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.     

An Early Ordovician Trilobite assemblage from the Lashkarak Formation, Damghan area, northern Iran

A low diversity Lower Ordovician trilobite assemblage is described from the middle part of the Lashkarak Formation at Simeh-Kuh, northwest of Damghan, northern Iran. Co-occurrence with conodonts of the lowermost Paroistodusproteus Biozone suggests an uppermost Tremadocian age. This assemblage includes four taxa, including one new genus and two new species, Damghanampyxginteri nov. gen. nov. sp. and Asaphellusfecundus nov. sp. The third diagnosable species from the formation is Taihungshaniamiqueli (Bergeron, 1894) which has previously been reported from southern France and the Turkish Taurides, suggesting north Gondwanan faunal affinities.     

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.     

Palynostratigraphy of Middle Cambrian to lowermost Ordovician stratal sequences in the High Zagros Mountains, southern Iran: Regional stratigraphic implications, and palaeobiogeographic significance

A palynological investigation of Cambro-Ordovician stratal sequences in the High Zagros Mountains of southern Iran permits the definition of a series of successive acritarch assemblage zones of chronostratigraphic significance, much improving the current knowledge of the Lower Palaeozoic stratigraphy of this important area for oil exploration. The five acritarch assemblage zones can be readily correlated with previously established palynostratigraphic schemes constrained by co-occurrence of independent age evidence, confirming the utility of organic-walled microfossils for the detailed biostratigraphic characterization of sedimentary units. The proposed biozonation will facilitate accurate dating of the southern Iranian Cambrian sequences during future drilling of deep test oil wells. Acritarch assemblage zone I (Middle Cambrian), occurs at the base of Member C of the Mila Formation; assemblages zone II (late Middle to earliest Late Cambrian) extends through the middle and upper part of the same lithostratigraphic unit; zone III (early Late Cambrian in age) characterizes the lower part of the Ilebeyk Formation; zone IV (middle Late Cambrian up to Cambrian/Ordovician transitional levels) occurs in the middle and upper part of the Ilebeyk Formation; finally, acritarch assemblage zone V ranges through the basal part of the Zardkuh Formation and proves an early Tremadocian age for the latter unit. The Mid-Late Cambrian acritarch associations show a marked Avalonian palaeobiogeographical affinity, also sharing a high proportion of taxa with typical Baltican and North Africa–Gondwanan assemblages; on the other hand, they are clearly different from known Laurentian (North America) fossil microphytoplankton suites. These results are in general agreement with current palaeogeographical models which place Avalonia, Baltica, and the North African part of Gondwana, all at relatively high southern palaeolatitudes, in contrast with the sub-equatorial position of Laurentia. However, the presence of many typical “Avalonian” taxa in the Iranian Mid-Late Cambrian assemblages would suggest a closer position of Iran to Avalonia than currently envisaged. The observed breakdown of acritarch biogeographic differentiation in earliest Ordovician times possibly represents a major disruption of oceanic current patterns and a lessened palaeolatitudinal thermal gradient.     

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).     

Exceptionally preserved soft parts in fossils from the Lower Ordovician of Morocco clarify stylophoran affinities within basal deuterostomes

The extinct echinoderm clade Stylophora consists of some of the strangest known deuterostomes. Stylophorans are known from complete, fully articulated skeletal remains from the middle Cambrian to the Pennsylvanian, but remain difficult to interpret. Their bizarre morphology, with a single appendage extending from a main body, has spawned vigorous debate over the phylogenetic significance of stylophorans, which were long considered modified but bona fide echinoderms with a feeding appendage. More recent interpretation of this appendage as a posterior “tail-like” structure has literally turned the animal back to front, leading to consideration of stylophorans as ancestral chordates, or as hemichordate-like, early echinoderms. Until now, the data feeding the debate have been restricted to evaluations of skeletal anatomy. Here, we apply novel elemental mapping technologies to describe, for the first time, soft tissue traces in stylophorans in conjunction with skeletal molds. The single stylophoran appendage contains a longitudinal canal with perpendicular, elongate extensions projecting beyond hinged biserial plates. This pattern of soft tissues compares most favorably with the hydrocoel, including a water vascular canal and tube feet found in all typical echinoderms. Presence of both calcite stereom and now, an apparent water vascular system, supports echinoderm and not hemichordate-like affinities.     

Tremadocian (Ordovician) biostratigraphy and graptolites at Dayangcha (Baishan,Jilin, NE China)

Based on a systematic study of Tremadocian graptolites from the Early Ordovician Yehli Formation at Dayangcha, Baishan of Jilin, NE China, a revised graptolite zonation is proposed for the Tremadocian rocks in the area. In ascending order, the graptolite zones include theRhabdinopora flabelliformis parabola Zone, theAnisograptus matanensis Zone, thePsigraptus jacksoni Zone and theAorograptus victoriae Zone. The first three zones are discussed with their definitions and durations clarified, whereas the last zone is introduced for the first time as a replacement of the formerAdelograptus-Clonograptus Zone. The entire Yehli Formation is considered to be older than the late Tremadocian Hunneberg Substage of Scandinavia, the latter being represented by the carbonate-dominated basal part of the overlying Liangchiashan Formation where no graptolites have so far been discovered. The Tremadocian succession in the Dayangcha area is regarded as being representative of the North China (Sino-Korea) Palaeoplate, and a correlation of the Tremadocian graptolite sequence with those of other continents is suggested. Nine species belonging toDendrograptus, Aspidograptus, Callograptus andAirograptus of the family Dendroidae, and eight species ofPsigraptus, Adelograptus, Aorograptus andKiaerograptus of the family Anisograptidae are described and some are revised in the present paper. Among the important revisions are the reconstructed proximal development ofPsigraptus jacksoni based on partially three-dimensional specimens from the area, and the complex thecal structures ofAspidograptus.      

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.     

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.     

BILLENGSELLIDE AND ORTHIDE BRACHIOPODS: NEW INSIGHTS INTO EARLIEST ORDOVICIAN EVOLUTION AND BIOGEOGRAPHY FROM NORTHERN IRAN

Abstract:  The eastern Alborz Mountains of Iran comprise a significant peri-Gondwanan terrane relevant to the early evolution of late Cambrian – early Ordovician brachiopods incorporated into the emerging benthic biota of the Paleozoic Evolutionary Fauna. A low diversity brachiopod assemblage from the late Tremadocian unit of the Lashkarak Formation contains six new species including the polytoechioideans Polytoechia and Protambonites and the orthoideans Paralenorthis, Ranorthis, Tarfaya and Xianorthis . The fauna preserves the earliest records of Polytoechia , unknown previously outside Laurentia and the Uralian margin of Baltica, and of Paralenorthis and Ranorthis , which were widespread along Gondwanan margins and in Baltica from the Floian (Arenig), plus Xianorthis , known hitherto only from the Floian of South China. The enigmatic Tarfaya has an impunctate shell fabric and setigerous perforations along the posterior margin, indicating placement within the Orthoidea in a new Family Tarfayidae. New species of Polytoechia , Protambonites , Paralenorthis, Ranorthis , Tarfaya , Xianorthis are described.