Guegoolithus

''Guegoolithus'' is an oogenus of fossil egg from the early Cretaceous of Spain. It is classified in the oofamily Spheroolithidae, and was probably laid by an ornithopod dinosaur.

Distribution

''Guegoolithus'' is known from the Early Cretaceous of Spain. It is common in the Maestrazgo Basin, dating to the lower Barremian. Fossils have been found in the Blesa, El Castellar, Camarillas, and Mirambel Formations. It has also been found at the slightly older El Hocajo, a part of the Cameros Basin which dates to the Valanginian or Hauterivian ages, up to 7 million years older than the Maestrazgo specimens.

Description

''Guegoolithus'' is known from over 400 fossil eggshell fragments, but no complete eggs have been found. They are very thin for spheroolithids, ranging from 0.42 to 1.5 mm in thickness.Moreno-Azanza, M., J.I. Canudo, and J.M. Gasca. (2014)
Spheroolithid eggshells in the Lower Cretaceous of Europe. Implications for eggshell evolution in ornithischian dinosaurs.
''Cretaceous Research'' 51: 75–87. The wide range of eggshell thickness is partially due to erosion on the inner and outer surfaces of the fragments.Amo Sanjuan, O., Canudo, J.I., Cuenca-Bescos, G. (2000)
First record of elongatoolithid eggshells from the Lower Barremian (Lower Cretaceous) of Europe (Cuesta Corrales 2, Galve Basin, Teruel, Spain).
In: Bravo, A.M., Reyes, T. (Eds.), ''First International symposium on Dinosaur eggs and babies. Extended abstracts. Isona i Conca Della,'' pp. 7–14. The size and shape of the complete egg is unknown.

Like other types of dinosaur eggs, the shell of ''Guegoolithus'' is made up of tightly packed crystalline units. The shell is single-layered, but the shell units appear to be have two distinct layers because they exhibit a radiating acicular crystal structure near the base (the inner edge of the eggshell), but form a tabular ultrastructure on the upper third of the eggshell. The surface of the eggshell exhibits sagenotuberculate ornamentation (nodes and ridges forming a net-like pattern, with pits and grooves in betweenCarpenter, K. 1999. Eggs, Nests, and Baby Dinosaurs: A Look at Dinosaur Reproduction (Life of the Past). Indiana University Press, Bloomington, Indiana.
). It has a prolatocanaliculate pore system, similar to ''Spheroolithus'', which means that the pores are irregularly shaped and vary in width along their length. Also like ''Spheroolithus'', it has a prolatospherulitic morphotype, with eggshell units fused together and horizontal growth lines on the lower portion of the eggshell. ''Guegoolithus'' has much more prominent ornamentation that ''Spheroolithus'', and a thinner eggshell than most ''Spheroolithus'' species.

History

''Guegoolithus'' was first discovered in 2000 by Spanish paleontologists Olga Amo-Sanjuán, José Ignacio Canudo, and Gloria Cuenca-Bescós, though it was described as an oospecies of ''Macroolithus''. In 2008, a paper suggested that ''"M." turolensis'' be used as an index fossil for fossil sites in the Sistema Ibérico, because they are easily identifiable and always found in rocks of Hauterivian-Barremian age.M. Moreno-Azanza, J. M. Gasca, and J. I. Canudo. (2008)
Macroolithus turolensis como fósil guia para el Hauteriviense superior-Barremiense basal de Teruel
In J I Ruiz-Omeñaca, L Piñuela and J C García-Ramos (eds), ''XXIV Jornadas de la Sociedad Española de Paleontología,'' 15–18 October 2008, Museo del Jurásico de Asturias (MUJA), Colunga, Spain, Libro de Resúmenes 43–44. An analysis in 2014 by Miguel Moreno-Azanza, José Ignacio Canudo, and José Manuel Gasca found that in fact, ''"M." turolensis'' is not an elongatoolithid, but instead belongs in Spheroolithidae, and named a new oogenus for it: ''Guegoolithus''. In 2016, Moreno-Azanza and several of his colleagues reported the discovery of a new fossil site in La Rioja, Spain, which included eggs assigned to ''Guegoolithus''. Since this site is somewhat older than the Barremian age of the previously reported ''Guegoolithus'' fossils, Moreno ''et al.'' rejected the idea of using them as index fossils.

Classification

While it was previously considered an oospecies of ''Macroolithus'' in the oofamily Elongatoolithidae, ''Guegoolithus'' is now considered a spheroolithid because of the similarities of microstructure, ultrastructure, and ornamentation. It contains a single oospecies: ''G. turolensis''. Unfortunately, relationships within Spheroolithidae are not well known because of the unclear definitions of ''Spheroolithus'' and Speroolithidae.

Paleobiology

''Guegoolithus'' is classified as a spheroolithid, which would indicate it was laid by an ornithopod dinosaur. It is very similar to the eggs of ''Maiasaura peeblesorum'' and other hadrosaurs, but these were not common in Europe during the Early Cretaceous. ''G. turolensis'' is most likely laid by a closely related ornithopods. Styracosterns (a clade of ornithopods that includes hadrosaurs and their close relatives) are, like ''G. turolensis'', incredibly common in the lower Cretaceous of Spain. Given that fossils of ''Guegoolithus'' span a larger time period than is likely for a single species, the fossils probably represent many different styracostern species with structurally indistinguishable eggshells. Because ''Guegoolithus'' remained unchanged for upwards of 5 million years and it is very similar to the late Cretaceous spheroolithids, Moreno-Azanza ''et al.'' (2017) concluded that the nesting behavior of hadrosaurs and their close relatives remained unchanged for 80 million years. The eggshell structure is apparently designed to be easily broken from the inside, indicating that these dinosaurs were altricial.

Its reclassification outside of Elongatoolithidae implies that no elongatoolithids are known from the early Cretaceous of Europe. However, elongatoolithids have been reported from the Late Cretaceous of France.

References

{{Ornithopoda, E.

Egg fossils
Dinosaur reproduction
Early Cretaceous dinosaurs of Europe
Ornithopods
Cretaceous Spain
Fossils of Spain
Camarillas Formation
Fossil parataxa described in 2014