PALAIOS, 2011, v. 26, p. 601606 Research Article DOI: 10.2110/palo.2011.

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DINOSAUR FOOTPRINTS AS ULTIMATE EVIDENCE FOR A TERRESTRIAL ENVIRONMENT IN THE LATE SINEMURIAN TRENTO CARBONATE PLATFORM
FABIO MASSIMO PETTI,* MASSIMO BERNARDI, ROSSANA TODESCO, and MARCO AVANZINI
Museo delle Scienze, Via Calepina 14, 38122 Trento, Italy, fabio.petti@mtsn.tn.it, massimo.bernardi@mtsn.tn.it, rossana.todesco@mtsn.tn.it, avanzini@mtsn.tn.it

ABSTRACT Three dinosaur footprints were found, preserved as natural casts, on the roof of a First World War tunnel in the Pasubio Massif (Trentino AltoAdige, northeastern Italy). The Austro-Hungarian Army used the tunnel until 1918, presumably to connect the front- with the back line. The tracks are tridactyl, two of which are organized in a trackway, and provide evidence for the presence of medium-sized theropods. The track-bearing horizon is situated at the base of the Rotzo formation (Calcari Grigi Group), within shallow-water black shale deposits characterized by oligotypic small bivalves, oligohaline ostracodes, and thecamebians. Vertebrate remains, represented by crocodyloform teeth and fish scales, were also identified from the same stratigraphic portion. The occurrence of dinosaur footprints from black shale levels sheds light on their controversial interpretation, confirming the occurrence of terrestrial environment in the Trento carbonate platform during the late Sinemurian. Dinosaur tracks indicate the emersion of a sector of the tidal flat during the deposition of the basal Rotzo formation, and suggest a complex paleonvironmental scenario in which tidal flats were associated with freshwater ponds and islands. INTRODUCTION On the 28th of June 1916, the First Regiment of the AustroHungarian Army was stationed in Serrada di Folgaria, after fighting a hard battlebetween 15 May and 27 June 1916against the Italian Army on the Altipiani Vicentini, the so-called Strafexpedition. Soldiers were ordered to reach the Pasubio Massif area. For the first time the Kaiserjager [Emperors hunters] troops of the Austro-Hungarian army set foot on the mountain that they kept stubbornly until the end of the First World Warsince then also known as Kaiserjager mountain. The Pasubio Massif eventually became a key sector, and, during 1916, the scene of one of the most fierce and bloody battles of the First World War. The Pasubio Massif is located in the southern sector of the Trentino Alto-Adige Region, near the town of Rovereto (Trento Province), between the Vallarsa and the Terragnolo valleys (Fig. 1). The Pasubio Massif is crosscut by a series of tunnels dug both by the Austro-Hungarian and Italian armies during the First World War. To the north of Cima Roite, several tunnels dug by the Austro-Hungarian Army constitute an impressive complex, known as the Stellung der Kaiserjager. During the exploration carried out with local speleologists from the Societa degli Alpinisti Tridentini (SAT), one of us (MA) ` noticed three bulges on the roof of one of these tunnels (Stol dei Campiluzzi), soon recognized as tridactyl dinosaur tracks preserved as natural casts. The track-bearing level belongs to the basal portion of the Lower Jurassic Rotzo formation, the most intensively studied formation of the Calcari Grigi Group (Fig. 2). The Rotzo formation is generally regarded as a shallow-water carbonate succession deposited from the Hettangian to the Pliensbachian on the Trento carbonate platform (Avanzini et al., 2007). Dinosaur tracks and trackways are widespread in the Calcari Grigi Group. To date, eleven dinosaur
* Corresponding author.

tracksites were discovered, grouped in three main track-bearing levels constrained between the upper Hettangian and upper Sinemurian (see Avanzini and Petti, 2008 for a review). Only three tracksites belong to the Rotzo formation (Bella Lasta, Marocche di Dro, Coste dellAnglone; Mietto et al., 2000; Avanzini et al., 2001; Petti et al., 2011). Early Jurassic dinosaur footprints were already described from the Pasubio Massif (Cima Palon, Malga Buse Bisorte); however, they belong to the oldest lithostratigraphic unit of the Calcari Grigi Group (Monte Zugna formation; Avanzini, 2001). Interestingly, Late Triassic (Norian) dinosaur footprints were recently described in the same area, preserved as convex hyporeliefs, on the roof of the 12th gallery of the so-called Strada delle Gallerie, dug in 1917 by the Italian Army, during the First World War (Belvedere et al., 2008). The aim of this paper is to document these tracks and to discuss their significance for paleoenvironmental and paleogeographical interpretation of the Lower Jurassic Trento carbonate platform. GEOGRAPHICAL AND GEOLOGICAL CONTEXT The Stol dei Campiluzzi tunnel, dug by the Austrian Army until 1918, is located below the Campiluzzi saddle, west of Monte Buso (Fig. 1; Pasubio Massif; latitude: 45u48954.870N; longitude: 11u1095.180E). The tunnel is about 250 m long, 2.5 m wide, and about 2 m high. The studied area is located on the Pasubio Massif, which corresponds to the southern sector of the Trento carbonate platform, one of the paleogeographic domains of the southern continental margin of the Alpine Tethys (Bosellini, 1973; Gaetani, 1975; Bernoulli et al., 1979; Winterer and Bosellini, 1981; Sarti et al., 1992). During the Mesozoic, this margin displayed a complex physiography characterized by carbonate platforms, pelagic plateaus, and basinal settings, strictly linked to the opening of the northern sector of the Central Atlantic and to the geodynamics of the Neotethys (Bertotti et al., 1993; Stampfli and Borel, 2004). During the Early Jurassic, the western area of the southern Alpine region was affected by a strong subsidence and various sectors of the Late Triassic carbonate platforms (Piedmont and Lombardy) rapidly drowned. On the contrary, the eastern part was still dominated by carbonate platform sedimentation and was articulated from the west to the east in three main paleogeographic units; the Trento Platform, the Belluno Basin, and the Friuli Platform. The base of the Pasubio Mesozoic sedimentary succession is characterized by the Dolomia Principale (Upper Triassic), which is overlain by the shallow-water carbonates of the Calcari Grigi Group (HettangianPliensbachian; see Avanzini et al., 2007 for a review of the unit). The drowning phase of the Trento Platform is recorded by the San Vigilio Oolitic Limestone (ToarcianAalenian) and by the Rosso Ammonitico Veronese Formation (upper BajocianTithonian). The Mesozoic succession ends with the well-stratified calpionellid limestone of the Maiolica Formation (TithonianCenomanian), which is overlain by scattered Quaternary continental deposits referable to the glacial phases. The structural setting of the area is characterized by transcurrent or transpressive faults that derive from Mesozoic extensional faults
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arid climate. Azonotriletes (40%) belonging to Pteridophyta indicates the presence of warm swamp or marsh vegetation, typical of a humid climate and freshwater influence. Chasmatosporites, belonging to Cycadales, reflecting subtropical zones to warm-humid climates, are relatively rare (2%; see also Avanzini et al., 2006). According to Bassi et al. (1999), Boomer et al. (2001), and Fugagnoli (2004), the laminated black shales of the Rotzo formation lie below the lower boundary of the Orbitopsella Zone (upper Sinemurian), below the first occurrence of the Lithiotis lithofacies and just above the boundary with the underlying Loppio Oolitic Limestone. The discovered track-bearing layer can be thus confidently attributed to the upper Sinemurian. THE STOL DEI CAMPILUZZI SECTION The base of the studied section (,19 m thick) is characterized by a massive oolitic grainstone 80 cm thick, marked by low-angle cross laminations (Loppio Oolitic Limestone). This portion is overlain by a thin reddish surface, evidence of prolonged subaerial exposure. The section continues with well-stratified, gray to yellow limestone-marl alternations, for a total thickness of ,13 m. In this portion, decimeterthick layers of wackestone-packstone with intraclasts and bioclasts (mainly bivalves) occur at various stratigraphic levels. Skeletal limestones represent calcareous tempestites due to normal or storm wave action (basal lag of the tempestites). Within the limestone-marl alternations, black shale deposits with small bivalves (Eomiodon sp. and Lithioperna sp.; Sprocatti, 2010) occur at four different stratigraphic levels (Fig. 3). The shales consist of laminated, dark-gray to black wackestones and packstones rich in organic matter. The bivalve assemblage is made of complete shells, disarticulated valves, and shell fragments (Fig. 4). The track-bearing horizon is placed between the second and the third black shale level at about 9.2 m from the base of the section. The marly calcareous alternation is overlain by 5 m of thinbedded limestone, intensively burrowed mainly by Thalassinodes isp. (interpreted as decapod crustacean burrows), giving to this portion the typical nodular aspect. MATERIAL AND METHODS Three medium-sized, functionally tridactyl, moderately well preserved pedal prints made by a bipedal trackmaker are recognizable on the track-bearing surface (Fig. 5). They are preserved as convex hyporelief (sensu Leonardi, 1987). Two tracks are arranged in a discrete trackway (Fig. 6). Footprints were mapped using the conventional method of tracing footprint outlines on transparent acetate overlays but also digitally documented with the aim of making three-dimensional prints of the uncollected specimens. The tracks were labeled with the prefix MTSN-SC (Museo Tridentino di Scienze Naturali, Stol dei Campiluzzi) (abbreviations for the following discussion: FL 5 Footprint length; FW 5 Footprint width; te 5 toe print extension; h 5 hip height; L 5 body length; Mb 5 body mass). MTSN-SC1 is a right pedal print, mesaxonic and slightly asymmetric, longer than wide (FL 5 26 cm; FW 5 24 cm). Digits are slender, similar in width (3.5 to 5 cm), distally tapering with rounded claw traces. Digit III is slightly longer than digit IV, which is clearly longer than digit II. Digit III is moderately projected beyond the line connecting the tip of digits II and IV (te 5 9.7 cm). Total divarication (IIIV) is 64u and the interdigital angle between digits II and III is smaller (25u) than the one between II and IV (39u). Three subcircular digital pad traces occur on digit IV, whereas only one is visible on digits II and III. The proximal pad of digit IV lies lateral to the digit III axis and behind the proximal pad trace of digit II. The pedal hypex between digits II and III is slightly higher than the one between III and IV. MTSN-SC2 is a left pedal print, less well preserved than MTSNSC1 with digits II and III faintly impressed. This track is tridactyl,

FIGURE 1Location map of the study area in Southern Alps (northeastern Italy) (figure was generated using the Global Multi-Resolution Topography [GMRT] synthesis, the default basemap in GeoMapApp version 3.1.2 [Ryan et al., 2009]).

reactivated and reversed during the Alpine compression (Avanzini and Masetti, 1992; Avanzini, 1994; Tobaldo et al., 2004). The Stol dei Campiluzzi tunnel was dug entirely into the lower portion of the Rotzo formation (Fig. 2; Bosellini and Broglio Loriga, 1971; Masetti et al., 1998), characterized by a heterogeneous lithological compositionooidal, peloidal, bioclastic and intraclastic limestone, marls, and claysand arranged into asymmetrical thickening and shallowing-upward meter-scale sequences (Masetti et al., 1998). This formation has been generally referred to a subtidal environment (Masetti et al., 1998) and can be subdivided into two main lithofacies: the lower is mainly represented by limestone-marl alternations, whereas the upper is characterized by the so-called Lithiotis beds or mounds (Masetti et al., 1998; Posenato and Avanzini, 2006). The lower lithofacies is essentially characterized by bioturbated mudstones-wackestones which are locally interbedded with microlaminated wackestones-packstones dark gray in color, slightly marly, strongly fissile, and with a high concentration of organic matter (Masetti et al., 1996, 1998; Bassi et al., 1999, 2008; Boomer et al., 2001; Monaco and Gianneti, 2001; Fugagnoli, 2004). These black shale deposits are characterized by a well-preserved paucispecific and oligohaline fauna of small thin-shelled bivalves (Eomiodon, ?Myrene, and rare isognomids; Bassi et al., 1999, 2005), ostracodes (Phraterfabanella with subordinated Klieana and Limnocythere; Boomer et al., 2001), as well as fossil testate amoebae (Difflugia, Pontigulasia, and Centropyxis; Bassi et al., 2008). Vertebrate remains have also been documented from these levels with crocodyloform (?thalattosuchian) teeth and semionotid fish scales (M. Bernardi, F. M. Petti, and M. Avanzini, unpublished data, 2011). In addition, these black shales contain abundant macrofloral remains, mostly composed of cuticle, rhizoliths in life position, and charcoal, suggesting the presence of a well-developed flora, including woody plants (Avanzini, 1998; Masetti et al., 1998). The palynological assemblage is largely dominated by Circumpolles forms (56%), usually attributed to a xerophitic vegetation of subtropical, warm, and rather

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FIGURE 2Stratigraphic column showing the lithological and paleontological composition of the Calcari Grigi Group. In the right column a close view of the measured section with the track-bearing horizon.

mesaxonic and slightly asymmetric, longer than wide (FL 5 25 cm; FW 5 22 cm). Digit III toe extension is low (te 5 8.0 cm) but the distal-most portion is not impressed. The divarication angle is 27u between digits II and III, 35u between II and IV and the total

divarication (IIIV) is 62u. Only digit IV preserves two digital pad traces and a rounded claw mark. No anatomical details can be discerned either on digit II or on IV. Pedal hypices seem to lie at the same distance from heel.

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FIGURE 5Stol dei Campiluzzi dinosaur tracks. The tracks are preserved as natural casts (convex hyporeliefs) on the sole surface of a limestone bed.

interdigital angle IIIIV being wider (34u) than IIIII (23u). Pedal hypices lie at the same height. A narrow and sinuous drag mark, less than 1 to 3 cm wide and about 26 cm long, was produced by the animal during the locomotion (Figs. 56). Digit III lies above digit III impression of MTSN-SC1, testifying to the subsequent trampling of the MTSN-SC3 producer. DISCUSSION Ichnotaxonomy and Trackmaker Attribution
FIGURE 3The freshwater black shale deposits containing a well-preserved benthic bivalve and ostracode assemblage, along one of the walls of the First World War tunnel.

The trackway MTSN-SC1-2 has a narrow gauge pattern, with an estimated pace angulation of ,180u. The calculated pace length is 116 cm. MTSN-SC3 is a left tridactyl and mesaxonic pedal print. It is symmetric, as long as wide (FL 5 FW about 26 cm). Only digits III and IV display anatomical features of the trackmakers pes. Digits III and IV are slender, tapering distally with short and pointed claw marks. Only the proximalmost and the distalmost digital pad impressions, both subcircular in shape, are preserved on digit III. Nevertheless, digit III shows some creases on its left lateral margin, possibly defining other two digital pad traces. Digit III is the longest, followed by digit IV, which is markedly longer than digit II. Total divarication is 57u with the

One or probably two individuals with equivalent pes proportions likely produced the tracks at different times. The trackmakers hip height (H) is 1.25 m according to Thulborns (1990) equation (h 5 3.06 3 FL1.14). The estimated body length is ,5.0 m (L 5 4H, Thulborn, 1990) and the estimated body mass value is 206 kilograms calculated following the formula developed by Weems (2006). The Calcari Grigi Group has yielded many tridactyl footprints, mainly from the Monte Zugna and Rotzo formations (Avanzini and Petti, 2008). The described footprints show analogies with several tridactyl specimens of the Lavini di Marco, Coste dellAnglone, and Bella Lasta tracksites assigned to the ichnogenus Kayentapus Welles 1971 (Leonardi and Mietto, 2000; Mietto et al., 2000; Piubelli et al., 2005; Avanzini et al., 2006; Piubelli, 2006; Petti et al., 2008, 2011). The examined footprints also resemble a tridactyl track discovered in the ski et al., 2004) and attributed to Early Jurassic of Poland (Gierlin Kayentapus minor (Weems, 1992). It shows similar values of the ratio between the footprint length and the projection of digit III beyond the lateral toes (FL/te), being 2.59 in the Polish specimen and 2.58 in the best-preserved specimens found in the Stol dei Campiluzzi tunnel. Nevertheless, the Polish specimen is larger (FL is ,39 cm), more symmetrical, and less divaricated (IIIV 5 48u). We conservatively assign the tracks described here to the ichnogenus Kayentapus isp. and attribute them to medium-sized theropods. This ichnogenus has been often associated with such ceratosaurian dinosaurs as Dilophosaurus sp. (Weems, 2006; Petti et al., 2011), whose skeleton is known only from the Lower Jurassic of Arizona. Paleoenvironmental Inferences The Rotzo formation has been traditionally referred to an inner lagoon environment protected westward by marginal oolitic bars and shoals (Bosellini and Broglio Loriga, 1971). Masetti et al. (1998) proposed a ramp-lagoon environment sheltered by oolitic barrier

FIGURE 4Close-up view of a slab exhibiting a concentration of broken and complete bivalve shells (Eomiodon sp.).

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importance for paleoenvironmental and paleogeographical interpretations. They suggest, together with other fossil evidence from the study interval, a complex environment for the Early Jurassic Trento Platform, in which tidal flats were probably associated with a mosaic of ponds and emergent land. ACKNOWLEDGMENTS
FIGURE 6Line drawing of the studied specimens, uncollected from Stol dei Campiluzzi locality.

islands located at the two Trento Platform margins. According to these authors, the shallowing-upward trend of the lower lithofacies can be attributed to a subtidal slope depositional environment gently dipping westward and northward, and connecting the deeper portion of the platform, where the limestone-marl alternations were being deposited, to the shallower areas (see also Masetti, 2002). Avanzini (1998) and Avanzini and Tomasoni (2001) proposed a fining-upward model in which the basal bioclastic and bioturbated levels, deposited in a subtidal and oxygenated environment, are covered by the marly calcareous alternations, deposited in a restricted and shallow environment with scarce or absent water circulation. According to these models the richness in organic matter is due to the relative decrease in carbonate sediment supply. Other authors (Bassi et al., 1999, 2005, 2008; Boomer et al., 2001; Fugagnoli, 2004), agreed with this latter interpretation, focusing their attention on the palaeontological content of the black shale interval. Fossil invertebrates (bivalves, ostracodes, thecamoebians) and plants indeed indicate a stressed freshwater environment with poorly oxygenated oligohaline waters with episodic and slow currents typical of a shallow eutrophic restricted aquatic environment (coastal pond) with low or absent water exchange with the open sea (Avanzini, 1998; Bassi et al., 1999, 2008; Boomer et al., 2001; Monaco and Giannetti, 2001, 2002). The fossil vertebrate assemblage composition (M. Bernardi, F. M. Petti, and M. Avanzini, unpublished data, 2011) is also consistent with a brackish pond eutrophic environment where semionotid fish were possibly preyed on by marine crocodylomorphs (?thalattosuchians), well adapted to life in marginal marine and shallow-water environments (Karl and Tichy, 2004; Karl et al., 2006). Within this context, the discovery of tracks conclusively provides support for a transitional marine-continental depositional environment for the black shales. The dinosaur footprints in the study interval suggest the emersion of part of the tidal flat during the sedimentation of the basal Rotzo formation (upper Sinemurian) and, together with other fossil evidence, indicate a complex environment for the Early Jurassic Trento Platform in which tidal flats were associated with an articulated complex of ponds close to emergent land. CONCLUSIONS The upper Sinemurian black shale deposits of the Rotzo formation, outcropping in northeastern Italy, include diverse and abundant fossil assemblages represented by a well-developed flora (cuticles, rhizoliths in life position, charcoal, and pollen) and fauna (oligotypic bivalves, monospecific ostracods, testate amoebians, crocodylomorph teeth, fish scales). Dinosaur tracks attributed to medium-sized theropods were found in a dark-gray marly bed situated between two distinct black shale levels. The new finding strongly contributes to the comprehension of the environmental conditions in which the Rotzo formation was deposited, particularly for its basal portion. Although in the last twenty years many authors agreed to consider the black shale levels as deposited in a freshwater environment, the dinosaur tracks provide the ultimate material evidence for this interpretation and another instructive example of their considerable

The authors thank the editors of PALAIOS and reviewers M.G. Lockley and V. Faria Dos Santos for their helpful comments, which greatly improved the manuscript. This research was funded by the Autonomous Province of Trento, as the sponsor of the DINOGEO post-doctoral research project (Scientific Responsibility: F.M. Petti; Scientific Coordinator: M. Avanzini) and of the OPENLOC research project under the call for proposals Major Projects 2006. Partners of the OPENLOC project are the E. Mach Foundation, the Manchester Institute of Innovation Research, the Trento Museum of Natural Sciences, the University of Bologna, and the University of Trento. REFERENCES
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ACCEPTED JULY 25, 2011