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Photogrammetry derived 3D models, digitally reconstructed using different convergent images. The generated dense surface models, shown here in colour-code mode, have a resolution of 0.5 mm. Color banding reflects topography. A) Specimen CA2-17 (light blue = highest; red = lowest) the maximal depth of the impression of 1.6 cm, scale bar 5 cm. B) Specimen CA15-11 (blue = highest; green = lowest) the maximal depth of the impression of 2.1 cm, scale bar 5 cm.
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Bipedal tridactyl dinosaur footprints dominate the Early Juras-sic Coste dell'Anglone tracksite, located on the eastern slope of the Mt. Biaina-Mt. Brento chain (Trentino Alto-Adige, NE Italy). The site yielded 544 tridactyl footprints, arranged in 20 long trackways, and belongs to the upper part of the Calcari Grigi Group (lower portion of the Rot...
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Abundant dinosaur tracks discovered near Thermopolis, Wyoming, are preserved on bedding planes within palustrine-lacustrine, crevasse-splay, and point-bar deposits of the Morrison Formation. We recognize nine vertebrate track types in the study area. We attribute the tracks to small theropod dinosaurs, turtles, possible small sauropod dinosaurs or...
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The Late Triassic-Early Jurassic Elliot Formation of South Africa is one of the most important geological formations worldwide for understanding the early evolution of sauropodomorph dinosaurs. However, many of the taxa currently recognized as valid within its lower strata remain either poorly understood, vaguely diagnosed, or both. The recent disc...
Sauropodomorpha were herbivorous saurischian dinosaurs that incorporate Sauropoda and early-diverging sauropodomorphs. The oldest sauropodomorph remains are known from Late Triassic deposits, most of them Gondwanan. The Laurasian record comprises some Triassic forms, but the bulk is Jurassic in age. Among the 14 Jurassic non-sauropodan sauropodomor...
Pathological or traumatic loss of teeth often results in the resorption and remodeling of the affected alveoli in mammals. However, instances of alveolar remodeling in reptiles are rare. A remodeled alveolus in the maxilla of the Chinese theropod Sinosaurus (Lower Jurassic Lower Lufeng Formation) is the first confirmed example of such dental pathol...
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... The track-bearing horizon belongs to a stratigraphic succession characterised by shallowing-upward cycles. Each cycle is characterised by the transition from a low energy subtidal unit (intensely bioturbated nodular limestone), with associated invertebrate traces (Thalassinoides isp., Chondrites isp.), to high energy subtidal deposits (oolitic grainstone with intraclasts and bioclasts) up into inter-and supra-tidal deposits (Petti et al., 2011b). ...
... In the upper part of the sequence, the track layer consists of poorly fossiliferous dark grey stromatolitic and peloidal mudstone Petti et al., 2011b). The section belongs to the lowermost portion of the Rotzo Formation (formerly Tovel Member sensu Castellarin et al., 2005) and the microfossil assemblages indicate a late Sinemurian to Pliensbachian age Petti et al., 2011b). ...
... In the upper part of the sequence, the track layer consists of poorly fossiliferous dark grey stromatolitic and peloidal mudstone Petti et al., 2011b). The section belongs to the lowermost portion of the Rotzo Formation (formerly Tovel Member sensu Castellarin et al., 2005) and the microfossil assemblages indicate a late Sinemurian to Pliensbachian age Petti et al., 2011b). ...
This article reports the state of the art of research on the Jurassic tetrapod footprints from Italy. Most dinosaur ichnosites are located in the Southern Alps (Northeastern Italy, Veneto, Trentino Alto-Adige) with the exception of Mattinata (Southern Italy, Gargano Promontory, Apulia) and the Burano River ichnosite (Central Italy, Central Apennines), where a marine tetrapod reptile trackway was found. The Southern Alpine sites occur in four distinct stratigraphic levels of the Lower Jurassic Calcari Grigi Group whereas the Mattinata site is provisionally assigned to the Upper Jurassic Sannicandro Formation (Kimmeridgian-Tithonian).
In this review, we provide geographical and geological setting, age, ichnotaxonomy and the possible identity of trackmakers for each site. Additionally, we report on the history of discovery, the state of the art in ichnological knowledge and the evolution of methodological approaches and techniques adopted through time.
... The tracks are associated with invertebrate burrows, mudcracks, and wrinkle structures. The latter can be interpreted as the result of ancient microbial mats that primarily may have facilitated the preservation of vertebrate tracks (Marty et al., 2009;Noffke, 2010;Petti et al., 2011). To our knowledge, this tetrapod footprint slab has only been mentioned and figured once (Newell et al., 1976, pp. ...
Tetrapod footprints have long been known from Permian outcrops of the Cheguimi Sandstone in southern Tunisia, but never described in detail. The only recovered specimen preserves vertebrate tracks of two different biospecies. Ichnomorphotype 1 shows affinity with mid-Permian tetrapod tracks from France (Merifontichnus Gand et al., 2000) and South Africa (unnamed tracks); ichnomorphotype 2 is most similar to Middle to Late Permian tracks of France (Planipes Gand et al., 1995) and Italy (Dicynodontipus Rühle von Lilienstern, 1944), respectively. Both kinds of the Tunisian tetrapod tracks can be referred to non-mammalian therapsid trackmakers. The ichnoassemblage is in agreement with the supposed late Middle to early Late Permian (∼Capitanian to Wuchiapingian) age of the Cheguimi Sandstone, although additional finds are necessary to obtain a more precise age control.
... This footprint, despite the poor preservation and the lack of the distal digit III, is similar to the morphotype described in the Coste dell'Anglone site (Petti et al., 2011) and assigned to Kayentapus Welles, 1971. Despite the similarities, however, we prefer not to assign an ichnotaxonomical attribution for Pelm.A1, which is therefore classified as an indeterminate theropod track. ...
Dinosaur footprints from the Lower Jurassic of northeastern Italy are well known and, since the first discoveries in the early 1990s, many sites have been described. Tracks are mostly found in the peritidal limestones of the Calcari Grigi Group, deposited on the Trento carbonate platform, now cropping out in the Southern Alps. In 2011, a group of speleologists discovered a new tracksite in the Lower Jurassic Calcari Grigi Group exposed almost at the top of Mt. Pelmo (Dolomites), 3037 m above sea level. Footprints are generally poorly preserved, but it proved possible to recognise some tridactyl footprints with theropodian features (i.e., elongated digit III and narrow interdigital angle) and some possible quadruped tracks whose configuration resembles that of a sauropodomorph trackmaker. Careful examination of the depressions excludes their inorganic origin (chemical weathering). Despite the poor quality of the traces, the Pelmo site is significant because it is the most easterly site ever found on the Trento Platform and the only one which is located north of the Valsugana Fault. This fault system is a major alpine tectonic lineament that separates the classical successions of the Calcari Grigi Group in the Italian Prealps from those located in the Dolomites. Moreover, the discovery of the Pelmo tracks considerably expands the documented area of movement of Early Jurassic terrestrial vertebrates in the northern part of the Trento Platform, extending the size of the Early Jurassic megatracksites of the Southern Alps. RIASSUNTO-[Orme di dinosauro dalla cima del Mt. Pelmo: nuovi dati sulla paleogeografia del Giurassico Inferiore delle Dolomiti (NE Italia)]-Le impronte di dinosauri del Giurassico Inferiore del nordest italiano sono ben note e, dopo la prima scoperta all'inizio degli anni Novanta, sono stati riconosciuti diversi siti. Le impronte si trovano nei calcari pertitidali del Gruppo dei Calcari Grigi, depositatosi sulla piattaforma di Trento e affiorante oggi nelle Alpi Meridionali. Nel 2011, un gruppo di speleologi scoprì un nuovo sito nel Giurassico Inferiore dei Calcari Grigi, situato a 3037 m sul livello del mare, quasi alla sommità del Mt. Pelmo (Dolomiti). La qualità di conservazione delle orme è generalmente scarsa, tuttavia è possibile riconoscere alcune orme tridattile con caratteristiche teropodiane (dito III molto allungato, angolo interdigitale stretto) e alcune orme quadrupedi con una configurazione simile a quella dei sauropodi (o sauropodomorfi). L'attenta analisi delle depressioni esclude la loro origine inorganica (erosione chimica, carsismo). Nonostante la scarsa qualità delle orme, la scoperta del Pelmo è significativa perché il sito è il più orientale mai rinvenuto e l'unico situato a nord della Linea della Valsugana, un importante lineamento tettonico che separa le classiche successioni dei Calcari Grigi nelle Prealpi da quelle delle Dolomiti. Grazie al sito del Pelmo l'area dove si riscontra la presenza dei vertebrati nel Giurassico Inferiore si allarga alla parte settentrionale della Piattaforma di Trento, estendendo le dimensioni del mega-sito ad impronte del Giurassico Inferiore delle Alpi Meridionali.
... Within this general defi nition, we can fi nd the classical ichnogenera Grallator, Anchisauripus and Eubrontes (Hitchcock, 1858; Lull, 1904; Olsen et al., 1998). These kinds of tracks have been described worldwide from several tracksites and deposits of the Late Triassic and Early Jurassic (Gierliński, 1991; Gierliński & Alhberg, 1994; Olsen et al., 1998; Thulborn, 2000; Gatesy et al., 1999; Lucas et al., 2001; Gaston et al., 2003; Milàn et al., 2004; Clark et al., 2005; Klein & Haubold, 2007; Petti et al., 2011). Furthermore, in recent years, grallatorid-like tracks have been described from younger deposits in the Middle Jurassic of Argentina (De Valais, 2011), the Late Jurassic and Early Cretaceous of Spain (Piñuela, 2000; Lockley et al., 2008; Pascual-Arribas & Hernández-Medrano, 2012; Avanzini et al., 2012; Piñuela, 2015), the Late Jurassic of Germany (Diedrich, 2011) and the USA (), and the Late Jurassic and Early Cretaceous of Asia (Lockley et al., 2013Lockley et al., , 2015). ...
The MUJA (Museo del Jurásico de Asturias, Jurassic Museum of Asturias) has an interesting collection of theropod tracks that show similarities with the ichnogenera assigned to the Eubrontes-Grallator plexus. In this paper we describe in detail the morphology of 21 specimens recovered from different localities on “The Dinosaur Coast” of Asturias, plus four specimens preserved in outcrops in the sea cliffs of Les Vinaes (Villaviciosa). All the specimens are from the outcrops of the Lastres Formation, which is Kimmeridgian in age. The general morphology of the tracks, the footprint length-width ratio, the mesaxony, low divarication of the digits (II-IV) and the absence of hallux and metatarsophalangeal impressions suggest that the tracks are more similar to Grallator than to any other theropod ichnotaxa. Geometric morphometric analysis (principal component analysis, PCA) based on 2D landmark techniques suggests that they differ from Kalohipus bretunensis (as yet the only Grallator-like ichnotaxon described in the Iberian Peninsula) mainly in the divarication angles and in the projection of digit III.
... Several other ichnotaxa are considered either behavioural/preservational variations (e.g., Gigandipus) or junior synonyms of the four commonly considered valid ichnogenera above. Size has long been the key separating criteria in the GrallatorAnchisauripusEubrontes or GAE allometric plexus (Olsen, Smith & McDonald, 1998; Rainforth, 2005; Petti et al., 2011), and synonymising the three ichnogenera into a single valid ichnogenus (Grallator or Eubrontes) has been long considered and debated (e.g., Olsen, 1980; Olsen, Smith & McDonald, 1998; Rainforth, 2005; Lucas et al., 2006). The ichnogenera Kayentapus and Anchisauripus have been incorporated into the GrallatorEubrontes spectrum largely because of their intermediate size between the two ichnogenera (Olsen, Smith & McDonald, 1998; Lockley, 1998; Milner et al., 2009). ...
Footprint morphology (e.g., outline shape, depth of impression) is one of the key diagnostic
features used in the interpretation of ancient vertebrate tracks. Over 80 tridactyl
tracks, confined to the same bedding surface in the Lower Jurassic Elliot Formation
at Mafube (eastern Free State, South Africa), show large shape variability over
the length of the study site. These morphological differences are considered here to
be mainly due to variations in the substrate rheology as opposed to differences in the
trackmaker's foot anatomy, foot kinematics or recent weathering of the bedding surface.
The sedimentary structures (e.g., desiccation cracks, ripple marks) preserved
in association with and within some of the Mafube tracks suggest that the imprints
were produced essentially contemporaneous and are true dinosaur tracks rather than
undertracks or erosional remnants. They are therefore valuable not only for the interpretation
of the ancient environment (i.e., seasonally dry river channels) but also
for taxonomic assessments as some of them closely resemble the original anatomy of
the trackmaker's foot. The tracks are grouped, based on size, into two morphotypes
that can be identified as Eubrontes-like and Grallator-like ichnogenera. The Mafube
morphotypes are tentatively attributable to large and small tridactyl theropod trackmakers,
possibly to Dracovenator and Coelophysis based on the following criteria: (a)
lack of manus impressions indicative of obligate bipeds; (b) long, slender-digits that
are asymmetrical and taper; (c) often end in a claw impression or point; and (d) the
tracks that are longer than broad. To enable high-resolution preservation, curation
and subsequent remote studying of the morphological variations of and the secondary
features in the tracks, low viscosity silicone rubber was used to generate casts of the
Mafube tracks.
... Matthews, 2008). For this reason DP has been used in many different areas of research, such as topography, architecture (e.g., Grussenmeyer et al., 2002), archaeology (e.g., Sumner AND Riddle, 2008), and of course also palaeontology/ichnology (e.g., Matthews, 2008;Petti et al., 2011;Rowland et al., 2014). The development of DP, in particular of the SFM (Structure from Motion) method, its integration with the methods of classic photogrammetric mapping and the possibility of post-processing and analysis GIS/CAD data, made DP an excellent tool for the production of clouds of points and textured 3D models. ...
Pleistocene carbonate eolianites crop out along the Portuguese Atlantic coast, in both mainland and Madeira islands in Eastern Atlantic. More continuous outcrops are found at the SW coast of Portugal including the only recorded Cenozoic vertebrate ichnosites from Portugal so far. The vertebrate paleoichnology of these formations have yielded at least 16 stratigraphic horizons distributed by 6 tracksites, with mammal and bird footprints and trackways that were found printed in the sandstone bedsets. Photogrammetric analysis and 3D modeling of the most relevant tracksites allowed revising and supporting previous ichnotaxonomic identifications and behavioral interpretations. Small and large mammals’ trackways and footprints attributed to Proboscipeda panfamilia, Bifidipes isp., Bestiopeda isp., Felipeda lynxi, and Leporidichnites malhaoi are described and discussed according to new and recent findings. Probably producers were, respectively, straight-tusked elephant, red deer, fox and wolf, Iberian lynx, and rabbit or hare. Tracks of birds found in two horizons at Pessegueiro island are now ascribed to two morphotypes of Charadriipeda isp.. These tracksites are dated from MIS6?-MIS3-2?, mostly Late Pleistocene.Among those footprints, elephant tracks are particularly important, since it is the first record of elephant footprints in the Pleistocene of mainland Europe and may represent some of the latest occurrences of Elephas antiquus before its final extinction. New data enable to detail further the importance of this vertebrate track record for the changing of the community dynamics, biogeographical isolation and extinctions in the transition for the Last Glacial.
... The depositional environment of the trackbearing bed is here interpreted as nearshore, supported by the presence of gastropod shell fragments and the occurrence, a few metres below, of the trace fossil Diplocraterium, representative of shallow-marine, high energy nearshore environment (Seilacher, 2007). Dinosaur footprints are occasionally preserved in microbial mats, typical of lagoonal and arid environments, which are believed to aid the track preservation (Marty et al., 2009;Noffke, 2010;Petti et al., 2011); the others Tunisian vertebrate tracksites occur in similar lagoonal to near shore environments (Contessi and Fanti, 2012b). ...
New dinosaur tracks discovered at the Chenini site, near Tataouine (southern Tunisia), are described. Isolated tracks as well as manus/pes couples of a quadrupedal dinosaur are referred to a hadrosaurian tracemaker and assigned to cf. Apulosauripus federicianus (Nicosia et al., 2000b), an ichnospecies originally described in the Upper Cretaceous carbonate platform deposits of southern Italy. This discovery thus represents the first report of hadrosaurian tracks in Africa.
The occurrence of tracks similar to A. federicianus in Tunisia also represents the first evidence of the presence of comparable ichnofauna in the Upper Cretaceous of both northern Africa and the carbonate platforms of the Mediterranean area. Supporting these new data, the Cenomanian theropod ichnofauna recently described from southern Tunisia is here compared with coeval theropod tracks from the peri-Adriatic carbonate platforms. This comparison documents a number of shared morphological features, suggesting possible affinities of the trackmakers and supporting the hypothesis of a subaerial connection linking the carbonate platforms of the Mediterranean area to the northern margins of Africa in the mid-Cretaceous.
... 3A and B). This feature, together with the early dolomitization, is indicative of arid tidal flat environments, and may have aided in the preservation of the tracks (Marty et al., 2009;Petti et al., 2011). Deeply impressed, non-avian theropod tracks occur on the block (Contessi and Fanti, 2012b), suggesting a high water content in the sediment. ...
... White arrows indicate shell hash and possible remains of benthic foraminifera. footprints and microbial mats has been observed in a number of track-bearing localities worldwide (Avanzini et al., 1997; Kvale et al., 2001; Marty et al., 2009; Noffke, 2010; Petti et al., 2011a; Contessi and Fanti, 2012a) and hypothesized to facilitate the preservation of tracks. The presence of dinosaur footprints within this setting supports the presence of nearby emerged landmasses during Oxfordian times, which otherwise would have been difficult to hypothesize. ...
... Saurischian track assemblages are relatively common in carbonate tidal flats deposits on a global scale (Meyer, 1993; Lockley and Hunt, 1994; Lockley et al., 1994; Meyer and Pittmann, 1994; Lockley et al., 1996; Marty, 2008; Lucas, 2009; Sacchi et al., 2009, Contessi and Fanti, 2012b, and references therein) supporting a systematic presence of largebodied dinosaurs in marginal marine ecosystems. In particular, dinosaur tracksites characterized by exclusively theropod tracks are relatively frequent in the Jurassic track record worldwide, and the predominant paleoenvironments in which these tracks occur are shorelines, tidal flats or shallow lagoons (Lockley et al, 2000; Conti et al., 2005; Leonardi and Dos Santos, 2006; Diedrich, 2008, 2011; Schulp et al., 2008; Boutakiout et al. 2009; Milner et al., 2009; Petti et al., 2011b; Xing et al., 2011). Differently, more variegate assemblages which include avian and nonavian theropod, sauropod, and ornithopod tracks are associated with a wider spectrum of depositional settings, such as carbonate platforms, floodplains, lacustrine, and lagoonal environments (Ga´zdzicka et al., 2001; Day et al., 2004; Milner et al., 2006; Avanzini, 2006; Leonardi et al., 2007; Marty, 2008; Mateus and M` ılan, 2009; Belvedere et al., 2010; Wagensommer et al., 2011). ...
In this study, we present new data on the Hojapil-Ata dinosaur
mega-tracksite located in the Koitendag National Park
of eastern Turkmenistan, central Asia. Accurate GPS mapping,
three-dimensional data on preserved tracks, and comparison with
previous studies provides now a detailed database for this site. The
track-bearing surface, Upper Jurassic in age, is discontinuously
exposed over an area of 28.500 square meters, with 24 trackways
and at least 913 footprints preserved. The collected data support
the previously identified co-occurrence of the two ichnogenera
Megalosauripus and Therangospodus and allow us to propose an
amended systematic description of M. uzbekistanicus. In addition,
a previously unreported site located a few km to the east of the
main tracksite is described. The track-bearing surface extends
over 24.000 square meters, and 34 trackways and 730 footprints
have been mapped. These two sites represent the sole evidence
of dinosaurs in Turkmenistan and they are among the largest
in the world. Data collected for this study document that both
localities are deteriorating at an increasing rate due to natural
erosion and human activities. High-resolution laser scanner and
digital photogrammetry allowed for the realization of comparable
digital 3D models of
... The track-bearing surface also preserves mud cracks and a distinctive texture interpreted to represent ancient microbial mats (Fig. 3A), virtually identical to those observed in modern tidal-flat environments (Fig. 3B). The co-occurrence of dinosaur footprints and microbial mats within shallow marine and lagoonal environments has been observed in other track-bearing localities (Kvale et al., 2001;Marty et al., 2009;Petti et al., 2011) and hypothesized to facilitate the preservation of the tracks avoiding weathering and erosion. The trampled surface Downloaded by [University of Calgary] at 05:37 07 January 2013 is overlain by fine, horizontally-laminated marls that have been largely eroded. ...
... The track-bearing surface occurs at the top of a 60-cm-thick, fining-upward sequence of wackestone and finely-laminated grainstone capped by a 3-cm-thick stromatolitic lamination. Thin sections of the track surface show shell fragments, extensive dolomitization, birdseye structures and a fenestral fabric, indicative of arid climatic conditions (Petti et al., 2011). As at the Beni Ghedir locality, the track-bearing surface shows a distinctive texture indicative of the presence of ancient microbial mats. ...
Four vertebrate tracksites from the Middle Jurassic and
Upper Cretaceous in the Tataouine basin of southern Tunisia
are described. Approximately 130 tridactyl footprints distributed
over an area of 200 square meters, preserved on Callovian beds
exposed at the Beni Ghedir site, represent the oldest evidence of a
dinosaur fauna in Tunisia. In addition, three tracksites—Chenini,
Ksar Ayaat, and Jebel Boulouha—have been discovered in the
Cretaceous beds of the upper Continental Intercalaire, previously
considered as a strictly marine depositional sequence. In addition
to dinosaur tracks, the Chenini tracksite (late Albian) includes
poorly preserved crocodilian tracks, and footprints assigned to a
pleurodiran turtle have been recovered at the Ksar Ayaat locality
(early Cenomanian). The Jebel Boulouha tracksite is dominated by
well-preserved tridactyl tracks referred to small-sized theropods.
Depositional settings of each tracksite have been defined on
stratigraphic and sedimentologic data, and tracks were ascribed to
different ichnocoenoses in relation to their paleoenvironments. This
new and differentiated track record gives important information
on how the fossil vertebrate fauna changed in southern Tunisia
during mid-Jurassic to mid-Cretaceous times. These data provide
a unique and useful census of tetrapod associations along the
southern margin of the peri-Mediterranean area.
