Contents lists available at ScienceDirect Palaeogeography, Palaeoclimatology, Palaeoecology journal homepage: www.elsevier.com/locate/palaeo Crocodylomorph, turtle and mammal tracks in dinosaur-dominated Middle–?Upper Jurassic and mid-Cretaceous ichnoassemblages of Morocco Hendrik Klein a, ⁎ , Abdelouahed Lagnaoui b , Gerard D. Gierliński c,d , Hafid Saber e , Jens N. Lallensack f , Mostafa Oukassou g , André Charrière h a Saurierwelt Paläontologisches Museum, Alte Richt 7, D-92318 Neumarkt, Germany b Laboratory of Stratigraphy of Oil and Gas Bearing Reservoirs, Department of Paleontology and Stratigraphy, Institute of Geology and Petroleum Technologies, Kazan (Volga Region) Federal University, Kremlyovskaya str. 18, 420008 Kazan, Russia c Państwowy Instytut Geologiczny - Państwowy Instytut Badawczy, ul. Rakowiecka 4, 00-975 Warszawa, Poland d Stowarzyszenie “Delta”, ul. Sandomierska 4, 27-400 Ostrowiec Św., Poland e Laboratory of Geodynamic and Geomatic, Department of Geology, Faculty of Sciences, Chouaïb Doukkali University, B.P. 20, El Jadida MA-24000, Morocco f Steinmann-Institut für Geologie, Mineralogie und Paläontologie, Rheinische Friedrichs-Wilhelm Universität Bonn, Nußallee 8, 53115 Bonn, Germany g Laboratory of Sedimentary Basins Dynamic and Geological Correlations, Department of Geology, Faculty of Sciences Ben M'sik, Hassan II University of Casablanca, B.P. 7955, Sidi Othman, Casablanca, Morocco h Toulouse III University, 13 Terrasses de la Figuière, F-30140 Anduze, France ARTICLE INFO Keywords: Crocodylopodus Hatcherichnus Emydhipus Mammaliamorph Central High Atlas High Moulouya ABSTRACT We report tetrapod traces from two stratigraphic intervals in Morocco, and discuss their palaeoecological im- plications. In the first interval, crocodylomorph footprints assigned to Crocodylopodus meijidei are described from Middle–?Upper Jurassic continental fluvial red beds of the Isli Formation in the Imilchil region, Central High Atlas, Morocco. These traces are associated with a diverse dinosaur-dominated ichnofauna, including the foot- prints of theropods, ornithischians, sauropods and pterosaurs, together with numerous invertebrate traces. In the second interval, we report crocodylomorph swim traces assigned to Hatcherichnus isp., turtle trackways assigned to Emydhipus isp. and isolated mammal footprints in the mid-Cretaceous Midelt Formation, in the High Moulouya region, Morocco. These traces are, also, associated with a diverse dinosaur-dominated ichnofauna including theropods and ornithopods as well as pterosaurs and abundant invertebrate traces. Together with plant remains, conchostracans and fishes, findings indicate a diverse community populating a fluvial-brackish en- vironment. 1. Introduction Jurassic–Cretaceous tetrapod footprint assemblages are essentially dominated by dinosaurs, with theropods and sauropods representing the major components of the Jurassic ichnofaunas (Lockley and Hunt, 1995; Lockley and Meyer, 2000). Less abundant are ornithischians, although they can be more frequent on some surfaces (Olsen and Rainforth, 2003). In Lower Cretaceous strata, the most abundant or- nithischian- and theropod-dominated ichnofaunas are known from North America, Europe and East Asia (e.g., Lockley, 1987; Pérez- Lorente, 2001, 2015; Lockley et al., 2006; Matsukawa et al., 2006; McCrea et al., 2014; Xing et al., 2015). Sauropod ichnofaunas can be more abundant locally (Farlow et al., 1989), as is the case for pterosaurs and birds (Lockley and Rainforth, 2002). Generally, minor components in both ichnofaunas include crocodylians, turtles and mammals. In the case of crocodylians and turtles, this scarcity is partly related to the semi-aquatic lifestyle of many Jurassic–Cretaceous groups and the low preservation potential in subaqueous and inundated substrates (Klein and Lucas, 2015). Mammal tracks are generally rare in the Mesozoic, and crocodylomorph tracks have mainly been reported from small terrestrial forms and by traces of swimming individuals, in particular from the Jurassic–Cretaceous of Europe and North America (Olsen and Padian, 1986; Lockley and Meyer, 2004). From the African continent, crocodylomorph tracks are scarcely known (Hadri et al., 2015; Mateus et al., 2017). Also, turtle tracks thus far have rarely been reported from this region (Belvedere et al., 2013). In recent years, Morocco has become one of the most important places for the study of Paleozoic–Mesozoic tetrapod footprint assemblages. Nearly complete successions with tetrapod footprints from this region https://doi.org/10.1016/j.palaeo.2018.02.028 Received 5 November 2017; Received in revised form 27 February 2018; Accepted 27 February 2018 ⁎ Corresponding author. E-mail addresses: Hendrik.Klein@saurierwelt.de (H. Klein), gerard.gierlinski@pgi.gov.pl (G.D. Gierliński), jens.lallensack@uni-bonn.de (J.N. Lallensack). Palaeogeography, Palaeoclimatology, Palaeoecology 498 (2018) 39–52 Available online 13 March 2018 0031-0182/ © 2018 Elsevier B.V. All rights reserved. T