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Exceptional crocodylomorph biodiversity of "La Cantalera" site (lower Barremian; Lower Cretaceous) in Teruel, Spain

Eduardo Puértolas-Pascual, Raquel Rabal-Garcés, and José Ignacio Canudo

Plain Language Abstract

The palaeontological site of La Cantalera in Teruel (Spain) can be characterized as one the greatest sites for vertebrate biodiversity during the Lower Cretaceous of the Iberian Peninsula. There are 32 different kinds of vertebrates: amphibians, lizards, turtles, crocodiles, pterosaurs, dinosaurs and mammals. The most common remains are the isolated teeth of crocodiles. Six different types of crocodile teeth have been distinguished, which may correspond to at least four different species of crocodiles, possibly more, living in the same area and time. Nowadays, no more than two or three crocodile species occur simultaneously in the same area. This palaeontological site presents a similar faunal assemblage to other sites from the Lower Cretaceous of Europe, with crocodile teeth that may correspond to the families Goniopholididae (crocodiles with a similar generalist lifestyle to modern crocodilians), Bernissartiidae (small crocodiles with diets of tough prey), Atoposauridae (terrestrial dwarf crocodiles with probably insectivorous diets) and several ziphodonts (terrestrial carnivorous crocodiles with serrated teeth). The crocodile assemblage of La Cantalera is characterized by the absence of large individuals and presents a wide variety of dental morphologies, adapted to diets ranging from generalist to insectivorous and durophagous (tough foods like mollusks and crustaceans). This rich biodiversity could be due to concentration of vertebrates into restricted flooded areas during dry seasons within a marsh ecosystem. Furthermore, La Cantalera could also be close to a nesting area, which would explain the abundance of small-sized individuals and the presence of crocodile eggshells.

Resumen en Español

Biodiversidad excepcional de crocodylomorfos del sitio "La Cantalera" (Barremiano inferior, Cretácico Inferior) en Teruel, España

El sitio paleontológico de La Cantalera en Teruel, España (Formación Blesa, Cretácico Inferior) puede caracterizarse como el sitio con la mayor biodiversidad de vertebrados (32 taxones) del Barremiano inferior de la Península Ibérica. Se han recuperado restos de anfibios, lagartijas, tortugas, cocodrilos, pterosaurios, dinosaurios y mamíferos. Los taxones más comunes son crocodylomorfos, y sus restos más abundantes son dientes pequeños aislados. Se han identificado seis morfotipos, los cuales corresponderían a por lo menos cuatro taxones de crocodylomorfos simpátricos. Este sitio presenta una asociación similar a otros sitios europeos del Cretácico Inferior, con morfotipos que corresponderían a las familias Goniopholididae, Bernissartiidae, Atoposauridae y múltiples crocodylomorfos zifodontes. La asociación de crocodylomorfos de La Cantalera se caracteriza por la ausencia de individuos grandes y presenta una amplia variedad de morfologías dentarias adaptadas a dietas desde generalistas hasta altamente especializadas. Esta biodiversidad tan rica podría deberse a la concentración de vertebrados en áreas inundadas restringidas durante estaciones secas dentro de un ecosistema de manglar. Más aún, La Cantalera podría estar cerca de una área de anidamiento, lo cual explicaría la abundancia de individuos de pequeño tamaño y la presencia de cáscaras de huevo crocodiloides fósiles.

Palabras clave: Cretácico, Inferior; Crocodylomorpha; dientes; paleoecología; paleobiodiversidad; Cordillera Ibérica

Traducción: Diana Elizabeth Fernández

Résumé en Français

Biodiversité exceptionnelle de crocodylomorphes du site de « La Cantalera » (Barrémien inférieur ; Crétacé inférieur) à Teruel, Espagne

Le site paléontologique de « La Cantalera » (formation de Blesa, Crétacé inférieur), situé à Teruel en Espagne, peut être caractérisé comme le site ayant la plus grande biodiversité de vertébrés (32 taxons) du Barrémien inférieur de la péninsule ibérique. Des restes d'amphibiens, de lézards, de tortues, de crocodiles, de ptérosaures, de dinosaures, et de mammifères y ont été mis à jour. Les taxons les plus communs sont des crocodylomorphes, et leurs restes les plus abondants sont de petites dents isolées. Six morphotypes ont été identifiés et pourraient correspondre au minimum à quatre taxons différents et sympatriques de crocodylomorphes. Ce site présente un assemblage similaire à celui d'autres sites du Crétacé inférieur d'Europe, avec des morphotypes qui pourraient correspondre aux familles Goniopholididae, Bernissartiidae, Atoposauridae, ainsi que de multiples crocodylomorphes ziphodontes. L'assemblage de crocodylomorphes de « La Cantalera » est caractérisé par une absence d'individus de grande taille et présente une large variété de morphologies dentaires adaptées à des régimes alimentaires allant de généralistes à hautement spécialisés. Cette riche biodiversité pourrait être due à une concentration des vertébrés dans des zones inondées limitées pendant les saisons sèches au sein d'un écosystème de marais. De plus, « La Cantalera » pourrait également être proche d'une zone de nidification, ce qui expliquerait l'abondance des individus de petite taille et la présence de coquilles d'œufs fossiles de crocodiloïdes.

Mots-clés : Crétacé inférieur ; Crocodylomorpha ; dents ; paléoécologie ; paléobiodiversité ; Cordillère ibérique

Translator: Antoine Souron

Deutsche Zusammenfassung

Außergewöhnliche Biodiversität an Crocodylomorphen in der "La Cantalera"-Fundstelle (unteres Barremium; untere Kreide) von Teruel, Spanien

Die paläontologische Fundstelle La Cantalera in Teruel, Spanien (Blesa Formation, untere Kreide) kann als die Fundstelle mit der größten Biodiversität an Vertebraten (32 Taxa) im unteren Barremium der spanischen Halbinsel charakterisiert werden. Es wurden Überreste von Amphibien, Eidechsen, Schildkröten, Krokodilen, Pterosauriern, Dinosauriern und Säugetieren gefunden. Die häufigsten Taxa sind Crocodylomorphe, die mit zahlreichen kleinen isolierten Zähnen vorkommen. Es wurden sechs Morphotypen identifiziert, die mit wenigstens vier verschiedenen sympatrischen crocodylomorphen Taxa korrespondieren dürften. Diese Fundstelle zeigt eine ähnliche Assemblage wie andere europäische unterkretaztische Fundstellen, mit Morphotypen die mit den Familien Goniopholididae, Bernissartiidae, Atoposauridae und verschiedenen ziphodonten Crocodylomorphen übereinstimmen dürften. Die crocodylomorphe Assemblage von La Cantelara ist durch die Abwesenheit von großen Individuen charakterisiert und präsentiert eine breite Palette an dentalen Morphologien, die an generalisierte bis hoch spezialisierte Ernährungsweisen angepasst sind. Die hohe Biodiversität könnte daran liegen, dass die Wirbeltiere innerhalb eines Sumpfgebietes während der Trockenzeit in abgetrennten überschwemmten Gebieten konzentriert waren. Darüber hinaus könnte La Cantelara auch nahe eines Nistgebietes gelegen haben, was die Häufigkeit von kleinen Individuen und das Vorhandensein von fossilen crocodiloiden Eischalen erklären würde.

Schlüsselwörter: Kreide, untere; Crocodylomorpha; Zähne; Paläoökologie; Paläobiodiversität; Iberisches Gebirge

Translator: Eva Gebauer

Arabic

 

Translator: Ashraf M.T. Elewa

 

 

FIGURE 1. Geographical and geological setting of La Cantalera site (early Barremian, Teruel, Spain). 1, Geological map of the Iberian Peninsula with detailed location of the site of La Cantalera and its position within the Maestrazgo Basin (Ol: Oliete, Pa: Las Parras, Ga: Galve, Mo: Morella, Pe: Perelló, Sa: Salzedella, Pg: Peñagolosa). 2, Geological map of the area of La Cantalera. 3, Local geological units. 4, Stratigraphic column and position of La Cantalera. Modified from Canudo et al., 2010.

figure1a

 

FIGURE 2. General view of La Cantalera site, showing the gray clays of the lower part of the Blesa Formation.

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FIGURE 3. Crocodylomorph teeth attributed to Goniopholididae? from La Cantalera site. 1-3, MPZ 2014/468, 469 and 470; photographs taken with a binocular microscope in mesial/distal, labial, lingual, apical (above) and radicular (below) views. 4, photographs taken with a scanning electron microscope in labial (above, MPZ 2014/471a) and mesial/distal (below, MPZ 2014/471b) views. Scale bars equal 1 mm.

figure3

FIGURE 4. 1-3, MPZ 2014/472, 473, 474a and 474b; crocodylomorph teeth from La Cantalera site attributed to Bernissartiidae?; photographs taken with a binocular microscope (1,2) in mesial/distal, labial, lingual, apical (above) and radicular (below) views; photographs taken with a scanning electron microscope (3) in lingual (MPZ 2014/474a, above) and apical (MPZ 2014/474b, below) views. 4-6, MPZ 2014/475, 476, 477a, 477b and 477c; crocodylomorph teeth from La Cantalera site attributed to Bernissartiidae?, which may correspond to teeth from the anterior or mid region of the tooth row; photographs taken with a binocular microscope (4,5) in mesial/distal, labial, lingual, apical (above) and radicular (below) views; photographs taken with a scanning electron microscope (6) in labial (MPZ 2014/477a, left), mesial/distal (MPZ 2014/477b, right) and apical (MPZ 2014/477c, below) views. Scale bars equal 1 mm.

figure4

FIGURE 5. Crocodylomorph teeth attributed to Theriosuchus sp. from La Cantalera site. 1-4, MPZ 2014/478, 479, 480 and 481; photographs taken with a binocular microscope in mesial/distal, labial, lingual, apical (above) and radicular (below) views. 5, MPZ 2014/482a, 2014/482b and 2014/482c; photographs taken with a scanning electron microscope in mesial/distal (2014/482a, above), lingual (2014/482a, middle) and apical (2014/482a, below) views. Scale bars equal 1 mm.

figure5

FIGURE 6. 1, MPZ 2014/483; ziphodont crocodylomorph tooth attributed to Mesoeucrocodylia indet. in mesial/distal, labial, lingual (with detail of the denticles), apical (above) and radicular (below) views. 2, MPZ 2014/484; bizarre crocodylomorph tooth attributed to Mesoeucrocodylia indet. in mesial/distal (arrows show the bifurcated keel), labial, lingual, apical (above) and radicular (below) views. All photographs taken with a binocular microscope. Scale bars equal 1 mm.

figure6

 

APPENDIX - Teeth measurements

Available in Excel and text format.

 

 

pascualEduardo Puértolas-Pascual
Grupo Aragosaurus-IUCA
Departamento de Ciencias de la Tierra
Facultad de Ciencias, Universidad de Zaragoza
Calle Pedro Cerbuna 12, 50009 Zaragoza
Spain
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Eduardo Puértolas Pascual has a degree in geology from the University of Zaragoza (2009), Master's in Introduction to Research in Geology (2010) and currently he is a Ph.D. student doing his doctoral thesis focused on the study of the systematics, biodiversity and paleobiogeography of the crocodylomorphs from the Cretaceous of Europe and Spain.

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rabal garcesRaquel Rabal-Garcés
Grupo Aragosaurus-IUCA
Departamento de Ciencias de la Tierra
Facultad de Ciencias, Universidad de Zaragoza
Calle Pedro Cerbuna 12, 50009 Zaragoza
Spain
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Dra. Raquel Rabal Garcés has a degree in geology from the University of Zaragoza (2006), Master's in Introduction to Research in Geology (2008) and PhD in Geology from the University of Zaragoza (2013). In her thesis, entitled "Estudio paleontológico de Ursus spelaeus Rosenmüller, 1794 del Pleistoceno Superior de Coro Tracito (Tella, Huesca, España)" conducted a detailed study of the taphonomy, systematic paleontology and paleoecology of fossil remains of Ursus spelaeus in order to characterize the population.

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canudoJosé Ignacio Canudo
Grupo Aragosaurus-IUCA
Departamento de Ciencias de la Tierra
Facultad de Ciencias, Universidad de Zaragoza
Calle Pedro Cerbuna 12, 50009 Zaragoza
Spain
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Dr. Jose Ignacio Canudo has a degree in Geology and a PhD in Sciences (Palaeontology) from the University of Zaragoza (UZ). Currently he is Full Professor in Paleontology (University of Zaragoza). In the last decade he has focused his research on the Mesozoic, with three lines of research. 1) Study of palaeobiodiversity of continental vertebrates from the Jurassic and Cretaceous of Iberia. 2) Paleobiogeography of dinosaurs from the Iberian Peninsula during the Jurassic and Cretaceous, including its relations with Gondwana (Argentina) and 3) Study of the extinction of the Cretaceous - Tertiary in the Iberian Peninsula. 

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TABLE 1. Faunal assemblage of La Cantalera. Colors show the ecological affinity of each taxon, aquatic in blue, amphibious in green and terrestrial in gray. Modified from Canudo et al., 2010 and Ruiz-Omeñaca et al., 1997.

Class Order/Superorder Sub/Infraorder Taxa
Charophyceae Charales Quinquespiralia Atopochara trivolvis triquetra Grambast, 1968
     

Pseudoglobator sp.

Gastropoda Caenogastropoda   Viviparidae indet.
  Basomatophora   Basomatophora indet.
Ostracoda Podocopida Podocopina Cypridea (Ulwellia) soriana Kneuper-Haack, 1966
Osteichtyes     Teleostei indet.
Lissamphibia     Albanerpetonidae indet.
  Anura   Discoglossidae indet.
“Reptilia” Chelonia   Pleurosternidae? indet.
  Crocodylomorpha   Goniopholididae?
      Bernissartiidae?
     

Theriosuchus sp.

      Mesoeucrocodylia indet.
  Squamata Lacertilia Paramacellodidae? indet.
  Pterosauria   Istiodactylidae? indet.
      Ornithocheiridae? indet.
  Ornithischia Ornithopoda Iguanodontoidea indet.
      Hadrosauroidea? indet.
      “Hypsilophodontidae” indet.
      Rhabdodontidae? indet.
    Thyreophora Polacanthidae indet.
  Saurischia Sauropoda Euhelopodidae indet
    Theropoda Baryonychinae indet.
      Carcharodontosauridae? indet.
      aff. Paronychodon sp.
      Maniraptora indet. 1
      Maniraptora indet. 2
      Maniraptora indet. 3
      Dromaeosaurinae indet. 1
      Dromaeosaurinae indet. 2
      Dromaeosaurinae indet. 3
      Velociraptorinae indet.
      Avialae? indet.
Mammalia Multituberculata   Cantalera abadi Badiola, Canudo and Cuenca-Bescós, 2008
      Eobaatar sp.
      “Plagiaulacida” indet.
      Plagiaulacidae/Eobaataridae indet.