Adynomosaurus

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Adynomosaurus
Temporal range: Late Cretaceous (Early Maastrichtian), 70  Ma
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Life reconstruction of Adynomosaurus arcanus.png
Life restoration
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Clade: Dinosauria
Clade:† Ornithischia
Clade:† Neornithischia
Clade:† Ornithopoda
Family:† Hadrosauridae
Subfamily:† Lambeosaurinae
Genus:† Adynomosaurus
Prieto-Márquez et al., 2019
Species:
†A. arcanus
Binomial name
†Adynomosaurus arcanus
Prieto-Márquez et al., 2019

Adynomosaurus is a genus of lambeosaurine dinosaur from the Late Cretaceous of what is now Catalonia, Spain. First discovered in 2012, it was named in 2019 with the type and only species being Adynomosaurus arcanus. It is only known from scant material, but is distinguished from other hadrosaurs by its weakly developed shoulder blade which would have had underdeveloped musculature, which lends it its scientific name, partially from the Greek word for "weak". Its exact relationships with other hadrosaurs remain unresolved, with it not consistently being recovered as a relative of any other specific genera, though some studies have allied it with Tsintaosaurini or even found it outside of Hadrosauridae. It would have lived as part of a diverse coastal estuary ecosystem, made up of meandering rivers and mud flats. The discovery of Adynomosaurus adds to the very incomplete fossil record of hadrosaurid dinosaurs in the Late Cretaceous of Europe, and it fits into a picture of major ecological turnover that was occurring during the Maastrichtian stage in the region.

Contents

Discovery and naming

Hadrosaur fossils from Serrat del Corb geologic locality, possibly representing Adynomosaurus Serrat-del-corb-hadrosaur.png
Hadrosaur fossils from Serrat del Corb geologic locality, possibly representing Adynomosaurus

The Costa de les Solanes locality of the Conques Formation was first discovered in 2012, by a wheat field in the village of Basturs in Catalonia; the site dates to the upper layers of the Maastrichtian. Numerous fossil sites preserving dinosaurs from the Maastrichtian are known from across this region of Spain. After being notified of the site by a local, researchers from the Institut Català de Paleontologia Miquel Crusafont and the Universitat Autònoma de Barcelona carried out excavations at the site during 2012 and 2013. The associated but disarticulated remains of a hadrosaur were discovered in this time, including a partial left dentary , a number of partial vertebrae , a left scapula , a left sternum , a partial rib, and numerous pelvic and partial hindlimb bones. Due to the presence of two left tibia, it was concluded the material belonged to at least two different individuals. The specimens are held at the Museu de la Conca Dellà in Lleida, Spain. [1] This would first be reported in an abstract for a presentation at the 78th annual meeting of the Society of Vertebrate Paleontology, with the remains then thought to pertain to a new species of the genus Pararhabdodon . [2]

These remains would be formally named and described by Albert Prieto-Márquez, Víctor Fondevilla, Albert G. Selles, Jonathan R. Wagner and Angel Galobart in the palaeontological journal Cretaceous Research, under the name Adynomosaurus arcanus. The distinctive scapula, with the specimen number MCD 7125, was selected as the holotype specimen for the taxon, due to being the most diagnostic element. The generic name is a composite of the Greek words ‘adýnamos’ (weak), ‘-mos’ (shoulder) and ‘sauros’ (lizard), referring to the taxon's shoulder blade being underdeveloped and likely supporting weaker musculature than other hadrosaurs. The specific name arcanus means "secret" or "occult", in reference to the scant and uninformative nature of the hadrosaur fossil record from the South-Central Pyrenean Basin. [1] An indeterminate hadrosaur specimen preserving a very complete pelvic girdle, MCD 4791, was described in 2013 by Albert Prieto-Márquez and colleagues. It hails from the Serrat del Corb locality of the Tremp Formation. [3] The paper describing Adynomosaurus noted multiple similarities shared between the specimen and their new taxon, but due to the lack of outright diagnostic traits from Adynomosaurus and a degree of stratigraphic separation, refrained from referring to the genus. [1]

Description

Scapulae of a standard lambeosaur; compared to this, the scapula of Adynomosaurus is underdeveloped, with reduced length and a small width on the bladed end (to the left) Scapulae of Olorotitan arharensis Godefroit, Bolotsky & Alifanov, 2003. Upper Cretaceous of Kundur, Russia.jpg
Scapulae of a standard lambeosaur; compared to this, the scapula of Adynomosaurus is underdeveloped, with reduced length and a small width on the bladed end (to the left)

In many respects, such as the vertebrae , humerus , femur , tibia , and metatarsals , the known anatomy of Adynomosaurus is indistinguishable from all other hadrosaurid dinosaurs. [1] As a member of the hadrosaur family, it would have been a quadrupedal animal, while bearing the ability to walk upon its hindlegs bipedally. It would have had a long skull, ending in a beak, and a large array of complex teeth; as a lambeosaur, it would have possessed a cranial crest made of the nasal bones, filled with hollow internal passages. [4] Despite its similarity to other hadrosaurs, some traits distinguish it from its relatives. Compared to other Spanish hadrosaurs, its dental anatomy stands out; the dentary of Koutalisaurus possesses tooth alveoli rotated slightly backwards, whereas in Arenysaurus and Blasisaurus they point upwards. Adynomosaurus possesses teeth intermediate to these conditions, rotated in the front half of the jaw but vertically oriented near the back. The tooth crown s are around three times taller than wide, also differing from the latter pair of genera, which have a more extreme ratio. [1]

Skull material of possible relative Tsintaosaurus; Adynomosaurus would have had a similar crest Tsintaosaurus spinorhinus.png
Skull material of possible relative Tsintaosaurus ; Adynomosaurus would have had a similar crest

In regards to the anatomy of the postcranium , the supraacetabular crest of the ilium is V-shaped, and is extended very far back, all the way to the connection point with the ischium . Such an extreme extension of the crest is not presented in most lambeosaurines, but can be observed in Parasaurolophus cyrtocristatus as well as the Serrat del Corb hadrosaur. Also shared with these two taxa are features of the iliac process of the ischium; its dorsal and ventral (i.e. top and bottom) margins are nearly parallel and it is stout, with a broad articulate facet. The most distinctive part of its anatomy is its scapula , or shoulder bone, which bears the only traits unique to it among hadrosaurs. Specifically, the scapula is generally underdeveloped; its length, proportionally, seems to be the shortest of all hadrosaurs, though this is unable to be definitively confirmed due to incompleteness of the bone amongst known specimens. More definitively, the scapular blade (the flattened end) is only 75% the width of the proximal end (base) of the bone, unlike in all other lambeosaurines where it is as wide or wider. Likewise, the proximal constriction of the scapula - the middle portion connecting the proximal end to the blade - is very thin, hardly half as deep at the proximal end and around eighty percent the depth of the deepest section of the blade. The deltoid ridge of the bone is heavily reduced. These aspects of scapular anatomy are known to not be variable with age in other hadrosaurs, ruling out that as an expalantion of the condition in Adynomosaurus. The scapulae of Pararhabdodon and the Basturs Poble hadrosaur, from similar times and places, display more conventional anatomy, distinct from that of Adynomosaurus. [1]

Based on the myology of modern crocodylians, it is surmised that the reduction of the scapulae in Adynomosaurus would have likewise meant reduction of the associated arm musculature. Thus the strength of abduction, reduction, and adduction of the humerus may have been lesser than other hadrosaurs. This cannot be known for certain, however, as it is possible the musculature size did not correlate with the size of the bone as might be expected. [1]

Classification

Adynomosaurus was described as a member of the hadrosaur subfamily Lambeosaurinae, but its relationships were not able to be determined more specifically; in a phylogenetic analysis it emerged in a soft polytomy alongside the genera Aralosaurus , Canardia , Jaxartosaurus , Tsintaosaurus , and Pararhabdodon as the base of the family. [1] Some studies have since found it to be outside of Lambeosaurinae, instead being a non-hadrosaurid hadrosauroid closely related to Nanningosaurus . [5] [6] Others, however, have maintained the lambeosaurine position. [7] [8] [9] One such study found it to be within a clade therein named Arenysaurini, alongside other European taxa and the moroccan genus Ajnabia . [8] Another study instead found it to be related to Tsintaosaurus and Pararhabdodon within Tsintaosaurini, with some other European taxa being more derived. [10] The resulting tree of the former study is seen on the left, while the tree of the latter is seen on the right:

Lambeosaurinae

Palaeoenvironment

Fossils indicate a diverse fauna of reptiles and amphibians lived alongside Adynomosaurus, such as this palaeobatrachid frog Fossilised frog.jpg
Fossils indicate a diverse fauna of reptiles and amphibians lived alongside Adynomosaurus, such as this palaeobatrachid frog

Adynomosaurus is known from the Conques Formation of the Tremp Group, which in the Cretaceous was part of the Ibero-Armorican island, the largest of several Europe was at the time divided into at the time. The distinct Basturs Poble lambeosaur as well as the Serrat del Rostiar tsintaosaurin hadrosaur are known from equivalent geological units. [1] It is well documented that a major change in the faunal composition of Late Cretaceous Europe occurred around the start of the Maastrichtian age, known as the "Maastrichtian Dinosaur Turnover". This saw the previously established dinosaurian herbivore fauna, composed of titanosaurs, rhabdodontids, and nodosaurids, go extinct in South-Western Europe and be replaced with different types of titanosaur as well lambeosaurine hadrosaurs, the latter of which go on to become overwhelmingly dominant across the region. Whether this was due directly to competition with lambeosaurs (which only arrive in Europe around the time of the turnover) or due to environmental changes that merely left a void for hadrosaurs is unknown. Despite the completeness of the change, there was a brief period of time wherein the pre- and post-turnover faunas coexist. [11] [12] [13] The Conques Formation ecosystem is one such example, being one of only two known occurrences of a rhabdodont, specifically Pareisactus , coexisting with hadrosaurs. [14] Contrastingly, the enormous Abditosaurus , also from the formation, is considered characteristic of post-turnover titanosaur faunas. [11] Fossils from temporally equivalent rocks of the Ibero-Armorican island include those tentatively referred to the theropods Arcovenator , Richardoestesia , Pyroraptor , intermediate remains referred to nodosaurid dinosaurs, indeterminate material from iguanid, anguid, Gekkota, and scincomorph lizards, indeterminate material from albanerpetontid amphibians, alytid and palaeobatrachid frogs, the bothremydid turtle Polysternon , and material tentatively referred to the crocodylomorph genera Allodaposuchus , Acynodon , Doratodon , and Sabresuchus . [12] [15]

Diagram of hadrosaur track depositional conditions, demonstrating the mudstone floodplains of meandering rivers (above) and braided rivers (below) hadrosaurs like Adynomosaurus lived in Hadrosaur tracks.png
Diagram of hadrosaur track depositional conditions, demonstrating the mudstone floodplains of meandering rivers (above) and braided rivers (below) hadrosaurs like Adynomosaurus lived in

The environment of the lower red mudstone unit (including the Conques Formation) of the Tremp Group has traditionally been considered fluvial floodplains. [16] [17] Sedimentary data associated with hadrosaur tracks preserved from the Tremp Group have been used to corroborate this. The ecosystem would have included abundant meandering rivers, with beds of either fine-grain or less often sand, as well as less common braided rivers, bedded with gravel. These rivers would have been interspersed across floodplains, possibly contiguous with estuarine conditions due to the proximity to marine environments (of the Tethys sea). [17] The abundance of mudstone, as well as other mineral and sediment features, indicates cyclic flooding and frequently fluctuating water levels in the ecosystem, which allowed the dinosaurs to leave tracks in the exposed wet mud which were then swiftly covered with water again to allow for preservation. [18] [17] In addition to the floodplains themselves, plant life would have colonized the abandoned rivers meanders and braid bars. [17] Carbon and oxygen isotope values from dinosaur eggshells also provided evidence of a wet environment, and found a mean air temperature for the ecosystem of 21 °C (70 °F). [19] A 2014 study proposed that the unit may have been more extensively marine-influenced than traditionally thought, something geologist Herbert Eisenscheer had previously proposed in a 1980s PhD thesis. In addition to sediment data, microfossils were investigated as an important source of evidence. Marine red algae and echinoderms among other marine microfossils were found to have been deposited into the river and mudflat environment. This was reasoned to have happened by tidal influences, transported along water channels after death; in some modern environments tidal forces influences rivers and transports material as much as 80 km inland. Sandy tidal flats may have been present, transitioning gradually into the mud sediments beyond the reach of high tide. [16] Thus the formation has been considered to represent a tide-dominated delta alongside contiguous inland tidal-influenced meandering river channels, acting as a transitional ecosystem between fully freshwater and fully marine environments. [18] [16] [20]

See also

Related Research Articles

<span class="mw-page-title-main">Hadrosauridae</span> Extinct family of dinosaurs

Hadrosaurids, or duck-billed dinosaurs, are members of the ornithischian family Hadrosauridae. This group is known as the duck-billed dinosaurs for the flat duck-bill appearance of the bones in their snouts. The ornithopod family, which includes genera such as Edmontosaurus and Parasaurolophus, was a common group of herbivores during the Late Cretaceous Period. Hadrosaurids are descendants of the Late Jurassic/Early Cretaceous iguanodontian dinosaurs and had a similar body layout. Hadrosaurs were among the most dominant herbivores during the Late Cretaceous in Asia and North America, and during the close of the Cretaceous several lineages dispersed into Europe, Africa, and South America.

<i>Aralosaurus</i> Extinct genus of dinosaurs

Aralosaurus was a genus of hadrosaurid dinosaur that lived during the Late Cretaceous in what is now Kazakhstan. It is known only by a posterior half of a skull and some post-cranial bones found in the Bostobe Formation in rocks dated from the Upper Santonian-Lower Campanian boundary, at about 83.6 Ma. Only one species is known, Aralosaurus tuberiferus, described by Anatoly Konstantinovich Rozhdestvensky in 1968. The genus name means Aral Sea lizard, because it was found to the northeast of the Aral Sea. The specific epithet tuberiferus means bearing a tuber because the posterior part of the nasal bone rises sharply in front of the orbits like an outgrowth. Aralosaurus was originally reconstituted with a nasal arch similar to that of North American Kritosaurus. For many years, Aralosaurus was thus placed in the clade of the Hadrosaurinae. This classification was invalidated in 2004, following the re-examination of the skull of the animal which allowed to identify in Aralosaurus many typical characters of Lambeosaurinae. In particular, this study revealed that Aralosaurus had a hollow bony structure located far in front of the orbits, which communicated with the respiratory tract. This structure being broken at its base, its shape and size remains undetermined. More recently, Aralosaurus has been identified as the most basal Lambeosaurinae, and placed with its close relative Canardia from the upper Maastrichtian of France in the new clade of Aralosaurini.

<i>Ampelosaurus</i> Extinct genus of dinosaurs

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<i>Amurosaurus</i> Extinct genus of dinosaurs

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<i>Tsintaosaurus</i> Hadrosaurid dinosaur from the Late Cretaceous

Tsintaosaurus is a genus of hadrosaurid dinosaur from China. It was about 8.3 metres (27 ft) long and weighed 2.5 tonnes. The type species is Tsintaosaurus spinorhinus, first described by Chinese paleontologist C. C. Young in 1958.

<i>Pararhabdodon</i> Extinct genus of dinosaurs

Pararhabdodon is a genus of tsintaosaurin hadrosaurid dinosaur, from the Maastrichtian-age Upper Cretaceous Tremp Group of Spain. The first remains were discovered from the Sant Romà d’Abella fossil locality and assigned to the genus Rhabdodon, and later named as the distinct species Pararhabdodon isonensis in 1993. Known material includes assorted postcranial remains, mostly vertebrae, as well as maxillae from the skull. Specimens from other sites, including remains from France, a maxilla previously considered the distinct taxon Koutalisaurus kohlerorum, an additional maxilla from another locality, the material assigned to the genera Blasisaurus and Arenysaurus, and the extensive Basturs Poble bonebed have been considered at different times to belong to the species, but all of these assignments have more recently been questioned. It was one of the last non-avian dinosaurs known from the fossil record that went extinct during the Cretaceous-Paleogene extinction event.

<span class="mw-page-title-main">Lambeosaurinae</span> Extinct subfamily of dinosaurs

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<i>Koutalisaurus</i> Extinct genus of dinosaurs

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<span class="mw-page-title-main">Tremp Formation</span>

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<i>Blasisaurus</i> Extinct genus of dinosaurs

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<i>Latirhinus</i> Extinct genus of reptiles

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<i>Canardia</i> Extinct genus of dinosaurs

Canardia is an extinct genus of lambeosaurine dinosaur known from the Late Cretaceous Marnes d'Auzas Formation of Haute-Garonne department, in Occitanie region, southwestern France. The type species Canardia garonnensis was first described and named by Albert Prieto-Márquez, Fabio M. Dalla Vecchia, Rodrigo Gaete and Àngel Galobart in 2013. It is only known from juvenile specimens. The name of the genus comes from “canard”, the French word for “duck”, an allusion to the fact that this animal belongs to the hadrosaurids which are also known as duck-billed dinosaurs. The specific epithet garonnensis refers to the Haute-Garonne department where this dinosaur has been found. Although universally recognized as a lambeosaurine, its precise position within them is debated. Some authors consider it as a close relative of the genus Aralosaurus from Central Asia with which it would form the tribe Aralosaurini, while others include it in a more derived clade, the Arenysaurini in which all lambeosaurines from Europe and North Africa are placed. Canardia was one of the last non-avian dinosaurs and lived between 67,5 and 66 my on the former Ibero-Armorican Island, which included much of France and Spain.

<span class="mw-page-title-main">Timeline of hadrosaur research</span>

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<span class="mw-page-title-main">Basturs Poble bonebed</span>

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<i>Ajnabia</i> Extinct genus of reptiles

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<i>Tamarro</i> Genus of troodontid dinosaur from the Late Cretaceous period

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References

  1. 1 2 3 4 5 6 7 8 9 Prieto-Márquez, Albert; Fondevilla, Víctor; Sellés, Albert G.; Wagner, Jonathan R.; Galobart, Àngel (2019). "Adynomosaurus arcanus, a new lambeosaurine dinosaur from the Late Cretaceous Ibero-Armorican Island of the European Archipelago". Cretaceous Research. 96: 19–37. Bibcode:2019CrRes..96...19P. doi:10.1016/j.cretres.2018.12.002. S2CID   134582286.
  2. ↑ "Society of Vertebrate Paleontology 78th Annual Meeting" (PDF). Vertpaleo.org. 2018. Retrieved 29 May 2022.
  3. ↑ Prieto-Márquez, A.; Dalla Vecchia, F. M.; Gaete, R.; Galobart, À. (2013). "Diversity, Relationships, and Biogeography of the Lambeosaurine Dinosaurs from the European Archipelago, with Description of the New Aralosaurin Canardia garonnensis" (PDF). PLOS ONE. 8 (7): e69835. Bibcode:2013PLoSO...869835P. doi: 10.1371/journal.pone.0069835 . PMC   3724916 . PMID   23922815.
  4. ↑ Horner, J.A.; Weishampel, D.B.; Forster, C.A. (2004). "Hadrosauridae". In Weishampel, David B.; Osmólska, Halszka; Dodson, Peter (eds.). The Dinosauria (Second ed.). University of California Press. pp. 438–463. ISBN   978-0-520-24209-8.
  5. ↑ Rozadilla, Sebastián; Brissón-Egli, Federico; Lisandro Agnolín, Federico; Aranciaga-Rolando, Alexis Mauro; Novas, Fernando Emilio (2022). "A new hadrosaurid (Dinosauria: Ornithischia) from the Late Cretaceous of northern Patagonia and the radiation of South American hadrosaurids". Journal of Systematic Palaeontology. 19 (17): 1207–1235. doi:10.1080/14772019.2021.2020917. S2CID   247122005.
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  7. ↑ Gates, Terry A.; Evans, David C.; Sertich, Joseph J. W. (2021). "Description and rediagnosis of the crested hadrosaurid (Ornithopoda) dinosaur Parasaurolophus cyrtocristatus on the basis of new cranial remains". PeerJ. 9: e10669. doi: 10.7717/peerj.10669 . PMC   7842145 . PMID   33552721.
  8. 1 2 Longrich, Nicholas R.; Suberbiola, Xabier Pereda; Pyron, R. Alexander; Jalil, Nour-Eddine (2020). "The first duckbill dinosaur (Hadrosauridae: Lambeosaurinae) from Africa and the role of oceanic dispersal in dinosaur biogeography". Cretaceous Research. 120: 104678. Bibcode:2021CrRes.12004678L. doi: 10.1016/j.cretres.2020.104678 . S2CID   228807024.
  9. ↑ Serrano-Brañas, Claudia Inés; Prieto-Márquezc, Albert (2022). "Taphonomic attributes of the holotype of the lambeosaurine dinosaur Latirhinus uitstlani from the late Campanian of Mexico: Implications for its phylogenetic systematics". Journal of South American Earth Sciences. 114: 103689. Bibcode:2022JSAES.11403689S. doi:10.1016/j.jsames.2021.103689.
  10. ↑ Ramírez-Velasco, Angel Alejandro (2022). "Phylogenetic and biogeography analysis of Mexican hadrosauroids". Cretaceous Research. 138: 105267. Bibcode:2022CrRes.13805267R. doi:10.1016/j.cretres.2022.105267. S2CID   249559319.
  11. 1 2 Vila, Bernat; Sellés, Albert; Moreno-Azanza, Miguel; Razzolini, Novella L.; Gil-Delgado, Alejandro; Canudo, José Ignacio; Galobart, Àngel (2022). "A titanosaurian sauropod with Gondwanan affinities in the latest Cretaceous of Europe". Nature Ecology & Evolution. 6 (3): 288–296. Bibcode:2022NatEE...6..288V. doi:10.1038/s41559-021-01651-5. PMID   35132183. S2CID   246650381.
  12. 1 2 Fondevilla, V.; Riera, V.; Vila, B.; Sellés, A.G.; Dinarès-Turell, J.; Vicens, E.; Gate, R.; Galobart, À. (2019). "Chronostratigraphic synthesis of the latest Cretaceous dinosaur turnover in south-western Europe". Earth-Science Reviews. 191: 168–189. Bibcode:2019ESRv..191..168F. doi:10.1016/j.earscirev.2019.01.007. S2CID   135231891.
  13. ↑ Vila, Bernat; Sellés, Albert G.; Brusatte, Stephen L. (2016). "Diversity and faunal changes in the latest Cretaceous dinosaur communities of southwestern Europe". Cretaceous Research. 57: 552–568. Bibcode:2016CrRes..57..552V. doi: 10.1016/j.cretres.2015.07.003 .
  14. ↑ Párraga, Javier; Prieto-Márquez, Albert (2019). "Pareisactus evrostos, a new basal iguanodontian (Dinosauria: Ornithopoda) from the Upper Cretaceous of southwestern Europe". Zootaxa. 4555 (2): 247–258. doi:10.11646/zootaxa.4555.2.5. PMID   30790960. S2CID   73469628.
  15. ↑ Pérez-Pueyo, Manuel; Cruzado-Caballero, Penélope; Moreno-Azanza, Miguel; Vila, Bernat; Castanera, Diego; Gasca, José Manuel; Puértolas-Pascual, Eduardo; Bádenas, Beatriz; Canudo, José Ignacio (2021). "The Tetrapod Fossil Record from the Uppermost Maastrichtian of the Ibero-Armorican Island: An Integrative Review Based on the Outcrops of the Western Tremp Syncline (Aragón, Huesca Province, NE Spain)". Geosciences. 11 (4): 162. Bibcode:2021Geosc..11..162P. doi: 10.3390/geosciences11040162 . hdl: 10362/145947 .
  16. 1 2 3 Díez-Canseco, D.; Arz, J.A.; Bentio, M.I.; Díaz-Molina, M; Arenillas, I. (2014). "Tidal influence in redbeds: A palaeoenvironmental and biochronostratigraphic reconstruction of the Lower Tremp Formation (South-Central Pyrenees, Spain) around the Cretaceous/Paleogene boundary". Sedimentary Geology. 312: 31–49. Bibcode:2014SedG..312...31D. doi:10.1016/j.sedgeo.2014.06.008.
  17. 1 2 3 4 Vila, Bernat; Oms, Oriol; Fondevilla, Víctor; Gaete, Rodrigo; Galobart, Àngel; Riera, Violeta; Canudo, José Ignacio (2013). "The Latest Succession of Dinosaur Tracksites in Europe: Hadrosaur Ichnology, Track Production and Palaeoenvironments". PLOS ONE. 8 (9): e72579. Bibcode:2013PLoSO...872579V. doi: 10.1371/journal.pone.0072579 . PMC   3760888 . PMID   24019873.
  18. 1 2 Puértolas-Pascual, E.; Arenillas, I.; Arz, J.A.; Calvín, P.; Ezquerro, L.; García-Vicente, C.; Pérez-Pueyo, M.; Sánchez-Moreno, E.M.; Villalaín, J.J.; Canudo, J.I. (2018). "Chronostratigraphy and new vertebrate sites from the upper Maastrichtian of Huesca (Spain), and their relation with the K/Pg boundary". Cretaceous Research. 89: 36–59. Bibcode:2018CrRes..89...36P. doi:10.1016/j.cretres.2018.02.016. S2CID   134852399.
  19. ↑ Riera, V.; Anadón, P; Oms, O.; Estraded, R.; Maestro, E. (2013). "Dinosaur eggshell isotope geochemistry as tools of palaeoenvironmental reconstruction for the upper Cretaceous from the Tremp Formation (Southern Pyrenees)". Sedimentary Geology. 294: 356–370. Bibcode:2013SedG..294..356R. doi:10.1016/j.sedgeo.2013.06.001.
  20. ↑ Gómez-Gras, D.; Roigé, M.; Fondevilla, V.; Oms, O.; Boya, S.; Remacha, E. (2016). "Provenance constraints on the Tremp Formation paleogeography (southern Pyrenees): Ebro Massif VS Pyrenees sources". Cretaceous Research. 57: 414–427. Bibcode:2016CrRes..57..414G. doi:10.1016/j.cretres.2015.09.010.
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