Magnetostratigraphy of the Sherwood Sandstone Group (Lower and Middle Triassic), south Devon, UK: detailed correlation of the marine and non-marine Anisian

The Sherwood Sandstone Group in south Devon consists of the Budleigh Salterton Pebble Beds and the Otter Sandstone Formation. The Otter Sandstone Formation comprises four lithostratigraphic sub-units, from the base labelled A-D, and contains tetrapods corresponding to the Perovkan land vertebrate faunachron, and is consequently a key European non-marine Anisian succession. A total of 181 palaeomagnetic specimens from 92 horizons were measured to determine the palaeomagnetic signal. The magnetic properties and characteristic remanence is predominantly carried by haematite, most of which probably formed early in the depositional and diagenetic history. The lower part of the Sherwood Sandstone Group (Budleigh Salterton Pebble Beds and Otter Sandstone Formation, unit A, lower part of unit B) has predominantly normal polarity (magnetozones BS1, BS2 and BS3n) with three thin reversed polarity magnetozones. The lower part of unit B to the lower part of unit C of the Otter Sandstone Formation is predominantly reverse polarity (magnetozones BS3r, BS4 and BS5), but includes three short normal polarity intervals. The upper ca. 50 m of unit C shows a mix of normal and reverse polarity (BS6, BS7) with normal dominating. The ca. 15-m-thick unit D is entirely of normal polarity (BS8n). The magnetostratigraphy allows unambiguous comparison with conodont and ammonite calibrated magnetostratigraphies from elsewhere in Europe. These show that the BS2 and BS3n magnetozones of the Otter Sandstone Formation are lower Aegean to middle Pelsonian (lower to middle Anisian). The BS3r to BS5r magnetozone interval, is middle Pelsonian to lower Illyrian (middle to upper Anisian), and the BS6 to BS8n magnetozone interval is middle Illyrian to lower Fassanian. The top of unit D is correlated to an interval in the middle of the Nevadites Zone (close to the Anisian-Ladinian boundary). Consequently, the Otter Sandstone Formation correlates with the Muschelkalk in the Polish Basin, but is equivalent to only part of the Germanic Basin Muschelkalk. The continuous magnetostratigraphy from the Sherwood Sandstone Group, provides additional details of the polarity pattern through the Pelsonian to Illyrian interval, which appears to be lacking in discontinuous marine sections. A synthesis of adjacent European sedimentary basins in combination with the magnetostratigraphy, suggests that the Budleigh Salterton Pebble Beds are probably Late Spathian to lower Aegean.     

The Permian–Triassic transition and the onset of Mesozoic sedimentation at the northwestern peri-Tethyan domain scale: Palaeogeographic maps and geodynamic implications

The main aim of this paper is to review Middle Permian through Middle Triassic continental successions in European. Secondly, areas of Middle–Late Permian sedimentation, the Permian–Triassic Boundary (PTB) and the onset of Triassic sedimentation at the scale of the westernmost peri-Tethyan domain are defined in order to construct palaeogeographic maps of the area and to discuss the impact of tectonics, climate and sediment supply on the preservation of continental sediment.At the scale of the western European peri-Tethyan basins, the Upper Permian is characterised by a general progradational pattern from playa-lake or floodplain to fluvial environments. In the northern Variscan Belt domain, areas of sedimentation were either isolated or connected to the large basin, which was occupied by the Zechstein Sea. In the southern Variscan Belt, during the Late Permian, either isolated endoreic basins occurred, with palaeocurrent directions indicating local sources, or basins underwent erosion and/or there was no deposition. These basins were not connected with the Tethys Ocean, which could be explained by a high border formed by Corsica–Sardinia palaeorelief and even parts of the Kabilia microplate. The palaeoflora and sedimentary environments suggest warm and semi-arid climatic conditions.At the scale of the whole study area, an unconformity (more or less angular) is observed almost everywhere between deposits of the Upper Permian and Triassic, except in the central part of the Germanic Basin. The sedimentation gap is more developed in the southern area where, in some basins, Upper Permian sediment does not occur. The large sedimentary supply, erosion and/or lack of deposition during the Late Permian, as well as the variable palaeocurrent direction pattern between the Middle–Late Permian and the Early Triassic indicate a period of relief rejuvenation during the Late Permian. During the Induan, all the intra-belt basins were under erosion and sediment was only preserved in the extra-belt domains (the northern and extreme southern domains). In the northern domain (the central part of the Germanic Basin), sediment was preserved under the same climatic conditions as during the latest Permian, whereas in the extreme southern domain, it was probably preserved in the Tethys Ocean, implying a large amount of detrital components entering the marine waters. Mesozoic sedimentation began in the early Olenekian; the ephemeral fluvial systems indicate arid climatic conditions during this period. Three distinct areas of sedimentation occur: a northern and southern domain, separated by an intra-belt domain. The latter accumulated sediments during the Early–Middle Permian and experienced erosion and/or no-deposition conditions between the Middle–Late Permian and the beginning of Mesozoic sedimentation, dated as Anisian to Hettangian. At the top of the Lower Triassic, another tectonically induced, more or less angular unconformity is observed: the Hardegsen unconformity, which is dated as intra-Spathian and is especially found in the North European basins. This tectonic activity created new source areas and a new fluvial style, with marine influences at the distal part of the systems. During the Anisian and Ladinian, continental sedimentation was characterised by a retrogradational trend. In other words, the fluvial system evolved into fluvio-marine environments, attesting to a direct influence of the Tethys Ocean in the southern and northern domains. Both at the end of the Olenekian (Spathian) and during the Anisian, the presence of palaeosols, micro- and macrofloras indicate less arid conditions throughout this domain.     

Oysters and oyster-like bivalves from the Middle Triassic Muschelkalk of the Germanic Basin

Middle Triassic marine deposits of the Germanic Basin (Muschelkalk) record a significant proliferation of cementing bivalves from different families. Based on previously undescribed, excellently preserved material from the Willebadessen Member (late Anisian, Illyrian) of the Upper Muschelkalk Trochitenkalk Formation of Willebadessen (Germany), we propose the new genus Noetlingiconcha, type species N. speculostreum sp. nov., for strongly plicate prospondylids lacking auricles. The new genus differs from Terquemia and Enantiostreon in being plicate rather than costate, and from Newaagia in the absence of auricles. We demonstrate that N. speculostreum was invariably attached by its right valve, in contrast to an externally similar species from the Lower Muschelkalk Freudenstadt Formation (lower Anisian, Bithynian) that was exclusively cemented by its left valve and thus represents the geologically oldest known oyster species. Previous reports of amphi-pleurothetic cemented bivalve species from the Muschelkalk probably result from lumping together these two externally similar species. The constancy of sinistral attachment in the geologically oldest Ostreidae suggests that left-pleurothetic valve orientation was already established in the ancestry of this family. Palaeontological data are therefore in accordance with genetic and larval shell morphology analyses that identified Pterioidea as the sister taxon of Ostreoidea, because Pterioidea contains several Permian-Triassic genera with an anatomically lower left valve.     

New fossil insects from the Anisian (Lower to Middle Muschelkalk) from the Central European Basin (Germany and The Netherlands)

The Palaeozoic–Mesozoic transition is characterized not only by the most massive Phanerozoic mass extinction at the end of the Permian period, but also its extensive aftermath and a prolonged period of major biotal recovery during the succeeding Middle to Late Triassic. Particularly, Anisian insect species from units of the Lower to Middle Muschelkalk from the Central European Basin are rare. The specimens described here originated from the Anisian Wellenkalk facies (Lower Muschelkalk), Vossenveld Formation of the Winterswijk quarry, The Netherlands, and from the orbicularis Member (lowermost Middle Muschelkalk, Anisian) of Esperstedt near Querfurt (Saxony-Anhalt). Thus, the described insect remains from Winterwijk and Esperstedt expand our knowledge about Middle Triassic terrestrial arthropod communities and their palaeodiversity. A new species of Chauliodites (C. esperstedti sp. nov) is introduced.     

The first specimen of the Middle Triassic Phalarodon atavus (Ichthyosauria: Mixosauridae) from South China,showing postcranial anatomy and peri‐Tethyan distribution

Phalarodon atavus from the Germanic Muschelkalk Basin was previously represented only by cranial elements. Here we report a nearly complete and articulated specimen of P. atavus from the Middle Triassic Luoping Biota, Yunnan, South China. This is the first specimen of P. atavus from outside the Germanic Basin. This discovery demonstrates a peri‐Tethyan distribution of P. atavus. The new specimen is also the first one preserving the postcranial anatomy of this species, providing the opportunity to evaluate its sustained swimming ability. Inferences made on its functional morphology suggest that this species was probably adapted for active foraging. Tooth crown morphology suggests that P. atavus may have preferred externally soft prey.     

Palynology of the marine Middle Triassic in the SW Germanic Basin (Upper Muschelkalk, Luxembourg): Evidence for an important latest Anisian-early Ladinian sea-level lowstand

A palynological study is presented of the open-marine Upper Muschelkalk from the SW Germanic Basin, a succession dominated by lagoonal fine-grained carbonates and terrigenous mudstones. The abundance and taxonomic composition of the marine and terrigenous palynomorphs shows a phase of strong terrigenous input, which interrupted the open-marine evolution. This suggests an important regression during latest Anisian (late Illyrian) to early Ladinian (Fassanian) time, which is approximately coeval with a strong regression in other parts of this basin. The documented sea-level lowstand probably has a strong eustatic component, which was overprinted by the regional tectonics in the Germanic Basin.     

The oldest nautiloid recorded from the Triassic Germanic Basin and the first ammonoid from the Aegean (Middle Triassic) of Poland

During the Triassic in Europe, the Germanic Basin extended from England in the west to the eastern border of Poland in the east. Although cephalopods are common in some Middle Triassic (Muschelkalk) horizons, there still persists a gap in the palaeontological record of the eastern part of the Germanic Basin, notably in the lowermost parts of the Lower Muschelkalk, spanning the latest Olenekian-Aegean interval. The present contribution attempts to fill this gap by presenting the first ammonoid (ceratitid Beneckeia sp.) and nautiloids (Germanonautilus cf. dolomiticus and G. cf. salinarius) from the Lower Gogolin Beds (Upper Silesia, Poland). These Germanonautilus are the oldest nautiloids found in the entire Germanic Basin.     

Le Lias et le Dogger du sud du Bassin parisien (France) : rôle de la tectonique syn-sédimentaire et reconstitutions paléogéographiques à l'échelle de la biozone d'ammonite

The Lias and Dogger series in the southern Paris Basin outcrops are precisely dated following the ammonite biozonation scale. In these series, the depositional environment context is accurately identified and thus various isopic zones can bee recognized. From the outcrop study, the depositional sequence and the tectonics events are also distinguished. The poorly dated subsurface data should be not anymore interpreted as diagraphic signatures, with the traditional peak to peak correlation. They now need to be analysed according to the geodynamic evolution of each isopic zone. The Liassic transgressive series (in the Armorican continent) and the Dogger platforms numerous carbonate bodies are correlated at biostratigraphic scale, throughout the whole southern Paris Basin. Subsequently, paleogeographic maps are established and based on depositional sequences. The basin evolution image changes radically. This shows clearly that the complexes carbonate bodies layout prohibits any depositional environment interpretation and any isopach mapping at geological stage level. If not, the carbonate platforms of different age would be amalgamated, without any connection with the reality. Over the time, paleogeographic maps show syn-sedimentary short wavelength deformations, more or less well expressed according to the local sedimentary context. During the Lias and the Dogger, three steps can be distinguished in the southern Paris Basin. Hettangian to Lower Aalenian, the progressive increase in accommodation drives to total immersion of the continental areas; the maximum of accommodation during the Middle Toarcian entails uniform sedimentary conditions. However, short wavelength deformations persist. The major accidents defining the isopic zones are mainly North-South oriented. Middle Aalenian to Middle Bathonian, isopic zones are contrasted, controlled by the North-South accidents but also the North West-South East ones ; the variable accommodation allowed the development of regional sedimentary gaps and shallow limestone progradations on a marly area (so called “Sillon marneux”) ; tectonics events correspond to the Tethyan and the North Sea riftings influences. Upper Bathonian to Upper Oxfordian, the North West-South East tectonics accidents are particularly active; this tectonic phase could be associated to the thermal event, described in the Paris Basin and announcing the Malm depositional environment.     

Oligocene and Early Miocene gastropods from Kutch (NW India) document an early biogeographic switch from Western Tethys to Indo-Pacific

Shallow marine gastropod assemblages from Chattian, Aquitanian and Burdigalian sections in the Indian Kutch Basin are described. They provide insight into the composition and biogeographic relations of the gastropod assemblages at this junction between the Western Tethys and Proto-Indo-Pacific Ocean. For the first time, an improved biostratigraphy allows a clear separation of the assemblages, especially for the hitherto undifferentiated Early Miocene faunas. Throughout the Oligocene, about one-third of the species are also frequently found in the Western Tethys, documenting a passable Tethyan Seaway for nearshore molluscs. A considerable provincialism is evident as well. The expected turnover during the Early Miocene, due to the closing of the Tethyan Seaway, is reflected in the Miocene assemblages. Surprisingly, however, the cut appears very early, i.e. already during the Aquitanian, when the West–East interrelation drops to zero despite the passage having been open during this interval. In contrast, the Burdigalian assemblages witness a minor re-appearance of Western Tethys taxa, suggesting the re-establishment of rather ineffective migration pathways prior to the final closure of the Tethyan Seaway. Cerithium bermotiense and Lyria (Indolyria) maniyaraensis are introduced as new species.     

Middle Triassic paleomagnetic data from northern Bulgaria: constraints on Tethyan magnetostratigraphy and paleogeography

Magnetostratigraphic and biostratigraphic data are presented from the Anisian (Middle Triassic) Peri-Tethyan Edivetur section of northwestern Bulgaria. A dual-polarity component of magnetization carried by magnetite delineates a magnetic stratigraphy of mainly reversed polarity. Magnetozones are dated by means of foraminifer and conodont biostratigraphy. Data from Edivetur are compared with data from Middle Triassic Tethyan limestone sections with the aim of contributing to the completion of the Middle Triassic magnetic polarity time scale. We also propose that paleomagnetic data from Edivetur can be used as proxy data for the paleogeographic position of the Moesian platform. The Moesian platform was located at 21–24°N along the southern margin of Europe. It was probably marginally separated, but not detached or rotated away from Europe by the North Dobrugea transtensional trough, which is interpreted as a back-arc basin resulting from the northward subduction of the Neo-Tethys (Vardar) or Paleo-Tethys ocean. Paleomagnetic data from this study and other minor tectonic elements are used to generate a paleogeographic sketch map of the Pangea-bounded western Tethys and Peri-Tethys at Middle/early Late Triassic time.     

Millions of reptile tracks—Early to Middle Triassic carbonate tidal flat migration bridges of Central Europe—reptile immigration into the Germanic Basin

Discoveries of vertebrate track sites in Central Europe have occurred in 75 localities in carbonate tidal flats of the Middle Triassic. In the Germanic Basin carbonate tidal flats were wide-span mapped, resulting in the finding of millions of small- to medium-sized reptile tracks. In the west of the basin the sediment of the Lower Muschelkalk to basal Upper Muschelkalk contains at least 21 track horizons, whereas in the eastern part more typically marine conditions were present. Here, tidal flats with additional track beds started earlier during the Upper Bunter and demonstrated marine ingression from the eastern Silesian gate. During low stands these tidal flats comprised inter-peninsula bridges, which allowed migration of reptiles. Only two medium to small prolacertilian reptiles, which were fully adapted to these environments, left any kind of track. A large thecodont reptile such as Euparkeria was the potential predator that may rarely have hunted the main small trackmakers Macrocnemus and Hescherleria.     

Middle Triassic horseshoe crab reproduction areas on intertidal flats of Europe with evidence of predation by archosaurs

A systematically excavated track site in a 243.5 Myr old Middle Triassic (Karlstadt Formation, Pelsonian, middle Anisian) intertidal carbonate mud‐flat palaeoenvironment at Bernburg (Saxony‐Anhalt, central Germany) has revealed extensive horseshoe crab trackways attributable to the Kouphichnium Nopsca, 1923 ichnogenus. The exposed track bed of a Germanic Basin‐wide spanned intertidal megatrack site is a mud‐cracked biolaminate surface on which detailed tracks have been preserved because of rapid drying and cementation as a result of high temperatures, followed by rapid covering with a protective layer of arenitic storm or tsunami sediments. The different trackway types and their orientations have allowed a tidal sequence to be reconstructed, with the initial appearance of swimming horseshoe crabs followed by half‐swimming/half‐hopping limulids under the shallowest water conditions. The Bernburg trackways, which have mapped lengths of up to 40 m, were all produced by adult animals and exhibit a variety of shapes and patterns that reflect a range of subaquatic locomotion behaviour more typical of mating than of feeding activities. The closest match to the proportions and dimensions of the horseshoe crab tracks at Bernburg is provided by the largest known Middle Triassic limulid Tachypleus gadeai, which is known from the north‐western Tethys in Spain. The horseshoe crab body fossils recognized in the German Mesozoic intertidal zones, instead, are from juveniles. The uniformly adult size indicated by the trackways therefore suggests that they may record the oldest intertidal reproductive zones of horseshoe crabs known from anywhere in the world, with the track‐makers having possibly migrated thousands of kilometres from shallow marine areas of the north‐western Tethys to reproduce in the intertidal palaeoenvironments of the Germanic Basin. Chirotherium trackways of large thecodont archosaurs also appeared on these flats where they appear to have fed on the limulids. With the tidal ebb, smaller reptiles such as Macrocnemus (Rhynchosauroides trackways) appeared on the dry intertidal flats, probably feeding on marine organisms and possibly also on horseshoe crab eggs. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 103 , 76–105.     

Facies and biota of Anisian to Carnian carbonate platforms in the Northern Calcareous Alps (Tyrol and Bavaria)

Summary During the Middle and early Late Triassic carbonate ramps and rimmed platforms developed at the northwestern margin of the Tethys ocean. In the Northern Calcareous Alps, Anisian stacked homoclinal ramps evolved through a transitional stage with distally steepened ramps to huge rimmed platforms of Late Ladinian to Early Carnian age. Middle Triassic to early Late Triassic facies and biota of basin, slope and platform depositional systems are described. Special emphasis is given to foraminifers, sponges, microproblematic organisms and algae. The Ladinian to early Carnian reef associations are characterized by the abundance of segmented sponges, microproblematica, biogenic crusts and synsedimentary cements. Among the foraminifers, recifal forms likeHydrania dulloi andCucurbita infundibuliformis (Carnian in age) are reported from the Northern Calcareous Alps for the first time. Some sphinctozoid sponges likeParavesicocaulis concentricus were known until now only from the Hungarian and Russian Triassic.     

贵州关岭上三叠统的楯齿龙类化石

记述的新铺中国豆齿龙（Sinocyamodus xinpuensis gen. et sp. nov.）代表了齿龙目的一新属。这是齿龙类化石在我国的首次发现,也是这类特殊的海生爬行动物在欧洲、北非和中东以外地区的首次记录。标本产自贵州关岭晚三叠世瓦窑组一段的泥晶灰岩中,与鱼龙类、幻龙类及海龙类等多种海生爬行类同时产出。这一化石组合表明,贵州关岭三叠纪海生爬行动物群的性质与西特提斯动物群（wstern-Tethyan fauna）的性质极为相似,这两个地区在晚三叠世早期有广泛的动物交流。     

On the first reliable find of the genus <Emphasis Type="Italic">Tholodus</Emphasis> (Reptilia: Ichthyopterygia) in the Asian peripheral area of the Panthalassic ocean

Until recently, Tholodus, a rare ichthyosaur with spherical tooth crowns of the pressing–crushing type, has only been known from the Middle Triassic (Anisian) beds of Western Europe. A record of Tholodus remains in the Karazin Formation (Middle Anisian) of southern Primorye (Russian Far East) significantly expands the range of these reptiles.     

Fossil reptiles from the Spanish Muschelkalk (mont‐ral and alcover,province Tarragona)

The marine tetrapods from the late Ladinian Spanish Muschelkalk deposits located between the towns of Mont‐ral and Alcover (province Tarragona) are reviewed. The fauna comprises an undetermined thalattosaur, a small cyamodontoid placodont probably referrable to Psephoderma, a possible pachypleurosaur with (if any) affinities to the Serpianosaurus‐Neusticosaurus clade, lariosaur specimens currently referred to Lariosaurus balsami, an incomplete specimen of Lariosaurus calcagnii, an endemic nothosaur (Nothosaurus cymatosauroides), and a possible pistosaur. Collectively, the faunal mix comprises endemic taxa as well as elements with affinities to the fauna of the Alpine and/or Germanic Triassic. The fauna suggests faunal interchange through the Burgundy Gate linking the southern branch of the developing Neotethys to the Germanic Basin.     

Conodonts from the Permian Jisu Honguer (Zhesi) Formation of Inner Mongolia, China

The Jisu Honguer Formation (“Zhesi Formation”) is a North China marine carbonate unit of Permian age containing a mixed fauna of Tethyan, Boreal and endemic elements. The age of the Jisu Honguer Formation has been thought to range from Artinskian to Kazanian based on previous studies using mostly benthic macrofossils. A typical Mesogondolella aserrata conodont fauna is reported from the lower part of the upper member of the Jisu Honguer Formation in the Ulanqub District, Zhesi area, of northern China. The fauna indicates a Middle Permian (Guadalupian) age, most likely late Wordian to early Capitanian, for the strata yielding conodonts. The whole Jisu Honguer Formation could be assigned to the Wordian to early Capitanian age. The overlying Yihewusu Formation is probably of Capitanian age. There are no Wuchiapingian marine deposits in the Zhesi area. As characteristic of the open sea Guadalupian conodont faunas of the Tethys, smooth Mesogondolella dominate the fauna. The view that all Tethyan Guadalupian conodont faunas consist exclusively of serrated Mesogondolella cannot be confirmed. Three new species are described: Mesogondolella neoprolongata C. - y. Wang, Mesogondolella mandulaensis C. - y. Wang and Wardlawella jisuensis C. - y. Wang.     

The cranial anatomy of Chinese placodonts and the phylogeny of Placodontia (Diapsida: Sauropterygia)

Placodonts are Triassic marine reptiles that inhabited the eastern and western margins of the Tethys Ocean (modern South China and Europe/Middle East). Although the crania of European taxa are relatively well understood, those of Chinese taxa have not been extensively studied, and most of them have not been incorporated into a comprehensive phylogeny. Here we present the first reconstructions of all known Chinese placodont holotype skulls using micro‐computed tomographic (μCT) scanning and/or detailed anatomical study. We also present the first phylogenetic analyses that incorporate all placodont genera using a general diapsid matrix that includes postcranial characters, and a placodont‐only cranial matrix. Results vary between the matrices; however, both support a monophyletic Placodontia with eastern taxa interspaced throughout, indicating no major separation between the eastern and western Tethyan realms. Support is strong for a western Tethyan origin of Placodontia, although the highly nested Placochelyidae first appear in the upper Middle Triassic of the eastern Tethys. Thus, all placodont clades appear to have originated in a period of intense speciation during the Middle Triassic. © 2015 The Linnean Society of London     

Postcranial morphology and growth of the pachypleurosaur Anarosaurus heterodontus (Sauropterygia) from the Lower Muschelkalk of Winterswijk, The Netherlands

Sauropterygia from the Muschelkalk are only found in lag deposits known as bone beds, and most of the material consists of isolated bones. Alpha taxonomy of Sauropterygia from the Germanic Basin which include Pachypleurosauria is thus based mainly on skull morphology of a few specimens. Articulated or associated postcranial material of pachypleurosaurs, associated with diagnostic skull material, is very rare in the Germanic Basin and currently occurs in larger numbers only in the Lower Muschelkalk of Winterswijk (Gelderland Province, The Netherlands), which continuously produces new material. For the first time, the morphology of several partially articulated skeletons of the pachypleurosaur Anarosaurus heterodontus is described and compared. Some of those specimens have skull material attached; others were identified as pachypleurosaurs on the basis of their long bone histology. The current study revealed that postcranial bones of A.?heterodontus feature a diverse morphology reflecting differences during ontogeny. Thus, A.?heterodontus specimens could be assigned to size classes (I?CIII). However, on the basis of morphology, histology, and maximal known size of isolated skulls and humeri, none of these specimens represent fully grown individuals. Growth mark counts of midshaft-femur samples, morphologically assigned to size class?III, document that this size class was reached within the first year of life. Size class?III continued into the second year of life, and then afterwards skeletal maturity was reached. Thus, a juvenile A.?heterodontus grew very fast, which is also indicated by its bone tissue type, composed of a high number of radial vascular canals and a fast-deposited bone matrix. The assignment of isolated bones from Lower to Middle Muschelkalk localities to A.?heterodontus is now possible with an extensive amended diagnosis of this taxon. This largely contributes to the understanding of taxonomical diversity and distribution. Morphological comparison of the postcranial skeleton of A.?heterodontus with that of the two other valid pachypleurosaurs from the Germanic Basin, Anarosaurus pumilio and Dactylosaurus, supports their close phylogenetic relationship. Furthermore, the skeleton of A.?heterodontus has no morphological or histological aquatic adaptation such as pachyostosis or pachyosteosclerosis and thus represents the least degree of aquatic adaptation within Pachypleurosauria.