Embryonic shell structure of Early–Middle Jurassic belemnites,and its significance for belemnite expansion and diversification in the Jurassic

Early Jurassic belemnites are of particular interest to the study of the evolution of skeletal morphology in Lower Carboniferous to the uppermost Cretaceous belemnoids, because they signal the beginning of a global Jurassic–Cretaceous expansion and diversification of belemnitids. We investigated potentially relevant, to this evolutionary pattern, shell features of Sinemurian–Bajocian Nannobelus, Parapassaloteuthis, Holcobelus and Pachybelemnopsis from the Paris Basin. Our analysis of morphological, ultrastructural and chemical traits of the earliest ontogenetic stages of the shell suggests that modified embryonic shell structure of Early–Middle Jurassic belemnites was a factor in their expansion and colonization of the pelagic zone and resulted in remarkable diversification of belemnites. Innovative traits of the embryonic shell of Sinemurian–Bajocian belemnites include: (1) an inorganic–organic primordial rostrum encapsulating the protoconch and the phragmocone, its non‐biomineralized component, possibly chitin, is herein detected for the first time; (2) an organic rich closing membrane which was under formation. It was yet perforated and possessed a foramen; and (3) an organic rich pro‐ostracum earlier documented in an embryonic shell of Pliensbachian Passaloteuthis. The inorganic–organic primordial rostrum tightly coating the protoconch and phragmocone supposedly enhanced protection, without increase in shell weight, of the Early Jurassic belemnites against explosion in deep‐water environment. This may have increased the depth and temperature ranges of hatching eggs, accelerated the adaptation of hatchlings to a nektonic mode of life and promoted increasing diversity of belemnoids. This study supports the hypothesis that belemnite hatchlings were ‘a miniature of the adults’.     

陷胸式蜂亚科（昆虫纲）化石在我国辽西上侏罗统的发现及其系统演化

描述产自辽宁北票上园地区黄半吉沟上侏罗统义县组下部膜翅目昆虫化石1新属新种Trematothoracoides liaoningensis gen.et sp.nov.,归入薤式产（Sepulcidae)中的陷胸茎蜂亚科（Trematothoracinae)。这是首次在我国发现可行的陷胸茎蜂化石,也是该亚科在侏罗纪的产次报道,分支分析结果表明Trematothoracinae为一单系类群。Thoracotrema与Prosyntexis进化程度较高,构成一对姊妹群,同时又是Trematothorax的姊妹群,Trematothoracoides最为原始,为Thoracotrema Prosymtexis Trematothorax的姊妹群。     

陷胸茎蜂亚科(昆虫纲)化石在我国辽西上侏罗统的发现及其系统演化

描述产自辽宁北票上园地区黄半吉沟上侏罗统义县组下部膜翅目昆虫化石1新属新种Trematothoracoides liaoningensis gen. et sp. nov., 归入葬茎蜂科(Sepulcidae)中的陷胸茎蜂亚科(Trematothoracinae)。这是首次在我国发现可靠的陷胸茎蜂化石,也是该亚科在侏罗纪的首次报道。分支分析结果表明Trematothoracinae为一单系类群。Thoracotrema与Prosyntexis进化程度较高,构成一对姊妹群,同时又是Trematothorax的姊妹群;Trematothoracoides最为原始,为Thoracotrema+Prosyntexis+Trematothorax的姊妹群。     

Farbmuster auf jurassischen Belemniten-Rostren

Colour patterns have been observed in belemnites of the genus Megateuthis from the middle Jurassic of North and South Germany. They are present only in adult specimens, appearing first on the dorsal and then on the ventral side, and, finally, on the flanks. This suggests that the rostrum at this stage was covered only by a thin (translucent) layer of soft tissue. More probably, the rostrum was temporarily uncovered, as indicated by the presence of borings of fungi or algae on several rostra. The regular distribution of these borings suggests pre-mortem infestation of the belemnites. Fossil bacteria have also been found within the individual colour patterns.     

New records of early Middle Jurassic belemnites in the French Subalpine Basin and their paleobiogeographic significance

A biostratigraphic and systematic study based on belemnites collected along with ammonites was performed on four sections in the Subalpine Basin (SE France): Lac du Castillon and La Baume (Castellane area), Galabrun and Grand Lara (Gap area). The specimens, originating from hemi-pelagic marl-limestone alternations in the lower part of the “Calcaires à Zoophycos” Formation, are dated from Middle Aalenian (Murchisonae Zone) to Lower Bajocian (Humphriesianum Zone). Five belemnite taxa (Megateuthis elliptica, Holcobelus munieri, H. trauthi, Pachybelemnopsis roettingensis, Hibolithes sp.) have been identified, and two more taxa are reported in an open nomenclature (Belemnitida incertae sedis sp. 1 and sp. 2). The biostratigraphic range of the belemnite fauna is established. The new findings contribute to a more detailed understanding of the paleobiogeography of holcobelid belemnites that flourished at the northern margin of the Tethys Ocean and formed a distinct sub-Mediterranean fauna. The association herein described is similar to the fauna of the Calabro-Peloritani Arc (Calabria, Italy), a further hint for the supposed paleogeographic position of the latter during the Middle Jurassic.     

Le jurassique supérieur et le Crétacé inférieurd'Ankilizato (bassin de Morandava,Madagascar)

Upper Jurassic and Lower Cretaceous from Beritsa, near Ankilizato, are an important landmark between well known cross sections of Northern (Antsalova) and Southern (Sikily) parts of the Morondava Basin. New ammonites and belemnites fauna point out an Upper Jurassic to Upper Valanginian age for this section. A paleogeographical scheme shows the dynamical structure of «Ankilizato gulf at that time.     

藏南喜马拉雅地区晚侏罗世-早白垩世双壳类雏蛤动物群

藏南喜马拉雅地区晚侏罗世地层中雏蛤化石丰富,其它化石仅有少量箭石和菊石,而且属种单一,多为地方种,无法进一步划分和对比,因此,雏蛤属(Buchia)的组合特征具有重要的地质意义。结合菊石Uhligites和Substeuraces,作者在藏南喜马拉雅地区上侏罗统中识别出:1)Buchia concentrica-Buchia spitiensis;2)Buchiamosquensis- Buchia rugosa;3)Buchia blanfordiana- Buchia piochii等3个雏蛤动物组合;在下白垩统中识别出Buchiasubokensis雏蛤动物组合,其中,Buchia mosquensis-Buchia rugosa组合为证明拉弄拉地区存在晚侏罗世晚基末利期至早提塘期地层提供了新的证据,并与斯匹提地区、北美等地区进行了动物群和地层的对比,初步探讨了本区雏蛤动物群在生物地层方面的研究价值。     

New fossil Progonocimicidae (Hemiptera: Coleorrhyncha: Progonocimicoidea) from the Upper Mesozoic of northeastern China,with a phylogeny of Coleorrhyncha

A new Progonocimicidae species, Cicadocoris anisomeridis sp.n. , with asymmetrical tegmina is described from the Middle Jurassic Jiulongshan Formation in northeastern China. This is the fifth report of Coleorrhyncha from China. A cladistic analysis based on a combination of fossil and extant taxa clarifies the phylogenetic status of the new fossils and allows the reconstruction of inter‐subfamily relationships within the suborder Coleorrhyncha. Coleorrhyncha is monophyletic and divided into two main clades. Progonocimicidae comprises a monophyletic lineage, to which the new fossils belong. The broadly conceived Progonocimicinae and Cicadocorinae, as recognized by earlier authors, are not supported. The monophyly of the family Karabasiidae is also not supported, and its two constituent subfamilies Hoploridiinae and Karabasiinae are raised to family rank. Hoploridiidae is found to be sister group to all extant moss bugs, and Karabasiidae is found to be the monophyletic sister group to Hoploridiidae + all extant moss bugs. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:30BC0498‐3B8A‐4650‐BC9D‐02A0C8D870B2 .     

The belemnites described by Baron Ernst Friedrich von Schlotheim (1764–1832)

E. F. v. Schlotheim described 46 species of Jurassic and Cretaceous belemnites in 1813,1820, and 1832. Their type specimens are in part housed at the Humboldt University of Berlin. The holotype ofBelemnites irregularis is here refigured. Lectotypes forB. acuarius, B. canaliculatus, B. clavatus, B. giganteus, andB. tripartitus are designated and figured for the first time. The lectotype ofBelemnites giganteus is designated to be the neotype ofBelemnites suevicus Klein 1773. 36 species are nomina nuda and nomina dubia, because none of their type specimens could be detected, among them the invalidB. paxillosus v. Schlotheim 1820, which is, according to its lectotype designated herein, likeBelemnites giganteus, a junior synonym ofMegateuthis suevica (Klein 1773). An original type specimenof Belemnites mucronatusv. Schlotheim 1813, for which a neotype had already been ruled by the ICZN in 1985, was re-discovered in thev. Schlotheim collection in 1996.v. Schlotheim’s belemnites were up to now designated to be type species of 12 genera.     

First record of a complete arm crown of the Early Jurassic coleoid Loligosepia (Cephalopoda)

The higher rank systematics of coleoid cephalopods are primarily based on the total number of arms. Completely preserved arm crows are, therefore, the most important character complex by which fossil representatives can be assigned to higher taxa. Whereas belemnoids—similar to modern Decabrachia—are well-known to have ten hooklet-bearing arms, recent studies have shown that Middle-Late Jurassic and Late Cretaceous gladius-bearing coleoids belong to the Vamyropoda, owing to having only eight hooklet-free arms. The arm number of Early Jurassic gladius-bearing coleoids was unclear for a long time. This paper, however, reports the first unequivocally complete arm crown from the Early Jurassic (Toarcian). It consists of eight strongly phosphatised arms of equal shape. Affinity with decabrachiate coleoids is therefore unlikely. The mantle sack of the specimen, which was identified as Loligosepia aalensis (Schübler in Zieten, 1832), has, furthermore, an unusual arrangement of numerous angular pieces. The irregular scattering of the mantle fragments was most probably caused by a scavenger. Finally, hooklets in the food residues in the posterior mantle indicate that Loligosepia preyed upon belemnites.     

Reconstructing the palaeoenvironment of the Middle Russian Sea during the Middle–Late Jurassic transition using stable isotope ratios of cephalopod shells and variations in faunal assemblages

Oxygen and carbon isotope data of well-preserved belemnite rostra and ammonite shells are presented from the Callovian–Oxfordian boundary (uppermost Lamberti to lowermost Cordatum zones) of the Dubki section near Saratov in the Russian Platform. Palaeotemperatures calculated for nektobenthic belemnites (averages of 5 °C and 8 °C for cylindroteuthids and belemnopseids, respectively) show the presence of cold bottom waters in the central part of the Middle Russian Sea during the studied interval. Palaeotemperatures calculated for ammonites, which are assumed to have lived in near-surface waters, are considerably higher (average 13 °C). The presented data show a vertical thermal gradient in the Middle Russian Sea. The belemnite oxygen isotope record and the relative abundances of ammonite families in the Dubki section do not correlate with each other probably as a result of different depth habitats of ammonites and belemnites. A review of literature isotope data shows the climatic zonation in European seas at the Middle–Late Jurassic transition. Despite the flux of cold polar waters to the Middle Russian Sea and the area of Scotland there is no evidence for glaciation at the Middle–Late Jurassic transition. Changes in water circulation during a sea-level highstand were likely a source of spreads of cold bottom waters and cardioceratid ammonite fauna in this time period.The belemnite isotope record of the Callovian–Oxfordian boundary in the Russian Platform is characterized by significant scatter of δ¹³C values. No temporal carbon isotope trend is observed. The δ¹³C values of Russian belemnite rostra average 2.6‰ VPDB being 1 to 2‰ higher than the values of coeval Lower Oxfordian belemnites from the area of the Submediterranean ammonite province. Higher (than Submediterranean) δ¹³C values of Russian belemnite rostra are likely related to high biologic productivity and/or high organic matter burial in semi-isolated Boreal–Subboreal marine basins.     

High northern palaeolatitude Jurassic-Cretaceous palaeotemperature variation: new data from Kong Karls Land, Svalbard

The traditional view of warm equable global climates in the Mesozoic with weakly developed latitudinal temperature gradients has been challenged by several recent studies. However, reliable high palaeolatitude palaeotemperature data is still rare. In this study palaeotemperature data, based on oxygen stable isotope palaeothermometry of well preserved mid Jurassic to Lower Cretaceous belemnites from Kong Karls Land, Svalbard, are presented. These data show cool high latitude marine isotopic palaeotemperatures for endemic species during the Lower to mid Valanginian (7.7°C), which may be compatible with the formation of high latitude ice. Middle Bathonian to Kimmeridgian samples give warmer isotopic palaeotemperatures (9.4°C), whilst samples from the Aalenian to Bajocian give isotopic palaeotemperatures which are warmer still (12.7°C). Comparison of the Callovian temperatures with those from a mid-latitude location shows them to be warmer than would be expected if previous latitudinal temperature gradients are correct. This may be due to the ameliorating effect of oceanic heat transport pole-wards from the Panthalassa Ocean via the South Anyui Ocean.     

Phylogenetic analyses elucidate the inter‐relationships of Pamphilioidea (Hymenoptera,Symphyta)

The phylogeny of the superfamily Pamphilioidea is reconstructed using morphology and DNA sequence data of living and fossil taxa by employing two phylogenetic methods (maximum parsimony and Bayesian inference). Based on our results, the monophyly of Pamphilioidea and Pamphiliidae are corroborated, whereas two extinct families, Xyelydidae and Praesiricidae, are not monophyletic. Because members of Praesiricidae together with Megalodontes form a monophyletic group, we propose that the paraphyletic Praesiricidae is synonymized under Megalodontesidae (syn. nov.). The origin of Pamphilioidea is hypothesized to be as early as the Early Jurassic. To better understand morphological evolution in the early lineages of Pamphilioidea, ancestral states of the first flagellomere and the first and second abdominal terga are reconstructed on the morphology‐based tree. In addition, three new genera (Medilyda, Brevilyda, Strenolyda) with five new species (Medilyda procera, M. distorta, Brevilyda provecta, Strenolyda marginalis and S. retrorsa) are described based on well‐preserved xyelydid fossils from the Middle Jurassic Jiulongshan Formation of north‐eastern China.     

Composition and ontogeny ofDictyoconites (aulacocerida,cephalopoda)

Dictyoconites from the middle Triassic Cassian Formation is a characteristic representative of the Aulacocerida. Embryonic development and construction of the phragmocone is like that of Jurassic belemnites. The siphuncular tube is double-walled with a long retrochoanitic mineralized septal neck continuing into an organic tube. The extended decoupling zone resembles that ofSpirula. Characteristic ofDictyoconites are the tubular »living chamber« and two layered deposits of the muscular mantle on the phragmocone. The Triassic coleoid was a slender squid with visceral mass and mantle cavity encapsuled in shell and the whole conch covered by muscular mantle extending in two lateral apical fins attached to the aragonitic rostrum.     

Comparison of carbonate C and O stable isotope records across the Jurassic/Cretaceous boundary in the Tethyan and Boreal Realms

Carbon and oxygen stable isotope records were compared for Jurassic/Cretaceous (J/K) boundary sections located in the Tethyan Realm (Brodno, Western Slovakia, and Puerto Escaño, Southern Spain; bulk limestones), and the Boreal Realm (Nordvik Peninsula, Northern Siberia, belemnites). Since a detailed biostratigraphic correlation of these Tethyan and Boreal sections is impossible due to different faunal assemblages, correlation of the isotope records was based on paleomagnetic data. This novel approach can improve our understanding of the synchroneity of individual isotope excursions in sections where detailed biostratigraphic correlation is impossible. No significant excursions in either the carbon or oxygen isotope records to be used for future Boreal/Tethyan correlations were found around the J/K boundary (the upper Tithonian and lower Berriasian; magnetozones M20n to M18n) in the studied sections. At the Nordvik section, where a much longer section (middle Oxfordian–basal Boreal Berriasian) was documented, the transition from the middle Oxfordian to the Kimmeridgian and further to the Volgian is characterized by a decrease in belemnite δ¹⁸O values (from δ¹⁸O values up to + 1.6‰ vs. V-PDB in the Oxfordian to values between + 0.3 and ? 0.8‰ in the late Volgian and earliest Boreal Berriasian). This trend, which has previously been reported from the Russian Platform and Tethyan Realm sections, corresponds either to gradual warming or a decrease in seawater δ¹⁸O. Supposing that the oxygen isotope compositions of seawater in the Arctic/Boreal and Tethyan Realms were similar, then the differences between oxygen isotope datasets for these records indicate differences in temperature. The Boreal/Tethyan temperature difference of 7–9 °C in the middle and late Oxfordian decreases towards the J/K boundary, indicating a significant decrease in latitudinal climatic gradients during the Late Jurassic. Two positive carbon isotope excursions recorded for the middle Oxfordian and upper Kimmeridgian in the Nordvik section can be correlated with a similar excursion described earlier for the Russian Platform. Minor influence of biofractionation at the carbon isotopes, and the influence of migration of belemnites to deeper, slightly cooler water at the oxygen isotopes, cannot be excluded for the obtained belemnite data.     

The oldest known mastotermitids (Blattodea: Termitoidae) and phylogeny of basal termites

Five finely preserved termites from the mid‐Cretaceous (Cenomanian) amber of Myanmar provide new information allowing a reanalysis of the phylogeny of basal termites. The Mastotermitidae family is recovered as monophyletic, and a redefined Hodotermitidae sensu lato is also monophyletic to include Archotermopsidae, Hodotermitidae and Stolotermitidae. Such a phylogenetic relationship agrees with the results from previous molecular phylogeny. Alongside these findings, there are many taxa that can only be shown to be termites with no other phylogenetically informative data. These form a comb of ‘grade groups’ emerging in the Late Jurassic. The new amber specimens are described as two new species. Anisotermes xiai gen. et sp.n. is described from multiple castes and has symplesiomorphic characteristics: large body size, a broad pronotum, well‐developed reticulated veins, and a large anal lobe of the hindwings. It shares wing features with the other new species, Mastotermes monostichus sp.n. Both new taxa are assigned to the Mastotermitidae, as they are shown to have synapomorphies that unite the family. This published work has been registered on ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:1AD5CECA‐27B7‐48D5‐88DC‐CEC5150962D7 .     

Signs of predation in the Middle Jurassic of south-central Poland: evidence from echinoderm taphonomy

Distinct faunal aggregates are described from the Middle Jurassic (uppermost Bajocian/lowermost Bathonian and Middle Bathonian) clay deposits of Częstochowa area, south-central Poland. These aggregates are composed of molluscs (scaphopods, gastropods, bivalves, ammonites and belemnites), articulate brachiopods and echinoderms (asteroids, crinoids and echinoids). A large percentage of the fossils, especially bivalves, are fragmented, but some fossils are complete. Although most of the fossils are crushed and fragmented, they are still identifiable to at least the genus level. Thorough statistical analysis of taphonomic features indicates that the preservation of asteroid marginal plates is distinct from the ossicles derived from the host clays. The high frequency of bite marks and the good state of preservation suggest that the accumulations are the products of predation activities and most probably are the effect of regurgitation. Taking into account the rich and diverse fauna, the predator was a bottom-feeding generalist. Possible predators include palaeospinacid sharks, a tooth of which was collected from the same bedding surface, but not associated with regurgitated remains. Although the bite marks on the asteroid ossicles point to sharks as potential producers of regurgitates, other vertebrates, like durophagous pycnodontiform fish, cannot be excluded.     

New teleostean fishes from the Jurassic of southern Germany and the systematic problems concerning the ‘pholidophoriforms’

A new genus,Siemensichthys, from the Upper Jurassic of southern Germany is described. The new genus includes two species,S. macrocephalus (Agassiz) which was formerly in the genusPholidophorus, andS. siemensi n. sp. The two species share synapomorphies such as only one supramaxillary bone covering the dorsal margin of the maxilla. Both species are described, and their phylogenetic position is analyzed. The phylogenetic analyses, based on 27 taxa and 141 characters, show thatAnkylophorus from the Kimmeridgian of Cerin,Siemensichthys andEurycormus from the Solnhofen Lithographic Limestone of Bavaria, form a monophyletic group. The new extinct clade (preliminarily identified as theSiemensichthys- group) is proposed as the sister-group ofPholidophorus s. str. plus more advanced teleosts. This sister-group relationship is supported by eight characters (e.g., supraoccipital bone extending forward in the roof of the otic region; articular bone fused with both the angular and retroarticular; presence of an elongated posteroventral process of quadrate; presence of dorsal processes at the base of the innermost caudal rays of upper lobe; mobile premaxillary bone). Comparisons with species ofPholidophorus s. str. provide a new understanding of the genusPholidophorus. At least four synapomorphies are proposed to support the monophyly ofPholidophorus. As a consequence of this new interpretation, the European Late Jurassic species previously assigned to the Pholidophoridae and to the genusPholidophorus (e.g., ‘Ph.’armatus, ‘Ph.’ falcifer, ‘Ph.’ micronyx, ‘Ph.’ microps) should be reexamined because they do not belong to the family nor to the genus. The order PholidophoriformesBerg is not a monophyletic group as currently constructed. Therefore, all so-called pholidophoriforms are in need of revision.     

A dated molecular phylogeny for the Chironomidae (Diptera)

We provide the first highly sampled phylogeny estimate for the dipteran family Chironomidae using molecular data from fragments of two ribosomal genes (18S and 28S), one nuclear protein‐coding gene (CAD), and one mitochondrial protein‐coding gene (COI), analysed using mixed‐model Bayesian and maximum likelihood inference methods. The most recently described subfamilies Chilenomyiinae and Usambaromyiinae proved elusive, and are unsampled. We confirm monophyly of all sampled subfamilies except Prodiamesinae, which contains Propsilocerus Kieffer, previously in Orthocladiinae. The semifamily Chironomoinae is confirmed only if Telmatogetoninae is included, which is closer to Brundin's original suggestion. Buchonomyiinae is excluded from Chironomoinae: it is a sister group to all remaining Chironomidae, conforming more to Murray and Ashe's argumentation. Semifamily Tanypodoinae is a grade and unsupported as monophyletic: the austral Aphroteniinae alone is sister to all Chironomidae (less Buchonomyiinae). Podonominae is weakly supported as the next sister group, in contrast to some estimates that place this subfamily as sister group to Tanypodinae alone. In Diamesinae, the southern African Harrisonini is confirmed as a member, but embedded within austral tribe Heptagiini, which is confirmed as sister to the undersampled Diamesini. Tribe Pentaneurini and ‘non‐Pentaneurini’ taxa are reciprocally monophyletic in Tanypodinae. Recent molecular findings concerning Podonominae are substantiated, with a monophyletic tribe Podonomini, Boreochlini forming a grade and Lasiodiamesa Kieffer placed as sister to all other Podonominae, but with uncertainty. In Orthocladiinae, a postulated two‐tribe system of Orthocladiini and Metriocnemini can be supported after exclusion of a Corynoneura group and a Brillia group, which is revealed as sister to Stictocladius Edwards. The marine Clunio Haliday and Thalassosmittia Strenzke & Remmert (given high rank in the past) are clearly embedded deep in Orthocladiinae. The finding of Shangomyia Sæther & Wang + Xyiaomyia Sæther & Wang as sister group to all other Chironominae justifies high rank, as their authors suggested. Pseudochironomini (untested by sampling shortfall) is sister to a monophyletic Tanytarsini (with a weakly supported inclusion of the enigmatic Nandeva Wiedenbrug, Reiss & Fittkau). The tribe Chironomini can be supported only by excluding Shangomyia + Xyiaomyia, and a postulated monophyletic clade comprising several taxa such as Microtendipes Kieffer, with six‐segmented larval antennae and alternate Lauterborn organs, that is sister group to Pseudochironomini + Tanytarsini. The tempo of diversification of the family, deduced by divergence time analysis (beast ), shows Permian origination with subfamily stem‐group origination from the mid–late Triassic to the early Cretaceous. Crown‐group origination ranged from Podonominae on a short stem originating in the mid Jurassic to long‐stemmed Aphroteninae from the late Cretaceous. Node dates allow inference of some vicariance via Gondwanan fragmentation, including certain nodes involving southern Africa.     

THE TAXONOMIC AND PHYLOGENETIC POSITION OF THE PLESIOSAUROIDEA FROM THE LOWER JURASSIC POSIDONIA SHALE OF SOUTH-WEST GERMANY

Abstract:  The two plesiosauroid species from the Posidonia shale of Holzmaden, ' Plesiosaurus ' guilelmiimperatoris and ' Plesiosaurus ' brachypterygius , do not belong to Plesiosaurus but form new monotypic genera. The new genus Hydrorion is erected for ' P .' brachypterygius , and the genus Seeleyosaurus is re-established for ' P .' guilelmiimperatoris . The recently described species Plesiopterys wildii is regarded as a junior synonym of S. guilelmiimperatoris . A short phylogenetic analysis shows that S. guilelmiimperatoris and Muraenosaurus are basal elasmosaurs. H. brachypterygius , Occitanosaurus tournemirensis from France and Microcleidus homalospondylus from England form a monophyletic clade, which is the sister taxon to the Cretaceous elasmosaurs. A palaeobiogeographical comparison of plesiosaur localities in the Lower Jurassic shows distinct palaeobiogeographical zones for the Toarcian, with different plesiosaur taxa in England, Germany and France.