Trilobite biofacies and sequence stratigraphy: an example from the Upper Ordovician of Oklahoma

There have been surprisingly few empirical investigations of the fundamental principle that the architecture of depositional sequences exerts considerable control on observed patterns of faunal distribution and replacement. In this paper, we examine trilobite associations in two sequences of the Upper Ordovician (Sandbian) Bromide Formation of southern Oklahoma. Cluster analysis and ordination of genus abundance data identified five lithofacies‐related biofacies that are also differentiated by diversity patterns. Biofacies of the transgressive system tract (TST) of successive sequences are more similar to each other than they are to biofacies in the highstand systems tract (HST) of the same sequence. This similarity likely records dominance of large, robust convex sclerites in taphonomically degraded samples from condensed, strongly winnowed grainstone and rudstone. Horizons with articulated exoskeletons of isoteline trilobites preserved by obrution deposits occur most commonly in the early HST and record behavioural aggregations. Grainstone and rudstone of the later HST are less winnowed than those of the TST and show less fragmentation and sorting of sclerites. These changes in taphonomic conditions preserve ecological patterns more clearly. In most biofacies, rarefied alpha diversity (samples) and gamma diversity (biofacies) of middle‐ and outer‐ramp HST deposits are greater than in the TSTs, and biofacies replace each other down ramp. Diversity patterns do not agree with model predictions and other data sets that indicate low beta and high alpha diversity in the TST, likely because of taphonomic degradation. Vertical replacement of biofacies is expressed by the appearance of peritidal facies in which trilobites are rare. Biofacies shifts also characterize sequence boundaries and are most profound in the inner‐ramp successions characterized by sharp facies offsets. Comparison with bathymetrically similar deposits in the Taconic foreland basin showed similar diversity trends along environmental gradients, with some differences in shallow‐water settings attributed to taphonomic differences.     

Paleoecology of Middle Ordovician stromatoporoid mounds in Vermont

Fossiliferous mounds of carbonate mud are a distinctive facies in the middle Chazy Group (Crown Point Formation) at Isle La Motte, Lake Champlain. The mounds are surrounded by bedded calcarenite of spar-cemented pelmatozoan debris. Channels which cut into the mounds during mound growth are filled with the same calcarenite. The mud-free intermound rocks and the mound biota suggest agitated, normal marine shallow-water environments. The principal lime-secreting organisms within the mounds are stromatoporoids, calcareous algae, tabulate corals, sponges, and bryozoans. Each mound is dominated in terms of biomass by one of three groups: stromatoporoids, calcareous algae, and bryozoans. Most of the mound biota first appear at the base of the Crown Point Formation. In the lower Crown Point Formation the organisms increase in number and species. Both changes in the biota are related to periods of shallowing of the Chazy sea which are also reflected in the character of the carbonate sands.     

Upper Ordovician chondrichthyan‐like scales from North America

Studies of Ordovician micromeric fish scales from the Sandbian of North America have identified a number of scale‐based taxa potentially referable to the chondrichthyans and therefore can be among the stratigraphically oldest representatives of the clade described to date. Two of these, Tezakia hardingensis gen. et sp. nov. and Canyonlepis smithae gen. et sp. nov., are formally described herein. Tezakia gen. nov. scales are composed exclusively of tubular dentine and possess polyodontocomplex crowns with a characteristically large primordial odontode. Similar scale crown architecture has been reported only in the reputed chondrichthyan Altholepis composita (Lower Devonian of Podolia, Ukraine), and on these grounds, the two are united within the newly erected Altholepidiformes ordo nov. Multiple odontocomplexes are also a feature of Canyonlepis gen. nov. scale crowns; however, the latter do not demonstrate prominent primordial odontodes and are supported by a base composed of acellular bone. Additional data suggest that both taxa possess a combination of characteristics (areal crown growth, scale symmetry, linear odontocomplex architecture and absence of enamel, osteons, cancellous bone and hard‐tissue resorption) previously documented to occur only in chondrichthyan scales. This study contributes to a growing body of evidence that reveals the presence of diverse tissue types (bone, tubular and atubular dentine) and morphogenetic patterns (odontocomplex and non‐odontocomplex type of scale crown growth) in the dermal skeleton of putative Ordovician chondrichthyans.     

Borings in trepostome bryozoans from the Ordovician of Estonia: two ichnogenera produced by a single maker, a case of host morphology control

The evolution of borings has shown that the morphology of borings is a function of both the borer and its substrate. This study investigated the effect of bryozoan internal skeletal morphology on the dimensions and distribution of borings. One hundred and forty-three trepostome colonies from the Middle and Upper Ordovician strata of northern Estonia were examined. Of these, 80% were matrix entombed, longitudinally sectioned ramose and hemispherical colonies, and 20% were matrix-free hemispherical colonies that allowed examination of the colony surfaces. Seventy-one percent of the ramose colonies were bored, whereas 88% of the hemispherical colonies were bored. On average, only 8% of colony surface areas were bored out. Borings were more randomly oriented in the hemispherical colonies. In contrast in the ramose colonies, the borings tended to more restricted to the thin-walled endozone and thus parallel to the branch axis. This is interpreted to be a function of the thick-walled exozones controlling to some extent where the borer could bore. Based on morphology, the borings in the hemispherical colonies are referred to Trypanites and those in the ramose colonies to Sanctum . Sanctum is revised to include two possible openings and to recognize that boring shapes were inherently constrained by the thick-walled exozones of the host bryozoan colonies. Both trace fossils were probably produced by a boring polychaete that used the tubes as domiciles.     

Borings in Quartzite Surf Boulders from the Upper Cambrian Basal Deadwood Formation,Black Hills of South Dakota

Shallow, semi-spherical borings occur in clusters with densities of 1–3.5/cm² in quartzite boulders and in vein quartz from the localized basal conglomerate of the Cambrian-Ordovician Deadwood Formation in the east-central Black Hills of South Dakota. Some borings are superimposed on primary but enigmatic semi-circular structures, 2.5–5 cm in diameter, which are soft-sediment trace fossils formed prior to lithification. The macroborings are the first to be recorded from quartzite and vein quartz. Host boulders were eroded from near vertically dipping Paleoproterozoic quartzites from several different stratigraphic units. The thin boulder conglomerate grades were abruptly changed to sandstone through a layer ≤2 m thick in the Marjuman transgression (regionally correlated to the Cedarina dakotaensis trilobite zone). This transgression occurred prior to the start of the globally recognized Upper Cambrian Paibian Stage. Physically similar rocky-shore settings are widely known from quartzite islands of Cambrian age in Wisconsin, Middle Ordovician age on Ontario's Manitoulin Island, Ordovician-Silurian age in Manitoba, and Devonian age in western Australia. Erosion of quartzite surf boulders of equal or larger size occurred in all those regions, but the Black Hills of South Dakota is the only region where borings in quartzite are documented.     

Upper Ordovician sequences of western Estonia

The Upper Ordovician (uppermost Caradoc-Ashgill) section of western Estonia consists of a series of seven open-shelf carbonate sequences. Depositional facies grade laterally through a series of shelf-to-basin facies belts: grain-supported facies (shallow shelf), mixed facies (middle shelf), mud-supported facies (deep shelf and slope) and black shale facies (basin). Locally, a stromatactis mud mound occurs in a middle-to-deep shelf position. Shallow-to-deep shelf facies occur widely across the Estonian Shelf and grade laterally through a transitional (slope) belt into the basinal deposits of the Livonian Basin.

Each sequence consists of a shallowing-upward, prograding facies succession. Sequences 1 (Upper Nabala Stage) and 2 (Vormsi Stage) record step-wise drowning of underlying shelf units (lower Nabala) that culminated in the deposition of the most basinal facies (Fjäcka Shale) in the Livonian Basin. Sequences 3–6 comprise the overlying Pirgu Stage and record the gradual expansion of shallow and middle-shelf facies across the Estonian Shelf. The Porkuni Stage (sequence 7) is bracketed by erosional surfaces and contains the shallowest-water facies of the preserved strata. The uppermost part of the section (Normalograptus persculptus biozone) is restricted to the Livonian Basin, and includes redeposited carbonate and siliciclastic grains; it is the lowstand systems tract of the lowest Silurian sequence 8. Sequence 7 and the overlying basinal redeposited material (i.e., the lowstand of sequence 8) correspond to the latest Ordovician (Hirnantian) glacial interval, and the bracketing unconformities are interpreted as the widely recognized early and late Hirnantian glacial maximums.

The sequences appear correlative to Upper Ordovician sequences in Laurentia. Graptolite biozones indicated that the Estonian sequences are equivalent to carbonate ramp sequences in the western United States (Great Basin) and mixed carbonate-siliciclastic sequences in the eastern United States (Appalachian Basin–Cincinnati Arch region). These correlations indicate a strong eustatic control over sequence development despite the contrasting tectonic settings of these basins.     

Paleoecology of epizoans and borings on some Upper Cretaceous chalk oysters from the Gulf Coast

Analysis was made of a hard substrate fauna found on right valve interiors and exteriors of the epifaunal reclining oyster Pycnodonte mutabilis from the Maastrichtian (Navarroan) Saratoga Formation (southwestern Arkansas). Comparison of boring and encrustation patterns on both sides of valves indicates that a major portion of colonization on valve exteriors occurred while host oysters were alive. Paleoautecologic information derived from such valve exterior patterns includes evidence of rheotropic orientation by encrusting juvenile P. mutabilis and preferential location of Trypanites sp. borings in surficial shell grooves. Valve exteriors supported a hard substrate paleocommunity which had the following non-interactive progressive colonization sequence: (1) Trypanites sp. and P. mutabilis juveniles; (2) Entobia sp., serpulid worm tubes, and Bullopora sp.; and (3) cheilostome bryozoans. This sequence could have been caused by low seasonality and ranked success of colonizing encrusters and borers. Colonization of valve interiors generally differed from exteriors only in that many interiors were first colonized by the clionid sponge that created Entobia sp., which had already occupied the exterior, and which quickly bored through the valve to occupy the interior upon the host's death. □ Trace fossils, epizoans, borings, Gryphaeidae, palaeoecology, communities, colonization sequence, Late Cretaceous, Maastrichtian, Navarroan, Arkansas.     

贵州剑河八郎寒武系清虚洞组的楔叶形虫Sphenothallus Hall,1847

贵州剑河八郎松山寒武系黔东统中部清虚洞组主要由灰岩、泥岩组成,下部以灰岩为主,上部为泥岩、灰岩互层,顶部为中层白云岩。上部含有大量的褶颊虫类和掘头虫类三叶虫及带软躯体动物多门类化石组合。本文报道的楔叶形虫Sphenothallus Hall,1847产于清虚洞组上部,分异度低,仅1属1相似种,即台江楔叶形虫(相似种)Sphenothallus cf.taijiangensis Zhu et al.,2000,位于Redlichia(R.)guizhouensis-Redlichia(R.)nobilis顶峰带。S.cf.taijiangensis化石保存较好,少数标本管壁之间保存有横纹,说明Sphenothallus为一具有锥管状外壳的虫体。剑河八郎松山清虚洞组的楔叶形虫是贵州继牛蹄塘组、凯里组之后寒武系另一个产出层位,它的发现不仅丰富了清虚洞组生物群的组成,同时为全球楔叶形虫Sphenothallus的研究提供了新的材料。初步研究表明全球Sphenothallus的演化趋势是个体增长,管壁加厚物宽度变宽,口部膨胀角变小。Sphenothallus与现代刺胞动物水螅虫类或钵水母有密切关系。     

An opportunistic Upper Ordovician trilobite assemblage from Missouri

The trilobite Ampyxina bellatula (Savage) occurs in monospecific accumulations within dolomitic strata of the shaly Upper Ordovician Maquoketa Group of northeastern Missouri. A dolomitic stratum containing 97 complete individuals and considerable amounts of pyrite exhibited no associated benthic megafauna. The strata appear to be equivalent to the Scales Formation of the Maquoketa Group of northern Illinois and eastern Iowa which have been interpreted as having been deposited in a muddy environment with severe circulation restrictions. The presence of the monospecific trilobite association in what otherwise would be an unfossiliferous stratigraphic sequence is interpreted as being the result of opportunistic strategy by this trilobite in a resource-rich unstable environmental setting. The association was apparently cat-astrophically buried, thus preserving the age structure of the holaspid population. A size-distribution histogram of the holaspid population indicates that reproduction may have occurred in periodic bursts with lesser reproductive events occurring during the intervening periods. Holaspid survivorship information indicates that mortality was low during the early holaspid stage and increased with age. Anomalous single bed trilobite accumulations of various age rocks have been documented by numerous authors. These may in part be analogous to the Ampyxina occurrence in the Maquoketa Group of Missouri, and be attributable to opportunistic strategies.     

Katian (Upper Ordovician) conodonts from Wales

Middle and upper Katian conodonts were previously known in the British Isles from relatively small collections obtained from a few localities. The present study is mainly based on 17 samples containing more than 17 000 conodont elements from an approximately 14‐m‐thick succession of the Sholeshook Limestone Formation in a road cut near Whitland, South Wales, that yielded a diverse fauna of more than 40 taxa. It is dominated by representatives of Amorphognathus, Aphelognathus/Plectodina and Eocarniodus along with several coniform taxa. Representatives of Decoriconus, Istorinus and Sagittodontina are reported from the Ordovician of UK for the first time. The fauna is a typical representative of the British Province of the Atlantic Realm and includes a mixture of taxa of North American, Baltoscandic and Mediterranean affinities along with pandemic species. Based on the presence of many elements of Amorphognathus ordovicicus and some morphologically advanced specimens of Amorphognathus superbus, the Sholeshook Limestone Formation is referred to the lower A. ordovicicus Zone. Most of the unit is also coeval with Zone 2 of the Cautleyan Stage in the British regional stage classification, and stage slice Ka3 of the middle Katian Stage in the global stratigraphical classification, an age assignment consistent with data from trilobites, graptolites and chitinozoans. The unusually large collection of M elements of Amorphognathus provides insight into the complex morphological variation in this element of some Katian species of this genus. The Sholeshook conodont fauna is similar to those of the Crûg and Birdshill limestones, but differs in several respects from the slightly older ones from the Caradocian type area in the Welsh Borderland. Although having some species in common, the Sholeshook conodont fauna clearly differs from coeval Baltoscandic faunas and is even more different in composition compared with equivalent North American Midcontinent faunas.     

新疆柯坪和甘肃平凉上奥陶统底部附近的牙形刺

新疆柯坪大湾沟和甘肃平凉官司庄剖面是Nemagraptusb gracilis带及上奥陶统底界的重要剖面,研究这两剖面的牙形刺具有十分重要的意义。本文描述了采自新疆柯坪大湾沟和甘肃平凉官司庄剖面上奥陶统底界附近的牙形刺共13属18种。     

Epithelial moulds from some Upper Ordovician acrotretide brachiopods of Ireland

Three new genera of acrotretid brachiopods are of general morphological interest in that they display polygonal mosaics on the internal shell surfaces, features which are believed to be moulds of the outer epithelial cells. The mosaic is further noted in a specimen of Eoconulus , a form which may also belong to the Acrotretacea. Such mosaics have not previously been recorded from the Acrotretacea. Polygonal mosaics, epithelial moulds, outer epithelium , Eoconulus, Acrotretacea, Ordovician.     

The graptolite correlation of the North American Upper Ordovician Standard

The North American Upper Ordovician reference standard, the Cincinnatian Series, contains rich shelly microfossil faunas in its type area in the Cincinnati Region but graptolites are uncommon in most of its shallow-water calcareous sediments. Consequently, the graptolite correlation of this key sequence has remained uncertain, in part, even controversial. A review of both previously published recently discovered graptolite Occurrences in the type Cincinnatian, combined with data from the important graptolite successions in Oklahoma New York-Quebec, has not only clarified the graptolite correlation of the Cincinnatian but also added new data on the morphology taxonomy, the vertical horizontal distribution, of several taxa. The information now at hand indicates that the Edenian Stage correlates with the C. spiniferus Zone, the Maysvillian Stage with the C. pygmaeus lower middle P. manitoulinensis Zone, the Richmondian Stage with the upper P. manitoulinensis , the D. complanatus , possibly part of the C. inuiti Zone. Comparison between graptolite conodont biostratigraphic evidence reveals no apparent conflict. Correlations are proposed between Upper Ordovician North American stages, graptolite conodont zones, successions in Texas, Oklahoma, Sweden, European graptolite zones, British series.     

The trace fossil Thalassinoides from the Upper Ordovician of Tuva

Traces of Thalassinoides (the tunnels of unknown burrowing organisms) are described from carbonates of the Khondelensky layers of the Upper Ordovician of Tuva. Hitherto, this fossil was unknown in the Ordovician of the USSR. They demonstrate great similarity with Thalassinoides from coeval deposits of the Great Basin, USA. The traces are assumed to have been made in terrigenous-carbonate sediments deposited on the areas of a gently sloping shelf in quiet water, below wave base. □ Upper Ordovician, terrigenous-carbonate sediments, burrows.     

Functional morphologies of the columns of Upper Ordovician Xenocrinus and Dendrocrinus

Two common Upper Ordovician crinoids, Xenocrinus and Dendrocrinus , had distinctive columns that showed marked contrasts of both form and function. Xenocrinus baeri (Meek) had a tetragonal facet geometry but functioned in the normal manner for a symplectially articulated column. Column flexure was by bending, the four directions of curvature being limited by the tetragonal arrangement of crenulae. In contrast, columns of Dendrocrinus casei Meek, with a characteristic pentastellate symmetry, were able to twist, a functional adaptation not previously reported from crinoids; this was controlled by the unusual geometry of the crenularium. This suggests that some or all crinoid columns may be subject to twisting stresses, perhaps associated with changes in the current direction, that crenulae may resist passively. Functional morphology, column, crinoids, X enocrinus , D endrocrinus     

Stratigraphic and paleogeographic distribution of the Upper Ordovician bryozoans of Mongolia

The taxonomic diversity and distribution of the Upper Ordovician bryozoans of Mongolia are analyzed. Five heterochronous assemblages of bryozoans are established: Chigertei and Tsagaan del in the Sandbian Stage and Bairim Ovoo, Uuregnur, and Sairin in the Katian Stage. The bryozoans are shown to be important for solving complex problems of stratigraphy of the Ordovician of Mongolia and for paleogeographic reconstructions.     

Foliomena Fauna (Brachiopoda) from the Upper Ordovician of Sardinia

The late Ordovician brachiopod assemblage from Sardinia is one of the youngest members of the deep-water Foliomena fauna and is characterized by the following core taxa: Christiania , Cyclospira , Dedzetina and Foliomena . The fauna also contains Epitomyonia , Leangella , Glyptorthis and Skenidioides , which are more typical of shallower-water environments during the late Ordovician but occupied deeper-water niches during the Silurian following the termination of the Foliomena fauna. The suprafamilial placement of the family Chrustenoporidae is discussed and the new species Dedzetina serpaglii and Leangella ( Leangella ) fecunda are established. In common with many mid-Ashgill Foliomena faunas the Sardinian assemblage shows significant differences from other faunal developments of this type, reflecting its geographical position and shallower water conditions than those of the classic early Ashgill Foliomena faunas. The brachiopods occur with abundant trilobites belonging to a variant of the cyclopygid fauna. The faunas developed on part of a complex of microcontinents derived from peri-Gondwana during the Ordovician.     

Magnetostratigraphy susceptibility of the Upper Ordovician Kope Formation,northern Kentucky

As a test of the utility of magnetic susceptibility (MS) measurements for correlation among lithified marine sequences, samples were collected from well-studied shale and limestone outcrops of the Upper Ordovician Kope Formation, northern Kentucky. The bulk (initial) low-field MS of these samples is compared among two litho- and biostratigraphically correlated sections containing the same beds. The results of this comparison demonstrated an excellent correlation among equivalent beds. Thermomagnetic susceptibility analysis supports previous XRD work indicating that illite is primary and is the main paramagnetic mineral responsible for the MS variations observed in this study. A composite of MS results for a sequence of 31 named shale–limestone couplets is used to build a composite section of Kope sequences for the area. These MS variations permit division of the various sections into a set of 28 MS cycles within the Kope Formation, covering ∼ 50 m of section. These cycles suggest a systematic control by climate on the influx of detrital material that formed these sedimentary sequences, and support other work indicating Kope climate cyclicity.     

New data on the morphology of Sphenothallus Hall: implications for its affinities

Van Iten, H., Cox, R. S. & Mapes, R. H. 1992 04 15: New data on the morphology of Sphenothallus Hall: implications for its affinities. Lethaia , Vol. 25, pp. 135–144. Oslo. ISSN 0024–1164.
Recent speculation on the phylogenetic relationships of Sphenothallus Hall, 1847 has focused on two alternatives: (1) affinity with hydrozoan or scyphozoan cnidarians, or (2) affinity with annelids or other 'worms'. We have found that some species of Sphenothallus formed branching clonal colonies, and that others produced a thin transverse wall, similar to the conulariid schott. Sphenothallus tests are composed of carbonate apatite and are built of numerous, extremely thin lamellae (< 1 μm) that parallel the surface of the test. The holdfast, long interpreted as consisting of a pair of nested cups, actually consists of a single closed, broadly conical expansion floored by a thin basal membrane. Many thecate hydrozoan and scyphozoan polyps form branching clonal colonies and produce a thin transverse wall, similar to that produced by Sphenothallus . Further, thecae of coronatid scyphozoans are built of submicron-thick lamellae that parallel the outer surface of the theca, and coronatid thecae possess a closed, broadly conical apical expansion that serves as a holdfast. No such combination of characteristics exists among annelids or other non-cnidarian taxa. The recent discovery of paired tentacles in Sphenofhallus from the Early Devonian Hunsrück Slate by Fauchald et al ., thought to indicate that Sphenorhallus was wormlike, does not militate against a hypothesis of affinity with cnidarians. We therefore favour the hypothesis of a cnidarian affinity for Sphenothallus. * Sphenothallus, vermiform problematica, phylogenetic inference, coloniality, Cnidoria, Hydrozoa, Scyphozoa, Annelida, conulariids, byroniids, Bear Gulch Limestone .