Cairanoolithus

''Cairanoolithus'' is an oogenus of dinosaur egg which is found in Southwestern Europe. The eggs are large ( in diameter) and spherical. Their outer surface is either smooth, or covered with a subdued pattern of ridges interspersed with pits and grooves. Multiple fossil egg clutches are known but the nest structure is unclear.

The parent of ''Cairanoolithus'' is probably some kind of non-ornithopod ornithischian, possibly the nodosaurid '' Struthiosaurus''.

The eggs were first named in 1994, when the two oospecies were classified in distinct oogenera as ''Cairanoolithus dughii'' and ''Dughioolithus roussetensis''. They are now considered to belong in a single oogenus, possibly even a single oospecies. Though it has been classified as a megaloolithid, ''Cairanoolithus'' is now placed in its own oofamily, Cairanoolithidae.

Description

''Cairanoolithus'' eggs are spherical and fairly large, measuring in diameter. The outer surface is smooth or covered with a subdued netlike pattern of ridges, interspersed with pits and grooves (sagenotuberculate ornamentation). The eggshells are made up of partially interlocking column-shaped shell units and range from thick.

Several egg clutches of ''C. dughii'' are known, containing as many as 25 fossilized eggs. However, taphonomical alterations (changes during the fossilization process) make it difficult to determine the original structure of the nest. Cousin (2002) hypothesized that ''Cairanoolithus'' eggs were laid on the surface of the ground, possibly buried beneath a mound of plant matter. Tanaka ''et al.'' (2015) noted that the shell had a high rate of water vapor conductance. Therefore, they concluded that ''Cairanoolithus'' nests were covered by organic or inorganic material, similar to modern eggs with high vapor conductance.

Oospecies

Two oospecies of ''Cairanoolithus'' have been described:
* ''Cairanoolithus dughii'' is the type oospecies. At , its eggshell is slightly thicker than that of ''C. roussetensis''. It has slender, partially fused columnar eggshell units. Their outer surface is almost without ornamentation, and the inner surface is covered with hollows once filled by organic cores. ''C. dughiis eggshell exhibits an angusticanaliculate pore system, i.e. its pores are long, narrow, and straight.
* ''Cairanoolithus roussetensis'', which was formerly classified in its own oogenus, ''Dughioolithus'', can be distinguished from ''C. dughii'' by its thinner eggshell (measuring thick), its broader eggshell units, and the relative prominence of its ornamentation. Like ''C. dughii'', ''C. roussetensis'' typically has an angusticanaliculate pore system, though some specimens have prolatocanaliculate pores, meaning they have variable diameter across their length.
Some authors consider the two oospecies to be synonymous. Cousin (2002) argued that the differences between them were due to intraspecific variation or due to taphonomy. He also described several eggshell fragments that possibly belong to an additional distinct oospecies of ''Cairanoolithus''; however these specimens were referred to ''C. roussetensis'' by Selles and Galobart (2015).

Classification

While it was formerly considered a megaloolithid, ''Cairanoolithus'' is now considered to belong its own monotypic oofamily, Cairanoolithidae. It belongs to the dinosauroid-spherulitic basic type, a group including sauropod eggs and ornithischian eggs, but paraphyletically excluding theropod eggs.

The cladistic analysis done by Selles and Galobart in 2015 recovered ''Cairanoolithus'' as a sister taxon to the clade of ornithopod eggs '' Guegoolithus'', '' Spheroolithus'', and '' Ovaloolithus''. Therefore, they considered it likely that ''Cairanoolithus'' belongs to a non-ornithopod ornithischian dinosaur.

Parentage

Since embryos are unknown in cairanoolithid eggs, the identity of their parent is uncertain. They have long been considered to be eggs of titanosaurs or ornithopods (like '' Rhabdodon''). However, numerous characteristics distinguish ''Cairanoolithus'' from sauropod eggs (oofamilies Megaloolithidae and Faveoloolithidae), even though they bear superficial similarities in size and shape. ''Cairanoolithuss columnar eggshell units are quite unlike the fan-shaped ones seen in '' Megaloolithus'', '' Faveoloolithus'', or '' Fusioolithus''. Also, its subdued ornamentation contrasts strongly with the heavily sculpted eggshells of sauropod eggs, and it has a different pore system. Eggs of ornithopods ( Spheroolithidae and Ovaloolithidae), on the other hand, show much closer similarity to cairanoolithids in ornamentation and pore system. However, ornithopod eggs are typically much smaller, and the crystal structure of their eggshell units is distinct.

The cladistic analysis by Sellés and Galobart in 2015 supported an ornithischian parentage. Late Campanian to early Maastrichtian ornithischians from Southwestern Europe are restricted to rhabdodontids and the nodosaurid '' Struthiosaurus''. When Sellés and Galobart analyzed the pelvises of ''Rhabdodon'' (the largest known rhabdodontid) and ''Struthiosaurus'', they found that ''Rhabdodon'' could not have laid eggs as big as ''Cairanoolithus''. On the other hand, even though ''Struthiosaurus'' was relatively small, the unique orientation of its ischia would have easily allowed it to lay eggs as large as a cairanoolithid egg. However, interpreting ''Cairanoolithus'' as the eggs of a nodosaur does raise the question of why ''Cairanoolithus'' or similar eggs have not been found in areas with a greater nodosaur abundance.

Distribution

''Cairanoolithus'' is native to Southwestern Europe, including southern France and northern Iberia. Its fossils date to the late Campanian to early Maastrichtian. They are usually found in the Aix-en-Provence Basin below the Rognac Limestone. ''C. dughii'' is from the La Cairanne site in Bouches du Rhône, France, from Roquehautes-Grand Creux and from the Villeveyrac Basin. ''C. roussetensis'' is found in the northern part of Iberia ( Tremp Group of Spain), and from southern France (in Rousset Village, Roquehautes-Crete du Marbre, the Villeveyrac Basin, and Argelliers-Montamaud).

Paleoecology

The Late Cretaceous ecosystems of Europe (which was then an island archipelago) show complex mixing of taxa originating from Africa, Asia, and North America. In Southwestern Europe, ''Cairanoolithus'' co-occurs with numerous other types of fossil eggs; ''Megaloolithus'' is particularly common, but theropod eggs such as ''Prismatoolithus'' and the ornithopod egg ''Guegoolithus'' are also present. Dinosaur body fossils are also common, including nodosaurids, rhabdodontids, titanosaurs, dromaeosaurids, basal iguanodontians, hadrosaurids, neoceratosaurians, and coelurosaurs. Other vertebrates include bony fish, squamates, cryptodiran turtles, alligatorids, and mammals.

History

The Aix Basin was first excavated for fossils in 1869 by French paleontologist Philippe Matheron. In the 1950s, Raymond Dughi and Francois Sirugue, a pair of French paleontologists working for the Museum d'Histoire Naturelle Aix-en-Provence, extensively studied the basin's fossil eggshells. They divided the eggs they had found into ten different types, but they did not describe them in detail. In the 1970s and 1980s, further work was done by the French paleontologist P. Kerourio and the German paleontologist H. K. Erben.M. Vianey-Liaud, P. Mallan, O. Buscail and C. Montgelard. (1994) "Review of French dinosaur eggshells: morphology, structure, mineral, and organic composition." ''Dinosaur Eggs and Babies,'' Cambridge University Press, Cambridge. pp 151-183

In his 1983 doctoral thesis, M. M. Penner devised one of the early classification schemes for egg fossils. He was the first to recognize the eggs now named ''Cairanoolithus'' as a distinct type; under his classification scheme, they were called "Group 2". In 1994, French paleontologists M. Vianey-Liaud, P. Mallan, O. Buscail and C. Montgelard described them under the modern parataxonomic system as ''Cairanoolithus dughii'' and ''"Dughioolithus" roussetensis''. They did not assign either of them to any oofamily, but both oogenera were classified in the oofamily Megaloolithidae by the Russian paleontologist Konstantin Mikhailov in 1996. Following further discoveries in 2001, Géraldine Garcia and Monique Vianey-Liaud synomized the two oogenera. In 2002, French paleontologist R. Cousin took a step further and synonymized the two oospecies.

In 2012, the first ''Cairanoolithus'' fossils discovered outside of France were first reported by Albert G. Selles in his PhD thesis at Universitat de Barcelona, in which he also proposed that ''Cairanoolithus'' be moved into its own oofamily. Three years later, Selles and Angel Galobart published a comprehensive reanalysis of ''Cairanoolithus'', in which they formally named the new oofamily, Cairanoolithidae, to contain ''Cairanaoolithus''. Contrary to Cousin's conclusions, Selles and Galobart separated the oospecies ''C. dughii'' and ''C. roussetensis''. Also, they demonstrated that ''Cairanoolithus'' was not the eggs of an ornithopod or sauropod and conjectured that it could be the eggs of a nodosaur.

References

{{Taxonbar, from=Q5017647

Dinosaur reproduction
Egg fossils
Ornithischia
Maastrichtian life
Late Cretaceous dinosaurs of Europe
Cretaceous France
Fossils of France
Cretaceous Spain
Fossils of Spain
Tremp Formation
Fossil parataxa described in 1994