Late Ordovician extinctions of bryozoans

An analysis of the final stratigraphic appearances of byrozoan species and genera, compiled in a world-wide bryozoan data base, revealed three discrete Late Ordovician extinctions. A Late Carddoc (Onnian) extinction was most pronounced on the plates of Baltica and Siberia. Endemic species and genera, confined to one plate and one lithotope were most affected and the extinction was coincident with increased migrations of bryozoan genera to Baltica and Siberia. The Late Caradoc extinction may be related to decreasing provinciality and competition between migrant and stenotopic taxa. Two major extinctions occurred in the Late Ashgill. The greatest of the two is recognized at the end of the Rawtheyan. and affected primarily taxa on the North American plate. The extinction at the end of the Hirnantian affected primarily Baltic taxa. The exact timing of the end-Rawtheyan extinction in North America cannot be established owing to incompleteness of the stratigraphic record. The Rawtheyan extinction occurred during a major glaciation centered in North Africa and a regression of epeiric seas. The large majority of North American survivors of the extinction are represented by Faunas preserved on Anticosti Island. which remained submerged during the regression. This evidence supports regression as a cause of the Rawtheyan extinctions in North America. The end-Hirnantian extinctions may be related to the ensuing transgression or to a wave of faunal migrations associated with the transgression. * Bryozoa, extinctions, Ordovician, Rawtheyan, Hirnantian, North America, Baltica .     

STROPHOMENIDE BRACHIOPODS FROM THE CHANGWU FORMATION (LATE KATIAN, LATE ORDOVICIAN) OF CHUN'AN, WESTERN ZHEJIANG, SOUTH-EAST CHINA

Abstract:  Late Ordovician strophomenide brachiopods (superfamilies Strophomenoidea and Plectambonitoidea) from the upper Changwu Formation (mid Ashgill, late Katian) of Jianglütang, Chun'an County, western Zhejiang Province, consist of ten genera and 12 species. Five new species of three new genera are recognized: Chunanomena triporcata , Chunanomena sembellina , Cheramomena subsolana , Lateriseptomena modesta , and Lateriseptomena rugosa . The strophomenide brachiopods from the upper Katian strata described in this study and those from the border region of Zhejiang and Jiangxi provinces reported in previous work contain 16 strophomenoid and 12 plectambonitoid genera, and most of the strophomenoids are endemic to South China. Numerical analysis of well-documented late Katian strophomenide brachiopod faunas indicates a strong provincialism, characterized by the highly distinct North American province (Laurentia), the South China-Kazakhstan province, and the Avalonia-Baltica province (Wales, Belgium and Sweden). Surprisingly, the Girvan district of Scotland, which was a peri-Laurentian terrane during the Ordovician, contains a late Katian brachiopod fauna that is more closely related to the contemporaneous brachiopods of Avalonia-Baltica than to those of North America.     

Bryozoan provinces and patterns of generic evolution and extinction in the Late Ordovician of North America

The Late Ordovician bryozoan genera of central and southeastern North America are geographically distributed in three biotic provinces, separated by boundaries reflecting major lithofacies differences. The central Cincinnati Province contains most of the North American endemic genera, and represents a narrow ecological zone separating the clastic wedges of the marginal Reedsville-Lorraine Province from the cratonic carbonate platform of the Red River-Stony Mountain Province. The provinces provided major life zones, or biomes, for each of the five bryozoan orders. Genera comprising the provinces differed as well in morphologic complexity, geochronologic survivorship, tiering, endemism and eurytopy. Regions on either side of the Cincinnati Province were dominated by inferred immigrants from Baltoscandia. Al-logenic provincial succession produced time-averaged mixed faunas in regions near the provincial boundaries. Although most generic originations took place within the Cincinnati Province, evolutionary novelties are associated with the Reedsville-Lorraine Province. The loss of the diverse Cincinnati Province, connected with global cooling and a eustatic lowering of sea level, may have been a chief factor in the Late Ordovician extinction of bryozoan genera. Genera from the Red River-Stony Mountain Province differentially survived into the Silurian.     

Late Ordovician extinction of North American and British crinoids

After reaching a diversity peak in the Caradocian, North American Ordovician crinoids underwent a gradual decline to a nadir in the early Ashgillian (Maysvillian). This interval, recording extinction of the Cleiocrinidae, Merocrinidae, Ottawacrinidae, Hybocystitidae, and several lineages of camerate crinoids, was apparently caused by major environmental shifts in seas of eastern North America resulting from a westward-prograding wedge of terrigenous clastics derived from the Taconic Highlands, possibly coupled with a marine transgression in the Maysvillian that allowed colder water slope biofacies to invade the craton. Crinoids suffered a major episode of extinction in the late Ashgillian (late Richmondian/Rawtheyan). This event, preceding the end of the Ordovician by at least one stage or 2 to 4 million years, resulted in extinction of 12 families of crinoids including the Xenocrinidae, Tanaocrinidae, Reteocrinidae, Archaeocrinidae, Anthracocrinidae, Cincinnaticrinidae, Iocrinidae, Anomalocrinidae, Carabocrinidae, Cupulocrinidae, Porocrinidae, and Hybocrinidae. Glacio-eustatic lowering of sea level may have triggered this crisis by partially draining the North American craton, resulting in changes in oceanic circulation, salinity, and temperature. Latest Ordovician (Hirnantian) carbonates of the North American mid-continent region contain pelmatozoan assemblages from which Silurian crinoids radiated. These taxa were largely unaffected by a minor extinction event at the Ordovician/Silurian boundary.     

Evolution of the Rhynchotrema–Hiscobeccus lineage: implications for the diversification of the Late Ordovician epicontinental brachiopod fauna of Laurentia

Multivariate analysis based on nine biometric characters of 171 Late Ordovician rhynchonellide specimens from nine upper Sandbian–upper Katian localities in North America supports the hypothesis that one of the diagnostic taxa of the North American epicontinental brachiopod fauna, Hiscobeccus, evolved from Rhynchotrema, which lived predominantly in peri‐cratonic settings. The oldest known Hiscobeccus, H. mackenziensis of early Katian age, exhibits transitional characteristics between Rhynchotrema and Hiscobeccus, and it clusters more closely with Rhynchotrema than with younger species of Hiscobeccus of mid–late Katian (Maysvillian–Richmondian) age. Diversification of the Hiscobeccus lineage in epicontinental seas was characterized by drastic increase in shell size, globosity and lamellosity, especially in palaeoequatorially located inland basins. Such morphological trends are interpreted as adaptation to relatively shallow, muddy substrates, moderate water turbulence, relatively low oxygen content and unstable supply of nutrients in generally overheated epicontinental seas with sluggish circulation.     

Response of Late Ordovician paleoceanography to changes in sea level, continental drift, and atmospheric pCO2: potential causes for long-term cooling and glaciation

We performed sensitivity experiments using an ocean general circulation model at two stages of the Late Ordovician (Caradoc, 454 Ma; Ashgill, 446 Ma) under a range of atmospheric pCO₂ values (8–18× PAL; pre-industrial atmospheric level) at high and low sea level.

The model results indicate that the long-term cooling trend during the Late Ordovician can be explained by progressive cooling of the global ocean in response to falling levels of atmospheric pCO₂, sea level change, and paleogeographic change. These results also explain the occurrence of low latitude cool-water carbonates in North America.

In all simulations, a drop in sea level led to a reduction in poleward ocean heat transport. This indicates a possible positive feedback that could have enhanced global cooling in response to sea level drop during the Late Ordovician. Alterations in poleward ocean heat transport linked to changes of atmospheric pCO₂ also indicate that there is a threshold of 10× PAL, above which ocean current change cannot be responsible for glaciation in the Late Ordovician. Continental drift could explain the observed global cooling trend in the Late Ordovician through a combined poleward ocean heat transport feedback and increased ice-albedo effect if atmospheric pCO₂ was low during the entire Late Ordovician.

The model results further indicate that the response of meridional overturning to changes in paleogeography, atmospheric pCO₂, and sea level is stronger than the response of surface circulation to these perturbations. Because the overturning circulation is so strong, meridional overturning was the dominant mechanism for described changes in heat transport in the Late Ordovician.     

Mountain uplift explains differences in Palaeogene patterns of mammalian evolution and extinction between North America and Europe

Patterns of late Palaeogene mammalian evolution appear to be very different between Eurasia and North America. Around the Eocene–Oligocene (EO) transition global temperatures in the Northern Hemisphere plummet: following this, European mammal faunas undergo a profound extinction event (the Grande Coupure), while in North America they appear to pass through this temperature event unscathed. Here, we investigate the role of surface uplift to environmental change and mammalian evolution through the Palaeogene (66–23 Ma). Palaeogene regional surface uplift in North America caused large-scale reorganization of precipitation patterns, particularly in the continental interior, in accord with our combined stable isotope and ecometric data. Changes in mammalian faunas reflect that these were dry and high-elevation palaeoenvironments. The scenario of Middle to Late Eocene (50–37 Ma) surface uplift, together with decreasing precipitation in higher-altitude regions of western North America, explains the enigma of the apparent lack of the large-scale mammal faunal change around the EO transition that characterized western Europe. We suggest that North American mammalian faunas were already pre-adapted to cooler and drier conditions preceding the EO boundary, resulting from the effects of a protracted history of surface uplift.     

The latest Ordovician Hirnantia Fauna (Brachiopoda) in time and space

The diachronous temporal and spatial distribution of the Hirnantia brachiopod fauna and the complicated pattern of terminal Ordovician events are documented through biostratigraphical analysis of the Ordovician-Silurian boundary strata in S China, Sibumasu, Xizang and elsewhere. The duration of these events (longer than the half Myr derived from isotopic excursions) indicates that they were not abrupt and instantaneous. The presence of some core taxa of the Hirnantia fauna in the upper P. pacificus Biozone (known from their earliest occurrence in China) signals the start of increased water ventilation due to the invasion of cool water across the Yangtze Basin. Low- and higher-diversity Hirnantia faunas related to onshore, shallow-water and to offshore, deeper-water environments, respectively, developed first in the basal and upper N. extraordinarius-N. ojsuensis Biozone. Disappearance of most of the fauna in the early N. persculptus Biozone suggests that the glacial maximum started to decline. The presence of the Hirnantia fauna in the upper N. persculptus to the lower P. acuminatus biozones indicates the continuation of cool water environments in some places. The diachronous disappearance of deteriorating environments (earlier in later Hirnantian and finally in the early Rhuddanian) is associated with geographical heterogeneity. Occurrences of atrypids, pentamerids and spiriferids along with key elements of the Hirnantia fauna in N Guizhou provide a link between the Late Ordovician radiation and Early Silurian recovery of these major brachiopod groups.     

简论相似性测度的选择——以奥陶纪末大灭绝后全球腕足动物古地理为例

灭绝事件对古生物地理格局的影响已引起关注,近期研究表明奥陶纪末大灭绝事件后多样性显著高于传统认识,而全球该时期腕足动物的古生物地理分布情况尚未见报道。本文基于已发表的和最新的资料及所掌握新数据的整理,建立全球腕足动物志留纪初鲁丹(Rhuddanian)早期（残存期）13个产地72属137个出现信息(occurrence...     

Conodontophorid biodiversification during the Ordovician in South China

Wu, R., Stouge, S. & Wang, Z. 2012: Conodontophorid biodiversification during the Ordovician in South China. Lethaia, Vol. 45, pp. 432–442. Analysis of the Ordovician conodontophorid diversity pattern for South China using normalized and total diversity measures reveals that diversity peaks occurred in the mid‐Tremadocian, mid‐late Floian, early Dapingian and mid‐Darriwilian periods. The conodontophorids radiated during the Floian, maintaining relatively high diversity into the early part of the Middle Ordovician until a significant diversity decrease occurred in the late Dapingian. A relatively low diversity level prevailed in the Late Ordovician. Three diversification intervals based on origination, extinction and turnover rates have been identified i.e. (1) Tremadocian to mid‐late Floian, (2) early Dapingian and (3) late Dapingian to early Darriwilian. Diversity curves for conodontophorids, brachiopods, graptolites, acritarchs and trilobites from South China are comparable during the Early Ordovician, although differences are apparent in the Middle and Late Ordovician. In South China, conodontophorid diversity reacted primarily to sea‐level changes during the Early and Middle Ordovician, when the peak of this biodiversification generally coincided with a transgression. Climate changes – especially the global cooling that occurred during the Late Ordovician glaciation – and sea‐water chemistry were also important controlling factors. □Biodiversification, conodonts, Ordovician, South China.     

Phylogenetic Clustering of Origination and Extinction across the Late Ordovician Mass Extinction

Mass extinctions can have dramatic effects on the trajectory of life, but in some cases the effects can be relatively small even when extinction rates are high. For example, the Late Ordovician mass extinction is the second most severe in terms of the proportion of genera eliminated, yet is noted for the lack of ecological consequences and shifts in clade dominance. By comparison, the end-Cretaceous mass extinction was less severe but eliminated several major clades while some rare surviving clades diversified in the Paleogene. This disconnect may be better understood by incorporating the phylogenetic relatedness of taxa into studies of mass extinctions, as the factors driving extinction and recovery are thought to be phylogenetically conserved and should therefore promote both origination and extinction of closely related taxa. Here, we test whether there was phylogenetic selectivity in extinction and origination using brachiopod genera from the Middle Ordovician through the Devonian. Using an index of taxonomic clustering (R_CL) as a proxy for phylogenetic clustering, we find that A) both extinctions and originations shift from taxonomically random or weakly clustered within families in the Ordovician to strongly clustered in the Silurian and Devonian, beginning with the recovery following the Late Ordovician mass extinction, and B) the Late Ordovician mass extinction was itself only weakly clustered. Both results stand in stark contrast to Cretaceous-Cenozoic bivalves, which showed significant levels of taxonomic clustering of extinctions in the Cretaceous, including strong clustering in the mass extinction, but taxonomically random extinctions in the Cenozoic. The contrasting patterns between the Late Ordovician and end-Cretaceous events suggest a complex relationship between the phylogenetic selectivity of mass extinctions and the long-term phylogenetic signal in origination and extinction patterns.     

Evolutionary and biogeographical shifts in response to the Late Ordovician mass extinction

The Late Ordovician mass extinction was an interval of high extinction with inferred low ecological selectivity, resulting in little change in community structure after the event. In contrast, the mass extinction may have fundamentally changed evolutionary dynamics in the surviving groups. We investigated the phylogenetic relationships among strophomenoid brachiopods, a diverse brachiopod superfamily that was a primary component of Ordovician ecosystems. Four Ordovician families/subfamilies sampled in the analysis (Rafinesquinidae, Strophomeninae, Glyptomenidae and Furcitellinae) were reconstructed as monophyletic groups, and the base of the strophomenoid clade that dominated the Silurian recovery was reconstructed as diversifying alongside these families during the Middle Ordovician. We time‐calibrated the phylogeny and used geographical occurrences to investigate biogeographical changes in the strophomenoids through time with the R package BiogeoBEARS . Our results indicate that extinction was higher in taxa whose ranges were constrained to tropical or subtropical regions. Furthermore, our results suggest important shifts in the diversification patterns of these brachiopods after the mass extinction. While most of the strophomenoid families survived the Late Ordovician event, ecologically abundant taxonomic groups during the Ordovician were either driven to extinction, reduced in diversity, or slowly died off during the Silurian. The new abundant strophomenoid taxa derived from one clade (consisting of Silurian–Devonian groups such as Douvillinidae, Strophodontidae and Amphistrophiidae) that diversified during the post‐extinction radiation. Our results suggest the selective diversification during the Silurian radiation, rather than selective extinction in the Late Ordovician, had a greater impact on the evolutionary history of strophomenoid brachiopods.     

Devonian conodont biogeography: Quantitative analysis of provinciality

Cluster analysis of conodont faunas from each of 17 Lower‐lower Upper Devonian zones and subzones (data as reported by Klapper and Johnson, 1980) reveal changing patterns of provinciality.

Provinciality, expressed by a differentiation into western Laurussian and proto‐Tethyan biogeographic regions, is moderate in the lower Lochkovian but is low or absent in the upper Lochkovian‐lower Pragian. Provincialism returns in the Pragian and reaches its maximum development during the Emsian. Most Australian faunas are distinct from those of western Laurussia and the proto‐Tethys. Conodont faunas from suspect terranes of western North America display no unusual biogeographic affinities. Provincialism declines during the Eifelian and is only weakly developed in Givetian‐lower Frasnian faunas.

Changing global sea level during the Devonian may explain the development of Devonian conodont provinciality. As proposed by Klapper and Johnson (1980), low provinciality is associated with low stands of sea level. Endemic faunas develop in isolated epeiric seas during intermediate stages of sea level rise. High stands of sea level ultimately drown barriers to faunal exchange and prompt a return to low provinciality conditions.     

Reef reconstruction after extinction events of the latest ordovician in the Yangtze platform,South China

Summary Early Silurian reef reconstruction on the Yangtze Platform, in the northern part of the South China Block, is preceded by a combination of regional and global processes. During most of Ashgill time (Late Ordovician), the area was dominated by Wufeng Formation deep water graptolitic black shales. Reefs largely disappeard in the middle of the Ashgill Stage, from the northwestern margin of Cathaysian Land (southeastern South China Block), in advance of the Late Ordovician glaciation and mass extinction, due to regional sea-level changes and regional uplift, unrelated to the mass extinction itselt. Late Ordovician microbial mudmound occurrence is also found in the western margin of the Yangtze Platform, its age corresponding to theDicellograptus complexus graptolite biozone of pre-extinction time. On the Yangtze Platform, a thin, non-reef-bearing carbonate, the Kuanyinchiao Formation (=Nancheng Formation in some sites), thickness generally no more than 1m, occurs near several landmasses as a result of Hirnantian regression. Reappearance of the earliest Silurian carbonates consisting of rare skeletal lenses in the upper part of Lungmachi Formation, are correlated to theacensus graptolite biozone, early Rhuddanian of Shiqian, northeastern Guizhou, near Qianzhong Land. Carbonate sediments gradually developed into beds rich in brachiopods and crinoids in the lower part of Xiangshuyuan Formation, middle Rhuddanian. In the middle part of Xiangshuyan Formation, biostromes, containing abundant and high diversity benthic faunas such as corals, crinoids and brachiopods, show beginnings of reconstruction of reef facies. Substantial reef recovery occurred in the upper part of Xiangshuyuan Formation, lower Aeronian, as small patch reefs and biostromes. During the late Aeronian, carbonate sediments, especially reefs and reef-related facies, expanded on the upper Yangtze Platform, and radiation of reefs occurred in Ningqiang Formation, upper Telychian. The long period of reef recovery, taking several million years, remains difficult to explain, because redistribution of any refugia faunas would be expected to take place soon after the extinction. Reefs and reef-related facies subsequently declined after Telychian time due to regional uplift of the major portion of the Yangtze Platform. Carbonate facies are therefore uncommon in South China during the rest of Silurian time.     

Late Devonian global ostracod palaeobiogeography

A global Late Devonian ostracod database is constructed, incorporating new materials from South China and Northwest China. Four palaeobiogeographical units (Cathaysia, North America, Europe and peri‐Gondwana) are recognized during the Frasnian and five palaeobiogeographical units (Cathaysia, North America, Europe, Siberia and Australia) in the Famennian. Three controlling factors (climatic zonation, geographical isolation and global sea‐level changes) are identified to have played roles in shaping the palaeogeographical regionalization of ostracod faunas in the Late Devonian. The ostracod palaeobiogeography in the Frasnian was mainly influenced by climatic zonation, while rapid changes in tectonic configuration in the Famennian drastically altered the global palaeobiogeography of ostracods. The palaeobiogeographical regionalization of ostracod faunas suggests that Laurussia and Gondwana continued to draw near during the Late Devonian, with the first collision occurring in Southern Central Europe in the Famennian. The South China plate drifted northward to the Kazakhstan plate away from the Australian plate, which gradually became isolated during the Famennian Stage.     

New Late Triassic gastropods from the wallowa Terrane (Idaho) and their biogeographic significance

Summary Two characteristic new species and one new genus are described from the Late Triassic of Idaho (Wallowa Terrane):Brochidiella idahoensis n. gen., n. sp. andPtychostoma ornata n. sp.Brochidiella is only known from western North America.Ptychostoma is present in the Carnian of the European Alps (Tethys) and is widespread in the western part of the North American continent (Panthalassa). Late Triassic gastropod faunas from the accreted terranes of North America are poorly known but hold a great potential for future palaeobiogeographic reconstructions.     

The last dicynodont: an Australian Cretaceous relict

Some long-forgotten fossil evidence reveals that a dicynodont (mammal-like reptile of the infraorder Dicynodontia) inhabited Australia as recently as the Early Cretaceous, ca. 110 Myr after the supposed extinction of dicynodonts in the Late Triassic. This remarkably late occurrence more than doubles the known duration of dicynodont history (from ca. 63 Myr to ca. 170 Myr) and betrays the profound impact of geographical isolation on Australian terrestrial faunas through the Mesozoic. Australia's late-surviving dicynodont may be envisaged as a counterpart of the ceratopians (horned dinosaurs) in Cretaceous tetrapod faunas of Asia and North America.     

Facies distribution patterns of some marine benthic faunas in early paleozoic platform environments

Worldwide Late Cambrian—Silurian lithofacies patterns indicate that the platforms of that time were sites of accumulation of two essentially different rocks suites: the platform carbonate rocks and the platform terrigenous rocks. Most of the platform rocks accumulated as sediments in shallow marine environments similar to those of the present but far more widely spread.Present-day marine benthic faunas are distributed in depth zones which are primarily controlled by temperature. Faunas tend to occur in substrate-related discrete clusters (communities) within each life zone; similar substrates in different depth zones commonly have different faunal associations. Individual phyletic stocks may encounter environmental optimum or near-optimum conditions in certain areas, that commonly are revealed by an abundance of species and individuals within species in each stock. Environmental optimum conditions depend upon availability of food that may be utilized, modes of feeding of the animals present, water motion, and substrate, among other factors. Organisms in past seas were distributed in patterns similar to those of the present.Carbonate platforms were particularly widespread during the latest Cambrian—Early Ordovician. Intertidal environments spread widely across those platforms during that time and characteristic faunal associations developed in them. Saukiid and related tribolites dominated latest Cambrian carbonate platform intertidal faunas. The Early Ordovician carbonate platform intertidal was dominated by archeogastropod-nautiloid cephalopod faunas. These animals were joined by tabulate corals and certain brachiopods during the latter part of the Ordovician and Silurian as prominent faunal elements in the carbonate platform intertidal—shallow subtidal. Cruziana and related trace fossils, bivalves, and certain tribolites (notably homalonotids and dalmanitids) dominated most terrigenous platform intertidal—shallow subtidal faunas of the Ordovician and Silurian.Articulate brachiopods (primarily orthoids, strophomenoids, and rhynchonelloids) appear to have been relatively prominent during the Early Ordovician in shallow subtidal environments on both carbonate and terrigenous platforms and to have spread down the bathymetric gradient into increasingly deeper subtidal areas of both platforms during the latter part of the Ordovician. Tribolites dominated faunas in relatively moderate to deep subtidal environments on both platforms during the early part of the Ordovician. They were gradually replaced by brachiopods in first the shallower, and later the deeper subtidal as dominant members of the faunas. Brachiopods (primarily pentameroids and spiriferoids) dominated nearly all Silurian warm-water subtidal environments from the shallow subtidal to the edges of the platforms.Platform uplifts in the Middle Ordovician and glacio-eustatic sea-level fluctuations in the Late Ordovician caused environmental changes across the platforms that were accompanied by marked replacements among marine benthic faunas in all environments. The distribution of Ordovician carbonate platforms and glacial deposits suggests that an Ordovician polar region may have been close to present-day equatorial Africa and that Ordovician warm temperate-tropical regions lay close to the present-day North Pole.     

Biostratigraphy, biodiversity and palaeoenvironments of the chitinozoans and associated palynomorphs from the Upper Ordovician of the Central Anti-Atlas, Morocco

The range, abundance and diversity of Upper Ordovician chitinozoans and of various other organic-walled microfossils (i.e., acritarchs, leiospheres, scolecodonts) have been documented in the Bou-Ingarf section, Central Anti-Atlas, Morocco. The goal of this study is to provide a refined biostratigraphic scheme for the Lower Ktaoua, the Upper Tiouririne, the Upper Ktaoua and the Lower Second Bani formations in order to locate the initial effects of the Late Ordovician glaciation on the composition of assemblages of organic-walled microfossils. Five Late Ordovician chitinozoan biozones belonging to the North Gondwana zonation are identified. During the late Caradoc and early Ashgill, no significant changes in chitinozoan assemblages are recorded that could be attributed to any dramatic climatic and/or environmental events. The almost absence of phytoplanktonic elements (acritarchs and leiospheres) and of cryptospores in the fistulosa/barbata and in the nigerica biozones are interpreted as resulting of lower offshore environmental conditions. The reappearance of a variety of palynomorphs in the lower part of the elongata biozone does not indicate drastic climatic changes but more likely a shallowing trend of the marine environment and an increased primary production, possibly related to a cooling of the surface water masses subsequently to early effects of a general climatic cooling. The first obvious evidence of the Late Ordovician glaciation occurs in the lower part of the elongata biozone, at the base of the sandy member topping the Lower Second Bani Formation. This sandy deposit is interpreted as the result of a dramatic fall of the sea level due to the development of the Hirnantian ice cap on Gondwana. This short-lived glaciation ended when the northern Gondwanan ice cap melted during the upper part of the elongata and the oulebsiri biozones. A biological crisis affected both the chitinozoans and the acritarchs just after this glaciation, i.e., in the latest Hirnantian.     

A re-evaluation of the ‘mid-Cretaceous sauropod hiatus’ and the impact of uneven sampling of the fossil record on patterns of regional dinosaur extinction

The mid-Cretaceous of North America and Europe has long been noted for the absence of sauropod dinosaurs, leading several authors to suggest that this depauperate interval is a consequence of an end-Albian sauropod extinction. This time period has become known as the ‘mid-Cretaceous sauropod hiatus’, with the subsequent presence of titanosaurian sauropods in the latest Cretaceous of North America and Europe interpreted as the result of dispersal of taxa from South America and Africa, respectively. However, several lines of evidence indicate that this hiatus is probably a sampling artefact. New fossil and trackway discoveries have considerably shortened the hiatus, reducing it to the Turonian–early Campanian in North America, and to just two short intervals in the late Cenomanian–early Turonian and late Coniacian–Santonian of Europe. Palaeoenvironmental analyses of sauropods demonstrate an inland terrestrial preference for titanosaurs, the dominant Late Cretaceous sauropods; however, during the hiatus there was a decline in inland deposits and increase in coastal sediments in Europe and North America, which would have greatly reduced the probability of preserving titanosaurs. Neither the decline in inland deposits, nor the ‘sauropod hiatus’, occurred elsewhere in the world. Statistical comparisons also demonstrate a significant positive correlation between fluctuations in inland deposits and sauropod occurrences during the mid–Late Cretaceous in Europe and North and South America. Lastly, cladistic analyses do not place latest Cretaceous North American and European titanosaurs within South American and African clades, contradicting the predictions of the ‘austral immigrant’ hypothesis. The latter hypothesis also receives little support from biogeographical analysis of dispersal among titanosaurs. Thus, the ‘sauropod hiatus’ of North America and Europe is most plausibly interpreted as the product of a sampling bias pertaining to the rarity of inland sediments and dominance of coastal deposits preserved in these two regions during the mid-Cretaceous. The presence of titanosaurs in these areas during the latest Cretaceous can be explained by dispersal from Southern Hemisphere continents, but this is no more probable than descent from Early Cretaceous incumbent faunas or dispersal from Asia.