La Huérguina Formation

The La Huérguina Formation (also known as the Calizas de La Huérguina Formation, La Huérguina Limestone Formation or as the Una Formation) is a geological formation in Spain whose strata date back to the Barremian stage of the Early Cretaceous.^[1] Las Hoyas is a Konservat-Lagerstätte within the formation, located near the city of Cuenca, Spain. The site is mostly known for its exquisitely preserved dinosaurs, especially enantiornithines.^[2] The lithology of the formation mostly consists of lacustrine limestone deposited in a freshwater wetland environment.

La Huérguina Formation
Stratigraphic range: Barremian ~125 Ma PreꞒ Ꞓ O S D C P T J K Pg N ↓
Lithographic limestones of the Las Hoyas locality
Type	Geological formation
Underlies	Contreras Formation or Unconformity with the Utrillas Group
Overlies	Tragacete Formation
Thickness	60–100 m (200–330 ft)
Lithology
Primary	Limestone, marl
Other	Conglomerate
Location
Coordinates	40.0897°N 1.8978°W / 40.0897; -1.8978
Region	Cuenca, Castile-La Mancha
Country	Spain
Extent	South Iberian Basin
Type section
Named for	Huérguina
La Huérguina Formation (Spain) Show map of Spain La Huérguina Formation (Castilla-La Mancha) Show map of Castilla-La Mancha

Las Hoyas

Taphonomy

As a Konservat-Lagerstätten, the preservation is exceptional. This may be a result of three factors: Microbial mats, Obruption and Stagnation.

Microbial mats may be responsible for the preservation of soft tissue in many fossils from Las Hoyas, like Pelecanimimus' crest. The iron carbonate depositions, a result from bacterial metabolism which covered the dinosaur's crest enhanced the preservation of those soft tissues. Evidence of these mats comes from the studies on microfacies and the fossils themselves.

Obruption is notable in the formation, due to the presence of highly articulated specimens. From actuotaphonomy studies on several different organisms it can be estimated that the burial of most entities was quick. Concornis may have been buried in less than 15 days, after a period of sub-areal exposition.

Paleoenvironment

Las Hoyas was an inland lacustrine environment which presents an important aquatic and terrestrial flora (with many specimens of Charophytes, Montsechia, Weischelia or Frenelopsis) and diverse fauna, with specimens of at least five or six Phyla: arthropods, molluscs, Chordata and many vermiform soft bodied animals which might be Nemertines or annelids.

Among vertebrates the most abundant and diverse group are fish. The presence of mostly articulated skeletons, exceptional preservation of tissue and lack of any other signs of transportation may indicate that these are demic and autochthonous entities (meaning that they lived and died in the same place where they fossilized).

Crocodylomorphs are the most abundant amniotes from Las Hoyas.

Dinosaurs from Las Hoyas (avian and non avian) are unique in many ways. The first ornithomimosaur dinosaur described in Europe, Pelecanimimus polyodon, shows some characters previously unknown in these dinosaurs which enhanced the knowledge on the evolution of the group, such as a high number of teeth. Concavenator corcovatus presents two unique features: very tall neural spines on the vertebrae near the hip, which look like a hump, and a structure on its forearm, which if homologous to quill knobs would push back the origin of feathers earlier in theropod evolution.

Las Hoyas birds are enantiornithes, the most diverse Cretaceous bird clade, which became extinct at the end of the period. Iberomesornis romerali shows both derived ("avian") and primitive ("dinosaurian") characters. Within the derived characters we can underline the presence of a pygostile, although it is still very large compared to that of modern neornithines and the presence of a quilled sternum. Eoalulavis hoyasi shows the first report of an alula or "bastard wing", which means it had a flight manoeuvrability analogous to that of modern birds.

Research

The Las Hoyas site has been studied for more than two decades by researchers from the Universidad Autónoma de Madrid, the Universidad Complutense de Madrid and the National University of Distance Education, in collaboration with the Museo de las Ciencias de Castilla-La Mancha, which is responsible for the fossil record from the area.

Fossil content

Color key Taxon Reclassified taxon Taxon falsely reported as present Dubious taxon or junior synonym Ichnotaxon Ootaxon Morphotaxon		Notes Uncertain or tentative taxa are in small text; crossed out taxa are discredited.

Crustaceans

Crustaceans
Genus	Species	Location	Stratigraphic position	Material	Notes	Images
Austropotamobius	A. llopisi	Las Hoyas		Numerous specimens[3]	A crayfish
Delclosia	D. martinelli				An atyid shrimp
Lahuerguinagrapta[4]	L. multicostata				A clam shrimp belonging to the family Afrograptidae
Skyestheria[4]	Indeterminate				A clam shrimp belonging to the family Polygraptidae
Pseudestherites[4]					A clam shrimp belonging to the family Antronestheriidae
Spinogriphus [5]	S. ibericus				A member of Peracarida belonging to Spelaeogriphacea

Chondrichthyes

Chondrichthyes
Genus	Species	Location	Stratigraphic position	Material	Notes	Images
Lonchidion[6]	Indeterminate	Las Hoyas		A full body impression of a juvenile	Hybodont

Osteichthyes

Osteichthyes
Genus	Species	Location	Stratigraphic position	Material	Notes	Images
Caturus	Caturus sp.	Las Hoyas			Amiiformes
Gordichthys	G. conquensis	Las Hoyas			Gonorynchiform
Hispanamia[7]	H. newbreyi	Las Hoyas			Amiid, originally described as Amiopsis
cf. "Holophagus"[8]	Indeterminate	Las Hoyas			Coelacanth
Hoyasotes	H. tanyrhis	Las Hoyas			Member of Lepidotidae, originally described as Lepidotes[9]
Lepidohyas	L. microrhis	Las Hoyas
Notagogus	N. aff. N. ferreri	Las Hoyas			Macrosemiid
Pleuropholis	P. sp.	Las Hoyas
Propterus	P. sp.	Las Hoyas			Macrosemiid
Rubiesichthys	R. gregalis	Las Hoyas			Gonorynchiform
Stenamara	S. mia	Las Hoyas			Pycnodont
Turbomesodon	T. praeclarus	Las Hoyas
Vidalamia	V. cf. catalunica	Las Hoyas			Amiid

Amphibians

Amphibians
Genus	Species	Location	Stratigraphic position	Material	Notes	Images
Albanerpeton	Indeterminate	Uña		Frontals	Albanerpetontid
Celtedens	C. ibericus	Las Hoyas
	C. megacephalus	Uña
Gracilibatrachus	G. avallei	Las Hoyas			Pipimorph frog
Hylaeobatrachus	Indeterminate	Las Hoyas			Salamander
Iberobatrachus	I. angelae	Las Hoyas			Frog
Valdotriton	V. gracilis	Las Hoyas			Salamander
Wealdenbatrachus	W. jucarensis	Uña			Frog

Turtles

Turtles
Genus	Species	Location	Stratigraphic position	Material	Notes	Images
Hoyasemys	H. jimenezi	Las Hoyas			A eucryptodire[10]
Pleurosternidae	Indeterminate	Uña
Pelomedusidae	Indeterminate	Uña

Squamates

Squamates
Genus	Species	Location	Stratigraphic position	Material	Notes	Images
Becklesius	Indeterminate, B. cataphractus	Uña			Paramacellodid lizard
Cuencasaurus	C. estesi	Uña
Hoyalacerta	H. sanzi	Las Hoyas
Jucaraseps	J. grandipes	Las Hoyas			Scleroglossa
Meyasaurus	M. diazromerali	Las Hoyas			teiioid lizard
	M. unaensis	Uña
Paramacellodus	P. sinuosus	Uña			Paramacellodid lizard
Scandensia	S. ciervensis	Las Hoyas

Mammals

Mammals
Genus	Species	Location	Stratigraphic position	Material	Notes	Images
Crusafontia	C. cuencana	Uña			Dryolestid
Eobaatar	E. hispanicus	Uña			Eobaatarid multituberculate
Galveodon	G. nannothus	Uña			Paulchoffatiid multituberculate
Spinolestes[11]	S. xenarthrosus	Las Hoyas		Exceptionally complete specimen, preserving soft tissues such as pelage, ears and internal organs.	Gobiconodont Eutriconodont

Crocodyliformes

Crocodyliformes
Genus	Species	Location	Stratigraphic position	Material	Notes	Images
Bernissartia	Indeterminate	Uña
Cassissuchus	C. sanziuami	Las Hoyas			Gobiosuchid
Montsecosuchus	Indeterminate	Las Hoyas
Sabresuchus	S.ibericus	Uña
Unasuchus	U. reginae	Uña
Bernissartiidae	Indeterminate	Las Hoyas

Ornithodirans

Ornithodirans
Genus	Species	Location	Stratigraphic position	Material	Notes	Images
Concavenator	C. corcovatus	Las Hoyas		"Nearly complete and articulated skeleton."	Carcharodontosaurid dinosaur[12]
Concornis[13]	C. lacustris[13]	Las Hoyas		"Postcranial skeleton."[14]	enantiornithean bird
Eoalulavis[13]	E. hoyasi[13]	Las Hoyas		"Thoracic region and forelimbs."[14]	enantiornithean bird
Euronychodon	Indeterminate	Las Hoyas			[15]
Europejara	E. olcadesorum	Las Hoyas			Tapejarid pterosaur
Iberomesornis	I. romerali	Las Hoyas		"Postcranial skeleton."	enantiornithean bird[13][16]
Mantellisaurus	M. atherfieldensis	Las Hoyas		"incomplete but fully-articulated right hindlimb"	Iguanodontian dinosaur[17]
Pelecanimimus	P. polyodon	Las Hoyas		"Skull and partial skeleton"	Ornithomimosaurian dinosaur[13][18]
Paronychodon	Indeterminate	Las Hoyas			[19]
Ricardoestesia	Indeterminate	Las Hoyas			[20]

Correlation

Early Cretaceous stratigraphy of Iberia

Ma	Age	Paleomap \ Basins	Cantabrian	Olanyà	Cameros	Maestrazgo	Oliete	Galve	Morella	South Iberian	Pre-betic	Lusitanian
100	Cenomanian		La Cabana	Sopeira	Utrillas		Mosquerela					Caranguejeira
			Altamira		Utrillas
			Eguino
125	Albian		Ullaga - Balmaseda	Lluçà						Traiguera
			Monte Grande		Escucha		Escucha			Jijona
			Itxina - Miono
	Aptian		Valmaseda - Tellamendi		Ol Gp. - Castrillo	Benassal		Benassal				Olhos
				Font	En Gp. - Leza	Morella/Oliete	Oliete	Villaroya	Morella	Capas Rojas		Almargem
			Patrocinio - Ernaga	Senyús	En Gp. - Jubela	Forcall				Villaroya	Upper Bedoulian	Figueira
	Barremian		Vega de Pas	Cabó	Abejar	Xert	Alacón	Xert		Huérguina		Assises
				Prada		Artoles				Collado	Moutonianum	Papo Seco
				Rúbies	Tera Gp. - Golmayo	Alacón/Blesa	Blesa	Camarillas	Mirambel
150	Hauterivian				Ur Gp. - Pinilla	Llacova		Castellar	Tera Gp. - Pinilla	Villares		Porto da Calada
							hiatus
						Huerva			Gaita
	Valanginian		Villaro		Ur Gp. - Larriba				Ped Gp. - Hortigüela
					Ped Gp. - Hortigüela				Ped Gp. - Piedrahita
					Peñacoba			Galve		Miravetes
	Berriasian		Cab Gp. - Arcera		Valdeprado	hiatus		Alfambra
					TdL Gp. - Rupelo			Arzobispo	hiatus	Tollo
					On Gp. - Huérteles Sierra Matute
	Tithonian		Lastres		Tera Gp. - Magaña		Higuereles			Tera Gp. - Magaña		Lourinhã
									Arzobispo
					Ágreda
Legend		Major fossiliferous, oofossiliferous, ichnofossiliferous, coproliferous, minor formation
Sources

See also

• List of dinosaur-bearing rock formations
• Paja Formation, contemporaneous Lagerstätte of Colombia
• Yixian Formation, contemporaneous Lagerstätte of China

References

1. Martínez et al., 2017
2. Weishampel et al., 2004, pp.556-563
3. Barrios-de Pedro, Sandra; Chin, Karen; Buscalioni, Ángela D. (June 2020). "The late Barremian ecosystem of Las Hoyas sustained by fishes and shrimps as inferred from coprofabrics". Cretaceous Research. 110: 104409. Bibcode:2020CrRes.11004409B. doi:10.1016/j.cretres.2020.104409. hdl:10486/695365. S2CID 213738761.
4. Gallego, Oscar F.; Monferran, Mateo D.; Zacarías, Iracema A.; Jiménez, Victoria C.; Buscalioni, Angela D.; Liao, Huanyu (June 2020). "Clam shrimp fauna (Diplostraca-Spinicaudata and Estheriellina) from the Lower Cretaceous of Las Hoyas, Cuenca (Spain)". Cretaceous Research. 110: 104389. Bibcode:2020CrRes.11004389G. doi:10.1016/j.cretres.2020.104389. S2CID 214213415.
5. Jaume, Damià; Pinardo-Moya, Eva; Boxshall, Geoff A. (January 2013). "A presumed spelaeogriphacean crustacean from an upper Barremian wetland (Las Hoyas; Lower Cretaceous; Central Spain)". Palaeontology. 56 (1): 15–28. Bibcode:2013Palgy..56...15J. doi:10.1111/j.1475-4983.2012.01144.x. ISSN 0031-0239.
6. Marugán-Lobón, Jesús; Martín-Abad, Hugo; Buscalioni, Ángela D. (2023-04-03). "The Las Hoyas Lagerstätte: a palaeontological view of an Early Cretaceous wetland". Journal of the Geological Society. 180 (3). doi:10.1144/jgs2022-079. hdl:10486/710760. ISSN 0016-7649.
7. MartÍn-Abad, Hugo; Poyato-Ariza, Francisco JosÉ (2017-06-05). "A new genus and species for the amiiform fishes previously assigned to Amiopsis from the Early Cretaceous of Las Hoyas, Cuenca, Spain". Zoological Journal of the Linnean Society. 181 (3): 604–637. doi:10.1093/zoolinnean/zlx010. ISSN 0024-4082.
8. Martín-Abad, Hugo; Martínez, Marian Fregenal- (2021-12-02). "The ecology of the Lower Cretaceous coelacanths from Las Hoyas Konservat- Lagerstätte (Cuenca, Spain): A new insight after the integration of palaeontological and sedimentological data". Spanish Journal of Palaeontology. 36 (2): 191–204. doi:10.7203/sjp.36.2.21966. ISSN 2660-9568.
9. Paiva, Hanna Carolina Lins de; Gallo, Valéria (2018-12-01). "Quasimodichthys gen. nov. (Neopterygii: Semionotiformes): A morphological and ontogenetic study". Journal of South American Earth Sciences. 88: 132–143. Bibcode:2018JSAES..88..132P. doi:10.1016/j.jsames.2018.08.010. ISSN 0895-9811. S2CID 134986658.
10. Pérez García et al., 2012
11. Martin et al., 2015
12. Ortega et al., 2010
13. "15.9 Provincia de Quenca, Spain; 4. Calizas de La Huergina Formation," in Weishampel et al., 2004, pp.561-562
14. "Table 11.1," in Weishampel et al. ,2004, p.212
15. Listed as "cf. Euronychodon sp." in "15.9 Provincia de Quenca, Spain; 4. Calizas de La Huergina Formation," in Weishampel et al., 2004, p.561
16. "Table 11.1," in Weishampel et al., 2004, p.213
17. Serrano et al., 2013
18. "Table 6.1," in Weishampel et al., 2004, p.138
19. Listed as "cf. Paranychodon sp." in "15.9 Provincia de Quenca, Spain; 4. Calizas de La Huergina Formation," in Weishampel et al., 2004, p.561
20. Listed as "cf. Ricardoestesia sp." in "15.9 Provincia de Quenca, Spain; 4. Calizas de La Huergina Formation," in Weishampel et al., 2004, p.561

Bibliography

• Martínez, Marian Fregenal; Meléndez, Nieves; Muñoz García, M. Belén; Elez, Javier; de la Horra, Raúl (2017), "The stratigraphic record of the Late Jurassic-Early Cretaceous rifting in the Alto Tajo-Serranía de Cuenca region (Iberian Ranges, Spain): genetic and structural evidences for a revision and a new lithostratigraphic proposal", Revista de la Sociedad Geológica de España, 30: 113–142, retrieved 2019-10-12
• Serrano, Mercedes Llandres; Vullo, Romain; Marugán-Lobón, Jesús; Ortega, Francisco; Buscalioni, Ángela D. (2013), "An articulated hindlimb of a basal iguanodont (Dinosauria, Ornithopoda) from the Early Cretaceous Las Hoyas Lagerstätte (Spain)", Geological Magazine, 150 (3): 572–576, doi:10.1017/S0016756813000095, ISSN 0016-7568, retrieved 2019-10-12
• Pérez García, Adán; Fuente, Marcelo S. de la; Ortega, Francisco (2012), "A New Freshwater Basal Eucryptodiran Turtle from the Early Cretaceous of Spain", Acta Palaeontologica Polonica, 57 (2): 285–298, doi:10.4202/app.2011.0031, ISSN 0567-7920, retrieved 2019-10-12
• Thomas Martin, Jesús Marugán-Lobón, Romain Vullo, Hugo Martín-Abad, Zhe-Xi Luo & Angela D. Buscalioni (2015). A Cretaceous eutriconodont and integument evolution in early mammals. Nature 526, 380–384. doi:10.1038/nature14905
• Ortega, F., Escaso, F. & Sanz, J. L. (2010): A bizarre, humped Carcharodontosauria (Theropoda) from the Lower Cretaceous of Spain Nature 467: 203–206. HTML abstract
• Weishampel, David B.; Dodson, Peter; Osmólska, Halszka (2004), The Dinosauria, 2nd edition, Berkeley: University of California Press, pp. 1–880, ISBN 0-520-24209-2, retrieved 2019-02-21
• Sanz, José L.; Chiappe, Luis M.; Pérez Moreno, Bernardino P.; Buscalioni, Ángela D.; Moratalla, José J.; Ortega, Francisco & Poyato Ariza, Francisco J. (1996): An Early Cretaceous bird from Spain and its implications for the evolution of avian flight. Nature 382(6590): 442–445. doi:10.1038/382442a0 (HTML abstract)
• Pérez Moreno, B. P., Sanz, J. L., Buscalioni, A. D., Moratalla, J. J., Ortega, F., and Raskin-Gutman, D. (1994). "A unique multitoothed ornithomimosaur from the Lower Cretaceous of Spain." Nature, 30: 363–367.
• Sanz, J. L. & Bonaparte, José F. (1992): A New Order of Birds (Class Aves) from the Lower Cretaceous of Spain. In: Jonathan J. Becker (ed.): Papers in Avian Paleontology Honoring Pierce Brodkorb. Natural History Museum of Los Angeles County Contributions in Science 36: 38–49.
• Sanz, J. L. & Buscalioni, A.D. (1992): A new bird from the Early Cretaceous of Las Hoyas, Spain, and the early radiation of birds. Palaeontology 35(4): 829–845. PDF fulltext

External links

• Las Hoyas site website Archived 2020-02-22 at the Wayback Machine