A catalogue of teleosauroids (Crocodylomorpha: Thalattosuchia) from the Toarcian and Bajocian (Jurassic) of southern Luxembourg

Abstract

Teleosauroids were a clade of semi-marine crocodylomorphs that attained near-global distribution during the Jurassic Period. They were particularly common during the Toarcian (Early Jurassic) and are well documented throughout the UK and Germany. However, Toarcian teleosauroids discovered in Luxembourg have been little studied and rarely discussed in the scientific literature. Here we present a comprehensive catalogue of Luxembourg thalattosuchian specimens, including nine teleosauroids (all from the Toarcian) and five Thalattosuchia indeterminate (four from the Toarcian and one from the Bajocian), many of which are noted in the literature for the first time. We describe these specimens and identify two distinct genera (Steneosaurus and Platysuchus) as present in the sample as well as three, or possibly four, distinct species. This represents a high diversity of teleosauroid taxa (both common and rare forms) from the Toarcian rarely seen elsewhere in the world.     

Ophiuroid remains from the Toarcian of Sainte-Verge (Deux-Sèvres, France): paleobiological perspectives

The Toarcian sediments exposed at Sainte-Verge (Deux-Sèvres, France) are especially rich in echinoderm remains. The present paper describes and illustrates the ophiuroids. On the basis of lateral arm plates, 13 species are distinguished, including two new ones: Sinosura fasciata sp. nov. and Sinosura extensa sp. nov. Most of the recognized species have been recorded previously from the Late Toarcian and Aalenian in Germany and, to a lesser extent, from the late Early Jurassic of England and Switzerland. High similarities between the faunas of northwest Europe suggest a boreal provincialism of ophiuroids. The recognition of 13 species is comparable to the diversity known from other stratigraphic levels (Jurassic and Cretaceous) or in the richest stations of recent oceans. The species association of the Toarcian of Sainte-Verge, with two Ophiolepididae, one Ophiacanthidae, four Ophioleucidae, two Ophiodermatidae, two Ophiuridae, and one Hemieuryalidae may be compared with species associations of recent shelf, offshore environments. Such persistence of components of diversity and ecological affinities of species suggests strong evolutionary conservatism of the ophiuroids, after a rapid radiation during the earliest Jurassic. © 2002 Éditions scientifiques et médicales Elsevier SAS. All rights reserved.     

First palynological data from the Jurassic South Xiangshan Formation (Nanjing area,China)

The outcrops of the Xiangshan Group have been studied since the beginning of the twentieth century, yielding a rich macroflora (the Xiangshan Flora), mainly from the lower part of the Xiangshan Group (South Xiangshan Formation). Nevertheless, no palynological data have been published so far from the South Xiangshan Formation. The present study provides the first palynostratigraphic data of the South Xiangshan Formation. More than 50 fossil taxa from 30 fossil genera have been identified, allowing for a more accurate dating. The most characteristic taxa are Polycingulatisporites triangularis, Quadraeculina anellaeformis, Manumia delcourtii, Ischyosporites variegatus, Callialasporites turbatus, C. trilobatus, C. minus, and Sestrosporites pseudoalveolatus, suggesting a late Toarcian-late Aalenian age for the South Xiangshan Formation. These results are consistent with previous studies suggesting a middle-late Early Jurassic age for this formation; consequently, the most probable age for the South Xiangshan Formation is late Toarcian (late Early Jurassic).     

Marine reptiles (Thalattosuchia) from the Early Jurassic of Lombardy (northern Italy)

The fossil remains of two small reptiles recently discovered in the Sogno Formation (Lower Toarcian) near Cesana Brianza (Lecco Province), represent the first mesoeucrocodylians reported for Lombardy and some of the few Jurassic reptiles from Italy. Due to the absence of diagnostic skeletal elements (the skulls are lacking), it is not possible to refer the new specimens at genus level with confidence. Although the well developed dermal armour would characterise Toarcian thalattosuchians of the genera Steneosaurus (Teleosauridae) and Pelagosaurus (Metriorhynchidae), the peculiar morphology of the osteoderms allow to tentatively refer the remains to the latter taxon (cf. Pelagosaurus sp.). The small size, along with the opening of the neurocentral vertebral sutures and, possibly, the non sutured caudal pleurapophyses, indicate that the specimens were morphologically immature at death. These “marine crocodiles” confirm the affinities between the fauna of the Calcare di Sogno Formation and coeval outcrops of central Europe that also share the presence of similar fishes and crustaceans.     

Global radiolarian zonation for the Pliensbachian, Toarcian and Aalenian

Jurassic radiolarians from 220 samples in Queen Charlotte Islands, B.C., Williston Lake, B.C., east-central Oregon, Baja California Sur, southern Spain, Austria, Slovenia, Turkey, Oman, Japan and Argentina were studied in order to construct global zonation for the Pliensbachian, Toarcian and Aalenian stages. Well-preserved faunas from continuous stratigraphic sections in Queen Charlotte Islands provide the most detailed record for this time interval, and all collections are tied to North American ammonite zones or assemblages. Collections from nearly all other areas lack independent dating except for early Toarcian carbon-isotope dating in Slovenia and late Aalenian ammonites in Spain.A database of 197 widely distributed updated taxonomic species was used to construct a Unitary Association (UA) zonation for the interval. A global sequence of 41 UAs was obtained for the top of the Sinemurian to the base of the Bajocian. The first and the last UAs represent the Late Sinemurian and the Early Bajocian respectively. The remaining 39 UAs were merged into nine zones (four Early Pliensbachian, one Late Pliensbachian, one Early Toarcian, one Middle-Late Toarcian, and two Aalenian) according to prominent radiolarian faunal breaks and ammonite data. The new zones are the Canutus tipperi - Katroma clara Zone (latest Sinemurian/earliest Pliensbachian); Zartus mostleri - Pseudoristola megaglobosa, Hsuum mulleri - Trillus elkhornensis and Gigi fustis - Lantus sixi zones (Early Pliensbachian); Eucyrtidiellum nagaiae - Praeparvicingula tlellensis Zone (Late Pliensbachian); Napora relica - Eucyrtidiellum disparile Zone (Early Toarcian); Elodium pessagnoi - Hexasaturnalis hexagonus Zone (Middle and Late Toarcian); Higumastra transversa - Napora nipponica Zone (early Aalenian); and Mirifusus proavus - Transhsuum hisuikyoense Zone (late Aalenian). These zones can be correlated worldwide and link previously established UA zonations for the Hettangian-Sinemurian and the Middle to Upper Jurassic. The new zonation allows high-resolution dating in the studied interval and provides a solid basis for analyzing faunal turnovers and the paleobiogeography of Jurassic radiolarians.     

The oldest giant lacewings (Neuroptera: Kalligrammatidae) from the Lower Jurassic of Germany

We describe the oldest Kalligrammatidae, two distantly related species of different subfamilies from the Lower Jurassic (lower Toarcian) of Germany: Liassopsychops curvatus Bode, 1953, sit. nov. (Liassopsychopinae n. subfam.) and Ophtalmogramma klopschari n. gen. n. sp. (Kallihemerobiinae). They lived in warm and relatively dry conditions. Subfamily Liassopsychopinae includes the Jurassic genera Liassopsychops and Huiyingogramma, sit. nov., which bear forewings characterized by Sc and RA not fused distally; MP, CuA, CuP, and A1 dichotomously branched; and a well-developed central eye-spot. The forewing of Ophtalmogramma n. gen. is most similar to that of Apochrysogramma Yang et al., 2011. The diversity of early Toarcian kalligrammatids indicates a late Triassic–earliest Jurassic origin of the family. A modified classification of the central eye-spots is proposed based on the presence/absence of four main components, i.e., the central pigmented disc (A), swellings (B), ocules (C), and outer rings (D). The function of the swellings is unclear, but they may have served as pheromone containers.     

Latest Toarcian-earliest Bajocian (Jurassic) Grammoceratinae (Hildoceratidae,Ammonitina) of the western Tethys: Their palaeobiogeographic and phylogenetic significance

Grammoceratinae (Hildoceratidae, Ammonitina) abound in the Toarcian strata of many western Tethyan localities, especially the Subbetic and Lusitanian basins (of southern Spain and central western Portugal, respectively). They decline through the Aalenian and disappear by the lowermost Bajocian. The genera Asthenoceras, Vacekia (subgenera Vacekia and Nadorites) and Fontannesia are traditionally considered as the last Grammoceratinae, with species of Osperleioceras occurring in the uppermost Toarcian. Grammoceratinae are common in the eastern Pacific, especially Oregon and Alaska where Asthenoceras is abundant. They also occur in the eastern Tethys (Thailand). Although studies of Toarcian to early Bajocian Subbetic and Lusitanian grammoceratins already exist, new material from these and other palaeogeographic areas (England, Portugal and Spain) is revised here. A new genus, Linaresites nov. gen. (for Fontannesia montillanensis Linares and Sandoval) and two new species (Vacekia striata Henriques, and Asthenoceras taverai Sandoval) are described. Another form, “Asthenoceras” sp. A is described and let in open nomenclature. Temporal analysis of Aalenian to early Bajocian Grammoceratinae demonstrates a progressively more evolute morphology through time, sometimes coupled with size reduction. Palaeogeographic evidence suggests that during the early Middle Jurassic, western Tethys and eastern Pacific were temporarily well connected, possibly through the Hispanic Corridor, as demonstrated by the similarity between Tethyan and eastern Pacific Grammoceratinae.     

Benthic foraminiferal bioevents in lower to upper Toarcian strata of Southern Beaujolais (SE France)

This paper documents changes in benthic foraminiferal assemblages compared with high resolution ammonite biozonation along the lower Toarcian to upper Toarcian marine succession of Southern Beaujolais in southeastern France. Eight ammonite and three benthic foraminiferal zones including five subzones are distinguished based on the occurrence of twelve foraminiferal events. Each benthic foraminiferal subzone is characterized by its taxonomic and morphogroup composition, which represents the paleoecological response of these taxa and morphotypes of benthic foraminifera in the Early Jurassic and early Middle Jurassic. Major changes in abundance and diversity occur at the end of the Toarcian Oceanic Anoxic Event (T-OAE) and near the Early-Middle Jurassic transition. The low-abundance foraminiferal assemblages recorded in the Serpentinus ammonite Zone are interpreted as reflecting adverse environmental conditions after the T-OAE. The later recovery and development of the foraminiferal assemblages is documented in the Bifrons up to the Aalensis zones and is attributed to improved bottom water oxygenation. Common occurrences of agglutinated foraminifera represented mostly by Trochammina pulchra Ziegler in the Dispensum Zone point to an influx of cooler water masses during the late Toarcian. The morphogroup analysis carried out on the foraminifera and their paleoecological interpretations shed light on the changes in the stratigraphic record at the end of the T-OAE up to the Toarcian/Aalenian boundary.     

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.     

A new species of glypheid lobster,Glyphea pisuergae (Crustacea,Glypheoidea), from the Early Jurassic of Palencia,Basque-Cantabrian Basin,Spain

A new glypheid lobster (Crustacea, Glypheidae), Glyphea pisuergae sp. nov. is described from the Early Jurassic (late Pliensbachian–early Toarcian) of Salinas de Pisuerga, Palencia, Spain. This species represents the second record for the genus in the Early Jurassic.     

The Early Jurassic palynostratigraphy of the Lusitanian Basin,western Portugal

A comprehensive investigation of the Early Jurassic stratigraphical palynology of the Lusitanian Basin in western Portugal was undertaken, with most emphasis placed on dinoflagellate cysts. A total of 214 samples from an upper Sinemurian to upper Toarcian composite section based on six successions were examined. The Sinemurian material examined was barren of dinoflagellate cysts; however, the Pliensbachian and Toarcian successions are characterised by relatively low diversities where Luehndea spinosa, Mancodinium semitabulatum, Mendicodinium microscabratum, Nannoceratopsis gracilis, Nannoceratopsis senex, and Scriniocassis priscus are relatively common and biostratigraphically significant. Luehndea spinosa dominates the lowermost Toarcian (Dactylioceras polymorphum ammonite Biozone), and is an index species. At the base of the Hildaites levisoni ammonite Biozone, the effects of the Toarcian Oceanic Anoxic Event (T-OAE) caused Luehndea spinosa to become extinct. At the same time, dinoflagellate cyst abundance and diversity markedly decreased. After the T-OAE, during the middle and late Toarcian, phytoplankton recovery was prolonged and slow in the Lusitanian Basin. The Luehndea spinosa and Mendicodinium microscabratum dinoflagellate cyst biozones are defined, both of which are subdivided into two dinoflagellate cyst subbiozones.     

Palynology of the Toarcian-Aalenian Global Boundary Stratotype Section and Point (GSSP) at Fuentelsaz (Lower-Middle Jurassic, Iberian Range, Spain)

The Fuentelsaz Section is located in the Castilian Branch of the Iberian Range (Guadalajara, Spain). Its exceptional Lower-Middle Jurassic transition outcrops led to its designation as the Global Boundary Stratotype Section and Point for the base of the Aalenian. The sediments of the Toarcian-Aalenian transition at Fuentelsaz are composed of marls with interbedded limestones in rhythmic alternation. Marls are dominant in the Toarcian and the Opalinum Zone up to the middle part of the Comptum Subzone. In the remaining Comptum Subzone, marly materials are scarce and limestones predominate. Sedimentation at Fuentelsaz took place over an extensive marine epeiric carbonate platform that was well connected with the open sea. In the Fuentelsaz Section, a total of 43 palynomorph taxa were recorded: 23 spore taxa, 13 pollen taxa, 4 acritarchs, 2 prasinophytes and 1 dinoflagellate cyst. The studied sediment samples were always dominated by terrestrial allochthonous miospores. In general, miospore assemblages are biased due to the transport of pollen from land or islands to the continental platform. Spheripollenites, Classopollis or indeed both pollens numerically dominate the assemblages. Other miospores appear in low numbers. Aquatic palynomorphs are also scarce; Micrhystridium lymensis is the most common. Five palynological assemblages (PA) were distinguished: PA1, PA2 and PA3 are Toarcian in age, PA4 is located at the boundary between the Toarcian and the Aalenian and PA5. This latter palynological assemblage which is lower Aalenian in age shows a strong reduction in palynomorph diversity and preservation in the Comptum Subzone. Palaeoecological analysis of the palynomorphs indicated the presence of calm, oligotrophic sea water. The palaeofloral communities of gymnosperms and vascular cryptogams—which grew in subtropical arid conditions—were poorly diversified.     

Discovery of Iliestheria (Crustacea: spinicaudatan) from the Lower Jurassic Sangonghe Formation in Junggar Basin,northwestern China

Iliestheria Li and Shen, 1995, was erected based on specimens of two co-occurring species from the Lower Jurassic Badaowan Formation in Nilka, Xinjiang Uygur Autonomous Region, northwestern China. Since then Iliestheria has not been reported in any other locality, partly because clam shrimp from the Lower Jurassic in western China have been reported with studies only based on the outline of the carapace, most of them as Euestheria or Palaeolimnadia components. Here we report a new species of Iliestheria from the Pliensbachian–Toarcian Sangonghe Formation, based on scanning electron microscopic characteristics of the carapace morphology. We also revise the diagnosis of the genus based on re-examination using SEM microscopy of the two species from the Badaowan Formation, to include spinicaudatans characterized by growth bands ornamented by radial lirae intercalated with 2–4 columns of puncta; radial lirae sporadically curved and branched to form fine reticulation on the growth bands in the anterior and upper parts of the carapace. This study indicates that the stratigraphic range of Iliestheria extends to late Early Jurassic.     

New bibionomorpha (Insecta: Diptera) from the Jurassic of Asia

The new species Protorhyphus rohdendorfi sp. nov. (Protorhyphidae), Mesorhyphus handlirschi sp. nov., and M. hennigi sp. nov. (Anisopodidae), Procramptonomyia ponomarenkoi sp. nov. and P. kovalevi sp. nov. (Procramptonomyiidae) are described from the Upper Jurassic deposits of Mongolia (Shar Teg); some features of Archirhyphus asiaticus Rohdendorf, 1964 are revealed. The latter two species are also known from the locality of Karatau (Kazakhstan, J_2/3). The genus Tega Blagoderov, Krzemi??ska, and Krzemi??ski, 1993 is transferred from the family Cramptonomyiidae into Anisopodidae (Teginae subfam. nov.).     

An eastern Tethyan (Tibetan) record of the Early Jurassic (Toarcian) mass extinction event

A record of the Early Jurassic mass extinction event is reported from eastern Tethyan (Tibetan) locations for the first time. In the Mount Everest region a thick Lower Jurassic carbonate formation, here named the Yungjia Formation, is developed within the predominantly clastic Triassic–Jurassic succession. Within the formation a sharp transition from peloidal packstones/grainstones to thin‐bedded, pyritic micrite‐shales interbeds records a sharp pulse of deepening and development of dysoxic bottom waters. Both the lithiotid bivalves and the lituolid foraminifera are important constituents of the lower Yungjia Formation but they disappear at this flooding surface or a short distance below it. This extinction event is comparable to that seen at the base of the Pliensbachian/Toarcian boundary in western Tethyan platform carbonates but the Tibetan events occurred late in the Toarcian Stage as indicated by nannofossil biostratigraphy and C isotope chemostratigraphy. The Early Jurassic extinction event (and the associated spread of oxygen‐poor waters) was therefore not synchronous throughout the Tethyan region.     

Systematics and palaeoecology of middle Toarcian Asteroidea (Echinodermata) from the “Seuil du Poitou”, Western France

Complete, articulated starfish fossils are rare. However, more frequently encountered dissociated skeletal elements (ossicles) permit reliable taxonomic identification, making them a valuable data source for diversity estimates. Nearly 300 asteroid ossicles, collected from the middle Toarcian marls in western France can be assigned to five species. Four species and two genera are described: Comptoniaster vrinensis nov. sp. (Goniasteridae), Poncetaster crateri nov. gen. nov. sp. (Stauranderasteridae), Galbaster recurrans nov. gen. nov. sp. (Goniasteridae) and Pentasteria? liasica nov. sp. (Astropectinidae). The known diversity of Early Jurassic asteroids is increased from 12 to 16 species. These taxa illustrate the diversification of crown-group asteroids early in the Jurassic, following the Permo-Triassic crisis. They also reflect bias of the fossil record, and imply the existence of numerous ghost lineages in the evolutionary trees of extant groups. Variation in asteroid diversity across the “Seuil du Poitou” was driven by ecological constraints. The relative frequency (abundance of ossicles and diversity) of goniasterids and stauranderasterids increases in shallower environments. The Benthopectinidae, represented by Plesiastropecten hallovensis, occurred primarily from deep-shelf sediments. Similar ecological patterns are observed for more recent fossil and extant relatives, which further supports the idea of conservative evolution in post-Palaeozoic starfishes since the Early Jurassic.     

First diagnostic marine reptile remains from the aalenian (middle jurassic): a new ichthyosaur from southwestern Germany

Background

The Middle Jurassic was a critical time in the evolutionary history of ichthyosaurs. During this time interval, the diverse, well-studied faunas of the Lower Jurassic were entirely replaced by ophthalmosaurids, a new group that arose sometime prior to the Aalenian-Bajocian boundary and by the latest middle Jurassic comprised the only surviving group of ichthyosaurs. Thus, the Middle Jurassic Aalenian-Bathonian interval (176–165 million years ago) comprises the time frame during which ophthalmosaurids not only originated but also achieved taxonomic dominance. However, diagnostic ichthyosaur remains have been described previously from only a single locality from this interval, from the Bajocian of Argentina.

Methodology/Principal Findings

In this paper, we describe a new species of ichthyosaur based on a partial articulated specimen from the Middle Jurassic of southwestern Germany. This specimen was recovered from the Opalinuston Formation (early Aalenian) and is referable to Stenopterygius aaleniensis sp. nov. reflecting features of the skull and forefin. The genus Stenopterygius is diverse and abundant in the Lower Jurassic of Europe, but its presence has not previously been confirmed in younger (Middle Jurassic) rocks from the northern hemisphere.

Conclusions/Significance

This specimen represents the only diagnostic ichthyosaur remains reported from the Aalenian. It bears numerous similarities in size and in morphology to the Lower Jurassic species of the genus Stenopterygius and provides additional evidence that the major ecological changes hypothesized to have occurred at the end of the Toarcian took place sometime after this point and most likely did not occur suddenly. There is currently no evidence for the presence of ophthalmosaurids in the northern hemisphere during the Aalenian-Bathonian interval.     

On Lower and Upper Jurassic dibranchiate cephalopods from Germany and England

Three dibranchiate cephalopod remains from different bituminous shales contribute to our knowledge of Jurassic representatives: 1. The recently re-discovered holotype ofLioteuthis problematica Naef, 1922, from the Lower Toarcian of southwestern Germany has been prepared and is found to be taxonomically and phylogenetically isolated of its strange morphology: 2. An exceptionally preserved ink sac of a Lower Toarcian squid [probablyBelopeltis aalensis (Zieten, 1832)] from southwestern Germany, shows the finest imprints of the blood vessels yet seen. 3. The Callovian speciesBelemnoteuthis antiqua Mantell, 1847, mentioned, but not figured from the middle Kimmeridge Clay of southern England, is figured here from these beds for the first time.     

A longirostrine Temnodontosaurus (Ichthyosauria) with comments on Early Jurassic ichthyosaur niche partitioning and disparity

Abstract: We describe an almost complete ichthyosaur skeleton from the middle Toarcian (Lower Jurassic) of the Beaujolais foothills near Lyon, France, and assign it to Temnodontosaurus azerguensis sp. nov. This new species exhibits cranial peculiarities such as a thin, elongated and possibly edentulous rostrum, as well as a reduced quadrate. These characters indicate dietary preferences that markedly differ from other species referred to Temnodontosaurus, a genus previously considered as the top predator of the Early Jurassic seas. Despite a conservative postcranial skeleton, we propose that Temnodontosaurus is one of the most ecologically disparate genera of ichthyosaurs, including apex predators and now a soft prey longirostrine hunter. Ammonites collected from the same stratigraphic level as the described specimen indicate that the new species is somewhat younger (bifrons ammonite zone) than the most known Toarcian ichthyosaurs and therefore slightly postdates the interval of severe environmental changes and marine invertebrate extinctions known as the Toarcian Oceanic Anoxic Event. The present study therefore raises the question of whether postcrisis recovery of vertebrate faunas, including the radiation of Temnodontosaurus into a new ecological niche, may have been a consequence of marine ecosystem reorganization across this event.