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.     

青藏高原白垩纪双壳类生物地理

青藏高原白垩纪沉积见于６条东西向延伸的条带内。双壳类主要发育于这些条带的Ｂｅｒｒｉａｓｉａｎ,Ａｐｔｉａｎ－Ａｌｂｉａｎ,Ｃｅｎｏｍａｎｉａｎ－Ｔｕｒｏｎｉａｎ,Ｃｏｎｉａｃｉａｎ－Ｓａｎｔｏｎｉａｎ和Ｃａｍｐａｎｉａｎ－Ｍａａｓｔｒｉｃｈｔｉａｎ５个时期的地层。雅鲁藏布江缝合线为白垩纪双壳类地理分布的主要控制界线。早白垩世期间,雅鲁藏布江缝合带以南的喜马拉雅地区的双壳类Ｐｅｔｒｏｃｅｒａｍｕｓ,     

The Early Cretaceous of North-East Greenland: A crossroads of belemnite migration

Recent field work in Lower Cretaceous successions of Traill Ø and Wollaston Forland, North-East Greenland, have resulted in c. 2350 belemnite guards collected bed-by-bed from the upper Ryazanian – Hauterivian. The most common belemnite genera observed, Acroteuthis, Pachyteuthis, and Cylindroteuthis are of boreal-arctic affinities and closely related to NW European and Siberian faunas. Other taxa, including Hibolithes (common), Pseudobelus (relatively common) and Duvalia (rare), show faunal links to both NW European and Mediterranean faunas. This paper describes and discusses these findings in their taxonomic, biostratigraphic, palaeobiogeographic, palaeoecologic and palaeoceanographic context. In particular, the occurrence of Pseudobelus which is common in the circum Mediterranean area, is remarkable since it is the first observation of this Tethyan genus in the entire Boreal Realm. The palaeoecological interpretation of these observations result in the recognition of four different palaeobiogeographic belemnite assemblages for the Boreal Realm: 1) North-East Greenland, 2) Spitsbergen, 3) NW Europe and 4) Siberia. In contrast to the other assemblages, the belemnite faunas of North-East Greenland consist of a) Boreal-Arctic elements, b) Boreal-European taxa, c) endemic belemnites of Tethyan ancestry, and d) Tethyan species. These findings make North-East Greenland part of an immigration route from the Tethyan Realm via the north Atlantic to the high Boreal. This allowed Tethyan species, which are otherwise unknown from the Boreal Realm, to reach North-East Greenland. The occurrence of the Tethyan genus Pseudobelus in North-East Greenland also supports the interpretation of this taxon as a hemipelagic dweller, capable of crossing major distances. The belemnite patterns further suggest the existence of a proto Gulf-stream, documenting a south-to-north flow of warm surface waters as far north as Greenland already in the earliest Cretaceous (Valanginian). This has substantial implications for the interpretation of Early Cretaceous climate and oceanic current systems, as well as for the palaeobiology of belemnites.     

An early Bathonian Tethyan ammonite fauna from Argentina

Evidence for the Lower Bathonian Substage in most of the Pacific area has been lacking or controversial because index ammonoids of the Tethyan Subrealm in southern Europe and North Africa were unknown. During the Bathonian, the East Pacific margin belonged to either the Boreal Realm or the East-Pacific Subrealm of the Tethyan Realm, each with faunas distinctly different from the Eurafrican Tethyan faunas that form the basis for the chronostratigraphical standard. The first representatives of Morphoceras from South America, M. gulisanoi sp. nov., here described from a single locality in southern Mendoza Province, clearly document the Lower Bathonian Substage, probably the Zigzag Zone. Associated ammonoids are the perisphinctoidean Procerites cf. schloenbachi de Grossouvre and the oppeliid Oxycerites ( O. ) cf. aspidoides (Oppel), also typical Tethyan elements but less useful for dating.     

Late Triassic (Late Norian) gastropods from the Wallowa Terrane (Idaho,USA)

A diverse Late Triassic (Late Norian) gastropod fauna is described from the Mission Creek Limestone of the Wallowa terrane (Idaho, USA). Sample standardization by rarefaction analysis indicates that the fauna is even more diverse than the Late Triassic gastropod fauna from the Pucara Formation (Peru) which represents the most diverse gastropod fauna from South America. The gastropod fauna consists of 66 species; several genera are reported for the first time from North America. A high percentage of the species are highly ornamented and several have distinct siphonal canals. This suggests that the appearance of truly Mesozoic elements among the gastropods began before the Mesozoic Marine Revolution in other clades. The fauna is dominated by high-spired strongly ornamented procerithiids, a group more characteristic for the Jurassic. Comparison of the present fauna and the Iranian Nayband Formation gastropod fauna show that the procerithiids underwent a first global radiation in the Late Triassic. The high number of new species in this fauna suggests that sampling of Late Triassic gastropod faunas is still incomplete and hinders palaeobiogeographic considerations. Previous suggesions that gastropod faunas from the Wallowa and Wrangellia terranes resemble each other and are distinct from those of Alexander, Chulitna, and Farewell terranes are basically corroborated. The gastropod fauna of the Mission Creek Limestone differs considerably from that of the western and central Tethys but shares several taxa with the Late Triassic gastropod fauna of the Pucara Formation in Peru. Thus, the Hispanic corridor was probably not present in the Norian but opened only in the Early Jurassic. The subfamily Andangulariinae is introduced and placed in the Zygopleuridae. The generaSpiniomphalus, Nodoconus, Gudrunella, Blodgettella, Idahospira, andSiphonilda and the subgenusCryptaulax (Wallowax) are introduced. 27 species are erected. A lectotype is designated forCryptaulax rhabdocolpoides Haas, 1953.      

The evolution of canaliculate rudists in the light of a new canaliculate polyconitid rudist from the Albian of the Central Pacific

A new polyconitid rudist, Magallanesia canaliculata gen. et sp. nov., of probably late Albian age, is described from the Pulangbato area, central Cebu Island, the Philippines in the western Central Pacific and Takuyo‐Daini Seamount, now located in the Northwest Pacific. It is similar to Praecaprotina Yabe and Nagao, 1926, a Japanese–Central Pacific endemic genus of late Aptian – early Albian age, but differs in having canals that developed by partitioning of the large ectomyophoral cavity in the posterior part of the left valve. Its discovery strengthens the evidence for Pacific endemism in Albian times. Several other clades of canaliculate rudists flourished or evolved at the same time in different regions of the Tethyan Realm, suggesting the presence of common global biological and/or environmental factors stimulating the evolution of the canals despite such endemism. Furthermore, the finding of a canaliculate polyconitid provides evidence in favour of the evolutionary hypothesis of a polyconitid origin for the Late Cretaceous canaliculate rudist family Plagioptychidae Douvillé, 1888.     

Stratigraphie et sédimentologie des « Couches rouges » continentales du Jurassique-Crétacé du Haut Atlas central (Maroc) : implications paléogéographiques et géodynamiques

In several synclines of the central High Atlas, the “Redbeds” following the closure of the marine Tethyan Atlasic trough during the Middle Jurassic are constituted by three successive formations or units of continental deposits dated recently with biostratigraphical elements. Some micropaleontological markers, mainly charophytes and ostracods, allow to precise the stratigraphy in agreement with a Bathonian-?Callovian assignment for the lower unit (Guettioua Formation), and in dating the middle and upper units. The Upper Jurassic, mainly the Kimmeridgian, is developed in the lower part of the middle unit (Iouaridene Formation). The Barremian has been recognized in this middle unit and in the upper unit (Jbel Sidal Formation). The Jurassic/Cretaceous boundary is thus delimited for the first time with micropaleontological data. These new data are very significant for the Atlasic history during the Mesozoic. The basaltic flows inserted in the continental Jurassic-Cretaceous deposits of the central High Atlas result from two separate events in the Middle Jurassic and in the Barremian. The tectogenesis in the basins is characterized by a polyphase process including notably a synsedimentary tectonic activity conspicuous in the Barremian. The evidence of marine to brackish intercalations allows moreover to date the first Cretaceous transgressive event on the NW boundary of the High Atlas during the Lower Barremian and to consider an Atlantic paleogeographical interaction. SW margin of the Tethyan trough in the Lower and Middle Jurassic, the central High Atlas is merged with the margin of the central Atlantic Ocean during the Lower Cretaceous.     

Paleobiogeography of Africa: How distinct from Gondwana and Laurasia?

Although Africa was south of the Tethys Sea and originally belonged to the Gondwana, its paleobiogeographical history appears to have been distinct from those of both Gondwana and Laurasia as early as the earliest Cretaceous, perhaps the Late Jurassic. This history has been more complex than the classical one reconstructed in the context of a dual world (Gondwana vs. Laurasia). Geological and paleobiogeographical data show that Africa was isolated from the Mid-Cretaceous (Albian-Aptian) to Early Miocene, i.e., for ca. 75 million years. The isolation of Africa was broken intermittently by discontinuous filter routes that linked it to some other Gondwanan continents (Madagascar, South America, and perhaps India), but mainly to Laurasia. Interchanges with Gondwana were rare and mainly “out-of-Africa” dispersals, whereas interchanges with Laurasia were numerous and bidirectional, although mainly from Laurasia to Africa. Despite these intermittent connections, isolation resulted in remarkable absences, poor diversity, and emergence of endemic taxa in Africa. Mammals suggest that an African faunal province might have appeared by Late Jurassic or earliest Cretaceous times, i.e., before the opening of the South Atlantic. During isolation, Africa was inhabited by vicariant West Gondwanan taxa (i.e., taxa inherited from the former South American-African block) that represent the African autochthonous forms, and by immigrants that entered Africa owing to filter routes. Nearly all, or all immigrants were of Laurasian origin. Trans-Tethyan dispersals between Africa and Laurasia were relatively frequent during the Cretaceous and Paleogene and are documented as early as the earliest Cretaceous or perhaps Late Jurassic, i.e., perhaps by the time of completion of the Tethys between Gondwana and Laurasia. They were permitted by the Mediterranean Tethyan Sill, a discontinuous route that connected Africa to Laurasia and was controlled by sea-level changes. Interchanges first took place between southwestern Europe and Africa, but by the Middle Eocene a second, eastern route — the Iranian route — involved southeastern Europe and southwestern Asia. The Iranian route was apparently the filtering precursor of the definitive connection between Africa and Eurasia. The relationships and successive immigrations of mammal (mostly placental) clades in Africa allow the recognition of five to seven phases of trans-Tethyan dispersals between Africa and Laurasia that range from the Late Cretaceous to the Eocene-Oligocene transition. These Dispersal Phases involve dispersals toward Laurasia and/or toward Africa (immigrations). The immigrations in Africa gave rise to faunal assemblages, the African Faunal Strata (AFSs). All successful and typical African radiations have arisen from these AFSs. We recognize four to six AFSs, each characterized by a faunal association. Even major, old African clades such as Paenungulata or the still controversial Afrotheria, which belong to the oldest known AFS involving placentals, ultimately originated from a Laurasian stem group. Africa was an important center of origin of various placental clades. Their success in Africa is probably related to peculiar African conditions (endemicity, weak competition). Although strongly marked by endemicity, the African placental fauna did not suffer extinctions of major clades when Africa contacted Eurasia. The present geographic configuration began to take shape as early as the Mid-Cretaceous. At that time, the last connections between Africa and other Gondwanan continents began to disappear, whereas Africa was already connected to Eurasia by a comparatively effective route of interchange.     

Un faux polymorphitiné: Dubariceras, nouveau genre d'Ammonitina du Carixien mésogéen

Most of the Tethyan species usually attributed to the genera Platypleuroceras, Uptonia and Dayiceras (Polymorphitidae) are in reality forms phylogenetically related to Metaderoceras, but which show convergence in their morphology. Their originality justifies the creation of a new genus, Dubariceras and two new species, D. dubari and D. freboldi.The palaeobiogeographic implications of this phylogenetic interpretation confirm the profound differences between Tethyan and Boreal faunas in the Carixian. The platycone Polymorphitidae (Platypleuroceras, Uptonia and Dayiceras) are almost completely restricted to the Boreal Realm, whereas the faunas of the Tethyan Realm largely comprise the Acanthopleuroceratidae (Tropidoceras), Metaderoceras and its allied genus Dubariceras.     

A basal thunnosaurian from Iraq reveals disparate phylogenetic origins for Cretaceous ichthyosaurs

Cretaceous ichthyosaurs have typically been considered a small, homogeneous assemblage sharing a common Late Jurassic ancestor. Their low diversity and disparity have been interpreted as indicative of a decline leading to their Cenomanian extinction. We describe the first post-Triassic ichthyosaur from the Middle East, Malawania anachronus gen. et sp. nov. from the Early Cretaceous of Iraq, and re-evaluate the evolutionary history of parvipelvian ichthyosaurs via phylogenetic and cladogenesis rate analyses. Malawania represents a basal grade in thunnosaurian evolution that arose during a major Late Triassic radiation event and was previously thought to have gone extinct during the Early Jurassic. Its pectoral morphology appears surprisingly archaic, retaining a forefin architecture similar to that of its Early Jurassic relatives. After the initial latest Triassic radiation of early thunnosaurians, two subsequent large radiations produced lineages with Cretaceous representatives, but the radiation events themselves are pre-Cretaceous. Cretaceous ichthyosaurs therefore include distantly related lineages, with contrasting evolutionary histories, and appear more diverse and disparate than previously supposed.     

A new sphenodontian (Lepidosauria: Rhynchocephalia) from the Late Triassic of Argentina and the early origin of the herbivore opisthodontians

Sphenodontians were a successful group of rhynchocephalian reptiles that dominated the fossil record of Lepidosauria during the Triassic and Jurassic. Although evidence of extinction is seen at the end of the Laurasian Early Cretaceous, they appeared to remain numerically abundant in South America until the end of the period. Most of the known Late Cretaceous record in South America is composed of opisthodontians, the herbivorous branch of Sphenodontia, whose oldest members were until recently reported to be from the Kimmeridgian–Tithonian (Late Jurassic). Here, we report a new sphenodontian, Sphenotitan leyesi gen. et sp. nov., collected from the Upper Triassic Quebrada del Barro Formation of northwestern Argentina. Phylogenetic analysis identifies Sphenotitan as a basal member of Opisthodontia, extending the known record of opisthodontians and the origin of herbivory in this group by 50 Myr.     

Foraminiferan tests and dasycladalean thalli as cryptic microhabitats for thaumatoporellacean algae from Mesozoic (Late Triassic–Late Cretaceous) platform carbonates

Thaumatoporellacean algae are widespread constituents in Middle Triassic–Cretaceous shallow-marine carbonates of the Tethyan realm. Based on various examples from Mesozoic limestones of Mediterranean platforms (e.g., Dinaric, Apenninic, Apulia) and rare records of Iberia (Pyrenees), Saudi Arabia and Mexico, it is shown that thaumatoporellaceans commonly dwelt as cryptoendoliths in the tests of larger benthic foraminifera and the thalli of dasycladalean algae. Their high morphological plasticity allowed the test invasion and the adaptation to the available interior spaces (chambers, apertures). The temporal distribution of cryptoendolithic thaumatoporellaceans with first records in the Late Triassic, shows acme intervals in Early–Middle Jurassic and Early–Late Cretaceous times. Within the foraminiferans, the thaumatoporellaceans were erroneously considered as an integral part of the test, respectively, phrenoteca-like structures (species Biokovina gradacensis) in the Lower Jurassic and trematophore (species Scandonea? mediterranea) in the Upper Cretaceous. Therefore, the presence of phrenoteca-like structures in the Biokovinidae, being part of the family diagnosis, is challenged. The comparably thin walls of the cryptoendolithic thaumatoporellacean algae are interpreted as an adaptation to the poorly illuminated microhabitats (photoadaptation) in order to maximize light capture for photosynthesis.     

The West Gondwanan Occurrence Of The Hybodontid Shark Priohybodus, And The Late Jurassic–Early Cretaceous Age Of The Tacuarembó Formation, Uruguay

Priohybodus cf. P. arambourgi is reported for the first time from the Tacuarembó Formation of Uruguay. This species is a hybodontid shark known previously only from Upper Jurassic and Lower Cretaceous deposits of Saharan Africa and the Arabian Peninsula. The material (22 isolated teeth and a dorsal fin spine) was found in a thin bone bed, associated with abundant bone fragments, scales and teeth of semionotiform fishes and theropod dinosaurs. Until now, the age of the Tacuarembó Formation has been difficult to determine because its fossil content lacked useful biostratigraphic indicators. The finding of Priohybodus cf. P. arambourgi in that unit greatly expands the palaeobiogeographic range of the species, and allows us to propose a Late Jurassic–Early Cretaceous age for the Tacuarembó Formation.     

Late Triassic palynofloras in the Sichuan Basin,South China: Synthesis and perspective

Mesozoic strata are well developed and exposed continuously across the Sichuan Basin, South China. In particular, the Upper Triassic strata yield diverse and abundant spore-pollen fossils, providing a significant reference for the study of palaeoenvironmental variations across the Triassic–Jurassic transition where mass extinctions were occurring. In this paper, we summarize the major progress on Late Triassic palynological studies in this basin. To date, 151 genera (454 species) of sporomorph fossils have been reported from the Late Triassic strata in the Sichuan Basin. Three palynological assemblages are distinguished for the Late Triassic in the Sichuan Basin. Late Triassic vegetation in the Sichuan Basin shows a remarkable predominance of ferns, followed by conifers and cycads/ginkgophytes, and conifers show a distinct increase in abundance in the latest Triassic. In general, the Late Triassic palaeoclimate in the Sichuan Basin was tropical-subtropical, humid and warm. A synthesis of the data shows that the Late Triassic did not have a constant palaeoclimate in the Sichuan Basin, several climatic events are recognized: two warm and humid climate events in Norian-Rhaetian time, coupled with a cooler and drier condition in the latest Late Triassic. Further investigations in higher resolution at more continuous sections in the Sichuan Basin are needed to better understand the Late Triassic vegetation response, climate changes, as well as palaeoecosystem variations across the Triassic–Jurassic boundary.     

Palaeobiogeography of late cretaceous belemnites of Europe

A North Temperate Realm, characterized by Belemnitellidae Pavlov, and a South Temperate Realm, characterized by Dimitobelidae WHITEHOUSE, existed throughout the Late Cretaceous, while Tethyan belemnites belonging to Belemnopseidae Naef existed only in the Cenomanian and disappeared afterwards. The North Temperate Realm may be subdivided into North European and North American Provinces. The latter province includes Greenland, Canada, the Western Interior Region of North America, and the Atlantic and Gulf coasts of North America. The belemnites from the North American Province, consisting of populations of the generaActinocamax Miller andBelemnitella d’Orbigny, are closely related to the belemnites of the North European Province and appear to have migrated from this province to North America via Greenland and Arctic Canada. The North European Province extends from Ireland to the Ural Mountains. Belemnites from this province belong to the following genera:Neohibolites Stolley,Parahibolites Stolley,Belemnocamax Crick,Actinocamax Miller,Belemnellocamax Naidin,Gonioteuthis Bayle,Belemnitella d’Orbigny,Belemnella Nowak, andFusiteuthis Kongiel. Two subprovinces within the North European Province have been recognized: the Central European and Central Russian Subprovinces. These subprovinces are well-defined in the late Coniacian-Early Campanian and are characterized by theGonioteuthis stock andBelemnitella stock, respectively. The two subprovinces are less distinct in other periods of the Late Cretaceous and may disappear completely.     

Mesozoic conifers

The Coniferophyta attained their greatest diversity and abundance during the Mesozoic Era. Many early forms died out without apparent involvement in the evolution of the modern types. Yet, all living coniferophyte families and a surprising number of their present-day genera are clearly in evidence in the Mesozoic. Generally, modern families are recognizable by the Late Triassic or Early Jurassic while certain contemporary genera make their appearance as early as the Middle Jurassic. The Pinaceae appears to lag behind other families in that it lacks unequivocal representation before the onset of the Cretaceous although certain Late Triassic and Jurassic remains may belong to the family. Modern coniferophyte families appear to have originated somewhat earlier than was formerly believed and this brings to light problems in envisioning their evolution from the known Voltziales. Seed cones of certain of the latter are now known from petrified material which show a greater modification than was formerly known. Various kinds of detached organs that cannot be assigned to existing families are described and discussed and the bearing of the fossil record in certain questions on coniferophyte systematics is evaluated.     

Episodic sedimentation on a peri-Tethyan ridge through the Middle–Late Jurassic transition (Villány Mountains, southern Hungary)

The Villány area, as a central part of the Tisza microcontinent/terrane along the European margin of Tethys, was characterized by intense subsidence in the Early and Middle Triassic, followed by a long interruption of subsidence in the Late Triassic to Middle Jurassic. During the Middle–Late Jurassic transition, marine sedimentation started with three distinct sedimentary episodes dated as Late Bathonian, Early Callovian, and Middle–Late Callovian, respectively. The succession is terminated by a thick limestone of Middle Oxfordian age. The sedimentary features, microfacies, and macroinvertebrate associations of these four stratigraphic units are documented and illustrated. The Middle to Late Jurassic sedimentary episodes of the Villány succession record an interplay of local and global factors and paleogeographical changes. At the beginning, local tectonic movements governed the main features of sedimentation, though the role of eustasy was also essential. From the mid-Callovian onwards, global climatic, biotic, and paleoceanographical changes controlled the nature and formation of the local carbonate sediments. The Callovian stromatolites are attributed to the activity of sulphate-reducing bacteria in a deep sublittoral, current-swept environment. Upwelling of eutrophic Tethyan waters is recorded by the prevalence of the Bositra filament microfacies in the Callovian. The long submarine hiatus at around the Callovian–Oxfordian transition mirrors a serious restriction of the carbonate budget, due to sudden cooling and a change in the oceanic current system (opening of a circumglobal Tethyan Passage), and to a higher amount of dissolved CO₂. In the Middle Oxfordian, the carbonate production considerably increased in accordance with the sudden global warming.     

Jurassic neptunian dikes at Mt Mangart (Julian Alps,NW Slovenia)

Jurassic neptunian dikes are common within Upper Triassic to Lower Jurassic platform limestone of the Julian Alps. At Mt Mangart, the following geometries were observed: irregular dissolution cavities, thin penetrative fractures, larger fractures with sharp sidewalls, and laterally confined breccia bodies. Inside a complex neptunian dike system two main generations of infillings were differentiated. The first generation is heterogeneous and consists of bioclastic limestones, representing uniquely preserved sediments subdivided into five different microfacies. The second generation is more common and typically consists of coarse-grained breccias with host-rock clasts and marly limestone matrix containing echinoderms. Fracture formation and void filling of the first generation of neptunian dikes is dated as Pliensbachian and is interpreted to be caused by the Julian carbonate platform dissection due to widely recognized Lower Jurassic Tethyan rifting. The timing of formation for the second generation is only broadly constrained, ranging from the Pliensbachian to the Late Cretaceous.     

Tethyan changes shaped aquatic diversification

The Tethys Ocean existed between the continents of Gondwana and Laurasia from the Triassic to the Pliocene. Analyses of multiple biogeographic and phylogenetic histories reveal that the subsequent breakup of the Tethys greatly influenced the distributions of many species. The ancestral Tethyan realm broke into five biogeographic provinces, including the present‐day East Pacific, West Atlantic, East Atlantic, Mediterranean Sea, and Indo‐West Pacific. Palaeogeographic maps illustrate the Mesozoic Atlantic opening, the Cenozoic closure of the Tethys, the Messinian Salinity Crisis, the mid‐Miocene closure of the Central American Seaway, and Quaternary geological changes. Further, we consider Cenozoic sea‐level changes and the formation of freshwater habitats. These reconstructions allow assessment of patterns of aquatic diversification for marine and freshwater animals, and comparison of vicariance and dispersal processes. Estimated divergence times indicate that fragmentation of the Tethys was responsible for the vicariant speciation of aquatic animals because these dates are consistent with associated tectonic events. The opening of the Atlantic Ocean during the Cretaceous is responsible for the earliest isolation between the West and East Atlantic. The mid‐Miocene closure of the Tethys, which blocked global equatorial currents, appears to have isolated the Atlantic/Mediterranean Sea and Indo‐West Pacific. Finally, formation of the Isthmus of Panama isolated East Pacific and West Atlantic marine organisms. Dispersals related to the Messinian Salinity Crisis and Quaternary sea‐level changes influenced population structuring. Tethyan changes affected marine habitats, created new freshwater habitats, inland caves and ancient lakes along the Alps and Himalayas, and influenced anchialine caves at the edge of the ancient sea. The extensive new habitats provided opportunities for colonisation and rapid diversification. Future work should focus on testing the biological impact of the series of Tethyan changes.