Delayed recovery of non-marine tetrapods after the end-Permian mass extinction tracks global carbon cycle

During the end-Permian mass extinction, marine ecosystems suffered a major drop in diversity, which was maintained throughout the Early Triassic until delayed recovery during the Middle Triassic. This depressed diversity in the Early Triassic correlates with multiple major perturbations to the global carbon cycle, interpreted as either intrinsic ecosystem or external palaeoenvironmental effects. In contrast, the terrestrial record of extinction and recovery is less clear; the effects and magnitude of the end-Permian extinction on non-marine vertebrates are particularly controversial. We use specimen-level data from southern Africa and Russia to investigate the palaeodiversity dynamics of non-marine tetrapods across the Permo-Triassic boundary by analysing sample-standardized generic richness, evenness and relative abundance. In addition, we investigate the potential effects of sampling, geological and taxonomic biases on these data. Our analyses demonstrate that non-marine tetrapods were severely affected by the end-Permian mass extinction, and that these assemblages did not begin to recover until the Middle Triassic. These data are congruent with those from land plants and marine invertebrates. Furthermore, they are consistent with the idea that unstable low-diversity post-extinction ecosystems were subject to boom-bust cycles, reflected in multiple Early Triassic perturbations of the carbon cycle.     

Trace fossil evidence for restoration of marine ecosystems following the end-Permian mass extinction in the Lower Yangtze region,South China

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.     

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.     

A New Genus of Rhynchonellid Brachiopod from the Lower Triassic of South China and Implications for Timing the Recovery of Brachiopoda After the End-Permian Mass Extinction

A new genus, Meishanorhynchia , is proposed based on new material from the Lower Triassic of the Meishan section, South China. It is of a late Griesbachian age based on both associated biozones (ammonoids and bivalves) and radiometric dates of the intercalated volcanic ash beds. Comparison with both Palaeozoic and Mesozoic–Cenozoic-related genera suggests that it may represent the first radiation of progenitor brachiopods in the aftermath of the end-Permian extinction. The lowest brachiopod horizon that contains the genus is estimated to be about 250.1 ± 0.3 Ma. This implies that the initial stage of recovery of Brachiopoda in the Early Triassic was probably about 1.3 ± 0.3 myr after the major pulse of the end-Permian mass extinction (dated as 251.4 ± 0.3 Ma). This is in agreement with Hallam's expectancy that biotic recovery typically begins within one million years or so of major mass extinctions, in contrast to current views on the end-Permian extinction event which propose that the recovery of most if not all biotic groups in the Early Triassic was severely delayed and only began about five million years after the end-Permian extinction.     

Survival and recovery of calcareous foraminifera pursuant to the end-Permian mass extinction

Ninety-one percent of calcareous foraminiferal genera became extinct during the end-Permian mass extinction. The Early Triassic Epoch was a survival phase characterized by a short-lived proliferation of disaster forms and then a prolonged interval of low diversity. The orders Miliolida and Lagenida experienced limited taxonomic re-diversification in Early and Late Anisian time, respectively. All fusulinoidean fusulinides became extinct in Late Permian time, and only two non-fusulinoidean genera persisted into the Early Triassic. Triassic fusulinides diversified to just five genera before the order became entirely extinct in Late Triassic time. Involutinides originated in Olenekian time from an unknown ancestor. They did not significantly diversify until Late Triassic time. To cite this article: J.R. Groves, D. Altiner, C. R. Palevol 4 (2005).     

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.     

Deciphering the exceptional preservation of the Early Triassic Paris Biota (Bear Lake County,Idaho, USA)

After the end-Permian mass extinction, the Early Triassic (∼251.9 to 247 Ma) is characterized by several biotic crises that particularly affected marine faunas; accordingly, marine ecosystems from this unstable interval have been often described as heavily depauperate. This assumption, however, may relate to a biased fossil record. The discovery of taphonomic windows, like Konservat-Lagerstätten, in the Early Triassic would help to better understand the composition and diversity of ecosystems at that time. The Paris Biota (Idaho, USA) is a highly diverse fossil assemblage from the earliest Spathian (early late Olenekian, ∼250.6 Ma), indicating a rapid rediversification for many groups after the end-Permian crisis and pointing toward a remarkably complex marine ecosystem ∼1.3 m.y. after the Permian-Triassic boundary. However, its detailed taphonomy has not yet been investigated. Here we present the mineral characterization of four of its most abundant taxa: discinoid and linguloid brachiopods, leptomitid sponges, and caridean shrimps. For this purpose, we combined data from Raman microspectroscopy, Fourier Transform InfraRed spectroscopy, and SEM-EDXS. Although all taxa were preserved in calcium phosphate, the morphology, structuring and size of crystals are highly dissimilar at a nano- to micrometric scale. In brachiopods, the ultrastructure of calcium phosphate shows unorganized bacillary-like crystals, while in crustaceans their size is considerably smaller and round-shaped. Similar small crystals are observed in sponges. However, the ultrastructure of calcium phosphate in sponges exhibits a well-defined preferential orientation. In addition, sponges show some compressed but preserved three-dimensional features, with an inner surface better preserved. Such analyses are essential to understand the taphonomic pathways enabling exceptional preservation. The further comprehension of preservation features would help to understand potential bias on observed diversity signals and their interpretation.     

Additional thylacocephalans (Arthropoda) from the Lower Triassic (upper Olenekian) Osawa Formation of the South Kitakami Belt,Northeast Japan

The Lower Triassic Osawa Formation in the South Kitakami Belt, Northeast Japan, consisting mostly of mudstone of shallow-marine environment, was deposited during the late Olenekian (ca. 250 Ma), and is an important unit through which to examine the biotic recovery process after the end-Permian mass extinction. The Osawa Formation is the only unit in Japan that yields thylacocephalans (Arthropoda). Three species belonging to three genera have been reported before: Ankitokazocaris bandoi, Kitakamicaris utatsuensis and Ostenocaris sp. In addition to the known species, some thylacocephalans, including one new genus and three new species, are described in the present paper: Ankitokazocaris tatensis n. sp., Concavicaris parva n. sp., Miyagicaris costata n. gen. n. sp. and Ostenocaris? sp. Although Thylacocephala have a rather long stratigraphic range (from Silurian to Cretaceous) and are known from a wide geographical region, there are only about thirty genera in this group. The Osawa thylacocephalan fauna comprises at least five genera, making it one of the most diverse in the world at the generic level. During the Triassic Period, the Thylacocephala diversified and spread widely throughout low-latitude regions.     

The earliest post-paleozoic freshwater bivalves preserved in coprolites from the karoo basin, South Africa

Background

Several clades of bivalve molluscs have invaded freshwaters at various times throughout Phanerozoic history. The most successful freshwater clade in the modern world is the Unionoida. Unionoids arose in the Triassic Period, sometime after the major extinction event at the End-Permian boundary and are now widely distributed across all continents except Antarctica. Until now, no freshwater bivalves of any kind were known to exist in the Early Triassic.

Principal Findings

Here we report on a faunule of two small freshwater bivalve species preserved in vertebrate coprolites from the Olenekian (Lower Triassic) of the Burgersdorp Formation of the Karoo Basin, South Africa. Positive identification of these bivalves is not possible due to the limited material. Nevertheless they do show similarities with Unionoida although they fall below the size range of extant unionoids. Phylogenetic analysis is not possible with such limited material and consequently the assignment remains somewhat speculative.

Conclusions

Bivalve molluscs re-invaded freshwaters soon after the End-Permian extinction event, during the earliest part of the recovery phase during the Olenekian Stage of the Early Triassic. If the specimens do represent unionoids then these Early Triassic examples may be an example of the Lilliput effect. Since the oldest incontrovertible freshwater unionoids are also from sub-Saharan Africa, it is possible that this subcontinent hosted the initial freshwater radiation of the Unionoida. This find also demonstrates the importance of coprolites as microenvironments of exceptional preservation that contain fossils of organisms that would otherwise have left no trace.     

High organic carbon content and a decrease in radiolarians at the end of the Permian in a newly discovered continuous pelagic section: A coincidence?

The greatest mass extinction occurred at the end of the Permian. Most records of the mass extinction are not from pelagic sediments, but from shallow-marine and terrestrial sediments. Although several pelagic sections that span the end-Permian mass extinction have been found, these sections contain few index fossils and are often discontinuous because of small faults. We found the index fossils Albaillella cf. triangularis (Radiolaria) in siliceous claystone beds, Hindeodus parvus (Conodont) in the overlying black claystone beds, and Neospathodus cf. cristagalli and Ns. waageni (Conodont) in the subsequent siliceous claystone beds in Akkamori section-2 in northern Japan. These fossils suggest that this section ranges from the late Permian to the Early Triassic, including the early Induan and Olenekian stages. Furthermore, the lithological changes in the section, i.e., starting from bedded chert through siliceous claystone and black claystone to siliceous claystone, are concordant with those of well-known Permian–Triassic pelagic sequences in Japan. There is no gap between each lithofacie of the Akkamori section-2. Critical lithological continuity between Upper Permian siliceous claystone beds and uppermost Permian to lowermost Triassic black claystone beds of the Akkamori section-2 was recognized by observing hand-polished specimens and thin sections. Such paleontological and sedimentological evidence implies that the Akkamori section-2 is a continuous pelagic section that records the end-Permian mass extinction event. The carbonaceous black claystone beds have high total organic carbon (TOC) concentrations (1.06–3.31 wt.%), suggesting oceanic anoxia at least deep and probably stable primary productivity. A decrease in radiolarian abundance from 26–563 to 0.27–20 specimens/cm² coincided with an increase in TOC content from 0.01–0.16 to 1.06–3.31 wt.% at the boundary of the siliceous claystone and the overlying black claystone beds near the top of the Permian, implying that radiolarian extinction occurred at the end of the Permian coinciding with oceanic anoxia. Although TOC contents decreased in the early Olenekian (Smithian), radiolarian abundance did not increase at that time, indicating that radiolarian recovery was delayed by > 1.5 m.y.     

中国三叠纪海生爬行动物化石研究的回顾与进展

中国三叠纪海生爬行动物化石研究始于20世纪50年代,近10年来取得了重要的进展。此类化石在华南分布广泛,已见于多个省区的十余处地点,涉及自下三叠统奥伦尼克阶至上三叠统诺利阶的至少7个层位,并显示出由东向西产出层位逐渐升高的趋势。我国的三叠纪海生爬行动物化石门类齐全,属种丰富,已知类群包括鱼龙类、海龙类、檐齿龙类、始鳍龙类、原龙类、初龙类和湖北鳄类,显示出典型的西特提斯动物群特征,同时也体现了与东太平洋动物群的某些联系,以及一定的地方色彩。这些化石为研究三叠纪海生爬行动物各个类群的起源、演化和绝灭以及海洋环境的变迁提供了新的材料。     

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.     

A capitosauroid from the Lower Triassic of South America (Sanga do Cabral Supersequence: Paraná Basin), its phylogenetic relationships and biostratigraphic implications

Surviving through the end-Permian mass extinction, stereospondyls reemerged reaching a widespread distribution during the Early Triassic. A well representative lineage of this clade, Capitosauroidea, became worldwide abundant from Early to Late Triassic, in which their first undoubtful representatives were recovered from Lower Triassic deposits. Here, we describe a new capitosauroid, Tomeia witecki gen. et sp. nov., from the Sanga do Cabral Supersequence (Paraná Basin). Although relatively incomplete, the material presents a particular combination of early and late-diverging capitosauroid characters. Supporting Tomeia witecki as a new capitosauroid, our phylogenetic analysis placed the taxon as part of a clade that comprises only Early Triassic capitosauroids, specifically as the sister-taxon of the madagascarian Edingerella madagascariensis and close to Watsonisuchus spp., from Australia, South Africa and Madagascar. The status of Tomeia witecki as a new capitosauroid from western Gondwana supports a continuous record of the Stereospondyl lineage, since their first appearance during the Middle Permian in this supercontinent. Additionally, the temporal range of the Sanga do Cabral Supersequence during the Early Triassic was specifically reinforced as Olenekian, mainly based on the overall faunal content previously reported to this unit, associated with the known temporal distribution from those taxa phylogenetically closer to Tomeia witecki.

http://zoobank.org/urn:lsid:zoobank.org:pub:7AA9F874-BED5-4A99-B098-B50656E706C1     

安徽巢湖地区早三叠世菊石序列

由于地处低纬度下扬子碳酸盐缓坡台地较深水区 ,安徽巢湖地区的下三叠统由一系列泥、灰岩旋回层组成 ,并含有丰富的菊石化石。通过对巢北平顶山和马家山一带上二叠统顶部和下三叠统多条剖面的系统研究 ,建立了在华南乃至低纬度特提斯地区具有代表性的、以属级分类单位为基础的下三叠统菊石生物地层序列 (从下而上 ) :Ophiceras Lytophiceras带、Gyronites Prionolobus带、Flemingites Euflemingites带、Anasibirites带、Columbites Tirolites带和Subcolumbites带。在提议的全球印度阶 奥伦尼克阶界线层型候选剖面———巢湖平顶山西坡剖面上 ,Flemingites Euflemingites带之底与牙形石Neospathoduswaageni首现点比较接近 ,可以作为本界线主要辅助标志之一。记述采自该区上二叠统顶部及下三叠统 ,尤其是印度阶 奥伦尼克阶界线附近的一些具有年代地层意义的菊石化石资料 。     

New beetles (Insecta,Coleoptera) from the Lower Triassic of European Russia

Fossil remains of beetles are described from two Lower Triassic localities: Entala (Induan) and Tikhvinskoe (Olenekian). Only one beetle fossil was previously known from the Lower Triassic of Tikhvinskoe. The fossils are rather few and poorly preserved, but they are worth describing as finds rare for the Lower Triassic. Five fossils from Entala most probably belong to beetles of the same species of the formal genus Pseudochrysomelites. Beetles of this genus are especially abundant in deposits close to the Permian–Triassic boundary and can be considered “disaster taxa.” There are no known cases, either in the Permian or in the Middle–Upper Triassic, of a random sample of five specimens belonging to a single species. This suggests that in the Entala oryctocenosis the species diversity of beetles is extremely low. All three beetle fossils found in Tikhvinskoe belong to beetles of different species, showing that diversity had already started to increase. However, it remained low, and all fossils belong to the formal family Schizocoleidae, and two of the three belong to the same genus, Pseudochrysomelites. The Khei-Yaga locality, which immediately follows Tikhvinskoe in time (topmost Olenekian or early Anisian), already contains beetles of the families Asiocoleidae and Permosynidae. In the Lower Anisian of the Buntsandstein, such typical Mesozoic beetles as Cupedidae and Coptoclavidae have been recorded. The appearance of such advanced beetles as early as the Lower Anisian suggests that the famous Permian–Triassic crisis was not as deep as it is usually believed, and many beetles survived it, disappearing, however, from the fossil record in the Early Triassic.     

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.     

Chondrichthyan teeth from the Early Triassic Paris Biota (Bear Lake County,Idaho, USA)

A new, diverse and complex Early Triassic assemblage was recently discovered west of the town of Paris, Idaho (Bear Lake County), USA. This assemblage has been coined the Paris Biota. Dated earliest Spathian (i.e., early late Olenekian), the Paris Biota provides further evidence that the biotic recovery from the end-Permian mass extinction was well underway ca. 1.3 million years after the event. This assemblage includes mainly invertebrates, but also vertebrate remains such as ichthyoliths (isolated skeletal remains of fishes). Here we describe first fossils of Chondrichthyes (cartilaginous fishes) from the Paris Biota. The material is composed of isolated teeth (mostly grinding teeth) preserved on two slabs and representing two distinct taxa. Due to incomplete preservation and morphological differences to known taxa, the chondrichthyans from the Paris Biota are provisionally kept in open nomenclature, as Hybodontiformes gen. et sp. indet. A and Hybodontiformes gen. et sp. indet. B, respectively. The present study adds a new occurrence to the chondrichthyan fossil record of the marine Early Triassic western USA Basin, from where other isolated teeth (Omanoselache, other Hybodontiformes) as well as fin spines of Nemacanthus (Neoselachii) and Pyknotylacanthus (Ctenachanthoidea) and denticles have been described previously.     

A new holocrinid (Articulata) from the Paris Biota (Bear Lake County,Idaho, USA) highlights the high diversity of Early Triassic crinoids

After the end-Permian crisis and the extinction of their four Paleozoic subclasses, crinoids rapidly recovered. This group is classically believed to have radiated from a small surviving clade and to have diversified during the Middle and Upper Triassic from two lineages. Nevertheless, recent findings suggested that several lineages of crinoids had already diversified during the Early Triassic, and that their diversity has been overlooked. Here we describe a new form of holocrinid, Holocrinus nov. sp., from the earliest Spathian (Early Triassic) of southeastern Idaho (USA). So far, the exceptional completeness of sampled specimens, with skeletal elements of arms and stem in connection, is unique for the Early Triassic. They show that derived morphological features had already evolved ∼1.3 million years after the Permian–Triassic boundary, supporting the scenario of a rapid Early Triassic diversification of crinoids.     

New insights into the diversity dynamics of Triassic conodonts

In this paper, we examine the diversity trends and the evolutionary patterns of Triassic conodonts through a newly powered large-scale data-set compiled directly from the primary literature. Paleodiversity dynamics analyses have been undertaken by working at the species level and using a system of time units based on biozone subdivisions for a fine temporal level resolution. The role of heterogeneous duration of taxa in diversity estimates has been evaluated through the probabilistic profiles. Results reveal three different stages in the diversity behaviour of Triassic conodonts from standing metrics delimited by two inflections at the mid-Anisian and mid-Carnian. Survivorship analysis supports this pattern. Origination–extinction metrics report a diversification pattern characterised by important fluctuations during the Lopingian–Induan (earliest Triassic), the early-middle Olenekian (Early Triassic) and the Anisian–Ladinian transitions (Middle Triassic), as well as in the early Late Triassic. In addition, two clear diversification peaks are observed in the late Carnian and in the end-Norian. Reported patterns are interpreted in the context of deep extinction and environmental instability by documenting the biological signal of the main diversification and turnover patterns observed from such records. This study emphasises the singularity behaviour of diversity trends derived from the conodont record.     

Early Stages of Recovery of the East European Tetrapod Fauna after the End-Permian Crisis

The succession of tetrapod communities, which arose in Eastern Europe after the end-Permian mass extinction, displays three major evolutionary episodes designated the Tupilakosaurus, Benthosuchus-Wetlugasaurus, and Parotosuchus faunas. They are dated Induan, Early Olenekian, and Late Olenekian, respectively. Each fauna comprises two successive groupings, with some of these showing further subdivisions. In terms of its biochronological importance, the Late Induan tetrapod grouping is currently regarded as the most ambiguous. The lists of taxa that characterize the composition of the distinguished faunal units are provided.