Late Early Triassic climate change: Insights from carbonate carbon isotopes,sedimentary evolution and ammonoid paleobiogeography

The late Early Triassic sedimentary–facies evolution and carbonate carbon-isotope marine record (δ¹³C_carb) of ammonoid-rich, outer platform settings show striking similarities between the South China Block (SCB) and the widely distant Northern Indian Margin (NIM). The studied sections are located within the Triassic Tethys Himalayan belt (Losar section, Himachal Pradesh, India) and the Nanpanjiang Basin in the South China Block (Jinya section, Guangxi Province), respectively. Carbon isotopes from the studied sections confirm the previously observed carbon cycle perturbations at a time of major paleoceanographic changes in the wake of the end-Permian biotic crisis. This study documents the coincidence between a sharp increase in the carbon isotope composition and the worldwide ammonoid evolutionary turnover (extinction followed by a radiation) occurring around the Smithian–Spathian boundary.Based on recent modeling studies on ammonoid paleobiogeography and taxonomic diversity, we demonstrate that the late Early Triassic (Smithian and Spathian) was a time of a major climate change. More precisely, the end Smithian climate can be characterized by a warm and equable climate underlined by a flat, pole-to-equator, sea surface temperature (SST) gradient, while the steep Spathian SST gradient suggests latitudinally differentiated climatic conditions. Moreover, sedimentary evidence suggests a transition from a humid and hot climate during the Smithian to a dryer climate from the Spathian onwards. By analogy with comparable carbon isotope perturbations in the Late Devonian, Jurassic and Cretaceous we propose that high atmospheric CO₂ levels could have been responsible for the observed carbon cycle disturbance at the Smithian–Spathian boundary. We suggest that the end Smithian ammonoid extinction has been essentially caused by a warm and equable climate related to an increased CO₂ flux possibly originating from a short eruptive event of the Siberian igneous province. This increase in atmospheric CO₂ concentrations could have additionally reduced the marine calcium carbonate oversaturation and weakened the calcification potential of marine organisms, including ammonoids, in late Smithian oceans.     

Biodiversity and biogeography of middle–late liassic ammonoids: implications for the early Toarcian mass extinction

In Western Tethyan areas, the Toarcian stage begins with two important evolutionary events in ammonite faunas: (1) the disruption of Tethyan–Boreal provinciality; (2) a biological crisis linked with the oceanic anoxic event OAE. The analysis of these events has been addressed by constructing curves of ammonoid diversity (species richness, origination and extinction rates) in the Late Pliensbachian (= Domerian)–Early Toarcian interval in selected localities. Two diversity drops are recognized. The first one is recorded at the end of the Dactylioceras mirabile subzone and reflects the disruption of Tethyan–Boreal provinciality, through the progressive extinction of the Boreal endemic family Amaltheidae that occupied the north-western European seas during the whole Pliensbachian on the one hand, and the extinction of Late Domerian Ammonitina endemic to the Mediterranean areas on the other hand. The Early Toarcian homogeneization of Mediterranean and north-western European ammonoid faunas was reached via elimination of both Boreal and Mediterranean endemics with differential rates of extinction in the two palaeogeographic domains and the subsequent geographical expansion of Tethyan-derived ammonoids. The second, dramatic drop in ammonite diversity in the upper part of the Dactylioceras semicelatum subzone coincided with the onset of OAE. It also affected epioceanic ammonoid clades like Phyllocerataceae and Lytocerataceae. These two drops are interpreted as two distinct extinctions and not as episodes of a single, stepwise event. Complex relations between ammonoid diversity and sea-level changes are suggested by trends in endemism, which may be reversed during either a single transgression or a single regression.     

Exploring the major depletions of conodont diversity during the Triassic

In this paper, we show that the Triassic fossil record reflects just two great depletions of conodont diversity before the Rhaetian, which occurred in the Smithian (Olenekian, Early Triassic) and in the Julian (Carnian, Late Triassic). By exploring this context, our results highlighted that they respond to different origination–extinction dynamics. Thus, while the Smithian diversity depletion can be interpreted as a consequence of elevated extinction, the Julian diversity depletion was triggered by fluctuations in origination regime. This evidence suggests that, despite the role of extinction on diversity losses, conodonts suffered crucial changes on the origination regimes during the Late Triassic which triggered these events. Notwithstanding, our results indicate that the end-Triassic diversity depletion of conodonts was produced by background extinction levels in a context of lower origination. This suggests that several biological factors, rather than a unique, environmental and/or cyclic cause, could have influenced the evolutionary history of conodonts during the Triassic.     

Ammonoid recovery from the Late Permian mass extinction event

Previous research indicated that ammonoid taxonomic diversity exploded after the Late Permian mass extinction, regaining pre-extinction levels by the Late Induan (Dienerian substage). From taxonomic analyses it had been inferred that ammonoids recovered rapidly, relative to other marine invertebrate groups. Complementing taxonomic metrics with morphologic and spatial data revealed more complex recovery dynamics. Morphological analysis indicated that ammonoids did not fully recover until the Spathian or Anisian. Taxonomic diversity is a poor predictor of disparity during the recovery. Spatial partitioning of taxonomic and morphological diversity revealed spatially homogeneous recovery patterns. Combining taxonomic, morphological, and spatial data refined interpretations of Triassic ammonoid recovery patterns and indicated that ecological, not intrinsic, factors were the probable control on ammonoid recovery rates. To cite this article: A.J. McGowan, C. R. Palevol 4 (2005).     

Global Taxonomic Diversity of Anomodonts (Tetrapoda,Therapsida) and the Terrestrial Rock Record Across the Permian-Triassic Boundary

The end-Permian biotic crisis (∼252.5 Ma) represents the most severe extinction event in Earth''s history. This paper investigates diversity patterns in Anomodontia, an extinct group of therapsid synapsids (‘mammal-like reptiles’), through time and in particular across this event. As herbivores and the dominant terrestrial tetrapods of their time, anomodonts play a central role in assessing the impact of the end-Permian extinction on terrestrial ecosystems. Taxonomic diversity analysis reveals that anomodonts experienced three distinct phases of diversification interrupted by the same number of extinctions, i.e. an end-Guadalupian, an end-Permian, and a mid-Triassic extinction. A positive correlation between the number of taxa and the number of formations per time interval shows that anomodont diversity is biased by the Permian-Triassic terrestrial rock record. Normalized diversity curves indicate that anomodont richness continuously declines from the Middle Permian to the Late Triassic, but also reveals all three extinction events. Taxonomic rates (origination and extinction) indicate that the end-Guadalupian and end-Permian extinctions were driven by increased rates of extinction as well as low origination rates. However, this pattern is not evident at the final decline of anomodont diversity during the Middle Triassic. Therefore, it remains unclear whether the Middle Triassic extinction represents a gradual or abrupt event that is unique to anomodonts or more common among terrestrial tetrapods. The end-Permian extinction represents the most distinct event in terms of decline in anomodont richness and turnover rates.     

Maxima im Tempo der Evolution karbonischer Ammonoideen

On the basis of evolutionary rates several maxima and minima in the diversity of ammonoids can be recognized during Upper Devonian and Carboniferous times. The development of new characters, the beginning and continuation of phylogenetic trends and the origination as well as the extinction of bradytelic genera are restricted to diversity maxima.     

AMMONOIDS ACROSS THE PERMIAN/TRIASSIC BOUNDARY: A CLADISTIC PERSPECTIVE

Abstract:  The rapid diversification of ceratitid ammonoids during the earliest Mesozoic has been taken at face value as an example of explosive radiation following the Permian/Triassic mass extinction. However, the validity of this interpretation has never been tested within a phylogenetic framework. A total evidence cladistic analysis of Mid–Late Permian and Induan (earliest Triassic) ammonoids confirms the monophyly of the Ceratitida. Partitioned phylogenetic analysis of suture line characters vs. shell shape and ornament characters confirms the importance of suture-line characters for resolving the higher taxonomy of ammonoids. When the cladogram is compared with the observed fossil record, the resultant tree implies that the divergence of a number of early Triassic lineages actually occurred during the latest Permian. If these range extensions are taken into account the ammonoid per-genus extinction rate across the Permian/Triassic boundary drops from c. 85 per cent to c. 60 per cent.     

Paedomorphosis of ammonoids as a result of sealevel fluctuations in the Late Devonian Wocklumeria Stufe

A remarkable diversification of several independent ammonoid lineages with high evolutionary rates occurred in the Late Devonian Wocklumeria Stufe. Many speciation events led to paedomorphic ammonoids that display a striking range of conch shapes, sculpture, and ornamentation. In the goniatite family Prionoceratidae, the transition from normal Mimimitoceras species to paedomorphic Balvia species provides an example of rapid size decrease combined with an early character developmental offset arising from progenesis. Adults of early Balvia species largely have the preadult ancestral morphology of Mimimitoceras , but later evolving species acquire distinct conch and ornamentation types. Progenetic ammonoid species also appeared within the clymeniid family Kosmoclymeniidae and probably in the Glatzielliidae. In the clymeniid family Parawocklumeriidae, evolution is characterized by the extension of tri-segmented and triangularly coiled whorls found only in juveniles of earlier species, to the adults of later species. This is interpreted as resulting from neoteny. The distribution of paedomorphic ammonoids in the Late Devonian Wocklumeria Stufe is closely correlated to relative sealevel changes. The regressive trend in the lower two-thirds of the Wocklumeria Stufe is interpreted as the cause of a diversification of the pelagic habitat during unstable conditions, and as an extrinsic factor inducing heterochronic change. Some ammonoids reacted by rapid maturation and faster reproductive rates, giving the opportunity to exploit a wider range of niches. The apparent consequence was the formation of several allopatric species. □ Ammonoidea, Late Devonian, evolution, heterochrony, sealevel changes.     

The two Early Toarcian (Early Jurassic) extinction events in ammonoids

The Early Toarcian (Early Jurassic) biological crisis was one of the ‘minor’ mass extinctions. It is linked with an oceanic anoxic event. Fossil data from sections located in northwestern European (epicontinental platforms and basins) and Tethyan (distal, epioceanic) areas indicate that Late Pliensbachian–Early Toarcian ammonoids experienced two extinction events during the Early Toarcian. The older one is linked with disruption of the Tethyan–Boreal provinciality, whereas the younger event correlates with the onset of anoxia and corresponds with the Early Toarcian mass‐extinction event. These two extinctions cannot be interpreted as episodes of a single, stepwise, event. Values of the net diversification, more than the number of extinctions, allow the two extinction events to be clearly recognized and distinguished. Values of regional net diversification for northwestern European and Tethyan faunas point to greater evolutionary dynamics in the epioceanic areas. The inclusion of Mediterranean faunas in the database proves that the ammonite turnover at the Early Toarcian mass‐extinction event was more important than previously thought. Progenitor (evolute Neolioceratoides), survivor (Dactylioceras, Polyplectus pluricostatus) and Lazarus (Procliviceras) taxa have been recognized. Different selectivity patterns are shown for the two events. The first one, linked to the disruption of the Tethyan–Boreal provinciality, has mainly affected ammonites adapted to epicontinental platforms. In the mass‐extinction event, no selectivity is recognized, because also Phylloceratina and Lytoceratina were deeply affected at species level, although their wide biogeographical distribution at clade level was a significant buffer against extinction. In contrast to Palaeozoic mass extinctions, ammonoid survivors and Lazarus taxa are characterized by complex sutures: Phylloceratina (long‐ranging ammonoids) and Polyplectus (relatively long‐ranging compared to other Ammonitina).     

Latest Smithian (Early Triassic) ammonoid assemblages in Utah (western USA basin) and their implications for regional biostratigraphy,biogeography and placement of the Smithian/Spathian boundary

The late Smithian extinction represents a major event within the Early Triassic. This event generally corresponds to a succession of two, possibly three successively less diverse, cosmopolitan ammonoid assemblages, which when present, provide a robust biostratigraphic framework and precise correlations at different spatial scales. In the western USA basin, known occurrences of latest Smithian taxa are rare and until now, have only been documented from northeastern Nevada. Based on these restricted basinal occurrences, a regional zone representing the latest Smithian was postulated but not corroborated, as representative taxa had not yet been reported from outside Nevada. Here we document two new ammonoid assemblages from distant localities in northern Utah, overlying the late Smithian Anasibirites beds and characterized by the unambiguous co-occurrence of Xenoceltites subevolutus and Pseudosageceras augustum. The existence of a latest Smithian zone in the western USA basin is therefore validated, facilitating the identification of the Smithian/Spathian boundary and intra-basin correlation. This zone also correlates with the latest Smithian zone recognized from southern Tethyan basins. Additionally, these new data support other observed occurrences of Xenoceltites subevolutus throughout most of the late Smithian.     

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.     

Quantitative biochronology of Devonian ammonoids from Morocco and proposals for a refined unitary association method

Monnet, C., Klug, C., Goudemand, N., De Baets, K. & Bucher, H. 2011: Quantitative biochronology of Devonian ammonoids from Morocco and proposals for a refined unitary association method. Lethaia, Vol. 44, pp. 469–489. Based on a rich dataset, the biostratigraphy of the late Emsian and the Eifelian (Early–Middle Devonian) ammonoids from the Moroccan Tafilalt is re‐evaluated. We processed this dataset (comprising 53 species from 15 sections) with the unitary association method (UAM), by means of the UA‐graph freeware. This led to the construction of a sequence of 17 UAs (maximal sets of actually or virtually coexisting taxa), which are grouped into 10 laterally reproducible association zones. This biostratigraphical subdivision of this interval is in some parts finer than the classically used empirical stratigraphical scheme and than a previous graphic correlation analysis. It enabled us to measure regional ammonoid diversity of this interval in detail. The UAM is a powerful biochronological method, which benefits from complementary tools to analyse conflicting inter‐taxon stratigraphical relationships inherent to complex biostratigraphical datasets. In cases of under‐constrained superpositional relationships between the taxa, the UAM can yield results, which are not parsimonious in terms of reconstructed virtual coexistences. We suggest several additions to complement the algorithmic steps of the method. The most important is the exhaustive or heuristic reconstruction of possible solutions resolving the observed biostratigraphical contradictions (conflicting inter‐taxon superpositional relationships and cycles between maximal cliques) and the selection among the solutions of the most‐parsimonious one(s) in terms of reconstructed virtual coexistences. Multiple equivalent results may then be processed with standard consensus techniques. Finally, the robustness of the results can be tested by bootstrapping methods to provide confidence estimates on the ranges and associations of studied taxa. □Ammonites, Anti‐Atlas, biostratigraphy, correlation, zonation, diversity.     

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.     

Evolutionary rates of the Triassic marine macrofauna and sea-level changes: Evidences from the Northwestern Caucasus,Northern Neotethys (Russia)

A diverse Triassic marine macrofauna from the Northwestern Caucasus sheds new light on the biotic evolution after the end-Permian mass extinction. In the early Mesozoic, the study area was located on the northern margin of the Neotethys Ocean. Data on stratigraphic ranges of 130 genera of brachiopods, bivalves, ammonoids, corals, and sponges have been used to calculate the changes in two evolutionary rates, namely faunal transformation rate (FTR) and rate of transformation of the taxonomic diversity structure (TTDSR). The FTR demonstrates the changes in the generic composition of assemblages through geologic time, whereas the TTDSR indicates changes in the generic control of the species diversity. The Triassic marine macrofauna of the Northwestern Caucasus was characterized by very high FTR and TTDSR during the Early Triassic through early Late Triassic. The FTR slowed in the Middle Triassic, and accelerated again in the Carnian–Norian. In contrast, the FTR was abnormally slow in the Norian–Rhaetian. A remarkable turnover among macrofauna occurred at the Carnian–Norian transition. Regional sea-level changes were similar to the global eustatic fluctuations. It is difficult to establish their direct connections with changes in the evolutionary rates, although the turnover at the Carnian–Norian boundary coincided with a prominent regressive episode. In general, high evolutionary rates reported for the Triassic marine macrofauna of the Northwestern Caucasus may be explained as a consequence of the devastating end-Permian mass extinction.     

Ecological interactions on macroevolutionary time scales: clams and brachiopods are more than ships that pass in the night

Competition among organisms has ecological and evolutionary consequences. However, whether the consequences of competition are manifested and measureable on macroevolutionary time scales is equivocal. Marine bivalves and brachiopods have overlapping niches such that competition for food and space may occur. Moreover, there is a long‐standing debate over whether bivalves outcompeted brachiopods evolutionarily, because brachiopod diversity declined through time while bivalve diversity increased. To answer this question, we estimate the origination and extinction dynamics of fossil marine bivalve and brachiopod genera from the Ordovician through to the Recent while simultaneously accounting for incomplete sampling. Then, using stochastic differential equations, we assess statistical relationships among diversification and sampling dynamics of brachiopods and bivalves and five paleoenvironmental proxies. None of these potential environmental drivers had any detectable influence on brachiopod or bivalve diversification. In contrast, elevated bivalve extinction rates causally increased brachiopod origination rates, suggesting that bivalves have suppressed brachiopod evolution.     

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 and extinction in the Cenozoic history of Caribbean reefs

Occurrences of reef corals are examined at Caribbean fossil localities to determine how biodiversity has changed within the region over the past 50 million years. Analyses of 294 species (66 genera) at 58 fossil localities show that Caribbean generic diversity rose to 44 between 50–22 Ma, ranged from 32–39 between 22–2 Ma, and dropped to 25 afterwards. Regional species diversity was high at 40–36 Ma, 28–22 Ma, and 5–2 Ma. Origination rates were elevated throughout each high diversity interval, but extinction was concentrated near the end of each interval. Regional highs of origination and extinction, therefore, differed in timing and duration, causing the observed regional diversity increases during the three remarkably long intervals of turnover. Highs of generic origination decreased in magnitude as immigration from the Mediterranean ceased, but speciation highs increased in association with emergence of the Central American isthmus. Peaks of extinction coincided with regional changes in climate and oceanic circulation. Maximum species diversities within assemblages increased to 40–60 between 50–36 Ma, and have remained relatively constant ever since. Assemblage compositions differed among localities having similar ages and environments, suggesting that the timing and pattern of turnover varied across the region. Stable diversities but variable compositions within assemblages suggest that dispersal and recruitment influenced the pattern of faunal change during turnover. Accepted: 22 August 1999     

Evolution and the latitudinal diversity gradient: speciation, extinction and biogeography

A latitudinal gradient in biodiversity has existed since before the time of the dinosaurs, yet how and why this gradient arose remains unresolved. Here we review two major hypotheses for the origin of the latitudinal diversity gradient. The time and area hypothesis holds that tropical climates are older and historically larger, allowing more opportunity for diversification. This hypothesis is supported by observations that temperate taxa are often younger than, and nested within, tropical taxa, and that diversity is positively correlated with the age and area of geographical regions. The diversification rate hypothesis holds that tropical regions diversify faster due to higher rates of speciation (caused by increased opportunities for the evolution of reproductive isolation, or faster molecular evolution, or the increased importance of biotic interactions), or due to lower extinction rates. There is phylogenetic evidence for higher rates of diversification in tropical clades, and palaeontological data demonstrate higher rates of origination for tropical taxa, but mixed evidence for latitudinal differences in extinction rates. Studies of latitudinal variation in incipient speciation also suggest faster speciation in the tropics. Distinguishing the roles of history, speciation and extinction in the origin of the latitudinal gradient represents a major challenge to future research.     

Molecular phylogenies and historical biogeography of a circumtropical group of gastropods (Genus: Nerita): implications for regional diversity patterns in the marine tropics

To determine how historical processes, namely speciation, extinction, and dispersal, have contributed to regional species diversity patterns across the marine tropics, we examined the biogeographical history of a circumtropical genus of intertidal gastropods. A species-level phylogeny of Nerita, representing approximately 87% of extant species, was developed from 1608bp of mitochondrial (COI and 16S) and nuclear (ATPSalpha) markers. Phylogenetic relationships generally corresponded to prior classifications; however, comprehensive sampling revealed a number of previously undetected ESUs. Using the resulting tree as a framework, we combined geographical distributions and fossil evidence to reconstruct ancestral ranges, produce a time-calibrated chronogram, and estimate diversification rates. Analyses revealed two monophyletic eastern Pacific+Atlantic (EPA) clades, each of which likely split from an Indo-West Pacific (IWP) sister clade prior to an early Miocene Tethys Seaway closure. More recent diversification throughout the IWP appears to have been driven by both vicariance and dispersal events; EPA diversity has been further shaped by speciation across the Central American Seaway prior to its closure and dispersal across the Atlantic. Despite the latter, inter-regional dispersal has been rare, and likely contributes little to regional diversity patterns. Similarly, infrequent transitions into temperate regions combined with reduced diversification rates may explain low diversity in West and South Pacific clades. Since origination, Nerita diversification appears remarkably constant, with the exception of a lag in the late Eocene-early Oligocene and elevated rates in the late Oligocene-early Miocene. However, a comparison among regions suggested that IWP clades have experienced, on average, higher rates of speciation. Fossil evidence indicates that the EPA likely witnessed greater extinction relative to the IWP. We propose that regional differences in species diversity in Nerita have been largely shaped by differential rates of speciation and extinction.     

Correlations in fossil extinction and origination rates through geological time

Recent analyses have suggested that extinction and origination rates exhibit long-range correlations, implying that the fossil record may be controlled by self-organized criticality or other scale-free internal dynamics of the biosphere. Here we directly test for correlations in the fossil record by calculating the autocorrelation of extinction [corrected] and origination rates through time. Our results show that extinction rates are uncorrelated beyond the average duration of a stratigraphic interval. Thus, they lack the long-range correlations predicted by the self-organized criticality hypothesis. In contrast, origination rates show strong autocorrelations due to long-term trends. After detrending, origination rates generally show weak positive correlations at lags of 5-10 million years (Myr) and weak negative correlations at lags of 10-30 Myr, consistent with aperiodic oscillations around their long-term trends. We hypothesize that origination rates are more correlated than extinction rates because originations of new taxa create new ecological niches and new evolutionary pathways for reaching them, thus creating conditions that favour further diversification.