Ammonite diversity and its palaeobiogeographical structure during the early Pliensbachian (Jurassic) in the western Tethys and adjacent areas

The early Pliensbachian (Early Jurassic) is known as a time of marked provincialism in the marine realm, notably between the Mediterranean Tethys and North–West Europe. In order to test this observation quantitatively, we compiled 104 locality-level species lists from those areas based on a comprehensive revision of early Pliensbachian ammonites. With this dataset, we also explore the relationship between ammonite richness and biogeography at the scale of the sub-chronozone during the early Pliensbachian. Using various multivariate statistics and rarefaction techniques, we show that: (i) there is a sharp contrast between the NW European (NWE) and the Mediterranean (MED) provinces, although there is some mixing in Austroalpine and Pontic ammonite faunas; (ii) species richness in the MED province is about twice that in the NWE province for each chronozone; (iii) ammonite species richness tends to decrease during the early Pliensbachian, especially at the Ibex–Davoei transition; and (iv) the NWE and MED sensu stricto provinces both record the same pattern of variations in richness despite the fact that their taxonomic compositions have virtually nothing in common at the species level. We suggest that the low ammonite richness of the Davoei chronozone may be related to a coeval warming of seawaters, but that this was insufficient to affect the sharp palaeobiogeographic contrast between the two provinces. This persistent compartmentalisation probably reflects a major palaeogeographical structure, such as an emerged or near-emerged barrier running from the Betic range to the Briançonnais ridge. Overall, it seems that the diversity and distribution of early Pliensbachian ammonite species were simultaneously controlled by climate, palaeogeography and eustasy.     

Aequiloboidea: A new Early Jurassic ammonite superfamily of the Mediterranean Tethys

A comprehensive outline of the Jurassic ammonoid superfamily Aequiloboidea is here presented: this group is known from? Rhaetian-early Hettangian to early Pliensbachian and it is exclusively found within the Mediterranean Tethys (Austrian Alps, Bavarian Prealps, Tuscany, Umbria-Marche and Abruzzi Apennines, Morocco, Tunisia, western Hungary). On the basis of the available documentation, it is composed by two families (Aequilobidae and Sinuiceratidae), to which five genera belong: Aequilobus Bilotta (previously informally indicated as “genus from Monte Bove”), Dudresnayiceras Rakús, Sinuiceras Venturi and Ferri, and Sphenoacanthites Venturi, Nannarone and Bilotta, plus Xenoloboceras nov. gen., which position at the family level is uncertain. All of them share a peculiar combination of morpho-structural characters, which was never observed in any form referable to known groups. Due to the particularities presented by the Aequiloboidea and to the scarce documentation on the earliest Jurassic in the Mediterranean regions, it is currently impossible to demonstrate that this taxon originated within any of the already-known Jurassic ammonoid orders (Phylloceratida and Psiloceratida). An at least partially independent derivation cannot be excluded, as suggested also by the results of a cladistic analysis.     

Victims of the Early Toarcian anoxic event: the radiation and extinction of Jurassic Koninckinidae (Brachiopoda)

The significant mass extinction attributed to the Early Toarcian anoxic event had a severe impact on the phylum Brachiopoda. Beyond a serious decrease of species diversity, the extinction of the orders Spiriferidina and Athyridida is connected with this episode. The order Athyridida was represented by the family Koninckinidae in the Early Jurassic. The stratigraphical and geographical distribution of the three Early Jurassic koninckinid genera ( Koninckella, Koninckodonta, Amphiclinodonta ) shows a definite radiative pattern. The number of their nominal species increased from 2 to 17 from the Sinemurian to Early Toarcian; in the same time interval, their area increased from the Alpine region to the whole Mediterranean and the NW-European domains. This radiative evolution can be explained as the result of different factors: (1) morphological adaptation to muddy bottoms, (2) fundamental changes in the current pattern in the Tethys/Laurasian Seaway, and, possibly, (3) utilization of methane-based chemosynthesis as alternative food source. The radiation of koninckinids, leading from the cryptic habitats of the Tethyan rocky floors to the extensive muddy bottoms of the open European shelves, was abruptly terminated by the anoxic event in the Early Toarcian Falciferum Zone. The main causes of the extinction might be: (1) the excessive warming of Tethyan deep waters by thermohaline circulation, (2) the anoxic event, which was not survived by the spire-bearers, handicapped by their stiff, calcareous spiralia. Brachiopoda, Early Jurassic, Europe, extinction, Koninckinidae, radiation, Tethys.     

Brachiopod faunal exchange through an epioceanic-epicontinental transitional area from the Early Jurassic South Iberian platform system

The La Mola region (eastern External Betic Zone) can be regarded as one of the easternmost complete Jurassic successions of the Betic Cordillera in the Iberian Peninsula, but the paleogeographical setting of their outcrops remains widely discussed. Analysis of brachiopod assemblages from the Lower Jurassic improves the accuracy of previous paleogeographical data, enabling identification of a mainly epioceanic transitional area in which influences of epicontinental habitats are also detected. Assemblage 1, mainly with a Mediterranean affinity but also sharing several constituents with the African and Northeastern Iberian basins, typifies the Sinemurian–Pliensbachian transition. Assemblage 2, as a whole, shows a transitional character between epioceanic and epicontinental habitats; it is subdivided into two successive and interrelated sub-assemblages: Ass. 2a (Demonense–Tenuicostatum Zones) reveals a free connection with the epioceanic Subbetic area, whereas Ass. 2b (Uppermost Pliensbachian–Lower Toarcian) shows a closer relationship with epicontinental environments. Assemblage 3 (Uppermost Spinatum–basal Serpentinum Zones) is commonly recorded in the peri-Iberian epicontinental platform system integrated within the NW-European bioprovince, but it can also be regarded as a marginal assemblage that is widespread in the westernmost Tethyan margin prior to the Early Toarcian extinction event. Assemblage 2 constitutes a suitable index for assessing the paleobiogeographical affinity of the La Mola region, as Ass. 2a is progressively replaced by Ass. 2b, thus triggering the arrival of epicontinental taxa to the more intra-epioceanic Subbetic environments, inferring a possible connection through the La Mola transitional slope. Consequently, this region enabled a faunal mixing and exchange between both environments, and La Mola likely remained as an area that would facilitate migration and an effective dispersal seaway or, at least, did not constitute an ecological filter-barrier for brachiopods. Biostratigraphical data from brachiopods and ammonites are correlated for the first time in La Mola, refining and calibrating biochronostratigraphical gaps in the pre-Domerian deposits where biochronological markers are usually scarce, and around the Pliensbachian–Toarcian boundary, a crucial timespan in which the Early Toarcian extinction event took place.     

Toarcian (Lower Jurassic) brachiopods in Asturias (Northern Spain): Stratigraphic distribution, critical events and palaeobiogeography

Brachiopod assemblages recorded in the Toarcian outcropping of the Asturian coast, between Gijón and Ribadesella, are described. In the Tenuicostatum Zone of the Early Toarcian, an assemblage dominated by Gibbirhynchia cantabrica nov. sp., which also includes some other species of North European affinities, has been recorded. Brachiopods disappear in the region at the end of this Zone, coinciding with a sedimentary episode of black shales, and they are not recorded again until the Variabilis Zone of the Middle Toarcian. Between this zone and the Aalensis Zone, several species appear with wide stratigraphic distribution but scarce representation, except for Soaresirhynchia renzi, which is very abundant, particularly in the Insigne Subzone. This assemblage differs from the ones recorded in other nearby Spanish basins and can be related to the ones described in part of the Lusitanian Basin (Portugal), Western Pyrenees and South-Western France. This similarity can be related to environmental characteristics, such as the existence in these cases of an external, relatively deep platform environment.     

New Early Jurassic brachiopods from the Western Tethys (Eastern Subbetic, Spain) and their systematic and paleobiogeographic affinities

Two new brachiopod genera (Alebusirhynchia and Praesphaeroidothyris), five new species (Alebusirhynchia jorali, Salgirella? goicoecheai, Praesphaeroidothyris cisnerosi, Lychnothyris lancisi, and Cincta peiroi) and a new combination (Securina plicata) have been recorded and established after the internal and external morphological study of populations attributed to Lower Jurassic carbonate successions of the South-Iberian Paleomargin (SE Spain). The analysis made enables us to detail the diagnostic characters of little-known genera such as Lychnothyris or Securina and to clarify the systematics of the group from new supraspecific definitions, establishing a new, strongly multicostate rhynchonellid genus (Alebusirhynchia) and a new smooth terebratulid genus (Praesphaeroidothyris). The palaeobiogeographic signature of these taxa has been analysed, revealing remarkable Mediterranean affinities. Thus, the previously restricted geographical distribution of these faunas can be extended to the westernmost Tethys areas, where the maximum degree of endemism is established for the brachiopod groups in the Subbetic domain around the Sinemurian-Pliensbachian transition.     

Patterns of biotic change in Middle Jurassic to Early Cretaceous Tethyan radiolaria

The rate of taxic turnover of nearly 400 radiolarian species/subspecies is analyzed in order to document long term biotic change of plankton during the Middle Jurassic to Early Cretaceous (Aalenian to Aptian). The pattern and dynamic of diversity change is described using four indices: rate of species first and last occurrence, rate of diversification and rate of turnover. Plots of cumulative sampling effort suggest that the analyzed data represent an adequate sample of total standing diversity for most examined stages. Rates of species first occurrence exceed rates of last occurrence for most of the Middle Jurassic, except for the middle Bajocian. In contrast, the Late Jurassic was a time of decreasing radiolarian diversity and the Kimmeridgian records the lowest rate of diversification. It is followed by a dramatic increase in first occurrences near the Jurassic–Cretaceous boundary with as a result the highest rate of diversification recorded in the late Tithonian. Regional radiolarian diversity was stable throughout most of the Early Cretaceous. A stratigraphic permutation test was performed to assess the influence of uneven sampling on the observed pattern of taxic turnover and identified the intervals for which randomly obtained patterns are significantly different from the observed pattern. The Kimmeridgian and late Tithonian events coincide with substantial climate-derived perturbations in water cycling, nutrient supply and oceanic productivity. They point to a negative relationship between radiolarian macroevolution and changes in the state of nutrient availability, although further work is needed to refine the temporal resolution of this relationship and to explore ecological aspects of its causal link with respect to radiolarian evolution.     

云南早侏罗世一新具嵴冠兽脚类恐龙

报道了具嵴冠的兽脚类恐龙一新属种,安龙堡双柏龙(Shuangbaisaurus anlongbaoensis gen.et sp.nov.).双柏龙发现于云南省楚雄彝族自治州双柏县下侏罗统冯家河组中,保存了部分头骨带下颌.双柏龙沿两侧眼眶背缘向上有嵴冠发育,这在其他兽脚类中未曾报道过.与其他早侏罗世体型较大且具一对矢状嵴冠的兽脚类(双嵴龙属和中国龙属)相比,双柏龙还独具一些特征组合,如相对较高的前颌骨体、抬高的前颌骨腹缘、后腹向延伸的轭骨后突及较小的上颞孔.比较研究表明,尽管中国“双嵴龙”(“Dilophosaurus”sinensis)可能应归入中国龙属(Sinosaurus),但未必属于模式种.双柏龙的发现将有助于研究基干兽脚类的演化,尤其是各类头骨骨饰在其中的作用.     

Spatial distribution of Late Pliensbachian (Early Jurassic) calcareous nannofossils within the Lusitanian basin (Portugal)

Quantitative analyses of Pliensbachian calcareous nannofossils have been carried out on a proximal-distal transect in the Lusitanian Basin (Portugal). The studied sections Vale Venteiro near Tomar and Peniche represent proximal and distal environments with respect to the emerged land of the Iberian Meseta (to the East). The upper portion of the Vale das Fontes Fm (Davoei and Margaritatus ammonite Zones) is studied in both sections and correlated by means of ammonite and nannofossil integrated biostratigraphy. A careful analysis of the preservation state of nannofossils is performed. Preservation state is moderate to good in the two settings; changes in nannofossil assemblages are therefore considered as primary. Samples were analysed for nannofossil absolute and relative abundances, species diversity and wt%CaCO₃. The analysis of calcareous nannofossils (absolute abundance, percentage, average abundance) shows changes in the community structure in space (i.e., from proximal to distal), and vertically within the water column. This pattern suggests a partitioning of habitats within the photic zone, and with respect to emerged lands. Proximal environments within the Lusitanian Basin were probably more eutrophic, because of the proximity of emerged lands to the East (Iberian Meseta). This was the source area for nutrients delivered into the basin by river transport. Distal environments were likely characterized by a greater water depth and by a deeper light penetration leading to a relatively more expanded photic zone. The assemblages recorded in the proximal setting exhibit high mean relative abundance of placolith-bearing coccolithophorids (Lotharingius spp., Biscutum spp. and Similiscutum spp.) and of Schizosphaerella spp., while distal assemblages are dominated by Crepidolithus crassus and Schizospharella spp. Crepidolithus crassus is interpreted as a deep-dweller coccolithophorid, inhabiting preferentially distal and deeper settings in the Lusitanian Basin. Placolith-bearing coccolithophorids were more abundant in proximal settings with respect to emerged lands, under relatively elevated trophic conditions. The probable calcareous dinocyst Schizospharella spp. proliferated in surface waters of both proximal and distal environments.     

Revision and re-evaluation of the Early Jurassic dinosaurian ichnogenus Otozoum

Otozoum moodii Hitchcock, 1847 is one of the classic Connecticut Valley ichnotaxa established by Edward Hitchcock between 1836 and 1865. The taxon is redescribed, and AC 4/1a re-established as the holotype specimen. Otozoum minus and O. caudatum are synonymised with O. moodii . The syntypes of Kalosauropus pollex ( nomen nudum ) are described as O. pollex sp. nov. Cladistic, quantitative, and comparative methods of trackmaker identification suggest that Otozoum was made by a prosauropod dinosaur. In quantitative analyses, only those phalanges likely to contribute to footprint morphology were considered; claws were also excluded due to their highly variable styles of impression. Several diagnostic characters distinguish Otozoum from Brachychirotherium , Chirotherium , Batrachopus , Tetrasauropus , and Pseudotetrasauropus ; these ichnotaxa are probably attributable to crurotarsan archosaurs. Otozoum is found primarily in Hettangian strata of the Hartford, Deerfield, and Fundy rift basins of eastern North America; it is also present in the Early Jurassic Navajo Sandstone (Colorado Plateau, USA) and Clarens Formation (Lesotho). The taxon is unknown in older strata; Triassic material previously referred to Otozoum (including O. grandcombensis ) instead belongs in Pseudotetrasauropus . Correlation of sediments and comparison of faunal assemblages, particularly within the Newark Supergroup, suggest that the Otozoum trackmakers were restricted to arid environments.     

Stalked crinoids from a Jurassic tidal deposit in western North America

This is the first systematic and paleoecological study of a crinoidal limestone (encrinite) from the Jurassic System of North America. The encrinite is part of a shallow-water tidal facies of the Middle Jurassic Carmel Formation located at Mount Carmel Junction (southwestern Utah, U.S.A.) and may represent one of the youngest shallow-water encrinites in the geological record. In the past, the crinoid at this locality was referred to as Pentacrinus asteriscus, a name used to describe almost all of the crinoid columnals found throughout the Jurassic of the U.S. western interior. However, systematic work indicates that the crinoid is Isocrinus nicoleti and is the first non-endemic crinoid to be reported from North American Jurassic strata. Although articulated pinnules and arms have been found, I. nicoleti occurs predominantly as well-preserved, partially articulated columnals. The crinoids occur within a tidal complex consisting of ooid shoal, tidal channel, and lagoonal facies. The unique environmental and ecological conditions which existed in the southernmost end of the Jurassic North America seaway may have allowed for the development of this crinoid colony and subsequent deposition of the encrinite.     

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.     

Evolutionary patterns in the terebratulid genus Caryona Cooper (Brachiopoda,Jurassic of France)

The study of ontogenetic and morphological changes in different species of the brachiopod genus Caryona COOPER (Terebratulidae) from the Lower Callovian‐Lower Oxfordian of northern France shows a succession (and an interference) of processes which vary according to the characters studied. This is an example of “mosaic evolution”;. The ontogenetic changes result from heterochrony, especially acceleration and hypermorphism. The main phylogenetic tendency is peramorphosis in most characters. This evolutionary pattern agrees with phyletic gradualism.

Moreover, analysis of the internal characters demonstrates the limited value of these in discrimination of the various genera.     

A new genus and species of fimbriid bivalve from the Kimmeridgian of the western Pontides, Turkey, and the phylogeny of the Jurassic Fimbriidae

A new lucinoidean genus and species, Cerkesia contiae gen. et sp. nov., from the Kimmeridgian carbonate platform deposits of the Çerkes area (western Pontides, Turkey), is described and placed in the family Fimbriidae. Morphological comparisons reveal similarities with Fimbria , Cyclopellatia and Sphaera . A phylogenetic analysis supports the monophyly of the family. Similarities between Cerkesia and Fimbria determine inclusion of these genera in the same clade. The unusual features characterizing Cerkesia closely relate it to Cyclopellatia . Sphaera belongs to a different clade and its similarities with Cerkesia are owing to convergent and plesiomorphic characters. Cerkesia was a slow and shallow burrower. The morphology of the anterior adductor scar suggests a mode of life comparable to that of Fimbria but other features indicate adaptive strategies typical of the Lucinidae. The associated fossils and facies reveal that Cerkesia lived in soft and sandy substrates of quiet and protected areas of the carbonate platform under stressful environmental conditions. The appearance of Cerkesia in the western Pontides marks the eastward expansion of a clade that developed principally in the European seas during the Late Jurassic while the Sphaeriola-Sphaera clade was confined to the southernmost Tethyan margin.     

Seafloor erosion and sea-level change: Early Jurassic Blue Lias Formation of central England

Minor bedforms within the mudstone-dominated Early Jurassic Hettangian Saltford Shale Member (Liasicus up to Angulata Chronozone) of the Blue Lias Formation in central England, indicate weak seafloor erosion in a mid to outer ramp setting. Distal storm flows below maximum storm wave base are proposed as the most likely generative mechanism for silty scour and gutter casts that enclose concentrations of well-preserved schlotheimiid ammonites and arthropod trace fossils. Within the upper part of the Saltford Shale (probably Angulata Chronozone), a discrete layer of reworked and bioencrusted limestone nodules signifies an episode of more persistent seafloor erosion. The immediately overlying strata, transitional to the Hettangian–Sinemurian Rugby Limestone Member, are relatively bioturbated and feature fossils of macrobenthos, as well as shell concentrations resembling relatively proximal storm beds. This suggests that the reworked nodule horizon marks sea-level fall, rather than stratigraphic condensation associated with sediment starvation. The biostratigraphic evidence raises the possibility that this erosional episode correlates with a mid-Angulata Chronozone hiatus documented from the Wessex Basin, southwest England. Equally however, it could be linked to contemporaneous movement on one or more nearby faults, affecting the southern part of the English East Midlands Shelf.     

The cranial morphology of Kayentachelys,an Early Jurassic cryptodire,and the early history of turtles

Gaffney, E.S. and Jenkins, F.A., Jr. 2010. The cranial morphology of Kayentachelys, an Early Jurassic cryptodire, and the early history of turtles. — Acta Zoologica (Stockholm) 91 : 335–368 The skull morphology of Kayentachelys aprix Gaffney et al., 1987 , a turtle from the Early Jurassic Kayenta Fm of northern Arizona, demonstrates the presence of cryptodiran synapomorphies in agreement with Gaffney et al. (1987, 1991, 2007) , and contrary to the conclusions of Sterli and Joyce (2007) , Joyce (2007) , Sterli (2008) , and Anquetin et al. (2008) . Specific characters found in Kayentachelys and diagnostic of cryptodires include the processus trochlearis oticum, the curved processus pterygoideus externus with a vertical plate, and the prefrontal–vomer contact, which are confirmed as absent in the outgroups, specifically the Late Triassic Proganochelys. The Joyce (2007) analysis suffers from the reduction of the signal from skull characters, with a consequently greater reliance on shell characters, resulting in pleurodires being resolved at various positions within the cryptodires. Kayentachelys reveals what a primitive cryptodire would be expected to look like: a combination of primitive and derived characters, with the fewer derived characters providing the best test of its relationships to other turtles. Although incompletely known, the Mid‐Late Jurassic Condorchelys, Heckeremys, and Eileanchelys may be early cryptodires close to Kayentachelys. We confirm the Late Triassic Proterochersis as a pleurodire, dating the pleurodire–cryptodire split as Late Triassic or earlier.     

Jurassic radiolarites in a Tethyan continental margin (Subbetic, southern Spain): palaeobathymetric and biostratigraphic considerations

A region of the pelagic Subbetic basin within the Southern Iberian Continental Margin is studied in lithostratigraphical and biostratigraphical detail. Jurassic radiolarites (Jarropa Radiolarite Formation, Bathonian–Oxfordian) interbedded with shallow-water marine limestones have been recognized. Underlying the radiolarites (Camarena Formation, Bajocian) are oolitic limestones showing shallowing-upward cycles with karstic surfaces on the top, corresponding to deposition on an isolated carbonate platform on volcanic edifices. The Milanos Formation (upper Kimmeridgian–Tithonian), overlying the radiolarites, contains calciclastic strata with hummocky cross-stratification, which indicate outer carbonate ramp deposition. In the Jarropa Radiolarite Formation some calcisiltite strata with hummocky cross-stratification have been found. The bathymetry of the Subbetic Jurassic pelagic sediments, including the radiolarites, is considered as moderate or shallow in depth. We suggest that the pelagic character of the Jurassic sediments in this margin and their equivalents in other Alpine domains is a consequence of distance from the continent (beyond the pericontinental platform) but not necessarily of depositional depth.     

First Early Jurassic Ornithischian and theropod footprint assemblage and a new ichnotaxon Shenmuichnus wangi ichnosp. nov. from Yunnan Province,southwestern China

The new ichnospecies, Shenmuichnus wangi ichnosp. nov., is the first evidence for the presence of large ornithischians in the Early Jurassic of Yunnan Province, whereas the known skeletal record documents small species only. Until now Shenmuichnus was known from a single locality in Shaanxi Province by the ichnospecies Shenmuichnus youngteilhardorum. Compared with the latter, Shenmuichnus wangi is larger and shows a different trackway configuration, particularly in the relative position of manus and pes imprints. Palecologically, the occurrence of Shenmuichnus wangi in a red bed facies indicates the preference of distinctive environments of trackmakers of both ichnospecies, questioning former hypotheses of exclusivity of ornithischians in more humid climates. By abundance both skeletons and footprints of ornithischians suggest their role as a minor component in Early Jurassic saurischian dominated dinosaur faunas in this region.     

Some Fossil Plants from Early Jurassic in Western Hubei With the Relative Problem Between Fossil Plants and Coal-Bearing

Coniopteris magnifica sp. nov., Ctenophyllum chenyuanense sp. nov., C. laxilobum sp. nov., Mironeura dakengensis Zhou, are derived from the Xiangxi Formation (s.s.), Western Hubei. According to the aspect and characteristic of the Xiangxi flora, author considers the Xiangxi Formation as Early Jurassic in age. In this paper, the relative problem between the coal-bearing and the fossil plants is briefly discussed also.