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Based on an exceptionally well-preserved articulated postcranial skeleton, the early amniote Cabarzia trostheidei gen. et sp. nov. is described. As the lack of the skull hampers its taxonomic assignment, a large sample of basal amniotes is included as part of an exhaustive comparison. Considering the slender, long-limbed proportions of the skeleton, several potential determinations are suggested in order to test for bolosaur, millerettid, araeoscelid, basal neodiapsid, or synapsid affinities. Numerous character conditions are re-evaluated regarding their distributions among early amniote subclades. The closest match to Cabarzia is found to be the middle Permian Mesenosaurus from Russia. The documentation for both genera provides the most complete postcranial descriptions of non-varanodontine varanopids. One of the main differences from Mesenosaurus is the curved ungual phalanx, indicating a use related to predatory behavior. An investigation of limb proportions, such as slender trunks, elongated hindlimbs, and relatively short forelimbs, may suggest the occurrence of facultative bipedalism in Mesenosaurus and Cabarzia. The oldest known mesenosaurine, C. trostheidei from the Asselian/Sakmarian, also pushes back the oldest evidence of bipedal locomotion by more than 15 Ma.

The early Permian Chemnitz Fossil Lagerstatte (Leukersdorf Formation, Chemnitz Basin, SE Germany) represents a diverse T 0 assemblage of a fossil forest ecosystem around the Sakmarian-Artinskian transition (290.6 ± 1.8 Ma), which was preserved by pyroclastic deposits of a multi-phased volcanic eruption. The multi-aged plant community consists of predominantly hygrophilous elements, which grew on an alluvial plain mineral substrate under sub-humid conditions, representing a wet spot environment. Strong seasonality triggered the formation of annual tree rings in arborescent woody plants, such as pycnoxylic gymnosperms, medullosan seed ferns and calamitaleans. From several hundred fossil trees, the 53 best-preserved specimens were selected and investigated in detail by measuring 2,081 tree rings in individual sequences of up to 77 rings. Ring sequences were analysed by standard dendrochronological methods to determine both annual growth rates and mean sensitivity. Morphological and statistical analyses on single tree rings reveal different tree-ring types according to the different plant groups. Pycnoxylic gymnosperms have distinct and regular tree rings, whereas medullosan seed ferns and calamitaleans show indistinct and regular tree rings as well as so called event rings. Results reveal differences between plant groups regarding their physiological reactions or adaptations to seasonal fluctuations. In comparison to pycnoxylic gymnosperms, both medullosan seed ferns and calamitaleans exhibit reduced growth rates and more sensitive reaction to environmental perturbances as water deficiency pointing to comparably lower adaptation to seasonally dry palaeoclimate. In this context, event rings are in many cases traced back to plant physiological stress during particularly severe drought periods. Altogether, these fossil trees serve as sensitive environmental archives, which shed light on growth conditions several decades back in time from the entombing eruption.

The Central European Basin is very suitable for high-resolution multistratigraphy of Late Permian to Early Triassic continental deposits. Here the well exposed continuous transition of the lithostratigraphic Zechstein and Buntsandstein Groups of Central Germany was studied for isotope-chemostratigraphy (δ13Corg, δ13Ccarb, δ18Ocarb), major and trace element geochemistry, magnetostratigraphy, palynology, and conchostracan biostratigraphy. The analysed material was obtained from both classical key sections (abandoned Nelben clay pit, Caaschwitz quarries, Thale railway cut, abandoned Heinebach clay pit) and a recent drill core section (Caaschwitz 6/2012) spanning the Permian-Triassic boundary. The Zechstein–Buntsandstein transition of Central Germany consists of a complex sedimentary facies comprising sabkha, playa lake, aeolian, and fluvial deposits of predominantly red-coloured siliciclastics and intercalations of lacustrine oolitic limestones. The new data on δ13Corg range from − 28.7 to − 21.7 ‰ showing multiple excursions. Most prominent negative shifts correlate with intercalations of oolites and grey-coloured clayey siltstones, while higher δ13Corg values correspond to an onset of palaeosol overprint. The δ13Ccarb values range from − 9.7 to − 1.3 ‰ with largest variations recorded in dolomitic nodules from the Zechstein Group. In contrast to sedimentary facies shifts across the Zechstein-Buntsandstein boundary, major element values used as a proxy (CIA, CIA*, CIA-K) for weathering conditions indicate climatic stability. Trace element data used for a geochemical characterization of the Late Permian to Early Triassic transition in Central Germany indicate a decrease in Rb contents at the Zechstein-Buntsandstein boundary.New palynological data obtained from the Caaschwitz quarry section reveal occurrences of Late Permian palynomorphs in the Lower Fulda Formation, while Early Triassic elements were recorded in the upper part of the Upper Fulda Formation. The present study confirms an onset of a normal-polarized magnetozone in the Upper Fulda Formation of the Caaschwitz quarry section supporting an interregional correlation of this crucial stratigraphic interval with the normal magnetic polarity of the basal Early Triassic known from marine sections in other regions. Based on a synthesis of the multistratigraphic data, the Permian-Triassic boundary is proposed to be placed in the lower part of the Upper Fulda Formation, which is biostratigraphically confirmed by the first occurrence date of the Early Triassic Euestheria gutta-Palaeolimnadiopsis vilujensis conchostracan fauna. Rare records of conchostracans reported from the siliciclastic deposits of the lower to middle Zechstein Group may point to its potential for further biostratigraphic subdivision of the Late Permian continental deposits.

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International audience; The Tajik basin archives the orogenic evolution of the Pamir hinterland. Stratigraphic‐sedimentologic observations from Cretaceous‐Pliocene strata along its eastern margin describe the depositional environment and basin‐formation stages in reaction to hinterland exhumation and basin inversion. During the Late Cretaceous‐Eocene (preorogenic stage: ~100–34 Ma), a shallow‐marine to terrestrial basin extended throughout Central Asia. An alluvial plain with influx of conglomerate bodies (Baljuvon Formation) indicates a first pulse of hinterland erosion and foreland‐basin formation in the late Oligocene‐early Miocene (synorogenic stage Ia: ~34–23 Ma). Further hinterland exhumation deposited massive alluvial conglomerates (Khingou Formation) in the early‐middle Miocene (synorogenic stage Ib: ~23–15 Ma). Westward thickening growth strata suggest transformation of the Tajik basin into the Tajik fold‐thrust belt in the middle‐late Miocene (synorogenic stage IIa: ~15–5 Ma). Increased water supply led to the formation of fluvial mega‐fans (Tavildara Formation). Latest Miocene‐Pliocene shortening constructed basin morphology that blocked sediment bypass into the central basin from the east (Karanak Formation), triggering drainage‐system reorganization from transverse to longitudinal sediment transport (synorogenic stage IIb: < ~5 Ma). Accelerated shortening (~27–20 Ma) and foreland‐directed collapse (~23–12 Ma) of Pamir‐plateau crust loaded the foreland and induced synorogenic stages Ia and Ib. Coupling of Indian and Asian cratonic lithospheres and onset of northward and westward delamination/rollback of Asian lithosphere (i.e., lithosphere of the Tajik basin) beneath the Pamir at ~12–11 Ma transformed the Tajik basin into the Tajik fold‐thrust belt (synorogenic stage IIa). The timing of the sedimentologically derived basin reconfiguration matches the thermochronologically derived onset of Tajik‐basin inversion at ~12 Ma.

The Early Pennsylvanian Xiaheyan entomofauna (Ningxia, China) is among the earliest assemblages of winged insects known so far, and therefore provides an essential input on deciphering the early diversification of this group. Despite its evolutionary significance, both the age and depositional environment remained poorly constrained. Here, we present high-resolution documentation of litho- and biofacies, biostratigraphy and geochronology of the up to 100 m thick type section. Accordingly, the insect-bearing strata are of latest Bashkirian (latest Duckmantian) to middle Moscovian (Bolsovian) age. The sediments, lithostratigraphically assigned to the Yanghugou Fm., represent a regressional marine sequence formed at the southern margin of the Qilian Inland Sea. Intercalations of crevasse channels and splays that laterally interlock with marine black shales and bioclastic limestones point to deposition in an interdelta bay. Ground-touching waves provoked the erosion of muds at shallow depths leading to mass mortalities among mollusc communities due to rising anoxia, turbidity and water turbulence. In addition, bottom currents transported mud intraclasts towards the basin, where sinking insects that reached the bay via winds and surface currents were buried sub-contemporaneously with bivalves below the storm wave base. Observed differences in insect preservation and assemblage composition across the sequence are found to correlate with lithology and are best explained by distance from land, and the action of events inducing insect carcasses to sink, such as storms. The proposed taphonomic model represents a new fossilization pathway from land to sea and provides new directions for prospecting Paleozoic deposits in the search for insects.

The end-Permian mass extinction marks the largest biotic crisis in the geologic record. The stratigraphic position of this boundary in continental deposits is still under discussion. In the present study, conchostracans from the Zechstein–Buntsandstein (Late Permian to Early Triassic) transition in central Germany have been taxonomically reinvestigated in order to better understand their utility for fine-scale biostratigraphy in continental Permian–Triassic boundary sections. The studied material was obtained from both collections and recent sampling activities in classical key sections and new outcrops of the Zechstein and Buntsandstein Groups. The sedimentary environments of the conchostracan occurrences are interpreted as lacustrine to fluvial facies depending on the paleogeographic position within the basin of the respective sections. The conchostracan fauna in the Zechstein–Buntsandstein transition consists of Euestheria gutta, Palaeolimnadiopsis vilujensis, Cornia germari, Estheriella marginostriata, Estheriella costata, Estheriella nodosocostata, Magniestheria mangaliensis, and Euestheria nordvikensis. Based on comparison with the Early Triassic conchostracan record in the Moscow syncline, the Palaeolimnadiopsis vilujensis–Euestheria gutta association in the upper part of the Fulda Formation indicates both its Early Triassic age and a position of the continental Permian–Triassic boundary within the Zechstein Group.

The Permian–Triassic boundary marks the greatest mass extinction in Earth's history. In order to understand the real causes of this severe extinction event, multidisciplinary investigations around the globe are required. Here, the terrestrial Permian–Triassic boundary sections in the Vladimir region, Central Russia, were sampled bed-by-bed for conchostracan study. In the Early Triassic intervals the following taxa were recognized for the first time: Cornia germari (Beyrich, 1857), Euestheria gutta (Lutkevitch, 1937), Magniestheria mangaliensis (Jones, 1862), Palaeolimnadiopsis vilujensis Varentsov, 1955, and Rossolimnadiopsis Novozhilov, 1958. The wide distribution of C. germari demonstrates their high value for biostratigraphy, since this species was also reported from the Lower Buntsandstein Subgroup in the Germanic Basin as well as from Early Triassic deposits in Hungary, Greenland and Siberia. The assumption of an Early Triassic age of the studied sections is also supported by associated Tupilakosaurus bone fragments, which point to the Tupilakosaurus wetlugensis Zone in the earliest Triassic.

Oribatid mites (Acari: Oribatida) are very diverse and important detritivorous and fungivorous micro-arthropods in modern forest ecosystems. Although the fossil record of oribatid mites can be traced to the Early Devonian, the paleoecology of oribatid mites during the deep geological past remains poorly understood. Remarkably good preservation of tunnel networks in a permineralized conifer wood specimen is described from the Early Permian of Germany. This fossil provides evidence for four aspects of oribatid mite feeding habits. First, there is preferred consumption of the more indurated tissues from growth-ring cycles. Second, tracheids were targeted for consumption. Third, feeding on tissues resulted in fecal pellet accumulations at the bottoms of tunnels. And fourth, the absence of feeding on ambient decomposing fungi such as necroses and rots, but rather the processing of pristine plant tissues, indicate the presence of a self-contained, microorganismic gut biota. These rather specialized feeding habits allowed oribatid mites a prominent role in the decomposition of digestively refractory plant tissues in Early Permian ecosystems.

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. Ju...

We present new palynological data from the Transantarctic Mountains that clarify the timing of sedimentary and magmatic processes in the transition from continental deposition of the Beacon Supergroup to emplacement of the Ferrar Large Igneous Province. Samples were collected from twenty-three Triassic and Jurassic sections in the southern area of north Victoria Land (NVL), East Antarctica. Recovered palynomorph assemblages are correlated with the widely used, although informal palynostratigraphic framework established for eastern Australia by Price. The associated Late Triassic–earliest Jurassic zone, APT5, is modified here with a proposed new subdivision: Lower APT5 (“APT5L”; middle–late Norian), Middle APT5 (“APT5M”; Rhaetian), and Upper APT5 (“APT5U”; Hettangian–earliest Sinemurian). We further propose a modification unifying the relevant formal eastern Australian and New Zealand palynostratigraphic zones, with a new Polycingulatisporites crenulatus Association Zone (new zonal status) that includes the P. crenulatus Association Subzone (new subzone; equivalent to APT5L) and the following Foveosporites moretonensis Association Subzone (new subzonal status; equivalent to APT5M). Our palynostratigraphic dating of the NVL assemblages demonstrates that the onset of sedimentation was diachronous in this part of the Transantarctic Basin, ranging from at least the Rhaetian to, in places, early Sinemurian. By lack of evidence for rocks containing APT5U assemblages and by analogy with the few coeval sections in Australia, we infer that the Hettangian interval in NVL is probably consumed by unconformity. Deposition of ashes from distal silicic volcanism commenced in the early Sinemurian and reached a peak phase beginning in middle Pliensbachian (ca 187 Ma), coinciding with the first major magmatic interval of the silicic Chon Aike Province in Patagonia and West Antarctica. Two major episodes of phreatomagmatic activity, driven by shallow-level sill intrusion into sandstone aquifers, occurred during the middle Pliensbachian and during the late Pliensbachian–early Toarcian. The latter episode was closely followed by the first pillow extrusion and local lava effusion. Contrary to some previous studies, we further conclude that all available palynological evidence is compatible with a short-lived emplacement of the plateau-forming Kirkpatrick Basalt at around 180 Ma during the early Toarcian.

Wei, D.D., Bethoux, O., Guo, Y.X., Schneider, J.W. & Ren D., 2013. New data on the singularly rare ‘cockroachoids’ from Xiaheyan (Pennsylvanian; Ningxia, China). Alcheringa 37, 547–557. ISSN 0311-5518.Additional material of stem-Dictyoptera, or ‘cockroachoids’, is described from the Early Pennsylvanian Xiaheyan locality (Zhongwei City, Ningxia Hui Autonomous Region, China). New specimens belonging to Qilianiblatta namurensis Zhang et al., 2012 are described, as are several specimens belonging to a new species (viz., Kinklidoblatta youhei sp. nov.). Three further specimens might represent additional species, to be better documented before they could be formally named. The occurrence of a ‘vannus’ in forewings of K. youhei sp. nov. is demonstrated, and discussed in the light of data on extant Dictyoptera. The low abundance and comparatively low diversity of the group in the locality is challenging, and might be a consequence of taphonomic bias.Dandan Wei [weidandanok@126.com], Yinxia Guo [kuaile422@163.com]...

The Dunkard Group of the Appalachian basin, Ohio–Pennsylvania–West Virginia, is a > 340 m thick sequence of primarily fluvial and lacustrine deposits that contain a diverse fossil record including a blattoid insect forewing identified as Sysciophlebia balteata (Scudder). This wing originates from the Cassville Shale in the Waynesburg Formation, at the very base of the Dunkard Group. S. balteata is a zone species of a Late Paleozoic spiloblattinid insect zonation that is calibrated by the co-occurrence of spiloblattinid zone species with conodont- and fusulinid-zone species in Kasimovian to Early Asselian mixed marine-continental profiles of New Mexico, USA, and the Donets basin in the Ukraine, as well as by the isotopic ages of volcanic intrusions into the Late Stephanian (Gzhelian) strata of the Saale basin in Germany. The base of the zone underlying the S. balteata Zone is defined by the FAD of Sysciophlebia ilfeldensis in the Late Gzhelian, and has an upper limit (LAD) of Early to Middle Asselian. An Early or Middle Asselian to earliest Sakmarian range of both the subzones of the S. balteata Zone can be inferred. Given that the mean duration of a spiloblattinid insect zone is about 1.5 to 2 Ma, the upper limit of the S. balteata Zone is Early Sakmarian. Hence, the base of the Dunkard Group should be maximally of Early Asselian or minimally of earliest Sakmarian age.

δ18OP values and 87Sr/86Sr ratios were determined on disarticulated xenacanthiform, hybodontid and ctenacanthid shark tooth material from several Early Permian (Sakmarian–Kungurian) continental bone beds of northern Texas and southern Oklahoma as well as from the marine Middle Permian (Roadian) of northern Arizona. The δ18OP values derived from the teeth of bone beds are in the range of 17.6–23.5‰ VSMOW, and are mostly depleted in 18O by 0.5–5‰ relative to proposed coeval marine δ18OP values. This indicates an adaptation to freshwater habitats on the Early Permian coastal plain by several sharks. Distinctly higher δ18OP values from two bone beds are attributed to significant evaporative enrichment in 18O in flood plain ponds. 87Sr/86Sr ratios of around 0.71077 are notably more radiogenic than 87Sr/86Sr of contemporaneous seawater. In contrast, the isotopic composition of teeth from the marine Kaibab Formation is characterised by low δ18OP values in the range of 13.4–15.6‰ VSMOW while 87Sr/86Sr ratios of a...

Plant remains with organic-matter preservation are uncommon in the Mississippian due to the general rareness of Mississippian terrestrial deposits. Moreover, the fossils are often strongly affected by post-sedimentary processes. However, bulk-macerated samples of the upper Visean of Chemnitz-Glosa (Germany) contain several well-preserved dispersed organic remains including cuticle fragments. Even though it is difficult to assign these remains to specific taxa and they probably represent only part of the vegetation, they provide important information on Mississippian floras. The cuticles are assigned to sphenopsids and gymnosperms. In addition, several bryophyte remains were found; nine morphologically different types are recognised, which represent at least five different taxa. This is remarkable because only a few bryophytes have been recorded from the Carboniferous to date.

Chondrichthyan egg capsules have been well known since the nineteenth century, although their systematic affinities have been controversial for much of this time. Currently, 10 chondrichthyan egg capsule morphotypes are distinguished but their phylogenetic signal and evolutionary traits have not yet been analysed. Here, we present an analysis of all extant and currently known fossil chondrichthyan egg capsule types, and use a purported placoderm egg capsule as an outgroup for character coding. The phylogenetic hypothesis based on discrete morphological characters indicates that the enigmatic Carboniferous egg capsule morphotypes Crookallia and Vetacapsula form a monophyletic group together with the egg capsule morphotype of chimaerid holocephalans. The elasmobranch egg capsule morphotypes are sister to the holocephalan types. Based on these results we conclude that the ancestral chondrichthyan egg capsule morphotype combined features of those found in callorhinchid holocephalans and placoderms. From this ...

The Sidi Said Maachou area in the Moroccan western Meseta preserves a succession, up to 400 m thick, of hitherto poorly studied continental Triassic deposits. Recent detailed geological mapping proposes a lithostratigraphic subdivision of the predominantly red-coloured siliciclastic deposits into three formations. Laminated mudstones and fine-grained sandstones in the upper part of the Oued Oum Er Rbiaa Formation have the most interesting fossil content including plant impressions, rhizoliths, fish scales, and invertebrate and vertebrate traces. These biogenic remains are partially associated with tool marks, microbially induced sedimentary structures, oscillation ripples, desiccation cracks, and halite pseudomorphs, suggesting sedimentation in a playa-like, fluvio-lacustrine system under semiarid conditions. All tetrapod footprints from these beds are assigned to Brachychirotherium parvum and indistinguishable from other occurrences of the ichnogenus in Central Europe and North America. Supposed trackmakers are archosaurs of the crocodile stem-group (Crurotarsi) that were widely spread over Triassic Pangaea. Because Brachychirotherium is only known from Late Triassic (Carnian–Rhaetian) deposits, the same age is attributed to the footprint horizon of the Oued Oum Er Rbiaa Formation. This is the first record of Brachychirotherium on the African continent and the first record of Triassic tetrapod footprints in Morocco outside of the High Atlas.

article i nfo Fossil shark remains occur in both marine and nonmarine Late Palaeozoic deposits, therefore their palaeo- ecology is controversial. The oxygen and strontium isotopic composition of biogenic fluorapatite in 179 teeth, scales and spines predominantly of hybodontid (Lissodus) and xenacanthiform (Orthacanthus, Xenacanthus, Bohemiacanthus, Triodus) sharks from various Late Carboniferous (Moscovian) to Early Permian (Artinskian) basins of Europe are used as ecological tracers to decipher diadromous or obligate freshwater lifestyle of the investigated taxa. The δ 18 OP values of the different shark teeth range from 11.7 to 20.2‰ within the different basins with mean values of 16.9±0.5‰ for the Bohemian Massif, 16.2±0.8‰ for eastern Germany, 18.2±1.0‰ for southwestern Germany, 18.5±0.7‰ for southern-central Spain, 17.6±0.4‰ for

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, alternati...

Die in der Stratigraphischen Tabelle von Deutschland 2016 (STD 2016) verwendete Zeitskala beruht mit kleinen Modifikationen auf den Komposit-Altern der Stratigraphischen Tabelle von Deutschland 2002 (STD 2002). Sie differiert von der ,,Geological Time Scale 2012" (GTS 2012). Die geochronologischen Einheiten zeigen auch die westeuropäischen Regionalstufen; die traditionelle deutsche Kulm-Gliederung wurde nicht mehr übernommen. Abfolgen des Mississippiums (unteres Karbon) sind in den deutschen Mittelgebirgen und im Untergrund Norddeutschlands weit verbreitet. Paralische Abfolgen des Pennsylvaniums (oberes Karbon) kommen übertage in der subvariszischen Saumsenke zwischen Aachen und Osnabrück sowie im Untergrund Norddeutschlands vor. Bis auf wenige Ausnahmen im Pennsylvanium einset- zende intramontane Abfolgen sind aus mehreren, zum Teil ausgedehnten Becken in Südwest- und Mitteldeutschland über- und untertage bekannt. Die vielgestaltige Fazies des deutschen Karbons wird von der nach Nordwesten progradierenden variszischen Orogenese und deren Ende im späten Westfalium gesteuert. Die Fazies reicht von Beckensedimenten und Flyschablagerungen sowie lateral anschließenden Plattformkalken im Mississippium (,,Kulm-Fazies" bzw. ,,Kohlenkalk- Fazies") zu paralischen und intramontanen, fast ausschließlich pennsylvanischen Molasse-Ablagerungen. Gegenüber der STD 2002 wurden die traditionellen Namen in den Standard-Profilen des Mississippiums von Aachen, Velbert und des Sauerlandes durch neue Formationsbezeichnungen ersetzt. Neu ist auch die Untergliederung des nordöstlichen und östlichen Rheinischen Schiefergebirges (Kellerwald, Lahn-Dill-Gebiet), wo durch Zirkon-Provenienz-Analysen rhenoherzynische und armorikanische Decken unterschieden wurden. Das Mississippium des Thüringisch-Vogtländischen Schiefergebirges und des Frankenwaldes wurde stärker differenziert und mit Formationsnamen belegt. Kleinere Änderungen gab es im Namurium der subvariszischen Saumsenke am NW-Rand des Rheinischen Schiefergebirges und in den intramontanen Pennsylvanium-Abfolgen der mitteldeutschen Becken. The Carboniferous timescale in the Stratigraphic Table of Germany 2016 (STD 2016) relies on slightly modified composite ages already used in the Stratigraphic Table of Germany 2002 (STD 2002). They differ from the Geological Time Scale 2012 (GTS 2012). Besides the international stages, the western European Mississippian regional stages are shown. The traditional German subdivision of the Kulm was discarded. Mississippian (lower Carboniferous) sedimentary successions are widespread in the mountainous regions and hills in the central part of Germany (Rhenish Mountains, Harz, Thuringian Forest, Franconian Forest and adjoining regions of Saxony). They are also widespread in the subsurface of Northern Germany. Paralic Pennsylvanian (upper Carboniferous) successions crop out in the Subvariscan Basin between Aachen and Osnabrück, and continue into the subsurface of Northern Germany. Intramontane successions, with few exceptions starting in the Pennsylvanian, occur in several, in part extended basins in southwestern and central Germany. They are known from outcrop and subsurface. The diversified facies of the Carboniferous in Germany is controlled by the northwestern progradation of the Variscan Orogeny and its finalisation during the late Westphalian. During the Mississippian, megafacies realms include deeper water basinal sediments and flysch deposits, and laterally adjoining shallow-water platform carbonates ("Kulm facies" and "Carboniferous Limestone facies", respectively). Locally starting in the later Mississippian (upper Viséan), paralic and purely continental intramontane molasse deposits prevail during the Pennsylvanian. The traditional lithostratigraphic terms of the Mississippian regional standard profiles of Aachen and the western and northern Rhenish Mountains (Velbert, Sauerland) were completely substituted by new formations. Lithostratigraphic terms of the northeastern and eastern Rhenish Mountains (Kellerwald, Lahn-Dill area) were completely revised, as provenance analyses of detrital zircons enabled the differentiation of Rhenohercynian and Armorican nappes. Also the Mississippian of the ThüringischFränkisches Schiefergebirge and the Frankenwald (Franconian Forest) are better differentiated; formations were introduced for most lithostratigraphic units. Minor modifications concern the Namurian of the Subvariscan Basin at the northwestern border of the Rhenish Mountains and the intramontane Pennsylvanian successions of several basins in central Germany.

A recently excavated locality in the Chemnitz Petrified Forest, lower Permian in age and occurring within the Leukersdorf Formation of the Chemnitz Basin, Germany, provides evidence for an outstanding fossil assemblage buried in situ by pyroclastics. The environment is interpreted as forested lowland that sheltered a dense hygrophilous vegetation of ferns, sphenophytes, and gymnosperms, as well as a diverse fauna of reptiles, amphibians, arthropods, and gastropods. A detailed measured section of the outcrop documents the early volcanic history of the Chemnitz fossil forest, including a paleosol that shows the root systems of Psaronius tree ferns, Arthropitys calamitaleans, and Medullosa and Cordaixylon gymnosperms in the same horizon. Fifty-three trunks are still standing upright and rooted at their place of growth, providing evidence that the top of the paleosol was the land surface on which the forest grew, thereby offering insights into the original plant community structure and density. Taphonomic analysis of both the petrified and adpression-fossil assemblages enable us to reconstruct the direction, estimate the violence and extent of the volcanic events, and their effects on the entire ecosystem. A complete dataset of three-dimensional coordinates resulting from three and one-half years of continuing excavation and study permits the recognition of organ connections and results in the first reconstructions of the excavation site, the floral elements, and the plant community as a whole.

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 Irohalene 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 ...

Bryophytes are found throughout the world in almost all terrestrial habitats. However, the fossil record of bryophytes is generally poor and fragmentary. In this contribution, we present the first unequivocal record of bryophytes from the Middle–Late Triassic Madygen Formation, in southwestern Kyrgyzstan, Central Asia. The bryophyte flora comprises two new species, the thalloid liverwort Ricciopsis ferganica and the leafy moss Muscites brickiae. In addition, a third form occurs, here described as Muscites sp. The dorsiventrally differentiated thallus of R. ferganica closely resembles that of the extant liverwort Riccia fluitans (Ricciaceae). The Central Asian finds represent the oldest known occurrence of the fossil genus Ricciopsis. The relative abundance of bryophytes in the Madygen fossil biota suggests that non-vascular plants played important role in special microhabitats within the Madygen palaeoecosystem and probably fostered the diversity of arthropods by providing favourable microhabitats for a range of aquatic and amphibiotic insects.

A spectacular new terrestrial Konzentratlagerstätte is introduced from the Turpan Basin of Xinjiang, China that probably belongs to the late Middle Jurassic Qigu Formation. It contains a mass accumulation of "xinjiangchelyid" turtles preliminarily identified as Annemys sp. In the zone with the highest turtle concentration, complete and articulated turtle skeletons are tightly packed at a density of up to 36 turtles per square meter. The fossiliferous layer is thickened here and shows an erosional base. This high concentration zone outcrops approximately 10 m in length and shows no decrease in turtle density after exposing 2 m of the layer into the hillside. Adjacent is a more expansive zone of at least 10 m by 30 m. In this region, the fossiliferous layer is evenly thick, and approximately five, fully disarticulated turtles are present per square meter. A conservatively estimated 1,800 turtles may, therefore, have been deposited at this site. It is likely that these aquatic turtles gathered in a retreating water hole in a riverine environment during a drought, much as some aquatic turtles will do today, but perished when the habitat dried up completely. A following catastrophic rainfall event caused a debris flow, possibly channelized in a dry river bed, which transported complete turtles, disarticulated turtles, and mudstone clasts and deposited them after a short distance. This taphonomic model is consistent with previous environmental reconstructions of the Turpan Basin during the late Middle Jurassic in predicting the episodic breakdown of regional monsoonal circulation resulting in a seasonally dry climate with severe episodic droughts.

The oxygen and strontium isotopic composition of 125 teeth of the hybodont sharks Lissodus and Hybodus from various late Triassic (Rhaetian) bone bed localities in the Central European Basin are used as environmental and ecological tracers for the shallow epicontinental Rhaetian Sea. The preservation of tooth enameloid was ascertained by cathodoluminescence microscopy. Mean δ18OP values differ regionally across the Rhaetian Sea with values of 18.6 ± 0.6‰ near the western gate, 16.1 ± 0.8‰ in the east, 15.4 ± 0.6‰ in the southeast, and 17.7 ± 0.9‰ in the south. The δ18OP values in the eastern Rhaetian Sea are 4–5‰ lower relative to Tethyan seawater, while teeth from the southern and western parts close to the marine gates are less fractionated. The strontium isotopic composition of teeth from most localities is more radiogenic in comparison to late Triassic seawater. The observed spatial δ18OP trend in the Rhaetian Sea indicates a shift from marine to brackish conditions in the south and west towards extensively brackish conditions with salinities of less than 16‰ in the east, thus confirming the existence of two marine gates. Brackish conditions persisted throughout early Rhaetian times, and were maintained by increased fluvial discharge from the Vindelician–Bohemian and Fennoscandian Highs as a result of enhanced atmospheric moisture availability in the course of the Rhaetian transgression. The δ18OP values of Lissodus and Hybodus indicate euryhaline behaviour of Rhaetian hybodonts by exceeding modern shark intraspecific variability. The lack of accordance in δ18OP values between different isotopically conspicuous localities in the Rhaetian Sea contradicts a basin-wide migration pattern. Differences between co-site taxa might indicate some degree of niche partitioning.

The Permian Ikakern Formation of the Argana Basin yielded important tetrapod skeletal remains but is still poorly known for any other kind of fossils. Here we present a moderately diverse assemblage of invertebrate and vertebrate traces from red beds in the upper part of the formation (Tourbihine Member, T2) near Timezgadiouine. Recorded ichnotaxa include Spongeliomorpha carlsbergi ( Bromley and Asgaard, 1979 ), Striatichnium natalis Walter, 1982 , Amphisauropus Haubold, 1970 , Hyloidichnus Gilmore, 1927 , Erpetopus Moodie, 1929 , and Dromopus Marsh, 1894 . The traces occur in laminated muddy siltstone and fine-grained sandstone that we interpret as slack-water deposits in an episodically high energy fluvial setting. Including arthropod burrows, arthropod grazing traces and tetrapod footprints, the ichnofossil assemblage corresponds to the Scoyenia ichnofacies. The described trace fossils provide evidence for various invertebrates and vertebrates hitherto unknown from the body fossil record of the study area. By extending the stratigraphic and geographic range of some of the recorded ichnotaxa, our finds strongly emphasise the importance of the Argana Basin for the reconstruction and understanding of Late Palaeozoic terrestrial ecosystems.

The holotype of Phylobktta gaudryi (Agnus, 1903), provided from the famous deposit of Commentry (Allier, France), exhibits a pair of forewings and a hind wing in connection, well preserved. Forewings show a network of veinlets in the area between M and CuA, and evidence of a fusion of some branches of M with CuA. It is concluded that the m-cua veinlets belong to M and that a reticulated fusion between M and CuA occurs. This fusion is considered as a homologue of the “dictyopteran forewing arculus”, but as an intermediate polymorphic state between “occurrence of a single discrete arculus“ and “no arculus”. The preserved hind wing is devoid of an m-cua arculus, has a multi-branched AA vein on the remigium, and has a vannus of limited extent. The new data on P. gaudryi might allow the phylogenetic position of Pennsylvanian stem-dictyopterans with respect to crown-dictyopterans to be better assessed.

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 λ = 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.

Habitat partitioning and site fidelity of spawning grounds are well-documented phenomena in extant selachians, but little is known about the reproductive strategies of their fossil relatives. Here we describe the selachian fauna of the Middle to Late Triassic Madygen Formation in southwestern Kyrgyzstan, Central Asia, based on several dozen tooth crowns and egg capsules. The material is assigned to three new taxa: Lonchidion ferganensis, sp. nov., and Palaeoxyris alterna, sp. nov., being teeth and egg capsules of hybodontid sharks, and Fayolia sharovi, sp. nov., being egg capsules of probable xenacanthids. Teeth of L. ferganensis, sp. nov., were almost exclusively found in pelecypod-rich shallow lacustrine mudstones and belong to juvenile individuals. Oxygen and strontium isotope data of tooth enameloid indicate freshwater conditions of the ambient water at the time of tooth mineralization. The egg capsules are common findings in near-shore lake deposits as well. Considering the mass co-occurrenc...

In terms of cumulative thickness and areal extent, the Khenifra Basin is one of the most important outcrops of Late Palaeozoic red-beds in central Morocco. Macro- and microfloral remains near the centre of the 1800 m-thick succession of interbedded conglomerates, sandstones, and mudstones are considered to be of middle to late Early Permian age. Here we give the first comprehensive analysis of the vertebrate ichnofossil record from the study area, based on 17 specimens of isolated footprints and incomplete step cycles collected at three localities that are lithostratigraphically equivalent to the plant-bearing horizons. The tetrapod ichnofauna comprises tracks of the plexus Batrachichnus Woodworth – Limnopus Marsh, Ichniotherium sphaerodactylum (Pabst), Dimetropus Romer and Price, and Dromopus Marsh which can be referred to temnospondyl, diadectomorph, synapsid (“pelycosaurian”) and early sauropsid trackmakers. This clearly Euramerican footprint assemblage, including the first occurrences of Ichniotherium and Dimetropus from outside Europe and North America, indicates a Late Carboniferous to Early Permian age of the fossiliferous strata. Judging from the relatively diverse ichnofauna and flora, the Khenifra Basin must have represented a well-established terrestrial ecosystem during that period. Its habitat could be specially important for the understanding of the phylogeny and dispersal of early tetrapods, inasmuch as we are able to report on an extremely rare type of diadectomorph footprint hitherto known only from the Early Permian of central Germany.

Footprints of Early Mesozoic terrestrial tetrapods are inadequately known from NW Africa despite pervasive and well-exposed continental strata of corresponding age in that region. Here, we report on the first occurrence of a tetrapod ichnofauna from the middle part of the Triassic Timezgadiouine Formation in the Argana Basin, Central Morocco that is outstanding in terms of abundance, preservation, and diversity. Footprints of this assemblage are assigned to the ichnotaxa Chirotherium barthii, Isochirotherium coureli, Synaptichnium isp., Atreipus–Grallator, Rotodactylus isp., Rhynchosauroides isp., and Procolophonichnium isp., suggesting early archosaurian, dinosauromorph, lepidosauromorph and possible therapsid or procolophonoid trackmakers. Based on comparison with the abundant record of Early Mesozoic vertebrate ichnofossils from Europe and North America, the ichnofauna presented here indicates a Middle Triassic (Anisian–Ladinian) age for the track-bearing horizon in the Argana Basin. Its clearly Euramerican affinity sheds light on the migration and dispersal of early archosaur-dominated tetrapod faunas considering that several ichnotaxa of the assemblage are documented from Africa for the first time.

The Rotliegend saline lake periodically covered wide areas of the Southern Permian Basin in Northwest Europe during the Permian. The sedimentology, mineralogy and geochemistry of lake deposits were studied to document very high frequency lake level fluctuations and to evaluate their triggers. Increased precipitation and marine ingressions into the basin resulted in lake extension. Increased run-off is documented by intercalated fluvial deposits in low-salinity deposits of the lake. Because lake deposits reflect mainly deposition in relatively wet climate phases, they are not correlatable to halite deposits in other basin areas. Decreased precipitation is followed by shrinkage of the lake, desiccation at its margins, and higher lake salinity due to concentration of the brine. Carbonate and anhydrite contents of lake deposits increased considerably before the areas fell dry, but halite is not preserved in the study area. Falling lake level is also reflected by the occurrence of wave ripples, reflecting decreased water depth and finally by desiccation and formation of evaporite crusts. Not only the mineralogical content, but also the colour of lake claystones changes with varying salinity. Anhydrite- and carbonate-free red claystones were replaced by violet to green and grey anhydritic and calcareous claystone or marls with increasing salinity. The amount of boron adsorbed on illite also corresponds to changes in salinity and can therefore be used as a palaeosalinity indicator. Rare earth element concentrations within lake deposits do not reflect variations in salinity but different palaeogeographical settings. Marine ingressions into the lake are neither reflected by the amount of boron adsorbed onto illite nor by rare earth element contents of the lake deposits. Highly saline deposits are partly characterised by intensive deformation, which can be interpreted as seismites or dissolution breccias.

During the mid-1980s, structural geological field work in the up to 300 m thick red-bed sequence of the Permian Tiddas Basin, central Morocco, revealed the first occurrence of Palaeozoic tetrapod footprints in NW Africa. Preservational ambiguity of part of the few published trace fossils and resulting taxonomic inconsistency necessitate a long-needed reanalysis of the type locality. Based on 110 recently collected specimens the revised tetrapod ichnofauna of the study area comprises tracks of Batrachichnus Woodworth, 1900 — Limnopus Marsh, 1894, Amphisauropus Haubold, 1970, Dimetropus Romer and Price, 1940, Varanopus Moodie, 1929 — Hyloidichnus Gilmore, 1927, and Dromopus Marsh, 1894 which can be referred to small and medium-sized temnospondyl, seymouriamorph, basal synapsid (‘pelycosaur’), captorhinomorph, and araeoscelid trackmakers. This assemblage is interpreted to represent a late Early Permian (Artinskian–Kungurian) tetrapod ichnofauna. Due to the relative abundance of footprints assigned to semiaquatic tetrapods and sedimentological evidence for pronounced humid palaeoclimatic conditions we propose correlation of the track-bearing strata with the late Artinskian/early Kungurian wet phase. Regarding the quality, quantity, and peculiar diversity of its fossils, including the earliest record of seymouriamorph and advanced captorhinomorph tracks from the African continent, the Tiddas Basin could be of great importance for the reconstruction of late Early Permian terrestrial ecosystems in proximity to the interface of Euramerican and Gondwanan biotic provinces.

Plant fossils recently collected from the Middle–Late Triassic Madygen Formation in SW Kyrgyzstan, Central Asia are described in detail. Based on the combination of gross morphology and epidermal/cuticular anatomy, four different taxa are identified, assigned to the Cycadales, i.e. Pseudoctenis lanei and Bennettitales, Pterophyllum pinnatifidum, P. firmifolium and Otozamites sp. All taxa are new for the Madygen Formation. Remarkable is the first occurrence of Pseudoctenis and two Pterophyllum species in the Triassic of Central Asia. The fossils are generally well preserved, but without cuticles. Nevertheless, epidermal and cuticular features can be recognized because the cuticles left behind imprints in the very fine-grained matrix. Silicon casts, examined under a stereomicroscope and with SEM, revealed xeromorphic features that are traditionally considered as indicators of arid conditions. However, comparisons of the functionality of the xeromorphic features in living plants show that these features occur in xerophytic, as well as in hygrophytic and halophytic plants. Sedimentological data from Madygen show that the fossil flora grew in alluvial plains, delta plains, and shallow lacustrine environments. The xeromorphic features are consequently interpreted as functional adaptations to environmental influences involved in the self-cleaning of the leaf surface, regulation of excessive radiation and leaf temperature and as an effective mechanical defense against phytophagous insects.

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 positio...

Tetrapod footprints from the Permian Ikakern Formation of the Argana Basin have long been known but never studied in detail. Here we present the first comprehensive description of recently discovered tracks and trackways that come from near the uppermost layers of an up to 1800 m thick red-bed sequence. Assigned to Hyloidichnus Gilmore, 1927 and Pachypes Leonardi et al., 1975 , these ichnofossils suggest a Middle to Late Permian age of the track-bearing strata. Potential trackmakers are medium to large-sized moradisaurine captorhinids and pareiasaurs. This strongly parallels the bodyfossil record inasmuch as the only known skeletal remains of terrestrial tetrapods from the study area belong to both of these groups. Based on this twofold evidence, captorhinids and Pareiasauria have to be considered as abundant faunal elements of the ancient floodplain environment that is represented by the upper part of the Ikakern Formation. As one of only a few places in the world with closely associated body and ichnofossils of Late Palaeozoic age, the Argana Basin has a high potential for allowing the reconstruction of terrestrial ecosystems during the early stages of the break-up of central Pangaea and just before Mesozoic diapsid reptiles gained ascendancy. Further research on this stimulating topic will significantly contribute to the understanding of still poorly known Middle and Late Permian tetrapod ichnofaunas.