerns a small skull roof specimen from the Campanian (Upper Cretaceous) of New Mexico, NMMNH P-33898, which was originally described as an indeterminate juvenile but later regarded as a mature adult and erected as the holotype of a new small-bodied species, Stegoceras novomexicanum. We restudied NMMNH P-33898 using computed tomography scanning, morphometric and phylogenetic analyses, and comparisons to growth series of other pachycephalosaurs (Stegoceras validum, Pachycephalosaurus wyomingensis). We conclude that two purported paratype specimens of Stegoceras novomexicanum cannot be referred to the same taxon as the holotype, that the holotype and controversial paratypes all belong to immature specimens and not aberrant small-bodied adults, but that current evidence cannot clearly determine whether NMMNH P-33898 is a juvenile belonging to its own diagnostic species (S. novomexicanum) or is a juvenile of Stegoceras validum, Sphaerotholus goodwini, or another known taxon. We review the pachycephalosaur record of New Mexico and demonstrate that pachycephalosaurs were important components of dinosaur faunas in the southern part of Western North America during the ∼15 million years before the end-Cretaceous extinction, just as they were in roughly contemporaneous northern localities.     

First report of gastroliths in the Early Cretaceous basal bird Jeholornis

Seeds preserved in association with the holotype of Jeholornis prima provided the first direct evidence of granivory in any Mesozoic bird. Although this long boney-tailed bird also displays several morphological indicators correlated with herbivory such as reduced dentition and a deep mandible, Jeholornis has not been previously reported to possess a gastric mill. However, this feature is commonly linked to herbivory in theropod dinosaurs and present in at least one sympatric ornithuromorph and the basal pygostylian Sapeornis, which also preserve direct evidence of granivory. Here we describe gastrolith masses preserved in five specimens of Jeholornis. The cluster of gizzard stones is nearly identical in each specimen, consisting of a tightly associated mass of proportionately small stones. Three previously undescribed specimens preserving seeds are also identified. Unlike in Sapeornis and a previously described ornithuromorph, no specimen of Jeholornis preserves both seeds and gastroliths. This may be due to the fact that, unlike in other Early Cretaceous birds, the seeds preserved in specimens of Jeholornis are found in the abdomen, suggesting that Jeholornis may have resembled extant ratites in lacking an esophageal crop. Consistent differences in the preservation and morphology of the gastrolith mass between Jeholornis and other early birds hint at subtle variations in alimentary function among basal lineages. Differences in ingested seed morphology among taxa in turn suggest these functional variations may be at least partially related to differences in diet.     

Utility of crinoid columnals in palaeontology illustrated by a new species: Clare Shale Formation (Carboniferous), Doolin,County Clare,western Ireland

Well-preserved specimens, such as complete individuals, crowns and cups, are the common focus for crinoid systematic research. Yet the majority of specimens are disarticulated ossicles which are essentially ignored. The incompleteness of the fossil record is even more so when we ignore potential sources of data. A new species of crinoid comes from a monospecific assemblage from the Pennsylvanian (Upper Carboniferous) of western Ireland. All specimens are from a float block of the Clare Shale Formation (Bashkirian stage) at Fisherstreet Bay, Doolin, County Clare, western Ireland. Heloambocolumnus (col.) harperi gen. et sp. nov. has a pentagonocyclic, heteromorphic column; the small, central lumen is in a shallow, circular claustrum; the articulation is radial symplectial; the crenulae are slightly swollen and peg-like close to the circumference; nodals have rounded, unsculptured epifacets; nodal articular facets are sunken and in which narrow internodals are situated; and circlets of tubercles on epifacet surround priminternodals. These columnals are associated with robust, uniserial brachial ossicles. This crinoid is most likely a cladid.     

New cranial remains of titanosaurian sauropod dinosaurs from the Late Cretaceous of Fox-Amphoux-Métisson (Var,SE France)

The Fox-Amphoux Syncline of the Aix Basin (SE France) has yielded a diverse Late Cretaceous vertebrate assemblage, including several taxa of dinosaurs. Here, we report on cranial material of titanosaurian sauropods, which consist of a partial braincase, two fragmentary skull roofs, and nine teeth, from the Métisson locality (Var Department). The braincase differs from those found previously in Europe (i.e., Ampelosaurus atacis, Lirainosaurus astibiae, and an unnamed juvenile skull from Romania) as well as from other titanosaurian braincases in having a groove that extends along the ventral surface of the occipital condyle neck (this feature may be autapomorphic). One of the fragmentary skull roofs may belong to the same taxon, whereas the other suggests the presence of a second titanosaur at Métisson very close to Ampelosaurus. Two dental morphologies are present in the sample; one of them includes teeth of different sizes. We suggest that this could be accounted for by age differences within a single taxon, or be due to different positions in the tooth row. The presence of a new, still unnamed titanosaurian taxon in the Ibero-Armorican Island supports previous works indicating a high titanosaurian diversity during the Campanian-Maastrichtian in southern Europe.     

Paleoecological implications of two closely associated egg types from the Upper Cretaceous St. Mary River Formation,Montana

Two closely associated egg types occur at the same locality in the Upper Cretaceous (Maastrichtian) St. Mary River Formation in north central Montana. These specimens represent the first fossil eggs described from this formation. At least fifteen small ovoid eggs or egg portions are scattered through a 25 cm interval of rock. Five significantly larger, round eggs overlie these smaller eggs and are in close proximity to one another on a single bedding plane. The best preserved egg of the smaller size measures 36 mm × 62 mm and exhibits the prismatic, two-layered eggshell structure of a theropod egg. The dispersed distribution and inconsistent angles of these small eggs likely resulted from disturbance by subsequent nesting activity and/or possibly nest predation. At least twelve additional small prismatic eggs also occur at this site. We assign the small eggs as a new oogenus and oospecies, Tetonoolithus nelsoni, within the Prismatoolithidae. The large round eggs measure 130 mm in diameter and the eggshell displays substantial diagenetic alteration. These eggs likely belonged to a hadrosaur due to their similarity in egg size, shape, and eggshell thickness to Maiasaura eggs from the stratigraphically lower Two Medicine Formation. Eggs at different stratigraphic levels at this site indicate that conditions favorable to both dinosaur species persisted for an extended period of time. However, determining whether these dinosaurs occupied the nesting site at the same or different years remains beyond the resolution of the rock record.     

Taphonomy of a Baurusuchus (Crocodyliformes,Baurusuchidae) from the Adamantina Formation (Upper Cretaceous,Bauru Basin), Brazil: Implications for preservational modes,time resolution and paleoecology

Upper Cretaceous vertebrate accumulations from the Adamantina Formation are known due to their high taxonomic diversity. On the other hand, taphonomic analyses still are rare, limiting the understanding of processes related to the biostratinomic and fossildiagenetic histories of this lithostratigraphic unit. In 2005, fossils were collected from an outcrop located at Jales municipality, state of São Paulo, southeastern Brazil. From this outcrop, a well-preserved Baurusuchus was recovered, which displays a peculiar set of taphonomic signatures. This paper identifies and interprets taphonomic features of a specimen of Baurusuchus (Crocodyliformes, Baurusuchidae; UFRJ DG 418-R) from the Adamantina Formation (Upper Cretaceous of the Bauru Basin), in Jales municipality, state of São Paulo. Brief taphonomic comparisons with other specimens previously studied (crocodiles and dinosaurs) and a lithofaciological analysis of the outcrop were undertaken in order to enhance the knowledge of the stratigraphy and paleoenvironment and improve the time resolution for the Adamantina Formation in the region of Jales. Furthermore, paleoecological data were interpreted based on the taphonomic analysis.The fossil is composed of an articulated segment of vertebral column, ribs, part of the pelvic girdle and gastralia. There is no hydraulic equivalence between both the specimen of Baurusuchus and the size of quartz grain predominant in the fossiliferous layer, suggesting death in situ or short transport as a “water carcass”. Teeth marks identified on the pubes were assigned to a small/juvenile baurusuchid crocodyliform or a theropod dinosaur. The repositioning of some elements (ribs and dorsal osteoderms) is suggestive of mummification. Desiccation marks were observed and attributed to the stage 1 of weathering. These features suggest subaerial exposure of the carcass prior to burial, however, probably after the mummification. On the other hand, the subaerial exposure was short, because the individual was not fully disarticulated. Furthermore, the degrees of articulation and preservation of the specimen nullify the hypothesis of reworking.Lithofaciological and taphonomic analyses suggest a well-drained floodplain as the burial environment and an arid or semi-arid climate in the region of Jales outcrop. In addition, the taphonomic signatures seem to indicate a time resolution about 10⁰–10⁴ years for the layer where the crocodyliforms were collected, but it seems to have, within this time limit, temporal-mixing among terrestrial crocodiles and dinosaurs collected from the same layer, suggesting time-averaging in this outcrop.     

Calcium isotopes in fossil bones and teeth — Diagenetic versus biogenic origin

We present the first systematic study of Ca isotopes (δ^44/40Ca) in Late Triassic to Late Cretaceous dinosaur bones and teeth (enamel and dentin) from sympatric herbivorous and carnivorous dinosaurs. The samples derive from five different localities, and data from embedding sediments are also presented. Additional δ^44/40Ca in skeletal tissues from modern reptiles and birds (avian dinosaurs) were measured for comparison in order to examine whether the original Ca isotopic composition in dinosaur skeletal apatite was preserved or might have changed during the diagenesis and fossilization process.δ^44/40Ca of fossil skeletal tissues range from −1.62‰ (Tyrannosaurus rex enamel) to +1.08‰ (Brachiosaurus brancai bone), while values in modern archosaur bones and teeth range from −1.63‰ (caiman enamel) to −0.37‰ (ostrich bone). The average δ^44/40Ca of the three types of fossil skeletal tissue analyzed - bone, dentin and enamel - show some systematic differences: while δ^44/40Ca in bone exhibits the highest values, while δ^44/40Ca in enamel has the lowest values, and dentin δ^44/40Ca falls in between. Values of δ^44/40Ca in the remains of herbivorous dinosaurs (0.1-1.1‰) are generally higher than those of bones of modern mammalian herbivores (−2.6‰ to −0.8‰) and from modern herbivorous archosaurs, which exhibit intermediate δ^44/40Ca (−0.8‰ to −0.4‰). These systematic isotopic shifts may reflect physiological differences between dinosaurs, mammals and reptiles representing different taxonomic groups of vertebrates.Systematic offsets in skeletal apatite δ^44/40Ca between herbivorous and carnivorous dinosaurs are not obvious, indicating a lack of a clear-cut Trophic Level Effect (TLE) shift between herbivores and carnivores in dinosaurs. This observation can be explained if the carnivorous dinosaurs in this study fed mainly on soft tissues from their prey and did not ingest hard (calcified) tissue to much extent. The most striking indication that the primary δ^44/40Ca is actually preserved in most of the fossil teeth is a difference in δ^44/40Ca of about 0.35 ± 0.10‰ (1SD) between dentin and enamel, based upon 11 of 16 analyzed dentin-enamel pairs. This difference is close to that found in modern reptiles (0.28 ± 0.05‰), and strongly suggests that this tell-tale signature is a primary feature of the fossilized dinosaur material as well. Furthermore, simple mass balance calculations show that changes of the original δ^44/40Ca in bones and teeth by diagenetically-formed calcium-bearing minerals are either small or would require implausible high original δ^44/40Ca values in the skeletal apatite.     

Ornithomimidae (Dinosauria: Theropoda) from the Bissekty Formation (Upper Cretaceous: Turonian) of Uzbekistan

The stratigraphically oldest remains of ornithomimid theropod dinosaurs are known from the Cenomanian Khodzhakul Formation and the Turonian Bissekty Formation of Uzbekistan. The ornithomimid from the Bissekty Formation is documented by more than 800 isolated bones that represent much of the skeleton. It shows at least three unambiguous synapomorphies of Ornithomimidae: length of anterior cervical centra three to five times greater than transverse width; low and rounded fibular crest of tibia; metatarsal III pinched between metatarsals II and IV; and proximal end of metatarsal III not visible in anterior view. Phylogenetic analysis, based on a dataset with 568 morphological characters and including all known ornithomimosaurian taxa, places the ornithomimid from the Bissekty Formation near the base of the ornithomimid radiation, between Archaeornithomimus asiaticus and Sinornithomimus dongi.     

A saltasaurine titanosaur (Sauropoda: Titanosauriformes) from the Angostura Colorada Formation (upper Campanian,Cretaceous) of northwestern Patagonia,Argentina

Saltasaurine titanosaurs are characterized by their relatively small size compared to other sauropods, extreme postcranial pneumaticity, and dermal armour covering the body. This group has been reported in the Upper Cretaceous of the Lecho, Allen, and Anacleto formations of Argentina. We describe here a new saltasaurine specimen (MACN-Pv RN 233) from the Campanian of the Angostura Colorada Formation (Río Negro Province) that is represented by eight caudal vertebrae and six osteoderms. This specimen is described in detail and its phylogenetic relationships with the other three known saltasaurines, as well as its implications to the knowledge of caudal vertebra and osteoderm anatomy, are discussed. Our results place MACN-Pv RN 233 more closely related to Saltasaurus loricatus and Rocasaurus muniozi than to Neuquensaurus australis. MACN-Pv RN 233 possesses a combination of features that differ from other saltasaurines, but because of the fragmentary nature of the specimen we decided for the sake of taxonomic stability to not erect a new taxon. This specimen shows the first unambiguous evidence of chevron pneumatisation for a sauropodomorph, implying a broader osteological invasion of the diverticula from the abdominal air sac than previously thought for this group of dinosaurs. MACN-Pv RN 233 preserves two osteoderm morphotypes, one similar to those reported for Neuquensaurus australis and Saltasaurus loricatus. This new specimen expands the distribution of the group to a new geological unit and increases the dinosaur diversity known for the Angostura Colorada Formation.     

A juvenile Edmontosaurus from the late Maastrichtian (Cretaceous) of North America: Implications for ontogeny and phylogenetic inference in saurolophine dinosaurs

The anatomy of an articulated juvenile specimen of the saurolophine hadrosaurid dinosaur Edmontosaurus annectens, LACM 23504, is described in detail. This individual consists of a partial skull and nearly complete articulated postcranium, collected from upper Maastrichtian strata of the Hell Creek Formation of Montana, USA. This study aims to help understand the ontogenetic morphological changes occurring in the cranium and postcranium of saurolophine hadrosaurids using E. annectens as a case study. Emphasis is placed on those morphological attributes that are ontogenetically variable via comparative osteology between the juvenile and the available adult specimens. It is observed that much of the cranial ontogenetic variation relates to the elongation of the skull and mandible. In the postcranium, most of the ontogenetic variation concentrates in the pectoral and pelvic girdles and the stylopodia. Not all the identified patterns of ontogenetic variation may be generalized to all hadrosaurids. The impact of ontogenetic variation on phylogenetically informative characters of saurolophine hadrosaurids is evaluated. It is concluded that, at least for Edmontosaurus annectens (and perhaps other saurolophine hadrosaurids), most characters used in phylogenetic inference of these animals are not affected by ontogeny. Thus, juvenile specimens are still a source of substantial character data suitable for use in phylogenetic analyses of saurolophine relationships. Nevertheless, it is recommended that ontogenetically variable characters are left as missing data in a character–taxon matrix when only juvenile material is available for a given saurolophine taxon. Scoring those characters based solely on juveniles is likely to decrease the accuracy of the phylogenetic inference.     

Diversity of hadrosauroid dinosaurs from the Late Cretaceous Ibero-Armorican Island (European Archipelago) assessed from dentary morphology

A substantial part of the European record of hadrosauroid ornithopod dinosaurs comes from the Maastrichtian (Upper Cretaceous) strata of the south-central Pyrenees (northern Catalunya and Aragón, northeastern Spain). Yet, our understanding on the taxonomy and phylogenetic systematics of these animals in this region is hindered by the lack of complete specimens, paucity of diagnostic material, and poor preservation of the fossils. The Basturs Poble site (Lleida Province, northern Catalunya) has yielded over 900 bones attributed to a hadrosauroid monodominant population. The dentary is the most common, and one of the better-preserved elements found in this locality. Here, we evaluate the utility of dentary morphology as an indicator of hadrosauroid taxonomic diversity. We do so via principal component analysis, cluster analysis, and mean comparison tests of various morphometric data from the available sample of dentaries collected so far in the Basturs Poble locality and other Maastrichtian sites in northeastern Spain. Three different dentary morphotypes are recognized: two different hadrosauroids, one more derived than the other, and one lambeosaurine type. One of the hadrosauroid morphotypes corresponds to relatively small individuals that may represent insular dwarf species. Concerning Basturs Poble locality, our results suggest the coexistence of hadrosauroid and lambeosaurine dinosaurs during the late Maastrichtian on the Ibero-Armorican Island.     

North Africa's first stegosaur: Implications for Gondwanan thyreophoran dinosaur diversity

Eurypoda, the major radiation of armoured dinosaurs, comprises the ankylosaurs and their sister group, the stegosaurs. As the earliest-branching major clade of ornithischian dinosaurs, the evolutionary history of Eurypoda is significant for understanding both the palaeobiology of bird-hipped dinosaurs and the composition of middle Mesozoic ecosystems. Eurypodans were diverse and abundant throughout the Late Jurassic and Cretaceous in Laurasia; in contrast, their remains are extremely rare in Gondwana. Herein, we describe a new genus and species of stegosaur from the Middle Jurassic of Morocco, Adratiklit boulahfa. Adratiklit is the first eurypodan from north Africa and the oldest definitive stegosaur from anywhere in the world. The genus is more closely related to the European stegosaurs Dacentrurus and Miragaia than it is to the southern African taxa Kentrosaurus and Paranthodon. Statistically significant correlations between the number of dinosaur-bearing formations, dinosaur-bearing collections, and eurypodan occurrences in Gondwana indicates that their fossil record is biased by both geological and anthropogenic factors. Tantalizing but fragmentary remains and trackways suggest that eurypodan diversity in Gondwana may have been as rich as that of Laurasia, and the prospects for future discoveries of new genera across Gondwana are therefore very good.     

Current situation of the ichnological locality of São Domingos from the Corda Formation (Lower Cretaceous), northern Tocantins state,Brazil

In the 80's, Leonardi treated the presence of a vertebrate ichnological locality from the Barremian Corda Formation, Parnaíba Basin, on the left bank of the Tocantins river, near of the São Domingos town, Itaguatins, State of Tocantins, Brazil. Originally, the record was composed of at least seven in situ trackways, accounting for fifty six tracks. Since 2011, the Hydroelectric Power Plant do Estreito has begun to work, causing the development of a water reservoir 160 km upstream to the ichnological site, causing periodic and highly energetic floods over the footprints-bearing level and altering it. The imprints are poorly to moderate preserved, but it is possible to distinguish the general morphology and the spatial arrangement of the footprints. The specimens are represented by pes imprints, mostly circular to subcircular, with no digital and claw impressions. No distinguishable manus imprints are present. The trackways are relative narrow with respect to the size of the tracks, so they are considered into the Parabrontopodus-like category. The São Domingos tracks have been originally assigned to iguanodontid dinosaurs, and posteriorly related to a sauropodian origin. This idea is herein accepted, particularly to a basal sauropod, basal macronarians, or diplodocoids. Up to date, the tracks from the São Domingos locality are the only vertebrate fossil record from the Corda Formation, meaning an important contribution to the Cretaceous ichnofauna from South America.     

Crinoid columns as hard substrates: Salthill Quarry (Mississippian,lower Carboniferous), Clitheroe,Lancashire, UK

Salthill Quarry, Clitheroe, Lancashire (Mississippian, Visean) is a Site of Special Scientific Interest (SSSI), in part based on its diverse fauna of invertebrates, particularly echinoderms. A small collection of crinoid pluricolumnals are described herein, collected from muddy horizons in the Hodder Mudstone Formation to typify the diversity and frequency of their encrusting and boring fauna. These specimens are infested by a range of episkeletozoans, namely a single occurrence of Sutherlandia parasitica (Phillips), four occurrences of Cladochonus sp. and eight of Oichnus paraboloides Bromley. Two variants of the pit (boring or embedment or both) O. paraboloides are recognised: those in which a living crinoid showed a growth reaction to pit formation; and those that did not and which were presumably dead at the time of infestation. The epizoozoan tabulate coral Cladochonus sp. is also common, including specimens intergrown between and within the columnals. Sutherlandia parasitica is relatively uncommon; the only specimen likely infested a dead pluricolumnal on the sea floor. A comparative collection of pluricolumnals infested by Cladochonus beecheri Grabau from the Mississippian Borden Group of the Midwest, USA, showed superior preservation to the Clitheroe Cladochonus sp. Cladochonus beecheri in the Borden Group infested platycrinitid crinoid stems, an association not noted from Salthill Quarry. Similar suites of borings-episkeletozoans, from two other sites – the Visean of Feltrim, Co. Dublin, Ireland, and the Permian of Timor – suggest that these were persistent associations for much of the Late Palaeozoic.     

Skeletal remains of Tyrannosauroidea (Dinosauria: Theropoda) from the Bissekty Formation (Upper Cretaceous: Turonian) of Uzbekistan

There exists a major gap in the fossil record of tyrannosauroid theropod dinosaurs spanning the early part of the Late Cretaceous. We report on skeletal remains referable to Tyrannosauroidea indet. from the Turonian Bissekty Formation at Dzharakuduk in the central Kyzylkum Desert of Uzbekistan. Phylogenetic analysis of the hypodigm places the Bissekty tyrannosauroid as a non-tyrannosaurid tyrannosauroid more basal than the Campanian tyrannosauroids Appalachiosaurus and Bistahieversor from North America.     

The rudist bivalve Pironaea milovanovici Kühn, 1935, a multiple-fold Hippuritidae,from south-eastern Spain. Taxonomic implications. Pore and canal system constructional morphology

The study of the external and internal shell characters of Pironaea milovanovici Kühn, as well as of their ontogenetic and intra-specific variability, observed on abundant, well preserved, and matrix-free specimens from Tavernes de la Valldigna, Valencia province, Spain, revealed that other Pironaea species (e.g. P. bacevicensis Milovanović, P. petkovici Milovanović, Sladić-Trifunović, and Grubić, P. postdalmatinica Pamouktchiev, P. signumpraeverta Pamouktchiev, and Pironaea sp. ex gr. polystyla in Lupu), based on one or a very small number of specimens, are junior synonyms of P. milovanovici Kühn. The pore and canal system of the left valve is described in detail, based on differently weathered valves and differently oriented sections; it is constrained by the presence of secondary pillars in both valves. Being the pore and canal system and its relation with the secondary pillars relatively simple and well understood in P. milovanovici, by comparison, an interpretation of these structures in more complex multiple-fold Hippuritidae such as those of the Americas is hypothesized.     

Spheroolithid eggshells in the Lower Cretaceous of Europe. Implications for eggshell evolution in ornithischian dinosaurs

Early Cretaceous spheroolithid eggshells have always been extremely scarce, unlike their putative egg layers, ornithopod dinosaurs, which are ubiquitous in Early Cretaceous ecosystems. In this paper, we re-evaluate eggshells previously assigned to the oofamily Elongatoolithidae, an oofamily related to derived theropod dinosaurs (including maniraptorans and oviraptorids), and reassign them to the oofamily Spheroolithidae, on the basis of the lack of a squamatic ultrastructure, and the presence of a prolatocanaliculate pore system and undulating extinction, typical of spheroolithid eggshells. We erect a new oogenus, Guegoolithus to include material previously reported as Macroolithus turolensis Amo-Sanjuan, Canudo and Cuenca-Bescós 2000, and new material from several lower Barremian localities of the Maestrazgo Basin in Spain. Guegoolithus eggshells exhibit an acicular radial ultrastructure in the lower part of the shell units, which develops into a tabular radial ultrastructure in the upper part, giving an appearance of a two-layered eggshell in thin sections that is not seen in SEM pictures. These features fill the gap between the two-layered prismatic eggshell of ceratopsians and the prolatospherulitic ornithopod eggshell of derived hadrosaurs.