Early Triassic Archosaur-Dominated Footprint Assemblage from the Argana Basin (Western High Atlas,Morocco)

An assemblage of abundant and well-preserved tetrapod footprints has been discovered in the Tanamert Member (T3) of the Triassic Timezgadiouine Formation (Argana basin, western High Atlas, Morocco). It is the first fossil record from T3. Surfaces from different localities show a uniform tetrapod ichnofauna that consists of chirotherian and small lacertoid forms. The chirotherians are assigned to the plexus Protochirotherium—Synaptichnium, their trackmakers interpreted as basal archosaurs. The lacertoid imprints show close affinities with Rhynchosauroides and may reflect archosauromorphs or lepidosauromorphs. Protochirotherium—Synaptichnium assemblages are characteristic of the Early Triassic and were known previously only from units of this age in central Europe. Biostratigraphically, the European record implies a wide-spread pre-Anisian Protochirotherium—Synaptichnium dominated assemblage preceding the first appearance of Chirotherium barthii near the Olenekian-Anisian boundary. The stratigraphic position of T3 between Late Permian (uppermost T2) and Middle Triassic (T4) and the European correlatives suggest an Early Triassic age of this unit. It is the first record of Early Triassic continental deposits in Morocco. The surfaces from T3 open up perspectives for further contributions to ecology, biogeography and locomotion of early archosaurs. Furthermore, excellent outcrops and quality of footprint preservation in the Argana basin offer a potential for clarification of ichnotaxonomic and biostratigraphic issues.     

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.     

Olenekian (Early Triassic) bivalves from the Salt Range and Surghar Range,Pakistan

Abstract: Based on newly collected material from the uppermost Smithian and lower to middle Spathian (Olenekian, Lower Triassic) of the Salt Range and Surghar Range (Pakistan), 15 bivalve species belonging to 11 genera are described, including two new genera, Eobuchia and Dimorphoconcha, and one new species, Palaeoneilo? fortistriata. Eobuchia gen. nov. is placed in a new subfamily, the Eobuchiinae, which differs from the Buchiinae in having an almost planar and only moderately inclined or offset right anterior auricle. Inclination of the right anterior auricle is proposed as a synapomorphy of the revised suborder Monotidina, which includes the Buchiidae, Monotidae, Oxytomidae and, tentatively, the Dolponellidae. The Pseudomonotidae, Chaenocardiidae and Claraiidae are discussed as candidate ancestors of the Monotidina. Dimorphoconcha gen. nov., provisionally placed in the Limidae, is a morphologically unusual genus characterized by a globose shell centre and a strongly plicate fringe. Permophorus costatus, which was previously known exclusively from Permian strata, is reported from the Spathian of the Surghar Range. This record extends the range of P. costatus for at least 8 Myr and makes it the first reported Lazarus species, with an outage of more than 2 Myr after the end‐Permian mass extinction. Ten of 15 species recognized in this study have not been reported from other regions, which may indicate increasing provincialism towards the end of the Early Triassic, or, alternatively, reflect the still insufficient knowledge of benthic faunas from the epoch that followed the greatest crisis in the history of life.     

Phylogenetic relationships of Early Triassic ammonoids (new data on the inner shell structure of some Olenekian ammonoids of Southern Primorye)

The inner shell structure for several ammonoid taxa (Inyoceras singularis, Anasibirites simanenkoi, A. nevolini, Yvesgalleticeras proximus, and Koninckitoides popovi) from the Olenekian of Southern Primorye is studied. The family-group level taxonomy of ammonoids is discussed.     

A new Late Olenekian (Triassic) ammonoid of low palaeolatitude affinity from Arctic Asia (Eastern Taimyr)

The new exotic ammonoid genusByrrangoceras is described from the Upper Olenekian of the Boreal Triassic (Arctic Siberia). The phylogenetic affinities and palaeobiogeographic distribution of the new genus are discussed.     

The oldest durophagous teleosauroid (Crocodylomorpha,Thalattosuchia) from the lower Bathonian of central High Atlas,Morocco

Marine crocodylomorphs were particularly abundant in Europe during the Middle Jurassic, but were very scarce in Africa. New finds of thalattosuchian cranial remains in Morocco suggest that this scarcity is probably related to poor sampling rather than original diversity. These remains pertain to the coastal thalattosuchians, the teleosauroids, and particularly to the clade grouping the blunt‐toothed ‘Steneosaurus’ obtusidens and the genus Machimosaurus. A new tribe is erected grouping these two taxa: Machimosaurini. Until now the machimosaurins have been known from the middle Callovian. The new material extends the presence of this group further back to the lower Bathonian, nearly 5 myr earlier. The machimosaurins are the only teleosauroid group that has been recently reviewed, and the difference between the revised diversity provided herein and that previously reported is large. A review of other teleosauroids and clear establishment of their phylogenetic relationships are also likely to have considerable impact on their observed diversity. So, until a complete review of the teleosauroids is carried out, the results of crocodylomorph diversity analyses should be treated with caution.     

Tempestitic shell beds formed by a new serpulid polychaete from the Bajocian (Middle Jurassic) of the Central High Atlas (Morocco)

Serpulid-dominated shell beds from the Lower Bajocian (Middle Jurassic) of the Central High Atlas (Morocco) are primarily formed by quadrangular tubes of the serpulid polychaete Nogrobs moroccensis sp. nov. The transverse tube ornament consisting predominately of wrinkles, the lack of narrow spirals and the shortness or absence of attached posterior tube portion represent the diagnostic features of this new species. Tubes of Nogrobs moroccensis co-occur with gastropods and oyster–crinoidal debris. We suggest that these serpulids were free-lying in their adult stage and formed benthic meadows on soft-bottom habitats. They frequently show parallel alignment in plane views in both graded and non-graded, few-cm-thick shell beds, corresponding to distal tempestites deposited close to and below storm wave base in an outer-ramp environment. The excellent preservation of serpulids implies short residence time on the seafloor, and the lack of compositional mixing with other communities and their restriction to outer-ramp habitats indicate that they represent parautochthonous assemblages. Analyses of serpulid orientation on the bottom bedding planes of shell beds revealed weak unidirectional arrangement of tubes, whereas orientation on the top bedding planes revealed strong unidirectional arrangement of tubes, with significantly different orientation relative to the bottom planes. This change could suggest a shift from predominantly a current component of the combined storm flow (with uneven seafloor surface and adhesion of skeletal grains to muddy substrate) to significant oscillatory component of the waning storm, as also suggested by bidirectional orientation of gastropods.     

The Black-necked Grebe (Podiceps nigricollis): an expanding species in the Middle Atlas wetlands,Morocco

The present study investigates the population trends of Black-necked Grebe Podiceps nigricollis (Podicipedidae: Podicidae) over 5 years (2009–2013) in three Middle Atlas wetlands (Aguelmam Afennourir, Dayet Aoua and Dayet Ifrah). Using generalized linear models with a Poisson distribution, we demonstrated that the number of Black-necked Grebe breeding pairs has varied significantly and positively over the 5 years and between the three study wetlands. The annual population growth rate of the three colonies was 0.48 (±0.01). This positive trend is consistent with the current Least Concern conservation status of the IUCN Red List. Further more detailed studies are, however, needed to improve our understanding on the mechanisms driving the population increase in this part of North Africa. This remains a prerequisite for proper population conservation and management.     

The limnology of Lac ďIfni (High Atlas Mountains, Morocco), an unusually productive mountain lake

• 1 Lac ?Ifni (surface area 30 ha, z_max 60 m, altitude 2300 m) lies in a catchment comprised of Precambrian igneous rocks in the High Atlas Mountains of Morocco. Previously described as a typical ohgotrophic lake, it was also reported to have a warm layer at the base of the hypolimnion which was ascribed to phreatic water supply and drainage.
• 2 Visits to the lake made in May/June and September 1990 showed strong thermal stratification but no hypolimnial warm layer. Drainage from the lake is subterranean, through a natural rubble dam, and the water level dropped by at least 7m through the summer. The flushing rate is about 0.6 year^?1.
• 3 The upper part of the metalimnion and the lower epilimnion were strongly supersaturated with oxygen in June, and there was a pH gradient from 7 to 10.5 between the hypolimnion and the epilimnion. These features were less marked in September when planktonic gross primary productivity was estimated to be about 145mgCm^?2 h^?1. In September the hypolimnion was 38% saturated with oxygen; the areal hypolimnetic oxygen deficit between June and September was 0.073mg O₂ cm^?2 day^?1. Total dissolved phosphate concentrations were 7–14 μgl^?1.
• 4 The open water faunal assemblage consisted of Brachionus calyciflorus and Filinia lotigiseta (Rotifera), Cyclops abyssorum (Copepoda), and stunted Salmo trutta (Pisces). Trout diets were comprised principally of adult copepods and copepodites, which showed diurnal vertical migration, and of algal material apparently scraped from rock surfaces. The dry biomass density of Cyclops was at least 2gm^?2 in June and at least 4.1 gm ^?2 in September.
• 5 Tubifex tubifex (Oligochaeta) dominated the benthos below depths of 40m. Mean dry biomass density was 2.8gm^?2, but ranged from 0.6–8.1 gm^?2 between samples. About 50% of the lake bottom is below 40m depth.
• 6 The persistent oxygen supersaruration of the euphoric zone, the field estimate of primary productivity, the hypolimnetic oxygen deficit, the biomass of zooplankton and the benthic biomass together indicate that Lac ?Ifni is an unusually productive mountain lake. This high level of productivity may be sustained by nutrient addition through contamination by dust originating outside the catchment, aided by efficient nutrient recycling in the euphoric zone.

     

Dispersal traits and dispersal patterns in an oro-Mediterranean thorn cushion plant formation of the eastern High Atlas, Morocco

This study describes the dispersal traits and dispersal patterns of 51 perennial plant species belonging to 19 families in an oro-Mediterranean thorn cushion plant formation on the High Atlas, Morocco. Diaspore type, mass, number, antitelechoric mechanisms and dispersal time were studied with respect to growth forms, dispersal modes and spatial dispersal. Species spanned 10⁵ range of diaspore mass, which coincided with those found in other high mountain regions. Diaspore mass was significantly higher in trees and shrubs than in semi-shrubs and perennial herbs. Barochorous and zoochorous species are more likely to have heavy diaspores, whereas anemochorous and ballistic species have a medium diaspore mass and semachorous and ombro-hydrochorous species have low diaspore mass. Diaspore number was significantly higher in trees and shrubs than in semi-shrubs and perennial herbs. The barochorous, ombro-hydrochorous and zoochorous species tended to produce higher diaspore numbers than species with other dispersal modes. Bradyspory was well-developed by trees and large shrubs dispersed by biotic vectors. Synaptospermy was represented by its long dispersal component. Myxospermy was significantly associated with semi-shrubs and perennial herbs with restricted spatial dispersal. It seems that ombro-hydrochory combined with myxospermy and a high number of light seeds is an efficient mechanism that ensures successful establishment of the most typical and endemic thorn cushion plant species, such as Alyssum, Vella and Ormenis. In our study area, the highest dispersal availability was synchronized with the dry summer season (July–August) and the beginning of the rainy months (September). The dispersal peak for the wind-dispersed species, which is the most effective primary dispersal mode, occurs during the dry season, while dispersal for the biotic-dispersed species takes place throughout the year.     

Additional thylacocephalans (Arthropoda) from the Lower Triassic (upper Olenekian) Osawa Formation of the South Kitakami Belt,Northeast Japan

The Lower Triassic Osawa Formation in the South Kitakami Belt, Northeast Japan, consisting mostly of mudstone of shallow-marine environment, was deposited during the late Olenekian (ca. 250 Ma), and is an important unit through which to examine the biotic recovery process after the end-Permian mass extinction. The Osawa Formation is the only unit in Japan that yields thylacocephalans (Arthropoda). Three species belonging to three genera have been reported before: Ankitokazocaris bandoi, Kitakamicaris utatsuensis and Ostenocaris sp. In addition to the known species, some thylacocephalans, including one new genus and three new species, are described in the present paper: Ankitokazocaris tatensis n. sp., Concavicaris parva n. sp., Miyagicaris costata n. gen. n. sp. and Ostenocaris? sp. Although Thylacocephala have a rather long stratigraphic range (from Silurian to Cretaceous) and are known from a wide geographical region, there are only about thirty genera in this group. The Osawa thylacocephalan fauna comprises at least five genera, making it one of the most diverse in the world at the generic level. During the Triassic Period, the Thylacocephala diversified and spread widely throughout low-latitude regions.     

Ammonoid age control of the Early Triassic marine reptiles from Chaohu (South China)

Here we described a series of ammonoid specimens from the Early Triassic of Chaohu, South China and recognized the occurrence of Procolumbites for the first time in this area. The Procolumbites layer is about one meter above the first appearance of Chaohusaurus, indicating that the oldest Chaohusaurus is within the Procolumbites Zone of middle Spathian age. This new age constrain is significantly older than the previously suggested Subcolumbites Zone assignment (early late Spathian). To date, Chaohusaurus is the oldest known ichthyopterygian.     

New eocene oysters and the final regression at the southern Rim of the central High Atlas (Morocco)

The oysters Ostrea (Ostrea) todraensis n. sp. and Ostrea (Ostrea) sidialiensis n. sp. are described from the uppermost marine beds south of the central High Atlas in two sections west of Tinerhir. The oysters and the codiacean alga Ovulites margaritula (Lamarck, 1801) Lamarck, 1816 indicate a final regression at the end of Middle Eocene times, i.e. earliest at the end of the Lutetian and latest at the end of the Bartonian. The oysters lived in muddy lagoonal environments sealed off from the sea by oolitic-bioclastic carbonate sand shoals and bordered in landward direction by sebkha-like redbed deposits. Taxonomy and systematic affinities of the oysters are discussed. Their shell structures and depositional history are described in respect to sedimentary infillings. Some selected aspects of biometry explain differences in shell morphometry both between different valves of the same species and equivalent valves from the different populations, illustrating the importance of size and location of the adductor muscle scar.     

Toarcian carbon isotope shifts and nutrient changes from the Northern margin of Gondwana (High Atlas, Morocco, Jurassic): Palaeoenvironmental implications

The Early Toarcian is marked by a global perturbation of the carbon cycle and major marine biological changes. These coincide with a general decrease in calcium carbonate production and an increase in organic carbon burial, and culminate in the so-called Toarcian Oceanic Anoxic Event. It is believed that the environmental crisis was triggered by the activity of the Karoo-Ferrar large igneous province. In order to further document the Early Toarcian palaeoenvironmental perturbations, carbon isotope, total organic matter, calcareous nannofossils and phosphorus content of the Amellago section in the High Atlas rift basin of Morocco were investigated. This section is extremely expanded compared to the well-studied European sections. Its position along the northern margin of the Gondwana continent is of critical importance because it enables an assessment of changes of river nutrient input into the western Tethyan realm. The carbon isotope curve shows two negative excursions of equal thickness and amplitude, at the Pliensbachian-Toarcian boundary and at the transition from the Polymorphum to the Levisoni Zone. This confirms the supra-regional nature of these shifts and highlights the possible condensation of the first “boundary” shift in European sections. Phosphorus content is used to trace palaeo-nutrient changes and shows that the two negative carbon isotope shifts are associated with increased nutrient levels, confirming that these episodes are related to enhanced continental weathering, probably due to elevated greenhouse gases in the atmosphere. In the High Atlas Basin, the increase in nutrient levels at the Pliensbachian-Toarcian boundary is moreover likely to be the main factor responsible for the coeval demise of the Saharan carbonate platform. A middle Toarcian event, centered on the boundary between the Bifrons and Gradata Zones, characterized by a positive carbon isotope excursion and nutrient level rise, is documented in the Amellago section.     

Palaeoecological significance of a new Griesbachian (Early Triassic) gastropod assemblage from Oman

A new Early Triassic (Griesbachian) gastropod fauna from the Al Jil Formation of Oman is described. Early Triassic faunas from elsewhere (e.g. the Italian Dolomites and the western USA) are typically of low diversity and high dominance, usually attributed to environmental stress in the immediate aftermath of the end-Permian mass extinction event. The new Oman fauna has, by contrast, relatively high diversity, low dominance and a more even spread of individuals between taxa. It is the most diverse Griesbachian fauna known to date. This is attributed to the favourable (i.e. well-oxygenated) conditions under which the fauna lived. This uncharacteristic Griesbachian gastropod fauna demonstrates that, in the absence of oceanic anoxia, biotic recovery after the end-Permian extinction event may occur surprisingly quickly (within one conodont zone). The fauna is also partially silicified, which has increased its preservation potential relative to other Griesbachian gastropod assemblages. However, only one reappearing Lazarus genus is present in the Oman fauna. This suggests that there was some other control on the abundance of Lazarus genera at this time, other than the absence of silicified faunas as previously suggested.