Conodonts from the Early Triassic microbialite of Guangxi (South China): implications for the definition of the base of the Triassic System

We describe a new Early Triassic (Griesbachian) succession of conodont faunas from a high‐resolution sampling of the basal Early Triassic microbial limestone and the base of the overlying unit at the Wuzhuan section (Nanpanjiang Basin, Guangxi, South China). The microbial limestone records the earliest phase of the Early Triassic biotic recovery after the end‐Permian mass extinction. For the first time, rich conodont faunas are reported from within the microbialite. The faunas from Wuzhuan are largely dominated by anchignathodontids, including several Isarcicella species, which were previously documented only from strata above the microbialite. A total of 14 conodont species assigned to three genera is recorded from the Wuzhuan section. Starting from the base of the microbialite upwards, several species are sequentially added to the conodont assemblage. The alpha diversity peaks at the top of the microbialite. The conodont record in the considered microbialite interval at Wuzhuan is presumably unaffected by local ecological changes. It therefore more likely represents an evolutionary rather than an ecological pattern. We compare the Wuzhuan's conodont record with a well‐supported phylogenetic model and suggest that the sequence of first occurrences at Wuzhuan is the closest to the ‘true’ sequence of evolutionary events that took place during this Griesbachian radiation of anchignathodontids. Based on comparisons with the GSSP section at Meishan, we suggest further that the first occurrence of Hindeodus parvus in Meishan does not correspond to its first appearance datum.     

Gondolelloid multielement conodont apparatus (Scythogondolella) from the Lower Triassic of Jiangsu,East China,revealed by high-resolution X-ray microtomography

The application of high-resolution X-ray microtomography on conodont natural assemblages has provided new information on the structure of the elements and enabled the three-dimensional reconstruction of apparatuses. We herein report four conodont natural assemblages from uppermost Lower Qinglong Formation, Longtan, Nanjing, East China. Using X-ray microtomography, we reconstructed the apparatus of the end-Smithian (Olenekian, Early Trassic) Scythogondolella milleri. Our result confirms that Scythogondolella has an octomembrate apparatus composed of 15 elements (a single S₀ element, two pairs of S_1–4, M and P_1–2 elements) like other gondolelloid apparatuses that have been tested by their corresponding natural assemblages, including Neogondolella, Novispathodus and Nicoraella. Element morphology of Scythogondolella closely resembles that of Neogondolella apparatus from the Illyrian (Anisian, Middle Triassic) of Monte San Giorgio: an alate (hibbardellan) S₀ element with two long lateral processes that meet at a denticle anterior of the cusp, a bipennate (hindeodellan) S₃ element with a bifurcated anterior process branching from the third denticle anterior of the cusp and an enantiognathiform S₂ element with two dissimilar processes of nearly equal in length. It differs from the latter in the length of the inner-lateral process of M element and the posterior process of S₀ and S_3–4 elements without considering the obvious morphological differences in P elements between them. The element positional homologues of Scythogondolella conforms to those of the standard 15-element plan shared primitively among ozarkodinin, prioniodinin and prioniodontid conodonts, and again confirms that the breviform digyrate elements of cypridodellan and enantiognathiform morphotypes occupy the S₁ and S₂ positions, respectively, within the superfamily Gondolelloidea.     

Phylogeny of the conodont genera Hindeodus and Isarcicella across the Permian–Triassic boundary

Jiang, H., Aldridge, R.J., Lai, X., Yan, C. & Sun, Y. 2010: Phylogeny of the conodont genera Hindeodus and Isarcicella across the Permian–Triassic boundary. Lethaia, Vol. 44, pp. 374–382. Several phylogenetic hypotheses have been put forward to explain the relationships between species of the conodont genera Hindeodus and Isarcicella across the Permian–Triassic boundary. These phylogenetic models can be tested by cladistic analysis. A data matrix of 25 P₁ element characters for 22 species of Hindeodus and Isarcicella has been analysed and the resulting cladogram converted to a phylogenetic tree using the ranges of taxa in the Meishan Section, China. The proposed phylogeny does not concur entirely with any of those forwarded in the previous literature, and Hindeodus parvus is not resolved as a direct descendant of Hindeodus praeparvus. Species assigned to the genus Isarcicella form a natural group, with Hindeodus species apparently representing a paraphyletic array. The Hindeodus + Isarcicella clade survived the end‐Permian extinction event well, with Hindeodus species that became extinct just above the boundary replaced by radiation within the Isarcicella clade. □Cladistics, conodonts, phylogeny, Permian, Triassic.     

RECONSTRUCTION OF AN APPARATUS OF NEOSTRACHANOGNATHUS TAHOENSIS FROM ORITATE,JAPAN AND SPECIES OF NEOSTRACHANOGNATHUS FROM OMAN

Abstract: The apparatus of an Early Triassic conodont Neostrachanognathus tahoensis Koike, 1998 from Oritate, Kumamoto Prefecture, Japan, and a species of Neostrachanognathus from Oman were reconstructed. On the basis of five natural assemblages from the Oritate area, the three‐dimensional apparatus model of N. tahoensis is interpreted as bilaterally symmetrical and composed of 14 elements consisting of pairs of P₁, P₂, P₃, S₁, S₂, S₃, and S₄ elements. The P₁ and P₂ elements are coniform elements, the P₃ elements are digyrate forms, and the S elements are bipennate ramiforms. The S elements are arranged rostrally in the apparatus and the pairs of the P₁, P₂, and P₃ elements are subvertically arranged caudally and ventrally to the S array. One of the natural assemblages was formed by rostrocaudal collapse of the apparatus on the sea floor, whereas the other assemblages indicate that conodont animals came to rest nearly parallel with the substrate prior to burial. A collection of isolated elements from Jabal Safra, Oman, includes a second species of Neostrachanognathus with a comparable apparatus.     

Cladistic tests of monophyly and relationships of biostratigraphically significant conodonts using multielement skeletal data –Lochriea homopunctatus and the genus Lochriea

Abstract: Since the 1960s, huge progress has been made in reconstructing the multielement skeletons of conodont species and developing a biologically defensible taxonomy. Nevertheless, a widespread prejudice remains that certain parts of the conodont skeleton, particularly the P₁ elements, are more informative than others with regard to taxonomy and evolutionary relationships. Here, we test these views. A new partial multielement reconstruction of the skeleton of the biostratigraphically significant conodont originally described as Gnathodus commutatus homopunctatus allows us to conduct a cladistic test of the alternative hypotheses of phylogenetic placement of this species. Our analysis also provides the first test of the hypothesis that Lochriea– species of which are markers for global correlation – is monophyletic and tests hypotheses concerning the origins of the genus. Our results demonstrate that homopunctatus is a species of Lochriea and that the genus is monophyletic. The widely held view that Lochriea arose from a species of Bispathodus is not supported. Our results show that it is difficult to predict a priori which parts of the conodont skeleton carry phylogenetic signal, and provide strong support for the hypothesis that similarity in the morphology of conodont P₁ elements alone is not a reliable guide to relationships and taxonomic groupings of conodont species. This is because P₁ elements with similar morphologies are convergently acquired in multiple conodont clades, because reliance on the characters of only one of the six or seven morphologically distinct elements of the conodont skeleton ignores phylogenetically significant data and because P₁ elements can lack characters that might seem to be diagnostic of a genus. Conodonts are no different to other organisms: ignoring data that have the potential to be phylogenetically informative is unlikely to produce the most reliable hypotheses of evolutionary relationships. We suggest that other biostratigraphically significant hypotheses of relationship between conodont taxa that are based on P₁ elements alone should be subject to cladistic testing.     

Conodont biostratigraphy of the Early Triassic in eastern Slovenia

The Early Triassic is a critical interval for the study of recovery from the terminal Permian mass extinction, as there are small-scale extinction events, which may have contributed to the delayed recovery. The systematic measuring and sampling of a 12-m-thick section at the Mokrice locality in eastern Slovenia has resulted in the recovery of a conodont fauna from the Olenekian beds. Four conodont zones have been recognized. These zones are in ascending order as follows: the Hadrodontina aequabilis Zone, Platyvillosus corniger Zone, Platyvillosus regularis Zone, and Triassospathodus hungaricus Zone. These conodont zones confirm the proposed conodont biozonation sequence in western Slovenia and have correlation value especially for the western marginal Tethys. Multielement conodont apparatuses of Triassospathodus hungaricus and Platyvillosus regularis have been reconstructed based on conodont elements that were recently obtained from the Slovenian sections. Although the S₂element was not found, the apparatus indicates that the conodont species “Spathognathodus” hungaricus should be assigned to the genus Triassospathodus.     

Testing hypotheses of element loss and instability in the apparatus composition of complex conodonts: articulated skeletons of Hindeodus

Knowledge of the conodont skeleton, in terms of the morphology of the elements and the positions they occupy, provides the foundation for understanding of homology, taxonomy and evolutionary relationships in conodonts. This knowledge also underpins analyses of conodont functional morphology and feeding. Direct evidence of skeletal anatomy and apparatus architecture comes from natural assemblages: fossils that preserve together the articulated remains of the conodont apparatus, either collapsed onto a bedding plane or as clusters of elements in which juxtaposed and overlapping elements have been fused together by diagenetic minerals. Here we describe six clusters of the biostratigraphically important conodont Hindeodus parvus from the Lower Triassic Shangsi section, Sichuan Province, South China. Five of these clusters represent the partial remains of articulated skeletons, providing direct evidence of the number and arrangement of elements in the apparatus. Combined with data from previously published natural assemblages this provides a test of the hypothesis that Triassic conodonts had a reduced dentition. Hindeodus parvus possessed a complete raptorial array of two M and nine S elements (unpaired S₀; symmetrically paired S₁, S₂, S₃, S₄); the paired P₁ locations were occupied by carminiscaphate elements, but the apparatus lacked P₂ elements. This is consistent with broader evidence for a particularly high degree of integration and constraint operating on the S–M array of morphologically complex conodonts, leading to conserved architecture of the array over a period of more than 250 million years. The loss of elements from the P domain implies a change in food processing ability and, given the predominance of data from P elements in conodont taxonomy and biostratigraphy, the hypothesis of element loss from the P domain has significant implications for the broader understanding of conodont diversity and evolutionary patterns.     

AnOistodus venustus-like conodont species from the Middle Ordovician of Baltoscandia

Ordovician conodont specimens resemblingOistodus venustus Stauffer, 1935 have been reported from many areas. There is increasing evidence, however, that several lineages with homeomorphic conodont elements have erroneously been referred to one and the same species. I have investigated Baltoscandian conodont elements of this kind in order to find out about their origins and phylogenetic relationships with morphologically comparable elements from other areas. A natural grouping of finds from the Middle and lower Upper Ordovician of Baltoscandia is here described as belonging to a new species of a new genus,Venoistodus balticus n. gen., n. sp. The new species probably evolved paedomorphically fromDrepanoistodus forceps (Lindström, 1955) in the Early Ordovician.     

New data from Oman indicate benthic high biomass productivity coupled with low taxonomic diversity in the aftermath of the Permian–Triassic Boundary mass extinction

A new Early Triassic marine fauna is described from an exotic block (olistolith) from the Ad Daffah conglomerate in eastern Oman (Batain), which provides new insights into the ecology and diversity during the early aftermath of the Permian–Triassic Boundary mass extinction. Based on conodont quantitative biochronology, we assign a middle Griesbachian age to the upper part of this boulder. It was derived from an offshore seamount and yielded both nektonic and benthic faunas, including conodonts, ammonoids, gastropods and crinoid ossicles in mass abundance. This demonstrates that despite the stratigraphically near extinction at the Permian–Triassic Boundary, Crinoidea produced enough biomass to form crinoidal limestone as early as middle Griesbachian time. Baudicrinus, previously placed in Dadocrinidae, is now placed in Holocrinidae; therefore, Dadocrinidae are absent in the Early Triassic, and Holocrinidae remains the most basal crown‐group articulates, originating during the middle Griesbachian in the Tethyan Realm. Abundant gastropods assigned to Naticopsis reached a shell size larger than 20 mm and provide another example against any generalized Lilliput effect during the Griesbachian. Whereas the benthic biomass was as high as to allow the resumption of small carbonate factories, the taxonomic diversity of the benthos remained low compared to post‐Early Triassic times. This slow benthic taxonomic recovery is here attributed to low competition within impoverished post‐extinction faunas.     

Palaeoecological significance of a new Griesbachian (Early Triassic) gastropod assemblage from Oman

A new Early Triassic (Griesbachian) gastropod fauna from the Al Jil Formation of Oman is described. Early Triassic faunas from elsewhere (e.g. the Italian Dolomites and the western USA) are typically of low diversity and high dominance, usually attributed to environmental stress in the immediate aftermath of the end-Permian mass extinction event. The new Oman fauna has, by contrast, relatively high diversity, low dominance and a more even spread of individuals between taxa. It is the most diverse Griesbachian fauna known to date. This is attributed to the favourable (i.e. well-oxygenated) conditions under which the fauna lived. This uncharacteristic Griesbachian gastropod fauna demonstrates that, in the absence of oceanic anoxia, biotic recovery after the end-Permian extinction event may occur surprisingly quickly (within one conodont zone). The fauna is also partially silicified, which has increased its preservation potential relative to other Griesbachian gastropod assemblages. However, only one reappearing Lazarus genus is present in the Oman fauna. This suggests that there was some other control on the abundance of Lazarus genera at this time, other than the absence of silicified faunas as previously suggested.     

An unusually diverse mollusc fauna from the earliest Triassic of South China and its implications for benthic recovery after the end-Permian biotic crisis

A single carbonate coquinoid lens from the Griesbachian (Early Triassic) of Shanggan, South China, yielded 11 bivalve species described in this study in addition to four gastropod and one ammonoid species reported elsewhere. This makes the Shanggan fauna one of the richest mollusc faunas from the early post-extinction interval after the end-Permian mass extinction event. Four of the present genera are long-term survivors, five are holdovers that went extinct at the end of the Griesbachian or later in the Early Triassic, and seven first appear in the Griesbachian. Three new bivalve species are described: Myalinella newelli nov. sp., Scythentolium scutigerulus nov. sp., and Eumorphotis shajingengi nov. sp. The genus Astartella, previously assumed to have vanished at the end of the Permian, is reported for the first time from the Early Triassic, which also removes Astartidae from Early Triassic Lazarus taxa. The small growth size of the Astartella specimens supports an earlier hypothesis that many of the Early Triassic Lazarus taxa did not survive in unknown refuges but were simply overlooked due to the scarcity of easily observable large-sized specimens. Ecologically, a comparatively high proportion of infaunal bivalve species (4/11) is remarkable for the early post-extinction interval, supporting the impression of a relatively advanced recovery state. Moreover, abundance-data of the bivalve-gastropod community reveal a remarkably low dominance index (D = 0.17) that is suggestive for advanced recovery and stable environmental conditions. It is proposed that the Shanggan fauna represents a late Griesbachian benthic recovery event that coincided with the appearance of similarly diverse benthic faunas in Oman and Primorye. A high proportion of genera that have previously not been reported from the Early Triassic indicate that the prevalence of poor preservation conditions is a major obstacle in identifying early phases of recovery from the greatest crisis in the history of metazoan life. The early recovery of benthic faunas reported in this study questions previous claims of a prolonged lag phase as a consequence of the extraordinary extinction magnitude or the persistence of adverse environmental conditions.     

Norigondolella n. gen., eine neue obertriassische Conodontengattung

The development of the genus concept for platform-bearing gondolellid conodonts is discussed. The collective genusNeogondolella Bender & Stoppel s. l. must be subdivided into several genera separated mostly by morphological differences of their platform elements. Only for these genera, but not forNeogondolella s. l., differential diagnoses againstGondolella Stauffer & Plummer s. l. can be given.Norigondolella n. gen., species of which were formerly placed intoParagondolella Mosher s. l.,Gondolella Stauffer & Plummer s. l. orNeogondolella Bender & Stoppel s. l., is a typical gondolellid platform conodont genus of topmost Triassic (Norian-Rhaetian) pelagic sediments. It has evolved from Carnian representatives ofParagondolella, but the lower surface of the platform element shows considerable similarities (homoeomorphy) to the Middle Carboniferous to Middle Permian genusMesogondolella Kozur.     

Middle Triassic conodont apparatus architecture revealed by synchrotron X-ray microtomography

The composition of conodont apparatuses is crucial for understanding the feeding mechanisms of these early vertebrates. However, the multielement apparatus reconstructions of most species remain equivocal because they have been inferred from loose element collections, guided by knowledge from rare articulated ‘bedding plane assemblages’ and fused clusters, often from distantly related taxa. Even these natural assemblages can be difficult to interpret because the component elements can be closely juxtaposed or embedded in matrix, making it hard to discern the morphology of each element and their relative positions within the architecture of the feeding apparatus. Here we report five exceptionally preserved conodont clusters from the Middle Triassic Luoping Biota, Yunnan Province, Southwest China. These materials were scanned using synchrotron radiation X-ray tomographic microscopy (SRXTM), revealing the morphology and positional homology of the component elements in the fused clusters. We confirm that the apparatus of Nicoraella was composed of eight types of elements, comprising a total of 15 elements. SRXTM reveals the positional homologies of the component elements, viz. a single alate element is located in the S₀ position, flanked successively abaxially by pairs of breviform digyrate S₁ and S₂ elements, bipennate S₃ and S₄ elements, and a pair of inwardly curved breviform digyrate M elements. Carminate elements occupy the P₁ and P₂ positions. The apparatus of Nicoraella is among the most completely characterised of all conodonts and serves as a template for the reconstruction of gondollellids.     

华南贵州宾夕法尼亚亚纪牙形类Neognathodus动物群:演化及地层对比意义

以华南斜坡相的罗甸纳庆剖面为代表,我国石炭纪宾夕法尼亚亚纪早–中期地层建立起了基于多个属种的牙形类分带,并可与全球其他地区同期地层进行广泛对比。Neognathodus属是这一时期一类常见的浅水相牙形类分子,在北美、东欧等地的浅水相地层中被广泛用作地层划分对比标志。该属分子在我国北方地区浅水相地层中是重要的带化石分子,而在华南斜坡相剖面中占比非常低。结合现有的生物地层框架,本文厘清了华南贵州罗甸附近3条斜坡相剖面中Neognathodus属分子的延限,尝试利用牙形类P1分子齿台的对称特征将其演化过程归纳为五个阶段,并提出阶段4中对称分子(如:N. bothrops, N. colombiensis, N. nataliae等)的再次出现可作为全球莫斯科阶底界的辅助对比标志。     

A new Devonian species of the enigmatic Carboniferous conodont Dollymae

Arrow-shaped P₁ elements of Dollymae are characteristic for the Tournaisian conodont faunas worldwide but the phylogeny of this conodont remains obscure and its origin is cryptic. Dollymae peregrina n. sp., from the upper Famennian (upper expansa Zone) strata in the Holy Cross Mountains (central Poland), is the oldest representative of the Dollymae lineage. The morphology of the new species’ P₁ element shows primitive features in comparison to other members of this genus and is transitional in the number of element processes. Thus D. peregrina n. sp. is most probably ancestral to the remaining species. The genus appears to be monophyletic, probably rooted in Pelekysgnathus within the Icriodontidae.     

A NEW EARLY ORDOVICIAN CONODONT GENUS FROM THE SOUTHERN MONTAGNE NOIRE, FRANCE

Abstract:  Two species of a new Tremadocian (Early Ordovician) conodont genus from the Saint Chinian Formation of the southern Montagne Noire, France, are erected: Hammannodus sarae gen. et sp. nov. and Hammannodus juliae gen. et sp. nov. They were found within a single storm-induced limestone nodule interbedded with offshore shales belonging to the regional Shumardia ( C .) pusilla (trilobite) Biozone, and to the Paltodus deltifer deltifer (conodont) Subzone. This conodont record is associated with the episodic development of carbonate productivity in temperate waters of the Montagne Noire platform, a process absent in neighbouring platforms of north-west Gondwana. The apparatus is composed of five coniform pyramidal elements occupying P and S positions and one bicostate element in the M position, having three or two sharp costae, respectively, with a subtriangular basal outline.     

Early Triassic conodont clusters from South China: revision of the architecture of the 15 element apparatuses of the superfamily Gondolelloidea

Abstract: Several fused clusters of conodont elements of the genera Neospathodus and Novispathodus were recovered from limestone beds at the Dienerian–Smithian and Smithian–Spathian boundaries, respectively, from several localities in Guangxi province, South China. Conodont clusters are otherwise extremely rare in the Triassic, and these are first described for the Early Triassic. The exceptional specimens partially preserve the relative three‐dimensional position and orientation of ramiform elements and are therefore extremely important for testing hypotheses on the architecture of apparatuses. These specimens partly confirm the previous reconstruction of the Novispathodus apparatus by Orchard. Within apparatuses of members of superfamily Gondolelloidea, elements previously identified as occupying the S₁ and S₂ positions instead occupy the S₂ and S₁ positions. Similarly, within apparatuses of members of the subfamily Novispathodinae, elements previously referred to S₃ and S₄ positions are reinterpreted to have occupied S₄ and S₃ positions, respectively.     

A 17‐element conodont apparatus from the Soom Shale Lagerstätte (Upper Ordovician), South Africa

Abstract:  Natural assemblages of a new conodont taxon, Notiodella keblon, from the Upper Ordovician Soom Shale Lagerstätte of South Africa contain 17 elements. This is the first time that a 17‐element apparatus plan has been unequivocally demonstrated in conodonts. The apparatus comprises paired P₁, P₂, P₃, M, S₁, S₂, S₃ and S₄ elements and an unpaired, axial S₀ element and provides a new template for use in the reconstruction of apparatuses from the collections of dispersed elements, particularly for those with icrion‐bearing P₁ elements and perhaps for other balognathids.     

A spathognathodont lineage of Mississippian conodonts

Evolutionary lineages of conodont elements and of some apparatuses are gradually being documented to increase the value of these fossils in stratigraphic interpretation and to further our understanding of conodont taxonomy. The well-documented faunas from Mississippian strata of the Illinois Basin and type Mississippian beds provide a great deal of information for the recognition of phylogenies. The disjunct element genus Spathognathodus is part of a basic and long-ranging type of conodont apparatus from which a number of other forms have been derived, and some of its Mississippian disjunct element species can now be placed in multielement species. A lineage involving spathognathodont Pa elements recognized in the Mississippian, then, is the basis of the present study. The phylogeny recognized started with Synprioniodina? crassidentata, and includes S.? regularis, S.? pulchra, S.? coalescens, S.? spicula, Hinduodus cristula, and H. minutus in the Mississippian. Aputiignathus scitulus and A. penrscitulus apparently represent a separate line, which also might include disjunct element Spathognathodus rexroadi.