Early Tremadocian (Ordovician) graptolite biostratigraphy and correlation

A revision of the early Tremadocian graptolites of the genus Rhabdinopora shows that Rhabdinopora proparabola from the Xiaoyangqiao section, Dayangcha, North China is the oldest species of the genus. Early Tremadocian species of Rhabdinopora can be differentiated easily by the construction of their nemata. The earliest species, Rhabdinopora proparabola, lacks a nema, while a multibranched nema is present in Rhabdinopora campanulatum (= Rhabdinopora parabola). The Rhabdinopora proparabola, Rhabdinopora campanulatum, Rhabdinopora flabelliformis (Anisograptus matanensis Biozone) and Rhabdinopora anglica biozones can be correlated worldwide. A short interval at the base of the Ordovician System does not contain any planktic graptolites.     

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.     

A Tremadocian (Early Ordovician) palaeoscolecidan worm from graptolitic shales in Hunan Province,South China

A new locality exposing Tremadocian (Early Ordovician) graptolitic shales in Hunan Province, South China, has yielded an exceptionally well‐preserved annulated worm. This palaeoscolecidan is described as Waflascolex changdensis gen. et sp. nov. and reveals extremely fine detail of the cuticle organization and plating array. The new taxon is characterized by three critical characters: incomplete plate rows that occur only on the posterior end of the worm (except in the posterior‐most area) and do not extend over the entire circumference or along the entire trunk; a regular rhomboidal array of platelets around intercalations; and reduced cuticular organization at the posterior termination. The unique cuticular organization and platelet ornamentation in the new taxon offer insight into functional differentiation of plates in the scleritome. Palaeoscolecid distribution through the early Palaeozoic is reviewed, showing that the worms were widespread in the Cambrian and Ordovician, but became more restricted during the Silurian. Ordovician palaeoscolecidans are diverse in scleritome architecture, and strikingly different from Cambrian taxa, indicating that this group diversified as part of the Great Ordovician Biodiversification Event.     

Tremadoc (earliest Ordovician) brachiopods from Purmamarca and the Sierra de Mojotoro, Cordillera Oriental of northwestern Argentina

The Late Tremadoc storm-dominated shoreface to inner platform deposits exposed west of the Purmamarca village (Coquena Formation) contain a considerably more diverse brachiopod fauna than previously reported. Coquinite horizons from the lower heterolitic succession have yielded monospecific associations of Nanorthis purmamarcaensis nov. sp. (formerly assigned to N. christianiae KJERULF), which is also reported from the Late Tremadoc rocks of the Cerro San Bernardo area. The fine-grained Upper Member of the Coquena Formation contains a more diverse fauna composed by Nanorthis brachymyaria nov. sp., Astraborthis quebradensis nov. sp. and the new plectorthid genus Lipanorthis (type species L. andinus nov. sp.). A different species of Lipanorthis (L. santalaurae nov. sp.) from the Mid Tremadoc Floresta Formation of the Sierra de Mojotoro is also described.     

Dendroid graptolites from the Lower Ordovician (Tremadocian) of the Yichang area, Hubei, China

The Yichang area of Hubei Province, China provides important information on Lower to Middle Ordovician fossil faunas and is a center for research on Ordovician graptolites due to the sometimes excellent preservation of the copiously available material. New material collected during a research project on Lower to Middle Ordovician biota in the Yichang Region provided among others specimens a number of dendroid (benthic) graptolites of the genera Acanthograptus, Aspidograptus, Dendrograptus, and Koremagraptus showing unexpected and unknown structural features, useful for structural and evolutionary investigations.     

Morphometric analysis of Tremadocian (earliest Ordovician) kirkocystid mitrates (Echinodermata, Stylophora) from the Taebaeksan Basin, Korea

Abundant isolated remains of stylophoran echinoderms (cornutes and mitrates) are reported for the first time in the late Tremadocian (Asaphellus Zone) Tumugol Formation of Korea. Mitrate remains include numerous adorals of Kirkocystidae. Several new important anatomical features have been observed on these adorals, as an internal calcitic layer that is associated to s₂ and possibly also to the palmar complex. This observation suggests that the palmar complex would be present not only in mitrocystitid mitrates, but also in peltocystitids. For the first time, several morphometric analyses have been undertaken based on isolated kirkocystid adorals, so as to explore the morphological diversity displayed by Korean adorals, but also in order to compare their morphology with that of other Gondwanan kirkocystids. Morphometric analyses indicate the occurrence of two contrasted morphologies within Korean adorals (morphotypes A and B), and of three distinct morphologies within European and North African forms (“Anatifopsis”, “Balanocystites”, and “escandei” morphotypes). Comparison of Korean adorals with those from Europe and North Africa shows that: (1) morphotypes B and “Anatifopsis” are equivalent; consequently, the two Korean specimens referred to morphotype B are assigned to the genus Anatifopsis; (2) morphology of most Korean adorals, which belong to morphotype A, is clearly distinct from that of all other described kirkocystids from Europe and North Africa. The small size, juvenile morphology, and great morphological variability observed in the morphotype A of the Korean adorals are suggestive of possible heterochronic processes (peramorphosis).     

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.     

Isolated Chewtonian (Lower Ordovician) graptolites from western Newfoundland

Isolated material of Chewtonian graptolites from the Cow Head Group of western Newfoundland shows some new and unexpected structural details. A high number of the two- and three-stipedPendeograptus fruticosus specimens show metasicular origin for th1¹ combined with a normal isograptid proximal development. Sigmagraptine sicular development includes a large parallel-sided prosicula and a parallel-sided metasicula. One species acquired an azygograptid-like proximal end, characterised by a loss of the initial branching, but kept the capacity to produce distal dichotomies.Maeandrograptus sinosus n. sp. andKeblograptus geminus n. sp. are described as new species.      

Ecostratigraphy, zonation and global correlation of earliest Ordovician planktic graptolites

The succession of earliest Ordovician planktic graptolites (graptoloids) in 11 key sequences around the world is reviewed and used to assess their distribution in depth facies, in paleolatitude and in time. An ecostratigraphic model shows the pattern of distribution of species across the shore-to-ocean profile in time. Fluctuations in this pattern are related to eustatic change; the Acerocare, Black Mountain, Ceratopyge and possibly, the Peltocare Regressive events are recognized. Surface-water temperature is found to have had little significant effect on distribution of Early Tremadoc species, and once the effects of facies is allowed for, their biostratigraphic utility can be assessed. A global suite of four chronozones is proposed for the Early Tremadoc and a further five zones comprise a provisional suite for the Late Tremadoc. The nine zones enable precise correlation of Tremadoc graptolitic sequences. The zones replace the previously used graptolite assemblages, are correlated with key conodont horizons and are, in upward sequence, the Zones of Rhabdinopora praeparabola, R. flabelliformis parabola, Anisograptus matanensis, R. f. anglica, Adelograptus, Paradelograptus antiquus, Kiaerograptus, Araneograptus murrayi-Araneograptus pulchellus , and Hunnegraptus copiosus.     

Early Ordovician bivalves from Dali, West Yunnan, China

A bivalve fauna of Early Ordovician (late Arenig) age is described from the Hsiangyang Formation of the eastern part of West Yunnan, China. The fauna contains elements in common with Early Ordovician faunas of southern Gondwanan areas such as the Montagne Noire and Morocco and with those of Avalonia, together with several previously undescribed taxa. The following taxa are new: Biseriodonta simplex gen. et sp. nov.; Glyptarca sinensis sp. nov.; Trigonoglyptarca magna gen. et sp. nov.; Erhaiconcha xiangyangensis gen. et sp. nov.; Fasciculodonta impressa gen. et sp. nov.; Yunnanoredonia laevis gen. et sp. nov.; Daliella gen. nov.; Goniophorina ( Goniophorina ) contracta sp. nov.; Haidongoconcha radialis gen. et sp. nov.; Eopterinea aequiconcha gen. et sp. nov. The bivalve fauna includes the most diverse glyptarcid fauna and the earliest nuculanid and paracyclid known hitherto.   Eastern West Yunnan belonged to the Indochina terrane and it is concluded that the bivalve fauna represents a high-latitude assemblage and that the Indochina terrane should be considered a component part of the Peri-Gondwanan continent in the Ordovician; its bivalve faunas contrast with those of neighbouring terranes which have affinities with those of lower latitudes.     

The oldest known bryozoan: Prophyllodictya (Cryptostomata) from the lower Tremadocian (Lower Ordovician) of Liujiachang,south‐western Hubei,central China

A new cryptostome bryozoan, Prophyllodictya simplex sp. nov., is described from the Nantzinkuan Formation (Lower Ordovician, lower Tremadoc) of Liujiachang, central China. This antedates the previously oldest known bryozoan by several million years. Colony morphology and the phylogenetic position of Prophyllodictya within Cryptostomata are explored. Phylogenetic analysis suggests that Cryptostomata (except Prophyllodictya) can be divided into two major groups, and that Prophyllodictya occupies a basal position in the cryptostome tree, which accords with its simple morphology and antiquity. A close relationship is evident between bryozoans from the South China palaeoplate and those from Baltica.     

Paleoecological and paleobiogeographic considerations of Ordovician graptolites from the Cordillera Oriental,Jujuy Province,Argentina

We analyse new paleoecological data from South American graptolite records to understand the patterns that influence the space-tempo distribution of the group. A cluster analysis including taxa from the Baltograptus cf. B. deflexus and Didymograptellus bifidus Zones is carried out to evaluate the paleogeographic relationships between north-western Argentina and the other regions in the world. Three different biofacies were recognised in the Acoite Formation. The first biofacies takes place in the lower part of the unit, corresponding to pelitic levels (Tetragraptus phyllograptoides and Tetragraptus akzharensis Zones), and it is represented by pandemic forms. The other two biofacies belong to the endemic or neritic shallow water environments, and develop in deposits corresponding to the middle part of the Acoite Formation (bottom of the Baltograptus cf. B. deflexus Zone) in the Sierra de Aguilar, and in the upper half of the unit exposed in the Los Colorados area (D. bifidus Zone), respectively. The statistical analysis highlight the paleobiogeographic relationship between north-western Argentina and south-western China during the middle-upper Floian (Baltograptus cf. B. deflexus and D. bifidus Zones), and it supports the hypothesis that during the Early Ordovician north-western Argentina was located in the middle to high latitudes, included in the cold water faunal realm.     

Ordovician Calcified Algae and Cyanobacteria, Northern Tarim Basin Subsurface, China

Boreholes between Kuqa and Korla, in the northern Tarim Basin, Xinjiang, penetrated Ordovician marine limestones at depths of 5–6.2km. From three boreholes 54 out of a total of 170 limestone samples contain calcified algae, cyanobacteria and associated Microproblematica. Calcified cyanobacteria (GirvanellaBotomaella, ?Subtifloria) account for 37 per cent of occurrences; green algae (Dasyporelleae and Vermiporella) 28 per cent; Microproblematica (NuiaBevocastriaRothpletzellaHalysis) 20 per cent; and ‘solenoporaceans’ 15 per cent GirvanellaNuia are common in the Early Ordovician deposits, and ‘solenoporaceans’ are abundant in the Mid‐Ordovician. Dasyporelleae and Vermiporella are most abundant in Mid–Late Ordovician samples. Calcified cyanobacteria are common throughout the limestone succession, but particularly in the Mid Ordovician part. MoniliporellaContextaPlexaTexturata, and Villosoporella, hitherto placed in the supposed red algal family Moniliporellaceae Gnilovskaya, are here regarded as dasycladalean green algae. Despite some omissions, this Tarim flora broadly resembles others from Kazakhstan, Baltica and North America, indicating the generally cosmopolitan nature of Ordovician calcified algae and cyanobacteria.     

Discovery of Anticostia uniformis from the Xiazhen Formation at Zhuzhai,South China and its stratigraphic implication

The Xiazhen Formation is an Upper Ordovician lithostratigraphic unit in the Jiangshan-Changshan-Yushan (JCY) area, which contains series of Late Ordovician reef successions. The reef successions of the Xiazhen Formation at Zhuzhai are critical for evaluation of the Late Ordovician marine diversity and palaeoecology. However, their age has long been uncertain and generally is regarded as of upper Katian, based on the occurrences of shelly fossils and correlation with the stratigraphic equivalent Changwu Formation. The newly discovered graptolite species Anticostia uniformis, in the Xiazhen Formation, together with the combined evidence of brachiopods and sedimentology, indicates an age range for the graptolite locality from the Dicellograptus complanatus Biozone to the Diceratograptus mirus Subzone of late Katian, but the graptolites do not rule out the possibility that it is Hirnantian.     

Early–Mid Ordovician Yangtzeella (Syntrophiidina, Brachiopoda) and its evolutionary significance

Abstract:  Re-examination of newly collected topotype material confirms that the type species of Yangtzeella , Y .  poloi , a widespread Early and Mid Ordovician syntrophiidine brachiopod in South China, has a finely costellate shell rather than being smooth as previously thought. Thus, the subgenus Yangtzeella ( Vadimella ) Nikitina et   al ., established on the basis of fine costellae, is invalidated. Among 15 species of Yangtzeella , five species are recognized as valid based on multivariate analyses: Y .  poloi , Y .  unsulcata , Y .  songziensis , Y .  kueiyangensis and Y .  igori , among which the type species was the oldest known. Six are synonymized: Y .  septata , Y .  reticulata , Y .  lensiformis , Y .  depressa , Y .  yichangensis and Y .  minuta . Four are rejected from Yangtzeella : Y .  extensa , Y .  similior , Y .  yohi and Y .  poloi var. minor . Regional biostratigraphy indicates that Yangtzeella first appeared in a relatively deep-water setting on the Lower Yangtze Platform (South China palaeoplate) during late Tremadoc time ( Scolopodus warendensis conodont biozone) and then expanded to the deeper Jiangnan Slope as well as to the shallower Upper Yangtze Platform. The genus experienced two episodes of heightened abundance and diversity on the Upper Yangtze Platform during late Dapingian and mid Darriwilian times, respectively. Outside South China, Yangtzeella occurs sporadically in a few microplates or terranes, such as Tarim, Chu-Ili (southern Kazakhstan) and Taurides (southern Turkey) during the Dapingian and Darriwilian. Worldwide, Yangtzeella became extinct by the end of the Darriwilian.     

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 base of the Middle Ordovician in China with special reference to the succession at Hengtang near Jiangshan, Zhejiang Province, southern China

Extensive work during the last decade has led to the recent decision by the International Subcommission on Ordovician Stratigraphy that the base of the Middle Ordovician Series should be placed at the base of the Baltoniodus triangularis Conodont Biozone in the Huanghuachang GSSP (Global Stratotype Section and Point) section near Yichang, Hubei Province, China. A review of the biostratigraphy of successions in many parts of China shows that for various reasons, it is currently difficult to recognize the precise boundary level in many regions, and additional studies are clearly needed. A newly exposed, previously poorly known, condensed section in deeperwater facies at Hengtang near Jiangshan in the Chiangnan (Jiangnan) belt has yielded a substantial number of important graptolites and conodonts through the boundary interval. It provides a more informative illustration of the relations between the ranges of several key taxa of these groups than any other section in China, and the level of the base of the Middle Ordovician appears to be within an about 1.8 m thick interval of Ningkuo Formation, and lies in the lower Isograptus caduceus imitatus Biozone.     

Early heliolitine tabulate corals from the Sandbian (Upper Ordovician) in the Yunnan-Sichuan border area,Southwest China

Heliolitine tabulate corals are extremely rare in pre-Katian (Upper Ordovician) rocks worldwide, hindering the understanding of their origin and early evolution. Here, we systematically study new heliolitine material from the upper member (Sandbian) of the Huadan Formation in the Yunnan-Sichuan border area, Southwest China, which includes Protaraea ningnanensis, Plasmoporella kiaeri, Wormsipora sp. and Heliolites spinosus. The findings indicate that Protaraea and Heliolites are ancestral forms of coccoserids and typical tubular heliolitines, respectively, and that the absence of trabecular septa is probably a primitive feature of plasmoporellids. The refined Ordovician stratigraphy of this area implies that the coccoserid Protaraea ningnanensis Lin, 1965 from the lower member (late Darriwilian) of the Huadan Formation is the earliest known heliolitine, which further indicates that all heliolitines might have evolved from Protaraea. The comparison of Protaraea with other early tabulate corals also suggests an independent heliolitine origin from auloporinids and sarcinulinids.     

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.