Eggs Embryos Nest

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Russo, J., Mateus O., Balbino A., & Marzola M. (2014).  Crocodylomorph eggs and eggshells from the Lourinhã Fm. (Upper Jurassic), Portugal. Comunicações Geológicas. 101, Especial I, 563-566. Abstractrusso_et_al_2014_crocodylomorph_eggs_and_eggshells_from_the_lourinha_fm_upper_jurassic_portugal.pdf

We here present fossil Crocodylomorpha eggshells from the Upper Jurassic Lourinhã Formation of Portugal, recovered from five sites: one nest from Cambelas with 13 eggs, and three partial eggs and various fragments from, Paimogo N (I), Paimogo S (II), Casal da Rola, and Peralta. All specimens but the nest were found in association with dinosaur egg material. Our research reveals that on a micro- and ultrastructural analysis, all samples present the typical characters consistent with crocodiloid eggshell morphotype, such as the shell unit shape, the organization of the eggshell layers, and the triangular blocky extinction observed with crossed nicols. We assign the material from the Lourinhã Formation to the oofamily Krokolithidae, making it the oldest crocodylomorph eggs known so far, as well as the best record for eggs of non- crocodylian crocodylomorphs. Furthermore, our study indicates that the basic structure of crocodiloid eggshells has remained stable since at least the Upper Jurassic.

Araújo, R., Castanhinha R., Martins R. M. S., Mateus O., Hendrickx C., Beckmann F., Schell N., & Alves L. C. (2013).  Filling the gaps of dinosaur eggshell phylogeny: Late Jurassic Theropod clutch with embryos from Portugal. Scientific Reports. 3(1924), , 2013/05/30/onlin: Macmillan Publishers Limited. All rights reserved Abstractaraujo_et_al_2013_filling_the_gaps_of_dinosaur_eggshell_phylogeny_late_jurassic_theropod_clutch_with_embryos_from_portugal.pdf

The non-avian saurischians that have associated eggshells and embryos are represented only by the sauropodomorph Massospondylus and Coelurosauria (derived theropods), thus missing the basal theropod representatives. We report a dinosaur clutch containing several crushed eggs and embryonic material ascribed to the megalosaurid theropod Torvosaurus. It represents the first associated eggshells and embryos of megalosauroids, thus filling an important phylogenetic gap between two distantly related groups of saurischians. These fossils represent the only unequivocal basal theropod embryos found to date. The assemblage was found in early Tithonian fluvial overbank deposits of the Lourinhã Formation in West Portugal. The morphological, microstructural and chemical characterization results of the eggshell fragments indicate very mild diagenesis. Furthermore, these fossils allow unambiguous association of basal theropod osteology with a specific and unique new eggshell morphology.

Carrano, M., Mateus O., & Mitchell J. (2013).  First definitive association between embryonic Allosaurus bones and prismatoolithus eggs in the Morrison Formation (Upper Jurassic, Wyoming, USA). Annual Meeting of Vertebrate Paleontology. 101.: Journal of Vertebrate Paleontology, Program and Abstracts, 2013 Abstractcarrano_mateus_mitchell_2013_allosaurus_embryos_morrison_svp_abstract.pdf

Despite more than a century of collecting, resulting in one of the best-studied vertebrate fossil records anywhere in the world, the Upper Jurassic Morrison Formation has produced surprisingly few examples of dinosaur eggs associated with embryonic remains. Even more puzzling, none of these seem to pertain to the theropod Allosaurus, one of the most common and best-understood dinosaur taxa in the formation. Here we
report on a dinosaur nest site that has produced both abundant prismatoolithid eggshell and embryonic (or perinatal) bones of Allosaurus from Fox Mesa, Wyoming. This represents the first such discovery for any theropod in the Jurassic of North America. The nest is heavily weathered but contains a few ellipsoid eggshell clusters that suggest an egg size of about 8 x 6.5 cm. Study of the eggshell morphology and microstructure confirms that a single egg type is present throughout, which is indistinguishable from Prismatoolithus coloradensis. All of the identifiable embryonic materials pertain to theropods, and two premaxillae specimens show the five alveoli diagnostic for Allosaurus among Morrison theropods. This confirms the theropod origin of Prismatoolithus eggs and implicates Allosaurus as the specific Morrison parent taxon. As a result, it is now possible to assign several previous discoveries of dinosaur eggs and potential nests to Allosaurus, including the isolated egg from the Cleveland-Lloyd Quarry. This discovery
also calls into question prior assignments of Prismatoolithus eggs to ornithopods, and suggests that more detailed study of such sites is warranted. Prismatoolithus eggshells are also associated with the Upper Jurassic theropod Lourinhanosaurus from Portugal, along with larger embryos that exhibit four premaxillary alveoli.

Ribeiro, V., Holwerda F., & Mateus O. (2013).  Theropod egg sites from the Lourinhã Formation, Portugal. Journal of Vertebrate Paleontology, Program and Abstracts, 2013. 198.ribeiro_et_al_2013_theropod_egg_sites_lourinha_svp_meeting_abstracts_213.pdf
Araújo, R., Castanhinha R., Mateus O., & Martins R. (2012).  Late Jurassic theropod embryos from Porto das Barcas, Lourinhã formation, Portugal. Journal of Vertebrate Paleontology, Program and Abstracts, 2012, ISSN 1937-2809 . 57. Abstractaraujo_et_al_mateus_2012_dinosaur_eggs_portugal_svp_2012_abstract_book_meeting_abstracts.pdf.pdf

A clutch of several crushed eggs and embryos from the Late Jurassic (near the Kimmeridgian-Tithonian
boundary), Lourinhã Formation, Portugal contains a complete maxilla, erupted and scattered teeth,
and presacral vertebrae. The maxilla bears four teeth separated by individualized interdental plates,
the dorsal process of the maxilla is confluent with the maxillary body, the ventral rim of the antorbital
fossa is parallel to the tooth row, and the anterior border of the maxilla forms a right angle with the
ventral margin. The teeth are conical and recurved distally with carinae on mesial and distal sides. The
vertebrae are amphiplatyan, with a ventral pair of neurovascular foramina and heavily pitted articular
facets. These fossils allow unambiguous association of basal theropod osteology (Megalosauroidea) with
a new eggshell morphotype. Synchrotron micro-computed tomographic scanning (SRμCT), scanning
electron microscopy, and thin-sections under polarized and normal light revealed that the outer
ornamentation of the eggshell is composed of anastomosing ridges and islets, the pores communicate
near the outer region of the eggshells, and in radial section they are irregular canals that ramify towards
the surface. Micro-proto induced x-ray emission (micro-PIXE) analysis of the eggshell (excluding pores)
revealed the presence of Mg, Fe, Mn (0.33%, 0.27% and 0.18%, respectively) and several trace elements,
with a corresponding loss of Ca (39.4% detected but 40.0% expected for calcite), which suggests minimal
eggshell diagenesis. The eggshells do not luminesce, which could imply that no diagenetic alteration
took effect. However, the quenching effect of Fe2+ has to be taken into consideration. Conversely,
luminescence is observed in the pores since they are filled with sediment, composed of phyllosilicates,
as revealed by SRμCT, micro-PIXE and x-ray diffraction analyses.

Martins, R. M. S., Beckmann F., Castanhinha R., Araújo R., O. Mateus, & Pranzas P. K. (2011).  Tomographic techniques for the study of exceptionally preserved dinosaur and crocodile fossils from the mesozoic of Portugal. VII Encontro Nacional de Biologia Evolutiva, Coimbra, 21 Dezembro 2011. 19. Abstractmartins_et_al_mateus_2011_tomographic_techniques_for_the_study_of_exceptionally_preserved_dinosaur_and_crocodile.pdf

Portugal is ranked within the ten countries with more dinosaur taxa and the Lourinhã Formation is known
by the Late Jurassic findings of dinosaurs and other fossils. Often, studies of the external morphological
characteristics of the fossils are not sufficient and, observations of internal structures, using non-
destructive techniques, are required. The fossils here presented belong to the Museum da Lourinhã
(Portugal) and comprise a lower jaw of a basal crocodilian (possibly a Tomistomidae), eggshells and
several vertebrae from the exceptionally well preserved in ovo remains of Late Jurassic theropod dinosaur
Lourinhanosaurus. Neutron Tomography (NT) experiments with this material has been carried out at the
Geesthacht Neutron Facility in Germany. Additionally, eggshell fragments and several vertebrae have
been studied by Synchrotron-Radiation based Micro-Computed Tomography (SRμCT) at the HARWI II and
BW2 beamlines, respectively. These beamlines are operated by the Helmholtz-Zentrum Geesthacht at the
storage ring DORIS III at the Deutsches Elektronen–Synchrotron DESY in Hamburg, Germany. In both cases
(NT and SRμCT) complete 3D recordings have been obtained using a non-destructive procedure. The high-
quality tomographic datasets can be effectively studied through interactive digital visualization. Hence,
these visualization methods provide precious information about the 3D internal micro morphology of
fossils, like the network of the eggshell pores, often invisible in more traditional techniques, and provide a
direct window into the evolutionary history of organisms.

Castanhinha, R., Araújo R., & Mateus O. (2009).  Dinosaur eggshell and embryo localities in Lourinhã Formation, Late Jurassic, Portugal. Journal of Vertebrate Paleontology, 29(3): . 76A. Abstractcastanhinhaetal2009dinosaureggshellp.pdf

Four different localities from the Late Jurassic of Lourinhã formation with eggshells and embryos were studied: Paimogo (lower Amoreira-Porto Novo member), Peralta (Praia Azul member), Porto das barcas (Bombarral member) and Casal da Rôla (Amoreira-Porto Novo member). All but Casal da Rôla have embryonic material. Preliminary results show that eggshells from Paimogo correspond to obliquiprismatic morphotype (0.92mm thick), similar to those from Morrison Formation. Within Paimogo site a different type of eggshell was discovered, having a radial section of 153 μm with a mammilary layer measuring 65 μm. Porto das Barcas eggshells represent a discretispherulitic morphotype (1,23 mm thick).
This locality presents a nest 60-cm diameter containing many eggshells but an indeterminate number of eggs. Some embryonic bones were discovered between the eggshells including teeth and skull bones showing that the eggs belong to a saurischian, tentatively a sauropod dinosaur. Peralta nest eggshells are preliminary ascribed to obliquiprismatic morphotype (column: 0,56mm and mammilla: 0,21mm) probably related to Paimogo’s nest taxon (Lourinhanosaurus). Peralta site bears embryonic bones namely small theropod teeth associated with bone fragments, and unidentifiable dinosaur vertebra. Only eggshells have been collected at Casal da Rôla (ML1194). The eggshells (0,78mm thick) are prismatic morphotype and it was impossible to determine the pore system, the outer surface is smooth with no ornamentation.
Lourinhã formation has the oldest sauropod and theropod nest with embryos known so far.

Castanhinha, R., Araujo R., & Mateus O. (2008).  Reptile Egg Sites From Lourinhã Formation, Late Jurassic, Portugal. Livro de Resumos de Tercer Congreso Latinoamericano de Paleontologia de Vertebrados. , Neuquén, Argentina Abstractcastanhinha_et_al_2008_reptile_egg_sites_from_lourinha_formation_late_jurassic_portugal.pdf

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Cunha, P. P., Mateus O., & Antunes M. T. (2004).  The sedimentology of the Paimogo dinosaur nest site (Portugal, Upper Jurassic). 23 rd IAS Meeting of Sedimentology. 93., Coimbra, Portugal Abstractcunha_p_p_2004_-_the_sedimentology_of_the_paimogo_dinosaur_nest_site_portugal10467.pdf

Sedimentological features of the Paimogo site, 6 km NNW of Lourinhã, western central Portugal are presented. More than one hundred theropod dinosaur eggs (some containing embryo bones) ascribed to Lourinhanosaurus antunesi Mateus 1998, three crocodilian eggs and some other fossils were found at the 32 m2 excavated area of the egg-bearing horizon (Mateus et al., 1998). The stratigraphic position of the site is the Praia Azul member (Lourinhã Formation), roughly corresponding to the Kimmeridgian-Tithonian boundary or, more likely lowermost Tithonian. The maximum flooding surface of the basinal transgressive event where the horizon is located corresponds to the base of the H depositional sequence defined by Pena dos Reis et al. (2000) and probably correlates to the base of sequence Ti1 identified within western European basins (Jacquin et al., 1998), dated as 150 Ma. Possibly during the normal river discharge, the theropods congregated in nesting colonies at the backswamp of an extensive flood plain with small meandering channels and freshwater ponds. There are no evidences that the nest was dug or the eggs buried. The eggs have probably been laid on a flat, shaded, muddy area near the bank of a large pond. It is probable that the eggs have not been actively incubated. The larger number of eggs suggest that they were laid near simultaneously by, at least, six females. The fossil record shows that crocodilians, mammals, gastropods and fish were also present. A flood event occurred when theropod embryos had attained a late stade of ontogenetic development, probably just before hatching. The overflow from a nearby channel flooded the plain, including the area where the eggs had been laid. The sheet flood flowing over the nest resulted into the scattering and breaking up of some dinosaur eggs. Eggshell and embryos skeletal parts fragments were displaced to an adjacent area where, due to hydrodynamic decline, the flow submerged other clutches and moderately dragged their eggs. The flooding caused the drowning of the embryos and covered the eggs with fine-grained sediment, hiding them from predators and scavengers. Hydrodynamic interpretation of the arrangement of the theropod eggs and egg-fragments suggests that the flow came from the NW. When the floodwaters receded, the fine-grained deposits became exposed to subaerial weathering. Although the sediment surface was often wet and small bodies of standing water may still have existed, the sediments were oxidized and plant remains have consequently been destroyed. Some carbonate cementation and redenning resulted from pedogenesis under alternating dry and moist conditions, in a semiarid/ sub-tropical climate under seasonal changing, contrasting conditions. The thick, stratigraphically above and below the nesting horizon mudrocks indicate a long persistence of periodic flooding, alternating with pedogenesis. During the early stages of diagenesis, vertical pressure crushed the eggs. Silt penetration into the inner part of each egg inhibited later flattening during the burial process.

de Ricqlès, A., Mateus O., Antunes M. T., & Taquet P. (2001).  Histomorphogenesis of embryos of Upper Jurassic Theropods from Lourinha (Portugal). Comptes Rendus De L Academie Des Sciences Serie Ii Fascicule a-Sciences De La Terre Et Des Planetes. 332, 647-656., Jan Abstractricqles_mateus_et_al_2011_histomorphogenesis_of_embryos_of_upper_jurassic_theropods_from_lourinha_portugal.pdfWebsite

Remains of dinosaurian embryos, hatchlings and early juveniles are currently the subject of increasing interest, as new discoveries and techniques now allow to analyse palaeobiological subjects such as growth and life history strategies of dinosaurs. So far, available ‘embryonic’ material mainly involved Ornithopods and some Theropods of Upper Cretaceous age. We describe here the histology of several bones (vertebrae, limb bones) from the tiny but exceptionally well preserved in ovo remains of Upper Jurassic Theropod dinosaurs from the Paimogo locality near Lourinhã (Portugal). This Jurassic material allows to extend in time and to considerably supplement in great details our knowledge of early phases of growth in diameter and in length of endoskeletal bones of various shape, as well as shape modelling among carnivorous dinosaurs. Endochondral ossification in both short and long bones involves extensive pads of calcified cartilages permeated by marrow buds. We discuss the likely occurrence of genuine cartilage canals in dinosaurs and of an avian-like ‘medullary cartilaginous cone’ in Theropods. Patterns of periosteal ossification suggest high initial growth rates (20 μ m·day−1 or more), at once modulated by precise and locally specific changes in rates of new bone deposition. The resulting very precise shape modelling appears to start early and to involve at once some biomechanical components.