A Unitary Association-based conodont biozonation of the Smithian-Spathian boundary (Early Triassic) and associated biotic crisis from South China.

The Smithian-Spathian boundary (SSB) crisis played a prominent role in resetting the evolution and diversity of the nekton (ammonoids and conodonts) during the Early Triassic recovery. The late Smithian nektonic crisis culminated at the SSB, ca. 2.7 Myr after the Permian-Triassic boundary mass extinction. An accurate and high-resolution biochronological frame is needed for establishing patterns of extinction and re-diversification of this crisis. Here, we propose a new biochronological frame for conodonts that is based on the Unitary Associations Method (UAM). In this new time frame, the SSB can thus be placed between the climax of the extinction and the onset of the re-diversification. Based on the study of new and rich conodont collections obtained from five sections (of which four are newly described here) in the Nanpanjiang Basin, South China, we have performed a thorough taxonomical revision and described one new genus and 21 new species. Additionally, we have critically reassessed the published conodont data from 16 other sections from South China, and we have used this new, standardized dataset to construct the most accurate, highly resolved, and laterally reproducible biozonation of the Smithian to early Spathian interval for South China. The resulting 11 Unitary Association Zones (UAZ) are intercalibrated with lithological and chemostratigraphical (δ13Ccarb) markers, as well as with ammonoid zones, thus providing a firm basis for an evolutionary meaningful and laterally consistent definition of the SSB. Our UAZ8, which is characterized by the occurrence of Icriospathodus ex gr. crassatus, Triassospathodus symmetricus and Novispathodus brevissimus, is marked by a new evolutionary radiation of both conodonts and ammonoids and is within a positive peak in the carbon isotope record. Consequently, we propose to place the SSB within the separation interval intercalated between UAZ7 and UAZ8 thus leaving some flexibility for future refinement and updating. Supplementary Information: The online version contains supplementary material available at 10.1186/s13358-022-00259-x.

The main stage of recovery after the end-Permian mass extinction: taxonomic rediversification and ecologic reorganization of marine level-bottom communities during the Middle Triassic.

The recovery of marine life from the end-Permian mass extinction event provides a test-case for biodiversification models in general, but few studies have addressed this episode in its full length and ecological context. This study analyses the recovery of marine level-bottom communities from the end-Permian mass extinction event over a period of 15 Ma, with a main focus on the previously neglected main phase during the Middle Triassic. Our analyses are based on faunas from 37 lithological units representing different environmental settings, ranging from lagoons to inner, mid- and outer ramps. Our dataset comprises 1562 species, which belong to 13 higher taxa and 12 ecological guilds. The diversification pattern of most taxa and guilds shows an initial Early Triassic lag phase that is followed by a hyperbolic diversity increase during the Bithynian (early middle Anisian) and became damped later in the Middle Triassic. The hyperbolic diversity increase is not predicted by models that suggest environmental causes for the initial lag phase. We therefore advocate a model in which diversification is primarily driven by the intensity of biotic interactions. Accordingly, the Early Triassic lag phase represents the time when the reduced species richness in the wake of the end-Permian mass extinction was insufficient for stimulating major diversifications, whereas the Anisian main diversification event started when self-accelerating processes became effective and stopped when niche-crowding prevented further diversification. Biotic interactions that might drive this pattern include interspecific competition but also habitat construction, ecosystem engineering and new options for trophic relationships. The latter factors are discussed in the context of the resurgence of large carbonate platforms, which occurred simultaneously with the diversification of benthic communities. These did not only provide new hardground habitats for a variety of epifaunal taxa, but also new options for grazing gastropods that supposedly fed from microalgae growing on dasycladaceans and other macroalgae. Whereas we do not claim that changing environmental conditions were generally unimportant for the recovery of marine level-bottom communities, we note that their actual role can only be assessed when tested against predictions of the biotic model.

Sedimentary organic matter from a cored Early Triassic succession, Georgetown (Idaho, USA).

The plant fossil record from Lower Triassic sedimentary successions of the Western USA is extremely meager. In this study, samples from a drill core taken near Georgetown, Idaho, were analyzed for their palynological content as well as their stable carbon isotope composition. The concentration of palynomorphs is generally low. The lowermost part of the drilled succession represents Dinwoody/Woodside Formation and contains spore and pollen assemblages with Permian and Early Triassic affinity. Representatives of lycophytes (Densoisporites spp., Lundbladisporites spp.) were found in the overlying Meekoceras Limestone, in agreement with middle Smithian assemblages elsewhere. Ammonoids and conodonts are extremely rare, but confirm a middle Smithian age. Bulk organic and carbonate carbon isotope composition provide a stratigraphic framework. Carbonate carbon isotope compositions are compatible with the Smithian-Spathian global trend, with a middle Smithian shift towards lower δ13C values followed by a late Smithian shift towards higher values. Bulk organic carbon isotope compositions have been influenced by changes in the constitution of organic matter. A comparison with other paired carbon isotope datasets from the same basin is difficult due to lithostratigraphic inconsistencies (Hot Springs, ID) or biochemical mediated disturbance of isotope signals (Mineral Mountains, UT).

Timing of global regression and microbial bloom linked with the Permian-Triassic boundary mass extinction: implications for driving mechanisms.

New high-resolution U-Pb dates indicate a duration of 89 ± 38 kyr for the Permian hiatus and of 14 ± 57 kyr for the overlying Triassic microbial limestone in shallow water settings of the Nanpanjiang Basin, South China. The age and duration of the hiatus coincides with the Permian-Triassic boundary (PTB) and the extinction interval in the Meishan Global Stratotype Section and Point, and strongly supports a glacio-eustatic regression, which best explains the genesis of the worldwide hiatus straddling the PTB in shallow water records. In adjacent deep marine troughs, rates of sediment accumulation display a six-fold decrease across the PTB compatible with a dryer and cooler climate as indicated by terrestrial plants. Our model of the Permian-Triassic boundary mass extinction (PTBME) hinges on the synchronicity of the hiatus with the onset of the Siberian Traps volcanism. This early eruptive phase released sulfur-rich volatiles into the stratosphere, thus simultaneously eliciting a short-lived ice age responsible for the global regression and a brief but intense acidification. Abrupt cooling, shrunk habitats on shelves and acidification may all have synergistically triggered the PTBME. Subsequently, the build-up of volcanic CO2 induced a transient cool climate whose early phase saw the deposition of the microbial limestone.

Severest crisis overlooked-Worst disruption of terrestrial environments postdates the Permian-Triassic mass extinction.

Generally Early Triassic floras are believed to be depauperate, suffering from protracted recovery following the Permian-Triassic extinction event. Here we present palynological data of an expanded East Greenland section documenting recovered floras in the basal Triassic (Griesbachian) and a subsequent fundamental floral turnover, postdating the Permian-Triassic boundary extinction by about 500 kyrs. This event is marked by a swap in dominating floral elements, changing from gymnosperm pollen-dominated associations in the Griesbachian to lycopsid spore-dominated assemblages in the Dienerian. This turnover coincides with an extreme δ(13)Corg negative shift revealing a severe environmental crisis, probably induced by volcanic outbursts of the Siberian Traps, accompanied by a climatic turnover, changing from cool and dry in the Griesbachian to hot and humid in the Dienerian. Estimates of sedimentation rates suggest that this environmental alteration took place within some 1000 years. Similar, coeval changes documented on the North Indian Margin (Pakistan) and the Bowen Basin (Australia) indicate the global extent of this crisis. Our results evidence the first profound disruption of the recovery of terrestrial environments about 500kyrs after the Permian-Triassic extinction event. It was followed by another crisis, about 1myrs later thus, the Early Triassic can be characterised as a time of successive environmental crises.

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.

Early Triassic marine biotic recovery: the predators' perspective.

Examining the geological past of our planet allows us to study periods of severe climatic and biological crises and recoveries, biotic and abiotic ecosystem fluctuations, and faunal and floral turnovers through time. Furthermore, the recovery dynamics of large predators provide a key for evaluation of the pattern and tempo of ecosystem recovery because predators are interpreted to react most sensitively to environmental turbulences. The end-Permian mass extinction was the most severe crisis experienced by life on Earth, and the common paradigm persists that the biotic recovery from the extinction event was unusually slow and occurred in a step-wise manner, lasting up to eight to nine million years well into the early Middle Triassic (Anisian) in the oceans, and even longer in the terrestrial realm. Here we survey the global distribution and size spectra of Early Triassic and Anisian marine predatory vertebrates (fishes, amphibians and reptiles) to elucidate the height of trophic pyramids in the aftermath of the end-Permian event. The survey of body size was done by compiling maximum standard lengths for the bony fishes and some cartilaginous fishes, and total size (estimates) for the tetrapods. The distribution and size spectra of the latter are difficult to assess because of preservation artifacts and are thus mostly discussed qualitatively. The data nevertheless demonstrate that no significant size increase of predators is observable from the Early Triassic to the Anisian, as would be expected from the prolonged and stepwise trophic recovery model. The data further indicate that marine ecosystems characterized by multiple trophic levels existed from the earliest Early Triassic onwards. However, a major change in the taxonomic composition of predatory guilds occurred less than two million years after the end-Permian extinction event, in which a transition from fish/amphibian to fish/reptile-dominated higher trophic levels within ecosystems became apparent.

Synchrotron-aided reconstruction of the conodont feeding apparatus and implications for the mouth of the first vertebrates.

The origin of jaws remains largely an enigma that is best addressed by studying fossil and living jawless vertebrates. Conodonts were eel-shaped jawless animals, whose vertebrate affinity is still disputed. The geometrical analysis of exceptional three-dimensionally preserved clusters of oro-pharyngeal elements of the Early Triassic Novispathodus, imaged using propagation phase-contrast X-ray synchrotron microtomography, suggests the presence of a pulley-shaped lingual cartilage similar to that of extant cyclostomes within the feeding apparatus of euconodonts ("true" conodonts). This would lend strong support to their interpretation as vertebrates and demonstrates that the presence of such cartilage is a plesiomorphic condition of crown vertebrates.

Allometries and the morphogenesis of the molluscan shell: a quantitative and theoretical model.

This article explores the close relationships between growth rate and allometries of molluscan shells. After reviewing the previous theoretical approaches devoted to the understanding of shell form and its morphogenesis, we present a free-form vector model which can simulate apertural shape changes and nonlinear allometries. Shell morphology is generated by iteratively adding a growth increment onto the last computed aperture. The first growth increment defines so-called growth vectors which are assumed to be constant in direction (relative to the last computed aperture position) during a simulation of a shell (ontogeny). These growth vectors are uniformly scaled at each time step according to various growth rate curves that are used to simulate the mantle growth over time. From the model, we derive morphometric variables that illustrate the ontogenetic trajectories in time-size-shape space. We investigate the effects of changing the growth curves types, growth rate parameters and growth vector maps on the direction, speed and patterns of ontogenetic allometries. Because this model focuses the issue on time, it highlights a plausible effect of growth rate on shell shape and illustrates some fundamental geometrical properties of the logarithmic spiral, in particular the close relationship between the size and the geometry of growth increments. This model could be used to develop a mathematically data-driven approach where experimentally obtained growth curves could be used as inputs in the model. More generally, our study recalls the role of growth rates in the generation of allometries.

Growth-dependent phenotypic variation of molluscan shells: implications for allometric data interpretation.

In recent years, developmental plasticity has received increasing attention. Specifically, some studies highlighted a possible association between shell shape and growth rates in intertidal gastropods. We use a growth vector model to study how hypothetical growth processes could underlie developmental plasticity in molluscs. It illustrates that variation in instantaneous shell growth rate can induce variability in allometric curves. Consequently, morphological variation is time-dependent. Basing our model parameters on a study documenting the results of transplants experiments of three gastropods ecomorphs, we reproduce the main aspects of the variation in size, shape, and growth rates among populations when bred in their own habitat or transplanted to another ecotype habitat. In agreement with empirical results, our simulation shows that a flatter growth profile corresponds to conditions of rapid growth. The model also allows the comparison of allometric slopes using different subdata sets that correspond to static and ontogenetic allometry. Our model highlights that depending on subdata sets, the "main effects" could be attributed to source population or environment. In addition, convergence or divergence of allometric slopes is observed depending on the subdata sets. Although there is evidence that shell shape in gastropods is to some extent growth rate dependent, gaining a general overview of the issue is challenging, in particular because of the scarcity of studies referring to allometry. We argue that the dynamics of development at the "phenotypic level" constitute a non-reducible level of investigation if one seeks to relate the observed amount of phenotypic variation to variability in the underlying factors.

Good genes and good luck: ammonoid diversity and the end-Permian mass extinction.

The end-Permian mass extinction removed more than 80% of marine genera. Ammonoid cephalopods were among the organisms most affected by this crisis. The analysis of a global diversity data set of ammonoid genera covering about 106 million years centered on the Permian-Triassic boundary (PTB) shows that Triassic ammonoids actually reached levels of diversity higher than in the Permian less than 2 million years after the PTB. The data favor a hierarchical rather than logistic model of diversification coupled with a niche incumbency hypothesis. This explosive and nondelayed diversification contrasts with the slow and delayed character of the Triassic biotic recovery as currently illustrated for other, mainly benthic groups such as bivalves and gastropods.