Recovery of benthic marine communities from the end‐Permian mass extinction at the low latitudes of eastern Panthalassa

Based on the quantitative community analysis using species‐level identifications, we track the restoration of benthic ecosystems after the end‐Permian mass extinction throughout the Lower Triassic of the western USA. New data on the palaeoecology of the Thaynes Group and Sinbad Formation are provided, which fill a gap between the recently studied palaeoecology of the Griesbachian–Dienerian Dinwoody Formation and the Spathian Virgin Formation. In the Sinbad Formation and Thaynes Group, 17 species (12 genera) of bivalves, 7 species and genera of gastropods and 2 species and genera of brachiopods are recognized. The new bivalve genus Confusionella (Pteriidae) is described. A comprehensive review of the whole Lower Triassic succession of benthic ecosystems of the western USA indicates that mid‐ and inner shelf environments show incipient recovery signals around the Griesbachian–Dienerian transition, during the Smithian and, most profound, during the early Spathian. Ecological data from youngest strata of the Dinwoody Formation as well as stratigraphic ranges of species suggest that the late Dienerian was likely a time interval of environmental stress for benthic ecosystems. Despite some evidence for short‐term environmental disturbances (e.g. shift of dominant taxa, transient drop in alpha‐diversity) during the Smithian–Spathian transition, benthic ecosystems did not show any notable taxonomic turnover at that time, in contrast to the major crisis that affected ammonoids and conodonts. Whereas alpha‐diversity of benthic communities generally increased throughout the Early Triassic, beta‐diversity remained low, which reflects a persistently wide environmental range of benthic species. This observation is in accordance with a recently proposed model that predicts a time lag between increasing within‐habitat diversity (alpha‐diversity) and the onset of taxonomic differentiation between habitats (beta‐diversity) during biotic recoveries from mass extinction events. The observation that beta‐diversity had not significantly increased during the Early Triassic might also provide an explanation for the comparably sluggish increase in benthic diversity during that time, which has previously been attributed to persistent environmental stress.     

Conodonts from the Early Triassic microbialite of Guangxi (South China): implications for the definition of the base of the Triassic System

We describe a new Early Triassic (Griesbachian) succession of conodont faunas from a high‐resolution sampling of the basal Early Triassic microbial limestone and the base of the overlying unit at the Wuzhuan section (Nanpanjiang Basin, Guangxi, South China). The microbial limestone records the earliest phase of the Early Triassic biotic recovery after the end‐Permian mass extinction. For the first time, rich conodont faunas are reported from within the microbialite. The faunas from Wuzhuan are largely dominated by anchignathodontids, including several Isarcicella species, which were previously documented only from strata above the microbialite. A total of 14 conodont species assigned to three genera is recorded from the Wuzhuan section. Starting from the base of the microbialite upwards, several species are sequentially added to the conodont assemblage. The alpha diversity peaks at the top of the microbialite. The conodont record in the considered microbialite interval at Wuzhuan is presumably unaffected by local ecological changes. It therefore more likely represents an evolutionary rather than an ecological pattern. We compare the Wuzhuan's conodont record with a well‐supported phylogenetic model and suggest that the sequence of first occurrences at Wuzhuan is the closest to the ‘true’ sequence of evolutionary events that took place during this Griesbachian radiation of anchignathodontids. Based on comparisons with the GSSP section at Meishan, we suggest further that the first occurrence of Hindeodus parvus in Meishan does not correspond to its first appearance datum.     

An unusually diverse mollusc fauna from the earliest Triassic of South China and its implications for benthic recovery after the end-Permian biotic crisis

A single carbonate coquinoid lens from the Griesbachian (Early Triassic) of Shanggan, South China, yielded 11 bivalve species described in this study in addition to four gastropod and one ammonoid species reported elsewhere. This makes the Shanggan fauna one of the richest mollusc faunas from the early post-extinction interval after the end-Permian mass extinction event. Four of the present genera are long-term survivors, five are holdovers that went extinct at the end of the Griesbachian or later in the Early Triassic, and seven first appear in the Griesbachian. Three new bivalve species are described: Myalinella newelli nov. sp., Scythentolium scutigerulus nov. sp., and Eumorphotis shajingengi nov. sp. The genus Astartella, previously assumed to have vanished at the end of the Permian, is reported for the first time from the Early Triassic, which also removes Astartidae from Early Triassic Lazarus taxa. The small growth size of the Astartella specimens supports an earlier hypothesis that many of the Early Triassic Lazarus taxa did not survive in unknown refuges but were simply overlooked due to the scarcity of easily observable large-sized specimens. Ecologically, a comparatively high proportion of infaunal bivalve species (4/11) is remarkable for the early post-extinction interval, supporting the impression of a relatively advanced recovery state. Moreover, abundance-data of the bivalve-gastropod community reveal a remarkably low dominance index (D = 0.17) that is suggestive for advanced recovery and stable environmental conditions. It is proposed that the Shanggan fauna represents a late Griesbachian benthic recovery event that coincided with the appearance of similarly diverse benthic faunas in Oman and Primorye. A high proportion of genera that have previously not been reported from the Early Triassic indicate that the prevalence of poor preservation conditions is a major obstacle in identifying early phases of recovery from the greatest crisis in the history of metazoan life. The early recovery of benthic faunas reported in this study questions previous claims of a prolonged lag phase as a consequence of the extraordinary extinction magnitude or the persistence of adverse environmental conditions.     

Recovery dynamics of benthic marine communities from the Lower Triassic Werfen Formation,northern Italy

The Lower Triassic Werfen Formation of northern Italy represents an important archive for Early Triassic ecosystems. Based on quantitative community analysis using species level identifications, we reconstruct the recovery of benthic ecosystems after the end‐Permian mass extinction throughout this unit. The analysis of benthic macrofossil communities shows that incipient recovery has taken place during the Griesbachian and the Griesbachian–Dienerian transition. A probable deterioration in environmental conditions is observed towards the end of the Dienerian. The Smithian part of the Werfen Formation is characterized by high siliciclastic input, which ceases around the Smithian‐Spathian boundary. The Spathian marks the definitive phase of recovery in the Werfen Formation. The comparison of this pattern with other palaoegeographical regions suggests that both the Griesbachian recovery and the Dienerian decline were of inter‐regional if not global extent, whereas the Smithian diversity low in the Werfen Formation is a local signal. In contrast to the recovery dynamics of ammonoids and conodonts, the Smithian–Spathian boundary interval was no caesura for benthic ecosystems. The Spathian recovery pulse is possibly also an inter‐regional event, at least in the palaeotropics. These results are in contrast with the previously proposed scenario of persistent hostile conditions during the Griesbachian time interval and highlight an initial recovery phase restricted to Griesbachian times. Instead, the apparently sluggish recovery of benthic ecosystems was at least partly shaped by set‐backs due to short‐term environmental perturbations during the Dienerian.     

Diversity partitioning in Permian and Early Triassic benthic ecosystems of the Western USA: a comparison

This paper compares the relative contributions of within-habitat diversity [alpha-diversity] and between-habitat-diversity [beta-diversity] to regional diversity [gamma-diversity] in marine benthic communities of the western US before and after the end-Permian mass extinction. We found that presumably cool-water faunas from the Permian Gerster Limestone and the Park City Formation had low alpha- and beta-diversities, comparable to those of low diverse faunas of the Early Triassic. In contrast, tropical Permian faunas had much higher alpha-diversities and a variable pattern of beta-diversity: Whereas faunas of space-limited bioherms show a positive correlation between beta-diversity and gamma-diversity, beta-diversity in level-bottom faunas is elevated only when gamma-diversity is very high (>250 species). This contrasting pattern probably reflects differential effects of interspecific competition on habitat partitioning. In low-competitive level-bottom faunas, species are able to coexist until competition forces species into their ecological optima, thereby increasing beta-diversity. This effect occurs at much lower gamma-diversities in more competitive reef-bound faunas, causing the observed positive correlation between beta- and gamma-diversity. We suggest that differences in the level of interspecific competition and hence diversity partitioning between Permian and Triassic benthic communities result from the higher average metabolic rates in the Mesozoic mollusc-dominated benthos in contrast to their Permian counterparts.     

Diversity and palaeoecology of a highly diverse Late Triassic marine biota from the Cassian Formation of north Italy

A sample of marine invertebrates from the Late Triassic Cassian Formation (north Italy) yielded one of the most diverse Early Mesozoic fossil assemblages ever reported (c. 170 species). The assemblage was found in basin clays, but was transported from nearby carbonate platforms as indicated by fragmentation, microbial encrustation and the presence of coated grains and ooids. Most of the specimens are small (< 1 cm) reflecting both, small adult sizes and size sorting during transport. Rarefaction analysis suggests that diversity of surface collection and bulk sampling is the same. However, rank abundance, species richness and taxonomic composition differ strongly according to sampling method. Low‐grade lithification of the sediments is the main reason that high diversity can be recognized, because it facilitates disaggregation and finding of small molluscs. Sample standardization shows that the studied assemblage is much more diverse than known Early Triassic assemblages. However, its diversity is similar to that of Anisian assemblages. This suggests that recovery from the end‐Permian mass‐extinction was quite advanced in the Middle Triassic and alpha‐diversity remained high until the Late Triassic. According to current models, Early Triassic and Anisian faunas match the niche overlap phase of recovery during which diversity is built up by increasing alpha‐diversity, whereas beta‐diversity rises slowly. Subsequently, habitat width of species contracts because of increasing competition, making beta‐diversity the principal drive of overall diversity increase. The diversity pattern of various Late Triassic Cassian associations meets the predictions for the transition from the niche overlap to the habitat contraction phase.: Triassic, Cassian Formation, palaeoecology, diversity, mollusc dominance.     

Permian–Triassic Osteichthyes (bony fishes): diversity dynamics and body size evolution

The Permian and Triassic were key time intervals in the history of life on Earth. Both periods are marked by a series of biotic crises including the most catastrophic of such events, the end‐Permian mass extinction, which eventually led to a major turnover from typical Palaeozoic faunas and floras to those that are emblematic for the Mesozoic and Cenozoic. Here we review patterns in Permian–Triassic bony fishes, a group whose evolutionary dynamics are understudied. Based on data from primary literature, we analyse changes in their taxonomic diversity and body size (as a proxy for trophic position) and explore their response to Permian–Triassic events. Diversity and body size are investigated separately for different groups of Osteichthyes (Dipnoi, Actinistia, ‘Palaeopterygii’, ‘Subholostei’, Holostei, Teleosteomorpha), within the marine and freshwater realms and on a global scale (total diversity) as well as across palaeolatitudinal belts. Diversity is also measured for different palaeogeographical provinces. Our results suggest a general trend from low osteichthyan diversity in the Permian to higher levels in the Triassic. Diversity dynamics in the Permian are marked by a decline in freshwater taxa during the Cisuralian. An extinction event during the end‐Guadalupian crisis is not evident from our data, but ‘palaeopterygians’ experienced a significant body size increase across the Guadalupian–Lopingian boundary and these fishes upheld their position as large, top predators from the Late Permian to the Late Triassic. Elevated turnover rates are documented at the Permian–Triassic boundary, and two distinct diversification events are noted in the wake of this biotic crisis, a first one during the Early Triassic (dipnoans, actinistians, ‘palaeopterygians’, ‘subholosteans’) and a second one during the Middle Triassic (‘subholosteans’, neopterygians). The origination of new, small taxa predominantly among these groups during the Middle Triassic event caused a significant reduction in osteichthyan body size. Neopterygii, the clade that encompasses the vast majority of extant fishes, underwent another diversification phase in the Late Triassic. The Triassic radiation of Osteichthyes, predominantly of Actinopterygii, which only occurred after severe extinctions among Chondrichthyes during the Middle–Late Permian, resulted in a profound change within global fish communities, from chondrichthyan‐rich faunas of the Permo‐Carboniferous to typical Mesozoic and Cenozoic associations dominated by actinopterygians. This turnover was not sudden but followed a stepwise pattern, with leaps during extinction events.     

Palaeoecological significance of a new Griesbachian (Early Triassic) gastropod assemblage from Oman

A new Early Triassic (Griesbachian) gastropod fauna from the Al Jil Formation of Oman is described. Early Triassic faunas from elsewhere (e.g. the Italian Dolomites and the western USA) are typically of low diversity and high dominance, usually attributed to environmental stress in the immediate aftermath of the end-Permian mass extinction event. The new Oman fauna has, by contrast, relatively high diversity, low dominance and a more even spread of individuals between taxa. It is the most diverse Griesbachian fauna known to date. This is attributed to the favourable (i.e. well-oxygenated) conditions under which the fauna lived. This uncharacteristic Griesbachian gastropod fauna demonstrates that, in the absence of oceanic anoxia, biotic recovery after the end-Permian extinction event may occur surprisingly quickly (within one conodont zone). The fauna is also partially silicified, which has increased its preservation potential relative to other Griesbachian gastropod assemblages. However, only one reappearing Lazarus genus is present in the Oman fauna. This suggests that there was some other control on the abundance of Lazarus genera at this time, other than the absence of silicified faunas as previously suggested.     

Survival patterns of macrobenthic marine assemblages during the end-Permian mass extinction in the western Tethys (Dolomites, Italy)

The ecological competition between brachiopods and bivalves is analysed by means of a quantitative palaeoecologic method applied on four assemblages located within a short stratigraphic interval, approximately 2 m thick, in the lower Tesero Member of the Werfen Formation (in the Southern Alps). The assemblages originate from the Tesero, Bulla and Sass de Putia sections. The analysed stratigraphic interval, uppermost Changhsingian in age, is located between the early and heaviest phase of the end-Permian mass extinction, which occurred across the Bellerophon/Werfen formational boundary (Event Boundary), and the Permian/Triassic boundary (Chronological Boundary), when nearly all the Permian stenotopic holdovers disappeared.These assemblages are characterised by small sized skeletons (“Lilliput effect”), which represent an adaptive survival strategy in stressed and harsh habitats resulting from the climatic and palaeoceanographic changes connected with the mass extinction. The Tesero assemblages are dominated by rhynchonelliform brachiopod Orbicoelia (bed CNT10) or Streptorhynchus (bed CNT11A), which were mostly attached at the top of shallow microbialitic mounds. These assemblages are again dominated by Permian stenotopic taxa and show a Palaeozoic structure. The Tesero habitat, which again permitted the survival of brachiopods, represented one of the last refuges in the western Tethys. On the contrary, the Bulla (BU9-10) and Sass de Putia (wPK13A) assemblages are bivalve-dominated, and thus show an ecologic structure typical of Early Triassic post-extinction marine benthic communities or Palaeozoic stressed marine communities. The bivalve-dominated assemblages proliferated in prevailing muddy siliciclastic substrates, with brief episodes of microbial algal growth. The most important environmental limiting factors and leading causes of end-Permian mass extinction are discussed in terms of palaeoautecologic and palaeosynecologic analysis.The different taxonomic composition and ecologic structure of the assemblages is related to palaeogeography, including water depth and connections with the open sea. The brachiopod-dominated assemblage, exclusive of the Tesero section, proliferated in microbial carbonate habitats in near-shore environments. The bivalve-dominated assemblages, which were more widespread than the brachiopod assemblages in the Dolomites and also occurred in other western Tethys localities, occur in more open and deeper marine environments. In the western Tethys margins, the local distribution of mixed faunas suggests that the extinction of Permian stenotopic taxa was caused by the onset of poisonous water on the shelves originating from deep marine environments.This extinction pattern appears to be a regional phenomenon and does not seem be applicable on a global scale. The extinction events were controlled by a complex network of interactive factors and the survival of faunal elements was probably stochastic.     

CALIBRATED DIVERSITY,TREE TOPOLOGY AND THE MOTHER OF MASS EXTINCTIONS: THE LESSON OF TEMNOSPONDYLS

Abstract: Three family‐level cladistic analyses of temnospondyl amphibians are used to evaluate the impact of taxonomic rank, tree topology, and sample size on diversity profiles, origination and extinction rates, and faunal turnover. Temnospondyls are used as a case study for investigating replacement of families across the Permo‐Triassic boundary and modality of recovery in the aftermath of the end‐Permian mass extinction. Both observed and inferred (i.e. tree topology‐dependent) values of family diversity have a negligible effect on the shape of the diversity curve. However, inferred values produce both a flattening of the curve throughout the Cisuralian and a less pronounced increase in family diversity from Tatarian through to Induan than do observed values. Diversity curves based upon counts of genera and species display a clearer distinction between peaks and troughs. We use rarefaction techniques (specifically, rarefaction of the number of genera and species within families) to evaluate the effect of sampling size on the curve of estimated family‐level diversity during five time bins (Carboniferous; Cisuralian; Guadalupian–Lopingian; Early Triassic; Middle Triassic–Cretaceous). After applying rarefaction, we note that Cisuralian and Early Triassic diversity values are closer to one another than they are when the observed number of families is used; both values are also slightly higher than the Carboniferous estimated diversity. The Guadalupian–Lopingian value is lower than raw data indicate, reflecting in part the depauperate land vertebrate diversity from the late Cisuralian to the middle Guadalupian (Olson’s gap). The time‐calibrated origination and extinction rate trajectories plot out close to one another and show a peak in the Induan, regardless of the tree used to construct them. Origination and extinction trajectories are disjunct in at least some Palaeozoic intervals, and background extinctions exert a significant role in shaping temnospondyl diversity in the lowermost Triassic. Finally, species‐, genus‐, and family trajectories consistently reveal a rapid increase in temnospondyl diversity from latest Permian to earliest Triassic as well as a decline near the end of the Cisuralian. However, during the rest of the Cisuralian family diversity increases slightly and there is no evidence for a steady decline, contrary to previous reports.     

On Permian and Triassic insect faunas in relation to biogeography and the Permian-Triassic crisis

The taxonomic diversity dynamics of pterygote insects in the Permian and Triassic at the family/age level are considered. Different metrics of taxonomic diversity are compared. Biogeographic and taphonomic aspects of changes in the composition of insect faunas in the Permian and about the P-T transition are discussed. Some changes in the Permian insect faunas are of a biogeographic nature and do not indicate global changes in diversity. Insects with aquatic immatures were rather common in the Permian and Early Triassic, but these immatures are well represented in only few localities.     

Trace fossil evidence for restoration of marine ecosystems following the end-Permian mass extinction in the Lower Yangtze region,South China

Unlike the high-abundance, low-diversity macrofaunas that characterize many Early Triassic benthic palaeocommunities, ichnofossils were relatively common in the aftermath of the end-Permian mass extinction worldwide. Ichnofossils therefore are a good proxy for ecosystem recovery after the end-Permian biotic crisis. This paper documents 14 ichnogenera and one problematic form from Lower Triassic successions exposed in the Lower Yangtze region, South China. Post-extinction ichnodiversity remained rather low throughout the Griesbachian–early Smithian period and abruptly increased in the late Smithian. However, several lines of evidence, including extent of bioturbation, burrow size, trace-fossil complexity, and tiering levels, indicate that diversification of ichnotaxa in the late Smithian did not signal full marine ecosystem recovery from the Permian/Triassic (P/Tr) mass extinction. Marine ichnocoenoses did not recover until the late Spathian in South China. The marginal sea provided hospitable habitats for tracemakers to proliferate in the aftermath of the end-Permian mass extinction.     

Olenekian (Early Triassic) bivalves from the Salt Range and Surghar Range,Pakistan

Abstract: Based on newly collected material from the uppermost Smithian and lower to middle Spathian (Olenekian, Lower Triassic) of the Salt Range and Surghar Range (Pakistan), 15 bivalve species belonging to 11 genera are described, including two new genera, Eobuchia and Dimorphoconcha, and one new species, Palaeoneilo? fortistriata. Eobuchia gen. nov. is placed in a new subfamily, the Eobuchiinae, which differs from the Buchiinae in having an almost planar and only moderately inclined or offset right anterior auricle. Inclination of the right anterior auricle is proposed as a synapomorphy of the revised suborder Monotidina, which includes the Buchiidae, Monotidae, Oxytomidae and, tentatively, the Dolponellidae. The Pseudomonotidae, Chaenocardiidae and Claraiidae are discussed as candidate ancestors of the Monotidina. Dimorphoconcha gen. nov., provisionally placed in the Limidae, is a morphologically unusual genus characterized by a globose shell centre and a strongly plicate fringe. Permophorus costatus, which was previously known exclusively from Permian strata, is reported from the Spathian of the Surghar Range. This record extends the range of P. costatus for at least 8 Myr and makes it the first reported Lazarus species, with an outage of more than 2 Myr after the end‐Permian mass extinction. Ten of 15 species recognized in this study have not been reported from other regions, which may indicate increasing provincialism towards the end of the Early Triassic, or, alternatively, reflect the still insufficient knowledge of benthic faunas from the epoch that followed the greatest crisis in the history of life.     

The biogeography of Early Triassic ammonoid faunas: Clusters, gradients, and networks

The aim of this paper is to quantitatively investigate the spatial and temporal biogeographical relationships of the recovery of ammonoid faunas after the Permian-Triassic mass extinction using three complementary numerical approaches among which is a new, non-hierarchical clustering strategy. The faunal data set consists of a taxonomically homogenised compilation of the spatial and temporal occurrences of ammonoid genera within 20 Early Triassic Tethyan and Panthalassic sites ranging from 40°S to 70°N in palaeolatitudes. In addition to hierarchical cluster analysis (hCA) and nonmetric multidimensional scaling (NMDS), we introduce a third, new non-hierarchical clustering technique allowing the visualisation of a nonmetric interassemblages similarity structure as a connected network constructed without inferring additional internal nodes. The resulting network, which we call a “Bootstrapped Spanning Network” (BSN), allows the simultaneous identification of partially or totally nested as well as gradational linear or reticulated biogeographical structures.The identified interlocalities relationships indicate that the very beginning of the Early Triassic (Griesbachian) corresponds to a very simple biogeographical context, representing a time of great cosmopolitanism for ammonoids. This context shifts rapidly to a more complex configuration indicative of a more endemic and latitudinally-restricted distribution of the ammonoids during the middle and late Early Triassic (Smithian and Spathian). From an evolutionary dynamic point of view, our results illustrate a very rapid (less than ca. 1.4 myr) Early Triassic recovery of the ammonoid faunas, in contrast to many other marine organisms. This recovery is linked with a marked increase in the overall biogeographical heterogeneity, and parallels the formation of a latitudinal gradient of taxonomic richness, which may be essentially controlled by the progressive intensification of the gradient of sea surface temperature. From a methodological point of view, we show that a BSN is a simple, intuitively legible picture of the nested as well as gradational taxonomic similarity relationships, hence providing a good synthesis (and additional insights) between hierarchical clustering and ordination in reduced space results.     

Proharpoceras Chao: a new ammonoid lineage surviving the end-Permian mass extinction

Based on new, bed-rock controlled material from northwestern Guangxi and Oman, the Early Triassic genus Proharpoceras Chao is shown to be a representative of Otocerataceae. Character analysis excludes a direct link with the Griesbachian Otoceratidae and favours a derivation of Proharpoceras from the late Permian Anderssonoceratidae. The biostratigraphic range of Proharpoceras is restricted to the Smithian and its biogeographic distribution comprises Oman, South China, and Primorye, thus indicating an essentially low palaeolatitudinal distribution. Proharpoceras has no apparent relatives among other Early and Middle Triassic Ceratitida and is thus considered to be the last representative of Otocerataceae. This offshoot of the late Permian Anderssonoceratidae implies that an additional ammonoid lineage survived the end Permian extinction and that it dwindled away for some 2 Myr before going extinct.     

Microbialites and trace fossils from a Middle Triassic restricted carbonate ramp in the Catalan Basin,Spain: evaluating environmental and evolutionary controls in an epicontinental setting

The timing of recovery after the end‐Permian mass extinction has been a matter of debate, with some authors favouring a more rapid faunal recovery during the Early Triassic and others considering a more protracted biotic reestablishment spanning until the Middle Triassic. In this work, we investigated the lowermost Middle Triassic (Ladinian) carbonate deposits in the Catalan Basin to evaluate the potential environmental mechanisms and evolutionary constrains involved in the kilometre‐scale predominance of microbialites and the low‐diversity and high‐density Planolites association in a low‐latitude epicontinental setting. The studied sedimentary succession records the development from a low‐gradient, homoclinal microbial‐dominated carbonate ramp evolving towards a slightly inclined swell‐dominated type. Sedimentological analysis suggests that facies heterogeneity was controlled by pulses of syn‐rift tectonic activity, which compromised Peri‐Tethyan basin connectivity, reducing palaeobathymetry gradients. Although the monospecific nature of the studied trace‐fossil association may reflect the delayed recovery after the end‐Permian mass extinction, this is inconsistent with widespread, relatively high‐diversity ichnofaunas in carbonates elsewhere in the region. Since other Ladinian basins were characterized by the recurrence of microbial carbonates, low‐diversity ichnoassemblages and limited skeletal production, we hypothesize that shallow and restricted carbonate ramp settings harboured limited ecological complexity and widespread opportunistic colonization of the sediment when compared to coeval open marine locations.     

Diverse earliest Triassic ostracod fauna of the non‐microbialite‐bearing shallow marine carbonates of the Yangou section,South China

Since diverse ostracod faunas in the immediate aftermath of the latest Permian mass extinction are mainly found within Permian–Triassic boundary microbialites (PTBMs), the idea of an ostracod ‘microbial‐related refuge’ has been proposed. Here, we report a diversified earliest Triassic ostracod fauna from the Yangou section in South China, where no PTBMs were deposited, providing evidence inconsistent with this ‘microbial‐related refuge’ hypothesis. In addition, a significant ostracod extinction is recorded, corresponding with the earliest Triassic mass extinction (ETME). This ETME of ostracods is associated with size increases and a length/height ratio (L/H) decrease, indicating varied evolutionary patterns of shape and size of ostracods through the Permian–Triassic (P‐Tr) extinction events. Although the nature of these biotic changes is somewhat unclear, the temporally varied ‘refuge zone’ scenario provides us with a window to reconstruct the environmental dynamics of ecosystem changes during the P‐Tr transition.     

Survival and recovery of calcareous foraminifera pursuant to the end-Permian mass extinction

Ninety-one percent of calcareous foraminiferal genera became extinct during the end-Permian mass extinction. The Early Triassic Epoch was a survival phase characterized by a short-lived proliferation of disaster forms and then a prolonged interval of low diversity. The orders Miliolida and Lagenida experienced limited taxonomic re-diversification in Early and Late Anisian time, respectively. All fusulinoidean fusulinides became extinct in Late Permian time, and only two non-fusulinoidean genera persisted into the Early Triassic. Triassic fusulinides diversified to just five genera before the order became entirely extinct in Late Triassic time. Involutinides originated in Olenekian time from an unknown ancestor. They did not significantly diversify until Late Triassic time. To cite this article: J.R. Groves, D. Altiner, C. R. Palevol 4 (2005).     

Competition in slow motion: the unusual case of benthic marine communities in the wake of the end‐Permian mass extinction

Changes of community structure in response to competition usually take place on timescales that are much too short to be visible in the geological record. Here we report the notable exception of a benthic marine community in the wake of the end‐Permian mass extinction, which is associated with the microbial limestone facies of the earliest Triassic of South China. The newly reported fauna is well preserved and extraordinarily rich (30 benthic macroinvertebrate species, including the new species Astartella? stefaniae (Bivalvia) and Eucochlis obliquecostata (Gastropoda)) and stems from an environmentally stable setting providing favourable conditions for benthic organisms. Whereas changes in the taxonomic composition are negligible over the observed time interval of 10–100 ka, three ecological stages are identified, in which relative abundances of initially rare species continuously increased at the cost of previously dominant species. Concomitant with the changes of dominant species is an increase in faunal evenness and heterogeneity. In the absence of both environmental and taxonomic changes, we attribute this pattern to the long‐term effects of interspecific competition, which acted at an unusually slow pace because the number of competing species and potential immigrants was dramatically reduced by the end‐Permian mass extinction. We suggest that these non‐actualistic conditions led to decreased rates of niche differentiation and hence to the delayed rediversification of benthos that characterizes the aftermath of the greatest Phanerozoic mass extinction event. A hyperbolic diversification model is proposed, which accounts for the positive relationship between the intensity of interspecific competition and the rate of niche differentiation and resolves the conundrum of delayed rediversification at a time when niche space was largely vacated.     

Asteriacites lumbricalis von schlotheim 1820: Ophiuroid trace fossils from the lower triassic thaynes formation,central Utah

Several trace fossils of burrowing ophiuroids, Asteriacites lumbricalis von Schlotheim 1820, occur on slabs of thinly‐bedded silty limestone in the lower Spathian part of the Lower Triassic Thaynes Formation in central Utah. This is the first record of Asteriacites in the Lower Triassic of North America. Their occurrence adds information to studies of the recovery of marine faunas following the Permian extinctions, including the oxygénation of benthic sediments.