Ecological signature of the end-Triassic biotic crisis: what do bivalves have to say?

In order to understand the causes underlying the Triassic-Jurassic (T/J) mass extinction, we tested different bivalve features for extinction selectivity, i.e. shell mineralogy, age at the Rhaetian and three main autoecologic traits (feeding mechanism, tiering and motility/attachment). Also, diversity and turnover rates throughout the Triassic and the Early Jurassic were analysed in detail. The dataset employed for this analysis was a precise database at genus level including data from Induan to Sinemurian times. Results point to a true mass extinction for bivalves around the T/J boundary. This extinction was not ageselective at the boundary. Certain analyses suggested that shell mineralogy was a character significantly increasing survival odds, but this relationship seems to reflect selectivity on autoecologic traits. There was no difference in extinction proportions between both feeding types (i.e. deposit feeders and filter feeders); among the other traits, deep burrowers, epifaunal-motile and endobyssate forms seem to have been favoured, while shallow burrowers (and probably reclined forms) were more heavily affected. This pattern suggests an environmental stress at the boundary with some particular issues affecting the different life modes. Models linking magmatism in the Central Atlantic Magmatic Province with the end- Triassic mass extinction are a plausible scenario for this kind of perturbation

Dinámica de la paleodiversidad de los Bivalvos del Triásico y Jurásico Inferior.

RESUMEN Los bivalvos son un grupo de invertebrados muy utilizado para el estudio de dinámicas evolutivas, dado que su concha los hace buenos candidatos para la fosilización y son abundantes en el registro fósil. El objetivo de esta tesis es el estudio de la paleodiversidad genérica de los bivalvos marinos en el intervalo Induense (Triásico Inferior) Sinemuriense (Jurásico Inferior) [In-Sin], que comienza con la recuperación de la extinción del límite Pérmico Triásico (P/T) y termina con la recuperación de la extinción del Triásico Jurásico (T/J). En primer lugar, se han eliminado, en la medida de lo posible, los errores de la práctica taxonómica mediante la elaboración crítica de una base de datos, para lo cual se han revisado todos los géneros citados en el intervalo de estudio. Todos los datos han sido tomados de trabajos que contuvieran imágenes y/o descripciones de los taxones. Se justifica la exclusión de los taxones no considerados presentes o válidos en el intervalo In-Sin. Para cada uno de los géneros se ha revisado su rango estratigráfico, su distribución paleobiogeográfica (representada en mapas paleogeográficos), su modo de vida y la mineralogía de su concha. En cuanto al rango estratigráfico, se ha elaborado una comparación con el Treatise on Invertebrate Palaeontology y el Compendium of Fossil Marine Animal Genera. En cuanto al modo de vida, se han tenido en cuenta la posición de vida con respecto a la columna de agua y al sustrato, la estrategia alimentaria, el grado de movilidad y/o de fijación. En segundo lugar, se han evaluado los sesgos tafonómicos que pueden afectar al grupo de los bivalvos usando tablas de contingencia complementadas con residuales ajustados para diferentes bases de datos. Hemos considerado el efecto de la rarefacción del registro con la edad geológica, la mineralogía de la concha, el carácter del mar (aragonítico o calcítico), el modo de vida y la amplitud de la distribución paleobiogeográfica aproximada por la longevidad de los géneros. Se concluye que tanto la mineralogía de la concha como la rarefacción afectan al registro de los bivalvos tanto durante el Triásico como para el total del registro fanerozoico. Con los datos resultantes se procedió al análisis de la paleodiversidad desde los puntos de vista taxonómico y paleoecológico. En el primer caso, se analiza la diversidad de los géneros surgidos durante el intervalo In-Sin y su posterior destino hasta la actualidad; se ha realizado un estudio detallado de las tasas de originación, extinción y crecimiento; y por último, se ha hecho uso del análisis de cohortes para el estudio de los patrones de extinción. Todo ello se ha realizado para el total de los géneros y para su desglose según modo de vida (epifaunal e infaunal). Para esto último, se han estudiado los cambios en el ecoespacio de los bivalvos a lo largo del intervalo In-Sin, poniendo especial énfasis en el impacto de la extinción T/J. Del análisis de los datos se concluye que: 1) a consecuencia de la extinción P/T, el escalonamiento ecológico se encuentra muy poco desarrollado durante el Triásico Inferior, pero se restablece durante el Triásico Medio y apenas se ve afectado durante la extinción T/J; 2) se observa un proceso de infaunalización, sobre todo a partir del Triásico Superior, que continúa durante el Jurásico Inferior, y 3) aunque la extinción del T/J tiene un impacto taxonómico importante (42%), debido a las tasas de extinción altas y de originación bajas, a nivel paleoecológico no se modifica el número de estrategias utilizadas, lo cual es interpretado como un desacoplamiento entre las intensidades de extinción taxonómica y ecológica que sufriría este grupo en dicho momento del tiempo. __________________________________________________________________________________________________The main goal of this PhD dissertation is the study of the marine bivalve generic palaeodiversity throughout the Induan (Lower Triassic) Sinemurian (Lower Jurassic) [In Sin] interval, which includes two recovery processes, namely those of the Permo-Triassic (P/T) and Triassic Jurassic (T/J) extinctions. A database was prepared from papers with graphic and/or descriptive information of taxa, excluding genera which were neither present nor valid throughout the interval. In addition, stratigraphic ranges (with comparison Treatise on Invertebrate Palaeontology and Compendium of Fossil Marine Animal Genera) and paleobiogeographic distributions as well as mode of life (guilds, mobile or attached, and position referred to the water column and substrate), and shell mineralogy have been re-examined. Taphonomic biases have been assessed by means of an analysis of contingency tables with adjusted residuals for different databases. In spite of having considered several biasing factors (mode of life, eurytopy or stenotopy, and so on) influencing preservation of bivalves throughout the [In Sin], we have only recognised the strong influence of mineralogy associated to rarefaction, as it occurs during the Fanerozoic. Through the filtered data, estimates of origination, extinction and growth rates, and perform cohort analysis have been obtained. Differences in rates produce fluctuations of taxonomic diversity throughout [In Sin]. Cohort analysis evidences detailed extinction patterns of genera arisen in each stage or the epifaunal in contrast with the infaunal genera. Finally, changes in the bivalve ecospace are studied during the [In Sin] interval, with special emphasis on the impact of the T/J extinction. The main conclusions are 1) ecological tiering shows a very low development during the Lower Triassic as a consequence of the P/T extinction, but it is recovered during the Middle Triassic; after the T/J extinction, bivalve tiering has been scarcely affected. 2) infaunal genera became more and more abundant; this trend began since the Upper Triassic and went on the Lower Jurassic. 3) 42% of genera become extinct during the T/J extinction; impoverishment was due to a high extinction rate jointly to low origination rate; however, the diversity of guilds was not altered; this suggests that taxonomic extinction was decoupled from ecological dynamics

Comprehensive database on Induan (Lower Triassic) to Sinemurian (Lower Jurassic) marine bivalve genera and their paleobiogeographic record

Marine bivalve genera that were described or mentioned for Triassic and Lower Jurassic deposits worldwide are reviewed in terms of their validity, stratigraphic range, paleogeographic distribution, paleoautecology, and shell mineralogy. Data were originally compiled at species level and are arranged systematically. A brief discussion for each genus includes synonymy, taxonomic status, and included subgenera, as well as current uncertainties about their validity and range. The distribution of each genus is also shown on paleogeographic maps. Type species and first and last appearances of each genus are also mentioned. We recognize as valid 281 genera and their included subgenera, and we further discuss 148 genera (arranged alphabetically) that were mentioned for the study interval but are not included for different reasons. The purpose of this paper is to provide an updated critical assessment of all available basic information for each genus, in order to obtain a sound database to study the generic paleodiversity of marine bivalves in the time interval from the Induan (Early Triassic) to the Sinemurian (Early Jurassic). This was a critical time for bivalve evolution and diversification, which began with the recovery from the Permian¿Triassic extinction and ended with the recovery from the Triassic¿Jurassic extinction

Early Triassic-Early Jurassic Bivalve Diversity Dynamics

Bivalves are a highly diversified molluscan class, with a long history dating from early Cambrian times (Cope, 2000). Although the group already showed a steady diversification trend during the Paleozoic, it only became highly successful and expanded rapidly from the Mesozoic onward. The Triassic was, for bivalves, first a recovery period and later a biotic diversification event. It was also the time bivalves first fully exploited their evolutionary novelties.Facultad de Ciencias Naturales y Muse

Early Triassic-Early Jurassic Bivalve Diversity Dynamics

Bivalves are a highly diversified molluscan class, with a long history dating from early Cambrian times (Cope, 2000). Although the group already showed a steady diversification trend during the Paleozoic, it only became highly successful and expanded rapidly from the Mesozoic onward. The Triassic was, for bivalves, first a recovery period and later a biotic diversification event. It was also the time bivalves first fully exploited their evolutionary novelties.Facultad de Ciencias Naturales y Muse

Triassic/Jurassic bivalve extinction and recovery in the Neuquén Basin, Argentina

The extinction event at the end of the Triassic is one of the “big five” global crisis in the history of life in the marine realm. Nevertheless, that extinction and the subsequent biotic recovery are not so well known as others, and most of the published analyses were based on data from the Northern Hemisphere. Bivalves are one of the best studied groups in relation to the recovery after the end-Triassic extinction event. We analyze the Late Triassic extinction and Early Jurassic recovery of bivalve faunas within marine environments in the Atuel river area of the Neuquén Basin, Argentina (Riccardi et al., 1988; Lanés, 2005). The nearly continuous presence of marine stenohaline major taxa such as cnidarians, rhynchonelliform brachiopods, echinoderms and cephalopods indicate normal salinity (Damborenea & Mancenido, 2005). Data were collected from a thick and exceptionally well-exposed latest Triassic-earliest Jurassic section of the Andes, which allows a highresolution reconstruction of the local diversity dynamics.Facultad de Ciencias Naturales y Muse

Biotic recovery after the end-Triassic extinction event: Evidence from marine bivalves of the Neuquén Basin, Argentina

We analyze the Late Triassic extinction and Early Jurassic recovery of bivalve faunas within marine environments in the Atuel River area of the Neuquén Basin, Argentina. Data were collected from a hundred samples with invertebrates in a well-exposed uppermost Triassic to lower Jurassic section in the Neuquén Basin (southern Mendoza Province, Argentina) and allow a high-resolution reconstruction of the local diversity dynamics. The nearly continuous presence of marine stenohaline major taxa such as cnidarians, rhynchonelliform brachiopods, echinoderms and cephalopods indicates normal salinity throughout. All bivalve species were identified, and each occurrence was recorded in meters above the base. To analyze the systematic diversity trends, diversity curves were calculated on the basis of the first and last occurrence data for each bivalve species, and both total diversity and boundary crossers diversity were used. As a result, four main phases were identified: a) Triassic equilibrium phase (Rhaetian), with relatively high origination and extinction rates; b) extinction phase (latest Rhaetian to earliest Hettangian), with high extinction rates and low origination rates; c) recovery phase (late Early to early Late Hettangian), with high origination rates and almost null extinction rates; and d) Jurassic equilibrium phase (Late Hettangian-Sinemurian), again with similar and relatively high origination and extinction rates. The extinction and recovery phases are separated by a gap of about 135 m without identifiable benthonic invertebrates but with early Hettangian ammonites. On the other hand, bivalve palaeoecologic diversity seems to have been more homogeneous along the section, being dominated by attached epifaunal species, though before the extinction epifaunal habits were slightly surpassed by infaunal ones. Slight differences observed include a) shallow burrowers were more diverse during the Rhaetian than during the earliest Jurassic and b) epifaunal free-lying and semi-infaunal attached bivalves were more diverse after the Rhaetian extinction.Facultad de Ciencias Naturales y Muse

De extinción en extinción: la vida se reinventa una y otra vez

Episodios de extinción y recuperación que afectaron el desarrollo de la vida en la Tierra, cada uno extendido por lapsos de millones de años, más largos que la historia completa de la especie humana. Se ilustran mediante ejemplos del registro fósil marino del Mesozoico temprano, centrándonos en los invertebrados marinos del tiempo que transcurrió entre el fin del Pérmico y el del Jurásico temprano, es decir, entre hace unos 252 y 174 Ma. Los fósiles encontrados en el actual territorio argentino, en sitios que entonces estuvieron bajo el mar, aportan información útil para analizar este lapso de tiempo que contiene dos de las grandes extinciones señaladas y también una menor

Population response during an Oceanic Anoxic Event: The case of Posidonotis (Bivalvia) from the Lower Jurassic of the Neuquén Basin, Argentina

Benthonic marine species show a wide range of biological reactions to seawater chemical changes through time, from subtle adjustments to extinction. The Early Toarcian Oceanic Anoxic Event (T-OAE) was recently recognized in the Neuquén Basin, Argentina, confirming its global scope. The event was identified chemostratigraphically on the basis of a relative increase in marine organic carbon and a characteristic negative carbonisotope excursion (δ13Corg) in bulk rock and fossil wood in the upper Pliensbachian-lower Toarcian interval in the Arroyo Lapa section (Neuquén). Simultaneously with collection of lithological samples, a high-resolution biostratigraphical survey was carried out, and the scarce benthonic fauna was collected in order to check the biotic response to changing marine geochemical conditions. We present here an analysis of size and abundance data from the T-OAE interval in the Neuquén Basin for the dominant bivalve species, the paper-clam Posidonotis cancellata (Leanza), and relate these data to geochemical proxies (%TOC and δ13Corg) obtained at the same locality. The abundance of P. cancellata increased when the rest of the benthos diminished, reaching a maximum at the onset level of the T-OAE, and then decreasing. Size-frequency distributions show a noteworthy lack of juvenile shells. Shell size shows a positive correlation with %TOC in the whole section, though over the T-OAE interval proper, it decreases below the level where the maximum %TOC value is attained and increases above it. Posidonotis cancellata shows features of opportunistic species, such as high tolerance to hypoxia, strong dominance in impoverished environments and a strong dependence on primary productivity, but at the same time had a reproductive strategy more similar to equilibrium species, with relatively low juvenile mortality rates. Several anatomical features suggest adaptation to permanently dysaerobic environments. The species disappeared just before the minimum negative carbon-isotope value was reached; and by the same time the genus became extinct worldwide

Biogeographic response to major extinction events: The case of Triassic bivalves

Mass extinctions decimate the biota causing the disappearance of existing palaeobiogeographical patterns, which is not surprising given that palaeobiogeographical units are usually established on the basis of endemic taxa and these are the most affected in global crisis events. This paper deals with the evolution of the biogeographic pattern shown by bivalve genera in response to the end-Permian extinction event. From a network analysis (based on Simpson´s and Bray-Curtis´ similarity indices) and an Occurrence Ratio Profile, the Early Triassic is recognized as a time of general biogeographic homogeneity. The Middle Triassic shows a clear differentiation, with the evolution of many new genera, and an increase in endemicity, which reaches a maximum by the Carnian. During the Norian-Rhaetian the main biogeographic biochoremas appear well differentiated. Unlike other invertebrates, like brachiopods, bivalves show generally high similarity values between biogeographic units, and in this particular case study, Early Triassic biogeographic units appear more related to each other than to the corresponding Middle or Late Triassic units.Fil: Echevarría, Javier. Universidad Nacional de La Plata. Facultad de Ciencias Naturales y Museo. División Paleozoología Invertebrados; Argentina. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - La Plata; ArgentinaFil: Ros Franch, Sonia. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - La Plata; Argentina. Universidad de Valencia; Españ