A Diverse Upper Jurassic Dinosaur Ichnofauna from Centralwest Portugal
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A diverse Upper Jurassic dinosaur ichnofauna from central-west Portugal OCTA´ VIO MATEUS AND JESPER MILA` N Mateus, O. & Mila`n, J. 2010: A diverse Upper Jurassic dinosaur ichnofauna from central-west Portugal. Lethaia, Vol. 43, pp. 245–257 A newly discovered dinosaur track-assemblage from the Upper Jurassic Lourinha˜ Forma- tion (Lusitanian Basin, central-west Portugal), comprises medium- to large-sized sauro- pod tracks with well-preserved impressions of soft tissue anatomy, stegosaur tracks and tracks from medium- to large-sized theropods. The 400-m-thick Lourinha˜ Formation consists of mostly aluvial sediments, deposited during the early rifting of the Atlantic Ocean in the Kimmeridgian and Tithonian. The stratigraphic succession shows several shifts between flood-plain mud and fluvial sands that favour preservation and fossiliza- tion of tracks. The studied track-assemblage is found preserved as natural casts on the underside of a thin bivalve-rich carbonate bed near the Tithonian–Kimmeridgian boundary. The diversity of the tracks from the new track assemblage is compared with similar faunas from the Upper Jurassic of Asturias, Spain and the Middle Jurassic York- shire Coast of England. The Portuguese record of Upper Jurassic dinosaur body fossils show close similarity to the track fauna from the Lourinha˜ Formation. h Dinosaur tracks, Lusitanian Basin, Portugal, skin impressions, Upper Jurassic. Octa´vio Mateus [[email protected]], Universidade Nova de Lisboa, Faculdades de Cieˆn- cias e Tecnologia, Monte de Caparica, Portugal; Museu da Lourinha˜, Rua Joa˜oLuisde Moura, 2530-157 Lourinha˜, Portugal; Jesper Mila`n [[email protected]], Department of Geography and Geology, Geology Section, University of Copenhagen, Øster Voldgade 10, DK-1350 Copenhagen K, Denmark; Geomuseum Faxe, Østsjællands Museum, Højerup bygade 38, DK-4660 Store Heddinge, Denmark; manuscript received on 16 ⁄ 12 ⁄ 2008; manuscript accepted on 08 ⁄ 06 ⁄ 09. TheMiddletoLateJurassiccarbonateplatformsof 1989). The Lourinha˜ Formation contains a highly Portugal have yielded several of the largest and most diverse vertebrate fauna comprising fish, amphibians, important dinosaur tracksites in Europe (e.g. Dantas turtles, pterosaurs, crocodiles, dinosaurs and mam- et al. 1994; Lockley et al. 1994a; Santos et al. 1994; mals (Antunes 1998; Antunes & Mateus 2003; Mateus Lockley & Meyer 2000), yet only recently the value of 2006; Mateus et al. 2006), as well as dinosaur nests, dinosaur tracks from Upper Jurassic siliciclastic eggs and embryos (Antunes et al. 1998; Mateus et al. deposits have been recognized. This paper discusses 1998). The formation is also known for its fossil vege- one such occurrence. tation, namely plant fragments and large fossilized tree The Lourinha˜ Formation is exposed in central-west logs (Pais 1998). Yagu¨e et al. (2006) proposed a litho- Portugal, mostly near to the town of Lourinha˜ stratigrapic scheme for the continental Upper Jurassic (Fig. 1). The formation was deposited in the Lusita- of the Lusitanian basin, which includes the Bombarral, nian Basin and comprises more than 400 m of mostly Sobral and Alcobac¸a Formations (including Conso- terrestrial sediments, with some coastal intercalations lac¸a˜o and Praia da Amoreira- Porto Novo Members) (Kullberg et al. 2006) deposited during the early within the Lourinha˜ Group. rifting stage of the Atlantic Ocean in latest Jurassic, The succession shows abundant shifts between Tithonian–Kimmeridgian times (Hill 1989). The sedi- unconsolidated flood-plain mud and fluvial sandstone ments consist predominantly of thick beds of red and that favour recognition of fossil tracks. Within the last green clay, interbedded with massive fluvial sandstone years, more than 30 well-preserved single tracks and bodies and heterolithic beds. The sandstone bodies are small trackways of sauropods, theropods, thyreopho- horizontally extensive, lenticular beds that in some rans and ornithopods have been discovered, both pre- instances are traceable for several kilometres in the served in-situ on the original sedimentary surfaces and exposed cliff sections along the coast. The sandstone on loose blocks fallen to the beach caused by the lenses have been interpreted as distal alluvial fan facies extensive coastal erosion (Antunes & Mateus 2003; originating from periods of extensive faulting (Hill Mila`n et al. 2005; Mateus & Mila`n 2008). DOI 10.1111/j.1502-3931.2009.00190.x Ó 2009 The Authors, Journal compilation Ó 2009 The Lethaia Foundation 246 O. Mateus & J. Mila`n LETHAIA 43 (2010) Fig. 1. Geological map of central-west Portugal (based on Alves et al. 2003) with main track localities depicted. The aim of this study is to describe a new multi- Thulborn 1990; Lockley 1991). Ornithopod tracks are taxon track assemblage found in the coast cliffs just generally wider than they are long, the digits are short south of the locality of Porto Dinheiro, to compare it and rounded, and when present the imprints of the with similar track faunas from the Middle Jurassic of claws are blunt and rounded. The divarication angle England and the Upper Jurassic of Spain, and to com- between digits II and IV are normally in excess of 60°, pare it with the known dinosaur skeletal record from their trackways are set in a broader pattern than the Lourinha˜ FormationandtheUpperJurassicof theropods, and the feet often show an outward rota- Portugal. tion (Moratalla et al. 1988; Thulborn 1990; Lockley 1991). Stegosaur tracks are poorly known in the fossil PortoDinheirotrackassemblage record, and some confusion exists about the identifi- cation of stegosaur tracks. (Thulborn 1990; Whyte & Identification of tracks Romano 1994, 2001; Lockley & Hunt 1998; Long 1998; Gierlinski & Sabath 2002; Lires et al. 2002; Theropod and ornithopod dinosaurs are both func- Garcı´a-Ramos et al. 2006, 2008; Whyte et al. 2007; tionally tridactyl, and both leave tridactyl tracks, that Lockley et al. 2008) The track named as Deltapodus at first glance can seem difficult to distinguish them. brodricki (Whyte & Romano 1994, 2001), however, is Theropod tracks are characterized by being longer a close fit for the flesh-out morphology of the stego- than they are wide, with long narrow, often tapering, saur pedal skeleton. Deltapodus is characterized by digits ending in long, sharp claw impressions (Mora- entaxonic, crescent-shaped manus impression that is talla et al. 1988; Thulborn 1990; Lockley 1991), and approximately twice as wide as long, and may have the divarication angle between digits II and IV is the impression of an inward-directed pollex claw. normally 50–60°,butcanbeaslowas35° andupto The pes of Deltapodus is generally triangular to sub- 75° (Thulborn 1990). Theropod trackways are set in a triangular in outline, tridactyl and mesaxonic, with narrow gauge pattern, with high pace angulations, impressions of short, bluntly rounded digits and a approaching 180° and a tendency for the feet to maximum width across the base of the digit impres- show slight inward rotation (Moratalla et al. 1988; sions (Whyte & Romano 1994). LETHAIA 43 (2010) Upper Jurassic dinosaur ichnofauna from central-west Portugal 247 Sauropods display an extreme degree of heteropody cliff face. The preservation of the tracks is unusual in in their tracks, the sauropod manus track is crescent that the tracks are located on the underside of a mar- shaped, normally without indications of free digits, ine limestone bed containing numerous large bivalves except for, in some genera, a prominent inward-direc- and large Thalassinoides burrows. The casts of the ted pollex claw. The pes track is elongated, entaxonic, tracks contain several of the large bivalves, and bival- and can display from three to five short outward- ves are present between the digit impressions in some rotated digit impressions (Thulborn 1990; Lockley of the tracks, suggesting that the tracks were emplaced 1991; Lockley et al. 1994b; Wright 2005). The manus- in shallow water or during a period of sub-aerial expo- pes size ratio varies from 1:2 (Santos et al. 1994) to sure. 1:5 (Lockley et al. 1994b), and sauropod trackways The first track to be discovered from the locality can be roughly divided into wide- and narrow-gauge was a large well-preserved cast of the right pes from a trackways (Lockley et al. 1994b). sauropod (Fig. 3), followed by several tridactyl thero- During field work in the summers of 2005 and pod tracks, a stegosaur track and a giant-sized thero- 2006, several casts of dinosaur tracks were discovered pod track were found within a 20 m stretch of fallen on the beach and in the cliff section approximately blocks on the beach. Later, several tracks and frag- 500 m north of the beach at Porto Dinheiro (Fig. 1). ments of tracks were recovered in loose blocks on the All the tracks were found as casts on the underside of cliff face. large blocks fallen from a laterally extensive marine carbonate bed near the top of the cliff section (Fig. 2). Sauropod tracks This carbonate bed is the upper of two distinct 0.5-m- thick carbonate layers, rich in the bivalves Isognomon Several natural casts of tracks was indentified as sauro- and Eomiodon securiformis, and represents short, mar- pod pes tracks as they range in size from 58 to 105 cm ine transgressions. The Isognomon ⁄Eomiodon layer in length, are entaxonic, elongated piriform, with with the tracks can be traced for several kilometres impressions of short, pointy, outward facing digits along the coastal cliffs, and the upper layer represent (Figs 3–5). Some specimens show evidence of short the interface between the Praia da Amoreira-Porto blunt claws and a division into digital pads (Figs 3, 4). Novo Member and the Sobral Formation (= Praia Theheelareaofthefootconsistsofalargerounded Azul Member), recognized as Upper Kimmeridgian to pad, separated from the digit impressions by a more Early Tithonian in age (Leinfelder 1987). The carbon- shallowly impressed area. ate bed displays abundant tracks, and seven blocks The casts of the individual digits, including the with tracks were found within a distance of 20 m from ungual insertion in digit II, are preserved exquisitely each other, as well as several loose scattered down the in the best-preserved cast (Fig.