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We describe early Tremadocian (Kainella meridionalis Biozone) cephalopods from the Cordillera Oriental, Jujuy, NW Argentina. They consist of numerous small specimens collected at the Quebrada de Arenal, Trancas section, near the town of Tilcara, in the Alfarcito Member of the Santa Rosita Formation. All but three specimens were assigned to a new species of Ellesmeroceras (Family Ellesmeroceratidae), E. humahuacaensis sp. nov., based on its slightly endogastric curvature, the characteristics of the siphuncle and chambers dimensions. Micro CT scanning of one specimen aided in the description of the apex and facilitated the construction of a 3D model of the species. A single, similar specimen was assigned to Ellesmeroceras sp. pending the availability of additional material. Two specimens differ from the rest, being exogastric with a lower angle of expansion. They are tentatively assigned to Bassleroceras sp. This material indicates that Cambrian and early Tremadocian cephalopods are not as different as previously thought. “Diversification” and “extinction” events during the late Cambrian may be attributed to taxonomic “over-splitting” and taphonomic and/or sampling biases, respectively. These specimens are currently the oldest recorded in the Central Andean Basin and of West Gondwana, and probably represent the first migration of cephalopods into the region, when the water column was still poorly colonized. During the middle Tremadocian, subsequent immigrations and originations of several cephalopod orders accounted for a rise in diversity and expansion into new niches during this interval. Some of these taxa persisted into the middle Floian, at which time, a second increase in diversity is recorded. Ellesmeroceras humahuacaensis sp. nov. is interpreted as a sub-vertical nektobenthic organism.

Graptolites from the early Tremadocian Anisograptus matanensis Biozone are identified from east-central Sonora, northwestern Mexico, within a carbonate-shelf succession deposited on the southwestern continental shelf margin of Laurentia. This is the second occurrence of typical anisograptid graptolites in Mexico, after its original record in the Oaxaca area, which belongs to a Gondwana-related paleogeographical realm. The graptolites from Sonora represent the second global occurrence of any member of the Rhabdinopora flabelliformis group in the Ordovician equatorial shelf region.

Tremadocian highly metalliferous black shales and associated grey shales from the Aseri PH012B drill core (NE Estonia) in the innermost part of the Baltic Palaeobasin were targeted to record their nitrogen and organic carbon isotope variance combined with the total organic carbon and total nitrogen record. The obtained molar C/N ratios of black shales from 26 to 52 indicate a considerable loss of N compared to primary biomass. The recorded δ15N values from â2.5 to 0.2â° likely evolved due to isotopic fractionation related to N2 fixation by primary producers, superimposed by later anoxic ammonium oxidation processes within the uppermost sediments. The high net primary productivity, which controlled the accumulation of organic-rich shallow-water complexes, was fuelled by the internal cycling of P in the sea basin and combined with intensive N exchange between marine and atmospheric pools.

The Erdaopuzi section in the Baishan area of Jilin Province, northeast China, contains abundant late Tremadoc graptolites. However, there are some taxonomic controversies regarding the characters of the hand specimens that are difficult to identify. In this paper, we present our restudy of isolated graptolites from acid residue and discuss the taxonomic problems of the subgenus Clonograptus (Neoclonograptus) Zhao and Zhang. Four genera and six species were identified, including Adelograptus tenellus (Linnarsson), Ancoragraptus gracilis (Zhao and Zhang), Dictyonema sp., Psigraptus jacksoni Rickards and Stait, Psigraptus lenzi Jackson, Psigraptus arcticus Jackson. According to the significant evolution of morphological characters, two graptolite zones (from bottom to top) can be identified in ascending order in the study area, i.e. Adelograptus Zone and the Psigraptus Zone.

Solutans are relatively common members of echinoderm assemblages in the Saint-Chinian Formation (upper Tremadocian) of the Montagne Noire (France). The revision of all available material confirms that most specimens can be assigned to Minervaecystis vidali, which is here reconstructed for the first time. The occurrence of Plasiacystis mobilis is evidenced based on a single, well-preserved dististele. The interpretation of the small-sized individual as a putative dendrocystitid is rejected: it corresponds to a juvenile specimen of M. vidali. Early Ordovician solutans from the Montagne Noire partly fill the gap between Laurentian midâlate Cambrian syringocrinids and Avalonian-Gondwanan EarlyâMiddle Ordovician taxa.

A block of stromatolitic limestone found on the Angara River shore near Kodinsk, Siberia, derived from the exposed nearby Ust-kut Formation, has yielded a sample of 146 ellesmeroceratid nautiloid specimens. A minor contribution to the fossil assemblage from bellerophontid and hypseloconid molluscs suggests a restricted abnormal salinity environment. The associated shallow-water low diversity assemblage of the conodonts Laurentoscandodus triangularis and Utahconus(?) eurypterus indicates an age close to the Furongian–Tremadocian boundary. Echinoderm sclerites, trilobite carapaces, and hexactinellid sponge spicules were found in another block from the transitional strata between the Ust-kut and overlying terrigenous Iya Formation; these fossils indicate normal marine salinity. The conodont L. triangularis is there associated with Semiacontiodus iowensis and Cordylodus angulatus. This means that the stromatolitic strata with cephalopods are older than the early Tremadocian C. angulatus Zone but not older than the Furongian C. proavus Zone. The sample of nautiloid specimens extracted from the block shows an unimodal variability in respect to all recognizable aspects of their morphology. The material is probably conspecific with the poorly known Ruthenoceras elongatum from the same strata and region.

Abstract In this research, Ordovician conodonts obtained from the sedimentary succession of the informal Katkoyeh formation is studied in a stratigraphic section at the south of Kuh-e-Bonorg, eastern flank of Kalmard Anticline in the west of Tabas (Central Iran). The formation which is partly equivalent to the succession of the Shirgesht Formation in Tabas Block is unconformably overlaid the Kalmard Formation (attributed to the Neoproterozoic) and disconformably overlain by the Carboniferous Gachal formation. The Katkoyeh formation with 214.8 meter thickness is divided into three lithostratigraphic members including the Lower Sandstone (mostly quartzarenite to litharenite), Middle Shale (red to green shales interbedded with some dolomitic and calcareous sandstones, in laminated wavy stromatolite structure at the base) and Upper Carbonate (limestones, dolostones and marls). The section was sampled bed by bed for conodont studies so that 80 samples have been selected, however only seven of them were productive. Within the selected samples, six genera and 14 species of coniform and ramiform conodont elements are determined, a few of them are reporting for the first time from Iran. The obtained conodonts demonstrate the faunal assemblages equal to the proteus, elegans-evae and diprion Zones which confirm the age of Early Ordovician (late Tremadocian–Floian) for the studied section. The whole conodont material display a faunal affinity with the Baltoscandic and Argentina paleobiogeographic provinces. Keywords: Conodont, Floian, Katkoyeh formation, Ordovician, Tremadocian. Introduction In spite of the moderately -exposed outcrops of the Ordovician successions throughout the country, most of the stratigraphical studies in Iran are focused on Alborz Range (e.g.; Gansser and Huber 1962, Muller 1973, Ghavidel-Syooki 2006, Ghobadi Pour et al. 2007, 2011; Jahangir et al. 2016) and the appropriate data from the other regions are restricted to a few published reports (e.g.; Ruttner et al. 1968, Aghanabati 1977, Hamedi 1995, Ghaderi et al. 2008). Especially our knowledge is very little about the conodont contents of the Ordovician rocks in Iran. The Ordovician sedimentary succession in the north of Tabas in central Iran have been introduced as the Shirgesht Formation by Ruttner et al. (1968). The rock sequences of this formation composed of brown, cream, and green to red limestones, shales and partly sandstones with hardly more than 800 m in thickness (Ghobadi Pour et al. 2006). Lithostratigraphically, the Ordovician sedimentary interval in adjacent Kalmard Block (Aghanabati 2010) is different, includes more siliciclastic rocks somewhat carbonate beds in upper part. These succession has been considered as the Katkoyeh formation by Hamedi and Wright (1992). The less known Katkoyeh formation as an informal stratigraphic rock unit in central Iran has been regarded to Arenigian to Ashgillian in the type area in Kerman region, but the age of the formation in the Kalmard type region is dissimilar in different studies. In the current research we have looked for the succession Katkoyeh formation in a section at the south of Kuh-e-Bonorg, eastern flank of the Kalmard anticline, from the age dating view, based on the conodont contents. Material & Methods In the Kuh-e-Bonorg section, 80 rock samples were collected bed by bed for conodont content while only seven of them were productive. For this purpose, sandy limestones, dolomitic limestones and pure limestones with about 3–5 kg were dissolved by 10% formic acid and 20% acetic acid following the standard procedure for conodont extraction (e.g., Jeppsson & Anehus, 1995). Residue washed on appropriate clean 18 and 230 mesh sieve-stack (0.85 to 0.063 mm). The extracted conodont collections stored in the Ferdowsi University of Mashhad, Iran under the collection name with the FUM#MC prefix. Discussion of Results & Conclusions The Katkoyeh formation in the south of Kuh-e-Bonorg section is unconformably overlaid the Neoproterozoic rocks attributed to the Kalmard Formation and disconformably overlain by the Carboniferous Gachal formation. The Katkoyeh formation of 214.8 meter thickness in the studied area is divided into three members such as the Lower Sandstone (mostly quartzarenite to litharenite), Middle Shale (red to green shale interbedded with some dolomitic and calcareous sandstone, in laminated wavy stromatolite structure at the base) and Upper Carbonate (limestones, dolostones and marls). Among the seven fossiliferous beds, we obtained the index conodonts of Lower Ordovician which belong to six genera and 14 species of coniform and ramiform elements. The assemblage is correlative with the same collections from the Baltoscandic and Argentina paleobiogeographic provinces (e.g.; Mannik and Virra 2012; Voldman ‌‌‌‌‌‌et al. 2017). The distribution of taxa in the stratigraphic succession is variable, in abundance of elements as well as in the number of taxa. The oldest conodonts in the section, Drepanodus arcuatus and Drepanoistodus costatus, come from the dolostones and dolomitic limestones of uppermost Tremadocian-early Floian proteus Zone (equivalent to Acodus apex Zone in NW Argentina Province) the sample MC208B. The next fossiliferous bed, MC232, comprises Acodus triangularis, Acodus cf. deltatus, Drepanoistodus cf. nowlani, Drepanoistodus cf. bassiovalis, Drepanoistodus bassiovalis and Gothodus costalatus which confirm Floian elegans - evae Zones (equivalent to Gothodus vetus–Gothodus andinus Zones in NW Argentina Province). The last productive beds, MC263–MC265, contain Trapezognathus diprion as the index taxon for upper evae Zone (equivalent to Trapezognathus diprion Zone in NW Argentina Province). This species confirm the age of late Floian for the uppermost part of the Katkoyeh formation in the Kuh-e-Bonorg section.

Marrellomorphs (class Marrellomorpha) are a group of Paleozoic arthropods with a very poor fossil record. Here we describe a new marrellomorph arthropod Mimetaster florestaensis sp. nov. from the Tremadocian (earliest Ordovician) of Argentina. The new species is characterized by the shape and direction of the three pairs of principal spines, and the existence of strong secondary spines only in the proximal two-thirds of the anterolateral spines. As a result of phylogenetic analysis the new species integrates a trichotomy with Mimetaster hexagonalis and a Moroccan unnamed marrellid as sister groups. This discovery increases the known diversity of Marrellomorpha and represents the first occurrence of this group in South America, expanding the spatial distribution of the clade.

The eoorthid brachiopod Apheoorthina is reported for the first time from the Lower Ordovician of NW Argentina. It is represented by a species similar to A. ferrigena from the Tremadocian of the Prague Basin, increasing the faunal affinities between the Central Andean Basin and the South European microcontinents, in particular the Bohemian region (Perunica). Nine out of the fourteen brachiopod genera reported from the Tremadocian of the Central Andean Basin (~64%) are shared with the Mediterranean region, four of which (~28%) have been recorded in the Prague Basin, and two (Kvania and Apheoorthina) are restricted to the Central Andes and Perunica. Dispersal pathways around Gondwana are analyzed in the light of major factors affecting large-scale distribution of brachiopods (environment, larval capacity for dispersal, oceanic currents). The presence in Apheoorthina aff. ferrigena of a well-preserved larval protegulum measuring 420 μm in width and 210 μm in length strongly suggests that this species had planktotrophic larvae capable of long-distance dispersal. According to recent ocean-atmosphere general circulation models for the Ordovician Period, the Central Andean margin was dominated by the cold-water Antarctica Current. Despite the complex non-zonal pattern produced by current deflections around the peri-Gondwanan microcontinents, the general westward circulation sense favoured larval dispersal from the Andean region to North Africa, Avalonia, the Armorican Terrane Assemblage, and Perunica. On the other hand, the eastwards flowing Gondwana Current connected the North Gondwana waters with the South American epicontinental seas, which could explain the reversed migration of some brachiopods.

The transition from the Cambrian to Ordovician in Morocco is known to be characterized by a frequent Furongian hiatus, restricted extension of the Tremadocian marine deposits, and frequent unconformities at the base of the transgressive upper Floian deposits. In the present work, we first highlight the occurrence of Fe- and mica-rich, red silty/sandy formations in the Central and Eastern High Atlas between the Middle Cambrian and Upper Floian sequences. In the Tislyt type-locality, a synsedimentary hemigraben structure is defined, within which the red beds show frequent slump folds, debris flows and internal unconformities. The correlation with several coeval series of the Meseta domain allows us to define a shallow marine, ferruginous clastic Atlas-Meseta Red Beds (AMRB) basin during the Tremadocian-early Floian. The AMRB basin extended between the Meseta coastal block and the Anti-Atlas domain, being limited by the fault zones that became the West Meseta shear zone and the South Meseta fault, respectively, in the Variscan orogen. The AMRB basin compares with the coeval rifted basins of the central Iberian and Armorican massifs. The red beds were likely sourced from the east, from both the Precambrian basement and Early Ordovician magmatic rocks, contrary to the Ordovician deposits of the Sahara platform sourced from the south. Subsidence of the AMRB and central Iberian-Armorican basins of the NW-Gondwana border aborted during the Floian, whereas the opening of the Rheic ocean went on more to the west.

Restudy of the Cambrian–Ordovician boundary beds, traditionally assigned to the Mila Formation Member 5 in Mila-Kuh, northern Iran, for the first time provides convincing evidence of the Early Ordovician (Tremadocian) age of the uppermost part of the Mila Formation. Two succeeding trilobite assemblages typifying the Asaphellus inflatus–Dactylocephalus and Psilocephalina lubrica associations have been recognized in the uppermost part of the unit. The Tremadocian trilobite fauna of Mila-Kuh shows close similarity to contemporaneous trilobite faunas of South China down to the species level, while affinity to the Tremadocian fauna of Central Iran is low. The trilobite species Dactylocephalus levificatus and brachiopod species Tritoechia tenuis from the Tremadocian of Mila-Kuh are new to science.

The high-resolution study of vertical geochemical variability of shallow-water Tremadocian black shales of the Türisalu Formation targeted two drill core sections from Suur-Pakri Island, NW Estonia. Altogether 374 samples from 4.6 m thick shale were analysed by XRF. The metalliferous and organic-rich black shales revealed significant centimetre-scale variation in the concentration of redox-sensitive trace metals – U, Mo and V. The V profiles show cyclic variations in half a metre- to metre-scale and the strongest correlation with loss on ignition (LOI) 500 °C (interpreted to reflect organic matter abundance). The abundance of Mo presents high values near the lower and upper contacts of black shale and otherwise moderate covariance with LOI. The distribution of U is not coupled with LOI, being characterized by irregular local enrichment anomalies in the profiles of both sections. This suggests that sequestration of U may have been time-dependent and possibly favoured by dissimilatory U-reduction at the sediment–water interface under iron-reducing conditions. Significant depositional variability of the studied organic-rich muds apparently supported dynamic physicochemical and biological microenvironments at the sediment–water interface and thus temporally and spatially diversified the paths and efficiency of synsedimentary redox-sensitive trace element enrichment.

Six linguliform and two rhynchonelliform brachiopods, including three new species Eurytreta ahmadii, Wahwahlingula kharbashi and Nanorthis bastamensis are described from Tremadocian strata (Paltodus deltifer deltifer conodont Biozone) in the Deh-Molla area southwest of Shahrud, Northern Iran. The fauna is dominated by micromorphic lingulides and acrotretides and shows distinct similarity to the contemporaneous micromorphic brachiopod association from Tremadocian chalcedonites of the Holy Cross Mountains, Poland. New data on the early ontogeny of the enigmatic lingulide Diencobolus show a very distinct pattern, including the presence of a metamorphic protegulum ornamented with flat-based pits and a single pair of larval setal bundles, which links this taxon to Paterula and suggests close phylogenetic relationships of both taxa to the Discinoidea.

New late Tremadocian (Early Ordovician) conodont and graptolite faunas from the eastern and central belts of the Eastern Cordillera (Jujuy Province, northwestern Argentina) are reported. The conodont fauna includes the guide species Paltodus deltifer pristinus, Paltodus deltifer deltifer, and Acodus deltatus (sensu lato), in association with other taxa, and the graptolites Aorograptus victoriae, Ancoragraptus cf. bulmani, and Adelograptus cf. altus. Overlapping ranges of the recorded species allow for a partial correlation between the Acodus deltatus- Paroistodus proteus and Aorograptus victoriae zones, and the Notopeltis orthometopa trilobite Zone. The conodont fauna includes a mixture of forms typical of the Baltoscandian and Laurentian provinces, respectively. The Early Ordovician basin of northwestern Argentina may correspond to the Shallow-Sea Realm and Cold Domain and probably records the development of a differentiated conodont province in the southern South American margin of Gondwana.

The Tremadocian graptolite and conodont faunas of the Gondwana Margin recorded at the Eastern Cordillera (EC) of Salta and Jujuy provinces (northwestern Argentina) are studied. The previous data and the new provided information indicate that a “no nominated interval” and several graptolite zones (Anisograptus matanensis, Rhabdinopora flabelliformis anglica, Bryograptus, Kiaerograptus, Kiaerograptus supremus, Araneograptus murrayi, and Hunnegraptus copiosus zones), as well as several conodont zones (Iapetognathus, Cordylodus angulatus, Paltodus deltifer, and Acodus deltatus - Paroistodus proteus zones) occur in the Ordovician successions studied. New or reappraised data from sections such as Angosto del Moreno, Angosto de Lampazar, Parcha-Incamayo-Incahuasi area, and Cajas range (EC western margin), and Alfarcito area and Mojotoro range (EC eastern margin), provide new information on the composition of the fossil faunas. A composite graptoliteconodont biostratigraphic framework, including records of key trilobites as external control group, is proposed for the Tremadocian of the South American margin of Gondwana. Both fossil groups are generally recorded from outer platform to ocean basin environments, showing significant similarities with the Baltoscandian faunas. Nevertheless, particular shallow-water facies include either specific forms or species associations that are common to the epeiric shallow-water seas of Laurentia. This fact demonstrates an important interplay between faunas of different regions of the Iapetus Ocean during the Tremadocian.