Here there were early Giants: The Aganane Formation (Pliensbachian, Morocco) and its Record Dinosaur ichnotaxa

^{_{Tracksites like the one Above located at Vallée des Aït Bougmez evidence the early trends to Gigantism of Saurischian and maybe Ornithischian Dinosaurs (Photo from Le Maroc Avant l´historie)}}

_{The initial evolutionary intervals of the Dinosauria group are interesting, specially those works focused on the initial, Triassic-Lower Jurassic trends seen in the Sauropodomorph group, having been subject of multiple studies related to the whole group (ex. Apaldetti et al., 2018; Botha et al, 2022; Lefebvre et al, 2022) those focused only on the Sauropod group (ex. Sander et al, 2011), both focused mostly on bones, less on Footprints. The Theropod group sees the opposite trend, with multiple works focused on the size increase based on both bones and mostly, Footprints (ex. Scisio et al., 2017; Dal Sasso, 2018). The overall sampling trends to show a general size increase of both groups after the Tria/Jura boundary, while Ornithischians trend to also follow this, as seen on the apperance of larger taxa such as stem Thyreophora (ex. Scelidosaurus). Here we will focus on a series of tracksites recovered from the Pliensbachian-aged Aganane Formation of the Central High Atlas of Morocco, that feature enormous specimens for it´s age, and what do these deposits tell us about the tendency to gigantism in Early Jurassic dinosaurs.}

This post is also in homage to Martin Lockley, who contributed to the study of these footprints and will be remembered for his general contribution to Ichnology.

The Aganane Formation contextualized

_{^{Fig.1 Simplified geology & stratigraphy of the central High Atlas, the Aganane Formation is included in the Lower-Upper Lias sequences (extracted from Teixell et al., 2003)}}

_{The Aganane Formation represents a proximal/circalittoral environment, being the major remnant of a local Bahamian/Floridian-alike carbonate Platform, whose best modern analoge is the Island of Andros in Bahamas, alocated mostly in the Demnate, Azilal, Beni Mellal and Zaouiat Ahansal (Fig.1; Fig2), evolving W-E/SE from nearshore intertidal or supratidal (lagoonal) environments to shallow marine ones (Fig.2).}

_{Located in the Pliensbachian at an almost tropical paleolatitude, the region constituted the western termination against the ancient massif of the «Atlas Gulf», open to the East on the Tethys Sea. North of the Demnate accident extended a vast subsiding Tidal flat where carbonates with gypsum ghosts and stromatolitic laminites were deposited, cut by desiccation polygons, while on the edge of the accident, active at this time, intercalations of Lignite, certainly resulting from the dismantling of forested areas located to the South as evidenced by the traces of roots of the sandstones at the base of the lateral west Aït-Bazzi Formation in Ait Toutline. Dinosaurs, both herbivorous and carnivorous, roamed this vast maritime marsh. In the East, excluding the Demnat region, on the edge of a low continent, gypsum precipitated in sebkhas Under a climate that was probably quite arid. Locally, more massive facies with large Lamellibranches separated the “tidal flat” from the open Ammonite Sea whose western extension borders the eastern edge of the Azilal Map.}

_{The supratidal area exhibits diverse deposits, including continental sediments with quartz-rich cricundant deposits, gypsum layers, and caliche crusts, suggesting a connection to river courses and coastal Sabkhas (Lee & Burgess, 1978; Septfontaine, 1984; 1985). Intertidal deposits feature algal laminates, bioturbated shales, and wackestones with storm sequences, resembling modern coastal environments like Australia´s Shark Bay. Subtidal deposits range from shallow water to coastal lagoons, with accumulations of skeletal lime packstones, oolitic tidal deltas, and coral reefs. Further east towards the Dadés Area, subtidal conditions transition to flint-bearing calcareous shales with sparse faunas, marking the appearance of ammonites. Changes in surface conditions were consistently reflected in rock sequences by diagenetic features, potentially correlating with major events like hurricanes (Burgess & Lee, 1978; Septfontaine, 1984; 1985).}

_{^{Fig.2 selected and simplified stratigraphical charts of the central High Atlas (extracted from Vergés, 2017)}}

_{The lithology of the formation displays varying lime cycles, with alternating sequences of impure limestones and red marls of decameter thickness (Septfontaine, 1985; Milhi, 1992). Light grey limestone, rich in fossils and sometimes rathy, forms thick banks ranging from decimeters to meters. These limestone/marl interbed sequences indicate continental influence, often exhibiting dry cracks. The banks, described as elementary sequences, form megasequences up to 20 meters thick, with tendril thickness decreasing from bottom to top (Milhi, 1992). Megasequences typically end with retic marl layers and stromatolitic layers accompanied by desiccation phenomena. Occasional tempestite horizons are present. In the Boutonnière of Talmest, the base of the Aganane Formation features a conglomerate horizon up to 60 meters thick, gradually thinning towards the southwest (Milhi, 1992; Milhi et al., 2002).}

_{The Aganane Formation shows a particularly high faunal diversity, dominated by thick-shelled Bivalve bioherms (Perna, Megalodontidae, Cochlearites, Lithiotis, Opisoma) exceptionally large (up to 1 m in size). They sometimes form wide biostromes with gastropods, echinoderms, brachiopods, ostracods, solitary corals and algae (Dasycladaceae & Solenoporaceae). Also widespread are lituolid foraminifera, cyanobacteria and thaumatoporellae (Septfontaine, 1985; Milhi, 1992). Locations such as Ait Athmane recover these «Reefs» with a strong zonation, starting with the bivalves Gervilleioperna and Mytiloperna, restricted to intertidal and shallow-subtidal, cross-bedded floatstones, later evolving to layers with evidence of subaerial exposure, including lagoonal marls, and bioturbated red mudstones with root traces and calcrete, all living in a sheltered lagoon in the interior of the local carbonate platform, similar to the Rotzo Formation of the Italian Trento Platform (Brame et al., 2019).}

_{The faunal diversity generally indicates normal salinity conditions and warm, well-lit water. The presence of Lituolids and palaeodasycladaceae are usually indicative of protected lagoon tides below the wave base (Septfontaine, 1984; 1985; Milhi, 1992; Fig.3). Microfacies analysis shows that each bank corresponds to an elementary regressive cycle, with megacycles (10-20 m) interpreted in connection with global sea-level fluctuations and regional tectonic development (Milhi, 1992). Reworked horizons and rearrangements indicate episodic storms (Milhi, 1992; Fig.3).}

_{Another indicator of emersion are the presence of Dinosaur tracks, described initally in the 30´s and latter revised in the 80´s from several locations in Demnate, Azilal, Ahansal, and then in the 2000´s also from the Dadés Gorges (Plateau et al., 1937; Jenny & Jossen, 1982; Ishigaki & Jossen, 1988; Masrour et al., 2015; Fig.3). They included back in 1988 up to 1300 tracks, the current number is unknow (Nouri, 2007).}

Fig.3: Paleogeographical reconstruction of the Pliensbachian Central High Atlas Area (After Jenny, 1988) with location of the tracksites and environments

The Tracksites & Tracks

_{a) Theropod Tracks}

_{The theropod tracks are locally the most diverse, made of imprints left by small, medium & large taxa. The Small sized theropod imprints include didactyl, tridactyl and tetradactyl theropod trackways (Fig.4). Of these, the main ones are know from the westernmost reported tracksite at Ait Blal, referred originally to «Coelurosaurs» (Jenny & Jossen, 1982). This tracks where revised by Ishigaki & Lockley (2010), who suggested the didactyl tracks may have been badly preserved tridactyl ones (or even dromeaosaurid-like foot morphologies) or maybe even swimming tracks (Milner et al., 2016), while the tetradactyl are probably just convergent with younger-aged coelurosaurs. Beyond this the tracksite records limping behaviour, with animals being forced to modify their gaits either by external substrate conditions, internal factors (such as injury) or both (Ishigaki & Lockley, 2010). The tracks have been recently suggested to belong to the ichnogenus Argoides, of possible Ceratosauroid or Coelophysoid affinities (Molina-Pérez & Larramendi, 2019), however they still need a proper re-revision, as some also resemble Achisauripus & Grallator (Nouri, 2007).}

Fig4. Selection of Small Theropod tracks from the Aït Blal tracksite, including photos from Jenny & Jossen (1982), Ishigaki & Lockley (2010) and ilustrations from Ishigaki & Jossen (1988).

_{Other small theropod tracks are know from the tracksites of Ibaqualiwn, Assif-n-Sremt, Ansous or Tabant, but lack a proper description (Jenny & Jossen, 1982; Nouri, 2007; Masrour et al., 2014; 2015).}

_{Medium-sized theropod tracks are the most featured in literature, yet only a few from Aït Bou Guemmez y adjacent tracksites where described by Ishigaki and Jossen (1988); Nouri (2007) and others have been quoted from tracksites in Ahansal & Dadés areas (Jenny & Jossen, 1982; Masrour et al., 2010; 2014; 2015; Fig.5). Of this ones, Ishigaki & Jossen (1988) & Nouri (2007) featured 3 types (Fig.5), attributed to indet. theropods. Of the featured tracks by Nouri, a few are of a sturdy, short type with a well pronounced claw on toe II, resembling certain Ceratosauroids and Abelisauroids, while other resembles the pes of Piatnitzkysaurus (Fig.5). Of the types presented by Ishigaki & Jossen, 2 resemble the ichnogenus Megalosauripus and 1 the typical Eubrontes tracks common in EJ strata (Molina & Larramendi, 2016). Overall seems like 4 morphotypes are present resembling the pes of younger taxa what can infer either convergence or preservation artifacts, or more unlikely the presence of Early representatives of the Ceratosaurian & Tetanuran clades, otherwise know from Sinemurian & Toarcian layers of Italy, Morocco and Argentina (Allain et al., 2007; Pol et al., 2012; Dal Sasso, 2018; Rauhut et al, 2019).}

Fig.5: Selection of Medium-sized theropod tracks from Ishigaki & Jossen (1988) & Nouri (2007). Ishigaki & Jossen´s type 5 and 8 resemble the ichnogenus Megalosauripus, while 6 is clearly an Eubrontes isp. Nouri´s type 1 resembles the pes of Piatnitzkysaurus, while the others have resemblance with certain ceratosaur pes.

_{Large Theropod tracks are the rarest of the collection, yet include the currently largest know african EJ theropod track, of about 53 cm (Fig.6), referred in recent literature as Carmelopodus isp., of ceratosaurian affinities, suggesting a taxon up to 8 m in length (Molina & Larramendi, 2016). Yet, based on the poor preservation that asignation should be taken with extreme caution, even if large ceratosaurs have been suggested to be present in the EJ of nearby areas (Dal Sasso et al., 2018). This tracks are know from Talsant and the Aït Bou Guemez area, with new specimens from previuosly unsampled localities in the last area such as at Akourbi, awaiting for description (Jenny & Jossen, 1982; Masrour et al., 2010).}

Fig.6 Large Theropod tracks featured in Ishigaki & Jossen (1988). N9 represents the recordholder as the largest Early Jurassic Theropod track from Africa

_{B) Sauropodomorph Tracks}

_{Sauropodomorph tracks are the most diverse & abundant (around 72%) in the Aganane tracksites (Jenny & Jossen, 1982; Nouri, 2007; Masrour et al., 2010; 2014; 2015). These footprints are also the most interesting, since they not only collect very varied morphotypes, but also include the largest examples of both sauropod and Sauropodomorph footprints from the Lower Jurassic recorded so far. Morphotypes referring to Sauropodomorphs include specimens reminiscent of the ichnotaxon cf.Pseudotetrasauropus and others not named, but are clearly dominated by the taxon Otozoum (Jenny & Jossen, 1982; Ishigaki & Jossen, 1988; Nouri, 2007; Masrour et al., 2010; 2014; 2015). Within Ichnotaxon Otozoum, footprints reminiscent of the Gondwanic sauropodomorph Lamplughsaura pes are found, and in locally reported specimens include 46 cm ones from Issil-Aït-Arbi attributed to O. moodii (Masrour et al., 2014; 2014b; Fig.7), and the most important, an enormous track with a reported size around 86 cm from Assif-n-sremt (Ishigaki & Jossen, 1988; Fig.7), what can suggest an animal up to 13-16 m (Molina & Larramendi, 2016). However the size of this massive track must be taken with caution and probably measures around ~60 cm, what is still the largest one reported on literature (Lockley et al., 2024; Fig.7).}

Fig.7 Large Sauropodomorph tracks from Aït-Arbi attributed to O. moodii (Masrour et al., 2014; 2014b) and gigantic one from Assif-n-sremt (Ishigaki & Jossen, 1988)

_{Traces referring to sauropods are the most common and also present a remarkable variability. These are dominated by the taxon Parabrontopodus (and maybe Eosauropus), asymmetrical and primitive, wide gauge with a shape that is rather different from those that are usually seen in well preserved sauropod hind foot tracks with the foot without toe rotation, some with a hand like a horseshoe, which includes in the area the largest sauropod track reported from the Lower Jurassic, measuring 85 cm (Ishigaki & Jossen, 1988; Fig. 8). More recent reports anticipate the presence of even larger specimens, approaching 87 cm (Masrour et al., 2010). If true, they would represent a taxon above 15 m, taking the taxon Tazoudasaurus from the same region as a reference, as where quoted in the past to resemble Vulcanodon while others have been quoted to resemble the more derived pes of Rhoetosaurus (Farlow, 1992; Fig.7). Other record ichnotaxa in the area include large posible Lavinipes(?)-like tracks resembling the pes of «specialized descendants of melanosaurids or blikanasaurids» (Avanzini et al., 2003; Farlow, 1992; Fig.7).}

_{Nouri (2007) reported an extremely anomalous track, superasymmetric, toe II longer than I narrow track, resembling the Neosauropod-atributted ichnogenus Breviparopus, yet the hand imprints are strange and their hand-foot relationship does not fit into the concept such ichnogenus (Fig.7). This outliner track type is extremely unusual for such an early age, and can imply the presence of a Non-Neosauropod with convergent evolved pes, or more unlikely, a stem Neosauropod. Currently the oldest material referred to Neosauropods is based only on teeth or remains very unstable phylogenically from the Toarcian of Argentina (Carballido et al., 2017).}

Fig.8 Selection of Sauropod tracks from the Aganane Formation: A) Issil-n-Aït Arbi tracks and it´s preservational features (Masrour et al., 2015); B) Eosauropus & Parabrontopodus-type tracks, including the largest sauropod track from the Early Jurassic (Ishigaki & Jossen, 1988; Nouri, 2007); C) Rhoetosaurus-alike tracks (Ishigaki & Jossen, 1988; Nouri, 2007); Bizarre Breviparopus resembling track (Nouri, 2007)

_{C) Ornithischian? tracks}

_{Currently, there have been no solid reports of Ornithischia footprints in the Aganane Formation deposits. Ishigaki & Jossen (1988) reported tracks similar in appearance to those left by Stegosaurs, but more recent literature has reclassified them as being made by Sauropodomorphs (Masrour et al., 2014). Recently, tracks reported from the Ansous area have been suggested to be from Thyreophorans, as they can´t be assigned to sauropods by the symmetry and position of the toes, the number of toes of the same same autopodium, the biconvex shape of the hand mark the biconvex shape of the hand mark and the shape of the fingers (Masrour et al., 2014). This asignation must be taken with extreme caution, as the features can be either convergent of a preservation artifact. If this tracks belong truly to a thyreophoran, it must have been an enormous taxon, as tracks measure 58×50 cm (Masrour et al., 2014). Currently the closest in age and provenance Thyreoporan tracks are those atributted to Moyenisauropus lusitanicus, from the Sinemurian Coimbra Formation of Portugal (Figueiredo et al., 2023).}

Conclusion

_{The footprint deposits of the Aganane Formation, apart from providing a rich faunal inventory of the Pliensbachian in North Africa, represent a diverse setting where different groups reached early gigantism in their history (Fig.9). They are a series of sites that require review and publication in greater detail, partly to inventory the footprints and partly to confirm measurements and proportions.}

_{The state of preservation of the tracks in the most recently sampled areas is either good (Ibaqalliwn or Issil-n-Aït Arbi), usually in well-consolidated limestone (Nouri, 2007; Masrour et al., 2010;2014; 2015), but some of the tracksites like Ait Blal suffering from weathering and in need of protection (Ishigaki & Lockley, 2010). Protection and restoration of these deposits is a matter of urgency, and several suggestions have been proposed to do it, including fences to make possible to appreciate the sites as closely as possible without trampling them by stepping on them and damaging them and shelters protect the deposits from insolation and erosion due to rain run-off (Nouri, 2007).}

Fig.9: Selected scaling of the previously featured Tracks from the Aganane Formation, highlighting the unusual amount of large bodied taxa in an Early Jurassic environment. Figures courtesy of Fabio Ale Romero (Dragonthunders), GunnarBivens, Eotyrannu5, Cisiopurple, Gregory S. Paul and PhyloPic. Note that the atributed sillouettes are speculative.

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