The trace fossil Sinusichnus from the Upper Ordovician of Anticosti Island,eastern Canada

Branched, sinusoidal burrows assigned to Sinusichnus cf. seilacheri Knaust et al. 2016; occur in Upper Ordovician (Hirnantian) carbonates on Anticosti Island. These are abundant and monospecific at the base of the Oncolite Bed of the uppermost Ellis Bay Formation, marking a regional discontinuity surface during the multiphase Ordovician mass-extinction event. In contrast to Mesozoic and Cenozoic records of Sinusichnus, commonly attributed to the activity of decapod and isopod crustaceans, the burrow systems from Anticosti Island were probably produced by other arthropods of unknown affinity. A combined dwelling, locomotion and feeding behaviour of their trace maker is assumed. This is the oldest record of the ichnogenus Sinusichnus, previously only known since the Triassic, and its first evidence from North America.     

Island benthic assemblages: With examples from the Late Ordovician of Eastern Australia

The Benthic Assemblage (B.A.) concept, developed by A.J. Boucot two decades ago principally for continental marine margins, is extended to offshore island settings. Characteristics of modern island biotas, including ecological displacement, and the effects of r and K selection, can be identified in Late Ordovician volcanic islands of central New South Wales. Two variants of B.A.1 are represented, a quiet‐water lingulide biofacies, and a rough‐water rhynchonellide biofacies. The quiet‐water Eodinobolus biofacies occupied a B.A.1–2 position, onshore of and sheltered by a Tetradium wave‐baffle (B.A.2). The offshore shelfal high diversity strophomenide biofacies is equivalent to B.A.3. Remnants of periplatformal B.A.4–5 communities are recognised in allochthonous limestone breccias which were displaced downslope into graptolitic B.A.6 sediments. Steeper offshore gradients, typical of islands, laterally compress the B.A. profiles, and also contribute to downslope slumping. Ecological displacement in island environments results in extension of the habitat range of species into adjacent B.A.s. Reefs complicate the usual B.A. profile by introducing distinct sheltered and turbulent water environments. These characteristics may have applicability in interpretation of islands throughout the Palaeozoic record.     

Late Ordovician-Early Silurian fluctuations in sea level from eastern Anticosti Island, Quebec

Over 1,000 m of Upper Ordovician to Lower Silurian mixed carbonate and clastic strata on Anticosti Island are nearly tectonically undisturbed, despite their proximity to the Northern Appalachians fronting Quebec's Gulf of St. Lawrence. Natural cliffs exposed along the coast and rivers in the eastern part of the island make a relatively conformable sequence belonging to the Ashgill and Llandovery Series. Fossil communities interpreted as depth-associated in life are especially repetitious in the Becscie, Gun River, Jupiter, and Chicotte Formations (Llandovery Series), and to a lesser degree in the Upper Vaureal and Ellis Bay Formations (Ashgill Series). Preliminary study of the pattern of changeovers in Eocoelia, Pentamerus and Stricklandia communities suggests that Anticosti seas deepened and shallowed three and a half times during the Early Silurian. High water peaks were reached during B₁-B₂, C₁-C₂ and C₄-C₅ times, with a final deepening trend beginning in late C₅ time. Age determinations of these events are based on the occurrence of graptolites (with some new records from Anticosti) keyed to the standard graptolite zones, and species of the Eocoelia lineage are also useful for correlation. The profile of the Anticosti sea-level curve compares well with other curves reconstructed from the Lower Silurian of New York, Michigan, and Iowa. Widespread synchronism in sea-level changes on the North American platform is thus corroborated.     

A Curious Rusophycus (Arthropod Ichnofossil) Assemblage from the Upper Ordovician of Ontario,Canada

The ichnogenus Rusophycus includes a wide range of short bilobate excavations generally attributed to variable feeding behaviors of arthropods, especially trilobites. An unusual Rusophycus assemblage from Upper Ordovician Georgian Bay Formation in Ontario departs radically from previously described examples and presents new challenges for understanding the behavior represented by these traces. This specimen is unique in the arrangement of multiple Rusophycus burrows in a circular, lens-shaped array (as opposed to a linear or random arrangement typical of other Rusophycus assemblages). The size and shape of the individual Rusophycus components are consistent with traces attributed to the coeval trilobite Flexicalymene. Multiple Rusophycus assemblages likely reflect aggregations of trilobites in response to a local concentration of food. The topology of this particular Rusophycus assemblage suggests that the trilobites opportunistically exploited a rich and narrowly restricted food source, perhaps the decaying remains of a buried organism.     

Late Ordovician vertebrates from the Bighorn Mountains of Wyoming, USA

Abstract:  Late Ordovician vertebrate faunas occur in clastic sedimentary units along the length of the Rocky Mountains from Colorado to Montana, and across the border into Canada. Most research has, however, been conducted on localities in the southern part of the outcrop belt, particularly the Harding Sandstone Formation of Colorado. Micropalaeontological sampling of the coeval South Piney Member (Winnipeg Formation) in the Bighorn Mountains of Wyoming has revealed an abundant vertebrate fauna. The vertebrate assemblage includes a low-abundance fauna of 13 conodont taxa that together indicate an undatus Chronozone age (mid-Mohawkian; mid-Caradoc; Late Ordovician). The pteraspidomorphs Astraspis desiderata Walcott and Eriptychius americanus Walcott are also present together with one new taxon, Eleochera glossa gen. et sp. nov., which is interpreted as a derived stem-gnathostome on the basis of its scale histology and morphology. The fauna bears a strong similarity to that of the Harding Sandstone but is of lower diversity. In particular, it lacks the fine-grained, deeper water component of the Harding Sandstone that contains, inter alia , thelodonts and stem-chondrichthyans.     

Late Ordovician brachiopods from Peru and their palaeobiogeographical relationships

A Sandbian brachiopod association from the Calapuja Formation, in the Peruvian Altiplano, north‐west of Lake Titicaca, has allowed a re‐examination of the palaeobiogeographical relationships between Gondwana and Avalonia during the Late Ordovician, when the palaeocontinents are considered to be already very distant from one another. The brachiopod fauna includes the new species Onnizetina calapujensis sp. nov., Horderleyella chacaltanai sp. nov., Drabovinella minuscula sp. nov. and Tasmanella curtiseptata sp. nov., as well as Caeroplecia sp., Dinorthis cf. flabellulum and Tunariorthis cardocanalis. In addition, Colaptomena expansa expansa and Heterorthis retrorsistria, known from the British Burrellian Stage of the Caradoc Series (late Sandbian) in Wales and the Welsh Borderlands, have also been identified. The brachiopod collection is the most diverse known from a single locality in the whole Central Andean Basin. Within it, forms with clear Gondwanan links occur, such as the new species of Onnizetina, Drabovinella and Horderleyella, and typical representatives of the Avalonian faunas, such as the Welsh Colaptomena expansa expansa and Heterorthis retrorsistria. The brachiopod species exchange between the Proto‐Andean margin of Gondwana and Avalonia, now believed to be possible during the late Sandbian, allows a reconsideration of the global taxonomic affinities of both regions. With this in mind, detrended correspondence analysis (DCA) and cluster analysis have been applied to an updated rhynchonelliformean brachiopod matrix consisting of presence/absence data. The scatter plot resulting from the DCA allows a vivid visualization of the grouping and geographical trends of the South American localities with respect to Avalonia–Baltica and the Mediterranean margin of Gondwana during the Sandbian. Our results agree with previous palaeogeographical reconstructions, depicting Avalonia very close to Baltica and already distant from Gondwana. As a few brachiopod species, with low dispersal potential, would have been able to migrate between those distant palaeocontinents, the existence of intermediate islands in the Rheic Ocean, permitting the transit by island hopping of eurythermal species, must be considered.     

Restudy of some Ordovician–Silurian boundary graptolites from Anticosti Island, Canada, and their biostratigraphic significance

The Ellis Bay Formation on Anticosti Island has long been recognized for the biostratigraphic importance of its latest Ordovician conodont, palynomorph, and shelly fossil assemblages. However, a sparse record of graptolites has made it difficult to correlate these assemblages with the graptolite biozonation. Restudy of some previously described and examination of newly collected normalograptid specimens from the Ellis Bay and lower Becscie formations, however, shows that biostratigraphically important taxa are present and these species provide important constraints on the age of the strata. Normalograptus parvulus and N. minor have been identified in the upper half of the Ellis Bay Formation, suggesting that these strata are late Hirnantian in age ( N. persculptus Biozone). Normalograptus imperfectus and Normalograptus sp. aff. N. acceptus occur in the basal beds of the Becscie Formation, indicating that these strata are earliest Silurian. These graptolite data support the hypothesis that the positive carbon isotope excursion seen in the uppermost strata of the Ellis Bay Formation is isochronous with that seen in the Hirnantian strata at Dob's Linn, Scotland, and does not span much of the lower to mid-Hirnantian, as is the case in Arctic Canada and Nevada, USA.     Anticosti Island , Hirnantian, graptolites, Ordovician, Silurian.     

Huemacaspis (Trilobita, Kerfornellinae) from the Late Ordovician of the Argentine Cordillera Oriental

The trilobite genus Huemacaspis P?IBYL and VANěK, 1980 is redescribed from well-preserved material from the Santa Gertrudis Formation (early Caradoc), Argentine Cordillera Oriental. The new species H. gallinatoensis is described and figured. Huemacaspis is widely distributed in the southern part of the Central Andean basin during the early Caradoc and it is a key taxon for the correlation of different units from the Cordillera Oriental and Sierras Subandinas of Argentina and the Cordillera Oriental of Bolivia. This taxon is one of the few records of trilobites of early Caradoc age found in the shallow shelf environments of southwestern Gondwana. Huemacaspis is accommodated within the Kerfornellinae, a subfamily otherwise known from southern Central Europe and northern Africa. The trilobite fauna from the early Caradoc of the Central Andean basin suggests a faunal exchange with the latter regions indicating the persistence of a broadly similar biogeographic pattern to that documented in earlier Ordovician times.     

Facies development of a Late Ordovician mixed carbonate-siliciclastic ramp proximal to the developing Taconic orogen: Lourdes Formation, Newfoundland, Canada

The Upper Ordovician (late Whiterockian to Mohawkian) Lourdes Formation represents a narrow (tens of kilometers), short-lived [∼5–7 million years (my)], open-ocean (high-energy) mixed siliciclastic-carbonate ramp that onlapped allochthonous strata along the orogen side of the local Taconic foreland basin. Platform development followed a 6–8 my hiatus during which weathering had concentrated chemically mature siliciclastics that were admixed with initial carbonate sediments. A cross-platform facies gradient contains paleokarst and peritidal carbonates and sandstones, shallow-ramp carbonate bioherms and skeletal shoals, and deeper ramp calcareous shales. Transgressive systems tracts are marked by ramp-wide sheets and shoals of skeletal grainstone and low accumulation rates, and highstand systems tracts are marked by significant admixture and interbedding of siliciclastics with cross-ramp carbonate facies. Platform demise coincides with increased siliciclastic input, which is likely tectonically influenced. The Lourdes platform is equivalent to epicontinental foreland ramps along eastern Laurentia, but its narrower width precluded formation of oceanographically restricted platform-interior facies.     

Tabulate Corals after the Frasnian/Famennian Crisis: A Unique Fauna from the Holy Cross Mountains,Poland

Famennian tabulate corals were very rare worldwide, and their biodiversity was relatively low. Here we report a unique tabulate fauna from the mid- and late Famennian of the western part of the Holy Cross Mountains (Kowala and Ostrówka), Poland. We describe eight species (four of them new, namely ?Michelinia vinni sp. nov., Thamnoptychia mistiaeni sp. nov., Syringopora kowalensis sp. nov. and Syringopora hilarowiczi sp. nov.); the whole fauna consists of ten species (two others described in previous papers). These corals form two assemblages—the lower, mid-Famennian with Thamnoptychia and the upper, late Famennian with representatives of genera ?Michelinia, Favosites, Syringopora and ?Yavorskia. The Famennian tabulates from Kowala represent the richest Famennian assemblage appearing after the F/F crisis (these faunas appear some 10 Ma after the extinction event). Corals described here most probably inhabited deeper water settings, near the limit between euphotic and disphotic zones or slightly above. At generic level, these faunas show similarities to other Devonian and Carboniferous faunas, which might suggest their ancestry to at least several Carboniferous lineages. Tabulate faunas described here represent new recruits (the basin of the Holy Cross mountains was not a refuge during the F/F crisis) and have no direct evolutionary linkage to Frasnian faunas from Kowala. The colonization of the seafloor took place in two separate steps: first was monospecific assemblage of Thamnoptychia, and later came the diversified Favosites-Syringopora-Michelinia fauna.     

黔北上奥陶统宝塔组头足类多样性初探*

上奥陶统宝塔组(桑比阶–凯迪阶)是华南分布广泛的标志性地层单元, 以发育特殊的网纹状灰岩并产出大量头足类化石为特征。黔北桐梓县尧龙山地区的宝塔组厚度较大, 出露极好, 产出数量丰富、保存精美和特征典型的头足类化石。本文在研究区酒店垭、集镇采坑和天域极地3个宝塔组剖面采集到大量原域埋藏的头足类化石, 共8属11种及少量属种未定类型, 以直形壳或盘形壳、体管很小且适合远洋深水游泳的类型为主, 整体上符合扬子台地深水陆棚Sinoceras–Michelinoceras–Disoceras生物相特征。基于壳体扩大率、壳径与体管直径比、气室高与隔壁颈长比、隔壁下凹度、壳径与气室高比等参数变量的主成分分析表明, 在外部特征不显著的情况下, Sinoceras和Michelinoceras的直观差别是壳体扩大率, 而Michelinoceras属不同种的差异主要体现在气室高与隔壁颈长之比, 其他特征的区分度并不明显。层面上单位面积的化石定量统计显示, 尧龙山地区宝塔组中头足类的多样性较高, 并以Sinoceras和Michelinoceras具有极高的个体丰度为特征。扬子板块在奥陶纪桑比期–凯迪期之交处于高水位的陆表海环境, 栖息环境均一化和生态系统复杂化可能是促进该时期头足类动物的复苏和多样性演化的重要原因。     

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.     

Influence of valve geometry, ornamentation, and microstructure on fractures in Late Ordovician brachiopods

Among Late Ordovician brachiopods from southeastern Indiana. strophomenids display a ratio of 4:1 parabolic to linear repaired fractures in contrast to the 1:2 ratio found for orthids and rhynchonellids. Additionally, only strophomenids display repaired elliptical fractures. The weakest parts of strophomenid and orthid-rhynchonellid shells are the regions of the adductor muscle scars and the sulcus, respectively. Fractured biplanate shells of strophomenids are commonly cleaved anteriorly to posteriorly, whereas fractures are localized on the anterior of strongly curved to geniculate conspecific specimens. Rugae on leptaenids, thickened anterior margins of the brachial valve of rafinesquinids, and the dense distribution of pseudopunctate in all strophomenids, functioned to localize anteriorly the (un)repaired linear and parabolic fractures. No sublethal fracture occurs on any biconvex shell where the height is greater than 14 mm, despite the fact that numerous specimens of certain species attained shell heights of 20 mm or more, an observation suggesting the upper limit in the gape of the crushing elements of the predator. Crushing experiments on valve 'models' reveal that the inflated equibiconvex, plicate shape of the shells of Plarysfrophia is the strongest. However, the architecture of the concave strophomenid valves is relatively stronger than the corresponding valves of many orthids and rhynchonellids when normalized for valve thickness.     

Late Ordovician carbon isotope trend in Estonia, its significance in stratigraphy and environmental analysis

Carbon isotope changes during most of Late Ordovician time (from the mid-Caradoc Kinnekulle K-bentonite until the beginning of the Silurian) were investigated. As the corresponding sequence of rocks is stratigraphically nearly complete in Estonia, an attempt was made to use it to elaborate the general pattern of carbon isotope changes in the Late Ordovician. Complications were caused by several local or regional hiatuses in the middle and late Caradoc and Hirnantian. A total of 385 whole rock samples were studied from eight drill cores in northern and central Estonia. The following positive carbon isotope events were observed: (1) the mid-Caradoc excursion (peak δ¹³C value 2.2‰) in the uppermost part of the Keila Stage, also known in Sweden; (2) the first late Caradoc excursion (1.9‰) in the lower part of the Rakvere Stage; (3) the second late Caradoc excursion (2.4‰) in the upper part of the Nabala Stage; (4) the early Ashgill excursion (2.5‰) in the lowermost part of the Pirgu Stage; (5) the widely known large Hirnantian excursion (in Estonia the peak value reaches 6.7‰) in the Porkuni Stage. The study interval comprises a long (10 Ma) period characterized by low-magnitude carbon isotope changes and a following brief (2 Ma) interval with large changes. No obvious lithological preference for hosting the positive shifts was recorded. In principle, the δ¹³C values exceeding the background values may occur in all types of rocks present in a sedimentary basin. Several δ¹³C positive excursions (values 1.5‰ to 3‰) in the Mohawkian of North America are evidence that the minor Caradoc and early Ashgill δ¹³C positive shifts in Baltoscandia may have counterparts in Laurentia. If correctly correlated, these shifts may have global significance. The Hirnantian excursion is usually linked to a major glacial event, even if some carbon cycling mechanisms are not completely understood. The environmental causes suggested for the earlier minor shifts range from global climatic and glacial events to very local changes in basin regime and sea level. Our study supports the primary role of climatic or climatically triggered oceanic processes.     

Traces and related chemical changes in a Late Ordovician paleosol,Glossifungites ichnofacies,southern Appalachians,USA

Traces interpreted as Skolithos sp. in a Glossifungites ichnofacies occur in the upper part of the Beans Gap Claystone paleosol, in association with a Late Ordovician marine erosional surface. Paleosol color and whole‐rock chemistry are distinctly different in the vicinity of burrows, which are enveloped by olive gray mottles, 1 to 10 cm in diameter, in which total iron (measured as Fe₂O₃) is 30% lower than in grayish red, “unaltered”; claystone.

Post‐pedogenic bioturbation increased permeability locally and permitted the introduction of marine fluids and coarser sediment into the top of the paleosol. Anaerobic microbial decay of organic matter in the burrows and surrounding claystone permitted reduction (and mobilization) of formerly oxidized iron, which then combined with marine‐derived sulfate to form pyrite, both within burrows and along oxidation‐reduction fronts in the claystone proximal to burrows. Pyrite was later converted to hematite as a result of outcrop‐related oxidation by meteoric water.

Paleosols formed in coastal marine settings are especially susceptible to early diagenetic alteration related to marine hydromorphism. Differentiating these ichnologic and diagenetic effects from true pedogenic ones is essential when interpreting paleosols in paralic sequences.     

Evolutionary and biogeographical shifts in response to the Late Ordovician mass extinction

The Late Ordovician mass extinction was an interval of high extinction with inferred low ecological selectivity, resulting in little change in community structure after the event. In contrast, the mass extinction may have fundamentally changed evolutionary dynamics in the surviving groups. We investigated the phylogenetic relationships among strophomenoid brachiopods, a diverse brachiopod superfamily that was a primary component of Ordovician ecosystems. Four Ordovician families/subfamilies sampled in the analysis (Rafinesquinidae, Strophomeninae, Glyptomenidae and Furcitellinae) were reconstructed as monophyletic groups, and the base of the strophomenoid clade that dominated the Silurian recovery was reconstructed as diversifying alongside these families during the Middle Ordovician. We time‐calibrated the phylogeny and used geographical occurrences to investigate biogeographical changes in the strophomenoids through time with the R package BiogeoBEARS . Our results indicate that extinction was higher in taxa whose ranges were constrained to tropical or subtropical regions. Furthermore, our results suggest important shifts in the diversification patterns of these brachiopods after the mass extinction. While most of the strophomenoid families survived the Late Ordovician event, ecologically abundant taxonomic groups during the Ordovician were either driven to extinction, reduced in diversity, or slowly died off during the Silurian. The new abundant strophomenoid taxa derived from one clade (consisting of Silurian–Devonian groups such as Douvillinidae, Strophodontidae and Amphistrophiidae) that diversified during the post‐extinction radiation. Our results suggest the selective diversification during the Silurian radiation, rather than selective extinction in the Late Ordovician, had a greater impact on the evolutionary history of strophomenoid brachiopods.     

Fustiglyphus annulatus from the Ordovician of Ontario,Canada, with a systematic review of the ichnogenera Fustiglyphus Vialov 1971 and Rhabdoglyphus Vassoievich 1951

The ichnogenera Rhabdoglyphus Vassoievich 1951 and Fustiglyphus Vialov 1971, both a source of taxonomic confusion and inconsistent usage, are re‐examined. Many specimens of Fustiglyphus have been incorrectly described as Rhabdoglyphus, both prior and subsequent to the formulation of the former by Vialov in 1971. Updated synonymies for both ichnogenera and their ichno‐species indicate that two ichnospecies of Rhabdoglyphus can be distinguished, namely R. grossheimi Vassoievich 1951 and R. spinosus Ksiazkiewicz 1977, while F. annulatus Vialov 1971 is the only acknowledged ichnospecies of Fustiglyphus. The significant feature distinguishing the two ichnogenera is regarded as the presence of invaginated segments on Rhabdoglyphus as opposed to well‐defined rings or knots with no invagination on Fustiglyphus.

Although Spongolithus annulatus Fritsch 1908 is technically the senior synonym of Fustiglyphus annulatus, it is recommended that the latter name continue to be used in the interests of maintaining nomenclatural stability. Previously unreported specimens of Fustiglyphus annulatus from the Ordovician of Ontario are described and illustrated; one of these suggests that the swellings in Fustiglyphus possibly represent brood chambers of the producing‐organism(s). We also suggest that Fustiglyphus "retreated”; from onshore to deep‐water offshore environments at the end of the Paleozoic.     

A new problematic Early Ordovician univalve mollusc from France

Lamaureriella vizcainoi gen. et sp. nov., from the Montagne Noire region of southern France, is a coiled, laterally compressed, bilaterally symmetrical mollusc in which prominent folds in the apertural margin produce a characteristic spiral plication across each lateral surface. Similar plications are present in technophorid rostroconchs but the prominent coiling and lack of a pegma suggest that Lamaureriella is a helcionelloid. It is the second such genus described from Early Ordovician rocks.     

A Revision of Early To Mid Ordovician Hyoliths from Sweden

Hyolith assemblages of early to mid Ordovician age in Sweden include the hyolithids Hyolithus bisulcatus Holm, H. concinnus Holm and H. innotatus Holm, which are confirmed as representatives of Hyolithes Eichwald. Dorsolinevitus Syssoiev is known from D. dispar (Holm), D. textilis (Holm) and D. vomer (Holm), and Carinolithes Syssoiev is represented by C. hospes (Holm) and C. triumvir (Holm). Sulcavitus caelatus (Holm), the type species that forms the basis for a family and even an order according to some, is redescribed; the order is suppressed. Hyolithus cymbium Holm is transferred to Stelterella gen. nov., and Theca crispata (Boll) is now the type species of Crispatella gen. nov. The unusual orthothecid Quadrotheca quadrangularis (Holm) is redescribed; the concept of Trapezotheca aemula (Holm) is emended, and Semielliptotheca dens (Holm) is transferred to Trapezotheca . Semielliptotheca Syssoiev is poorly known from only one species, S. rosmara (Holm), and that genus is now considered to be unrecognizable. Hyolithus exaratus Hadding is identified as closely related to the gastropod Lytospira norvegica Koken, and specimens of Hyolithus sp. nos 11, 12 and 13 Holm and Hyolithus sp. indet. Wiman, remain to be located.