High‐resolution conodont biostratigraphy from the Darriwilian Stage (Middle Ordovician) of the Argentine Precordillera and biodiversity analyses: a CONOP9 approach

Ordovician conodonts have been extensively documented in the Argentine Precordillera, providing a robust database for a diverse set of palaeontological studies. Despite the numerous studies, the published taxonomy and stratigraphical ranges, data remain contradictory for particular time intervals. Data from a new conodont collection from the Gualcamayo Formation exposed at the Cerro La Chilca section, and new information on the occurrence and ranges of species from the Las Chacritas and Las Aguaditas sections are presented herein. We used a computer‐assisted numerical sequencing program (CONOP9) to construct a composite stratigraphical range chart from data of 57 conodont species in four sections from the Argentine Precordillera. The identified zones of Lenodus variabilis, Yangtzeplacognathus crassus, Eoplacognathus pseudoplanus and Eoplacognathus suecicus allowed us to verify and adjust the biostratigraphical scheme for the Darriwilian of the Central Precordillera. Additionally, species of the genera Histiodella, namely Histiodella sinuosa, Histiodella holodentata, Histiodella kristinae and Histiodella bellburnenisis, enable a reasonable correlation between the Histiodella‐based zonation and the Baltoscandian zonation. Conodont diversity is evaluated using conventional measures (total diversity and normalized diversity) and an interval‐free approach with CONOP9 software. Our data show a positive pattern in conodont diversification, increasing rapidly through the L. variabilis to the Y. crassus zones and reaching a peak in the E. pseudoplanus Zone. When analysing diversity fluctuations with respect to the environmental changes within the depositional basin, migrations in and out of the basin related to local sea level fluctuations appear to be an important process driving the conodont diversity pattern in the Precordillera.     

The Middle Caradoc Facies and Faunal Turnover in the Late Ordovician Baltoscandian palaeobasin

A Late Ordovician episode of remarkable biotic, climatic, sea level and facies changes, named here as the Middle Caradoc Facies and Faunal Turnover, took place in the Baltoscandian area. This paper presents an integrated overview of these changes in the critical middle Caradoc interval. Data are given on carbonate rock composition, distribution and grain-size composition of the siliciclastic material and the carbon isotopic composition of whole-rock carbonates in cores of Estonia and Sweden.

The Middle Caradoc Facies and Faunal Turnover can be described as a succession of related environmental changes. The turnover began with a positive excursion in carbonate δ¹³C and continued with sea level changes that led to a sedimentary hiatus on the shelf and a change from carbonate-dominated to siliciclastic-dominated sedimentation in the basin. The turnover ended with an extinction event and associated microfaunal crisis.

The middle Caradoc turnover in Baltoscandia is comparable to a similar succession of changes in North America. The turnover affected two palaeocontinents, and reflects a widespread, possibly global environmental change. Onset of glaciation on Gondwana and/or increased orogenic activity might have initiated the changes in ocean circulation and led to the initial carbon isotope excursion. The following sea level rise and faunal changes affected several different continents.     

The oldest known occurrence of the Foliomena fauna in the uppermost Darriwilian (Middle Ordovician) of South China

The oldest Foliomena fauna was, until now, known from the middle–upper Miaopo Formation (Nemagraptus gracilis Biozone, lower Sandbian, basal Upper Ordovician) of South China. In this study, the oldest record of the fauna is set back to the latest Darriwilian (upper Hustedograptus teretiusculus Biozone), represented by Foliomena jielingensis and some typical constituents of the Foliomena fauna from the basal Miaopo Formation at Jieling, northern Yichang, western Hubei Province, central China. The Miaopo Formation is characterized by its organic‐rich dark‐grey shale facies, unique in its localized distribution on the Yangtze Platform, and distinguished by its rich and diverse benthic and graptolitic faunas. This suggests an origin of the Foliomena fauna in periodically oxygen‐starved local depressions on the Yangtze Platform during the Middle–Late Ordovician transition.     

Palaeoenvironmental significance of cool‐water microbialites in the Darriwilian (Middle Ordovician) of Sweden

Well‐developed oncoids and centimetre‐sized stromatolites are reported for the first time from the Darriwilian (Middle Ordovician) cool‐water ‘orthoceratite limestone’ at Kinnekulle, Västergötland, Sweden. The characteristics and stratigraphical distribution of these microbialites show an apparent relationship to fluctuations in relative sea level. The most abundant and well‐developed oncoids occur in stratigraphical intervals that are characterized by notable sea‐level lowstands. Stromatolites, which share many compositional characteristics with the oncoids, are essentially confined to a single bed associated with an especially prominent lowstand. Stromatolite‐like lamination also occurs in the uppermost part of the studied succession, but this feature may be of abiogenic origin. The microbialites appear to be originally calcareous, but synsedimentary iron‐ and/or phosphate‐enriched laminae are conspicuous, and secondary substitution by coarse calcite and barite is common. Iron staining is most prominent in poorly preserved specimens. Diagenesis has occluded the identity of the producers of these microbialites, but characteristics of associated endolithic borings suggest that they were formed in photic waters. The laminated fabrics of the documented microbialites record a depositional environment sensitive to high‐frequency environmental change. Most significantly, the microbialites have provided important information about the depositional environment of their enigmatic host limestone, and the collective observations challenge the notion that the studied strata were deposited in a deep shelf to basinal environment – rather, it appears that they are to a large extent, shallow‐water deposits, formed in waters only a few tens of metres deep.     

Allometry in Anisian (Middle Triassic) segminiplanate conodonts and its implications for conodont taxonomy

Conodonts are a clade of chordates and are valuable indicator fossils for biostratigraphy. The segminiplanate (neogondolelliform) conodonts represent a major morphological group ranging from upper Carboniferous to Upper Triassic marine sediments. However, the morphological similarity of segminiplanate P₁ elements generates problems for taxonomy, especially in the Permian and Triassic clades. This paper represents the first study of morphological variation in Triassic segminiplanate conodonts using a geometric morphometric approach. The laminar microstructures observed in conodont cross‐sections indicate that, within our analysed specimens, smaller conodonts with fewer laminae are generally from an earlier ontogenetic stage while larger conodonts with more laminae are from a later stage of ontogeny. Using linear regressions between relative warp scores from both upper and lateral views and conodont length, we demonstrate strongly allometric growth patterns for the species Paragondolella bifurcata Budurov & Stefanov. Our results indicate that the species‐group taxon Pg. praeszaboi bystrickyi (Kovacs et al.) is an early growth stage of Pg. bifurcata and thus synonymous. We suggest that the allometry of conodonts should be considered seriously, especially when there are numerous transitional morphologies between large‐ and small‐sized conodonts. Reconstructing the ontogenetic series and using larger‐sized conodonts within the numerous transitional morphologies in the population of a rock sample for the definition of new species are suggested for future studies.     

Conodont biostratigraphy and carbon isotope chemostratigraphy of the Middle to Upper Ordovician on the western Yangtze Platform,South China

Middle and Late Ordovician strata in the nearshore area of the western Yangtze Platform are investigated. In total, 241 δ¹³C_carb samples and 108 conodont samples from the Songliang and Tuanjie sections in the northeastern Yunnan Province were collected for chemostratigraphic and biostratigraphic studies. Altogether, 21 species belonging to 16 conodont genera were recovered from the two sections, among which Ansella jemtlandica, Drepanoistodus basiovalis, Hamarodus brevirameus, and Microzarkodina hagetiana are important age indicators within the Dapingian to early Katian interval. Four carbon isotope shifts are recognized in the studied sections: a negative shift in the early Darriwilian, the middle Darriwilian carbon isotope excursion (MDICE), the early Katian Guttenberg carbon isotope excursion (GICE), and a further positive carbon isotope shift later in the early Katian which is a new record of this event in South China. Integrated carbon chemostratigraphy and conodont biostratigraphy indicate early Darriwilian and early Katian ages for the base and top, respectively, of the Huadan Formation at the Songliang section, and a Darriwilian to early Katian age for the Shihtzupu Formation at the Tuanjie section.     

Billingen (Lower Arenig/Lower Ordovician) acritarchs from the East European Platform and their palaeobiogeographic significance

Billingen (Lower Arenig/Lower Ordovician) sediments of the St. Petersburg region, northwest Russia and the Leba area, northern Poland of the East European Craton yield acritarch assemblages, which are largely homogenous though displaying minor compositional differences that probably reflect a gradient from inner to outer shelf environments. Comparison with coeval acritarch microflora from the Yangtze Platform, South China, shows an overall similarity between Baltoscandian and South Chinese phytoplankton. The widespread uniformity in the fossil microphytoplankton may be related to the extensive global 'evae' sea-level transgression, which characterized the Billingen time. This suggests that during the Tremadoc through early Arenig times, acritarch assemblages displayed essentially an undifferentiated cold-water and oceanic character along the whole margin of Perigondwana in the South, as well as on the South Chinese and Baltic platforms, at middle latitudes (Mediterranean oceanic Realm). Despite this overall similarity, however, some typical taxa of the high-latitude Mediterranean Province (Arbusculidium, Coryphidium and Striatotheca) occur in South China, but are absent in Baltica. This discrepancy is explained as caused by differences in climatic and physiographic conditions that prevailed at the two palaeocontinents at this time. The inferred pattern of oceanic circulation during the Lower Ordovician is consistent with the palynological evidence of a prevailing warmer climate in Baltica than in South China, although the two palaeocontinents occupied the same palaeolatitudinal position.     

The Darriwilian Hiswah fauna of western Gondwana (Jordan): Biostratigraphy,palaeogeography and palaeoecology

The fauna and facies of the Hiswah Formation in southern Jordan were investigated. The formation indicates deposition under low energy, open-marine, siliciclastic shelf conditions and reflects the maximum of the first regional post-Cambrian marine transgression on the northern edge of the Arabian-Nubian Shield. This eustatic signal accompanied by a maximum flooding surface can be traced over the entire Arabian Plate and Turkey. The lower part of the Hiswah Fm. consists of pelagic mudstones with only occasional thin beds of rippled, hummocky cross-bedded siltstone and mudstone concretions, and contains a distinct fauna of low diversity. The upper part of the formation is characterized by sandstone-siltstone alternations with some hummocky cross-stratification in its lower portion and trough cross-bedding and ripple marks above indicating somewhat shallower conditions. A very detailed re-investigation of the fauna from the lower part of the Hiswah Fm. and extensive new fossil material indicates that the most remarkable faunal element is the planktonic graptolite Didymograptus murchisoni (Beck in Murchison, 1839in) which, in the past, has been determined erroneously as D. bifidus (Hall in Berry, 1962). Numerous specimens of small obolid brachiopod Palaeoglossa sp. cf. P. attenuata (Sowerby, 1839) represent the most common benthic element within the Hiswah fauna. The exceptional arthropod Hanadirella cf. bramkampi El-Khayal, 1985, is reported for the first time from Jordan, representing its fifth known region of occurrence in addition to Saudi Arabia, Algeria, Spain, and France. For the first time, cephalopod remains are described from the Ordovician of Jordan by the occurrence of ?Dideroceras sp. and a second, indeterminate species. The biostratigraphic age of the Hiswah fauna is revised to late Darriwilian (Dw 3, upper Middle Ordovician) according to our re-investigation of the graptolites. The Hiswah fauna represents the oldest body fossils of the Ordovician marine sedimentation in southern Jordan and is palaeogeographically strongly related to northern and northeastern Saudi-Arabia and to high/mid-latitude Gondwana margins in general.     

The earliest known strophomenoids (Brachiopoda) from early Middle Ordovician rocks Of South China

The upper Daguanshan Formation (middle Expansograptus hirundo graptolite biozone, Dapingian, early Middle Ordovician) of the Shuanghe area, Changning County, southern Sichuan Province, contains three new genera and species of strophomenoids: Ochyromena plana, Shuangheella elongata, and Primotimena globula, which are attributed to the Strophomenidae, Rafinesquinidae and Glyptomenidae respectively. These are the earliest known strophomenoids from the South China palaeoplate, and also the oldest rafinesquinid and glyptomenid brachiopods worldwide. Global review of the superfamily Strophomenoidea of Middle Ordovician age suggests that the first diversity peak at the species level occurred in late Darriwilian (Llanvirn) time, mainly as a result of the rapid diversification of the family Strophomenidae. The first appearance datum (FAD) of strophomenoids and their possible westward dispersal were from North China (latest Floian) and/or South China (early Dapingian), through the Chu‐Ili terrane of Kazakhstan, Iran, and Baltica (early Darriwilian), to Avalonia and Laurentia (late Darriwilian). This points to the existence of early diversification hotspots of the strophomenoid superfamily in the North and South China palaeoplates during the early Middle Ordovician in generally shallow water (corresponding to BA2) environments. The high degree of similarity in the external morphology and ventral interior of the three new genera indicates that the early diversification of strophomenoids began with differentiation of the cardinalia, especially in the configuration of the bilobed cardinal process, a key evolutionary novelty for the strophomenoids.     

The Upper Tremadocian (Ordovician) graptolite Bryograptus: taxonomy, biostratigraphy and biogeography

Abstract:  The taxonomy, biostratigraphical and palaeogeographical distribution of the Lower Ordovician graptolite genus Bryograptus is evaluated. Bryograptus is recognized as a distinct triradiate anisograptid with a multiramous, pendent rhabdosome. The species of the genus Bryograptus can be interpreted as shallow water faunal elements with a strongly limited biogeographical distribution to the Atlantic Faunal Realm. Bryograptus is restricted to a narrow interval in the Upper Tremadocian, the Bryograptus Biozone of Scandinavia and South America (Argentina), making it a taxon with a high potential for precise biostratigraphical correlations. The proximal end development can be used to differentiate the genus Bryograptus from other pendent multiramous graptoloid genera with a homoplastic rhabdosome development. Characteristics of the proximal end development and structure easily differentiate these genera in relief specimens, but not in flattened material.     

Earliest ontogeny of Middle Ordovician rhynchonelliform brachiopods (Clitambonitoidea and Polytoechioidea): implications for brachiopod phylogeny

New data on the earliest ontogeny of Mid-Ordovician Baltoscandian clitambonitoid ( Apomatella , Neumania and Oslogonites ) and polytoechioid ( Antigonambonites and Raunites ) brachiopods reveal significant differences in the life history of the taxa belonging to these two superfamilies. The Polytoechioidea and probably other members of the Billingsellida had planktotrophic larvae, in which the dorsal and ventral mantle lobes formed separately and without reversion. The 'pedicle sheath' in Antigonambonites is secreted by a section of modified ventral mantle and thus this 'pedicle' is not homologous within the pedicle of rhynchonellate brachiopods. It is likely that polytoechioids and other members of the strophomenate clade had the same type of ontogeny and mode of attachment. In contrast, the ontogeny and mode of attachment of clitambonitoids are similar to that of recent rhynchonellates: their mantle lobes and both valves formed simultaneously, and the pedicle most likely formed from the larval pedicle lobe. Evidence for the lecithotrophic nature of clitambonitoid larva is discussed. This confirms that the Clitambonitoidea, unlike the Polytoechioidea, represents an ingroup within the Rhynchonellata.     

High‐resolution δ13Corg chemostratigraphy links the Decorah impact structure and Winneshiek Konservat‐Lagerstätte to the Darriwilian (Middle Ordovician) global peak influx of meteorites

The precise age of the Winneshiek Shale, a recently discovered Konservat‐Lagerstätte located in a very unusual depositional setting inside the Decorah impact structure, has remained uncertain in the absence of biostratigraphically highly diagnostic fossils. This chemostratigraphical study, based on δ¹³C_org data from 36 drill core samples through the shale, shows that the age ranges from the upper part of a small unnamed δ¹³C excursion in the Dw1 Stage Slice of the Darriwilian Global Stage to the lower part of the MDICE excursion in Stage Slice Dw2 of the same stage. This Dw1–Dw2 interval has an isotopic age of ~464–467 Ma. The gradational contact between the Winneshiek Shale and the underlying, rapidly deposited, impact breccia indicates minimal time difference between the impact event and the Winneshiek Shale. New age data show that the Decorah impact event was coeval with the early Darriwilian abnormally high influx of micrometeorites and meteorites recorded in sections in Baltoscandia, Russia and China and that the Decorah crater can be included among the unusually large number of meteorite craters formed during Middle and early Late Ordovician time. As is commonly the case in black shale deposits, the partly uniquely preserved Winneshiek Shale crater fauna is impoverished taxonomically and adds numerically relatively little to the conspicuous and much discussed Darriwilian global biodiversification increase.     

New data on perittocrinids and hybocrinids (Crinoidea, Echinodermata) from the Middle Ordovician of the Baltic region

Age variation of radial plates of perittocrinids and aberrant cups of the hybocrinid genus Hoplocrinus from the Middle Ordovician of the St-Petersburg region are examined. Three patterns for the development of folds on the radial plates of perittocrinids are recognized. The differences between patterns are of species level. It is proposed here that the folded structures of perittocrinids (as well as the pore-structures of other pelmatozoans) were not only used for respiration, but also very likely for feeding. Tetracionocrinus transitor is a young individual of a species of the genus Perittocrinus or a paedomorphic species of this genus. The study of aberrant cups of Hoplocrinus suggests that its ancestor could have had biserial arms. Phylogenetic implications of this interpretation are discussed, and homologies within the phylum Echinodermata, reassessed. The three-armed genus Baerocrinus may be an aberrant form of a species of the genus Hoplocrinus.     

Evenness and diversity in Upper Cambrian – Lower Ordovician trilobite communities from the Central Andean Basin (Cordillera Oriental,Argentina)

Community evenness has recently received much attention, either because it is related to ecosystem functioning or because it may affect estimation of diversity. Temporal and environmental trends in diversity and evenness of trilobite communities during the Late Cambrian – Early Ordovician of the Cordillera Oriental (north‐western Argentina) are here analysed. Richness and evenness increase through time in both deep subtidal (between fair‐weather and storm wave base) and offshore (below storm wave base) communities. Two significant patterns are superimposed on this general trend: (1) the magnitude of the increase in evenness is much more pronounced in deep than in shallower settings, and (2) richness and evenness trajectories are decoupled (while a significant rise in evenness is recorded in the middle Tremadocian (Tr2), an increase in richness is delayed until the late Tremadocian (Tr3)). In contrast to expectations, a single family (Olenidae) is dominant in samples associated with this earlier rise in evenness relative to richness. Hence, this trend is explained neither by the number of families present in the communities nor by the familial identity of the most abundant taxon. Large‐scale comparisons of the timing and geographical components of these trends are restricted to the patterns recognized in Laurentian North American studies. Results from the Cordillera Oriental mirror those of Laurentia regarding the rise in both metrics in deep marine settings. Nevertheless, the timing of this increase in richness and evenness is delayed in the Cordillera Oriental, supporting the idea that palaeogeographical regions differed in the nature and timing of ecological changes. Finally, the rise in trilobite alpha‐diversity through the Late Cambrian – Early Ordovician of the Cordillera Oriental supports the idea that trilobite alpha‐diversity did not decline worldwide, suggesting that the relative decline in trilobite alpha‐diversity is most probably caused by the dilution effect.     

Late Ordovician (Turinian–Chatfieldian) carbon isotope excursions and their stratigraphic and paleoceanographic significance

Five positive carbon isotope excursions are reported from Platteville–Decorah strata in the Upper Mississippi Valley. All occur in subtidal carbonate strata, and are recognized in the Mifflin, Grand Detour, Quimbys Mill, Spechts Ferry, and Guttenberg intervals. The positive carbon isotope excursions are developed in a Platteville–Decorah succession in which background δ¹³C values increase upward from about −2‰ at the base to about 0‰ Vienna Pee Dee belemnite (VPDB) at the top. A regional north–south δ¹³C gradient, with lighter values to the north and heavier values to the south is also noted. Peak excursion δ¹³C values of up to +2.75 are reported from the Quimbys Mill excursion, and up to +2.6 from the Guttenberg excursion, although there are considerable local changes in the magnitudes of these events. The Quimbys Mill, Spechts Ferry, and Guttenberg carbon isotope excursions occur in units that are bounded by submarine disconformities, and completely starve out in deeper, more offshore areas. Closely spaced chemostratigraphic profiles of these sculpted, pyrite-impregnated hardground surfaces show that they are associated with very abrupt centimeter-scale negative δ¹³C shifts of up to several per mil, possibly resulting from the local diagenetic effects of incursions of euxinic bottom waters during marine flooding events.     

Integrated radiolarian and conodont biostratigraphy of the Middle Permian Gufeng Formation (South China)

Radiolarians are usually abundant in chert sequences and they have thus been widely used for the biostratigraphy of deep-water sediments. However, there are many difficulties in the correlation of radiolarian biostratigraphic schemes with the standard conodont zones. In this study, 21 radiolarian species were extracted from the Gufeng Formation that crops out in the Luojiaba (LJB) section (western Hubei, China), together with 5 co-occurring conodont species. In this way, it is the first time that the Pseudoalbaillella globosa, Follicucullus monacanthus and F. scholasticus radiolarian zones can be directly correlated with the Jinogondolella nankingensis gracilis, J. aserrata and J. postserrata conodont zones. Accordingly, the 3 radiolarians zones are now firmly correlated with the Roadian to middle Capitanian interval (Middle Permian).     

Astogenetic aspects of the dendroid graptolite Airograptus (Late Cambrian-Early Ordovician) and their phylogenetic implications

Based on a Backscatter Electron (BSE) examination of partially 3-dimensional graptolite specimens of Airograptus from the Tremadocian (Early Ordovician) Yehli Formation at Dayangcha, Baishan of Jilin Province, NE China, the thecal structure and astogenetic aspects of this genus (except for the missing proximal part) are described and illustrated in detail. In the present collection, no furcating but only peltate thecal apertural processes are confirmed. A comparison between the presumably benthic dendroid Airograptus and Rhabdinopora, the earliest planktonic graptolite, indicates that there are remarkable similarities in astogenetic aspects between them, although there are also differences in the development of dissepiments and thecal apertural processes. A possible ancestor-descendent relationship between the two genera is herein proposed.     

Magnesian calcite and chamositic ooids forming shoals peripheral to Late Ordovician (Ashgill) muddy siliciclastic shores: southern Ontario

In southern Ontario, ooids are associated with two distinct facies associations in the Queenston Formation, the final stage of Late Ordovician (Ashgill) Taconic basin fill. One facies consists of thin ooid and bioclastic grainstones interbedded with mudrock, and lies near the base of the formation, and, in southwestern Ontario, also forms a local NW-thickening wedge near the middle of the formation. Ooids have radial-fibrous and radial-concentric fabrics (Type A), with chamosite, illite, and Fe-oxide laths at intercrystalline sites. Vertical lithologic and ooid abundance patterns indicate that thresholds to carbonate production were sensitive to changes in terrigenous sediment supply, sea level, circulation, accommodation space, and tectonism.

Ooids in the second facies association are admixed with abraded fragments of open-marine biota, or occur burrow fills, within a <30-cm-thick interval of mudrock near the top of the preserved Queenston succession, a few metres below the Ordovician–Silurian unconformity. Ooids have radial concentric and crosscutting patchy microcrystalline fabrics (Type B). This unit may represent a transgressive or stillstand deposit modified by bioturbation.

The extent of preserved fabric suggests that both ooid types were originally magnesian calcite, but Type A ooids underwent greater burial alteration. This is shown by crystalline mosaics that cross-cut relict primary fabrics; δ¹³C values (−1.82‰ to +0.67‰) and δ¹⁸O values (−4.46‰ to −10.57‰) more negative than marine calcite of similar age; Mn and Fe concentrations more elevated above expected marine values; and a luminescence similar to that of intergranular cements. Burial meteoric diagenesis was likely promoted by excellent permeability of the host sand. We interpret authigenic chamosite and Fe-oxide to reflect diagenesis of iron-bearing and clay detritus trapped during ooid growth. Type B ooids suffered less alteration: δ¹³C (+1.1‰ to +6.64‰) and δ¹⁸O (−3.04‰ to −4.81‰) values overlap the expected marine range, including ¹³C enrichment that occurs within the Hirnantian (latest Ordovician) excursion. Although Mn and Fe values are still higher than those of modern calcitic ooids, negligible luminescence suggests that recrystallization occurred in the presence of marine-derived pore fluids. Further burial alteration was inhibited due to low permeability of the host mud.

Type A ooid facies in the Queenston Formation forms an ancient analogue for lesser known Quaternary ooid shoals peripheral to tropical deltaic systems. The facies of Type B ooids, while more enigmatic, may preserve a geochemical herald of latest Ordovician climate change. The presence of minor chamosite in Type A ooids defines a possible distal facies of the well-known oolitic ironstones of similar age in the mid-continental USA.