the lilliput effect in the aftermath of the end permian extinction event early triassic animal body fossils and trace fossils are small relative to those in older and younger intervals size decreases sharply through the end permian extinction event and permian triassic boundary and the smallest sizes are encountered in the parvus and isarcica zones of the earliest induan animals appearing within these two zones are also exceedingly small compared to younger congenerics and conspecifics temporary dramatic size decrease of surviving taxa in the immediate aftermath of the extinction event is an example of the lilliput effect coined by urbanek a 1993 biotic crises in the history of upper silurian graptoloids a palaeobiological model historical biology 7 29 50 body size increases somewhat from the carinata zone mid induan but remains depressed for the duration of the early triassic and pre extinction sizes are not commonly recorded until at least the middle triassic marine and terrestrial faunas appear to be similarly affected the lilliput effect and longer term size reduction could be the result of several factors environmental parameters such as marine anoxia due to low atmospheric concentrations of oxygen at this time coupled with sluggish ocean circulation in a greenhouse world and food shortage are the likely proximal causes for the early triassic lilliput effect no single cause can explain all the observations and a combination of factors are likely to be involved

postapocalyptic greenhouse paleoclimate revealed by earliest triassic paleosols in the sydney basin australia the permian triassic boundary in the sydney basin of australia is coincident with a pronounced decrease in 13 c isotopic values of organic carbon the last coals anywhere in the world for all of the early triassic ca 6 m y time and extirpation of the glossopteris flora at the top of the illawarra and newcastle coal measures coal bearing paleosols of the latest permian represent extensive swamplands of the seasonally deciduous glossopteris flora in a humid cold temperate lowland southwest of an andean style volcanic arc stone rolls large ribs of floor rock up into the coal within some of the uppermost coals of the permian coal measures can be interpreted as string bogs of a kind now found in cold climates at latitudes of 68 70 which is compatible with a paleomagnetically estimated paleolatitude of 65 85 s for the sydney basin paleolatitude was not much different for earliest triassic time but paleosols of that age include inceptisols and entisols showing substantial chemical and textural weathering more like soils now forming at latitudes of 40 58 than those within polar circles this anomalous high latitude warmth set in at the permian triassic boundary sedimentation rates increased at the boundary marked by geochemically unusual acidification of clay and a dramatic carbon isotopic excursion these changes in environments and ecosystems can be explained by soil erosion following deforestation implied by the plant extinctions and abundant fungal remains at the permian triassic boundary evidence from paleosols can now be added to that from paleontological and isotopic studies showing that disruption of the carbon cycle at the permian triassic boundary resulted in a co 2 or ch 4 post apocalyptic greenhouse paleoclimate

end permian extinction and subsequent recovery of the ophiuroidea echinodermata a worldwide stratigraphical survey of late palaeozoic and early mesozoic ophiuroids echinodermata reveals unique extinction and recovery patterns during and after the end permian mass extinction event the ophiuroidea suffered the loss of all palaeozoic genera however this clade profited greatly from the event by undergoing a significant increase in species number in the postextinction oceans although both sampling and taphonomic biases may have affected ophiuroid diversity changeover patterns across the permian triassic p tr transition to some extent both biases cannot explain adequately the abrupt increase in species numbers and geographically wide distributions of ophiuroids in the early triassic oceans the ophiuroidea recovery follows erwin s model of a logistic increase in diversity beginning immediately after the end of the mass extinction without the survival interval morphological innovation i e emergence of advanced canal systems diversification of life habitats and biogeographical dispersal over the palaeotethyan and gondwanan regions enabled the ophiuroidea to occupy efficiently vacant ecospace created by the end permian extinction enabling the clade to recover and radiate in the early triassic oceans previously reported ophiuroids from the upper permian to lower triassic of china are also taxonomically revised these ophiaulaxoides bijieensis feng huangzhishania gulinensis feng ophiolepis shanxiensis yang and ophioderma schistovertebrata yang were described from the lower triassic while aganaster qingchangensis feng and syntomospina kaiyangensis feng came from the upper permian

δ13c evidence that high primary productivity delayed recovery from end permian mass extinction euxinia was widespread during and after the end permian mass extinction and is commonly cited as an explanation for delayed biotic recovery during early triassic time this anoxic sulfidic episode has been ascribed to both low and high productivity states in the marine water column leaving the causes of euxinia and the mechanisms underlying delayed recovery poorly understood here we use isotopic analysis to examine the changing chemical structure of the water column through the recovery interval and thereby better constrain paleoproductivity the 13c of limestones from 5 stratigraphic sections in south china displays a negative gradient of approximately 4 from shallow to deep water facies within the lower triassic this intense gradient declines within spathian and lowermost middle triassic strata coincident with accelerated biotic recovery and carbon cycle stabilization model simulations show that high nutrient levels and a vigorous biological pump are required to sustain such a large gradient in 13c indicating that early triassic ocean anoxia and delayed recovery of benthic animal ecosystems resulted from too much productivity rather than too little

multiple early triassic greenhouse crises impeded recovery from late permian mass extinction the late permian mass extinction was not only the most catastrophic known loss of biodiversity but was followed by unusually prolonged recovery through the early triassic opinion has been divided on whether delayed recovery was a legacy of especially profound ecological disruption or due to additional environmental perturbations new records from the sydney basin in southeastern australia now reveal five successive late permian and early triassic spikes of unusually high atmospheric co2 and profound chemical weathering these successive atmospheric co2 greenhouse crises coincided with unusually warm and wet paleoclimates for a paleolatitude of 61 s successive transient greenhouse crises punctuated long term cool dry and low co2 conditions and may account for the persistence of low diversity and small size in early triassic plants and animals

palaeoenvironments and faunal recovery after the end permian mass extinction abstract strata of the lower triassic werfen formation dolomites northern italy record deposition in a wide range of palaeoenvironments from peritidal to outer ramp settings this enables accurate assessment of faunal recovery following the end permian mass extinction by eliminating facies related effects extinction of the shelly taxa appears to be rapid and occurs in the latest permian tesero oolite horizon disappearance of the various trace fossil taxa appears to be more gradual following the extinction the lower and middle griesbachian are characterised by laminated sediments containing low diversity bedding plane assemblages ichnological geochemical and sedimentological data show evidence of widespread low oxygen conditions normally oxygenated waters return in the late griesbachian and there is a moderate increase in tiering above and below the surface macrofaunal and ichnofossil diversity decrease markedly during the smithian with the appearance of more terrigenous strata possibly reflecting an increased run off and more brackish conditions complete recovery begins in the basal spathian with an increase in tiering above and below the substrate crinoids and ammonoids appear for the first time in the werfen formation and the ichnogenus rhizocorallium re appears global correlation is hindered by poor biostratigraphy in the dolomites region however on a broad scale it appears as though many of the palaeoenvironmental changes are repeated worldwide current available data suggest that the re appearance of some trace fossil taxa e g diplocraterion rhizocorallium may be globally synchronous

anomalously diverse early triassic ichnofossil assemblages in northwest pangea a case for a shallow marine habitable zone early triassic trace fossil assemblages from the northwest margin of pangea record a diverse suite of postextinction infauna these ichnofossil assemblages occurred within well oxygenated shallow marine refuges in a panthalassa ocean otherwise characterized by widespread anoxia we propose an environmentally controlled model for their distribution in which wave aeration enhanced by frequent storms gave rise to an optimal zone for benthic colonization within this habitable zone extinction pressures were ameliorated and postextinction recovery duration was minimized

global coal gap between permian triassic extinction and middle triassic recovery of peat forming plants early triassic coals are unknown and middle triassic coals are rare and thin the early triassic coal gap began with extinction of peat forming plants at the end of the permian ca 250 ma with no coal known anywhere until middle triassic 243 ma permian levels of plant diversity and peat thickness were not recovered until late triassic 230 ma tectonic and climatic explanations for the coal gap fail because deposits of fluctuating sea levels and sedimentary facies and paleosols commonly found in coal bearing sequences are present also in early triassic rocks nor do we favor explanations involving evolutionary advances in the effectiveness of fungal decomposers insects or tetrapod herbivores which became cosmopolitan and much reduced in diversity across the permian triassic boundary instead we favor explanations involving extinction of peat forming plants at the permian triassic boundary followed by a hiatus of some 10 m y until newly evolved peat forming plants developed tolerance to the acidic dysaerobic conditions of wetlands this view is compatible not only with the paleobotanical record of extinction of swamp plants but also with indications of a terminal permian productivity crash from delta sup 13 c sub org and total organic carbon of both nonmarine and shallow marine shales 205 refs 3 figs more 3 tabs less

early triassic recovery of the brachiopod faunas from the end permian mass extinction a global review abstract brachiopod faunas in lower triassic deposits from spitzbergen primorye of russia japan mangyslak of kazakhstan alpine europe the caucasus the middle east the himalaya south china new zealand and western usa are taxonomically and stratigraphically reviewed they comprise survivors of the end permian extinction and taxa originating in the early triassic origination of the mesozoic type taxa represents the recovery of early triassic brachiopods the initial recovery of brachiopod faunas began in the late griesbachian and the time interval between the end of extinction and the onset of the recovery of brachiopods is much shorter than previously suspected the early triassic recovery of brachiopod faunas is characterized by widespread brachiopod dispersal multiprovincialism and the presence of rare lazarus genera at that time taxonomic selectivity of the recovery brachiopod faunas favors the rhynchonellids the re population of post extinction brachiopods varies geographically there is a preference for regions either previously barren of latest permian taxa or regions where the pre extinction changhsingian and surviving brachiopods are very rare this unique biogeographic selectivity is probably partly responsible for the impoverished nature of brachiopod faunas in the triassic even mesozoic oceans five intervals of extinction survival survival recovery recovery dispersal and radiation are recognized based on variations in brachiopod faunas from the latest permian to the early triassic a dramatic reduction in brachiopod diversity at the end permian mass extinction is followed by several stepwise declines in diversity in the survival interval the time during which the surviving brachiopods are dominated by geographically widespread generalist faunas that adapted to a wide variety of environments the survival interval was followed by a slow re population of new lineages dominated by progenitor taxa

u pb zircon geochronology and tempo of the end permian mass extinction the mass extinction at the end of the permian was the most profound in the history of life fundamental to understanding its cause is determining the tempo and duration of the extinction uranium lead zircon data from late permian and early triassic rocks from south china place the permian triassic boundary at 251 4 0 3 million years ago biostratigraphic controls from strata intercalated with ash beds below the boundary indicate that the changhsingian pulse of the end permian extinction corresponding to the disappearance of about 85 percent of marine species lasted less than 1 million years at meishan a negative excursion in 13 c at the boundary had a duration of 165 000 years or less suggesting a catastrophic addition of light carbon

transient metazoan reefs in the aftermath of the end permian mass extinction microbes were thought to be the dominant reef constructors following the end permian mass extinction sponge microbe reef deposits formed in the early triassic from the western united states suggest that instead metazoan reef building continued immediately following the extinction wherever marine conditions allowed

trace fossil evidence for restoration of marine ecosystems following the end permian mass extinction in the lower yangtze region south china abstract unlike the high abundance low diversity macrofaunas that characterize many early triassic benthic palaeocommunities ichnofossils were relatively common in the aftermath of the end permian mass extinction worldwide ichnofossils therefore are a good proxy for ecosystem recovery after the end permian biotic crisis this paper documents 14 ichnogenera and one problematic form from lower triassic successions exposed in the lower yangtze region south china post extinction ichnodiversity remained rather low throughout the griesbachian early smithian period and abruptly increased in the late smithian however several lines of evidence including extent of bioturbation burrow size trace fossil complexity and tiering levels indicate that diversification of ichnotaxa in the late smithian did not signal full marine ecosystem recovery from the permian triassic p tr mass extinction marine ichnocoenoses did not recover until the late spathian in south china the marginal sea provided hospitable habitats for tracemakers to proliferate in the aftermath of the end permian mass extinction

paleophysiology and end permian mass extinction physiological research aimed at understanding current global change provides a basis for evaluating selective survivorship associated with permo triassic mass extinction comparative physiology links paleontological and paleoenvironmental observations supporting the hypothesis that an end permian trigger most likely siberian trap volcanism touched off a set of physically linked perturbations that acted synergistically to disrupt the metabolisms of latest permian organisms global warming anoxia and toxic sulfide probably all contributed to end permian mass mortality but hypercapnia physiological effects of elevated pco2 best accounts for the selective survival of marine invertebrates paleophysiological perspectives further suggest that persistent or recurring hypercapnia global warmth also played a principal role in delayed triassic recovery more generally physiology provides an important way of paleobiological knowing in the age of earth system science 2007 elsevier b v all rights reserved

abundance anomalies of aromatic biomarkers in the permian triassic boundary section at meishan china evidence of end permian terrestrial ecosystem collapse several aromatic biomarkers detected in solvent extracts and kerogen pyrolysates from samples of the global stratotype section and point gssp for the permian triassic boundary at meishan china show distinctive relative abundance anomalies that may be related to the end permian biotic crisis on land enrichment of dibenzofuran homologues across the permian triassic boundary may corroborate the concept of excessive soil erosion following loss of rooted vegetation enrichment of alkylphenol entities occurs just below the main marine mass extinction horizon and prior to the characteristic end permian 13c decline the alkylphenols were probably derived from lignin and may provide a molecular record of dieback of woody vegetation highly decreased relative abundances of dibenzothiophenes could be indicative of excessive supply of fe2 it is suggested that fe2 was land derived and may have originated from soils in addition to the permian triassic boundary interval biomarker abundance anomalies also occur around the wuchiapingian changhsingian boundary in the basal part of the meishan section

sm nd systematics of hydrothermal scheelite from the mount charlotte mine kalgoorlie western australia an isotopic link between gold mineralization and komatiites

recovery tempo and pattern of marine ecosystems after the end permian mass extinction high resolution sampling of more than 10 000 microfossils from seven late permian middle triassic paleoequatorial sections in south china refutes claims for a 5 m y recovery delay after the end permian mass extinction we show that level bottom seafloor diversity began to recover in the early smithian little more than 1 m y after the mass extinction while recovery of reef building metazoans began 4 m y later in the anisian a further mass extinction in the late smithian identified in the pelagic fossil record is weakly manifest as a temporary pause in diversification among benthic communities in the early triassic of south china the offshore diversity increase began before then in shallower settings the recovery from the end permian mass extinction in south china was therefore significantly more rapid and environmentally more complex than hitherto known

the prelude of the end permian mass extinction predates a postulated bolide impact the mass extinction at the permian triassic boundary ptb is said to have been abrupt and probably caused by an extraterrestrial impact however evidence from the global stratotype section and point gssp of the base of the induan at meishan china shows that the biotic crisis began prior to the level in beds 25 and 26 at which the postulated impact event occurred evidence of such an earlier biotic crisis occurs in other sections in south china and in central and western tethyan regions this event is characterized by the extinction of a range of faunas including corals deep water radiolarians most fusulinids and pseudotirolitid ammonoids and many permian brachiopods in all sections this extinction level is usually a few decimeters to meters below that of the main mass extinction in the event beds 25 and 26 at meishan and their correlatives elsewhere this earlier extinction event happened before the postulated bolide impact at the level of beds 25 and 26 and constrains interpretation of the mechanisms that brought about this greatest mass extinction

radiolaria and pollen records from 0 to 50 ka at odp site 1233 continental and marine climate records from the southeast pacific abstract site 1233 drilled during leg 202 of the ocean drilling program provides a detailed record of marine and continental climate change in the southeast pacific and south american continent splits from over 500 samples taken at 20 cm intervals for quantitative analysis of radiolarian and pollen populations yield a temporal resolution of 200 400 years in each sample 39 pollen taxa and 40 radiolarian species and genera were evaluated age control is provided by 25 ams 14c dates lamy f kaiser j ninnemann u hebbeln d arz h w stoner j 2004 science 304 1959 1962 multivariate statistical analyses of these data allow us to conclude the following 1 during the past 50 ka the region of the central chile coast is not directly influenced by polar water from the antarctic region 2 changes in ocean conditions off central chile during this time interval primarily reflect north south shifts in the position of the south pacific transition zone 3 changes in chilean vegetation reflect comparable latitudinal shifts in precipitation and the position of the southern westerlies 4 the first canonical variate of radiolarian and pollen records extracted from site 1233 are remarkably similar to each other as well as to temperature records from the antarctic which suggests that marine and continental climate variability in the region is tightly coupled at periods longer than 3000 years 5 the phase coupling of these climate records which lead variations of continental erosion based on iron abundance at the same site are consistent with a hypothesis that erosion is linked to relatively long i e few thousand years response times of the patagonian ice sheet and thus is not a direct indicator of regional climate

conodont diversity and evolution through the latest permian and early triassic upheavals abstract a study of latest permian and early triassic multielement conodonts identifies eight steps in their evolutionary history during an interval when earth s biosphere was highly stressed these steps are 1 gradual decline of families and genera through the changhsingian up to the late griesbachian 2 conodont biofacies change and extinction of tethyan endemic species close to the permian triassic boundary ptb 3 faunal turnover with extinction and origination in the late griesbachian 4 initial radiation in gondolellids and diversification in apparatuses during the dienerian 5 explosive radiation in the early middle smithian 6 major extinction in the late smithian 7 major radiation early in the spathian 8 gradual turnover and decline in the late spathian through early anisian conodontophorids were reduced from five families at the beginning of the changhsingian to three by the ptb and to two late in the griesbachian from the changhsingian to the late griesbachian there was a single multielement apparatus represented in the gondolellidae whereas by the olenekian there were at least twelve given absolute age constraints the conodont recovery in the aftermath of the ptb was extraordinary conodont evolution during the dienerian remains obscure but was very significant in terms of changes in multielement apparatuses in spite of induan extinctions generic diversity generally increased from the ptb up to the late smithian in terms of species genera and multielement apparatuses the acme of triassic conodonts was in the middle smithian and the biggest extinction was in the late smithian after this extinction generic and apparatus diversity quickly returned to high levels marking a significant spathian recovery gradual decline characterized the late spathian and persisted into the early anisian a synthetic summary of the temporal distribution and proposed relationships amongst all known lower triassic conodonts provide an improved foundation for biostratigraphic and biochronologic studies extinction and radiation trends correlate well with sea level changes and sequence boundaries faunal turnovers correspond to lowstands in the late griesbachian late smithian and late spathian and radiations correspond to transgressions in the early griesbachian early smithian and early spathian the trends also correlate with perturbations in the carbon cycle with lower triassic 13c minima corresponding to extinction and faunal turnover and positive values generally occurring during radiations however middle triassic recovery of marine benthos follows a positive maxima and an overall decline in conodont diversity new taxa borinella chowadensis b megacuspa neogondolella griesbachensis and n mongeri are described

trace fossils and the aftermath of the permo triassic mass extinction evidence from northern italy abstract the duration of early triassic anoxia in the carbonate ramp settings of the werfen formation dolomites northern italy has been constrained using ichnofabric and sedimentological data this includes fine lamination and abundant pyrite framboids commonly reworked into lags by storm events the anoxic event spanned the entire griesbachian stage and extended from deep waters into extraordinarily shallow water settings evidence for oxygen deficiency can be found in mid and inner ramp facies the extent and duration of this event may at least partially explain the magnitude of the permo triassic mass extinction normal benthic conditions returned rapidly in the dienerian stage and pervasively bioturbated distal ramp facies testify to the ventilation of the deeper water areas of the region in post griesbachian times a post mass extinction radiation is not particularly well displayed in either trace fossil or body fossil diversity compilations from the dolomites moderate low levels of trace fossil diversity are maintained throughout the post griesbachian werfen formation whilst the body fossil data shows a moderate increase from the mid smithian stage there is thus a curious lag period spanning the dienerian to mid smithian when environmental conditions were apparently normal but the marine fauna failed to show any recovery

altered river morphology in south africa related to the permian triassic extinction the permian triassic transition in the karoo basin of south africa was characterized by a rapid and apparently basin wide change from meandering to braided river systems as evidenced by preserved sedimentary facies this radical changeover in river morphology is consistent with geomorphic consequences stemming from a rapid and major die off of rooted plant life in the basin evidence from correlative nonmarine strata elsewhere in the world containing fluvial permian triassic boundary sections suggests that a catastrophic terrestrial die off of vegetation was a global event producing a marked increase in sediment yield as well as contributing to the global 13 c excursion across the permian triassic boundary

gastropod evidence against the early triassic lilliput effect comment brayard et al 2010 1 assert that their study of late early triassic gastropods provides evidence against the post extinction lilliput effect regrettably their data provide no such evidence the lilliput effect as defined by urbanek 1993 2 is a quantifiable temporary decrease in the

the end permian mass extinction and its aftermath on an equatorial carbonate platform insights from ichnology terra nova 22 195 202 2010 abstract a semi quantitative analysis of the endobenthic activity across the permo triassic boundary ptb was carried out on a 445 m thick well core interval from the persian gulf by logging the degree of bioturbation trace fossil distribution and ichnodiversity id this analysis enables for the first time the evaluation of recovery of endobenthic organisms after the end permian mass extinction by direct comparison of post extinction lower triassic with pre extinction upper permian units the results indicate that the studied shallow marine carbonate platform within an equatorial setting acted as oxygenated refugee for the biotic recovery traces of suspension and deposit feeders reappear simultaneously shortly after the extinction event but are reduced in size macroborings so far unknown from the lower triassic occur where suitable firm or hard substrate was available recovery took place in two phases early induan 50 pre extinction id and 20 30 bioturbation and late induan 75 id and almost 90 bioturbation re established

new trace fossil evidence for an early recovery signal in the aftermath of the end permian mass extinction abstract new ichnological data from the lower triassic werfen formation dolomites italy revealed an unexpectedly diverse and complex ichnofauna in mixed siliclastic carbonate shelf sediments of the western tropical tethys shortly after the end permian mass extinction event common elements are thalassinoides palaeophycus and planolites other associated ichnogenera include spongeliomorpha rhizocorallium lockeia catenichnus helminthopsis and taenidium one structure is tentatively determined as curvolithus biostratigraphic data conodonts and bivalves of the trace fossil interval lower seis siusi member suggest a late griesbachian age less than 1 ma after the end permian mass extinction event similarly diverse ichnofossil communities from this time interval have recently been described from the boreal realm but this is the first record of a diverse griesbachian ichnoassemblage from the tropics indicating that the early recovery of trace fossil producers was not latitudinally restricted as previously proposed accordingly relatively advanced recovery stages were reached on a global scale much earlier than the spathian as is commonly acknowledged the early and interregional peak in ichnofaunal recovery implies subsequent ecological setbacks in post griesbachian times that explain the overall delay of benthic recovery until the spathian rather than persistent catastrophic conditions during the griesbachian a global and synchronous succession of crises and relaxation phases after the griesbachian is proposed as an explanatory model for the recovery pattern of benthic ecosystems

smithian spathian boundary event evidence for global climatic change in the wake of the end permian biotic crisis one of the most important carbon cycle perturbations following the end permian mass extinction event straddles the smithian spathian boundary ssb olenekian early triassic this anomaly is characterized by a prominent positive carbon isotope excursion known from tethyan marine rocks its global signifi cance is established here by a new high paleolatitude record spitsbergen paleontological evidence such as boreal palynological data barents sea norway and global patterns of ammonoid distribution indicates a synchronous major change in terrestrial and marine ecosystems near the ssb the reestablishment of highly diverse plant ecosystems including the rise of woody gymnosperms and decline of the formerly dominating lycopods is interpreted as an effect of a major climate change this hypothesis is supported by modeling of ammonoid paleobiogeography the distribution patterns of which are interpreted as a proxy for sea surface temperatures sst the latest smithian thus appears to have been a time of a warm and equable climate as expressed by an almost fl at pole to equator sst gradient in contrast the steep spathian sst gradient suggests latitudinally differentiated climatic conditions we propose that this drastic climate change and the global carbon cycle perturbation were triggered by a massive end smithian co2 injection the ssb event could therefore represent one of the causes for stepwise and delayed recovery of marine and terrestrial biotas in the wake of the end permian biotic crisis

terrestrial marine teleconnections in the collapse and rebuilding of early triassic marine ecosystems abstract the latest permian mass extinction lpe just prior to the permian triassic boundary at 252 ma resulted in the disappearance of 90 of skeletonized marine taxa and the replacement of the paleozoic fauna by the modern fauna in the immediate aftermath of the extinction shallow marine ecosystems were dominated by microbial communities and diminutive disaster taxa lilliput faunas and the post crisis period in general was characterized by high abundance low diversity marine communities occupying a reduced ecological space the recovery of marine ecosystems was a protracted process lasting throughout the 5 myr long early triassic some clades failed to recover and subsequently went extinct dead clades walking while others recovered only to suffer secondary crises during the early triassic profound environmental changes accompanied these biotic developments deep oceans experienced sustained hypoxia or anoxia for at least several million years following the boundary event anoxia was also widespread in shallow marine areas especially at the time of the lpe but it was more episodic and abated more rapidly than in the deep ocean making it an unlikely factor in the delayed recovery of early triassic marine ecosystems the development of anoxia was due at least in part to a sharp temperature rise following the lpe but may have been stimulated by changes in marine nutrient inventories and productivity rates as well these conditions may have been the result of a concurrent increase in subaerial weathering rates that was sustained into the early triassic in addition to increasing nutrient fluxes to marine areas rapid weathering caused a massive influx of sediment to shallow marine systems subjecting marine biotas to siltation stress this scenario implies close teleconnections between terrestrial and marine environments during the permian triassic boundary crisis with perturbations of the terrestrial realm contributing to the marine crisis

gastropod evidence against the early triassic lilliput effect reply size reduction in the aftermath of the permian triassic mass extinction event has repeatedly been described for various marine organisms including gastropods the lilliput effect a smithian gastropod assemblage from utah usa reveals numerous large sized specimens of different genera as high as 70 mm the largest ever reported from the early triassic other gastropods reported from serbia and italy are also as large as 35 mm size frequency distributions of the studied assemblages indicate that they were not unusually small when compared with later mesozoic and modern faunas the occurrence of large sized gastropods less than 2 ma after the permian triassic mass extinction refutes the lilliput hypothesis in this clade at least for the last 75 of the early triassic

pattern of marine mass extinction near the permian triassic boundary in south china the meishan section across the permian triassic boundary in south china is the most thoroughly investigated in the world a statistical analysis of the occurrences of 162 genera and 333 species confirms a sudden extinction event at 251 4 million years ago coincident with a dramatic depletion of 13ccarbonate and an increase in microspherules

organic matter and palaeoenvironmental signals during the early triassic biotic recovery the salt range and surghar range records latest permian to the middle triassic mixed siliciclastic carbonate shelf deposits of the northern gondwana margin have been studied in four sections nammal chhidru chitta landu and narmia in the salt range and surghar range of pakistan sedimentological and palynofacies patterns combined with a high resolution ammonoid based age control have been used to assess environmental changes such as sea level change distance from the shore and oxygenation conditions of the sections in the aftermath of the end permian mass extinction the base and the top of the early triassic are marked by second order sequence boundaries srt1 srt8 within the early triassic two third order sequence boundaries could be delineated by means of palynofacies analysis and sedimentology one near the dienerian smithian srt2 and the second one near the smithian spathian boundary srt5 the extinction event at the smithian spathian boundary seems to be closely associated to the latter globally recorded sea level low stand five additional sequences of undetermined order srt3 srt 4 srt5 1 srt6 and srt7 are reflected in the sedimentological record of the studied sections the observed changes in the composition of the particulate organic matter pom indicate a general shallowing upward trend which is modulated by smaller transgressive regressive cycles supporting the sedimentologically defined sequences the pom is mostly dominated by terrestrial phytoclasts and sporomorphs the strongest marine signal is reflected by increased abundance of amorphous organic matter aom in the lower part of the ceratite marls at nammal late dienerian and chhidru earliest smithian and the lower ceratite limestone at chitta landu late dienerian aom of marine origin is characteristic for deeper distal basinal settings and is preferentially preserved under dysoxic and anoxic conditions indicating reduced oxygen conditions during these intervals up section transgressive events are reflected by increased numbers of acritarchs reaching up to 50 of the pom well oxygenated conditions and low total organic carbon contents toc continue up to the top of the early triassic mianwali formation the most pronounced terrestrial influx is expressed in the middle triassic organic carbon isotope data parallel the carbonate carbon isotope records from the tethyan realm therefore they reflect real global changes in the carbon cycle independent of the om composition the biomarker study of the apolar hydrocarbons of three samples from the nammal section indicates an enhanced bacterial productivity especially in the smithian and spathian reflected in high relative abundances of hopanes pom toc data and redox sensitive biomarkers together with high resolution biostratigraphy demonstrate that well oxygenated environmental conditions prevailed in the early triassic with the exception of the dienerian to earliest smithian interval the pom assemblages of late permian to late griesbachian age indicate well oxygenated conditions during this time interval there is no evidence in support of an anoxic event in the late griesbachian in these sections

sulfur isotopic evidence for chemocline upward excursions during the end permian mass extinction the latest permian was a time of major change in ocean chemistry accompanying the greatest mass extinction of the phanerozoic to examine the nature of these changes samples from two well studied marine sections that span the permian triassic boundary have been analyzed the meishan and shangsi sections located in southern china isotopic analysis of the carbonate associated sulfate in these samples provides a detailed record of several isotopic shifts in 34scas approaching and across the ptb ranging from 30 to 15 vcdt with repeated asynchronous fluctuations at the two locations we interpret the patterns of isotopic shifts in conjunction with other data to indicate a shallow unstable chemocline overlying euxinic deep water which periodically upwelled into the photic zone these chemocline upward excursion events introduced sulfide to the photic zone stimulating a bloom of phototrophic sulfur oxidizing bacteria we hypothesize that elemental sulfur globules produced by these organisms and 34s depleted pyrite produced in the euxinic water column were deposited in the sediment later oxidation led to incorporation as cas this created the large changes to the 34scas observed in the latest permian at these locations

the timing and pattern of biotic recovery following the end permian mass extinction the aftermath of the great end permian period mass extinction 252 myr ago shows how life can recover from the loss of 90 species globally the crisis was triggered by a number of physical environmental shocks global warming acid rain ocean acidification and ocean anoxia and some of these were repeated over the next 5 6 myr ammonoids and some other groups diversified rapidly within 1 3 myr but extinctions continued through the early triassic period triassic ecosystems were rebuilt stepwise from low to high trophic levels through the early to middle triassic and a stable complex ecosystem did not re emerge until the beginning of the middle triassic 8 9 myr after the crisis a positive aspect of the recovery was the emergence of entirely new groups such as marine reptiles and decapod crustaceans as well as new tetrapods on land including eventually dinosaurs the stepwise recovery of life in the triassic could have been delayed either by biotic drivers complex multispecies interactions or physical perturbations or a combination of both this is an example of the wider debate about the relative roles of intrinsic and extrinsic drivers of large scale evolution from a much more devastated planet and biota than the others with only some 10 of species surviving the epme was much harsher than the other mass extinctions during which global species diversity reduced to only about 50 of the pre extinction total 1 2 24 26 this means that the triassic recovery has two profound implications first it may show qualitative as well as quantitative differences from the other post extinction recoveries and second it can act as an exemplar of what to expect at its most extreme when global biodiversity is pushed to the brink there are obvious implications for current concerns about biodiversity loss and recovery resulting from human impacts 27 28 in the past ten years attention has focused on the sedimentary successions in south china these are enormously laterally extensive with some formations extending more than 2 000 km from the zhejiang to yunnan provinces the huge exposures length of the sections and improving dating open up the opportunity to explore physical environmental and biotic changes through the extinction and recovery times in varied marine habitats and compare these with patterns elsewhere in the world fig 1 a fine scale forensic analysis of this extraordinary time in earth s history now becomes possible the end permian mass extinction the epme killed 80 96 of marine animal species and 70 of terrestrial vertebrate species

the origin and early radiation of dinosaurs abstract dinosaurs were remarkably successful during the mesozoic and one subgroup birds remain an important component of modern ecosystems although the extinction of non avian dinosaurs at the end of the cretaceous has been the subject of intense debate comparatively little attention has been given to the origin and early evolution of dinosaurs during the late triassic and early jurassic one of the most important evolutionary radiations in earth history our understanding of this keystone event has dramatically changed over the past 25 years thanks to an influx of new fossil discoveries reinterpretations of long ignored specimens and quantitative macroevolutionary analyses that synthesize anatomical and geological data here we provide an overview of the first 50 million years of dinosaur history with a focus on the large scale patterns that characterize the ascent of dinosaurs from a small almost marginal group of reptiles in the late triassic to the preeminent terrestrial vertebrates of the jurassic and cretaceous we provide both a biological and geological background for early dinosaur history dinosaurs are deeply nested among the archosaurian reptiles diagnosed by only a small number of characters and are subdivided into a number of major lineages the first unequivocal dinosaurs are known from the late carnian of south america but the presence of their sister group in the middle triassic implies that dinosaurs possibly originated much earlier the three major dinosaur lineages theropods sauropodomorphs and ornithischians are all known from the triassic when continents were joined into the supercontinent pangaea and global climates were hot and arid although many researchers have long suggested that dinosaurs outcompeted other reptile groups during the triassic we argue that the ascent of dinosaurs was more of a matter of contingency and opportunism dinosaurs were overshadowed in most late triassic ecosystems by crocodile line archosaurs and showed no signs of outcompeting their rivals instead the rise of dinosaurs was a two stage process as dinosaurs expanded in taxonomic diversity morphological disparity and absolute faunal abundance only after the extinction of most crocodile line reptiles and other groups

survival strategies of brachiopod faunas from the end permian mass extinction abstract global reviews reveal that 102 brachiopod species including 40 indeterminate species belonging to 43 genera within 27 families survived the end permian mass extinction in south china the himalayan regions southern tibet nepal salt range and kashmir southern alps italy arctic canada western usa and western australia of these the surviving brachiopods are most abundant and widespread in south china where they occur in 32 of the 42 permian triassic p t boundary sections around the world brachiopod faunas are the most abundant and diverse post extinction survivors and their appearance above the end permian mass extinction horizon somewhat masks recognition of the end permian extinction these surviving brachiopods occur in three stratal intervals corresponding to the mixed fauna beds mfb hereafter 1 3 of the p t boundary beds a sharp drop in diversity of the brachiopod faunas coincides with the widely accepted end permian mass extinction horizon the post extinction brachiopods were also affected by a subsequent crisis corresponding to the boundary between mfb 2 and mfb 3 so that most survivors were extinct approximately 0 7 ma after the end permian mass extinction the surviving brachiopods were mainly productida followed by spiriferida in particular small chonetids chonetid like productellids and smooth thin shelled ambocollid spiriferids are most abundant lingula is extraordinarily abundant and widespread approximately 0 7 ma after the greatest extinction when many regions became devoid of articulate brachiopods widespread broadly adapted and small sized taxa preferentially survived frequent intrageneric speciation of widespread widely adapted generalist genera enabled survival brachiopods to occupy rapidly and efficiently vacant ecospace in the aftermath of the end permian extinction the ecological habitats of the post extinction brachiopods are subdivided into nearshore epeiric sea restricted carbonate platform open carbonate platform ramp and outer shelf environments of these the survival brachiopods are most diverse in the open platform environments the nearshore zone provided ideal habitats for the nonarticulated brachiopods to occupy whereas the survivors in the outer shelf are predominantly chonetids and chonetid like productellids the most successful survivors the productid brachiopods are widely distributed in six types of ecological habitats and thus have no apparent preference for specific environments the end permian extinction resulted in the movement of biogeographical provinces towards the palaeo equator the brachiopods survived mostly in the palaeo equatorial provinces and along the southern margins of the palaeo tethys ocean