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Vertebrate footprints from the Kem Kem beds (Morocco): A novel ichnological approach to faunal reconstruction
Abstract
Abstract The first survey of vertebrate fossil tracks and trackways from the well-known Kem Kem beds of southern Morocco is here presented. The aims are to: (1) discuss the diversity of vertebrate trackmakers represented in two trace-yielding units at three localities, and (2) apply the novel data to ongoing debates about vertebrate ecological associations within the Kem Kem. The majority of the tracks we recorded from the Kem Kem are tridactyl theropod footprints; other dinosaurian track records include those of possible ornithopod dinosaurs, extremely rare as body fossils within Kem Kem collections. Traces of swimming turtles are reported, alongside the first tracks of crocodyliforms and possible pterosaurs to be recorded from the Moroccan Cretaceous (both unsurprising given their abundance as body fossil records in this region). Differences between the collected ichnological sample and faunal reconstructions made on the basis of skeletal evidence are discussed; as trace fossils record the same environment in which the trackmaker lived (i.e. not subject to post-mortem transportation), data of this type arguably provide a more precise palaeoecological sample than the heavily re-worked and usually fragmentary body fossils from the Kem Kem.
... More recently, Vila et al. (2015) identified and analyzed scratch marks made by buoyant crocodiles, thus allowing them to provide significant data on the paleoecology and ichnofacies of the latest Cretaceous environments in south-western Europe. In North Africa, there is a limited description of Mesozoic crocodylian swim traces, which have been reported to date from the 'mid'-Cretaceous Midelt Formation, in the High Moulouya region (Klein et al., 2018), and from the Upper Cretaceous, Cenomanian Kem Kem Group (Belvedere et al., 2013;Ibrahim et al., 2014) of Morocco. In Algeria, despite the abundance of crocodylian body fossils and coprolites among the Mesozoic vertebrate fauna (Bendella et al., 2023;Benyoucef et al., 2015Benyoucef et al., , 2022Benyoucef et al., , 2023, their trackways have been unknown so far. ...
... The crocodylian ichnogenus Hatcherichnus was originally introduced with the ichnospecies H. sanjuanensis by Foster & Lockley (1997) based on trackways from the Upper Jurassic Morrison Formation of North America and was described as a tetradactyl pes impression associated with a tridactyl manus impression. Moreover, similar traces have also been described from the Dakota Group (Lower Cretaceous) of North America (Kukihara et al., 2010;Lockley, 2010;Lockley et al., 2010), the Upper Jurassic of Asturias, Spain , and from 'Mid'-Upper Cretaceous deposits of Morocco (see Belvedere et al., 2013;Ibrahim et al., 2014;Klein et al., 2018). Usually, in more complete Hatcherichnus, a tetradactyl manus, as well as a body and tail trace, are frequently associated (Foster & Lockley, 1997). ...
... In North Africa, Mesozoic crocodylomorph swim traces are uncommon, being described so far only from Morocco (e.g. Belvedere et al., 2013;Ibrahim et al., 2014;Klein et al., 2018). There are also similarities with the traces of swimming pterosaurs, although these imprints vary largely by the number and orientation of claw impressions (Lockley & Wright, 2003). ...
We report tetrapod traces and associated ichnofauna from two stratigraphic, dominantly terrestrial levels of the Tiout Formation (Valanginian-latest Albian to lower Cenomanian) in Laghouat and Brezina areas, Central Saharan Atlas (Djebel Amour), northwestern Algeria and discuss their paleoecological implications. The assemblages display abundant footprints that show similarities with crocodylian swim traces assigned to Hatcherichnus known from Jurassic–Cretaceous deposits of North America and Morocco. However, there is also some similarity in shape with traces attributed to swimming pterosaurs by different authors. Because of the isolated materials from Algeria and the lack of distinct trackways, we leave the decision on the tracemaker open. The majority of the traces are tridactyl to tetradactyl imprints consisting of parallel furrows left by the claws of swimming or buoyant individuals. Tetrapod swim traces are identified, described, and figured herein for the first time from the Mesozoic of Algeria. These vertebrate fossil traces are associated with a low-diversity invertebrate marine ichnofauna, including cf. Bergaueria isp., Phycodes isp., Sinusichnus cf. seilacheri, and Thalassinoides suevicus. Together with body fossil data, including abundant fishes and non-avian dinosaurs, they indicate a diverse animal community populating a fluvial system environment with marine influence. Paleoecological and paleoenvironmental features of the Tiout Formation add new information to the ichnoassemblages previously reported from the ‘mid’-Cretaceous of North Africa.
... The Kem Kem Groups two formations, the Ifezouane Formation, and overlying Aoufous Formation, differ in their lithology, but the transition between these two formations is gradational, and not easy to pin point. The Ifezouane Formation is characterised by red-brown fluvial sandstones, with high-angle, metres-thick cross-bedded units representing broad channel fills, with intercalations of pedogenised overbank mudstones (Belvedere et al., 2013), pinkish sandstones and conglomerates with distinctive amber coloured quartz pebbles (Cavin et al., 2010). Of particular note are thin (generally <1 m) event horizons of mudstone intraclasts in chaotically bedded ossiferous sandstones, the source of the majority of Kem Kem Group vertebrate fossils. ...
... The Aoufous Formation is characterised by variegated mudstones and marls, with calcitic palaeosols, thin intercalations of evaporites, detrital sandstone and micro-conglomerates, and carbonate-cemented mudstones (Cavin et al., 2010;Belvedere et al., 2013). The vast majority of vertebrate fossils (including all pterosaurs) come from intraformational, mud-flake conglomerates in the upper Ifezouane Formation (Martill et al., 2018). ...
... Rare, short-lived shallow marine environments occur in some northern localities as at Tarda and Zrigat (Beevor et al., 2021) and in the Atlas fold belt (Adardor et al., 2021). Overall, the sequence records a fluvial to marine transgression, terminating in a thick sequence of fully marine carbonates, represented by the Cenomanian-Turonian Akrabou Formation (Ettachfini and Andreu, 2004;Ibrahim et al., 2010;Belvedere et al., 2013). However, there is a sharp contact between the KKG and Akrabou Formation, with no discernible brackish facies in between, suggesting a possibly significant time hiatus between the two formations. ...
... Although a partial edentulous mandibular symphysis (MN 7054-V) was originally regarded as a pteranodontid by Kellner et al. (2007a), it was later interpreted as a tapejarid by Ibrahim et al. (2020), and more recently a referred specimen of Afrotapejara zouhrii by Smith et al. (2022). The Kem Kem Group comprises the Ifezouane and Aoufous formations, and has been interpreted to represent a fluvial to deltaic setting (Belvedere et al., 2013;Martill and Ibrahim, 2015;Wellnhofer and Buffetaut, 1999a). For more precise information regarding the stratigraphy see Ibrahim et al. (2020), and Rodrigues et al. (2011) and references therein. ...
... Possible pterosaur manus tracks have been reported from the Kem Kem Group by Belvedere et al. (2013). As in other pterosaur manus tracks, digit I is positioned anteriorly and short, whereas digit III is the longest and directed laterally (Belvedere et al., 2013). ...
... Possible pterosaur manus tracks have been reported from the Kem Kem Group by Belvedere et al. (2013). As in other pterosaur manus tracks, digit I is positioned anteriorly and short, whereas digit III is the longest and directed laterally (Belvedere et al., 2013). Given that the tracks are poorly preserved and do not form a trackway, Belvedere et al. (2013) did not assign these to an ichnotaxon. ...
The Gondwanan pterosaur record is scarce when compared with that of Laurasia and is reviewed here. The majority of Gondwanan pterosaur remains are derived from South America; however, the relative richness of the South American record compared with other Gondwanan continents is largely a result of the ‘Lagerstätten’ effect. Nevertheless, the South American pterosaur assemblage represents the most speciose and diverse known from Gondwana, with several lineages represented, including the Raeticodactylidae, Rhamphorhynchoidea, Darwinoptera, Ctenochasmatidae, Gnathosaurinae, Nyctosauridae, Ornithocheiridae, Tapejaridae, Thalassodromidae, Dsungaripteridae, Chaoyangopteridae and Azhdarchidae. Gondwanan pterosauromorphs are known only from South America. From Africa rhamphorhynchids, archaeopterodactyloids, pteranodontians, nyctosaurids, ornithocheirids, tapejarids, dsungaripteroids, chaoyangopterids, and azhdarchids have been reported. The Arabian Peninsula has produced nyctosaurids, an istiodactyliform, ornithocheirids and azhdarchids. Non-pterodactyloid pterosaurs have been reported from India. A possible azhdarchid has been reported from Madagascar and rhamphorhynchids are known from isolated teeth. The Antarctic pterosaur assemblage also comprises isolated remains of indeterminate pterodactyloids, and a possible indeterminate rhamphorhynchoid. The pterosaur record from East Gondwana comprises ornithocheirids, azhdarchids and a possible ctenochasmatoid from Australia, as well as azhdarchids from New Zealand. Although our understanding of Gondwanan pterosaurs has greatly improved within the last three decades, the discovery and description of more specimens, particularly from Antarctica and East Gondwana, will enhance our understanding of pterosaurian biodiversity and palaeobiogeography.
... Most of these remains are currently under study, and among these, four pterosaur manus tracks are the main goal of this contribution. The Gondwanan pterosaur ichnological record is still poorly documented [27][28][29][30][31][32], whereas the bone record is very abundant, especially in South America, and particularly in Argentina and Brazil, with some records in Chile, Peru and Venezuela [33][34][35][36][37][38][39][40][41][42][43][44][45][46][47][48][49][50][51][52]. The aims are to describe, compare and analyze the new pterosaur tracks found at the ANPM Paso Córdoba and then to discuss the important role that they have in the reconstruction of the paleocommunity of the Anacleto Formation and for the latest Cretaceous of Gondwana. ...
... The pterosaurian track record, spanning from the Middle Jurassic to the Late Cretaceous [30], is relatively scarce in comparison with those linked to other groups of Mesozoic vertebrates such as dinosaurs ( [110,111] but see [30] for a different opinion). This scarcity is particularly evident in Gondwana, where only five pterosaur track-bearing lithostratigraphic units were known before the present study, four in Morocco [28][29][30][31] and one in Argentina [27,32]. If they are described in chronological order, among the Moroccan sites, brief mentions about pterosaur tracks assigned to Pteraichnus are made in the Middle-?Upper Jurassic Isli Formation [31,112]. ...
... If they are described in chronological order, among the Moroccan sites, brief mentions about pterosaur tracks assigned to Pteraichnus are made in the Middle-?Upper Jurassic Isli Formation [31,112]. Some poorly preserved tracks putatively assigned to pterosaurs were identified in the Cenomanian Kem Kem beds [28], but the affinity of this material has subsequently been questioned [30]. In Argentina, two pterosaur track-bearing outcrops are present in the Cenomanian Candeleros Formation of Patagonia. ...
The Campanian Anacleto Formation holds an abundant and diverse ichnofossil and body-fossil vertebrate record. Despite the striking diversity of this record, pterosaur fossils had never been described from the unit. Here, we report four pterosaur manus tracks from fluvial red beds cropping out in the Área Natural Protegida Municipal Paso Córdoba (Río Negro Province, northern Patagonia, Argentina). Tracks are longer than wide, tridactyl with digit impressions of different lengths (I < II < III), anteriorly directed and laterally asymmetrical. Being on loose slabs and lacking direct examination of pes morphology, the material is classified as undetermined pterosaur tracks. The new find represents the first occurrence of pterosaurs from the lower–middle Campanian of Argentina and one of the few evidences from South America for this time interval. In addition, it is one of the few ichnological pterosaur records from Gondwana, thus shedding light on the palaeobiogeography of this clade during the latest Cretaceous. Pterosaur tracks from the Anacleto Formation allow us to integrate the body-fossil record from the unit and to add a new component, along with birds, to the flying archosaur fauna coexisting with non-avian dinosaurs, notosuchians, chelonians, squamates and mammals in the Campanian of northern Patagonia.
... transjuranicus) from Central High Atlas (Belvedere et al., 2010); xxvii) Boutakioutichnium atlasicus from Central High Atlas (Nouri et al., 2011); xxviii) GSB6 (cf. Megalosauripus) from Kem Kem beds (redrawn from Belvedere et al., 2013); xxix) morphotype "Ensemble 1" (cf. Megalosauripus) from Saharan Atlas (Bessedik et al., 2019; xxx) morphotype "Ensemble 2" (cf. ...
... The palaeogeographic position of the peri-Adriatic region close to the northern Gondwana margin suggests a comparison between the Molfetta specimens and those known from Northern Africa. Several theropod tracks were reported from Upper Jurassic to Upper Cretaceous deposits of Morocco (Boutakiout et al., 2009(Boutakiout et al., , 2018Belvedere et al., 2010Belvedere et al., , 2013Belvedere et al., , 2019Nouri et al., 2011;Masrour et al., 2013;Ibrahim et al., 2014Ibrahim et al., , 2020 and Algeria (Bensalah et al., 2005;Mahboubi et al., 2007;Bessedik et al., 2008;Chabou et al., 2015;Mammeri, 2018). Several dinosaur tracks are known from the Iouarid ene Fm. (Central High Atlas, Morocco), dated ?Oxfordian-Kimmeridgian. Boutakiout et al. (2009) and Belvedere et al. (2010) described giant theropod footprints (FL > 50 cm; type specimen: 23IGR1.7; ...
... Other theropod tracks in Morocco come from the Upper Unit of the Kem Kem beds (Cenomanian), coeval to the numerous body fossils discovered in the same Group (Ibrahim et al., 2020 and references therein). Belvedere et al. (2013) analyse several tetrapod natural casts (attributed to theropods, ornithopods and turtles) from Gara Sbaa (southern Morocco). Despite poor preservation, the bestpreserved tridactyl track (i.e., GSB 6 specimen; Fig. 15xxviii) was classified as cf. ...
The lower Albian track-bearing surface of the San Leonardo quarry (Molfetta, Apulia) is characterised by more than 800 footprints, produced by both quadrupedal and bipedal dinosaurs. Six well-preserved bipedal trackways, composed of tridactyl footprints are attributed to medium-to large-sized theropod dinosaurs. Only one clear but poorly preserved trackway and numerous isolated manus-pes couples have been attributed to quadrupedal dinosaurs. The tridactyl ichnoassemblage, analysed using both traditional methods and close-range photogrammetry, is represented by weakly mesaxonic and robust specimens. Morphological comparison with Upper Jurassic and Lower Cretaceous theropod tracks from surrounding areas, supported by morphometric analyses, points out a highest affinity with the specimens from Switzerland and North Africa. Nevertheless, a set of unique characters appears to justify the establishment of a new ichnospecies, Jurabrontes melphicticus. Additionally, the photogrammetric models of the quadrupedal trackway and four isolated manus-pes sets suggest they belong to the same morphotype, represented by asymmetrical tetradactyl pes and highly digitigrade tetra- or pentadactyl manus. These tracks share numerous morphological characters with both the ichnogenera Tetrapodosaurus and Metatetrapodus and thus can be attributed to a medium-sized ankylosaurian trackmaker.
... 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. ...
... Lagnaoui, 2014;Lagnaoui et al., 2016). Middle Jurassic-Cretaceous (Bathonian-Maastrichtian) assemblages mostly yield theropod, sauropod and ornithischian tracks as well as the footprints of pterosaurs (Ishigaki, 1989;Gierliński, 2016;Gierliński et al., 2009Gierliński et al., , 2017aGierliński et al., , 2017bMarty et al., 2010;Hadri and Pérez-Lorente, 2012;Belvedere et al., , 2013Ibrahim et al., 2014;Masrour and Pérez-Lorente, 2015;Masrour et al., 2013Masrour et al., , 2017aMasrour et al., , 2017bMasrour et al., , 2018. However, most of those works focused on the western and central part of the Central High Atlas, leaving the studied area in the eastern part, the Imilchil and Midelt region, less recognized ichnologically. ...
... Crocodyliform swim traces have also been reported from the Kem Kem beds (Upper Cretaceous, Cenomanian) of Morocco (Belvedere et al., 2013;Ibrahim et al., 2014). Here, they co-occur with the footprints of theropod and ornithopod dinosaurs, pterosaurs, sauropods and those of possible turtles. ...
... Most of these tracks correspond to marks on wet sediments (e.g. Haubold, 1971aHaubold, , 1984Bernier et al., 1982;Foster et al., 1999;Mazin et al., 2000;Wright and Lockley, 2001), while turtle swimming traces are scarce (Gaillard et al., 2003;Dentzien-Dias et al., 2010;Belvedere et al., 2013). The Mesozoic record of turtles in the Iberian Pensinsula contains no record of fossil remains during the Triassic, the oldest remains being those asigned to Hispaniachelys prebetica (Slater et al., 2011), a marine turtle from the Upper Oxfordian of the Prebetic (Betic Cordillera, South Spain). ...
... The pes is functionally digitigrade and tetradactyl, outwardly rotated (e.g. Belvedere et al., 2013). Potential swimming tracks of crocodiles are not congruent due to the fact that they use their tails more than their feet for swimming. ...
... Emydhipus is known from the Upper Jurassic and Lower Cretaceous of Spain (e.g. Fuentes Vidarte et al., 2003;Avanzini et al., 2005;Moratalla and Hernán, 2009) and Morocco (Belvedere et al., 2013), and the Early Cretaceous of Xinjiang, China (Xing et al., 2014). The incomplete preservation of manus-pes couples and the lack of a trackway make it impossible to give a definitive ichnotaxonomic assignment for the Cortes de Pallas and Domeño outcrops. ...
Some of the earliest European records of fossil turtle footprints (Late Triassic, Middle Carnian, ~227–237 Ma)
are interpreted from 46 footprints from three outcrops, Domeño, Quesa and Cortes de Pallas, located in the
Iberian Range (eastern Spain). The samples were obtained from Upper Triassic rocks in Keuper Facies. They are
characterized in the studied area by two well-defined evaporitic sequences, separated by a detrital stratigraphic
interval, constituting the Manuel Sandstones Formation in which the studied fossil footprints were recorded.
These fluvial deposits are correlatable with the Carnian Humid Episode. The footprints are tridactyl and tetradactyl,
mainly digitigrade, and elongated scratch marks are common. The digit traces present triangular and
curved claws. The morphology of these marks suggests they were produced by the autopodium of a tetrapod
animal while swimming near the bottom, and they are consistent with the anatomical features of a turtle's
manus. Other footprints were produced by turtles walking in wet medium- to coarse-grained sand in a subaerial
environment. A short trackway is identified, sharing similarities with both the ichnogenera Emydhipus and
Chelonipus. The observed tracks support a freshwater semi-aquatic habit for some stem turtles during the early
Late Triassic. These are remarkably ancient turtle tracks, close to the origin of the group, as indicated by skeletal
fossils; they confirm that the earliest forms were narrowly related to aquatic (fluvial) environments.
... has allowed us to maintain this ichnogenus and ichnospecies, and reaffirm that they constitute the first footprints of pterosaurs ever described three years before the report of Stokes (1957) from the western USA. It is possible that there is another site (Kem-Kem, Cenomanian/Turonian) with pterosaur footprints, cited by Belvedere et al. (2013). ...
... This is the third confirmed discovery of pterosaur tracks in Morocco. The morphological characters of the Kem-Kem pterosaur prints are similar to those of the turtle footprints that were also described by Belvedere et al. (2013) from the same site. This exceptional circumstance and the fact that they are the first ones found from the Coniacian-Santonian age, make us consider it appropriate to publish, describe and relate them with those already known, both from the Upper Cretaceous age and those of earlier times. ...
... The other possible pterosaur footprints cited in Morocco (Kem Kem site, Belvedere et al., 2013), show very unclear pterosaurian characteristics (Fig. 8), resembling some turtle species. Turtle tracks are very similar to those considered to pterosaur, so we doubt that they can be classified as such. ...
Cretaceous pterosaurs tracksites are very rare worldwide. Until now,only one African Cretaceous site withtracks of (Agadirichnus elegans and Pteraichnus) was known. This makes the discovery of a new outcrop in the Upper Cretaceous of Anza (Morocco) the third manifestation of this type of footprint in Africa, extending the existence of such traces from the Coniacian-Santonian to the Maastrichtian. The site contains only manus tracks, which can be explained as a result of erosion of pes prints. The lack of pes prints and the morphometric characteristics of the manus prints only allow us to relate these prints to Agadirichnus, Pteraichnus or maybe to a new ichnogenus. It is possible that the trackmakers are related to Ornithocheiroidea or Azhdarchoidea superfamilies whose fossil bones have been found from the Late Cretaceous in Morocco.
... For example, synchrotron X-ray micro-tomography (Laloy et al., 2013;Pradel et al., 2009;Rücklin et al., 2014) enables us to see through rock and to have access to internal structures. Conversely, three dimension (3D) surface analyses are generally acquired with surface scanners (e.g., Clarac et al., 2015;Falkingham, 2012) or photogrammetry (e.g., Belvedere et al., 2013;Brassey et al., 2015;Castanera et al., 2013;Falkingham, 2012;Petti et al., 2008). Photogrammetry is a method for reconstructing an object in 3D from photographs. ...
... Photogrammetry is already commonly used in some research fields like geology, for monitoring earth deformation from aerial or satellite photographs (e.g., Lane et al., 2000;Mora et al., 2003;Wolf and Dewin, 2000), in archeology to improve in situ topological or taphonomic data acquisition (De Reu et al., 2013, or for surveys of the cultural heritage (e.g., Guarnieri et al., 2004;Salonia et al., 2007Salonia et al., , 2009. In paleontology, photogrammetry has only been used for a few years Breithaupt et al., , 2004Matthews and Breithaupt, 2001), but 3D digitizing of both specimens and surfaces of excavation, in particular for ichnofossils, is increasingly used (e.g., Belvedere et al., 2013;Brassey et al., 2015;Castanera et al., 2013;Falkingham, 2012;Petti et al., 2008). There is no theoretical limitation of specimen size for using photogrammetry. ...
... Moreover, this software is used by paleontologists (e.g., Brassey et al., 2015;Castanera et al., 2013). Agisoft PhotoScan, also recently used in paleontology (e.g., Belvedere et al., 2013), is considered the most efficient (L. Cazes, pers. ...
Since the late 20th century, new technologies have provided powerful ways to digitize biological structures in three dimensions (3D). Among those, photogrammetry is a low cost and non-destructive method, which has become increasingly used since the development of the digital camera. Recent studies have demonstrated that reconstructions of isolated elements can be of as high quality as those obtained with laser scanners. Here, we wanted to test the performance of photogrammetry for the quantitative analysis of mounted specimens in museum exhibitions. Indeed, access to material can be an issue in comparative anatomy and, especially, in paleontology. This is notably the case for large, impressive specimens. We performed reconstructions based on acquisitions done under various conditions and also tested the reconstruction performance of two software programs. The resulting 3D models were then compared to a reference object corresponding to the bone of interest digitized with a cutting-edge surface scanner. Our results show that photogrammetry enables quality reconstruction of the almost entire surface of the mounted bone of interest. Photogrammetry thus appears a reliable method perfectly suited to study large specimens exposed in museum gallery.
... part consists predominantly of continental sediments (Ifezouane Formation), characterized by reddish fluvial detritic sandstones, with high angled, cross-bedding representing channel fills, intercalations of pedogenic overbank mudstones (Belvedere et al. 2013), and thin conglomerates with quartz pebbles (Cavin et al., 2010). The overlying claystones (Aoufous Formation) are characterized by variegated mudstones and marls, with calcitic palaeosols, thin intercalations of evaporites, detrital sandstone, micro-conglomerates, and carbonate cemented layers (Cavin et al., 2010;Belvedere et al., 2013). ...
... part consists predominantly of continental sediments (Ifezouane Formation), characterized by reddish fluvial detritic sandstones, with high angled, cross-bedding representing channel fills, intercalations of pedogenic overbank mudstones (Belvedere et al. 2013), and thin conglomerates with quartz pebbles (Cavin et al., 2010). The overlying claystones (Aoufous Formation) are characterized by variegated mudstones and marls, with calcitic palaeosols, thin intercalations of evaporites, detrital sandstone, micro-conglomerates, and carbonate cemented layers (Cavin et al., 2010;Belvedere et al., 2013). ...
The Cretaceous marine sediments of Morocco and adjacent coastal basins provide an outstanding archive of environmental diversity from extended shelf seas and marginal basins along the Atlantic and Tethyan margins to deep oceanic basins of the western Tethys and eastern Atlantic Ocean. The geological highlights of Morocco's fascinating landscape include records of Lower Cretaceous Tethyan marginal and deep water clastic sequences in the Rif mountain chain (submarine fan systems of the Massylian and Mauretanian flysch units), as well as siliciclastic sedimentary sequences in subsiding coastal basins (TanTan Delta), which extend offshore along the Northwest African Atlantic margin. Vestiges of the Aptian to Turonian greenhouse climate, sea-level highstands and oceanic anoxic events are exceptionally well-preserved in Tethyan marginal and deep water sedimentary successions of the Rif, in Atlantic coastal basins and as transgressive pulses on the Moroccan Meseta, Sahara platform and High Atlas rift system. Furthermore, sedimentary expressions of the tectonic movements between the African and European plates associated with the end Cretaceous climate and sea-level changes are documented in the Rif mountain chain, in coastal basins, and in the massive marginal marine phosphorite sedimentation in the Middle and High Atlas, on the Moroccan Meseta and Sahara platform. In this review we provide a brief history of geological investigations and an overview of Cretaceous sedimentary archives, as well as a selection of research highlights and outstanding questions concerning the Cretaceous system in Morocco. The Cretaceous sedimentary archives from Morocco and adjacent coastal basins still retain untapped potential to further contribute to our understanding of global eustatic sea-level changes and the response of the oceans and marine biota in upwelling driven oxygen minimum zones under greenhouse climate conditions.
... The Kem Kem Group's two formations, the Ifezouane Formation and overlying Aoufous Formation, differ in their lithology. The Ifezouane Formation is characterised by reddish fluvial detritic sandstones, with high angled, metrethick cross-bedding representing channel fills, with intercalations of pedogenic overbank mudstones (Belvedere et al. 2013), pinkish sand, and thin conglomerates with quartz pebbles . The overlying Aoufous Formation is characterised by variegated mudstones and marls, with calcitic palaeosols, thin intercalations of evaporites, detrital sandstone and micro-conglomerates, and carbonate cemented layers Belvedere et al. 2013). ...
... The Ifezouane Formation is characterised by reddish fluvial detritic sandstones, with high angled, metrethick cross-bedding representing channel fills, with intercalations of pedogenic overbank mudstones (Belvedere et al. 2013), pinkish sand, and thin conglomerates with quartz pebbles . The overlying Aoufous Formation is characterised by variegated mudstones and marls, with calcitic palaeosols, thin intercalations of evaporites, detrital sandstone and micro-conglomerates, and carbonate cemented layers Belvedere et al. 2013). The vast majority of vertebrate material comes from intraformational, mud-flake conglomerates in the upper Ifezouane Formation ). ...
The pterosaur assemblage of the mid-Cretaceous Kem Kem Group of Morocco is reviewed. This analysis examines their
taxonomy, palaeoecology and palaeobiology with comments on taphonomy. New material permits the rediagnosis of the
azhdarchoids Alanqa saharica and Afrotapejara zouhrii. Several specimens are reported that do not ft within the paradigms of previously named taxa. They represent three distinct jaw morphotypes, but are not assigned to new taxa here. The
assemblage is highly diverse, including four tooth-bearing taxa assigned to Ornithocheiridae and fve named taxa and three
additional morphotypes assigned to Azhdarchoidea. The Kem Kem Group assemblage is the most diverse for any pterosaurbearing fuvial deposit and one of the most diverse of any pterosaur assemblage. The assemblage is heavily biased in terms of preservation with an as yet unexplained high abundance of jaw fragments. We highlight the importance of fragmentary
material in pterosaur studies.
... The Kem Kem beds that have yielded the best specimens of Spinosaurus are indeed unusual in their prevalence of aquatic species and numerous theropod dinosaurs but lack of herbivores, and this has been commented on repeatedly in the literature (e.g., McGowan and Dyke, 2009;Belvedere et al., 2013;Läng et al., 2013). However, there is no particular reason to think that Spinosaurus was exploiting only aquatic prey (and of that, only fish) over other available prey, or would not scavenge at least on occasion. ...
... Specimens of Spinosaurus have been recovered from a wide variety of localities and environ-ments and this perhaps argues against an animal that is in some way unique in its association with the Kem Kem. Indeed, the recently suggested synonymisation of numerous genera and species within Spinosaurus aegyptiacus by Ibrahim and colleagues (Ibrahim et al., 2020b;Smyth et al., 2020) would greatly extend the temporal and geographic range of this species meaning that the oddity of the Kem Kem (which itself may be less strange that often depicted e.g., Belvedere et al., 2013) would not potentially remain an outlier and would in itself not explain the unusual morphology of Spinosaurus. ...
The giant theropod Spinosaurus was an unusual animal and highly derived in many ways, and interpretations of its ecology remain controversial. Recent papers have added considerable knowledge of the anatomy of the genus with the discovery of a new and much more complete specimen, but this has also brought new and dramatic interpretations of its ecology as a highly specialised semi-aquatic animal that actively pursued aquatic prey. Here we assess the arguments about the functional morphology of this animal and the available data on its ecology and possible habits in the light of these new finds. We conclude that based on the available data, the degree of adaptations for aquatic life are questionable, other interpretations for the tail fin and other features are supported (e.g., socio-sexual signalling), and the pursuit predation hypothesis for Spinosaurus as a “highly specialized aquatic predator” is not supported. In contrast, a ‘wading’ model for an animal that predominantly fished from shorelines or within shallow waters is not contradicted by any line of evidence and is well supported. Spinosaurus almost certainly fed primarily from the water and may have swum, but there is no evidence that it was a specialised aquatic pursuit predator.
... Consequently, it should be expected to find non-pterodactyloid trackways alongside pterodactyloid ones in diverse ichnosites, at least during the Late Jurassic. In fact, while pterodactyloid tracks and trackways appear to be quite common in numerous sites worldwide (Lockley et al., 2008;Barrett et al., 2008;Belvedere et al., 2013), non-pterodactyloid traces remain unknown, which does not reflect the stratigraphic range and the known geographical distribution of the pterosaurian fossil record. Indeed, no tracks or trackways clearly referable to non-pterodactyloid pterosaurs were known, except cautious reports in Mazin et al. (1997Mazin et al. ( , 2001aMazin et al. ( , 2001bMazin et al. ( , 2003 from the Late Jurassic of Crayssac (France), as well as Mickelson et al. (2004) from the Late Jurassic of Utah (USA). ...
... Among the vertebrate ichnites, the pterodactyloid tracks are so far the most abundant, with thousands of isolated manus-and pes-prints, as well as more than twenty trackways referred to the ichnogenus Pteraichnus Stokes, 1957. This assemblage of trackways is undoubtedly one of the finest pterosaurian ichnological record in the world (Barrett et al., 2008;Lockley et al., 2008;Belvedere et al., 2013). ...
New discoveries on the ichnological site known as “the Pterosaur Beach of Crayssac” (lower Tithonian, Upper Jurassic; south-western France) answer the question of terrestrial capabilities of non-pterodactyloid pterosaurs. If the terrestrial type of locomotion of pterodactyloid pterosaurs has been solved from ichnological evidence for more than twenty years, no tracks and trackways referable to non-pterodactyloid pterosaurs have ever been described. Thus, the debate on terrestrial capabilities of these non-pterodactyloids was based on morpho-functional studies, with the main conclusion that those pterosaurs were arboreal dwellers and bad walkers. Six trackways referable to three non-pterodactyloid new ichnotaxa, maybe closely related to Rhamphorhynchidae, are described in this work. Their study leads to the conclusion that grounded non-pterodatyloids, at least during the Late Jurassic, were quadrupedal with digitigrade manus and plantigrade to digitigrade pes. They were clearly good walkers, even if hindlimbs are supposed to be hampered by the uropatagium, what could have constrained the terrestrial agility of these animals. Thus, from ichnological evidence and contrary to the current hypotheses, non-pterodactyloid pterosaurs seem to have been good walkers even though their trackways are very rare or unidentified to date. This rarity could be due to behaviour rather than to functional capacities, many non-pterodactyloids being considered both littoral fishers and arboreal or cliff dwellers. However, the concept of non-pterodactyloid “good climbers and bad walkers” has to be modified to “good climbers and rare walkers”, unless many non-pterodactyloid ichnites have yet to be discovered.
... In this regard, a combination of both sources of data can represent the census for a paleoecological reconstruction of the unit, thus indicating whether the ichnological record is consistent or not with the skeletal record (Lockley, 1991). Some examples have come to light where the skeletal and the ichnological record represent a similar faunal assemblage (e.g., Lockley et al., 1986) while in other cases there are considerable differences between the two records (e.g., Belvedere et al., 2013). Thulborn (1990) warns of possible preservational biases produced in this kind of census (e.g. ...
... These discrepancies between the ichnological and the skeletal record might be due to an TABLE 2. Sites with osteological remains reported from the Mirambel Formation. ecological signal or a preservational bias (Belvedere et al., 2013). ...
Up to now, the ichnological vertebrate record from the Barremian Mirambel Formation (NE Spain) has
remained completely unknown despite the fact that osteological findings have been reported in recent years. Here we provide an overview of 11 new dinosaur tracksites found during a fieldwork campaign in the year 2011. The majority of these tracksites (seven) preserve small- to medium-sized tridactyl tracks here assigned to indeterminate theropods. Only one footprint presents enough characters to classify it as Megalosauripus isp. Ornithopod tracks identified as Caririchnium isp. and Iguanodontipodidae indet. and sauropod tracks are recorded at two tracksites. The footprints are preserved in a variety of paleoenvironmental conditions and thus display different kinds of preservation (true tracks, shallow undertracks, natural casts and undertrack casts). The ichnological record from the Mirambel Formation seems to be theropod dominated. This is a clear discrepancy with the osteological record identified in this formation, which shows a predominance of ornithopod dinosaurs.
... The Kem Kem Group is generally considered to be Cenomanian in age but may date back to the Aptian in places (Cavin et al., 2010;Ibrahim et al., 2020a). Two formations are present: the vertebrate fossil-rich lower Ifezouane Formation, representing a braided fluvial system; and the upper Aoufous Formation, rich in ichnofossils, representing a coastal lagoonal system (Belvedere et al., 2013;Cavin et al., 2010;Ibrahim et al., 2020a;Mannion & Barrett, 2013;Fig. 1B). ...
The Kem Kem Group of Southeastern Morocco, North Africa, is well known for theropod remains, especially
isolated teeth. Here, a collection of isolated theropod teeth is assessed for diversity using a combination of linear discriminant,
phylogenetic, and machine learning analyses for the first time. The results confirm earlier studies on Kem Kem theropod
diversity, with teeth referred to Abelisauridae, Spinosaurinae, and Carcharodontosauridae. A single tooth is ascribed to a
non-abelisauroid ceratosaur or a megaraptoran and may represent the enigmatic averostran Deltadromeus. Spinosaurine
teeth are clearly differentiated by all three methodologies, whereas abelisaurid and carcharodontosaurid teeth could only
be distinguished by the machine learning and phylogenetic analyses. This study shows that a combination of independent
methods is most effective at providing strong evidence on theropod dental diversity in a particular assemblage, and that
cladistic and machine learning analyses are the most reliable approaches to identify isolated dinosaur teeth. The
methodology used here is likely to yield results in other dinosaur assemblages where isolated teeth are more abundant
than body fossils.
... Sauropod tracks are much less common in the Cretaceous of Algeria than theropod tracks (Bassoulet 1971, Bessedik et al. 2008, Draoui et al. 2022. Belvedere et al. (2013) reviewed the vertebrate tracks in the Kem Kem Beds (Cenomanian/Turonian) of southern Morocco and did not recognize any sauropod tracks. They hypothesized various explanations including: (1) failure to recognize sauropod tracks; (2) sporadic sauropod activity in the area and sauropod bones were transported in; and (3) possibly paucity of vegetation. ...
For the first time, a complete study of sauropod trackways has been carried out in the lower part of the Djoua series in the In Amenas region (southeastern Algeria). Several locomotion tracks have been discovered in this area. In the present work, we report the discovery of sauropod tracks exposed on a bedding surface of the Lower Formation attributed to the early Cenomanian that we assign to cf. Brontopodus isp. The tracks are located in two different areas approximately 2 km apart and are oriented N00° and N330° at the first and second sites, respectively. These tracks were formed when dinosaurs crossed an emergent or lightly submerged river plain. At the second site, trackways show several tracks generated by sauropods. They are sub-parallel in orientation and provide the first ichnological evidence of gregarious behavior in Cretaceous sauropods in Africa. ARTICLE HISTORY
... The Kem Kem Group outcrops for~250 km, roughly paralleling the Moroccan-Algerian border (Ibrahim et al., 2020). It is a continental deposit formed by~200 m of fluvial-to-deltaic deposits, mainly fine sandstones lower down, and red, variegated mudstones above (Cavin et al., 2010;Belvedere et al., 2013;Klein et al., 2017). The succession is split into the lower Ifezouane Formation and upper Aoufous Formation (Cavin et al., 2010;Ibrahim et al., 2010Ibrahim et al., , 2020. ...
Plesiosaurs were a long-lived and widespread group of marine reptiles, with a worldwide distribution and a temporal range from the Late Triassic to the Late Cretaceous. Most occur in marine deposits, but some occur in low-salinity, brackish to freshwater environments. We report plesiosaurs from the freshwater fluvial deposits of the mid-Cretaceous (?Albian-Cenomanian) Kem Kem Group of Morocco. Remains include numerous shed teeth, vertebrae, and a humerus. The humerus represents a young juvenile; vertebrae likely belong to sub-adults. Teeth show heavy wear, similar to teeth of co-occurring spinosaurids. While coeval plesiosaurs from the Bahariya Formation of Egypt are members of Polycotylidae, the Kem Kem fossils show features of Leptocleididae, small-bodied plesiosaurs that were widely distributed in nearshore and non-marine settings in the Early Cretaceous. These fossils are the first freshwater plesiosaurs from Morocco, and are among the youngest representatives of Leptocleididae. The Kem Kem leptocleidids could have been infrequent visitors from the sea, freshwater-tolerant, or even freshwater-adapted, as in modern river dolphins. The abundance of shed teeth in the Kem Kem Group supports the hypothesis that they had some degree of freshwater tolerance. Furthermore, leptocleidids occur almost exclusively in shallow nearshore, brackish, or freshwater environments, suggesting adaptation to shallow, low-salinity environments. Other plesiosaur groups and other Mesozoic marine reptiles, including teleosaurids and mosasaurids, also occur in freshwater settings, suggesting plesiosaurs and other marine reptiles frequently exploited non-marine environments.
... Perhaps the most spectacular "unbalanced" dinosaur fauna comes from the Late Cretaceous Kem Kem beds of North Africa (Belvedere et al., 2013;Benyoucef et al., 2015;Ibrahim et al., 2020;Läng et al., 2013;McGowan & Dyke, 2009). Large theropods here are more abundant than big herbivores, in both the skeletal and footprint samples. ...
Counts of the number of skeletal specimens of “adult” megaherbivores and large theropods from the Morrison and Dinosaur Park formations—if not biased by taphonomic artifacts—suggest that the big meat‐eaters were more abundant, relative to the number of big plant‐eaters, than one would expect on the basis of the relative abundance of large carnivores and herbivores in modern mammalian faunas. Models of megaherbivore population density (number of individuals per square kilometer) that attempt to take into account ecosystem productivity, the size structure of megaherbivore populations, and individual megaherbivore energy requirements, when combined with values of the large theropod/megaherbivore abundance ratio, suggest that large theropods may have been more abundant on the landscape than estimates extrapolated from the population density versus body mass relationship of mammalian carnivores. Models of the meat production of megaherbivore populations and the meat requirements of “adult” large theropods suggest that herbivore productivity would have been insufficient to support the associated number of individuals of “adult” large theropods, unless the herbivore production/biomass ratio was substantially higher, and/or the large theropod meat requirement markedly lower, than expectations based on modern mammals. Alternatively, or in addition to one or both of these other factors, large theropods likely included dinosaurs other than megaherbivores as significant components of their diet.
... Tale procedura di rilievo è estremamente vantaggiosa perché le immagini fotografiche, se ben riprese, contengono un'enorme quantità d'informazioni utili sia in termini geometrici sia in termini di dettagli; perché fotografare un oggetto è molto meno rischioso e molto più comodo e veloce che misurarlo direttamente; perché immagini d'archivio possono rivelarsi utili per la ricostruzione visuale e geometrica di ogni oggetto, anche di quelli non più esistenti; perché per la realizzazione di modelli 3D a partire da fotogrammetrie sono sufficienti anche attrezzature hardware e software di poco costo e, quindi, alla portata di tutti (Petti et al., 2008;Belvedere et al., 2013;Mallison & Wings, 2014). ...
... These faunas have been the subject of several studies (e.g. Dutheil, 1999;Cavin & Forey, 2004;Rage & Dutheil, 2008;Belvedere et al., 2013;Mannion & Barrett, 2013;Ibrahim et al., 2020). ...
We present the first known cranial remains of the fossil batoid †Onchopristis numidus. Based on two exceptionally well-preserved specimens collected from the ‘Kem Kem Beds’ (Albian–Cenomanian), south-east Morocco, an almost complete description of the rostral and cranial portions of the genus †Onchopristis is provided, together with new observations regarding the development and arrangement of the rostral denticle series for this genus. The comparison between the rostrum length of the specimens of †O. numidus with those of extant pristids, reveals a relatively large batoid species with an estimated total length between two and four metres. Overall, the cranial morphology of †Onchopristis resembles that of other sclerorhynchoids. Its robust hypertrophied rostrum, with the characteristic wood-like mineralization covering the inner layer of tessellate cartilage at the centre of the rostrum, in addition to the thick lateral layers of densely porous cartilage on the sides of the rostral cartilages, resembles that observed in †Ischyrhiza and †Schizorhiza, and differentiates †Onchopristis from other sclerorhynchoids (e.g. †Libanopristis, †Micropristis and †Sclerorhynchus). Aiming to determinate the phylogenetic relations of †Onchopristis within sclerorhynchoids, a cladistic analysis was carried out based on the remains described here; its results suggest a new taxonomic arrangement within the sclerorhynchoids.
... A good example is presented by Belvedere et al. (2017) in the description of the Aoufous Formation (Kem Kem beds, Late Cretaceous, Morocco) ichnofauna. There is a large number of theropods, turtles, crocodiles and possible pterosaurs tracks preserved in a siliciclastic succession interpreted as a coastal mudflat, close to the shoreline, with sabkhas and marginal ponds (Belvedere et al., 2013). ...
The origin and preservation of a track are related to many distinct environmental factors, concerning especially the substrate cohesiveness, plasticity, grain size, texture and water content. Then, the environment, through the sedimentation processes, plays a role that enhances the origin and quality of the tracks and their preservation.
Three distinct contexts - tidal flats, aeolian, fluvial-lacustrine paleoenvironments, that encompass the majority of
fossil footprints occurrences are analyzed. Footprints as biosedimentary structures, due to their close relationships with physical and chemical processes that control their formation, represent an important clue to paleoenvironmental interpretation. The present study is mainly based on the direct examination of ichnosites that allow us to evaluate the aspects of Mesozoic tracks from different regions of the paleocontinent Gondwana, currently correspondent to Argentina, Australia, Bolivia, Brazil, Congo, Iran, India, Madagascar and Morocco as sedimentary structures and their use in paleoenvironmental interpretations.
... Lovelace and Lovelace 2012;) as well as other tracks attributable to turtles (e.g. Avanzini et al. 2005;Belvedere et al. 2013). ...
A proximal distributary channel deposit in the Turonian Ferron Sandstone of central Utah contains a trace-fossil assemblage comprised of turtle tracks (Chelonipus), vertebrate swim trace fossils (Characichnos), repichnial traces of mollusks (Archaeonassa), Rhizocorallium, and Teredolites. Mayflies are abundant producers of horizontal, spreiten, U-shaped burrows in modern freshwater channels, but rarely ascribed as producers of Rhizocorallium in the rock record. The in situ trace fossils studied are interpreted as a freshwater assemblage. Teredolites occurs within allochthonous debris reworked from stratigraphically lower brackish-water deposits. This Ferron Sandstone assemblage provides an example of the juxtaposition of freshwater ichnocoenose with exhumed brackish-water clasts. Caution must be exercised in environmental interpretation when taking into consideration both adjacent floodplain and the sedimentary structures surrounding the clasts. Complex trace-fossil relationships such as these may play a role in the scarcity of Rhizocorallium reports from freshwater deposits in the rock record, and may be more common than presently recognized. This study illustrates the importance of addressing the detailed ichnology above and below the base of channel deposits in progradational environments (e.g., delta plain), where reworked, brackish-water deposits (and bored clasts) may be entrained within overlying lower salinity facies.
... Each model has been scaled on the basis of a metallic scale-bar. Finally, in respect of the latest scientific protocols (Falkingham et al., 2018) and according to the procedure of Belvedere et al. (2013) some detailed contour maps and high-resolution depth maps have been created through Kitware Paraview. Each parameter of the figured models is shown in Table 1 according to the method of Lockley et al. (2015). ...
This report aims to give notice of and provide a more detailed dataset and detailed remarks on what can be considered a one-of-a-kind hominin fossil walking pattern: Trackway B of the Foresta ichnological site (Tora e Piccilli, Caserta, Central Italy). Although the site is known since 2003, only recently has the study been performed by means of the newest photogrammetric and experimental techniques of collection, analysis and interpretation of ichnological data. The results obtained enable us to depict an astonishing movie printed in rock, describing some body features and common moments of the everyday movements of a hominin who lived about 350 ka. In particular, some up-to-now absolutely unique fossil prints of body parts of a Pleistocene hominin (calf, ankle, and gluteus), which have simply been mentioned in the ichnological fossil record, are here quantitatively described for the first time. The data coming from this research will provide scientists with new valuable elements thus far undetected anywhere else in the world.
... 1C). Tracks pertain to medium-and large-sized theropods and, much less frequently, large-sized ornithopods and sauropods (Sereno et al. 1996, Belvedere et al. 2013, Ibrahim et al. 2014a. Tracks occur as impressions and natural casts, the latter the more common and usually occurring in the rare siltstone and sandstone beds. ...
The geological and paleoenvironmental setting and the vertebrate taxonomy of the fossiliferous, Cenomanian-age deltaic sediments in eastern Morocco, generally referred to as the “Kem Kem beds”, are reviewed. These strata are recognized here as the Kem Kem Group, which is composed of the lower Gara Sbaa and upper Douira formations. Both formations have yielded a similar fossil vertebrate assemblage of predominantly isolated elements pertaining to cartilaginous and bony fishes, turtles, crocodyliforms, pterosaurs, and dinosaurs, as well as invertebrate, plant, and trace fossils. These fossils, now in collections around the world, are reviewed and tabulated. The Kem Kem vertebrate fauna is biased toward largebodied carnivores including at least four large-bodied non-avian theropods (an abelisaurid, Spinosaurus, Carcharodontosaurus, and Deltadromeus), several large-bodied pterosaurs, and several large crocodyliforms.
No comparable modern terrestrial ecosystem exists with similar bias toward large-bodied carnivores. The Kem Kem vertebrate assemblage, currently the best documented association just prior to the onset of the Cenomanian-Turonian marine transgression, captures the taxonomic diversity of a widespread northern African fauna better than any other contemporary assemblage from elsewhere in Africa.
Keywords
Africa, Cretaceous, dinosaur, Gara Sbaa Formation, Douira Formation, paleoenvironment, vertebrate
... Since their first discovery in Morocco (Plateau et al., 1937), dinosaur tracks, including those of theropods, ornithischians and sauropodomorphs, have attracted the interest of scientists (e.g. Lapparent, 1945;Dutuit and Ouazzou, 1980;Jenny et al., 1981;Monbaron et al., 1985;Ishigaki, 1985Ishigaki, , 19861988Pérez-Lorente et al., 2006;Hadri et al., 2007;Gierliński et al., 2009Gierliński et al., , 2017bIshigaki and Matsumoto, 2009a;Belvedere and Mietto, 2010; Díaz-Martínez et al., 2010;Marty et al., 2010;Nouri et al., 2011;Hadri and Pérez-Lorente, 2012;Belvedere et al., 2013;Ibrahim et al., 2014;Masrour et al., 2013Masrour et al., , 2015Enniouar et al., 2014;Boudchiche et al., 2017;Lallensack et al., 2019). ...
A new dinosaur tracksite is reported from continental red beds of the Jurassic (Late Bathonian-?Callovian) Isli Formation along the northern flank of the Aït Ali ou Ikkou Syncline of the Imilchil area, Central High Atlas, Morocco. The succession was deposited in a fluvio-lacustrine environment, and contains at least fourteen track-bearing levels. The diverse dinosaur-dominated ichnofauna includes the footprints of crocodylomorphs, pterosaurs, theropods, sauropods and ornithischians, together with numerous invertebrate traces. Here, we focus on a long sauropod trackway, which comprises nine consecutive manus-pes sets preserved as concave epireliefs. The low heteropody and asymmetry of manus prints with a large digit I (pollex) impression oriented medially, and a large triangular claw trace, which is posteriorly oriented, are characteristic of the ichnogenus Polyonyx. Different from typical Polyonyx is the narrow gauge pattern compared to the wide gauge observed in the type trackway from Portugal. Additional material from Morocco, similar to Polyonyx, comprises the trackway of a very small (?juvenile) individual found close to the main trackway, as well as a short trackway from a different locality in the Isli Formation. Our data from the Moroccan High Atlas indicates the presence of basal eusauropods in the Middle Jurassic-Early Cretaceous interval in the northwestern part of Gondwana for the first time, and suggests they inhabited a lacustrine paleoenvironment.
... In some cases, e.g., in Portugal (Mateus et al., 2006;Mateus and Milàn, 2010), Spain Rauhut et al., 2018), Germany (Lallensack et al., 2015) skeletal sites and tracksites are in close geological proximity allowing a more precise trackmaker identification. Dinosaur footprints have proved to be a valuable tool for palaeoecological analyses and reconstructions of faunal associations (e.g., Lockley, 1986;Belvedere et al., 2013). Despite the uncertainty of trackmaker identification, ichnotaxonomy would be a good tool to pursue these purposes, at least to determine which groups of animals where present in a certain environment. ...
Late Jurassic theropod tracks are very common both in North Africa and Europe. Two recently described ichnotaxa Megalosauripus transjuranicus and Jurabrontes curtedulensis from the Kimmeridgian of Switzerland show the coexistence of two apex predators in the same palaeoenvironment. Similar tracks can be found in tracksites from the Iberian Peninsula and from Morocco. Here, we further explore the similarities among the Swiss ichnotaxa and the other tracks from Germany (Kimmeridgian), Spain (Tithonian-Berriasian), Portugal (Oxfordian-Tithonian) and Morocco (Kimmeridgian) through novel three-dimensional data comparisons. Specimens were grouped in two morphotypes: 1) large and gracile (30 < Foot Length<50cm) and 2) giant and robust (FL > 50cm). The analyses show a great morphological overlap among these two morphotypes and the Swiss ichnotaxa (Megalosauripus transjuranicus and Jurabrontes curtedulensis, respectively), even despite the differences in sedimentary environment and age. This suggests a widespread occurrence of similar ichnotaxa along the western margin of Tethys during the Late Jurassic. The new data support the hypothesis of a Gondwana-Laurasia faunal exchange during the Middle or early Late Jurassic, and the presence of migratory routes around the Tethys.
... Since their first discovery in Morocco (Plateau et al., 1937), dinosaur tracks, including those of theropods, ornithischians and sauropodomorphs, have attracted the interest of scientists (e.g. Lapparent, 1945;Dutuit and Ouazzou, 1980;Jenny et al., 1981;Monbaron et al., 1985;Ishigaki, 1985Ishigaki, , 19861988Pérez-Lorente et al., 2006;Hadri et al., 2007;Gierliński et al., 2009Gierliński et al., , 2017bIshigaki and Matsumoto, 2009a;Belvedere and Mietto, 2010; Díaz-Martínez et al., 2010;Marty et al., 2010;Nouri et al., 2011;Hadri and Pérez-Lorente, 2012;Belvedere et al., 2013;Ibrahim et al., 2014;Masrour et al., 2013Masrour et al., , 2015Enniouar et al., 2014;Boudchiche et al., 2017;Lallensack et al., 2019). ...
Horseshoe crabs (Chelicerata, Xiphosura) from Mesozoic deposits are extremely rare in the fossil record of the African continent. Here we report new evidence of the occurrence of xiphosurans in North Africa. These are horseshoe crab traces, including the ichnogenera Kouphichnium (repichnia) and Selenichnites (fodinichnia and/or domichnia), which have been discovered in Middle Jurassic strata of the Imilchil area (Central High Atlas, Morocco). They are preserved on upper and lower bed surfaces of sandy limestones and marls in the upper parts of the Tislit and Imilchil formations (late Bajocian-early Bathonian).
The Selenichnites traces, measuring up to 30 cm in width, co-occur with theropod and birdlike tracks. The reported Kouphichnium is the first record from Mesozoic deposits of Morocco, while Selenichnites is the second occurrence in Jurassic strata of Africa. These ichnogenera evidence different behaviour of horseshoe crabs, i.e. Selenichnites is the result of burrowing activity when searching for food in the sediment, and Kouphichnium is a regular locomotion trace left on the sediment surface, occasionally leaving an impression of the telson.
The combination of sedimentological data and ichnological analysis indicates a shallow-water subtidal depositional environment preceding the Bathonian regression of the Atlas domain.
Palaeobiogeographically, the discoveries indicate the presence of horseshoe crabs at the southern margin of the Tethys. Furthermore, they enhance our knowledge of their previously scarcely documented distribution in Gondwana during the Mesozoic.
... The area recording the elephant fossil footprints was georeferenced with a Garmin Etrex 10 detector (accuracy ±3 m) and photographed through Sony Nex6 camera with Sony 16-50 lens. A detailed 3D model was then created by processing the pictures with Agisoft Photoscan Pro and Kitware Paraview software following the procedures proposed by Mallison & Wings (2014) and Belvedere et al. (2013). ...
This research aims to give notice of and provide preliminary information on some elephant footprints recently identified at Foresta ichnological site (Tora-Piccilli, Caserta, central Italy). The elephant-footprint area was previously poorly investigated because partially affected by anthropic modifications during historical times. The footprints, ascribed to the Proboscipeda panfamilia ichospecies, were left by a young strait-tusked elephant, passing the top of the slope formed during the deposition of the Roccamonfina Brown Leucitic Tuff (BLT), close to the renowned prehistorical Devil's path. The Foresta elephant footprints are the unique conceivably belonging to Palaeoloxodon recorded in Italy and the first elephant tracks reported from the continental Italian territory. © 2010 AIQUA - Associazione Italiana per lo Studio del Quaternario e EMMEVI - Servizio Congressi SPA.
... A. elegans Ambroggi and Lapparent 1954 can be regarded as a valid pterodactyloid ichnotaxon from the Late Cretaceous, different from the other known Jurassic and Cretaceous pterodactyloid ichnotaxa. It was the first pterosaurian ichnotaxon ever described in Africa and the world, but remained ignored for a long time, which lead several authors to assume the Agadir tracks were not pterosaurian (Belvedere et al. 2013;Calvo and Lockley 2001;Calvo and Moratalla 1998), or to question its validity Lockley et al., 2008;S anchez-Hern andez et al. 2009). ...
Discovered in 1954 by Ambroggi and Lapparent, the Tagragra tracksite (Maastrichtian, Agadir, Morocco) yielded tracks of theropod dinosaurs, birds and enigmatic forms tentatively attributed to Lacertilia under the name Agadirichnus elegans. The original specimens are today considered to be lost, and the status of A. elegans has been questioned, even though these footprints can retrospectively be referred to pterodactyloid pterosaur tracks. In order to discuss the status of these foot prints, the historical site has been actively sought and recently rediscovered by one of us (MdD). New material confirms the presence of pterodactyloid footprints, with two morphotypes, in two different facies, respectively associated with bird and theropod tracks. Morphotype I is tentatively conferred to Pteraichnus Stokes 1957 Stokes, W. L. 1957. Pterodactyl tracks from the Morrison Formation. Journal of Palaeontology, 31:952–4.[Web of Science ®] [Google Scholar], which extends the stratigraphical distribution of this ichnotaxon to the Maastrichtian. Morphotype II corresponds to the original ichnospecies A. elegans Ambroggi and Lapparent 1954 Ambroggi, R., and A. F. de Lapparent 1954. Les empreintes de pas fossiles du Maestrichtien d'Agadir. Notes du Service Géologique du Maroc, 10:43–6. [Google Scholar], which is here reassessed as a valid ichnotaxon, with the designation of a neotype. Thus, A. elegans Ambroggi and Lapparent 1954 Ambroggi, R., and A. F. de Lapparent 1954. Les empreintes de pas fossiles du Maestrichtien d'Agadir. Notes du Service Géologique du Maroc, 10:43–6. [Google Scholar] is historically the first pterosaurian ichnotaxon ever described.
... A. elegans Ambroggi and Lapparent 1954 can be regarded as a valid pterodactyloid ichnotaxon from the Late Cretaceous, different from the other known Jurassic and Cretaceous pterodactyloid ichnotaxa. It was the first pterosaurian ichnotaxon ever described in Africa and the world, but remained ignored for a long time, which lead several authors to assume the Agadir tracks were not pterosaurian (Belvedere et al. 2013;Calvo and Lockley 2001;Calvo and Moratalla 1998), or to question its validity Lockley et al., 2008;S anchez-Hern andez et al. 2009). ...
ABSTRACT
Discovered in 1954 by Ambroggi and Lapparent, the Tagragra tracksite (Maastrichtian, Agadir, Morocco) yielded tracks of theropod dinosaurs, birds and enigmatic forms tentatively attributed to Lacertilia under the name Agadirichnus elegans. The original specimens are today considered to be lost, and the status of A. elegans has been questioned, even though these footprints can retrospectively be referred to pterodactyloid pterosaur tracks. In order to discuss the status of these foot prints, the historical site has been actively sought and recently rediscovered by one of us (MdD). New material confirms the presence of pterodactyloid footprints, with two morphotypes, in two different facies, respectively associated with bird and theropod tracks. Morphotype I is tentatively conferred to Pteraichnus Stokes 1957 Stokes, W. L. 1957. Pterodactyl tracks from the Morrison Formation. Journal of Palaeontology, 31:952–4.
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, which extends the stratigraphical distribution of this ichnotaxon to the Maastrichtian. Morphotype II corresponds to the original ichnospecies A. elegans Ambroggi and Lapparent 1954 Ambroggi, R., and A. F. de Lapparent 1954. Les empreintes de pas fossiles du Maestrichtien d'Agadir. Notes du Service Géologique du Maroc, 10:43–6.
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, which is here reassessed as a valid ichnotaxon, with the designation of a neotype. Thus, A. elegans Ambroggi and Lapparent 1954 Ambroggi, R., and A. F. de Lapparent 1954. Les empreintes de pas fossiles du Maestrichtien d'Agadir. Notes du Service Géologique du Maroc, 10:43–6.
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is historically the first pterosaurian ichnotaxon ever described.
... Since the first discovery in the 19th century (Tagart, 1846), "large-sized" ornithopod footprints have been reported from all continents except Antarctica (e.g., Sternberg, 1932;Leonardi, 1984;Dong et al., 2003;Romillio and Salisbury, 2011;Belvedere et al., 2013). They were traditionally interpreted as iguanodontid or hadrosaurid footprints (Thulborn, 1990). ...
Two ichnogenus of “large-sized” ornithopods are found from the Lower Cretaceous (Aptian) Kitadani Formation in central Japan. Caririchnium isp. is characterized by the longer pes print than its width with strong mesaxony. Amblydactylus isp. is characterized by the wider pes print than its length with weak mesaxony. In the northern hemisphere, Caririchnium-type footprints are known from the Berriasian–Cenomanian strata, whereas Amblydactylus-type footprints are known from the Barremian–Maastrichtian strata. It is consistent with the temporal and geographic distribution of non-hadrosauroid iguanodontians and basal hadrosauroids. It suggests that footprint length-width ratio and mesaxony are important factors to indicate trackmakers (basal iguanodontian or hadrosauroid). Two “large-sized” ornithopod ichnogenus from the Kitadani Formation is also consistent with two iguanodontians from same site. It indicates a high diversity of the Kitadani ichnofauna and its importance to elucidate the ecosystem of the Kitadani Formation.
... The pictures were processed with Agisoft Photoscan Pro and then scaled to obtain a photogrammetric 3D model of the surface, following the procedures of Mallison & Wings (2014). The mesh was then imported in Kitware Paraview to create high resolution depth maps, following the procedure of Belvedere et al. (2013). The parameters of each figured model are shown in Tab. 1, according to the method suggested by Lockley et al. (2015). ...
Fossil footprints of Middle Pleistocene hominids, locally known as “Ciampate del Diavolo”, were discovered in 2001 on the North-Eastern slope of the Roccamonfina volcano (central Italy, municipality of Tora and Piccilli, Caserta). The imprints occur on a surface of Brown Leucitic Tuff (BLT), dated to 349 ± 3 ka. After this discovery, e this tuffaceous formation was investigated in more detail on the North-Eastern, Eastern and South-Eastern side of the Roccamonfina volcano, with the aim of locating other fossil footprints. In 2003 another succession of nine hollows fully compatible with human fossil footprints for stratigraphic position, pattern and dimensional range was found on the same ridge but at a higher level. This second site, known locally as Cantarelle di Carangi, is positioned at a distance of about 1.8 km from the first fossiliferous area (municipality of Marzano Appio, middle-central Italy, Caserta province). Further surveys and archaeological excavations were carried out in the Cantarelle di Carangi site during 2008 to better understand, contextualize and confirm the potential ichnological value of the findings of 2003. Ichnological and stratigraphical analyses, combined with findings during the excavations and historical data, however, tell a different story. What was supposed to be a new ichnosite with Pleistocene fossil footprints, is very likely the result of anthropic digging, erosion made by animals, and weathering. In any case, the discovery shows how careful researchers have to be in interpreting possible fossil footprints, especially in conditions of poorly preserved traces.
... Other tracks assigned to Megalosauripus from the Late Jurassic [96] and Early Cretaceous of Morocco [111], Arizona and Utah [3,112], Poland [113], and Germany [13,18,114], and from the Middle Jurassic of Madagascar [115] and China [10] either display subequal phalangeal pads when preserved (Arizona, Utah, Germany, Morocco), lack a discrete phalangeal pad formula (Madagascar, England, Poland) or are too poorly preserved for a sound comparison (China). ...
A new ichnospecies of a large theropod dinosaur, Megalosauripus transjuranicus, is described from the Reuchenette Formation (Early–Late Kimmeridgian, Late Jurassic) of NW Switzerland. It is based on very well-preserved and morphologically-distinct tracks (impressions) and several trackways, including different preservational types from different tracksites and horizons. All trackways were excavated along federal Highway A16 near Courtedoux (Canton Jura) and systematically documented in the field including orthophotos and laserscans. The best-preserved tracks were recovered and additional tracks were casted. Megalosauripus transjuranicus is characterized by tridactyl tracks with clear claw and digital pad impressions, and notably an exceptionally large and round first phalangeal pad on the fourth digit (PIV1) that is connected to digit IV and forms the round heel area. Due to this combination of features, M. transjuranicus clearly is of theropod (and not ornitho-pod) origin. M. transjuranicus is compared to other Megalosauripus tracks and similar ichno-taxa and other unassigned tracks from the Early Jurassic to Early Cretaceous. It is clearly different from other ichnogenera assigned to large theropods such as Eubrontes–Grallator from the Late Triassic and Early Jurassic or Megalosauripus–Megalosauropus–Bueckebur-gichnus and Therangospodus tracks from the Late Jurassic and Early Cretaceous. A second tridactyl morphotype (called Morphotype II) is different from Megalosauripus trans-juranicus in being subsymmetric, longer than wide (sometimes almost as wide as long), with blunt toe impressions and no evidence for discrete phalangeal pad and claw marks. Some Morphotype II tracks are found in trackways that are assigned to M. transjuranicus, to M.? transjuranicus or M. cf. transjuranicus indicating that some Morphotype II tracks are intra-trackway preservational variants of a morphological continuum of Megalosauripus transjura-nicus. On the other hand, several up to 40 steps long trackways very consistently present Morphotype II features (notably blunt digits) and do not exhibit any of the features that are typical for Megalosauripus (notably phalangeal pads). Therefore, it is not very likely that these tracks are preservational variants of Megalosauripus transjuranicus or PLOS ONE | https://doi.
... 3.98.1.64) was used to generate high resolution depth maps, following the procedure of Belvedere et al. (2013). Photogrammetry has been preferred to field laser scanning because of the difficulties in accessing the Table 1, which is filled according to the method suggested by Lockley et al. (2015). ...
Some human footprints-like hollows in the tuffaceous ground of an ancient quarry and of a recent anthropic path, created by the inhabitants no more than 30 years ago near the small village of Carangi (Caserta province, Southern Italy), allowed scientists to study some of the most misleading alteration processes of the volcanic tuff formations. The closeness of this site to the “Ciampate del Diavolo” ichnosite, bearing some of the oldest human fossil footprints, and the common stratigraphical and sedimentological background permitted accurate comparisons between actual human fossil footprints and footprint-like depressions, which can be easily mistaken for actual fossil evidence. This comparative study has permitted the reporting of some useful features and advice on distinguishing actual fossil footprints from natural and/or anthropic pseudo-tracks.
... Recent reports for Morocco consist of Barremian-Aptian dinosaur footprints that include two quadrupedal trackways assigned to the ichnogenus Brontopodus (Masrour et al., 2013). Cretaceous theropod footprints were found on isolated slabs from the Kem Kem beds of Morocco (Belvedere et al., 2013). As for the rest of North Africa, around 300 footprints including tridactyl and circular footprints were reported from the Albian-Cenomanian of Niger (Ginsburg et al., 1966). ...
Dinosaur trackways have rarely been reported in Cretaceous strata across the African continent. To the exception of ichnological occurrences in Morocco, Tunisia, Niger and Cameroon, our knowledge on the composition of Cretaceous dinosaur faunas mostly relies on skeletal evidence. For the first time, we document several dinosaur trackways from the Cretaceous of the Mamfe Basin in western Cameroon. Small and medium-size tridactyl footprints as well as numerous large circular footprints are present on a single horizon showing mudcracks and ripple marks. The age of the locality is considered Cenomanian-Turonian and if confirmed, this ichnological assemblage could be younger than the dinosaur footprints reported from northern Cameroon, and coeval with or younger than skeletal remains reported from the Saharan region. These trackways were left in an adjacent subsiding basin along the southern shore of the Benue Trough during a time of high-sea stand when the Trans-Saharan Seaway was already disconnecting West Africa from the rest of the continent. We predict that other similar track sites may be occurring along the margin of the Benue Trough and may eventually permit to test hypotheses related to provincialism among African dinosaur faunas.
... Some examples have come to light elsewhere where the skeletal and the ichnological record represent a similar faunal assemblage (Lockley et al., 1986), while in other cases there are considerable differences between the two records (e.g. Belvedere et al., 2013). The ichnological record of the Mirambel Formation is theropod-dominated. ...
... In addition, the digit traces II and IV terminate in sharp claw impressions. These features exclude crocodyliforms because they present marked heteropodia, the manus is much smaller than the pes, and the pes is tetradactyl and outwardly rotated (e.g., Belvedere et al., 2013) and crocodiles usually do not paddle with their limbs while swimming, they use their tail for propulsion. However, Foster and Lockley (2006) and Vila et al. (2015) describe swim tracks (Hatcherichnus) of crocodiles from Upper Jurassic of North America and Upper Cretaceous of Europe, respectively. ...
Twenty-six enigmatic footprints composed of three elongated parallel depressions (two lateral ones and a shorter central one located forward with respect to the others) are described from Middle Triassic red beds of the Tabular Cover in southern Spain. The purpose of this work is the interpretation of these enigmatic tracks. The footprints range from 28 to 48 cm in length and 23 to 44 cm in width. Limestones on top of the footprint-bearing sandstone contain invertebrate traces such as Planolites, Thalassinoides, and Rhizocorallium, and bivalves (Pleuromya, Trigonodus, and Unionites), typical of marine and brackish environments. They indicate that these footprints developed in a coastal or nearly coastal environment. Whereas the enigmatic trace fossils correspond to tridactyl footprints, the digital marks are parallel and do not show any divarication as in typical walking footprints of archosauria. These traces were left when the trackmaker was swimming in a waterbody deep enough for floating only occasionally touching the ground with the digit tips. The tracks formed as a result of the backward sweeping of the tips when the digits came in contact with the bottom while the animal was buoyed by water. They differ from typical swim traces of other archosaurs (such as theropods) that show three longitudinal scratches because in studied tracks the axial digit (III) of the bipedal tridactyl archosaur sinks in the sediment and gives a short and deep scratch. The other digit scratches (from digits II and IV) would correspond to the upper surface of the toes. The studied tracks correspond to Characichnos ichnofacies (swimming tracks composed by parallel scratch marks) and they were made by a bipedal tridactil archosaur or some functionally tridactyl, chirothere tracemaker.
... A main difference with the Lourinhã Formation is, however, the low diversity of herbivorous dinosaurs and the scarcity of their remains in the Kem Kem beds [32], which seems to support an unbalanced food chain given such a diversity of carnivorous dinosaurs. Yet, with two sauropod taxa, at least one ornithopod taxon, and a large taxonomic diversity of fish, amphibians, crocodiles, turtles, and pterosaurs (see [5,32,37,39,[110][111][112][113][114][115][116] and references therein), prey was not rare in the Kem Kem ecosystems and seems to have been in a sufficient quantity to feed six predatory dinosaurs of various size. As for Spinosauridae, the presence of two taxa is not surprising given the opportunistic nature of their feeding strategies ( [89,117]; see below). ...
Six quadrate bones, of which two almost certainly come from the Kem Kem beds (Cenomanian, Upper Cretaceous) of south-eastern Morocco, are determined to be from juvenile and adult individuals of Spinosaurinae based on phylogenetic, geometric morphometric, and phylogenetic morphometric analyses. Their morphology indicates two morphotypes evidencing the presence of two spinosaurine taxa ascribed to Spinosaurus aegyptiacus and? Sigilmassasaurus brevicollis in the Cenomanian of North Africa, casting doubt on the accuracy of some recent skeletal reconstructions which may be based on elements from several distinct species. Morphofunctional analysis of the mandibular articulation of the quadrate has shown that the jaw mechanics was peculiar in Spinosauridae. In mature spinosaurids, the posterior parts of the two mandibular rami displaced laterally when the jaw was depressed due to a lateromedially oriented intercondylar sulcus of the quadrate. Such lateral movement of the mandibular ramus was possible due to a movable mandibular symphysis in spinosaurids, allowing the pharynx to be widened. Similar jaw mechanics also occur in some pterosaurs and living pelecanids which are both adapted to capture and swallow large prey items. Spinosauridae, which were engaged, at least partially, in a piscivorous lifestyle, were able to consume large fish and may have occasionally fed on other prey such as pterosaurs and juvenile dinosaurs.
Pterosaurs are powered flight vertebrates that first emerged in the Late Triassic and persisted until the end of the Cretaceous. Recent studies on the ecological niches and habitats of pterosaurs, based on discoveries of Cretaceous bone beds in China, Brazil, Morocco, and other locations, suggest a decline in pterosaur diversity during the mid-Cretaceous. Various hypotheses have been proposed to interpret this phenomenon. This study focuses on newly discovered pterosaur tracks at the Upper Cretaceous (Truonian) Hwasun Seoyuri tracksite in Korea. The analysis is based on pterosaur tracks from in situ and ex situ specimens that originate from a variety of stratigraphic levels. The evidence found suggests that the Hwasun Seoyuri tracksite was occupied by small pterosaurs over an extended period, with indications of recurrent activity across at least six levels. Furthermore, the footprint sizes observed across these multiple layers follow a non-skewed normal distribution. Considering the known logarithmic growth characteristics of pterosaurs, this result suggests a predominantly immature population at this site. The prevalence of mid-Cretaceous non-marine pterosaur tracksites on the Korean Peninsula may reflect broader ecological shifts for pterosaurs during this period. This evidence is considered alongside the habitat shifts suggested by other non-marine pterosaur fossil sites in China and Brazil, as well as with indications of the decline in Late Cretaceous non-marine basins.
The footprints left by the Palaeolithic hominins at the ca. 350 ka old Foresta "Devil's Trails" ichnosite (Tora-Piccilli, central Italy) are rather variable, even in a single trackway. The peculiar characteristics of the deposit and the acclivity of the soft, slipping slope the hominins were walking on, which forced trackmakers to change pace and walking direction, likely account for this variability. As a result, determining whether the footprints were left by distinct trackmakers, as it would be logical to hypothesize based on the main settings of the trackways, or by a single individual who descended the slope more than once in a short time span, is difficult. To try to answer the question, we have analysed the Foresta/"Devil's Trails" footprint sample by means of various statistical methods with the double aim of quantitatively defining the minimum number of hominin trackmakers who walked on the ignimbrite deposit's slope and scrutinizing to what extent the acclivity of the substrate and the position of each footprint on the slope may affect their dimensions and proportions. The obtained results suggest that four trackmakers (A, B, C, and E) walked on the ignimbrite slope of the deposit. Individuals A, B, and C most likely had similar foot sizes, whereas individual E had larger one. Conversely, more solid data are needed to support the hypothesis that a fifth individual, smaller in size, left the footprints of short sequence D. Furthermore, the results underline how much the coarse, soft, and slippery substrate, along with the slope acclivity, influenced the direction of walking and its changing, the velocity, the length of the stride, the pace stability, and the way in which the foot rests on the substrate slope and, in turn, the shape and size of the footprints. The synergetic action of these factors influenced the footprint proportions, which differ in dimensions even within the same trackway.
This work describes the new vertebrate tracksite named El Barrancazo, located in Cortes de Pallás (Eastern Iberia). This new site turns this area into the one that concentrates the largest number of vertebrate footprints from the Upper Triassic of Iberia. The footprints are presented as natural casts distributed in isolated blocks belonging to the Manuel Formation (K2, Keuper facies), deposited during the Carnian Pluvial Episode (Late Triassic). The blocks come from two beds of the same outcrop. The tracks are made up of sub-parallel digital impressions, displaying occasional curved traces, often without forming a complete outline, and showing a set of features such as lateral and posterior overhangs, longitudinal striations or digital reflectures, among others. These features, taken together with the sedimentological analysis carried out, suggest that animals moving in totally or partially buoyant conditions in a fluvial environment produced most of these tracks. The size and morphology of these paleoichnological impressions suggest that they were probably produced, in some cases, by turtles, adding to the understanding of the paleoenvironmental and paleofaunistic conditions of the Late Triassic of Iberia during the Carnian, a key period in the evolution of different groups of continental vertebrates. In this paper we show the comparative study of the three sites described to date in Cortes de Pallás and an analysis of the interplay of trackmaker organism, animal behavior and substrate in the resulting morphology of tracks made under subaqueous conditions.
This work describes the new vertebrate tracksite named El Barrancazo, located in Cortes de Pallás (Eastern Iberia). The footprints are presented as natural casts distributed in isolated blocks belonging to the Manuel Formation (K2, Keuper facies), deposited during the Carnian Humid Episode (Late Triassic).
The blocks come from two beds of the same outcrop. The tracks are made up of sub-parallel digital impressions, displaying occasional curved traces, often without forming a complete outline, and showing a set of features such as lateral and posterior overhangs, longitudinal striations or digital reflectures, among others. These features, taken together with the sedimentological analysis carried out, suggest that animals moving in totally or partially buoyant conditions in a fluvial environment produced most of these tracks. The size and morphology of these paleoichnological impressions suggest that they were probably produced, in some cases, by turtles, adding to the understanding of the paleoenvironmental and paleofaunistic conditions of the Late Triassic of Iberia during the Carnian, a key period in the evolution of different groups of continental vertebrates.
The area of Cortes de Pallás concentrates the largest number of vertebrate footprints from the Upper Triassic of Iberia, which allows, by means of the comparative study of the three sites described to date, the analysis of the interaction of the trackmaker organism, animal behavior and substrate in the resulting morphology of the tracks made under subaqueous conditions.
The Upper Cretaceous 'upper' Winton Formation of Queensland, Australia is world famous for hosting Dinosaur Stampede National Monument at Lark Quarry Conservation Park, a somewhat controversial tracksite that preserves thousands of tridactyl dinosaur tracks attributed to ornithopods and theropods. Herein, we describe the Snake Creek Tracksite, a new vertebrate ichnoassemblage from the 'upper' Winton Formation, originally situated on Karoola Station but now relocated to the Australian Age of Dinosaurs Museum of Natural History. This site preserves the first sauropod tracks reported from eastern Australia, a small number of theropod and ornithopod tracks, the first fossilised crocodyliform and ?turtle tracks reported from Australia, and possible lungfish and actinopterygian feeding traces. The sauropod trackways are wide-gauge, with manus tracks bearing an ungual impression on digit I, and anteriorly tapered pes tracks with straight or concave forward posterior margins. These tracks support the hypothesis that at least one sauropod taxon from the 'upper' Winton Formation retained a pollex claw (previously hypothesised for Diamantinasaurus matildae based on body fossils). Many of the crocodyliform trackways indicate underwater walking. The Snake Creek Tracksite reconciles the sauropod-, crocodyliform-, turtle-, and lungfish-dominated body fossil record of the 'upper' Winton Formation with its heretofore ornithopod-and theropod-dominated ichnofossil record.
The morphological study of geological hand samples has a wide variety of applications in the geosciences, which is conventionally accomplished by measuring the distance between features of interest on the sample's surface. Close-range three-dimensional (3D) laser scanners provide an opportunity to study the form and shape of geological samples in a digital environment and has been increasingly utilized in fields such as paleontology, rock mechanics and sedimentology, with some uptake in planetary sciences and structural geology. For paleontological studies, primary applications are in quantitative analysis of fossil morphology and integration into 3D animated models for understanding species movements. In the field of rock mechanics, typical uses of 3D digital geological hand sample models include quantifying joint roughness coefficient (JRC), determining the surface roughness of rock samples, and assessing morphological changes over time due to processes such as weathering. In the field of sedimentology, such models are incorporated to characterize the shape of sediment particles and to calculate key parameters such as bulk density. This paper aims to provide a comprehensive review of established literature that includes substantial use of digital geological hand samples acquired from 3D close-range (<1 m target distance) triangulation laser scanners in an effort to identify opportunities for future progress (such as global data sharing) as well as challenges specific to the nature of geological samples (e.g., translucency) and geoscientific workflows (on and off-site).
Abelisaurid theropods first appear in the fossil record in the early Jurassic and survived at least until the end of the Mesozoic. They were known to have dominated South America, India and Madagascar but were not so abundant in North America or Asia. Much less is known about their presence in Africa, although there has been several recent discoveries of abelisaurid material in Morocco. Here we add a partially preserved ilium to a growing body of evidence that suggests abelisaurs might also have dominated Africa.
The Cretaceous Kem Kem beds of Morocco and equivalent beds in Algeria have produced a rich fossil assemblage, yielding, amongst others, isolated sauropod teeth, which can be used in species diversity studies. These Albian-Cenomanian (∼113–93.9 Ma) strata rarely yield sauropod body fossils, therefore, isolated teeth can help to elucidate the faunal assemblages from North Africa, and their relations with those of contemporaneous beds and geographically close assemblages. Eighteen isolated sauropod teeth from three localities (Erfoud and Taouz, Morocco, and Algeria) are studied here, to assess whether the teeth can be ascribed to a specific clade, and whether different tooth morphotypes can be found in the samples. Two general morphotypes are found, based on enamel wrinkling and general tooth morphology. Morphotype I, with mainly rugose enamel wrinkling, pronounced carinae, lemon-shaped to (sub)cylindrical cross-section and mesiodistal tapering towards an apical tip, shows affinities to titanosauriforms and titanosaurs. Morphotype II, characterized by more smooth enamel, cylindrical cross-section, rectangular teeth with no apical tapering and both labial and lingual wear facets, shows similarities to rebbachisaurids. Moreover, similarities are found between these northwest African tooth morphotypes, and tooth morphotypes from titanosaurs and rebbachisaurids from both contemporaneous finds from north and central Africa, as well as from the latest Cretaceous (Campanian–Maastrichtian, 83.6 Ma–66.0 Ma) of the Ibero-Armorican Island. These results support previous hypotheses from earlier studies on faunal exchange and continental connections between North Africa and Southern Europe in the Cretaceous.
Le plateau des Hamadas, situé au sud du Haut Atlas marocain, contient de nombreux
fossiles de vertébrés qui constituent une succession de faunes provenant d’environnements divers
sur une période de près de 10 millions d’années à la base du Crétacé supérieur. La base de la
série crétacée est constituée des formations Ifezouane et Aoufous, correspondant à des plaines
alluviales et des deltas pour la première, et à des milieux paraliques et de sebkhas pour la seconde.
Les fossiles de l’assemblage composite des « Kem Kem beds » se trouvent le long d’une falaise de
250 km située le long de la frontière algérienne et au nord-est de la ville d’Erfoud. Les origines
géographique et stratigraphique de nombreuses découvertes scientifiquement importantes
sont imprécises car les fossiles sont souvent découverts par les personnes de la région qui les
exploitent dans un but commercial. La majorité des spécimens semblent provenir du sommet
de la formation Ifezouane. Cet assemblage, comprenant des requins, des poissons osseux, des
amphibiens, des squamates, des tortues, des crocodiles, des dinosaures sauropodes et théropodes
ainsi que des ptérosaures, fait partie d’un ensemble de faunes continentales rencontrées dans
une grande partie de l’Afrique du Nord dans la série nommée « Continental intercalaire ». Les
différents assemblages du « Continental intercalaire » sont d’âges variables, s’échelonnant de la
base du Crétacé inférieur (peut-être même du Jurassique) jusqu’à la base du Crétacé supérieur.
L’assemblage composite des « Kem Kem beds » peut être rapporté au Cénomanien inférieur sur
la base de similitudes taxonomiques avec la faune bien datée de Bahariya en Égypte. Plusieurs
traits écologiques caractérisent cet ensemble : 1) une abondance de taxons, en particulier chez les
poissons et les dinosaures, représentés par des individus de grande taille ; 2) la présence quasi exclusive de dinosaures saurischiens et 3) la surreprésentation des restes de carnivores par rapport
aux herbivores.
Au-dessus de la formation Ifezouane, la formation Aoufous est surmontée de la formation
Akrabou, constituée de sédiments marins déposés lors de la grande transgression du Cénomano-
Turonien. La faune d’Agoult, située près de cette limite cénomano-turonienne sur le lieu même où
Lavocat découvrit des restes de dinosaures dans les années 1950 (Gara Sbâa), contient des poissons
qui vivaient dans un environnement marin restreint, calme et peu profond. Plus au nord, dans la
région de Goulmima, un assemblage de poissons, essentiellement des téléostéens, et de reptiles
marins (mosasauroïdes, polycotylidés, elasmosauridés, protostegidés) est connu dans les niveaux
du Turonien inférieur.
D’un point de vue palaéobiogéographique, l’assemblage composite des « Kem Kem beds »
est proche d’autres faunes d’Afrique du nord et du nord-est du Brésil, indiquant l’existence
d’écosystèmes relativement homogènes s’étendant sur de gigantesques régions. La faune d’Agoult
montre aussi de fortes affinités avec les faunes de la Téthys centrale et du Nord du Mexique, alors
que l’assemblage de Goulmima est proche des faunes de la Téthys occidentale et de la Western
Interior seaway et, dans une moindre mesure, de l’Atlantique Sud.
Sigilmassasaurus brevicollis is an enigmatic theropod dinosaur from the early Late Cretaceous (Cenomanian) of Morocco, originally based on a few isolated cervical vertebrae. Ever since its original description, both its taxonomic validity and systematic affinities were contentious. Originally considered to represent its own family, Sigilmassasauridae, the genus has variously been suggested to represent a carcharodontosaurid, an ornithischian, and, more recently, a spinosaurid. Here we describe new remains referrable to this taxon and re-evaluate its taxonomic status and systematic affinities. Based on the new remains, a re-evaluation of the original materials, and comparisons with other spinosaurids, the holotype of Sigilmassasaurus brevicollis is identified as an anterior dorsal, rather than a cervical vertebra, and differences between elements referred to this taxon can be explained by different positions of the elements in question within the vertebral column. Many characters used previously to diagnose the genus and species are found to be more widespread among basal tetanurans, and specifically spinosaurids. However, the taxon shows several autapomorphies that support its validity, including the presence of a strongly rugose, ventrally offset triangular platform that is confluent with a ventral keel anteriorly in the mid-cervical vertebral centra and a strongly reduced lateral neural arch lamination, with no or an incomplete distinction between anterior and posterior centrodiapophyseal laminae in the posterior cervical and anterior dorsal vertebrae. We argue furthermore that Spinosaurus maroccanus, also described on the basis of isolated cervical vertebrae from the same stratigraphic unit and in the same paper as Sigilmassasaurus brevicollis, is a subjective synonym of the latter. Both a detailed comparison of this taxon with other theropods and a formal phylogenetic analysis support spinosaurid affintities for Sigilmassasaurus. However, we reject the recently
Although pterosaurs from Africa are still rare, in recent years several specimens have been described from the Kem Kem beds (Upper Cretaceous, Cenomanian) of Morocco. Here we describe four additional specimens from this informal lithostratigraphic unit: a jaw fragment, two mid-cervical vertebrae, and a humerus. All these specimens show three-dimensional preservation, differing much from the flat condition found in most pterosaur material. The vertebrae are particularly well preserved, and allow accurate observations on the pneumatization of the neural arch. Based on comparable material, we show that at least two edentulous pterosaur species were present in this informal lithostratigraphic unit, thus adding to the growing evidence of considerable pterosaur diversity in northwestern Africa during the "middle" Cretaceous. So far, the Kem Kem beds have the most diverse pterosaur fauna in this continent, with the presence of anhanguerids, azhdarchids, pteranodontids, and tapejarids.
Although tracks of dinosaurs are well known from Upper Jurassic sediments, tracks of non-dinosaurian vertebrates are fairly rare. The Upper Jurassic Lastres Formations of Asturias in northern Spain contain many vertebrate tracksites that include footprints and trackways of non-dinosaurian tetrapods. Several of these tracks are natural casts of pentadactyl to tridactyl footprints with digits connected by arched structures. The digits are short with deep scratch marks oriented anteriorly. The Asturian tracks show a high degree of morphological similarity to other specimens previously described as possible turtle tracks. Observations from extant turtle trackways show some surprising similarities with the fossil material. The tracks are here interpreted as having been made by turtles partially buoyed by water or by turtles walking in a slightly wet subaerial environment.
Diverse crocodyliforms have been discovered in recent years in Cretaceous rocks on southern landmasses formerly composing Gondwana. We report here on six species from the Sahara with an array of trophic adaptations that significantly deepen our current understanding of African crocodyliform diversity during the Cretaceous period. We describe two of these species (Anatosuchus minor, Araripesuchus wegeneri) from nearly complete skulls and partial articulated skeletons from the Lower Cretaceous Elrhaz Formation (Aptian-Albian) of Niger. The remaining four species (Araripesuchus rattoides sp. n., Kaprosuchus saharicus gen. n. sp. n., Laganosuchus thaumastos gen. n. sp. n., Laganosuchus maghrebensis gen. n. sp. n.) come from contemporaneous Upper Cretaceous formations (Cenomanian) in Niger and Morocco.
Isolated theropod teeth, from the Cenomanian Kem Kem beds of the Tafilalt region, southern Morocco, are described. Some of these teeth are referred to the species Carcharodontosaurus saharicus, and others to the family Dromaeosauridae. This is the first record of this family in northwestern Africa. Whereas Carcharodontosaurus seems to have Gondwanan affinities, the Dromaeosauridae can be considered as northern hemisphere faunal elements, which indicate either an episode of faunal interchange between Laurasia and Gondwana in the Cretaceous, or derivation from an earlier (Jurassic) faunal assemblage with a vast Laurasian-Gondwanan distribution
We report an isolated frontal of a large-bodied theropod from the Cenomanian “Kern Kern beds” of Morocco with an unusual morphology that we refer to a new carcharodontosaurid distinct from the sympatric Carcharodontosaurus. The specimen shows an unique combination of plesiomorphic and potentially autapomorphic features: very thick and broad bone with a complex saddle-shaped dorsal surface, and a narrow vertical lamina between the prefrontal and lacrimal facets. This study supports the hypothesis that a fourth large theropod was present in the Cenomanian of Morocco together with Carcharodontosaurus, Deltadromeus, and Spinosaurus.
Two different theropod pedal unguals from the Upper Cretaceous (Cenomanian) of Tafilalt, Morocco are described. One of them belongs to a slender kind of abelisauroid theropod. Comparison with other members of this group reveals that abelisauroid unguals show high morphological diversity in congruence with what is known on other parts of the skeleton. Another large-sized ungual resembles that described as "Spinosaurus B" by Stromer (1934), and is here referred with caution to Theropoda, although ignoring to which theropod lineage it belongs. Comparisons between northern (e.g., Sahara, Africa and Brazil) and southern (e.g., Patagonia) Gondwanan Cretaceous dinosaur faunas reveal some differences leading to the conclusion that Gondwanan faunas were not uniform across this supercontinent.
Some small and medium-sized crocodylomorph footprints are described from the Late Jurassic (Kimmeridgian) coastal and deltaic units of the northern Spain (Asturias). There are at least four footprint morphotypes. Three of them, with well preserved trackways, are included in the ichnogenus Crocodylopodus (Crocodylopodus isp. and Crocodylopodus meijidei); the fourth one, documented by some isolated large footprints, is referable to the ichnogenus Hatcherichnus. This ichnoassociation confirms the presence of small crocodilians in palaeoenvironments apparently dominated by dinosaurs. The presence of Hatcherichnus seems to confirm the affinity between the Iberian and North American ichnofaunas.
Dinosaur tracks are biogenic, sedimentary structures
and not body fossils or biological objects in the common
sense. They result from the complex interaction of the
kinematics of the trackmaker, its foot anatomy, and the
substrate properties, and from taphonomic processes
acting prior to the incorporation of the tracks into the
sedimentary record. The objective of this work is an
interdisciplinary study of a large sample of dinosaur
tracks and trackways linking sedimentology with vertebrate
ichnology, palaeontology, and palaeoecology.
Excellent conditions are provided by the Late
Jurassic (Kimmeridgian) Chevenez—Combe Ronde
tracksite, which is one of several tracksites located
on the future course of the Transjurane highway
near Porrentruy (Canton Jura, NW Switzerland).
Here, eight superimposed dinosaur track-bearing
surfaces were systematically excavated level-by-level
within a 0,65 m thick laminite interval, unearthing
almost 1400 dinosaur tracks. The main track level,
located at the base of the interval, is the most diverse
ichnoassemblage composed of 14 trackways of tiny
(Pes Length < 25 cm) and small (25 cm < PL < 50 cm)
sauropods and 43 trackways of minute (PL < 10 cm),
small (10 cm < PL < 20 cm), and medium-sized
(20 cm < PL < 30 cm) bipedal, tridactyl dinosaurs.
The main issues are: (1) identification of true tracks,
undertracks, and overtracks, and their relationships with
substrate properties, their link with the exposure index,
and their utility in the reconstruction of the palaeoenvironment;
(2) implications of the main track level
ichnoassemblage for dinosaur behaviour, the terrestrial
palaeoecosystem, and vertebrate ichnofacies; (3) relationships
between variability in trackway patterns and
configurations with locomotion speed, behaviour, and
substrate properties as well as implications for locomotion
capabilities; (4) Quantification and relevance
of sauropod trackway gauge; and (5) interpretation of
manus-dominated and pes-only sauropod trackways.
The approach is first actualistic by studying
human footprints and processes acting during their
formation and preservation on modern tidal-flats.
In these highly structured environments, microbial
mats are ubiquitous, strongly facies-specific, and
occupy a key position during and after footprint formation.
Undertracks readily form in biolaminated
sediment, whilst underprints and deep tracks are
common in unlaminated, water-saturated sediment.
Most consolidated vertebrate tracks are affected
by taphonomic processes, including renewed
and/or repeated growth of microbial mats leading
to the formation of modified true tracks, internal
overtracks (track fills), and overtracks.
The sauropod tracks and the encasing laminite
interval of the Combe Ronde site are then subject
of detailed sedimentological and taphonomical
analyses. This discloses the sediment properties at
the time of track formation and reveals the processes
modifying the tracks during subaerial exposure
and integrating them into the sedimentary record.
Track morphology, associated track features,
and sedimentary features can be linked with the
exposure index, identifying the palaeoenvironment
as a supratidal flat not located in close proximity
to a coastline. These flats were susceptible for
track recording only during short periods after
wetting due to a rainy period or due to occasional
storms. Longer periods of subaerial exposure prior
to burial are indicated by the presence of internal
overtracks and/or overtracks, and rapid covering
up is indicated by the lack of overtracks on top
of tracks with large displacement rims. Crosssections
of sauropod tracks provide insight into
the consolidation history of the substrate prior to
track formation and into the walking dynamics
of dinosaurs, confirming that sauropods put their
hindfeet in a pronounced plantigrade way on the
ground.
The level-by-level superimposition of the studied
surfaces enables to identify true tracks, undertracks,
and overtracks. The best-defined true tracks
(anatomical morphotypes) of the main track level
are then used for ichnotaxonomy and trackmaker
identification, and the detailed analyses of trackway
parameters, including trackway gauge, provide
insight into the locomotion capabilities of dinosaurs.
The best-defined minute and small tridactyl tracks can
be assigned to the ichnogenus Carmelopodus, extending
it from the Middle Jurassic into the Late Jurassic.
These tracks were likely left by a small theropod dinosaur
similar in size to Compsognathus or Juravenator.
The medium-sized tridactyl tracks of morphotype II
exhibit some of the typical features of the ichnogenus
Therangospodus (attributed to large and robust theropods)
but also some of ornithopod ichnotaxa.
The sauropod trackways show a wide range of patterns
and configurations but are all medium- to widegauge.
Therefore, they are assigned tentatively to the
ichnogenus Brontopodus attributed to derived “brachiosaurid”
or “titanosaurid” dinosaurs. The variability
of the trackways reflects the general locomotion
capabilities of the trackmakers and is an expression of
individual walking style and behaviour, which may be
related to substrate properties. Trackway patterns (the
degree of manus overprinting by the pes) and different
trackway configurations including trackway gauge are
not only related to locomotion speed, and they provide
no evidence of a relationship with ontogeny.
The gauge of sauropod trackways can be
quantified with the pes trackway ratio and the here
defined [WAP/PL]-ratio (Width of the pes Angulation
Pattern / Pes Length). Gauge is possibly related to
the substrate and the behaviour of the trackmaker
adapting to it, but this does not change the overall
medium-gauge to wide-gauge appearance of the
trackways. The manus-dominated and pes-only
sauropod trackways of the Combe Ronde site are
explained by trackmakers exerting more pressure on
the manus than the pes, and by overprinting of the
manus by the pes, respectively.
The alignment of trackways on the main track
level shows no evidence of a nearby shoreline and
of interactions between the different groups of
dinosaurs. It indicates gregarious behaviour amongst
tiny and small sauropods, and suggests that minute
and small bipedal dinosaurs were frequent visitors on
the supratidal flats.
The ichnoassemblage of the main track level is
the first one found in the Jura Mountains displaying
abundant minute and small tridactyl tracks. This is
also typical for the other Ajoie ichnoassemblages,
which further exhibit tracks of tiny to large (up to 1,1
m PL) sauropods, and tracks of medium-sized to large
(up to 0,8 m PL) bipedal dinosaurs. Sauropod
trackways include narrow-gauge and wide-gauge
trackways indicating the presence of “basal” and
derived sauropods. This suggests that dwarfed
insular animals can be excluded as trackmakers of
the tiny and small sauropod trackways of the Ajoie
ichnoassemblages and the Combe Ronde tracksite
and that the Jura carbonate platform was connected
with the landmasses of the London-Brabant Massif
and the Massif Central during periods of emersion.
Dinosaurs used the Jura carbonate platform for the
establishment of in situ, predominantly saurischian
dinosaur populations, but also as a migration corridor
between the massifs.
Because the Ajoie ichnoassemblages are dominated
by small tridactyl tracks, they differ from other
Jurassic tetrapod ichnofacies in carbonate settings,
notably from the Brontopodus ichnofacies. In the
case of those ichnoassemblages commonly attributed
to the Brontopodus ichnofacies, the lack or rareness
of small tridactyl tracks may indicate the absence
of small trackmakers in those palaeoenvironments
or unsuitable conditions for the formation and
preservation of small tracks.
This study highlights the benefits of systematic
and interdisciplinary analyses of dinosaur tracks,
which disclose variations related to behaviour and
to differences in substrate. This allows recognizing
anatomical morphotypes and trackway configurations
representative of typical trackmaker behaviour. The
latter can then also be used in ichnotaxonomical
classification. Similar approaches should be in the
focus of future work and performed on the other
tracksites and ichnoassemblages of the Ajoie.
Together with the evidence from other tracksites of the
Jura Mountains, this will contribute towards a better
understanding of the terrestrial palaeoenvironments
and palaeogeography, and of dinosaur palaeoecology
and palaeobiogeography on the Jura carbonate
platform.
Late Cretaceous (Cenomanian) fossils discovered in the Kem Kem region of Morocco include large predatory dinosaurs that inhabited
Africa as it drifted into geographic isolation. One, represented by a skull approximately 1.6 meters in length, is an advanced
allosauroid referable to the African genus Carcharodontosaurus. Another, represented by a partial skeleton with slender proportions, is a new basal coelurosaur closely resembling the Egyptian
genus Bahariasaurus. Comparisons with Cretaceous theropods from other continents reveal a previously unrecognized global radiation of carcharodontosaurid
predators. Substantial geographic differentiation of dinosaurian faunas in response to continental drift appears to have arisen
abruptly at the beginning of the Late Cretaceous.
The Late Jurassic Iouaridène tracksite has been studied for decades and is well-known for the reference trackway of Breviparopus taghbaloutensis. These siliciclastic flood-plain deposits bear probably more than 1500 tracks, and at least 21 trampled levels: they yield tracks of medium to very large sauropods, possible stegosaurs and theropods. The first accurate description of the footprint association made by biped trackmakers is proposed herein. More than six hundred foot prints and more than a hundred trackways has been mapped and analysed; this led to the definition of four tridactyl and two tetradactyl morphotypes, mainly produced by small to very large theropods, while probable small ornithopod tracks are also present. The bipedal footprint association is dominated by medium-large theropods, which are also the most abundant type. The taxonomical attribution of the morphotypes is made difficult by the poor preservation of many specimens. Further more, for the most abundant theropod tracks, those with "megalosaurian" affinity, there is also a complex ichnotaxonomical situation, that makes the attributions yet more challenging; however, it was possible to recognize the great affinity of the tridactyl specimens with the Megalosauripus tracks from the Iberian Peninsula and North America. Three-di mensional models were generated from the moulds of the best-preserved specimens to render a more detailed description and for easier access to the specimens.
Fossils of vertebrates have been found in great abundance in the continental and marine early Late Cretaceous sediments of Southeastern Morocco for more than 50 years. About 80 vertebrate taxa have so far been recorded from this region, many of which were recognised and diagnosed for the first time based on specimens recovered from these sediments. In this paper, we use published data together with new field data to present an updated overview of Moroccan early Late Cretaceous vertebrate assemblages. The Cretaceous series we have studied encompasses three Formations, the Ifezouane and Aoufous Formations, which are continental and deltaic in origin and are often grouped under the name “Kem Kem beds”, and the Akrabou Formation which is marine in origin. New field observations allow us to place four recognised vertebrate clusters, corresponding to one compound assemblage and three assemblages, within a general temporal framework. In particular, two ammonite bioevents characterise the lower part of the Upper Cenomanian (Calycoceras guerangeri Zone) at the base of the Akrabou Formation and the upper part of the Lower Turonian (Mammites nodosoides Zone), that may extend into the Middle Turonian within the Akrabou Formation, and allow for more accurate dating of the marine sequence in the study area. We are not yet able to distinguish a specific assemblage that characterises the Ifezouane Formation when compared to the similar Aoufous Formation, and as a result we regard the oldest of the four vertebrate “assemblages” in this region to be the compound assemblage of the “Kem Kem beds”. This well-known vertebrate assemblage comprises a mixture of terrestrial (and aerial), freshwater and brackish vertebrates. The archosaur component of this fauna appears to show an intriguingly high proportion of large-bodied carnivorous taxa, which may indicate a peculiar trophic chain, although collecting biases alter this palaeontological signal. A small and restricted assemblage, the OT1 assemblage, possibly corresponds to a specific, localised ecosystem within the Kem Kem beds compound assemblage. Microfossils and facies from the Aoufous Formation, corresponding to the top of the compound assemblage, provide evidence of extremely abiotic conditions (hypersalinity), and thus of great environmental instability. At the base of the Akrabou Formation the first ammonite bioevent, Neolobites, corresponds to the onset of the marine transgression in the early Late Cenomanian while the Agoult assemblage (Late Cenomanian?) contains a variety of small fish species that have Central Tethyan affinities. Finally, the youngest Mammites bioevent in the late Early Turonian corresponds to a deepening of the marine environment: this sequence is isochronous with the Goulmima assemblage, a diverse collection of fish and other marine taxa, and shows affinities with taxa from the South Atlantic, the Central Tethys and the Western Interior seaway of North America, and further highlights the biogeographical importance of these North African Late Cretaceous assemblages.
Crocodyliforms were one of the most successful groups of Mesozoic tetrapods, radiating into terrestrial, semiaquatic and marine environments, while occupying numerous trophic niches, including carnivorous, insectivorous, herbivorous, and piscivorous species. Among these taxa were the enigmatic, poorly represented flat-headed crocodyliforms from the late Cretaceous of northern Africa. Here we report a new, giant crocodyliform from the early Late Cretaceous (Cenomanian) Kem Kem Formation of Morocco. Represented by a partial braincase, the taxon has an extremely long, flat skull with large jaw and craniocervical muscles. The skull roof is ridged and ornamented with a broad, rough boss surrounded by significant vascular impressions, likely forming an integumentary structure unique among crocodyliforms. Size estimates using endocranial volume indicate the specimen was very large. The taxon possesses robust laterosphenoids with laterally oriented capitate processes and isolated epipterygoids, features allying it with derived eusuchians. Phylogenetic analysis finds the taxon to be a derived eusuchian and sister taxon to Aegyptosuchus, a poorly understood, early Late Cretaceous taxon from the Bahariya formation. This clade forms the sister clade of crown-group Crocodylia, making these taxa the earliest eusuchian crocodyliforms known from Africa. These results shift phylogenetic and biogeographical hypotheses on the origin of modern crocodylians towards the circum-Tethyean region and provide important new data on eusuchian morphology and evolution.
The most recent account of Bueckeburgichnus maximus Kuhn 1958, a distinctive theropod dinosaur track from the Lower Cretaceous of Germany, is shown to be based on a referred specimen mistakenly identified as the holotype and the correct name of this taxon is deemed to be Megalosauripus maximus (Kuhn 1958). This minor revision has important consequences for nomenclature of the many European, Asian, North American and Australian dinosaur tracks attributed to megalosaurian theropods. Many of those tracks were named Megalosauripus, but that name has a confusing multiplicity of meanings and it should be restricted to the highly characteristic dinosaur track formerly identified as Bueckeburgichnus. Other tracks named "Megalosauripus”; (in its several other senses) will require new nomenclature, despite their extensive and repeated revision since 1996. It is recommended that future revision should adopt conventions of the International Code of Zoological Nomenclature. Although previous revisions expressed an intention to adhere to those conventions, these were not put into practice, with the unfortunate result of multiplying the problems that surround the nomenclature of megalosaur tracks. Introduction of the name Megalosauripus maximus (Kuhn 1958) eliminates those burgeoning problems and permits the introduction of new and objective nomenclature for presumed megalosaur tracks.
The Kem Kem beds in South Eastern Morocco contain a rich early Upper (or possibly late Lower) Cretaceous vertebrate assemblage. Fragmentary remains, predominantly teeth and jaw tips, represent several kinds of pterosaur although only one species, the ornithocheirid Coloborhynchus moroccensis, has been named. Here, we describe a new azhdarchid pterosaur, Alanqa saharica nov. gen. nov. sp., based on an almost complete well preserved mandibular symphysis from Aferdou N'Chaft. We assign additional fragmentary jaw remains, some of which have been tentatively identified as azhdarchid and pteranodontid, to this new taxon which is distinguished from other azhdarchids by a remarkably straight, elongate, lance-shaped mandibular symphysis that bears a pronounced dorsal eminence near the posterior end of its dorsal (occlusal) surface. Most remains, including the holotype, represent individuals of approximately three to four meters in wingspan, but a fragment of a large cervical vertebra, that probably also belongs to A. saharica, suggests that wingspans of six meters were achieved in this species. The Kem Kem beds have yielded the most diverse pterosaur assemblage yet reported from Africa and provide the first clear evidence for the presence of azhdarchids in Gondwana at the start of the Late Cretaceous. This, the relatively large size achieved by Alanqa, and the additional evidence of variable jaw morphology in azhdarchids provided by this taxon, indicates a longer and more complex history for this clade than previously suspected.
Pterosaur tracks from the Candeleros Member of the Rio Limay Formation (Albian–Cenomanian) at Lake Ezequiel Ramos Mexı́a, Neuquén Province, Argentina, are the first pterosaurian tracks reported from South America and from the Southern Hemisphere. Although the individual footprints are not well preserved, some trackway segments and manus-pes sets are preserved, suggesting that the tracks can be assigned to cf. Pteraichnus ichnosp. indet. The palaeoenvironmental setting of the track beds is a lake shoreline, where dinosaur tracks also occur. The discovery is consistent with reports of well known pterosaur-bearing Lagerstätten in the Lower Cretaceous of South America.
The 'Gres rouges infracenomaniens' of southern Morocco, possibly of Albian age, contain evidence of one of the most diversified dinosaur assemblages known from Africa, including a relatively long-necked species of Spinosaurus and abundant but isolated bones of a peculiar theropod ('Spinosaurus B' of Stromer 1934). Also preserved are the oldest records of abelisaurids and among the oldest records of titanosaurids in Africa. Bones of infantile didnosaurs are present. The assemblage resembles that of the Bahariya Formation more than that of Gadoufaoua, possibly because of a trophic dependence upon large, freshwater fishes. It was more closely linked zoogeographically to South America than to North America.
The maximum body size attained by a predator species reflects a balance between two conflicting demands. Population densities of huge predators must be very low to prevent their over-exploiting their food base of prey species. At the same time, however, total population sizes must be large enough to prevent change extinction of the predator species. Calculations suggest that a dinosaur-sized predatory mammal might find it difficult or even impossible simultaneously to maintain low enough population densities and large enough population sizes. Flesh-eating dinosaurs were able to get around this size constraint due to one, or more likely some combination, of the following factors: larger population densities and/or faster population turnover rates of their herbivorous dinosaur prey than expected for dinosaur-sized herbivorous mammals, oviparity in dinosaurs, differences in diet between juvenile and adult theropods, and lower mass-specific food consumption rates of carnivorous dinosaurs. -from Author
Fragmentary cranial bones of dinosaur origin have been recently recovered from the Kern Kern beds (Upper Cretaceous, Cenomanian) of Morocco. They include two incompletely preserved maxillary bones evidencing diagnostic features of abelisaurid theropods. These new finds provide further evidence of Abelisauridae in the Late Cretaceous of Morocco.
An unusually high proportion of large-bodied carnivorous theropod dinosaurs has been reported from the Moroccan Late Cretaceous Kem Kem Formation, a well-known package of North Africa vertebrate fossil-bearing sediments. We investigate whether recorded proportions of predator and prey taxa in Kem Kem sediments are real, or an artifact generated by collecting biases, by comparing field data to counts of fossil vertebrates from Moroccan fossil shops. The application of common techniques for standardizing ecological survey data confirms that previous workers have been misled by the acquisition by museums of specimens from commercial collectors rather than from detailed field surveying. Claims that an unusual number of theropod dinosaurs were present in North Africa Late Cretaceous ecosystems are likely the result of biases due to both commercial activity and collectorship biases.
We recognize three fundamental terms in ichnology: (1) ichnoassemblage, which is an assemblage of ichnofossils conceptually equivalent to an assemblage of body fossils; (2) ichnocoenosis, which is a trace fossil assemblage produced by a biological community that can be characterized by morphological criteria; and (3) ichnofacies, which refers to recurrent ichnocoenoses that represent a significant portion of Phanerozoic time. There are two different kinds of ichnofacies, ethoichnofacies (mostly invertebrate ichnofacies) and biotaxonichnofacies (mostly tetrapod ichnofacies). Nonmarine invertebrate ichnologists now recognize five archetypal ichnofacies (Mermia, Skolithos, Scoyenia, Coprinisphaera, Psilonichnus) to which we add the Octopodichnus ichnofacies. We propose a coherent and consistent classification and nomenclature for tetrapod ichnofacies. We name five archetypal vertebrate ichnofacies for nonmarine environments: Chelichnus, Grallator, Brontopodus, Batrachichnus and Characichnos ichnofacies.
Do dinosaurs from the Moroccan Kem Kem formation provide evidence for an ecosystem dramatically different from anything seen today? More likely the common palaeontological problem of time-averaging has had a part to play.
At least 50 pterosaur track sites have been reported from Late Jurassic through Late Cretaceous localities in North America, Europe, East Asia and South America, plus one possible site from north Africa. Tracks from these sites have been assigned to 11 ichnospecies in four ichnogenera. Of these, Pteraichnus is by far the most prevalent, well-preserved, and represented by multiple (presently eight) ichnospecies. The majority of Pteraichnus tracksites are Late Jurassic or earliest Cretaceous (Berriasian) in age. In contrast, the other three ichnogenera - Purbeckopus, Haenamichnus and possibly Agadirichnus- are all represented by single ichnospecies from single localities and are based on relatively poorly-preserved, earliest through latest Cretaceous material. At least 16 Late Jurassic, well-preserved Pteraichnus samples from marginal marine deposits in western North America are dominated by small tracks (pes length ~2-10 cm), often in trackways. The two ichnospecies from this region (P. saltwashensis and P. stokesi) are both based on trackway segments and differ in style of preservation. Other Late Jurassic trackways from marginal marine deposits in France and Poland are also small (pes length typically ∼3-5 cm); much larger tracks (pes length 18 cm) are known from a single locality in Asturias, Spain. It is unclear whether most Pteraichnus represent pterodactyloid or "rhamphorhynchoid" pterosaurs because the diagnostic impressions of pedal digit V are rarely clearly and unambiguously impressed. Six basal Cretaceous (Berriasian) Pteraichnus ichnospecies from a least a dozen sites in Soria, Spain are also mostly small (footprint length ∼1-5 cm) and based on as yet insufficiently described, isolated footprints, not trackways. As a result, several of these ichnospecies are probably nomina dubia. The contemporaneous Purbeckopus from England is much larger (foot length -19-22 cm). Although small, Pteraichnus-like tracks have been reported sporadically from the post-Berriasian Cretaceous, most are much larger (foot length 10-20 cm) and, in the case of Haenamichnus from Korea, reach 30-33 cm. It is unclear whether Agadirichnus from the Late Cretaceous of Morocco (foot length 10-12 cm) is pterosaurian. Most Cretaceous sites represent lacustrine, not marginal marine, habitats. Both Jurassic and Cretaceous assemblages often contain very high track densities and a range of track sizes associated with invertebrate traces. This suggests that diverse pterosaurian flocks may have congregated in large numbers to feed. Some assemblages reveal swim tracks that suggest pterosaurs floated in shallow water, touching the submerged substrate with only their hind feet. These swim track assemblages also contain possible beak traces that may indicate feeding.
A varied ichnocoenosis in the Cretaceous Kem Kem beds of Morocco. 71st meeting of the Society of Verte-brate Paleontology, Abstract Book
- N.-E Jalil
- M Belvedere
- A Breda
- G Gattolin
- G Dyke
Jalil, N.-E., Belvedere, M., Breda, A., Gattolin, G., Dyke, G., 2011. A varied ichnocoenosis in the Cretaceous Kem Kem beds of Morocco. 71st meeting of the Society of Verte-brate Paleontology, Abstract Book, Las Vegas, Nevada, USA, 2–5.11.2011. Journal of Vertebrate Paleontology 3, 167 (Special Issue)