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Russo, J., Mateus O., Marzola M., & Balbino A. (2017).  Two new ootaxa from the late Jurassic: The oldest record of crocodylomorph eggs, from the Lourinhã Formation, Portugal. PLOS ONE. 12, 1-23., 03, Number 3: Public Library of Science Abstractrusso_et_al_2017_two_new_ootaxa.pdfWebsite

The Late Jurassic Lourinhã Formation is known for its abundant remains of dinosaurs, crocodylomorphs and other vertebrates. Among this record are nine localities that have produced either dinosaur embryos, eggs or eggshell fragments. Herein, we describe and identify the first crocodiloid morphotype eggs and eggshells from the Lourinhã Formation, from five occurrences. One clutch from Cambelas, composed of 13 eggs, eggshell fragments from Casal da Rola and Peralta, one crushed egg and eggshells from Paimogo North, and four crushed eggs as well as eggshell fragments from Paimogo South. We observed and confirmed diagnostic morphological characters for crocodiloid eggshells and which are consistent with a crocodylomorph affinity, such as the ellipsoidal shape, wedge-shaped shell units, triangular extinction under cross-polarized light, and tabular ultrastructure. This material is distinctive enough to propose two new ootaxa within the oofamily Krokolithidae, Suchoolithus portucalensis, oogen. and oosp. nov., for the material from Cambelas, the most complete clutch known for crocodiloid eggs, and Krokolithes dinophilus, oosp. nov., for the remaining material. These are the oldest crocodylomorph eggs known, extending the fossil record for this group to the Late Jurassic. Furthermore, except for the clutch from Cambelas, the material was found with theropod eggs and nests, in the other four occurrences, which seem to suggest some form of biological relationship, still unclear at this point.

Mateus, O., Callapez P. M., & Puértolas-Pascual E. (2017).  The oldest Crocodylia? a new eusuchian from the Late Cretaceous (Cenomanian) of Portugal. Journal of Vertebrate Paleontology, Program and Abstracts. 2017, 160. Abstractmateus_et_al_2017_oldest_crocodylia_svp_2017_abstract.pdf


Marzola, M., Russo J., & Mateus O. (2015).  Identification and comparison of modern and fossil crocodilian eggs and eggshell structures. Historical Biology. 27(1), 115-133. Abstractmarzola_et_al_2015_identification_and_comparison_of_modern_and_fossil_crocodilian_eggs_and_eggshell_structures.pdfWebsite

Eggshells from the three extant crocodilian species Crocodylus mindorensis (Philippine Crocodile), Paleosuchus palpebrosus (Cuvier's Smooth-fronted Caiman or Musky Caiman) and Alligator mississippiensis (American Alligator or Common Alligator) were prepared for thin section and scanning electron microscope analyses and are described in order to improve the knowledge on crocodilian eggs anatomy and microstructure, and to find new apomorphies that can be used for identification. Both extant and fossil crocodilian eggs present an ornamentation that vary as anastomo-, ramo- or the here newly described rugosocavate type. The angusticaniculate pore system is a shared character for Crocodylomorpha eggshells and some dinosaurian and avian groups. Previously reported signs of incubated crocodilian eggs were found also on our only fertilised and hatched egg. Paleosuchus palpebrosus presents unique organization and morphology of the three eggshell layers, with a relatively thin middle layer characterised by dense and compact tabular microstructure.

Young, M. T., Hua S., Steel L., Foffa D., Brusatte S. L., Thüring S., Mateus O., Ruiz-Omeñaca J. I., Havlik P., Lepage Y., & de Andrade M. B. (2015).  Addendum to ‘Revision of the Late Jurassic teleosaurid genus Machimosaurus (Crocodylomorpha, Thalattosuchia)’. Royal Society Open Science. 2, , Number 2: The Royal Society Abstractyoung_et_al_2015_addendum_to_revision_of_the_late_jurassic_teleosaurid_genus_machimosaurus_crocodylomorpha_thalattosuchia.pdfWebsite


Marzola, M., Mateus O., Russo J., & Milàn J. (2014).  Comparison of modern and fossil Crocodylomorpha eggs and contribution to the oophylogeny of Amniota. XII Annual Meeting of the European Association of Vertebrate Palaeontologists. , p. 192, Regione Piemonte: European Association of Vertebrate Palaeontologists. Museo Regionale di Scienze Naturalimarzola_et_al._2014_comparison_of_modern_and_fossil_crocodylomorpha_eggs_and_contribution_to_the_oophylogeny_of_amniota-_eavp_2014.pdf
Russo, J., Mateus O., Marzola M., & Balbino A. (2014).  Eggs and eggshells of crocodylomorpha from the Late Jurassic of Portugal. Journal of Vertebrate Paleontology. Program and Abstracts, 2014, 218.russo_et_al_2014eggs_crocodylomorpha_portugal.pdf
Young, M. T., Hua S., Steel L., Foffa D., Brusatte S. L., Thüring S., Mateus O., Ignacio-Ruiz Omeñaca J., Lepage Y., Havilk P., & Andrade M. B. (2014).  Revision of the Late Jurassic teleosaurid genus Machimosaurus (Crocodylomorpha, Thalattosuchia). Royal Society Open Science. 1(140222), 1-42.young_et_al_2014_machimosaurus_crocodylomorph_revision.pdf
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.

Mateus, O. (2013).  Crocodylomorphs from the Mesozoic of Portugal and a new skull of eusuchian from the Late Cretaceous. 2013 Hwaseong International Dinosaurs Expedition Symposium, pp.66-67.. , Hwaseong, South Korea Abstractmateus_2013_crocodylomorphs_portugal_new_skull.pdf

The diversity of fossil crocodylomorphs in Portugal is high, with occurrence as old as Mystriosaurus (=Steneosaurus) bollensis from the Lower Jurassic. The Late Jurassic forms are the better documented, and include the following taxa: Machimosaurus hugii, Lisboasaurus estesi Seiffert, 1973, Lusitanisuchus mitrocostatus Seiffert, 1975; Schwarz & Fechner 2004, Theriosuchus guimarotae Schwarz and Salisbury 2005, Cf. Alligatorium, Goniopholis baryglyphaeus, and a crocodylomorph-like eggs in dinosaur nests (Mateus et al., 1998; Ricqlès et al., 2001). From the Lower Cretaceous were reported a few dinosaurs but its record is strangely scarce in crocodylomorphs (Mateus et al., 2011). The Upper Cretaceous crocodiles show a large diversity, but it is mostly based in fragmentary material that require revision, such as “Crocodylus” blavieri? Grey from the Upper Campanian - Maastrichtian of Viso, near Aveiro (initially reported by Sauvage 1897-98), Goniopholis cf. crassidens Owen 1841 and Oweniasuchus pulchelus Jonet 1981. Moreover there is a fascinating, but poorly understood, crocodylomorph diversity in the Cenomanian of Portugal, documented by fragmentary specimens that have been doubtfully assigned to Thoracosaurus Leidy 1852 of the Middle Cenomanian of Cacém, to the nomen dubium Oweniasuchus lusitanicus Sauvage 1897-98 (interpreted as a mesosuchian goniopholid) based in a fragmentary mandible from the Campanian-Maastrichtian, and also from the Middle Cenomanian of Portugal, Buffetaut and Lauverjat (1978) report an fragmentary unidentified possible dyrosaurid from Nazaré. All this specimens are too incomplete to be compared with the specimen here described. In contrast, Cenozoic crocodiles of Portugal are often known after complete skulls and several individuals. The taxa list include Iberosuchus macrodon (Lower to Middle Eocene), Tomistoma calaritanus (Early Miocene) and T. lusitanica (Burdigalian-Helvetian), and Diplocynodon sp. (Antunes, 1961, 1987, 1994).
At least, two different morphotypes of crocodylomorph eggs from the Late Jurassic of Lourinhã Formation are also known.
A new specimen here reported of crocodile based in a partial skull and mandible (ML1818) from the Uppermost Middle Cenomanian platform carbonates of Baixo Mondego, west central Portugal (Tentúgal Fm., Callapez, 2004). The taxon is phylogenetically positioned as a basal Eusuchia, due to the choanae enclosed by the pterygoid, and closely related with stem Crocodylia and Borealosuchus. This specimen represents the only well documented and valid eusuchian species in the Cenomanian of Europe and is the oldest representative of an eusuchian crocodylomorph, with the exception for the Barremian Hylaeochampsa vectiana.