The two Early Toarcian (Early Jurassic) extinction events in ammonoids

The Early Toarcian (Early Jurassic) biological crisis was one of the ‘minor’ mass extinctions. It is linked with an oceanic anoxic event. Fossil data from sections located in northwestern European (epicontinental platforms and basins) and Tethyan (distal, epioceanic) areas indicate that Late Pliensbachian–Early Toarcian ammonoids experienced two extinction events during the Early Toarcian. The older one is linked with disruption of the Tethyan–Boreal provinciality, whereas the younger event correlates with the onset of anoxia and corresponds with the Early Toarcian mass‐extinction event. These two extinctions cannot be interpreted as episodes of a single, stepwise, event. Values of the net diversification, more than the number of extinctions, allow the two extinction events to be clearly recognized and distinguished. Values of regional net diversification for northwestern European and Tethyan faunas point to greater evolutionary dynamics in the epioceanic areas. The inclusion of Mediterranean faunas in the database proves that the ammonite turnover at the Early Toarcian mass‐extinction event was more important than previously thought. Progenitor (evolute Neolioceratoides), survivor (Dactylioceras, Polyplectus pluricostatus) and Lazarus (Procliviceras) taxa have been recognized. Different selectivity patterns are shown for the two events. The first one, linked to the disruption of the Tethyan–Boreal provinciality, has mainly affected ammonites adapted to epicontinental platforms. In the mass‐extinction event, no selectivity is recognized, because also Phylloceratina and Lytoceratina were deeply affected at species level, although their wide biogeographical distribution at clade level was a significant buffer against extinction. In contrast to Palaeozoic mass extinctions, ammonoid survivors and Lazarus taxa are characterized by complex sutures: Phylloceratina (long‐ranging ammonoids) and Polyplectus (relatively long‐ranging compared to other Ammonitina).     

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.     

Multivariate hierarchical analyses of Early Jurassic Ostracoda assemblages

Palaeobiogeographic patterns of Early Jurassic ostracods from the northern and southern hemispheres (96 sections located in Europe, North Africa, Western Australia and North and South America) based on 243 species-level records document global patterns of distribution that can be compared to those previously published on ostracods from the European Epicontinental Sea and Tethyan and South Panthalassa areas. All described records of ostracods from both hemispheres spanning the Hettangian to Early Toarcian have been compiled and verified, and their patterns of origin and distribution have been interpreted. Jaccard coefficient of similarity was used to asses similarities among European, American and Tethyan ostracod shelf faunas. The numerical analysis shows a progressive longitudinal gradient in provincialism through the Early Jurassic, consistent with the northward drift of Tethyan ostracod faunas towards the European Epicontinental Sea and the southward movement of European taxa into Tethys and Panthalassa oceans. The spread of cosmopolitan species and extinction of endemic species, allied to the disappearance of geographical barriers, warmer climate conditions and rising sea levels can explain the reduction in ostracod diversity and the east-west provincialism throughout the Early Jurassic. Interchange between hemispheres, including bipolar distributions, are recognized from the Sinemurian time, pointing out that for most of the studied period, the climate worldwide was warm and tropical.     

特提斯喜马拉雅地区早侏罗世双壳类Lithiotis生物岩礁及其扩散

藏南聂拉木地区首次发现的早侏罗世Ｌｉｔｈｉｏｔｉｓ生物岩礁,证明特提啼马拉雅地块于早侏罗世后期已开始向北运移。并且至少在早托尔斯已接近的或达到能够适应Ｌｉｔｈｉｏｔｉｏｓ繁衍的低纬度区域。     

Biodiversity and biogeography of middle–late liassic ammonoids: implications for the early Toarcian mass extinction

In Western Tethyan areas, the Toarcian stage begins with two important evolutionary events in ammonite faunas: (1) the disruption of Tethyan–Boreal provinciality; (2) a biological crisis linked with the oceanic anoxic event OAE. The analysis of these events has been addressed by constructing curves of ammonoid diversity (species richness, origination and extinction rates) in the Late Pliensbachian (= Domerian)–Early Toarcian interval in selected localities. Two diversity drops are recognized. The first one is recorded at the end of the Dactylioceras mirabile subzone and reflects the disruption of Tethyan–Boreal provinciality, through the progressive extinction of the Boreal endemic family Amaltheidae that occupied the north-western European seas during the whole Pliensbachian on the one hand, and the extinction of Late Domerian Ammonitina endemic to the Mediterranean areas on the other hand. The Early Toarcian homogeneization of Mediterranean and north-western European ammonoid faunas was reached via elimination of both Boreal and Mediterranean endemics with differential rates of extinction in the two palaeogeographic domains and the subsequent geographical expansion of Tethyan-derived ammonoids. The second, dramatic drop in ammonite diversity in the upper part of the Dactylioceras semicelatum subzone coincided with the onset of OAE. It also affected epioceanic ammonoid clades like Phyllocerataceae and Lytocerataceae. These two drops are interpreted as two distinct extinctions and not as episodes of a single, stepwise event. Complex relations between ammonoid diversity and sea-level changes are suggested by trends in endemism, which may be reversed during either a single transgression or a single regression.     

Pleurotomaria DeFrance, 1826 (Gastropoda,Mollusca) from the Lower Bajocian (Middle Jurassic) sediments of Luxembourg,with considerations on its systematics,evolution and palaeobiogeographical history

Abstract: Pleurotomaria species from lower Bajocian (Middle Jurassic) sediments of south‐western Luxembourg housed in the National Natural History Museum of Luxembourg are described. Seven species are recognized, one of which is new, Pleurotomaria faberi sp. nov. A more detailed definition of the diagnostic characters of the genus is proposed and the morphological continuity between Talantodiscus and Pleurotomaria is demonstrated, suggesting that the former cannot be considered as a distinct taxon. The palaeoecology, evolution and palaeobiogeographical history of Pleurotomaria are outlined. Pleurotomaria presumably first appeared in late Middle Triassic of New Zealand where it underwent a relative diversification up to the Hettangian (Early Jurassic). From early Hettangian, most of its evolutionary history took place in Europe and western Tethys. In the European epicontinental seas, Pleurotomaria experienced two important radiations. The first occurred in the Early Jurassic, with a peak in the late Pliensbachian, and was marked by an expansion of the distribution to the central part of western Tethys. After a collapse in species diversity, probably related to the early Toarcian anoxic event, a second radiation occurred. This culminated in the early Bajocian and was mainly confined in a region encompassing southern England, Paris Basin and southern Germany. Low‐spired species, formerly attributed to Talantodiscus, probably originated independently and iteratively during the history of Pleurotomaria. The facies and associated benthic faunas suggest that Pleurotomaria probably lived on shallow soft bottoms composed of mixed calcareous–siliciclastic sediments. The two main Early Jurassic and early Middle Jurassic radiations of the genus took place in these environments. Records of the genus in Jurassic carbonate platform deposits are very few and concern mainly post‐Bajocian species.     

Microbes,mud and methane: cause and consequence of recurrent Early Jurassic anoxia following the end‐Triassic mass extinction

The end‐Triassic mass extinction (c. 201.6 Ma) was one of the five largest mass‐extinction events in the history of animal life. It was also associated with a dramatic, long‐lasting change in sedimentation style along the margins of the Tethys Ocean, from generally organic‐matter‐poor sediments during the Triassic to generally organic‐matter‐rich black shales during the Jurassic. New core material from Germany provides biomarker evidence of persistent photic‐zone euxinia during the Hettangian, the onset of which is associated with a series of both negative and positive carbon isotope excursions. Combined inorganic and organic geochemical and micropalaeontological analyses reveal strong similarities between the Hettangian and the better‐known Toarcian anoxic event. These events appear to be the most clearly expressed events within a series of anoxic episodes that also include poorly studied black shale intervals during the Sinemurian and Pliensbachian. Both the Hettangian and Toarcian events are marked by important changes in phytoplankton assemblages from chromophyte‐ to chlorophyte‐dominated assemblages within the European Epicontinental Seaway. Phytoplankton changes occurred in association with the establishment of photic‐zone euxinia, driven by a general increase in salinity stratification and warming of surface waters. For both events, the causes of large negative carbon isotope excursions remain incompletely understood; evidence exists for both variation in the δ¹³C of atmospheric CO₂ and variation in the sources of organic carbon. Regardless of the causes of δ¹³C variability, long‐term ocean anoxia during the Early Jurassic can be attributed to greenhouse warming and increased nutrient delivery to the oceans triggered by flood basalt volcanism.     

Extinction pattern of marine Ostracoda across the Pliensbachian-Toarcian boundary in the Cordillera Ibérica, NE Spain: Causes and consequences

This paper discusses the extinction pattern of the Pliensbachian-Toarcian boundary (PTB) ostracod assemblages at the Almonacid de la Cuba section (Cordillera Ibérica, NE Spain), which has been recently proposed as auxiliary boundary stratotype for the PTB. The ostracod record shows that the main Early Jurassic ostracod extinction event occurred not at the end of the Pliensbachian, but near the top of the Mirabile ammonite Subzone, Tenuicostatum ammonite Zone (Early Toarcian). On the basis of the evaluation of PTB ostracod record, a new causal explanation for the Early Toarcian ostracod turnover is proposed. This paper suggests that a reorganization of surface and deep-water circulations caused by the opening of the Hispanic Corridor could have generated a mild cooling episode, finally affecting the survival of healdioid ostracods.     

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.     

First record of a gladius-bearing coleoid Teudopsis bollensis Voltz (Cephalopoda, Coleoidea) in the Toarcian of the Western Carpathians (Slovakia)

A fairly complete and relatively well-preserved gladius of the vampyropod coleoid cephalopod Teudopsis bollensis Voltz is recorded from the Lower Toarcian succession of the Kysuca Unit in the Pieniny Klippen Belt of Slovakia. These sediments are represented by dark-grey and black shales laid down in an oxygen-depleted environment during the Toarcian Oceanic Anoxic Event and potentially represent the first Jurassic ??Konservat-Lagerst?tte?? in the Western Carpathians. The first reliable record of the Early Jurassic genus Teudopsis from this area notably extends its palaeogeographic distribution to the northwestern continental margin of the Tethys Ocean. The systematics, palaeogeography and stratigraphy are briefly discussed in a European context.     

Relationships between the early Toarcian anoxic event and organic-walled phytoplankton in central Italy

The integration of palynological and geochemical data from three lower Toarcian successions in central Italy reveals that the composition of organic-walled phytoplankton assemblages were strongly affected by palaeoecological conditions related to bituminous sedimentation which accompanied the global anoxic event. The marked compositional variations of dinoflagellate cysts and prasinophytes, together with geochemical variations, have been linked to changes in surface water habitats during the lower Toarcian transgression. On the basis of the relationships between total organic carbon (TOC) and marine palynomorph assemblage composition, the lower Toarcian evolution of the Umbria-Marche Basin, central Italy, has been divided into four phases. Total organic carbon values rose significantly during the early Toarcian (Lower-middle Dactylioceras tenuicostatum ammonite Zone), and this can be linked to certain dinoflagellate cyst datums, for example the temporary disappearance of Mancodinium semitabulatum and the extinction of Luehndea spinosa. The presence of Umbriadinium mediterraneense and Valvaeodinium spp. accompany these moderately high TOC values. Subsequently, TOC levels increased to over 2% and prasinophytes became abundant in the Middle-upper D. tenuicostatum ammonite Zone. Mancodinium semitabulatum reappeared when TOC values eventually decreased in the Upper D. tenuicostatum ammonite Zone. This analysis has allowed the different sunlight requirements and life strategies of the early Toarcian Tethyan dinoflagellates to be modelled. Due to the cosmopolitan nature of the early Toarcian anoxic event, the principal marine palynological signals observed have been interpreted as sequence stratigraphical and palaeoecological indices. The Transgressive Systems Tract (TST) is accompanied by an increase in dinoflagellate cyst species diversity and a decrease in abundance. The succeeding maximum flooding surface (mfs) corresponds with a prasinophyte acme. During the Highstand Systems Tract (HST), the phytoplankton shows an increase in abundance and a decrease in diversity. The range top of Luehndea spinosa appears to characterise the early Toarcian TST.     

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.     

Paleobiogeography of Mesozoic brachiopod faunas from Andean–Patagonian areas in a global context

During the Mesozoic, the Andean region has played a hinging role between high- and low-latitude faunas, which are, respectively, characterized by stocks that display long-term fidelity. This paper is aimed at providing an updated review of Late Triassic to Late Cretaceous South American articulated brachiopods in the light of previous knowledge at worldwide scale. Late Triassic brachiopods from the Argentine–Chilean Andes show unmistakable Maorian (or Notal) faunal elements alongside some more cosmopolitan genera, with certain influence of Eastern Pacific taxa. By Early Jurassic times, differentiation of Tethyan and Boreal Realms became progressively evident in Europe. In South America, Hettangian–Sinemurian brachiopod faunules from the Argentinian Andes are somewhat impoverished, with mostly cosmopolitan genera showing certain affinities to Maorian species, and with the addition of some endemics later. Increasingly, diverse Pliensbachian Andean brachiopods denote close relationships to Celto-Swabian taxa, then by Domerian times, a certain degree of endemism was developed, though somewhat delayed Tethyan influences, and persistent links with New Zealand are subordinately recognizable, too; most Toarcian assemblages reveal basically Celto-Swabian and Iberian affinities as well. East-west austral links across the Pacific may have been favored by migratory routes fringing the Gondwana margin, whereas faunal exchange with the western end of the Tethys appears to reflect an intermittent shallow-marine connection through the Hispanic Corridor. During the Middle Jurassic, distinction of Tethyan and Boreal Realms was maintained in the northern Hemisphere, and the differentiation of an Ethiopian or Southern Tethyan fauna became better characterized. Aalenian and Bajocian brachiopods of the Andes display generic affinities mainly with those from western Europe, with some minor endemic developments; brachiopods recorded from the Bathonian–Callovian of Argentina (and Chile) also occur along the northern Tethyan margin, yet with some genera extending into Indo-Ethiopian areas. During the Late Jurassic, Boreal faunas from high-latitudes became even more strongly differentiated from low-latitude, Tethyan ones. Oxfordian and Tithonian brachiopods from the Andes apparently belong to genera of cosmopolitan or northern Tethyan affiliation, yet there are few elements in common with other eastern Pacific areas, such as Mexico. Early Cretaceous brachiopods, in addition to Andean basins of Chile and western Argentina, are known also from Patagonia and Tierra del Fuego. They belong mostly to widely distributed, mainly Tethyan genera, with some quasi-cosmopolitan and circum-Pacific components (some shared with Antarctica become noticeable). Late Cretaceous brachiopods from northern Patagonia show significant affinities to Maastrichtian ones of northwest Europe and central Asia, which calls for further assessing the potential role that may have played the trans-Saharan passageway in such dispersal. Broad aspects of Mesozoic brachiopod paleobiogeography are fairly well understood, yet details of ranking and naming of certain units are still in need of more agreement.     

Uppermost Pliensbachian-Lowermost Toarcian ostracod assemblages from the Western Tethys: Comparison between Traras Mountains (Algeria), Subbetic (Spain), and Algarve (Portugal)

Major changes in ostracod assemblages have been recorded in the Lower Jurassic of Western Tethys related to biotic crises of the Pliensbachian-Toarcian boundary and the Polymorphum-Levisoni boundary (Toarcian Oceanic Anoxic Event). The study is focused on the ostracods assemblages from Benzerka section (Traras Mountains, Algeria) and their comparison with sections from External Subbetic and Algarve Basin (South Iberian Palaeomargin). Three stratigraphic intervals coincident with the ammonite zones have been differentiated with important fluctuations in the composition of ostracod assemblages. In the ostracod assemblages from the Traras Mountains the turnover related to this benthic biotic crisis is shown by assemblages previous to the biotic crisis (uppermost Pliensbachian) mainly composed by families Healdiidae and Cytherellidae to assemblages just after the T-OAE (upper part of the Levisoni Zone) with assemblages dominated by families Protocytheridae and Polycopidae and the extinction of family Healdiidae (suborder Metacopina). The Toarcian Oceanic Anoxic Event (T-OAE) probably resulted in restricted conditions but a barren interval due to oxygen-depleted conditions in this setting is not recorded. In this context, the change of components of ostracod assemblages looks gradual. In the Southiberian Palaeomargin, represented by the External Subbetic the turnover of ostracods assemblages is also recorded but the families involved are different than in Traras Mountains. Therefore, different phases have been differentiated in the ostracod turnover occurring from Upper Pliensbachian (assemblages dominated by families Pontocyprididae, Healdiidae and Bairdiidae) to end of Lower Toarcian (assemblages dominated by family Cytherellidae and the extinction of family Healdiidae). In the Algarve Basin the families Healdiidae and Bairdiidae were dominant previously to the Toarcian Oceanic Anoxic Event. Differences in the composition of ostracod assemblages between Traras Mountains and External Subbetic may be related to the fragmentation of these palaeomargins related to early stages of rifting in the Western Tethys. Moreover, the incidence of oxygen-depleted conditions was higher in External Subbetic than in Traras Mountains where some ostracod genera survive during the biotic crisis.     

Morphology and environment in the Jurassic Nucleatidae (Brachiopoda) from Western Tethys

Colás, J. & García Joral, F. 2011: Morphology and environment in the Jurassic Nucleatidae (Brachiopoda) from Western Tethys. Lethaia, Vol. 45, pp. 178–190. Nucleatidae (Brachiopoda, Terebratulida) are commonly considered as typical members of the Mediterranean assemblages during the Jurassic. However, nucleatids occasionally also occur in the margins of Western Tethys (Northwestern European shelf). Some of these occurrences in northeastern Spain are analysed, detecting a relation between nucleatid morphology and palaeoenvironment during the Jurassic. The validity of this relationship for the whole of the representatives of the group in the Western Tethys during the Jurassic is tested by means of morphofunctional analyses (principal components analysis and discriminant analysis), concluding that reduction of the lateral expansion of the shell is the key morphological feature whereby epioceanic and epicontinental taxa differ. This change can have functional significance because it can be interpreted as a mechanism to reduce the area exposed when the valves gape, increasing protection against harmful particles in environments characterized by higher terrigenous input. Some modifications on the systematics of the Nucleatidae at the genus level are proposed on the basis of the temporal and biogeographical distribution of these adaptations. □Brachiopods, Nucleatidae, functional morphology, adaptation, systematics, Jurassic, Spain.     

Pliensbachian gastropods from Venetian Southern Alps (Italy) and their palaeobiogeographical significance

Abstract: The Pliensbachian gastropods described by De Toni in 1912, coming from an isolated boulder at the foot of Mt Vedana (eastern margin of Trento Platform, Venetian Southern Alps, Italy) are revised. The fauna consists of 13 species representing nine families and eight superfamilies. Despite the low number of species, the assemblage represents the most diverse Early Jurassic gastropod fauna known for the Venetian Southern Alps. The boulder yielding the material was thought to derive from the upper part of the Early Jurassic Calcari Grigi Group, a carbonate platform unit extensively cropping out in the Mt Vedana area. The sedimentological analysis indicates a prevalently bioclastic wackstone‐floatstone, reflecting a lime‐muddy deposit undergone to an early consolidation. This and the high content of ammonoids, which is unusual for the Calcari Grigi Group, are typical aspects of a condensed pelagic sediment, presumably a fissure filling at the top of the carbonate platform succession. Palaeobiogeographical comparisons show that the fauna is composed of species occurring exclusively in pelagic limestones of the western Tethys. By contrast, it shows no relationships with the coeval faunas of the adjacent Trento platform and of the other western Tethyan carbonate platforms. These lines of evidence and the facies analysis would testify to the Pliensbachian drowning of the eastern margin of the Trento platform. In the wider context of the palaeobiogeographical history of Early Jurassic western Tethyan gastropods, the species from Vedana belong to a faunal stock which is typical for pelagic, mainly postdrowning sediments. Thus, appearance and diffusion in space and time of this stock were probably regulated by the direction, rate and pattern of the Neotethyan rifting. A new subgenus, Proarcirsa (Schafbergia) subgen. nov., and three new species, namely Ataphrus (Ataphrus) cordevolensis sp. nov., Guidonia pseudorotula sp. nov. and Proarcirsa (Schafbergia) zirettoensis sp. nov. are erected.     

Diversity and density characteristics of Pliensbachian-Toarcian molluscan and brachiopod faunas of the North Atlantic margins

Pliensbachian and Toarcian molluscs and brachiopods of Morocco, Iberia, England, and Greenland are analysed in terms of diversity and density. With regard to local variations, Substrate and food supply are thought to be the major factors controlling the predominance of suspension-feeding bivalves in shallow water deposits while the greater physical stability of deeper water favoured the ammonites; high diversity of the gastropods relates to firm substrates. On a regional scale, both bivalve density and diversity tend to increase from the Tethyan to the Boreal redm, while there is a Variable tendency for a northward diversity reduction in other groups. The bivalve data are incompatible with either temperature or salinity control of faunal provinciality. To provide a better account of this, a model is developed which lays emphasis on the physical instability of a boreal inland sea as compared with the Tethys. Some broader implications for endemism in Jurassic faunas are briefly discussed.     

Controls on migration of conodont fauna in peripheral oceanic areas. An example from the Middle Triassic of the Northern Peri-Tethys

Middle Triassic conodont assemblages in the semi-closed Peri-Tethyan Germanic Basin comprise endemic forms, which evolved within the basin, and immigrants from the Tethys Ocean. Migration from the Tethys proceeded diachronously through three tectonically predisposed pathways (gates) and was controlled by relative sealevel changes recorded in depositional sequences of the Muschelkalk. In the Early and Middle Anisian predominant were the forms that originated in the Eastern Tethys and in southern basins of the Western Tethys. They migrated through the East Carpathian Gate and Silesian-Moravian Gate. In the late Anisian and early Ladinian forms from the Western Tethyan basins prevailed; their migration pathway led through the Western and East Carpathian Gates. The latter gate was active only at the turn of Illyrian and Fassanian. Episodes of migration of the Tethyan forms and dispersal of conodont fauna throughout the German Basin coincide with the transgressive or highstand phases of a particular depositional sequence (as defined by Szulc, 1999). The most significant proportion of the Tethyan species appears to correlate with the maximum flooding surface of the A₃ sequence, (Lower Muschelkalk) later they decrease in diversity. In the Upper Muschelkalk the most numerous Tethyan species correspond to the transgressive phase of the L₁ sequence. Above the mfs of the L₁ sequence only the Germanic forms remain.