PBDB Taxon

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Macroelongatoolithus carlylei

Reptilia - Elongatoolithidae

Taxonomy

Oolithes carlylensis was named by Jensen (1970). It is considered to be a form taxon. Its type specimen is BYU-E 200, an egg, and it is a 3D body fossil. Its type locality is Castle Dale, Jensen Locality 1, which is in a Cenomanian terrestrial shale in the Cedar Mountain Formation of Utah.

It was recombined as Boletuoolithus carlylensis by Bray (1998); it was misspelled as Macroelongatoolithus carleylei by Krumenacker et al. (2022); it was recombined as Macroelongatoolithus carlylei by Zelenitsky et al. (2000), Jin et al. (2007), Bonde et al. (2008), Kim et al. (2011), Tanaka et al. (2011), Huh et al. (2014), Tanaka et al. (2016), Krumenacker et al. (2017), Simon et al. (2019), Hedge et al. (2025).

Synonyms

Longiteresoolithus xixiaensis was named by Wang and Zhou (1995). It is considered to be a form taxon. It is a 3D body fossil. Its type locality is Liuying, Yangcheng, which is in a Cenomanian/Turonian terrestrial siltstone in the Gaogou Formation of China.

It was recombined as Macroelongatoolithus xixiaensis by Jin et al. (2007), Jin (2008), Wang et al. (2010), Kim et al. (2011), Wang et al. (2012), Huh et al. (2014); it was misspelled as Macroelongatoolithus xixia by Cousin (2002), Zhou and Feng (2002), Liang et al. (2009); it was synonymized subjectively with Macroelongatoolithus carlylei by Zelenitsky et al. (2000), Simon et al. (2019), Hedge et al. (2025).
Megafusoolithus qiaoxianensis was named by Wang et al. (2010). It is considered to be a form taxon.

It was synonymized subjectively with Macroelongatoolithus carlylei by Simon et al. (2019), Hedge et al. (2025).
Macroelongatoolithus goseongensis was named by Kim et al. (2011). It is considered to be a form taxon. Its type specimen is KNUE 10101, an egg, and it is a 3D body fossil. Its type locality is Ddabakseom egg site, which is in a Campanian floodplain mudstone in the Goseong Formation of South Korea.

It was synonymized subjectively with Macroelongatoolithus carlylei by Simon et al. (2019), Hedge et al. (2025).

Synonymy list

Year	Name and author
1970	Oolithes carlylensis Jensen pp. 62-63 figs. 5, Pl. 1:1-2
1995	Longiteresoolithus xixiaensis Wang and Zhou p. 262
1998	Boletuoolithus carlylensis Bray p. 221
2000	Macroelongatoolithus carlylei Zelenitsky et al. p. 131–132
2002	Macroelongatoolithus xixia Cousin p. 44
2002	Longiteresoolithus xixiaensis Zhou and Feng p. 69
2007	Macroelongatoolithus carlylei Jin et al. p. 82
2007	Macroelongatoolithus xixiaensis Jin et al. p. 82
2008	Macroelongatoolithus carlylei Bonde et al. p. 255
2008	Macroelongatoolithus xixiaensis Jin p. 53
2009	Longiteresoolithus xixiaensis Liang et al. p. 1591
2010	Macroelongatoolithus xixiaensis Wang et al. p. 96
2010	Megafusoolithus qiaoxianensis Wang et al. p. 97
2010	Megafusoolithus qiaoxianensis Wang et al.
2011	Macroelongatoolithus carlylei Kim et al. p. 18
2011	Macroelongatoolithus goseongensis Kim et al. p. 18 figs. 2, 5
2011	Macroelongatoolithus xixiaensis Kim et al. p. 18
2011	Macroelongatoolithus carlylei Tanaka et al. p. 45
2012	Macroelongatoolithus xixiaensis Wang et al. p. 403
2012	Megafusoolithus qiaoxianensis Wang et al. p. 404
2014	Macroelongatoolithus goseongensis Huh et al. p. 218
2014	Macroelongatoolithus carlylei Huh et al. p. 220
2014	Macroelongatoolithus xixiaensis Huh et al. pp. 220-221
2016	Macroelongatoolithus carlylei Tanaka et al. p. 353
2017	Macroelongatoolithus carlylei Krumenacker et al. p. 170
2019	Macroelongatoolithus carlylei Simon et al. p. 7–8
2022	Macroelongatoolithus carleylei Krumenacker et al. p. 1339
2025	Macroelongatoolithus carlylei Hedge et al. pp. 6-7

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Rank	Name	Author
kingdom	Animalia	()
—	Bilateria
—	Eubilateria	Ax 1987
—	Deuterostomia
phylum	Chordata	Haeckel 1874
subphylum	Vertebrata
superclass	Gnathostomata
—	Osteichthyes	()
subclass	Sarcopterygii	()
subclass	Dipnotetrapodomorpha	(Nelson 2006)
subclass	Tetrapodomorpha	()
—	Tetrapoda
—	Reptiliomorpha
—	Anthracosauria
subclass	Amphibiosauria	Kuhn 1967
—	Cotylosauria	()
—	Amniota
—	Sauropsida

Rank	Name	Author
class	Reptilia
subclass	Eureptilia	()
—	Romeriida
—	Diapsida	()
—	Archosauromorpha	(Huene 1946)
—	Crocopoda
—	Archosauriformes	Gauthier 1986
—	Eucrocopoda
—	Archosauria	()
informal	Avemetatarsalia
—	Ornithodira
—	Dinosauromorpha
—	Dinosauriformes
—	Dinosauria	()
—	Theropoda	()
family	Elongatoolithidae
genus	Macroelongatoolithus	Li et al. 1995
species	carlylei	()

If no rank is listed, the taxon is considered an unranked clade in modern classifications. Ranks may be repeated or presented in the wrong order because authors working on different parts of the classification may disagree about how to rank taxa.

†Macroelongatoolithus carlylei Jensen 1970

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Invalid names: Longiteresoolithus xixiaensis Wang and Zhou 1995 [synonym], Macroelongatoolithus goseongensis Kim et al. 2011 [synonym], Megafusoolithus qiaoxianensis Wang et al. 2010 [synonym]

Diagnosis

Reference	Diagnosis
D. K. Zelenitsky et al. 2000	Same as for oogenus.
J. Y. Kim et al. 2011 (Macroelongatoolithus goseongensis)	Large, elongate eggs, 39 cm in length. Elongation index 3.4. Shell thickness 2.3-3.1 mm. Continuous to mammillary layer ratio 2.5:1-4.8:1. Eggs arranged within ring-like clutch.
M. Huh et al. 2014 (Macroelongatoolithus xixiaensis)	Large, elongate eggs 39 – 52 cm long and 13 – 18 cm wide. Elongation index ranges from 2.6 to 3.9. Continuous layer to mammillary layer ratio variable, with a distinct, undulating boundary between layers. Eggshell thickness ranges from 1.5 to 4.45 mm including surface ornamentation. Shell surface covered with coarse lineartuberculate ornamentation at equator grading to dispersituberculate at the poles. Eggs arranged in pairs within single-layered ring-shaped clutch with a central space devoid of eggs (modified after Jin et al. 2007).
D. J. Simon et al. 2019	Same as for oogenus.
J. Hedge et al. 2025	Large, elongate eggs exceeding 250 mm and up to 610 mm long. TST, including ornamentation, is 1.38–4.75 mm. Abrupt, undulating boundary separates the CL and ML. CL:ML ratio ranges widely from 2:1 to 8:1. Crystal splaying across the ML/CL boundary occurs at random intervals. Ornamentation is variable both in type and relief and may grade from dispersituberculate at the poles to lineartuberculate along the middle portion of the length of the egg; smooth patches may occur near poles. Clutches consist of paired eggs in a ring-shaped configuration with an external clutch diameter of up to 3 m.