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Journal Pre-Proof Journal Pre-proof Gregarious behaviour among non-avian theropods inferred from trackways: A case study from the Cretaceous (Cenomanian) Candeleros Formation of Patagonia, Argentina Arturo Miguel Heredia, Ignacio Díaz-Martínez, Pablo José Pazos, Marcos Comerio, Diana Elizabeth Fernández PII: S0031-0182(19)30695-9 DOI: https://doi.org/10.1016/j.palaeo.2019.109480 Reference: PALAEO 109480 To appear in: Palaeogeography, Palaeoclimatology, Palaeoecology Received date: 31 July 2019 Revised date: 15 November 2019 Accepted date: 15 November 2019 Please cite this article as: A.M. Heredia, I. Díaz-Martínez, P.J. Pazos, et al., Gregarious behaviour among non-avian theropods inferred from trackways: A case study from the Cretaceous (Cenomanian) Candeleros Formation of Patagonia, Argentina, Palaeogeography, Palaeoclimatology, Palaeoecology (2018), https://doi.org/10.1016/ j.palaeo.2019.109480 This is a PDF file of an article that has undergone enhancements after acceptance, such as the addition of a cover page and metadata, and formatting for readability, but it is not yet the definitive version of record. This version will undergo additional copyediting, typesetting and review before it is published in its final form, but we are providing this version to give early visibility of the article. Please note that, during the production process, errors may be discovered which could affect the content, and all legal disclaimers that apply to the journal pertain. © 2018 Published by Elsevier. Journal Pre-proof Gregarious behaviour among non-avian theropods inferred from trackways: A case study from the Cretaceous (Cenomanian) Candeleros Formation of Patagonia, Argentina Arturo Miguel Heredia1, 2, *, Ignacio Díaz-Martínez3, Pablo José Pazos1, 2, Marcos Comerio4, Diana Elizabeth Fernández1, 2 1- Universidad de Buenos Aires. Facultad de Ciencias Exactas y Naturales. Departamento de Ciencias Geológicas. Buenos Aires, Argentina. 2- CONICET - Universidad de Buenos Aires. Instituto de Estudios Andinos Don Pablo Groeber (IDEAN). Buenos Aires, Argentina. 3- CONICET - Universidad Nacional de Río Negro. Instituto de Investigación en Paleobiología y Geología (IIPG). General Roca, Río Negro, Argentina. 4- CONICET - Centro de Tecnología de Recursos Minerales y Cerámica (CETMIC). *Corresponding author. E-mail address: [email protected] (A.M. Heredia) ABSTRACT Gregarious behaviour among dinosaurs has been inferred from several lines of evidence (monospecific bonebeds, skeletal morphology, phylogenetic inferences, comparison with modern ecosystems and parallel trackways with particular characteristics), but is relatively poorly documented for non-avian theropods. Here, we report five parallel theropod trackways of large track size (average length of 28 cm) in the Cretaceous (Cenomanian) Candeleros Formation from northwestern Patagonia, Argentina. The tracks are provisionally assigned to aff. Asianopodus pulvinicalx and considering the autopod morphology of the theropod taxa documented in the Candeleros Formation, abelisaurid theropods are suggested as trackmakers. JournalThe trackways possess Pre-proof similar stride, speed estimation, direction and preservational features, and track show uniform depth and do not overlap. Physical barriers (i.e., large fluvial channels or perennial lake) that could influence the direction of the theropod trackmaker gaits were not recognised in the section. Taking into account these considerations, a gregarious behaviour for the abelisaurid theropod trackmakers is proposed. The tracks are preserved as shallow undertracks in a medium-grained sandstone bed deposited in a floodplain setting. The tracking surface is interpreted as the overlaying layer of muddy-siltstone. The occurrence of swelling clays and microbial 1 Journal Pre-proof mats in the track-bearing level could have contributed to the substrate stabilisation and their role in the preservation is discussed. Key words: Theropod footprint; gregariousness; dinosaur behavior; Abelisauridae; Upper Cretaceous; Neuquén Basin 1. Introduction Since the beginning of dinosaur research, some relevant palaeoecological aspects, such as social strategies, have been matter of debate (e.g. Ostrom, 1972). For instance, gregariousness has been attributed to a number of purposes, such as protecting individuals from common aggressors, providing effective access to food sources, defending and raising youngs, or increasing the efficiency of breeding or success of migration (Currie and Eberth, 2010). Tracks and trackways, considered as the product of animal behaviour, are excellent indicators to evaluate the possible social relationships among different dinosaur groups. Currie and Eberth (2010) proposed that parallel trackways, as well as skeletal morphology, bonebed assemblages, phylogenetic inferences and comparison with modern ecosystems, are evidences of gregariousness. Thulborn and Wade (1984) suggested that the trackways should have similarJournal preservation, uniform Pre-proof depth, lack of superposition, and indicate similar speeds to support the gregariousness hypothesis. However, some parallel trackways have been explained as solitary individuals who would pass at different times through the same place and direction due to a geographical barrier (Getty et al., 2015, 2017). Dinosaur gregarious behaviour has been inferred from several worldwide tracksites focused mainly in sauropods and ornithopods (e.g., Ostrom, 1972; Thulborn and Wade, 1984; Farlow et al., 1989; Lockley et al., 1994a; Lockley and Hunt, 1995; Matsukawa et al., 2001; Lockley et al., 2002; Day et al., 2004; Myers and Fiorillo, 2009; Castanera et al., 2011, 2014; Lockley et al., 2012; Piñuela et al., 2 Journal Pre-proof 2016). Parallel theropod trackways as an evidence of gregariousness were addressed by some authors (e.g., Moreno et al., 2012; García-Ortiz and Pérez-Lorente, 2014; McCrea et al., 2014; Lockley et al., 2015). Nevertheless, the social nature of theropod dinosaurs is still discussed (e.g. Barco et al., 2006; Roach and Brinkman, 2007). In Patagonia (Argentina, southern South America), a high diversity of saurischian and ornithischian dinosaurs was found in the Upper Cretaceous continental deposits of the Mesozoic– Cenozoic Neuquén Basin (Leanza et al., 2004). The Cenomanian Candeleros Formation, characterised mainly by fluvio-aeolian interaction deposits, constitutes the basal unit of the Neuquén Group deposited in the foreland basin of the Neuquén Basin (Garrido, 2010; Fennell et al., 2017). The Candeleros Formation presents one of the richest dinosaur fossil records from the basin and includes both, diplodocoid and titanosaurid sauropods (Otero and Salgado, 2015, and references therein), and a great diversity of theropods: abelisaurids, carcharodontosaurids, basal coelurosaurs, alvarezaurids and dromaeosaurids (Coria and Salgado, 1995; Calvo et al., 2004; Makovicky et al., 2005; Makovicky et al., 2012; Novas et al., 2012, 2013; Canale et al., 2016). Taking into account the published data, the Candeleros Formation presents the highest track ichnodiversity from the basin and includes: sauropod, theropod, iguanodontid and other ornithopod, and also pterosaur tracks and trackways (Calvo, 1991, 1999; Calvo and Lockley, 2001; Calvo and Mazzetta, 2004; Calvo and Rivera, 2018; Candia Halupczok et al., 2018; Heredia et al., 2019). While theropod tracksJournal are abundant for thePre-proof Cretaceous (Cenomanian) worldwide (e.g. Mezga and Bajraktarević, 1999; Lockley et al., 2014 and references therein; Romilio and Salisbury, 2014; Calvo and Rivera, 2018), ichnological evidence for theropod gregariousness from this stage has been poorly documented. In this work, several parallel trackways of tridactyl tracks found in a new tracksite from the Cenomanian Candeleros Formation (Northern Patagonia) will be analysed. The aim of this paper is fourfold: i) to describe in detail the surface bearing the tracks focusing on several parallel tridactyl trackways as well as the accompanying sauropod tracks and the substrate properties; ii) to discuss the role played by the substrate in track morphology and preservation; iii) to provide an 3 Journal Pre-proof ichnotaxonomical analysis and compare this record with similar theropod trackways worldwide; and, iv) to discuss the possible trackmakers and their palaeobiological implications focusing on gregarious behaviour. 2. Geological setting The Neuquén Basin is located in west-central Argentina and it extends from 32° to 40° S latitude along the Andean foothills (Fig. 1A). It covers an area of about 120,000 km² and includes a continuous Late Triassic–Palaeogene stratigraphic record of marine to continental facies, more than 7000 m thick (Legarreta and Gulisano, 1989; Yrigoyen, 1991; Vergani et al., 1995). In the Late Cretaceous– Palaeogene interval, the Neuquén basin represents a foreland basin mainly composed of continental deposits with minor marine intervals (Howell et al., 2005; Aguirre-Urreta et al., 2011). The Cenomanian–Campanian Neuquén Group lies above the Rayoso Formation (Bajada del Agrio Group) overlying an angular unconformity known as Patagonídica; it is 1 km thick and comprises fluvial, aeolian and lacustrine deposits (Stipanicic et al., 1968; Leanza, 2009; Garrido, 2010). The Neuquén Group is divided into three units (Fig. 1B) known as Río Limay, Río Neuquén and Río Colorado subgroups (Cazau and Uliana, 1973; Ramos, 1981). However, the stratigraphy of the group is not uniform throughout the entire basin, and the tripartite division is only recognizable between
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