Impact of the Late Triassic mass extinction on functional diversity and composition of marine ecosystems

Mass extinctions have profoundly influenced the history of life, not only through the death of species but also through changes in ecosystem function and structure. Importantly, these events allow us the opportunity to study ecological dynamics under levels of environmental stress for which there are no recent analogues. Here, we examine the impact and selectivity of the Late Triassic mass extinction event on the functional diversity and functional composition of the global marine ecosystem, and test whether post‐extinction communities in the Early Jurassic represent a regime shift away from pre‐extinction communities in the Late Triassic. Our analyses show that, despite severe taxonomic losses, there is no unequivocal loss of global functional diversity associated with the extinction. Even though no functional groups were lost, the extinction event was, however, highly selective against some modes of life, in particular sessile suspension feeders. Although taxa with heavily calcified skeletons suffered higher extinction than other taxa, lightly calcified taxa also appear to have been selected against. The extinction appears to have invigorated the already ongoing faunal turnover associated with the Mesozoic Marine Revolution. The ecological effects of the Late Triassic mass extinction were preferentially felt in the tropical latitudes, especially amongst reefs, and it took until the Middle Jurassic for reef ecosystems to fully recover to pre‐extinction levels.     

Body length of bony fishes was not a selective factor during the biggest mass extinction of all time

The Permo‐Triassic mass extinction devastated life on land and in the sea, but it is not clear why some species survived and others went extinct. One explanation is that lineage loss during mass extinctions is a random process in which luck determines which species survive. Alternatively, a phylogenetic signal in extinction may indicate a selection process operating on phenotypic traits. Large body size has often emerged as an extinction risk factor in studies of modern extinction risk, but this is not so commonly the case for mass extinctions in deep time. Here, we explore the evolution of non‐teleostean Actinopterygii (bony fishes) from the Devonian to the present day, and we concentrate on the Permo‐Triassic mass extinction. We apply a variety of time‐scaling metrics to date the phylogeny, and show that diversity peaked in the latest Permian and declined severely during the Early Triassic. In line with previous evidence, we find the phylogenetic signal of extinction increases across the mass extinction boundary: extinction of species in the earliest Triassic is more clustered across phylogeny compared to the more randomly distributed extinction signal in the late Permian. However, body length plays no role in differential survival or extinction of taxa across the boundary. In the case of fishes, size did not determine which species survived and which went extinct, but phylogenetic signal indicates that the mass extinction was not a random field of bullets.     

Body size changes in bivalves of the family Limidae in the aftermath of the end‐Triassic mass extinction: the Brobdingnag effect

Reduction in body size of organisms following mass extinctions is well‐known and often ascribed to the Lilliput effect. This phenomenon is expressed as a temporary body size reduction within surviving species. Despite its wide usage the term is often loosely applied to any small post‐extinction taxa. Here we assess the size of bivalves of the family Limidae (Rafineque) prior to, and in the aftermath of, the end‐Triassic mass extinction event. Of the species studied only one occurs prior to the extinction event, though is too scarce to test for the Lilliput effect. Instead, newly evolved species originate at small body sizes and undergo a within‐species size increase, most dramatically demonstrated by Plagiostoma giganteum (Sowerby) which, over two million years, increases in size by 179%. This trend is seen in both field and museum collections. We term this within‐species size increase of newly originated species in the aftermath of mass extinction, the Brobdingnag effect, after the giants that were contemporary with the Lilliputians in Swift's Gulliver's Travels. The size increase results from greater longevity and faster growth rates. The cause of the effect is unclear, although it probably relates to improved environmental conditions. Oxygen‐poor conditions in the Early Jurassic are associated with populations of smaller body size caused by elevated juvenile mortality but these are local/regional effects that do not alter the long‐term, size increase. Although temperature‐size relationships exist for many organisms (Temperature‐Size Rule and Bergmann's Rule), the importance of this is unclear here because of a poorly known Early Jurassic temperature record.     

Effect of end-Triassic CO<Subscript>2</Subscript> maximum on carbonate sedimentation and marine mass extinction

Correlation of stratigraphic sections from different continents suggests a worldwide interruption of carbonate sedimentation at the Triassic–Jurassic boundary, which coincided with one of the most catastrophic mass extinctions in the Phanerozoic. Both events are linked by a vulcanogenic maximum of carbon dioxide, which led to a temporary undersaturation of sea water with respect to aragonite and calcite and a corresponding suppression of carbonate sedimentation including non-preservation of calcareous skeletons. Besides the frequently cited climatic effect of enhanced carbon dioxide, lowering the saturation state of sea water with respect to calcium carbonate was an additional driving force of the end-Triassic mass extinction, which chiefly affected organisms with thick aragonitic or high-magnesium calcitic skeletons. Replacement of aragonite by calcite, as found in the shells of epifaunal bivalves, was an evolutionary response to this condition.     

Recovery from the most profound mass extinction of all time

The end-Permian mass extinction, 251 million years (Myr) ago, was the most devastating ecological event of all time, and it was exacerbated by two earlier events at the beginning and end of the Guadalupian, 270 and 260 Myr ago. Ecosystems were destroyed worldwide, communities were restructured and organisms were left struggling to recover. Disaster taxa, such as Lystrosaurus, insinuated themselves into almost every corner of the sparsely populated landscape in the earliest Triassic, and a quick taxonomic recovery apparently occurred on a global scale. However, close study of ecosystem evolution shows that true ecological recovery was slower. After the end-Guadalupian event, faunas began rebuilding complex trophic structures and refilling guilds, but were hit again by the end-Permian event. Taxonomic diversity at the alpha (community) level did not recover to pre-extinction levels; it reached only a low plateau after each pulse and continued low into the Late Triassic. Our data showed that though there was an initial rise in cosmopolitanism after the extinction pulses, large drops subsequently occurred and, counter-intuitively, a surprisingly low level of cosmopolitanism was sustained through the Early and Middle Triassic.     

Bivalves and evolutionary resilience: old skills and new strategies to recover from the P/T and T/J extinction events

Diversity dynamics among bivalves during the Triassic and Early Jurassic provides the opportunity to analyse the recovery patterns after two mass extinctions: Permian/Triassic and Triassic/Jurassic (T/J). The results presented here are based on a newly compiled worldwide genus-level database and are contrasted to the main morphological characters of the different taxonomical (orders and their constituent families and genera) and ecological groups. Many of such morphological characters are innovations appearing during the time span considered. Diversity and evolutionary rates were assessed and compared between these groups. During the Early Triassic there was a slow recovery, dominated by epifaunal taxa, the order Pectinida being the most diverse. The major post-Permian radiation took place during the Anisian, with several morphological and ecological innovations appearing and/or diversifying. The Late Triassic was a time of great diversification and ecological specialisation. Although the T/J was a true mass extinction for bivalves, it was not indiscriminate as its impact was stronger on specialised orders and not all ecological categories were equally affected. Recovery during earliest Jurassic was fast, confirming the high-evolutionary resilience of bivalve molluscs, except for groups with thick shells and tropical distribution, probably because of a biocalcification crisis.     

Oceanic redox evolution across the end-Permian mass extinction at Penglaitan section,South China

The end-Permian mass extinction (EPME) was known as the most severe biocrisis of the past 600 Ma. In order to explore the redox state of deep water environments, and the causal relationship between anoxia/euxinia and the EPME, this study selected the Penglaitan section in Guangxi, China, and measured the iron speciation and concentrations of trace elements and major elements. Several redox proxy approaches, including both iron speciation proxies and the excess fractions and enrichment factors of redox sensitive elements (RSEs), indicate that anoxia in the Penglaitan depositional environment was sporadic, with the majority of the section deposited under oxic conditions. This includes the interval immediately prior to and during the EPME; euxinia did not occur at the Penglaitan section around the EPME. Thus, anoxia was not the primary driver for the EPME at the Penglaitan section, and the accumulation of free H₂S in the oceanic water-column could not have been the only proximal kill mechanism for the EPME at the Penglaitan section.     

Archosauromorph extinction selectivity during the Triassic–Jurassic mass extinction

Many traits have been linked to extinction risk among modern vertebrates, including mode of life and body size. However, previous work has indicated there is little evidence that body size, or any other trait, was selective during past mass extinctions. Here, we investigate the impact of the Triassic–Jurassic mass extinction on early Archosauromorpha (basal dinosaurs, crocodylomorphs and their relatives) by focusing on body size and other life history traits. We built several new archosauromorph maximum‐likelihood supertrees, incorporating uncertainty in phylogenetic relationships. These supertrees were then employed as a framework to test whether extinction had a phylogenetic signal during the Triassic–Jurassic mass extinction, and whether species with certain traits were more or less likely to go extinct. We find evidence for phylogenetic signal in extinction, in that taxa were more likely to become extinct if a close relative also did. However, there is no correlation between extinction and body size, or any other tested trait. These conclusions add to previous findings that body size, and other traits, were not subject to selection during mass extinctions in closely‐related clades, although the phylogenetic signal in extinction indicates that selection may have acted on traits not investigated here.     

Geographical distribution and extinction risk: lessons from Triassic–Jurassic marine benthic organisms

Aim To evaluate the influence of geographical distribution on the extinction risk of benthic marine invertebrates using data from the fossil record, both during times of background extinction and across a mass‐extinction episode. Total geographical range is contrasted with proxies of global abundance to assess the relationships between the two essential components of geographical distribution and extinction risk. Location A global occurrence data base of fossil benthic macro‐organisms from the Triassic and Jurassic periods was used for this study. Methods Geographical distributions and biodiversity dynamics were assessed for each genus (all taxa) or species (bivalves) based on a sample‐standardized data set and palaeogeographical reconstructions. Geographical ranges were measured by the maximum great circle distance of a taxon within a stratigraphic interval. Global abundance was assessed by the number of localities at which a taxon was recorded. Widespread and rare taxa were separated using median and percentile values of the frequency distributions of occurrences. Results The frequency distribution of geographical ranges is very similar to that for modern taxa. Although no significant correlation could be established between local abundance and geographical range, proxies of global abundance are strongly correlated with geographical range. Taxon longevities are correlated with both mean geographical range and mean global abundance, but range size appears to be more critical than abundance in determining extinction risk. These results are valid when geographical distribution is treated as a trait of taxa and when assessed for individual geological stages. Main conclusions Geographical distribution is a key predictor of extinction risk of Triassic and Jurassic benthic marine invertebrates. An important exception is in the end‐Triassic mass extinction, which equally affected geographically restricted and widespread genera, as well as common and rare genera. This suggests that global diversity crises may curtail the role of geographical distribution in determining extinction risk.     

Phanerozoic survivors: Actinopterygian evolution through the Permo‐Triassic and Triassic‐Jurassic mass extinction events

Actinopterygians (ray‐finned fishes) successfully passed through four of the big five mass extinction events of the Phanerozoic, but the effects of these crises on the group are poorly understood. Many researchers have assumed that the Permo‐Triassic mass extinction (PTME) and end‐Triassic extinction (ETE) had little impact on actinopterygians, despite devastating many other groups. Here, two morphometric techniques, geometric (body shape) and functional (jaw morphology), are used to assess the effects of these two extinction events on the group. The PTME elicits no significant shifts in functional disparity while body shape disparity increases. An expansion of body shape and functional disparity coincides with the neopterygian radiation and evolution of novel feeding adaptations in the Middle‐Late Triassic. Through the ETE, small decreases are seen in shape and functional disparity, but are unlikely to represent major changes brought about by the extinction event. In the Early Jurassic, further expansions into novel areas of ecospace indicative of durophagy occur, potentially linked to losses in the ETE. As no evidence is found for major perturbations in actinopterygian evolution through either extinction event, the group appears to have been immune to two major environmental crises that were disastrous to most other organisms.     

The Luoping biota: exceptional preservation, and new evidence on the Triassic recovery from end-Permian mass extinction

The timing and nature of biotic recovery from the devastating end-Permian mass extinction (252 Ma) are much debated. New studies in South China suggest that complex marine ecosystems did not become re-established until the middle–late Anisian (Middle Triassic), much later than had been proposed by some. The recently discovered exceptionally preserved Luoping biota from the Anisian Stage of the Middle Triassic, Yunnan Province and southwest China shows this final stage of community assembly on the continental shelf. The fossil assemblage is a mixture of marine animals, including abundant lightly sclerotized arthropods, associated with fishes, marine reptiles, bivalves, gastropods, belemnoids, ammonoids, echinoderms, brachiopods, conodonts and foraminifers, as well as plants and rare arthropods from nearby land. In some ways, the Luoping biota rebuilt the framework of the pre-extinction latest Permian marine ecosystem, but it differed too in profound ways. New trophic levels were introduced, most notably among top predators in the form of the diverse marine reptiles that had no evident analogues in the Late Permian. The Luoping biota is one of the most diverse Triassic marine fossil Lagerstätten in the world, providing a new and early window on recovery and radiation of Triassic marine ecosystems some 10 Myr after the end-Permian mass extinction.     

The influence of denitrifying seawater on graptolite extinction and diversification during the Hirnantian (latest Ordovician) mass extinction event

A continuous trench exposure within the uppermost type Vinini Formation at Vinini Creek, Roberts Mountains, Nevada, provides an unparalleled opportunity to examine the fate of graptolites, prominent Paleozoic zooplankton, during most of the Hirnantian mass extinction event. On the basis of a detailed biostratigraphic and sedimentological dataset, the relatively complete extinction record is examined in the context of ecological constraints, and it is found to reflect an ecological collapse driven by glacio-eustatic sea-level fall and associated changes in oceanic circulation. Diverse graptolite populations of the Dicranograptidae-Diplograptidae-Orthograptidae (DDO) fauna, which flourished in denitrifying waters within the oceanic oxygen-minimum zone (OMZ) during sea-level highstand, largely vanished with the loss of these conditions during glacio-eustatic sea-level fall. However, populations of one clade, the normalograptids, which inhabited the oxygenated waters of the photic zone, not only survived but diversified. These survivors gave rise to rapid recolonization and diversification with re-establishment of the oxygen-minimum and denitrifying conditions during post-Hirnantian sea-level rise. This ecological model also applies globally to other well-documented coeval stratigraphic intervals, representing both oceanic and platform sea settings.     

Primate extinction risk and historical patterns of speciation and extinction in relation to body mass

Body mass is thought to influence diversification rates, but previous studies have produced ambiguous results. We investigated patterns of diversification across 100 trees obtained from a new Bayesian inference of primate phylogeny that sampled trees in proportion to their posterior probabilities. First, we used simulations to assess the validity of previous studies that used linear models to investigate the links between IUCN Red List status and body mass. These analyses support the use of linear models for ordinal ranked data on threat status, and phylogenetic generalized linear models revealed a significant positive correlation between current extinction risk and body mass across our tree block. We then investigated historical patterns of speciation and extinction rates using a recently developed maximum-likelihood method. Specifically, we predicted that body mass correlates positively with extinction rate because larger bodied organisms reproduce more slowly, and body mass correlates negatively with speciation rate because smaller bodied organisms are better able to partition niche space. We failed to find evidence that extinction rates covary with body mass across primate phylogeny. Similarly, the speciation rate was generally unrelated to body mass, except in some tests that indicated an increase in the speciation rate with increasing body mass. Importantly, we discovered that our data violated a key assumption of sample randomness with respect to body mass. After correcting for this bias, we found no association between diversification rates and mass.     

Evolutionary and biogeographical shifts in response to the Late Ordovician mass extinction

The Late Ordovician mass extinction was an interval of high extinction with inferred low ecological selectivity, resulting in little change in community structure after the event. In contrast, the mass extinction may have fundamentally changed evolutionary dynamics in the surviving groups. We investigated the phylogenetic relationships among strophomenoid brachiopods, a diverse brachiopod superfamily that was a primary component of Ordovician ecosystems. Four Ordovician families/subfamilies sampled in the analysis (Rafinesquinidae, Strophomeninae, Glyptomenidae and Furcitellinae) were reconstructed as monophyletic groups, and the base of the strophomenoid clade that dominated the Silurian recovery was reconstructed as diversifying alongside these families during the Middle Ordovician. We time‐calibrated the phylogeny and used geographical occurrences to investigate biogeographical changes in the strophomenoids through time with the R package BiogeoBEARS . Our results indicate that extinction was higher in taxa whose ranges were constrained to tropical or subtropical regions. Furthermore, our results suggest important shifts in the diversification patterns of these brachiopods after the mass extinction. While most of the strophomenoid families survived the Late Ordovician event, ecologically abundant taxonomic groups during the Ordovician were either driven to extinction, reduced in diversity, or slowly died off during the Silurian. The new abundant strophomenoid taxa derived from one clade (consisting of Silurian–Devonian groups such as Douvillinidae, Strophodontidae and Amphistrophiidae) that diversified during the post‐extinction radiation. Our results suggest the selective diversification during the Silurian radiation, rather than selective extinction in the Late Ordovician, had a greater impact on the evolutionary history of strophomenoid brachiopods.     

Large ecosystems in transition: Bifurcations and mass extinction

We propose a model of multispecies populations surviving on distributed resources. System dynamics are investigated under changes in abiotic factors such as the climate, as parameterized through environmental temperature. In particular, we introduce a feedback between species abundances and resources via abiotic factors. This model is apparently the first of its kind to include a feedback mechanism coupling climate and population dynamics. Moreover, we take into account self-limitation effects. The model explains the coexistence of many species, yet also displays the possibility of catastrophic bifurcations, where all species become extinct under the influence of abiotic factors. We show that as these factors change there are different regimes of ecosystem behavior, including a possibly chaotic regime when abiotic influences are sufficiently strong.     

Bias in phylogenetic measurements of extinction and a case study of end‐Permian tetrapods

Extinction risk in the modern world and extinction in the geological past are often linked to aspects of life history or other facets of biology that are phylogenetically conserved within clades. These links can result in phylogenetic clustering of extinction, a measurement comparable across different clades and time periods that can be made in the absence of detailed trait data. This phylogenetic approach is particularly suitable for vertebrate taxa, which often have fragmentary fossil records, but robust, cladistically‐inferred trees. Here we use simulations to investigate the adequacy of measures of phylogenetic clustering of extinction when applied to phylogenies of fossil taxa while assuming a Brownian motion model of trait evolution. We characterize expected biases under a variety of evolutionary and analytical scenarios. Recovery of accurate estimates of extinction clustering depends heavily on the sampling rate, and results can be highly variable across topologies. Clustering is often underestimated at low sampling rates, whereas at high sampling rates it is always overestimated. Sampling rate dictates which cladogram timescaling method will produce the most accurate results, as well as how much of a bias ancestor–descendant pairs introduce. We illustrate this approach by applying two phylogenetic metrics of extinction clustering (Fritz and Purvis's D and Moran's I) to three tetrapod clades across an interval including the Permo‐Triassic mass extinction event. These groups consistently show phylogenetic clustering of extinction, unrelated to change in other quantitative metrics such as taxonomic diversity or extinction intensity.     

Patterns of evolution and extinction in proetid trilobites during the late Devonian mass extinction event,Canning Basin,Western Australia

Abstract: In the early Late Devonian, terminal Frasnian proetid trilobites have previously only been known from Europe and North Africa. For the first time, a rich fauna of late Frasnian proetids is described from the Virgin Hills Formation, Canning Basin, Western Australia. Seventeen species in six genera are described, of which three are new: Rudybole gen. nov., Palpebralina gen. nov. and Canningbole gen. nov. A new subgenus, Chlupaciparia (Australoparia) subgen. nov. is also described. Fourteen of the species are new: Palpebralia initialis sp. nov., P. pustulata sp. nov., ?P. sp. nov. A, Rudybole depressa sp. nov., Palpebralina pseudopalpebralis sp. nov. (comprising the subspecies P. pseudopalpebralis pseudopalpebralis subsp. nov. and P. pseudopalpebralis ultima subsp. nov.), P. minor sp. nov., P. ocellifer sp. nov., Canningbole latimargo sp. nov., C. henwoodorum sp. nov., C. macromma sp. nov., Pteroparia extrema sp. nov., Chlupaciparia (Chlupaciparia) planiops sp. nov., Chlupaciparia (Australoparia) australis sp. nov. and C. (Australoparia) lata sp. nov. The subspecies Rudybole adorfensis angusta subsp. nov. is also described. The proetids range through conodont Zones 11–13b and terminate at the Upper Kellwasser Event, which marks the terminal Frasnian mass extinction event. Three of the six proetid lineages, Palpebralia, Canningbole and Pteroparia, show evolutionary trends of eye reduction. Two of the remaining lineages, Rudybole and Palpebralina, consist exclusively of blind taxa. The last, Chlupaciparia, also comprises forms with reduced eyes. The proetids show a stepped pattern of extinction during the late Frasnian, which correlate with two Kellwasser biocrises documented in European/North African Frasnian sections. The highest diversity preceded the Lower Kellwasser event that occurred at the end of conodont Zone 12 and saw the extinction of all species present in that zone. However, only one genus, Pteroparia, locally became extinct. A major higher‐level taxonomic mass extinction at the top of Zone 13b initiated the Upper Kellwasser extinction event. This included extinction at the generic level, with all five remaining genera becoming extinct, and at the family level, with the loss of the Tropidocoryphidae.     

Nature and timing of biotic recovery in Antarctic benthic marine ecosystems following the Cretaceous–Palaeogene mass extinction

Taxonomic and ecological recovery from the Cretaceous–Palaeogene (K–Pg) mass extinction 66 million years ago shaped the composition and structure of modern ecosystems. The timing and nature of recovery has been linked to many factors including palaeolatitude, geographical range, the ecology of survivors, incumbency and palaeoenvironmental setting. Using a temporally constrained fossil dataset from one of the most expanded K–Pg successions in the world, integrated with palaeoenvironmental information, we provide the most detailed examination of the patterns and timing of recovery from the K–Pg mass extinction event in the high southern latitudes of Antarctica. The timing of biotic recovery was influenced by global stabilization of the wider Earth system following severe environmental perturbations, apparently regardless of latitude or local environment. Extinction intensity and ecological change were decoupled, with community scale ecological change less distinct compared to other locations, even if the taxonomic severity of the extinction was the same as at lower latitudes. This is consistent with a degree of geographical heterogeneity in the recovery from the K–Pg mass extinction. Recovery in Antarctica was influenced by local factors (such as water depth changes, local volcanism, and possibly incumbency and pre‐adaptation to seasonality of the local benthic molluscan population), and also showed global signals, for example the radiation of the Neogastropoda within the first million years of the Danian, and a shift in dominance between bivalves and gastropods.     

Survival patterns of macrobenthic marine assemblages during the end-Permian mass extinction in the western Tethys (Dolomites, Italy)

The ecological competition between brachiopods and bivalves is analysed by means of a quantitative palaeoecologic method applied on four assemblages located within a short stratigraphic interval, approximately 2 m thick, in the lower Tesero Member of the Werfen Formation (in the Southern Alps). The assemblages originate from the Tesero, Bulla and Sass de Putia sections. The analysed stratigraphic interval, uppermost Changhsingian in age, is located between the early and heaviest phase of the end-Permian mass extinction, which occurred across the Bellerophon/Werfen formational boundary (Event Boundary), and the Permian/Triassic boundary (Chronological Boundary), when nearly all the Permian stenotopic holdovers disappeared.These assemblages are characterised by small sized skeletons (“Lilliput effect”), which represent an adaptive survival strategy in stressed and harsh habitats resulting from the climatic and palaeoceanographic changes connected with the mass extinction. The Tesero assemblages are dominated by rhynchonelliform brachiopod Orbicoelia (bed CNT10) or Streptorhynchus (bed CNT11A), which were mostly attached at the top of shallow microbialitic mounds. These assemblages are again dominated by Permian stenotopic taxa and show a Palaeozoic structure. The Tesero habitat, which again permitted the survival of brachiopods, represented one of the last refuges in the western Tethys. On the contrary, the Bulla (BU9-10) and Sass de Putia (wPK13A) assemblages are bivalve-dominated, and thus show an ecologic structure typical of Early Triassic post-extinction marine benthic communities or Palaeozoic stressed marine communities. The bivalve-dominated assemblages proliferated in prevailing muddy siliciclastic substrates, with brief episodes of microbial algal growth. The most important environmental limiting factors and leading causes of end-Permian mass extinction are discussed in terms of palaeoautecologic and palaeosynecologic analysis.The different taxonomic composition and ecologic structure of the assemblages is related to palaeogeography, including water depth and connections with the open sea. The brachiopod-dominated assemblage, exclusive of the Tesero section, proliferated in microbial carbonate habitats in near-shore environments. The bivalve-dominated assemblages, which were more widespread than the brachiopod assemblages in the Dolomites and also occurred in other western Tethys localities, occur in more open and deeper marine environments. In the western Tethys margins, the local distribution of mixed faunas suggests that the extinction of Permian stenotopic taxa was caused by the onset of poisonous water on the shelves originating from deep marine environments.This extinction pattern appears to be a regional phenomenon and does not seem be applicable on a global scale. The extinction events were controlled by a complex network of interactive factors and the survival of faunal elements was probably stochastic.