Cladistic analysis of the Middle Jurassic ammonite radiation

Cladistic analysis of the Middle Jurassic ammonite radiation shows how the Aalenian-age ancestral stock, known variously as the Hammatocerataceae or Hammatoceratinae, gave rise to numerous new groups in Bajocian times. Cladistic analysis shows this ancestral stock is not a single lineage but two: the ‘Erycites group’, which gave rise to the Stephanocerataceae, and the ‘Hammatoceras group’, from which all other Bajocian groups derived.

Foreword for the thematic volume of the 8ISCPP. Recent advances in present and past cephalopod studies.

3 pages, éditorial.; International audience

Le visage du monde au Sinémurien.

9 pages; National audience

The flourishing diversity of models in theoretical morphology: from current practices to future macroevolutionary and bioenvironmental challenges.

17 pages; International audience; For decades, theoretical morphological studies of different groups of organisms have been successfully pursued in biological, paleontological, and computational contexts, often with distinct modeling approaches and research questions. A regular influx of new perspectives and varied expertise has contributed to the emergence of a veritable multidisciplinary outlook for theoretical morphology. The broadening of this discipline is reflected in a substantial increase in the number of models, leading to a bewildering diversity that has yet to be scrutinized. In this work, we tackle this issue in a synthetic fashion, with a quantitativemeta-analysis that allows a…

Le débat macroévolutif : apports de la disparité morphologique

Resume La theorie macroevolutive est abordee sous des aspects historiques, puis sous un de ses aspects plus novateurs, qui aboutit a l’elaboration d’un programme de recherche initie par differentes equipes depuis une dizaine d’annees : il consiste a decoupler les composantes taxinomiques (diversite) et morphologiques (disparite) de la biodiversite, lors de l’etude de ses fluctuations. Plusieurs etudes suggerent des decouplages temporels frequents entre ces deux metriques au cours de l’histoire des clades. La confrontation des deux apparait comme une demarche feconde qui a permis d’explorer en termes de patterns , mais egalement de processus, les fluctuations de la biodiversite a large echel…

Biogeographic patterns of belemnite body size responses to episodes of environmental crisis

Body size changes have been investigated through episodes of environmental crisis among several groups of organisms but the relative contribution of within-lineage size changes, selective extinction and origination of taxa on these patterns is still being debated. Rapid warming, anoxia, and perturbations of the carbon cycle linked with volcanic activity, as well as their impact on marine diversity are well documented for the Pliensbachian-Toarcian (Pli-Toa) boundary and for the Toarcian Oceanic Anoxic Event (T-OAE). Belemnites were a very abundant and successful cephalopod group in the Mesozoic oceans playing a paramount role in the oceanic trophic webs. Belemnites have mainly been studied …

Geometric Measurement Analysis Versus Fourier Series Analysis for Shape Characterization Using the Gastropod Shell (Trivia) as an Example

Varied and efficient methods have been developed to describe and quantify natural objects. The most common ones use superimposition techniques (e.g. Procrustes methods; Bookstein, 1991), decomposition into harmonics (Fourier series and functions, wavelets; Anstey and Delmet, 1973; Christopher and Waters, 1974; Gevirtz, 1976; Lestrel, 1997; Toubin and others, 1999; Verrecchia, Van Grootel, and Guillemet, 1996; Younger and Ehrlich, 1977), analysis of spiral functions (e.g. Raup parameters; Raup, 1961, 1966; Tursch, 1998), and combinations of parameters from elementary geometry (e.g. circularity index, lengthening; Coster and Chermant, 1989; Schmidt-Kittler, 1986; Viriot, Chaline, and Schaaf, …

Early–middle Jurassic lytoceratid ammonites with constrictions from Morocco: palaeobiogeographical and evolutionary implications.

13 pages; International audience; The ammonite genus Alocolytoceras Hyatt, 1900 is an uncommon lytoceratid with distinctive shell ornament. A set of 58 specimens, recently collected at Amellago in the central High Atlas (Morocco), has enabled us to trace a succession of three species over eight biozones from the Toarcian to the Aalenian. Two specimens from the Lusitanian Basin are added for comparison. Following a review of the genus, based on original specimens and data from the literature, seven species are considered valid. A palaeobiogeographical synthesis of 13 regions demonstrates irregular distribution patterns over time, with a constant presence in the south-west Tethys and an insta…

Morphometrics of hard structures in cuttlefish

7 pages; International audience; Cuttlefishes exhibit several hard structures that have been characterised using morphometric analysis. Most of these data come from cuttlebones, although statoliths and beaks are also used. It appears that morphometric techniques are mainly used for taxonomic purposes. However, some analyses have emphasised functional morphology and macroevolution. Morphological features (including the inner shell) of cuttlefishes and their availability for study (numerous specimens, culture in the laboratory) make them a first rank model for numerous studies in life sciences that require morphometrics, and especially in the field of evolutionary biology.

Testing a portable laser-induced breakdown spectroscopy system on geological samples.

9 pages; International audience; This paper illustrates the potentialities of a home-made portable LIBS (laser-induced breakdown spectroscopy) instrument in Earth sciences, more particularly in geochemically recognizing (i) tephra layers in lacustrine sediments and (ii) fossilization processes in ammonites. Abundances for selected lines of Al, Ca, Fe, Ti, Ba and Na were determined in lacustrine chalk sediments of the Jura, where the Laacher See Tephra (LST) layer is recorded. A statistical treatment of elemental maps produced from the section of a sedimentary column containing the LST event allows instrumental conditions to be optimized. Accumulating spectra from close shot positions gives …

Embryonic Ammonoid Shell Features: Intraspecific Variation Revisited

Two samples of ammonoids belonging to the Oppeliidae, Sublunuloceras virguloidesHecticoceras (Brightii) canaliculatum, are analyzed to estimate the intraspecific variability of embryonic shell features. The study of embryonic shell characters reveals two main shapes of protoconch, flattened and round. Prosiphons may be straight or slightly curved. New parameters for area are added to the linear parameters commonly found in the literature. Prosiphon length and caecum area vary greatly whereas protoconch and ammonitella diameter vary only slightly, and the ammonitella angle is almost constant. The protoconch-to-ammonitella size ratio behaves differently in each species, suggesting different p…

Le cadre géographique et géologique.

2 pages; National audience

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, …

Size patterns through time: the case of the Early Jurassic ammonite radiation

The shell size of 1236 ammonite species representing all known Early Jurassic faunas is analyzed. Size patterns are studied for the entire period and then at the biozone scale for the first four stages of the Jurassic (28 Myr), during which ammonites recovered from the crisis at the Triassic/Jurassic (T/J) boundary. Our analysis reveals that (1) a size continuum (normal distribution from “dwarfs” to “giants”) exists for all Early Jurassic ammonites; (2) although there are no sustained trends (e.g., no Cope's rule), the succession is not monotonous and patterns may differ conspicuously from one biozone to the next; and (3) increases and decreases in size range are the most frequent evolution…

Stratotype Sinémurien.

320 pages; National audience

Deep-Time Phylogenetic Clustering of Extinctions in an Evolutionarily Dynamic Clade (Early Jurassic Ammonites)

7 pages; International audience; Conservation biologists and palaeontologists are increasingly investigating the phylogenetic distribution of extinctions and its evolutionary consequences. However, the dearth of palaeontological studies on that subject and the lack of methodological consensus hamper our understanding of that major evolutionary phenomenon. Here we address this issue by (i) reviewing the approaches used to quantify the phylogenetic selectivity of extinctions and extinction risks; (ii) investigating with a high-resolution dataset whether extinctions and survivals were phylogenetically clustered among early Pliensbachian (Early Jurassic) ammonites; (iii) exploring the phylogene…

Mollusca.

Heterochronic differentiation of sexual dimorphs among Jurassic ammonite species

Ontogenetic and then heterochronic approaches are used here to analyze sexual differentiation within two well-known Jurassic dimorphic species. This analysis compares two ways of determining the relative biological age of ammonites, one using size (diameter) and the other the number of septa as a proxy of age. The shape standard is established from factor analysis of morphological and growth parameters. Size-age-shape relationships are analyzed on the basis of a new heterochronic representation. When diameter is used as a proxy of age, microconch morphs are globally considered to be progenetic compared with macroconch morphs. When size and age are determined separately and shape is included…

La notion d'espace morphologique,outil d'analyse de la morphodiversité des organismes

Resume L'analyse de la morphologie des organismes est une base essentielle dans la mise en evidence des changementsevolutifs. La morphologie est maintenant frequemment abordee par des procedures quantitatives, dont certaines permettent de construire de veritables cartes morphologiques ou les dissemblances et ressemblances de formes entre les organismes sont directement lisibles et quantifiees. La notion d'espace morphologique, illustree ici par differents exemples, se revele particulierement feconde pour comprendre comment la diversite morphologique d'un ensemble d'organismes (stades ontogenetiques, populations, clades) se structure, et par consequent pour interpreter en termes evolutifs ou…

Les Ammonoïdes (Mollusca, Cephalopoda) : avancées et contributions récentes à la paléobiologie évolutive.

12 pages; Les ammonoïdes qui perdurent 335 Ma constituent un modèle de choix dans l'analyse du fait évolutif. Cet article de synthèse se propose d'aborder (1) : leur place phylogénétique au sein des céphalopodes et le choix d'un modèle actuel de référence ; (2) : la construction d'espaces phénotypiques qui offrent un angle d'étude pertinent de l'évolution biologique ; (3) : les concordances et discordances entre reconstruction phylogénétique et registre fossile et (4) : les reconquêtes postcrises, modèles d'étude de l'évolution à grande échelle. Il ressort que les ammonoïdes peuvent servir d'appui à de nombreuses thématiques paléontologiques (dynamique de la biodiversité, analyse phylogénét…

Extraire et paraître : le Sinémurien à Saint-Romain.

9 pages; National audience

Les ammonites traditionnellement regroupées dans la sous-famille des Bouleiceratinae (Hildoceratidae, Toarcien) : aspects phylogénétiques, biogéographiques et systématiques

Resume Les genres d’ammonites, traditionnellement regroupes dans la sous-famille des Bouleiceratinae, appartiennent en fait a des lignees differentes de la famille des Hildoceratidae. Une approche utilisant des arguments paleogeographiques, stratigraphiques et morphologiques d’une part et une approche cladistique d’autre part ont permis de distinguer 3 groupes se succedant au cours du temps et a repartitions paleogeographiques differentes : • les Bouleiceratinae s.s. (genres Bouleiceras, Nejdia et Kohaticeras) du Toarcien inferieur ; • les Leukadiellinae (genres Leukadiella et Renziceras) du Toarcien moyen ; • et les Paroniceratinae (genres Frechiella, Paroniceras et Oxyparoniceras) du Toar…

The Use of Landmarks to Describe Ammonite Shap

The use of landmarks and geometrical morphometric techniques is illustrated with examples from the Harpoceratinae (Ammonitina). This approach is useful in analysing morphological variations of characters such as rib pattern or ventral shape on the scale of the genus or species. It can be used to map forms so that similarities and dissimilarities between organisms can be read and quantified directly.

Allometric space and allometric disparity: a developmental perspective in the macroevolutionary analysis of morphological disparity.

8 pages; International audience; Here, we advance novel uses of allometric spaces--multidimensional spaces specifically defined by allometric coefficients--with the goal of investigating the focal role of development in shaping the evolution of morphological disparity. From their examination, operational measures of allometric disparity can be derived, complementing standard signals of morphological disparity through an intuitive and process-oriented refinement of established analytical protocols used in disparity studies. Allometric spaces thereby become a promising context to reveal different patterns of evolutionary developmental changes and to assess their relative prevalence and import…

Ammonite paleobiogeography during the Pliensbachian-Toarcian crisis (Early Jurassic) reflecting paleoclimate, eustasy, and extinctions.

14 pages; International audience; The Pliensbachian-Toarcian crisis (Early Jurassic) is one of the major Mesozoic paleoecological disturbances when ca. 20% of marine and continental families went extinct. Contemporaneously, profound paleobiogeographical changes occurred in most oceanic domains including a disruption of ammonite provincialism during the Early Toarcian. Here, we quantitatively reappraise the structure and evolution of paleobiogeographical patterns displayed by ammonite faunas before, during, and after the biological crisis, over a time-interval including 13 biochronozones. The high-resolution study presented here involves the use of hierarchical Cluster Analyses, non-metric M…

Existe-t-il une crise au passage Lias-Dogger chez les ammonites? Approche morphometrique par quantification de la disparite morphologique

Abstract Morphological changes observed in ammonites at the Lias-Dogger (Toarcian-Aalenian) boundary are analyzed here. The study covers all known ammonites worldwide over some 2 m.y. (the last two biozones of the Toarcian: Pseudoradiosa and Aalensis and the first biozone of the Aalenian: Opalinum). Ammonite taxonomy at this boundary is still open to question, the main disagreements being over groupings at family or super-family level. The evolutionary schema developed by Tintant and Mouterde [1981] shows two main sets of Ammonitina for the time: (1) the dominant Hammatocerataceae (Graphoceratidae+Hammatoceratidae) and (2) the final representatives of the Hildocerataceae. Two conservative a…

Les contraintes.

Les ammonites du Sinémuriensupérieur du Jebel-Bou-Hamid (Haut-Atlas central, Rich, Maroc). Approches paléontologique et biostratigraphique

Resume L'etude des faunes d'ammonites - plus de 25 taxons - et de leur succession biostratigraphique dans unecoupe exceptionnellement complete et fossilifere du Haut-Atlas central de la region de Rich (Maroc) a permis la mise en evidence de 10 unites biochronologiques elementaires (=assemblages) pour une periode debutant au sein de la partie terminale du Sinemurien inferieur et s'achevant avec le Sinemurien superieur. Pour les zones a Oxynotum et Raricostatum, il s'agit des premieres donnees biostratigraphiques collectees avec une telle precision dans le Nord de l'Afrique. Pour le Sinemurien inferieur et la zone a Obtusum, les nombreuses donnees nouvellement acquises, completent, precisent …

Spatial patterns of disparity and diversity of the Recent cuttlefishes (Cephalopoda) across the Old World

Aim Diversity and disparity metrics of all Recent cuttlefishes are studied at the macroevolutionary scale (1) to establish the geographical biodiversity patterns of these cephalopods at the species level and (2) to explore the relationships between these two metrics. Location Sampling uses what is known about these tropical, subtropical and warm temperate cephalopods of the Old World based on a literature review and on measurements of museum specimens. Some 111 species spread across seventeen biogeographical areas serve as basic units for exploring diversity and disparity metrics in space. Methods Landmarks describe the shape of the cuttlebone (the inner shell of the sepiids) and difference…

La Dalle aux ammonites de Digne‑les‑Bains : un patrimoine d’âge Sinémurien de réputation mondiale.

6 pages; National audience

Jacques Thierry (1941-2014).

10 pages; International audience

The shell matrix and microstructure of the Ram's Horn squid: Molecular and structural characterization.

20 pages; International audience; Molluscs are one of the most diversified phyla among metazoans. Most of them produce an external calcified shell, resulting from the secretory activity of a specialized epithelium of the calcifying mantle. This biomineralization process is controlled by a set of extracellular macromolecules, the organic matrix. In spite of several studies, these components are mainly known for bivalves and gastropods. In the present study, we investigated the physical and biochemical properties of the internal planispiral shell of the Ram's Horn squid Spirula spirula. Scanning Electron Microscope investigations of the shell reveal a complex microstructural organization. The…

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 …

Les ammonites.

Jean-Jacques Collenot (1814-1892).

3 pages; National audience

Disparity as a complement to taxonomy and phylogeny in biogeographic studies: present and past examples from the cephalopods.

33 pages; International audience

The space-time relationship of taxonomic diversity and morphological disparity in the Middle Jurassic ammonite radiation.

14 pages; International audience; The Middle Jurassic ammonite radiation (from the late Aalenian to the end of the mid-Bathonian) is traced using combined analyses of morphological disparity and taxonomic diversity. The global signals of disparity and diversity are compared. These signals are then broken down by paleogeographical provinces to detect any heterogeneity in the radiation. An examination of the global signals reveals three biodiversity crises (discordances between signals) where morphological disparity grows while taxonomic diversity declines. The subdivision of the signals indicates the radiation was heterogeneous between provinces: the global signal is an aggregate of signals …

World distribution of middle Jurassic ammonites (Upper Aalenian to Middle Bathonian): relationships between biodiversity and palaeogeography

Abstract The break up of the Pangea takes place in the Jurassic; the palaeoceanographic consequences are the opening of seaways, particularly at the place of the future Atlantic and Indian oceanic areas. During the Toarcian, and from the late Aalenian to the middle Bathonian, the so-called “Hispanic corridor” (or “Atlantic seaway”) exists between the “western Tethys” and the “American Pacific border”, through the “Caribbean Tethys”. Two additional seaways which play as by-passes of the Pangea are proposed, one along the northern border of Laurasia (Boreal sea), and a second along the southern border of Gondwana (South Pacific Sea); however, if these two last could be effectively used for fa…

Les grands auteurs du stratotype du Sinémurien.

1 page; National audience

Ammonoid morphological signal versus sea-level changes

The morphological diversity, considered as a biological signal, of a series of four Upper Callovian (Middle Jurassic) ammonite populations of Côte-d’Or (France) is quantified. These populations fall within two third-order sequences of a second-order transgression. A sampling method that adheres as closely as possible to the morphological characteristics of the populations is established. It is valuable in that it eliminates the subjectivity related to taxonomy to allow more objective comparisons between the biological signal (shell morphology) and the physical signal (eustasy).

Gastropods from upper Pliensbachian–Toarcian (Lower Jurassic) sediments of Causses Basin, southern France and their recovery after the early Toarcian anoxic event

AbstractA gastropod fauna has been studied from upper Pliensbachian – upper Toarcian deposits of two sections of the Causses Basin (southern France) in order to investigate the mode of recovery after the early Toarcian anoxic event. The fauna consists of 15 species, one of which is new (Bathrotomaria kronzwilmesorumsp. nov.). Their stratigraphical distribution shows two peaks of diversity – in the Bifrons Zone (Bifrons Subzone) and in the Aalensis Zone (Mactra Subzone) – which reflect brief times during which the oxygen content and bottom consistency favoured the settlement of a relatively diversified fauna. In the Variabilis–Pseudoradiosa zones, gastropods are only represented by two speci…

Faunal invasions as a source of morphological constraints and innovations? The diversification of the early Cardioceratidae (Ammonoidea; Middle Jurassic)

Abstract Multivariate analysis of shell characters and quantification of morphological diversity (morphospace occupation and disparity) are used here to investigate the modes of morphological diversification of ammonites. We define five events in early cardioceratid history that connect geographical changes causing emigration or immigration phases with biodiversity dynamics: (1) the initial colonization of the Arctic Basin by the Cardioceratidae at the end of the Bajocian, Middle Jurassic; (2) the first appearance of the Kosmoceratidae clade in the Boreal Realm during the Bathonian; (3) the ensuing expansion phase of this clade in the Boreal Realm; (4) the first phase of migration of the Ca…

Phylogenetic practices among scholars of fossil cephalopods, with special reference to cladistics.

12 pages; International audience

Les sites complémentaires.

6 pages; National audience

La gryphée du Sinémurien : un fossile emblématique.

6 pages; National audience

Combining ontogenetic and evolutionary scales of morphological disparity: a study of early Jurassic ammonites

SUMMARY Two major research themes in Evolutionary Developmental Biology and in Paleobiology, respectively, have each become central for the analysis and interpretation of morphological changes in evolution: the study of ontogeny/ phylogeny connections, mainly within the widespread and controversial framework of heterochrony; and the study of morphological disparity, the morphological signal of biodiversity, describing secular changes in morphospace occupation during the history of any given clade. Although enriching in their respective fields, these two themes have remained rather isolated to date, despite the potential value of integrating them as some recent studies begin to suggest. Here…

Le site stratotypique.

6 pages; National audience

Les contraintes.

Water mass exchange and variations in seawater temperature in the NW Tethys during the Early Jurassic: evidence from neodymium and oxygen isotopes of fish teeth and belemnites.

10 pages; International audience; Oxygen and neodymium isotope analyses performed on biostratigraphically well-dated fish remains recovered from the Hettangian to Toarcian of the Paris Basin were used to reconstruct variations of Early Jurassic seawater temperature and to track oceanographic changes in the NW Tethys. Our results indicate a strong correlation between δ18O trends recorded by fish remains and belemnites, confirming the paleoenvironmental origin of oxygen isotope variations. Interestingly, temperatures recorded by pelagic fishes and nektobenthic belemnites and bottom dwelling fishes are comparable during the Late Pliensbachian sea-level lowstand but gradually differ during the …

Le calcaire à gryphées arquées dans la construction : de la Bourgogne au Beaujolais.

4 pages; National audience

Evolutionary trends within Jurassic ammonoids.

An evolutionary trend is a pattern of traits shifting in a given direction over a sufficiently long period of time to be detected. In this study, we examine trends among Jurassic ammonoids. First, a compilation of selected literature on evolutionary patterns and trends leads to the conclusion that such trends have rarely been studied among Jurassic ammonoids and even more rarely are they formally studied using statistical tests. Second, a preliminary survey of trends for the entire Jurassic at the family level (based on four published articles) suggests two sustained trends occurred each lasting nearly 10 myr: the Early Jurassic and Middle Jurassic radiations, which were related to the firs…

Developmental aspects of morphological disparity dynamics: a simple analytical exploration

15 pages; International audience; We devised a simple model for assessing the role of development in shaping the evolution of morphological disparity. Disparity of a clade at any given time is expressed in terms of the developmental dynamics that lead to the variety of adult morphotypes observed. We use assumed phenotypic manifestations of developmental processes, as they could be detected from allometric characterizations, to distinguish a few, nonexclusive types of evolutionary changes in ontogeny. On the basis of this formalization, we describe the diversification of hypothetical clades, using the standard curve of adult morphological disparity, the curve of juvenile disparity, and the c…

La biodiversité au Sinémurien.

3 pages; National audience

Le Sinémurien de la région de Semur-en-Auxois (Jurassique inférieur, Sud-Est du bassin de Paris, France), localité type historique de l’étage Sinémurien de d’Orbigny.

Introduction.

L’état actuel du patrimoine géologique du Sinémurien.

1 page; National audience

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 …

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

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…