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The Spathian (late Early Triassic) Virgin Formation of south−western Utah (USA) yields a comparatively diverse benthic fauna that flourished ~2 Ma after the end−Permian mass extinction. In this study, we present quantitative palaeoecological data, which are analysed in the context of depositional environments. This integrated approach helps to discriminate between effects of the end−Permian mass extinction event and local environmental factors on alpha diversity and ecological structure of the Virgin Fauna. Shallow subtidal environments yield the highest species richness and lowest dominance val− ues as recorded in two benthic associations, the Eumorphotis ericiusAssociation and the Protogusarella smithi Association, both ofwhich contain 20 benthic species (bivalves, gastropods, brachiopods, echinoderms, and porifers). Tidal inlet deposits yield a low diverse fauna (Piarorhynchella triassica Association) with a very high dominance of filter feeders adapted to high energy conditions.Another comparably low diverse fauna is recorded by the Bakevellia exporrecta Association, which occurs in deposits of the offshore transition zone,most likely reflecting unconsolidated substrates. A single sample contain− ing five bivalve species (Bakevellia costata Assemblage) is recorded from a marginal−marine setting. The Virgin fauna yields a bulk diversity of 30 benthic species (22 genera) of body fossils and 14 ichnogenera and, thus, represents the most di− verse marine bottom fauna known so far from the Early Triassic. Our results suggest that oceanographic conditions during the early Spathian enabled ecosystems to rediversify without major abiotic limitations. However, taxonomical differentia− tion between habitats was still low, indicating a time lag between increasing within−habitat diversity (alpha diversity) and the onset of taxonomical differentiation between habitats (beta diversity). We suggest that taxonomical habitat differentia− tion after mass extinction events starts only when within−habitat competition exceeds a certain threshold, which was not yet reached in the Spathian of the investigated area. This interpretation is an alternative to previous suggestions that the preva− lence of generalistic taxa in the aftermath of mass extinction events reflects protracted environmental stress. The onset of in− creasing beta diversity is a potential criterion for distinguishing two major recovery phases, the first ending with habitat satu− ration and the second ending with the completion of ecosystem differentiation.
Ammonoid biodiversity changes from shallow to offshore environments across the Cenomanian–Turonian (C–T) boundary are reconstructed in the Yezo Group, Hokkaido, Japan. This group was probably deposited at approximately 35–45ºN along a westward subduction margin in the northeastern Asian continent. Temporal changes in species richness in the Yezo Group, which show persistently high values during the middle Cenomanian and then decline stepwise from near the middle–late Cenomanian boundary, resemble those in Europe, but not those in Tunisia and the Western Interior. These differences suggest that the Cenomanian–Turonian “mass extinction” was not a global event for ammonoids but was restricted to mid−palaeolatitudinal regions (Europe and Japan). Sea level and climate changes probably influenced ammonoid faunas in the Yezo Group as well as those in Europe. However, it is unlikely that a single, simple cause led to the C–T boundary “mass extinction” because various abiotic changes in the Cenomanian–Turonian transition have been detected, and biotic and abiotic change are interrelated.
To evaluate stratigraphic evidence for the time of origin of the clade of extant amphibians (Lissamphibia), we attempt to establish a confidence interval on the lower bound of the stratigraphic range of this clade. This is based on the stratigraphic distribution of 1207 fossiliferous localities that have yielded lissamphibians, the relative area of sedimentary rocks from various periods (upper Paleozoic to present) exposed on the continents, and ten exponential−growth models of lissamphibian diversity that differ by the assumed effects of three major biological crises and the assumed starting times of lissamphibian diversification. The results suggest a more recent origin of Lissamphibia than advocated in most recent molecular studies. They are also more compatible with monophyly than with polyphyly of the extant amphibians, but heavily depend on poorly constrained assumptions about lissamphibian extinction rates during biological crises. Counts of lissamphibian diversity through time that consider ghost lineages and stage durations show moderate declines across the Cretaceous–Paleogene and Oligocene–Miocene boundaries.
Taxonomic diversity of NW Caucasus brachiopods changed cyclically in the Early–Middle Jurassic. Diversifications took place in the Late Sinemurian–Early Pliensbachian, Middle–Late Toarcian and Late Aalenian–Early Bajocian, while diversity decreases occured in Late Pliensbachian–Early Toarcian, Early Aalenian and Late Bajocian. Outstanding diversity decline in the Late Pliensbachian–Early Toarcian corresponds to a global mass extinction interval, whose peak has been documented in the Early Toarcian. Similar diversity changes of brachiopods are observed in other Tethyan regions, including the well−studied Bakony Mountains, although in NW Caucasus the recovery after demise have begun earlier. The causes of Pl−To mass extinction in the studied region are enigmatic. Probably, it could be linked to anoxia, but its correspondence to the beginning of transgression is not coincident with the global record, so eustatic causes seem to be doubtful for this region.
This paper describes fourteen brachiopod species in eleven genera from the Late Permian Wuchiapingian Coal Series (Lungtan Formation) of South China. Of these, the shell bed fauna from the basal Lungtan Formation is interpreted to represent the onset of the recovery of shelly faunas in the aftermath of the Guadalupian/Lopingian (G/L) mass extinction in South China. The post−extinction brachiopod faunas in the Wuchiapingian are characterized by the presence of numerous Lazarus taxa, survivors, and newly originating taxa. These elements capable of adapting their life habits were relatively more resistant to the G/L crisis. The post−extinction faunas, including survivors and the elements originating in the recovery period, have no life habit preference, but they were all adapted to a variety of newly vacated niches in the Late Permian oceans. Two new species, Meekella beipeiensis and Niutoushania chongqingensis, are described, and two Chinese genera, Niutoushania and Chengxianoproductus, are emended based on re−examination of the type specimens and new topotype materials from the Lungtan Formation.
For the past three decades, the Alvarez impact theory of mass extinction, causally related to catastrophic meteorite impacts, has been recurrently applied to multiple extinction boundaries. However, these multidisciplinary research efforts across the globe have been largely unsuccessful to date, with one outstanding exception: the Cretaceous–Paleogene boundary. The unicausal impact scenario as a leading explanation, when applied to the complex fossil record, has resulted in force−fitting of data and interpretations (“great expectations syndrome”). The misunderstandings can be grouped at three successive levels of the testing process, and involve the unreflective application of the impact paradigm: (i) factual misidentification, i.e., an erroneous or indefinite recognition of the extraterrestrial record in sedimentological, physical and geochemical contexts, (ii) correlative misinterpretation of the adequately documented impact signals due to their incorrect dating, and (iii) causal overestimation when the proved impact characteristics are doubtful as a sufficient trigger of a contemporaneous global cosmic catastrophe. Examples of uncritical belief in the simple cause−effect scenario for the Frasnian–Famennian, Permian–Triassic, and Triassic–Jurassic (and the Eifelian–Givetian and Paleocene–Eocene as well) global events include mostly item−1 pitfalls (factual misidentification), with Ir enrichments and shocked minerals frequently misidentified. Therefore, these mass extinctions are still at the first test level, and only the F–F extinction is potentially seen in the context of item−2, the interpretative step, because of the possible causative link with the Siljan Ring crater (53 km in diameter). The erratically recognized cratering signature is often marked by large timing and size uncertainties, and item−3, the advanced causal inference, is in fact limited to clustered impacts that clearly predate major mass extinctions. The multi−impact lag−time pattern is particularly clear in the Late Triassic, when the largest (100 km diameter) Manicouagan crater was possibly concurrent with the end−Carnian extinction (or with the late Norian tetrapod turnover on an alternative time scale). The relatively small crater sizes and cratonic (crystalline rock basement) setting of these two craters further suggest the strongly insufficient extraterrestrial trigger of worldwide environmental traumas. However, to discuss the kill potential of impact events in a more robust fashion, their location and timing, vulnerability factors, especially target geology and palaeogeography in the context of associated climate−active volatile fluxes, should to be rigorously assessed. The current lack of conclusive impact evidence synchronous with most mass extinctions may still be somewhat misleading due to the predicted large set of undiscovered craters, particularly in light of the obscured record of oceanic impact events.
Throughout their history, species had to face environmental variations spatially and temporally. How both levels of variation interact will be of key importance in conditioning their response to major perturbations. We addressed this question by focusing on a period in Earth’s history marked by dramatic environmental and faunal changes, the Late Devonian Frasnian/Famennian boundary. From a paleogeographic point of view, this period is characterized by a cosmopolitanism of the faunas across a large ocean, the Prototethys. We considered the biotic reaction at a seldom considered scale, namely within a single subgenus of conodont, Palmatolepis (Manticolepis). Patterns of spatial and temporal differentiation were quantified using morphometrics of its platform element. The recognized cosmopolitanism of the faunas was confirmed at this scale of variation since temporal records gathered in distant areas around the Prototethys, including the seldom documented regions located nowadays in South−East Asia, displayed similar morphological trends in response to the major F/F crisis. Beyond this overall cosmopolitanism, subtle geographic structure was evidenced but was not stable through time. Geographic differentiation was maximal shortly before the F/F crisis, suggesting that despite high sea−level, tectonics leaded to complex submarine landscapes promoting differentiation. In contrast any geographic structure was swamped out after the crisis, possibly due to a global recolonization from few favorable patches.
Diverse atrypid brachiopod faunas characterize very late Givetian-early Frasnian deposits of Devonian Transgressive-Regressive (T-R) cycle IIb in North America which feature species of Desquamatia (Seratrypa), Desquamatia (Independatrypa), Pseudoatrypa, Radiatrypa, Spinatrypina (S.), S. (Exatrypa), Spinatrypa (S.), Davidsonia, and possibly Iowatrypa Middle Frasnian faunas are not well documented in much of North America. Middle Frasnian deposits of T-R cycle IIc in the Great Basin, Iowa, and the southern Northwest Territories and Mackenzie shelf feature species of Neatrypa, Pseudoatrypa, Radiatrypa, D. (Seratrypa), Spinatrypa, and possibly Costatrypa. Radiatrypa does not carry over into late Frasnian rocks of T-R cycle IId in North America. Genera common to late Frasnian deposits of T-R cycle IId-1 in central and western North America include widespread species of Pseudoatrypa, Spinatrypa, Costatrypa, Iowatrypa. D. (Seratrypa) and Neatrypa were restricted to the tropical platforms of western Canada at that time. Very late Frasnian brachiopod faunas of T-R cycle IId-2 yield species of Pseudoatrypa, Spinatrypa, Iowatrypa, and Pseudoatrypa? (southwest US only). Available data on Late Frasnian brachiopod records in North American subtropical platforms (New Mexico and Iowa) indicate that two successive stepped late Frasnian extinction events affected those faunas, coinciding with the Lower Kellwasser Event and the Frasnian-Famennian (F-F) Event. Over half of the atrypid genera represented in late Frasnian North American faunas survived the first wave of extinctions (Lower Kellwasser Event). The surviving species, recorded in very late Frasnian deposits of Devonian T-R cycle IId-2, became extinct during the final crisis associated with the F-F Event.
The common Early Triassic (Olenekian) gastropod Turbo rectecostatus from the upper Werfen Formation of the Alps is placed in the new genus Werfenella. Elimination of the wrong or outdated generic assignments of Late Palaeozoic and Early Mesozoic gastropods to archetypical genera such as Turbo, Trochus, or Natica (all with Recent type species) represents an important step toward understanding the evolutionary history of the gastropods across the Permian/Triassic mass−extinction event. The first appearance of Werfenella in the Olenekian, as well as the origination of other groups of gastropods, suggests an early turnover in the aftermath of the end−Permian mass extinction event. The relatively large size of Werfenella (up to 35 mm) sheds doubt on assertions that all Early Triassic gastropods are microgastropods (Lilliput effect). The new genus is placed in the caenogastropod family Purpurinidae and represents its earliest occurrence. However, a placement of Werfenella in the Archaeogastropoda (Vetigastropoda) is also possible because it resembles the paraturbinid genus Chartronella. The characteristic Werfenella rectecostata–Natiria costata gastropod association from the Werfen Formation is not found in the approximately contemporaneous Sinbad Limestone of the Moenkopi Formation (Utah, USA) nor elsewhere outside Europe. This suggests that the similarities between Olenekian gastropod faunas from the Tethys and western North America are more limited than previously thought.
Late Frasnian Atrypida (Brachiopoda) from the South Urals, South Timan and Kuznetsk Basin in Russia (east Laurussian and south Siberian shelf domains in Devonian time) reveal significant generic and specific diversity in the broadly defined Frasnian-Famennian (F-F) bio-crisis time. Eighteen species of atrypid brachiopods have been recorded, representing 4 subfamilies and 10 genera. The new genus Gibberosatrypa Markovskii & Rzhonsnitskaya, and the new subgenus Spinatrypa (Plicspinatrypa) Rzhonsnitskaya are proposed. Four new species Spinatrypina (Spinatrypina) sosnovkiensis Yudina, Spinatrypa (Plicspinatrypa) rossica Rzhonsnitskaya, Iowatrypa nalivkini Rzhonsnitskaya & Sokiran, and Cartnatina(?) biohermica Yudina are described. The representatives of the Variatrypinae (including especially common Desquamatia (Desquamatia) alticoliformis), Spinatrypinae (Spinatrypina) and Atypinae (Pseudoatrypa, ?Costatrypa) are widely distributed in the studied regions. The Pseudogruenewaldtiinae are represented by Iowatrypa and Pseudogruenewaldtia, of which the first is distributed worldwide, whereas the only undoubted species of the second is restricted to South Timan, and probably represents a localized latest Frasnian descendant of Iowatrypa. The decline phase of atrypid development was controlled by a variety of environmental factors tied to the global Kellwasser events, although it was not directly triggered by anoxic conditions. The investigated atrypid brachiopods, which were all confined to lower latitudes, disappeared during the F-F mass extinction, independently of their environmental and biogeographic settings.
Four major microfacies have been recognized in the Psie Górki section and the bioclastic content indicates an open marine environment in the photic zone close to an algal shole. Sedimentological studies point to a regressive episode starting close to the Frasnian–Famennian boundary. The regressive microfacies pattern is revealed by the presence of semirestricted algal microbreccias that compose all of the lower part of the Famennian. The regression was accompanied by meteoric water invasion as the sea level fell. Seventy−six ostracod species are recorded. The ostracod assemblage, dominated by podocopids, belongs to the Eifelian ecotype and is indicative of a well−oxygenated marine environment below fair−weather wave base in the Frasnian part of the section, and of shallower environments in the base of the Famennian. No ostracod assemblage characteristic of hypoxic or semi−restricted water conditions has been recorded. The rate of extinction of ostracod species (>70%) close to the Frasnian–Famennian boundary is comparable with that known on the same level in several other sections investigated in the world. Five new ostracod species are proposed by J.−G. Casier and F. Lethiers: Selebratina vellicata, Samarella? minuta, Bairdiocypris ventrorecta, Acratia pentagona, and “Bairdia” psiegorkiensis.
The rhynchonellid species, Pammegetherhynchus kowalaensis sp. n., occurs in the late Frasnian (Early to Late Palmatolepis rhenana, and possibly early Palmatolepis linguiformis conodont zones) marly-bituminous succession at Kowala (various outcrops) in the Gałęzice Syncline, south of Kielce in the Holy Cross Mountains, Poland. The only other known species of this genus is the type species, Pammegetherhynchus merodae Sartenaer, 1977, from the late Frasnian (somewhere in the Early and Late Palmatolepis rhenana Zones) of the French Fagne (dark shales of 'Matagne' aspect), and, probably, of the Eifel ('Büdesheimer Goniatitenschiefer'). P. kowalaensis sp. n. occurred in level-bottom pioneer assemblages, thriving in reef downslope, mostly poorly-oxygenated habitats of the Kellwasser interval. The species finally disappeared near the Frasnian-Famennian boundary. The genus Pammegetherhynchus seems to be particularly suited to stressed deep-water shelf environments in the European part of the Laurussian shelf, widely distributed in this crisis time.
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The last representatives of the order Atrypida on the southern flank of the Dinant Synclinorium (Vaulx-Nismes area) in Belgium belong to Costatrypa, Spinatrypa, Spinatrypina (?Spinatrypina), Spinatrypina (Exatrypa), Iowatrypa, ?Waiotrypa, Desquamatia (Desquamatia) and Desquamatia (?Seratrypa). Among the thirteen described taxa, five are new: Spinatrypa tumuli sp. n., Iowatrypa circuitionis sp. n., ?Waiotrypa pluvia sp. n., Desquamatia (Desquamatia) quieta sp. n. and Desquamatia (?Seratrypa) derelicta sp. n. Supposed lissatrypid 'Glassia drevermanni' Maillieux, 1936 from the late Frasnian Matagne shales is assigned to the Rhynchonellida. On the southern flank of the Dinant Synclinorium and in the Philippeville Massif, the Atrypida become extinct in the Palmatolepis rhenana Zone, significantly below the Frasnian-Famennian (F-F) boundary. Their extinction coincides with the first appearance of the green and black shales of the late Frasnian Matagne Formation, recording a transgressive-hypoxic event. Based on conodont data, this event takes place earlier on the southern flank of the Dinant Synclinorium than in the Philippeville Massif.
Late Frasnian Atrypida (Brachiopoda) from the Holy Cross Mountains, Poland, include 15 taxa and were widely distributed in foreslope habitats of the declining Dyminy Reef complex. The Palmatolepis semichatovae transgression, followed by the transgressive/hypoxic Lower Kellwasser (KW) Event during the Palmatolepis rhenana Zone did not have catastrophic effects for atrypid faunas, but were rather associated with the appearance of a new species group comprising Iowatrypa, Waiotrypa, Costatrypa, Spinatrypina, Desquamatia and Radiatrypa. Stepdown demise of the biota started during the inter-KW regression, and culminated as a result of increasing stress during the Upper Kellwasser Event in the late Palmatolepis linguiformis Zone, mainly due to catastrophic sea level changes and anoxia, possibly linked to oceanic thermal changes (cooling) and nutrification pulses. The extinction pattern was diachronous and facies-controlled in this area, and the last atrypid survivors reached the Frasnian-Famennian (F-F) boundary. Increasing expansion from the adjacent deeper-water environment of the more resistant assemblages, with productids, cyrtospiriferids, athyridids and schizophoriids, occurred in the final crisis interval. This brachiopod fauna profusion characterized the earliest Famennian survival and early recovery phases of the mass extinction in this part of the Laurussian shelf, as well as the continuity of the deeper-water rhynchonellid-inarticulate biofacies across the F-F boundary. Spinatrypina (Exatrypa) relicta sp. n. is proposed as new.
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In South China latest Frasnian (Palmatolepis linguiformis Zone) the representatives of the order Atrypida (Brachiopoda) are most common in central Hunan Province. They are relatively rare in other parts of South China due to unfavourable ecologic conditions. Unlike most previously reported sections, including some sections in South China, the four Frasnian-Famennian (F-F) boundary sections examined here do not show any evidence for black shale. Atrypids are abundant and relatively diverse about 15 to 20 m below the F-F boundary (six species), and very rare about 1-2 m below the boundary (with only two species). It seems that the disappearance of most atrypids occurred well before the F-F boundary. Nine species (including Iowatrypa? qidongensis sp. n.), assigned to six genera, are discussed and described.
The Frasnian-Famennian (F-F) mass extinctions saw the global loss of all genera belonging to the tropically confined order Atrypida (and Pentamerida): though Famennian forms have been reported in the literafure, none can be confirmed. Losses were more severe during the Givetian (including the extinction of the suborder Davidsoniidina, and the reduction of the suborder Lissatrypidina to a single genus), but origination rates in the remaining suborder surviving into the Frasnian kept the group alive, though much reduced in biodiversity from the late Early and Middle Devonian. In the terminal phases of the late Palmatolepis rhenana and P. linguiformis zones at the end of the Frasnian, during which the last few Atrypidae dechned, no new genera originated, and thus the Atrypida were extirpated. There is no evidence for an abrupt termination of all lineages at the F-F boundary, nor that the Atrypida were abundant at this time, since all groups were in decline and impoverished. Atypida were well established in dysaerobic, muddy substrate, reef lagoonal and off-reef deeper water settings in the late Givetian and Frasnian, alongside a range of brachiopod orders which sailed through the F-F boundary: tropical shelf anoxia or hypoxia seems implausible as a cause for atrypid extinction. Glacial-interglacial climate cycles recorded in South America for the Late Devonian, and their synchronous global cooling effect in low latitudes, as well as loss of the reef habitat and shelf area reduction, remain as the most likely combined scenarios for the mass extinction events.
The radiolarian species Astroentactinia paronae, A. stellata, Trilonche echinata, T. grandis, T. nigra, Haplentactinia inaudita, and H. rhinophyuosa are common in late Frasnian to early Famennian rhythmic, calcareous−marly sequence of the southern Holy Cross Mts., Poland. They are known also from coeval abundant siliceous biota assemblages from the carbonate shelf of East European Platform including more than 150 taxa of radiolarians. However, in ecological terms, the moderately diverse Polish microfaunas (34 species of 12 genera) are more similar to these from Kolyma and Alaska, also marked by abundance of sphaerical entactiniids and near−absence of bilateral−symmetric Ceratoikiscidae and Palaeoscenididae. A succession of two distinctive siliceous sponges associations is established in the incipiently submerged Holy Cross carbonate platform: from an ephemeral, diverse, mostly rigid−skeletal lithistid−hexactinosan foreslope assemblage (initial phase of the late Frasnian Kellwasser Crisis), to long−lasting, basinal loose−skeletal hexactinellid−demosponge faunas (appearing abundantly just prior the Frasnian–Famennian boundary in the late Palmatolepis linguiformis Zone). Such regional blooms of marine siliceous biotas, parallel to temporary retreat of calcareous biota, are demonstrated worldwide for the Kellwasser Crisis. These suggest probable causal links with cooling pulses and at least regional, volcanically induced eutrophication.
Late Frasnian representatives of the order Athyridida from the Holy Cross Mountains, Poland, support the idea that the Laurussian basins were the places of origin and radiation of the subfamilies Athyridinae and Meristinae during the middle and early late Paleozoic. At least three new species have been identified from two localities (Łgawa Hill and Kowala) in the Gałęzice Syncline. Of these, one was probaby endemic (Merista rhenanensis sp. n.; maybe also ?Zonathyris sp. A), and two (Athyris postconcentrica sp. n. and Pachyplaxoides postgyralea gen. et sp. n.) were more widely distributed in this part of the Laurussian shelf, being known also from the East European Platform and Rheinisches Schiefergebirge, respectively. This confirms an intermediate biogeographic position of the Holy Cross Mountains area, belonging to an important centre of brachiopod origin and diversification. In contrast to other articulate brachiopods, athyridids reveal a higher rate of diversification, especially at the species (and partly also generic) level, during the global Kellwasser Crisis.
Preliminary review of taxonomy of the brachiopod order Atrypida and its stratigraphic distribution in the late Frasnian Kellwasser Crisis of several regions of Laurussia, western Siberia and South China point to their moderate diversity and stepdown but irregular extinction pattern. The distinctive character of the late Frasnian atrypid fauna is emphasised by several relict genera, marked by recurrent and possibly aberrant characters (mainly in ornamentation types), tendency to size reduction and homeomorphy in some taxa. The transgressive/hypoxic Lower Kellwasser Event and preceding eustatic changes during the Palmatolepis rhenana Zone had only a regional destructive effect, and were linked rather to an enhanced dispersal of the last generic set of atrypids. The Variatrypinae, Spinatrypinae and Iowatrypa-group seem to belong to the latest surviving atrypids. The final demise of the remaining atrypids (and some other articulate brachiopods, e.g., gypidulids) coincided with the transgressive/hypoxic Upper Kellwasser Event, followed by catastrophic eustatic fall during the late Palmatolepis linguiformis Zone (F-F Event). This was probably exacerbated by accelerated submarine volcano-hydrothermal activity, and consequent progressive regional eutrophication, and climatic destabilization. The level-bottom rhynchonellid-inarticulate biofacies crosses the fatal F-F boundary horizon without major changes. No reliable data exist for the presence of atrypids in the Famennian survival and recovery biota, even for the smooth lissatrypid Peratos. Sustained competition from radiating and diversifying productid-cyrtospirifrid-athyrid faunas may have provide an additional biotic factor in the collapse of the Frasnian shelly benthos at the time of sfress, as well as in a post-extinction offshore repopulation from inner shelf habitats.
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