Emmanuel Fara

High-resolution dynamics of Early Jurassic marine extinctions: the case of Pliensbachian-Toarcian ammonites (Cephalopoda)

Abstract: The Pliensbachian–Toarcian interval was marked by major environmental disturbances and by a second-order mass extinction. Here, we reappraise the taxonomic, spatiotemporal and selective dynamics of extinctions over the whole interval, by analysing a high-resolution dataset of 772 ammonite species from NW Tethyan and Arctic domains. On average, 40–65% of ammonite species disappeared during each subchronozone, but higher extinction pulses (reaching 70–90%) prevailed from the Margaritatus to the Dispansum Chronozone. The main extinctions, corresponding to the Gibbosus, Pliensbachian–Toarcian boundary, Semicelatum, Bifrons–Variabilis, and Dispansum events, differed in their dynamics, suggesting episodes of ecological stress related to climate change, regression, disturbance in the carbon cycle or anoxia. The multi-pulsed volcanic activity in the Karoo–Ferrar province could well have triggered these ecological changes. In addition, ammonites experienced a morphological bottleneck during the Gibbosus event, 1 Ma before the Early Toarcian diversity collapse. Typically, drops in richness were related both to high extinctions and to declines in origination rates. This feature could result from strengthened ecological stresses related to the temporal overlap of environmental disturbances. After the Early Toarcian crisis, the recovery of ammonites was rapid (2 Ma) and probably influenced by a coeval marine transgression. Supplementary material: Figures showing a comparison of extinction and origination patterns based on different datasets, and variations of the morphospace occupation are available at http://www.geolsoc.org.uk/SUP18381.

The Fossil Record of Cretaceous Tetrapods

Abstract The fossil record of the Cretaceous is critical for understanding the evolution of modern tetrapods. Using a measure of relative completeness of the fossil record—the Simple Completeness Metric (SCM)—quality of the fossil record and diversity during the Cretaceous appear to be closely related, suggesting an artifactual component. The SCM calculations also show that knowledge of the fossil record has improved in the last ten years. Recent proposals that modern orders of birds and mammals originated early in the Cretaceous are rendered unlikely by four arguments: (1) the SCM calculations indicate that the fossil record of Cretaceous birds and mammals is relatively good; (2) it is unlikely that all modern orders, independently, would have remained cryptic throughout the Cretaceous; (3) control samples of exquisitely preserved tiny Cretaceous tetrapods lack any specimens of modern groups of birds and mammals; and (4) the suggestion that the undiscovered ancestors of modern groups are to be found in unsampled parts of the Earth is not supported by cladistic evidence.

Smithian ammonoid faunas from Utah: implications for Early Triassic biostratigraphy, correlation and basinal paleogeography

Intensive sampling of the lower portion of the Thaynes and Moenkopi Groups (Lower Triassic) at separate localities within the Confusion Range, Pahvant Range, Mineral Mountains, Star Range, Kanarraville, Cedar City, Torrey and San Rafael Swell areas (mainly central and southern Utah, USA) leads to the recognition of a new key regional Smithian ammonoid succession. The new biostratigraphical sequence, which is more precise than the long-recognized Meekocerasgracilitatis and Anasibiriteskingianus Zones, comprises twelve subdivisions, thus resulting in a sequence with much higher resolution that can be correlated not only with other western USA sites, but also with major worldwide localities as well. Middle and late Smithian faunas contain many taxa with wide geographic distribution, thus enabling long-distance correlation with faunal successions from other regions (e.g., British Columbia, Canadian Arctic, South China, Spiti and Oman). New assemblages from the lowermost beds are the least diversified and poorest preserved; they represent the earliest early/middle Smithian ammonoid faunas reported from the western North American basin. They highlight (a) the sudden Smithian advancement of the marine transgression within this epicontinental sea, (b) that this event is diachronous, and (c) that the paleotopography of the basin most likely was highly irregular. The newly obtained ammonoid succession also allows us to date and follow the transgression from the northern and central part of the basin to the southwesternmost and southeasternmost parts, which were reached during the late Smithian (Anasibiriteskingianus beds). In addition, we briefly discuss the now-limited previous regional biozonation in the light of these new results. One new genus (Minersvillites) and nine new species (Kashmirites utahensis, Kashmirites confusionensis, Kashmirites stepheni, ?Xiaoqiaoceras americanum, Minersvillites farai, Inyoites beaverensis, Meekoceras olivieri, Meekoceras millardense, Vercherites undulatus) are also described.

Sea-level variations and the quality of the continental fossil record

Theoretical and empirical studies suggest that sea-level change was a fundamental parameter determining the distribution of fossil marine organisms. Although a similar relationship is generally assumed for the continental fossil record, this intuition remains speculative. Quantitative evidence reveals no correlation between sea-level variations and the distribution of continental tetrapods in the Late Jurassic–Eocene interval. The quality of the continental fossil record depends on many more factors than just eustatic changes.

Unexpected Early Triassic marine ecosystem and the rise of the Modern evolutionary fauna.

A new marine fossil assemblage from the Early Triassic shows unexpected phyletic diversity and functional complexity. In the wake of the end-Permian mass extinction, the Early Triassic (~251.9 to 247 million years ago) is portrayed as an environmentally unstable interval characterized by several biotic crises and heavily depauperate marine benthic ecosystems. We describe a new fossil assemblage—the Paris Biota—from the earliest Spathian (middle Olenekian, ~250.6 million years ago) of the Bear Lake area, southeastern Idaho, USA. This highly diversified assemblage documents a remarkably complex marine ecosystem including at least seven phyla and 20 distinct metazoan orders, along with algae. Most unexpectedly, it combines early Paleozoic and middle Mesozoic taxa previously unknown from the Triassic strata, among which are primitive Cambrian-Ordovician leptomitid sponges (a 200–million year Lazarus taxon) and gladius-bearing coleoid cephalopods, a poorly documented group before the Jurassic (~50 million years after the Early Triassic). Additionally, the crinoid and ophiuroid specimens show derived anatomical characters that were thought to have evolved much later. Unlike previous works that suggested a sluggish postcrisis recovery and a low diversity for the Early Triassic benthic organisms, the unexpected composition of this exceptional assemblage points toward an early and rapid post-Permian diversification for these clades. Overall, it illustrates a phylogenetically diverse, functionally complex, and trophically multileveled marine ecosystem, from primary producers up to top predators and potential scavengers. Hence, the Paris Biota highlights the key evolutionary position of Early Triassic fossil ecosystems in the transition from the Paleozoic to the Modern marine evolutionary fauna at the dawn of the Mesozoic era.

Diversity of Callovian-Ypresian(Middle Jurassic-Eocene) tetrapod families and selectivity of extinctions at the K/T boundary

Abstract The diversity of tetrapod families increased through the Cretaceous, punctuated by three majorextinction events at the Jurassic/Cretaceous, Cenomanian/Turonian, and Cretaceous/Tertiary boundaries. Extinction and survivorship at these times are analysed with regard to taxonomy and ecological categories of diet, habitat and size. Whereas no constant selectivity pattern is found for diet and size through the period, freshwater tetrapods seem to have been less extinction-prone than terrestrial and marine ones. The Cretaceous/Tertiary extinction had less effect on small, omnivorous, and freshwater families than other ecological categories. This strong selectivity signal is consistent with other work made at higher taxonomic and stratigraphic levels of resolution, thus legitimising the use of fossil families in global studies of diversity and palaeoecology.

Smithian shoreline migrations and depositional settings in Timpoweap Canyon (Early Triassic, Utah, USA)

In Timpoweap Canyon near Hurricane (Utah, USA), spectacular outcrop conditions of Early Triassic rocks document the geometric relationships between a massive Smithian fenestral-microbial unit and underlying, lateral and overlying sedimentary units. This allows us to reconstruct the evolution of depositional environments and high-frequency relative sea-level fluctuations in the studied area. Depositional environments evolved from a coastal plain with continental deposits to peritidal settings with fenestral-microbial limestones, which are overlain by intertidal to shallow subtidal marine bioclastic limestones. This transgressive trend of a large-scale depositional sequencemarks a long-term sea-level rise that is identified worldwide after the Permian-Triassic boundary. The fenestral-microbial sediments were deposited at the transition between continental settings (with terrigenous deposits) and shallow subtidal marine environments (with bioturbated and bioclastic limestones). Such a lateral zonation questions the interpretation of microbial deposits as anachronistic and disaster facies in the western USA basin. The depositional setting may have triggered the distribution of microbial deposits and contemporaneous marine biota. The fenestral-microbial unit is truncated by an erosional surface reflecting a drop in relative sea level at the scale of a medium depositional sequence. The local inherited topography allowed the recording of small-scale sequences characterized by clinoforms and short-distance lateral facies changes. Stratal stacking pattern and surface geometries allow the reconstruction of relative sea-level fluctuations and tracking of shoreline migrations. The stacking pattern of these small-scale sequences and the amplitude of corresponding high-frequency sea-level fluctuations are consistent with climatic control. Large-andmedium-scale sequences suggest a regional tectonic control.

Ammonoid recovery after the Permian–Triassic mass extinction: a re-exploration of morphological and phylogenetic diversity patterns

The explosive ammonoid rediversification after the Permian–Triassic mass extinction is now well understood in terms of taxonomic richness and biogeography. Using an updated dataset of Early Triassic ammonoids, we compare morphological disparity and taxonomic richness patterns at the regional and global scales. Disparity evolved similarly at both scales, suggesting a global influence of abiotic factors. Morphological diversification occurred early in the Smithian and a marked contraction of the morphospace took place during the end-Smithian extinction. We confirm that trends in disparity and richness were decoupled during the Griesbachian and Dienerian. Three macroevolutionary processes may be involved: (1) a nonselective extinction at the Permian–Triassic boundary; (2) a Dienerian constrained radiation with several homeomorphic genera; (3) a potential deterministic extinction during the end-Smithian crisis. We also demonstrate a superfamily imprint upon disparity for the Spathian when most superfamilies occupied a restricted part of the morphospace. Sphaerocones were the most affected by the Dienerian and end-Smithian extinction, but explanations remain elusive. On the one hand, this may be linked to widespread harsh conditions at those times. On the other hand, as the sphaerocones occurred episodically during the Early Triassic, this might be explained by a relaxing of ecological constraints or simply by convergent evolution. Supplementary materials: The database, including measurements of specimens illustrated in previously published plates and of unpublished specimens from Utah, South China and Spiti, as well as the number of genera present in each studied substage of the Early Triassic, are available at www.geolsoc.org.uk/SUP18571.

Spine anatomy reveals the diversity of catfish through time: a case study of Synodontis (Siluriformes)

Synodontis (Mochokidae, Siluriformes) is a freshwater catfish endemic to Africa. The 118 extant species are present in almost all hydrographic basins. Some species are restricted to a single stream, whereas others have a vast distribution. Synodontis is known in the fossil record since the Miocene, and its history depends on the connections among African basins through time. The identification of species in the fossil record is essential to reconstruct this historical pattern. Catfish pectoral and dorsal spines are robust, they preserve well and they form most of the fossil remains for the genus Synodontis. Unfortunately, the criteria for the identification of extant Synodontis species are not applicable to fossil specimens. Here, we define 11 original morphological characters that permit to discriminate four extant species from the Chad-Chari hydrographic system. Six of these characters are defined on pectoral spines and five on dorsal spines. We then show that these characters can be used successfully for identifying fossil specimens. In particular, we present a case study in which we identify Synodontis cf. schall and Brachysynodontis cf. batensoda in the hominid-bearing sector Toros-Menalla (Late Miocene, northern Chad). We show that spine anatomy can be a powerful tool to recognise catfish species through time and thus to identify historical diversity pattern.

Paleorhinus magnoculus from the Upper Triassic of Morocco: a juvenile primitive phytosaur (Archosauria)

The holotype and only specimen of Paleorhinus (= Arganarhinus) magnoculus is most likely a juvenile, as demonstrated by absolute size and skull proportions such as large orbits, a short snout and a narrow rostrum in comparison with other primitive (non-phytosaurid) phytosaurs. The specimen is the most complete juvenile phytosaur known, and may shed light on the poorly known ontogenetic development of phytosaurs. On the basis of the data available at present, P. magnoculus, and therefore Arganarhinus, cannot be regarded as valid taxa.