Trigonoolithus
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''Trigonoolithus'' is an oogenus of dinosaur egg, representing a basal prismatoolithid. Its eggshell, like avian eggs, is composed of three structural layers, but cladistic analysis suggests that its parent was a non-avian theropod.


History

Fossil eggshells now assigned to ''Trigonoolithus'' were first discovered in
2009 File:2009 Events Collage V2.png, From top left, clockwise: The vertical stabilizer of Air France Flight 447 is pulled out from the Atlantic Ocean; Barack Obama becomes the first African American to become President of the United States; 2009 Iran ...
by Miguel Moreno-Azanza, José Manuel Gasca, and José Ignacio Canudo, three paleontologists from Universidad de Zaragoza.Moreno-Azanza, M., Gasca, J.M., and Canudo, J.I. (2009)
A high-diversity egg shell locality from the Hauterivian–Barremian transition of the Iberia Peninsula.
'' Journal of Vertebrate Paleontology'' 29 (Supplement to No. 3): 151A.
They recognized that the fossils represented a new oogenus of prismatoolithids, but the description would not be completed until
2014 File:2014 Events Collage.png, From top left, clockwise: Stocking up supplies and personal protective equipment (PPE) for the Western African Ebola virus epidemic; Citizens examining the ruins after the Chibok schoolgirls kidnapping; Bundles of wat ...
, when they published a description of the new oogenus and oospecies ''Trigonoolithus amoae'' in the paleontology journal '' Acta Palaeontologica Polonica''.


Distribution

Fossils of ''Trigonoolithus'' are found in abundance at the La Cantera site of the Blesa Formation in Teruel, Spain. This site is dated to the early
Barremian The Barremian is an age in the geologic timescale (or a chronostratigraphic stage) between 129.4 ± 1.5 Ma (million years ago) and 121.4 ± 1.0 Ma). It is a subdivision of the Early Cretaceous Epoch (or Lower Cretaceous Series). It is preceded ...
age.


Description

''Trigonoolithus'' is known from numerous eggshell fragments, but no complete or near-complete eggs. The whole egg of ''T. amoae'' was probably highly elongated, similar to other prismatoolithids. The shell fragments vary between 330 and 1040 μm in thickness, including the prominent triangular protuberances ornamenting their outer surface. Its eggshell is made up of three structural layers, with gradual boundaries between them. The middle layer, known as the prismatic layer, has a squamatic texture and the prismatic structure characteristic of Prismatoolithidae. It is three to four times thicker than the innermost layer (the mammillary layer), and two to three times thicker than the external layer. Circular pores 10 μm in diameter cut through ''Trigonoolithuss shell to allow for gas exchange. They are angusticanaliculate (meaning the pores are long, straight, and narrow), similar to the pore systems of ''
Prismatoolithus ''Prismatoolithus'' is an oogenus of dinosaur egg from the Cretaceous (Hauterivian-Maastrichtian) and possibly also the earliest Paleocene. They likely belonged to troodontids Troodontidae is a clade of bird-like theropod dinosaurs. During m ...
'', ''
Sankofa (pronounced ''SAHN''-koh-fah) is a word in the Twi language of Ghana meaning “to retrieve" (literally "go back and get"; - to return; - to go; - to fetch, to seek and take) and also refers to the Bono Adinkra symbol represented either w ...
'', and ''
Protoceratopsidovum ''Protoceratopsidovum'' is an oogenus of dinosaur egg from Mongolia. Despite its name (which means "eggs of ''Protoceratops''"Zelenitsky, D., and Currie, P. (2004) "A Cladistic Analysis of Theropod Ootaxa." ''Journal of Vertebrate Paleontology.'' ...
''.Carpenter, K. (1999). Eggs, Nests, and Baby Dinosaurs: A Look at Dinosaur Reproduction (Life of the Past). Indiana University Press, Bloomington, Indiana.


Parataxonomy

''Trigonoolithus'' is classified in the oofamily Prismatoolithidae, alongside ''
Preprismatoolithus ''Preprismatoolithus'' is a Late Jurassic oogenus. The species ''P. coloradensis'' is described by John Foster as being "of the prismatic basic type," with subspherical eggs about 10 cm (4 inches) in diameter."Eggs," Foster (2007) page ...
'', ''Prismatoolithus'', ''Protoceratopsidovum'', ''Sankofa'', and '' Spheruprismatoolithus''. Moreno-Azanza ''et al.'' performed multiple cladistic analyses to determine the phylogenetic position of ''Trigonoolithus''. Because no complete ''Trigonoolithus'' eggs are known, its position was slightly unstable, but ''Trigonoolithus'' was consistently placed as the
basalmost In phylogenetics, basal is the direction of the ''base'' (or root) of a rooted phylogenetic tree or cladogram. The term may be more strictly applied only to nodes adjacent to the root, or more loosely applied to nodes regarded as being close to th ...
member of Prismatoolithidae, or in a polytomy with other non-avian theropods and
birds Birds are a group of warm-blooded vertebrates constituting the class Aves (), characterised by feathers, toothless beaked jaws, the laying of hard-shelled eggs, a high metabolic rate, a four-chambered heart, and a strong yet lightweigh ...
.


Paleobiology

Prismatoolithids were previously hypothesized to be eggs of hypsilophodonts or ceratopsians, but later research found that they in fact are the eggs of theropods, based on analysis of preserved embryos of '' Prismatoolithus levis'' which showed them to be '' Troodon formosus''. Therefore, ''Trigonoolithus'' was also probably laid by a theropod. Based on its phylogenetic position, Moreno-Azanza ''et al.'' concluded it was most likely a non-dromaeosaurian, non-oviraptorid coelurosaur theropod dinosaur. Of the theropods found at La Cantalera (so far represented only by teeth), only aff. '' Paronychodon'' sp. and an indeterminate maniraptoran are possible parents of ''Trigonoolithus''.


References

{{Reflist Egg fossils Barremian life Early Cretaceous animals of Europe Cretaceous Spain Fossils of Spain Fossil parataxa described in 2014