Abdelouahed Lagnaoui

COMPLEX TETRAPOD BURROWS FROM MIDDLE TRIASSIC RED BEDS OF THE ARGANA BASIN (WESTERN HIGH ATLAS, MOROCCO)

ABSTRACT Although burrowing ability has been widespread in tetrapods for more than 300 million years, subsurface dwelling structures that indicate communal behavior are poorly evidenced from pre-Cenozoic strata. Here we present recently discovered tetrapod burrows from Middle Triassic red beds of the Argana Basin in central Morocco, whose complexity suggests an origin by gregarious animals. The well-preserved burrows occur in interbedded mudstones and sandstones interpreted as channel and overbank deposits of ephemeral, braided streams. All burrows originate from the top of thick-bedded sandstones and descend as moderately inclined (10°–30°), partially spiral tunnels to laterally extended, branched chambers in underlying mudstones. Tunnel segments are biconvex to planoconvex in cross section, up to 20 cm wide and 12 cm in maximum height and exhibit transverse scratch marks along the ceilings and sidewalls. Distinctive burrow characteristics include a laterally sinuous geometry (wavelength &lgr;  =  38–45 cm; amplitude A  =  5–10 cm) of the tubelike passages and the presence of grouped alcoves in terminal chambers. We attribute the burrows to procolophonids or therapsids based on closely associated tetrapod tracks and the limited diameter of the excavations. Our findings represent the second oldest record of communal fossorial behavior by tetrapods and the oldest example from low-latitude areas. Beyond providing refuge from predators, these elaborate underground structures probably functioned as a buffer against diurnal or seasonal variations of air temperature and humidity in a semiarid habitat that was situated just north of the paleoequator.

Late Triassic Tetrapod-Dominated Ichnoassemblages from the Argana Basin (Western High Atlas, Morocco)

Diverse tetrapod track assemblages with Scoyenia invertebrate traces were discovered in the Triassic Timezgadiouine and Bigoudine formations of the Argana Basin (Western High Atlas, Morocco). The ichnofossils occur in alluvial plain sandstones and mudstones of the Irohalène Member (T5) and Tadart Ouadou Member (T6) considered Carnian-Norian in age by vertebrate remains and palynomorphs. Tetrapod footprints are assigned to Apatopus, Atreipus-Grallator, Eubrontes isp., Parachirotherium, cf. Parachirotherium postchirotherioides, Rhynchosauroides ispp., and Synaptichnium isp. They can be referred to lepidosauromorph/ archosauromorph, basal archosaur, and dinosauromorph trackmakers. Apatopus, represented by 11 tracks of a more than 4 m long trackway, is recorded for the first time outside of North America and Europe. The assemblage concurs with the proposed Late Triassic age of the track-bearing beds by the occurrence of Apatopus, Atreipus-Grallator, and Eubrontes. If this is accepted, the stratigraphic range of Synaptichnium and Parachirotherium, hitherto known only from Early or Middle Triassic deposits, has to be extended to the Carnian-Norian. The occurrence of Eubrontes in the Irohalene Member (T5) provides further evidence for large theropods in pre-Jurassic strata. All assemblages are referred to the Scoyenia ichnofacies indicating continental environments with alternating wet and dry conditions.

The Tetrapod Ichnogenus Protochirotherium Fichter and Kunz 2004, a Characteristic Early Triassic Morphotype of Central Pangea

Early Triassic chirotherian footprint assemblages from Poland, Germany, and Morocco are important for understanding archosaur evolution in the aftermath of the Permian-Triassic crisis. However, their ichnotaxonomy is confusing because various authors have interpreted their diversity differently. After an analysis and ichnotaxonomic re-assessment, the presence of the ichnogenera Brachychirotherium, Isochirotherium, and Chirotherium in these assemblages is not supported. Distant similarities with these ichnotaxa are functions of extra morphological variation and substrate-related factors. Instead, Early Triassic chirotherian footprints described under these names are assigned here to the ichnogenus Protochirotherium and to a more slender morphotype identified as Synaptichnium. In particular, Protochirotherium appears to be more widely distributed in central Pangea as a characteristic morphotype reflecting a distinct stage in archosaur evolution. Trackmakers were nonarchosaurian archosauriforms or, alternatively, stem-group crocodylians. Morphologically and temporally these footprints match the hypothetical ancestor of the Chirotherium barthii trackmaker. Chirotherium barthii appears by the beginning of the Middle Triassic. Because of its restricted stratigraphic range, and its wider distribution in central Pangea, Protochirotherium also has biostratigraphic significance for this region and can be considered as an indicator of Early Triassic-aged strata.

First Occurrence of Tetrapod Footprints from Westphalian Strata of the Sidi Kassem Basin, Central Morocco

The Sidi Kassem Basin is the only limnic basin of Westphalian age in Morocco. It is built up of 1,250 m of alluvio-fluvial to lacustrine deposits that have so far yielded plant fossils and invertebrate remains only. Recent exploration for fossil tetrapod footprints in floodplain-deposits of the basin revealed a moderately diverse vertebrate ichnofauna composed of tracks assigned to cf. Batrachichnus Woodworth, 1900; cf. Hylopus Dawson, 1882; Dimetropus Romer and Price, 1940; and Notalacerta Butts, 1891. The tracks can be referred to temnospondyl, anamniote reptiliomorph, non-therapsid synapsid (pelycosaur), and captorhinomorph trackmakers. The described ichnoassemblage is important in at least three aspects: (1) It suggests an Early to mid-Pennsylvanian age for the footprint-bearing strata of the study area. (2) It is the oldest association of tetrapod footprints from Africa. (3) It is the first evidence of the relatively rare ichnogenera cf. Hylopus and Notalacerta outside of North America and Europe. Judged from the variety of tetrapod tracks and previously collected floral remains, the Sidi Kassem Basin must have represented a well-established continental ecosystem during Pennsylvanian time. Further exploration for trace and body fossils of Palaeozoic vertebrates in this basin may be important for the reconstruction of early tetrapod evolution.

Late Carboniferous Tetrapod Footprints from the Souss Basin, Western High Atlas Mountains, Morocco

ABSTRACT The Late Carboniferous Souss Basin of south-central Morocco exhibits an approximately 1,800 m thick succession of fluvial and lacustrine deposits that have yielded diverse fossil remains of plants, insects, conchostracans, ostracods, jellyfish, fishes, and few tetrapod footprints. Recent exploration of ichnofossils of the Souss Basin led to the discovery of several trampled surfaces including tetrapod footprints assigned to the plexus Batrachichnus (Woodworth, 1900). Limnopus (Marsh, 1894), Dimetropus (Romer and Price, 1940), and Ichniotherium (Pohlig, 1892). These footprints can be referred to temnospondyl, basal synapsid (“pelycosaurian”), and diadectomorph trackmakers. The moderately diverse tetrapod footprint assemblage from the Souss Basin is important because it is the second-oldest record of tetrapod footprints from Africa and only the second record of the well-known ichnogenus Ichniotherium from outside of North America and Europe. Based on the variety of tetrapod tracks and previously collected floral and insect remains, the Souss Basin must have represented a well-established continental ecosystem during the Late Carboniferous.

First International Congress on Continental Ichnology (ICCI), El Jadida, Morocco, April 21–25, 2015

Hafid Saber, Hendrik Klein, and Abdelouahed Lagnaoui a,c Laboratory of Geodynamic and Geomatic, Department of Geology, Faculty of Sciences, Choua€ıb Doukkali University, El Jadida, Morocco; Saurierwelt Pal€aontologisches Museum, Neumarkt, Germany; Laboratory of Stratigraphy of Oil-and-Gas Bearing Reservoirs, Department of Paleontology and Stratigraphy, Institute of Geology and Petroleum Technologies, Kazan (Volga Region) Federal University, Kazan, Russia

Unusual archosaur trackway and associated tetrapod ichnofauna from Irohalene member (Timezgadiouine formation, late Triassic, Carnian) of the Argana Basin, Western High Atlas, Morocco

ABSTRACT An archosaur trackway consisting of 10 successive pes imprints has been recovered from the Late Triassic Irohalene Member of the Timezgadiouine Formation (Argana Basin, Morocco). Footprints are tetradactyl-pentadactyl, and show an elongated ‘heel’, probably enclosing the trace of digit V. The trackway lacks a manus impression, which could be due to bipedal progression of the trackmaker or be a preservational phenomenon. It is assigned to cf. Parachirotherium isp. based on similarities with the ichnogenus that has been described from the same stratigraphic unit. Other footprints of this assemblage are assigned to the ichnotaxa Grallator-Eubrontes (dinosauromorphs/crocodilian-stem archosaurs), Brachychirotherium isp. (crocodilian-stem archosaurs), Rhynchosauroides isp. (lepidosauromorphs/archosauromorphs), and indeterminate ichnotaxa. Associated with the footprints are numerous invertebrate traces such as Scoyenia gracilis. The ichnofauna of the Irohalene Member supports a cosmopolitan distribution of pentadactyl but functionally tridactyl chirotheres (Parachirotherium) and grallatorids across the Ladinian-Carnian boundary. Similar assemblages are known from the Germanic Basin and from Eastern North America. Furthermore, it documents the occurrence of very large Eubrontes trackmakers in the early Carnian. The depositional environment of the lower part of the Irohalene Member is interpreted as wet red beds of a flood basin transitional to a distal braid plain.

MESOZOIC BASALTIC MAGMATISM OF THE SIDI SAÏD MAACHOU BASIN (WESTERN MESETA, MOROCCO): PETROGRAPHY, GEOCHEMISTRY AND GEODYNAMIC IMPLICATIONS.

Abdelkbir Hminna 1 , * Hafid Saber 2 and Abdelouahed Lagnaoui 2 . 1. Department of Geology, Sidi Mohamed Ben Abdellah University, Dhar El Mahraz, B.P. 1796, Fès-Atlas, 30003, Morocco. 2. Department of Geology, Chouaïb Doukkali University, Faculty of Sciences, B.P. 20, El Jadida, 24000, Morocco. ...................................................................................................................... Manuscript Info Abstract ......................... ........................................................................ Manuscript History