Volume 2 Quaternary Chapter 11 Neogene https://doi.org/10.32685/pub.esp.36.2019.11 Dinosaur Footprints from the Lower Cretaceous, Published online 25 November 2020 Batá Formation, Colombia (South America), and the Possible Interchange of Large Paleogene Ornithopods between Southern Laurasia and Northern Gondwana 1 [email protected] Universidad de los Andes Cretaceous Facultad de Ciencias Leslie F. NOÈ1* , Marcela GÓMEZ–PÉREZ2 , José Vicente RODRÍGUEZ3 , Departamento de Geociencias Carrera 1 n.° 18A–12 4 5 Alejandro CORRALES–GARCÍA , and William G. CARANTON–MATEUS Bogotá, Colombia 2 [email protected] Servicio Geológico Colombiano Abstract Dinosaur remains from northwestern South America are rare, with only ex- Jurassic Dirección de Geociencias Básicas tremely scarce fossil evidence recovered from Colombia. Here we report six dinosaur Diagonal 53 n.° 34–53 footprints preserved on a sub–vertical bedding plane of the upper Valanginian – lower Bogotá, Colombia 3 [email protected] Hauterivian Batá Formation, Santa María, Boyacá Department, Colombia. The Batá Universidad de los Andes Formation consists of a thick succession of conglomerates and sandstones with shale Facultad de Ciencias Departamento de Geociencias Triassic intercalations interpreted as deposited along the palaeoshoreline of an epicontinental Carrera 1 n.° 18A–12 seaway. Four of the footprints form a trackway made by a single dinosaur, which is Bogotá, Colombia 4 [email protected] interpreted as a sub–adult ornithopod, estimated at 8 m in length, weighing around Universidad de los Andes 2.5 metric tons, and travelling at an average walking speed of almost 5 km/h. The foot- Facultad de Ciencias Departamentos de Geociencias y Biología prints are assigned to the ichnogenus Iguanodontipus, and were probably produced Carrera 1 n.° 18A–12 Permian Bogotá, Colombia by an iguanodontian dinosaur. Prior to this work, Iguanodontipus was considered an 5 [email protected] exclusively European taxon, making this a unique record of the ichnogenus in Gondwa- Universidad de los Andes Facultad de Ciencias na. The presence of Iguanodontipus in northern South America suggests an Early Cre- Departamento de Geociencias taceous sweepstake, with dinosaurs crossing Tethys Ocean into modern–day northern Carrera 1 n.° 18A–12 Bogotá, Colombia Africa, and migrating along the northern shores of Gondwana into modern–day South Carboniferous * Corresponding author America. Range extension of iguanodontian ornithopods southwards into Gondwana during the Early Cretaceous was apparently prevented by the Central Gondwana Desert Belt, possibly as a result of the palaeoecology of these dinosaurs, which seem to have had an affinity for environments rich in water and lush vegetation. A migration route Devonian across Tethys and the Central Gondwana Desert Belt helps explain similarities between northern Gondwanan and southern Laurasian dinosaurs, and the differences between northern and southern Gondwanan faunas, during the Early Cretaceous. Keywords: dinosaur, ichnofossils, Lower Cretaceous, Gondwana, Laurasia, faunal interchange. Silurian Resumen Los restos de dinosaurio del noroeste de Suramérica son raros, con muy pocas evidencias fósiles recuperadas en Colombia. Aquí reportamos seis huellas de dinosaurio preservadas en una capa subvertical del Valanginiano superior–Hauteriviano inferior de la Formación Batá, Santa María, departamento de Boyacá, Colombia. La Formación Ordovician Citation: Noè, L.F., Gómez–Pérez, M., Rodríguez, J.V., Corrales–García, A. & Caranton–Mateus, W.G. 2020. Dinosaur footprints from the Lower Cretaceous, Batá Formation, Colombia (South America), and the possible interchange of large ornithopods between southern Laurasia and northern Gondwana. In: Gómez, J. & Pinilla–Pachon, A.O. (editors), The Geology of Colombia, Cambrian Volume 2 Mesozoic. Servicio Geológico Colombiano, Publicaciones Geológicas Especiales 36, p. 375–401. Bogotá. https://doi.org/10.32685/pub.esp.36.2019.11 375 Proterozoic NOÈ et al. Batá consiste en una secuencia espesa de conglomerados y areniscas con intercalacio- nes de lodolitas interpretadas como depósitos de la línea de costa de un antiguo mar epicontinental. Cuatro de las huellas forman una pista dejada por un único dinosaurio, interpretado como un ornitópodo subadulto, con una longitud estimada de 8 m, un peso de 2,5 toneladas métricas y que viajaba a un ritmo normal de casi 5 km/h. Las huellas se asignaron al icnogénero Iguanodontipus, y fueron probablemente hechas por un dino- saurio tipo iguanodontiano. Antes de este trabajo, Iguanodontipus se consideraba como un taxón exclusivamente europeo, por lo que este registro sería el único en Gondwana. La presencia de Iguanodontipus en el norte de Suramérica sugiere la existencia de una comunicación terrestre durante el Cretácico Temprano, con dinosaurios cruzando el océano Tetis hacia el norte de África actual, y migrando a lo largo de la costa norte de Gondwana hasta lo que hoy es Suramérica. La extensión del rango de los ornitópodos iguanodontes hasta el sur de Gondwana durante el Cretácico Temprano no ocurrió de- bido a la presencia del Cinturón del Desierto de Gondwana Central, posiblemente como un resultado de la paleoecología de los ornitópodos, los cuales tenían afinidad por el agua y la vegetación exuberante. Una ruta de migración a través del Tetis y una barrera en el Cinturón del Desierto de Gondwana Central explicarían las similitudes entre los dinosaurios del norte de Gondwana y el sur de Laurasia, y las diferencias entre las fau- nas de norte y sur de Gondwana, durante el Cretácico Temprano. Palabras clave: dinosaurio, icnofósiles, Cretácico Inferior, Gondwana, Laurasia, intercambio faunístico. 1. Introduction Berlin Society for Geography) on the 17 March 1839 (Buffetaut, 2000; Degenhardt in Mahlmann, 1840). Although originally Considering the first evidence for dinosaurs from South correctly cited as originating from Colombia, subsequent re- America came from Colombia (Buffetaut, 2000; Degenhardt ports erroneously indicated the footprints had been discovered in Mahlmann, 1840), body and trace fossils of these large ter- in México (Degenhardt, 1840; Winkler, 1886; see also Buffe- restrial Mesozoic reptiles are tantalizingly rare in this country. taut, 2000). These tracks were never figured, and were origi- Most South American dinosaurs are known from Argentina nally described as the footprints of giant birds (Degenhardt in and Brasil (e.g., Bittencourt & Langer, 2011; Costa da Silva Mahlmann, 1840). However, this was prior to Richard OWEN et al., 2012; de la Fuente et al., 2007; Díaz–Martínez et al., coining the term Dinosauria (Owen, 1841), so the Colombian 2016; Francischini et al., 2015; Leonardi, 1989; Pazos et al., footprints were almost certainly traces of dinosaurs (Buffetaut, 2012; Weishampel et al., 2004), and indicate taxonomic dis- 2000). These ichnofossils, now lost, came from a Lower Cre- similarity from Laurasian dinosaurs (Cox, 1974; Gheerbrant & taceous locality near Oiba, Santander Department, Colombia Rage, 2006; Canudo et al., 2009). This leaves a considerable (Degenhardt in Mahlmann, 1840), and remain one of only a gap in our understanding of the dinosaurs from northwestern handful of reports of dinosaur body and trace fossils from Co- Gondwana, and their relationships with the more southerly lombia (Table 1). Gondwanan and southern Laurasian realms. Here, we report Skeletal remains of Colombian dinosaurs are rare, with con- on the discovery of six footprints from the Lower Cretaceous firmed reports limited to Late Jurassic/Early Cretaceous sau- Batá Formation of Colombia, northern South America, four of ropods (Carballido et al., 2015; Langston, 1953; Langston & which are attributed to a large ornithopod dinosaur. The Batá Durham, 1955) and Late Cretaceous theropods (Table 1; Ezcu- Formation footprints apparently demonstrate affinities with rra, 2009). However, dinosaur footprints, although rare, are Laurasian iguanodontian ichnofossils, rather than with dino- more common, with reports (in addition to those of Degen- saur footprints from southern South America, therefore these hardt in Mahlmann (1840)) of unidentified footprints from the findings have important implications for the palaeobiogeogra- Jurassic; titanosaurid sauropod, theropod, and rare ornithopod phy of Early Cretaceous ornithopod dinosaurs. footprints from the uppermost Jurassic/lowermost Cretaceous; and theropod footprints from the Lower Cretaceous (Moreno– 1.1. Dinosaurs in Colombia Sánchez & Gómez–Cruz, 2013; Moreno–Sánchez et al., 2011). Hence, most dinosaurs reported from Colombia have been at- The first evidence for dinosaurs in South America were foot- tributed to Saurischia, either members of Theropoda or Saurop- prints reported in the Gesellschaft für Erdkunde zu Berlin (the oda (although some of these taxonomic assignments have been 376 Dinosaur Footprints from the Lower Cretaceous, Batá Formation, Colombia (South America) Table 1. Tabulation of dinosaur remains discovered in Colombia, showing the year of discovery, fossil type, material discovered, location of discovery, the geological formation, and geological age of the find, and the published reference to the discovery. Year Type Material Locality Fm/Age Reference(s)/Notes Summit of ‘Cuchillas de las 1839 T ‘Bird footprints’ ?Theropod. Pezuñas del Venado’, Oiba, Cretaceous, probably Paja Fm. [1, 2] Santander. 1937 B ?Theropod, teeth. Huila and Payandé, Tolima. Cretaceous, unspecified Fm. [3], considered