Late Ordovician extinction of North American and British crinoids

After reaching a diversity peak in the Caradocian, North American Ordovician crinoids underwent a gradual decline to a nadir in the early Ashgillian (Maysvillian). This interval, recording extinction of the Cleiocrinidae, Merocrinidae, Ottawacrinidae, Hybocystitidae, and several lineages of camerate crinoids, was apparently caused by major environmental shifts in seas of eastern North America resulting from a westward-prograding wedge of terrigenous clastics derived from the Taconic Highlands, possibly coupled with a marine transgression in the Maysvillian that allowed colder water slope biofacies to invade the craton. Crinoids suffered a major episode of extinction in the late Ashgillian (late Richmondian/Rawtheyan). This event, preceding the end of the Ordovician by at least one stage or 2 to 4 million years, resulted in extinction of 12 families of crinoids including the Xenocrinidae, Tanaocrinidae, Reteocrinidae, Archaeocrinidae, Anthracocrinidae, Cincinnaticrinidae, Iocrinidae, Anomalocrinidae, Carabocrinidae, Cupulocrinidae, Porocrinidae, and Hybocrinidae. Glacio-eustatic lowering of sea level may have triggered this crisis by partially draining the North American craton, resulting in changes in oceanic circulation, salinity, and temperature. Latest Ordovician (Hirnantian) carbonates of the North American mid-continent region contain pelmatozoan assemblages from which Silurian crinoids radiated. These taxa were largely unaffected by a minor extinction event at the Ordovician/Silurian boundary.     

The maintenance of evolutionary equilibrium in Late Ordovician benthic marine invertebrate faunas

An upper evolutionary limit on species equilibrium number (S of MacArthur & Wilson) has been postulated but never demonstrated. Results of the present study indicate that an S of about 10 was attained very rapidly in a Late Ordovician benthic marine invertebrate faunal succession. This S value was then maintained for perhaps five million years within a marine environment that did not remain uniform but changed progressively from relatively deep to shallow water. Most importantly, significant taxonomic and presumed trophic changes that accompanied the gradual habitat alteration seem to have had a negligible effect on the plateau-like limit to species equilibrium number. We conclude from these preliminary results that the maintenance of evolutionary equilibrium in an open ecosystem strongly suggests a long-term component of biotic resiliency, at least within this one marine ecosystem, and perhaps in many others. The reasons for this resiliency remain to be explored.     

Biogeographical analysis of Late Silurian brachiopod faunas, chiefly from Asia and Australia

Shallow water benthic marine invertebrates (such as brachiopods from Benthic Assemblages [BA] 1–3) have usually played a much more important role than deeper ones (BA 4–5 or even deeper) in evaluating biogeographical provincialism in geological history. The Silurian brachiopod Retziella Fauna, characterized by the common presence of Retziella in association with various provincial taxa and many common North Silurian Realm genera, is known from southwest Tienshan, North China, South China, North Vietnam, and East Australia. It is possibly also present in North Korea, the central Pamirs, Afghanistan, and New Zealand. The coeval Tuvaella Fauna occurs only in the southern marginal belt of the Siberian Plate. Synecologically, both faunas inhabited a normal, shallow-water, level-bottom environment, usually with a low-diversity community (commonly 3–8 genera); assignment to BA 2–3 is indicated. Their mutual exclusiveness is of biogeographical significance: subdivisions of the Uralian-Cordilleran Region can be based on them, with the Tuvaella Fauna being included in a redefined Mongolo-Okhotsk Province. A Sino-Australian Province is established and defined herein for the area occupied by the Retziella Fauna during the Ludlow-Pridoli and probably the Wenlock. Two subdivisions of the province can be recognized, a Sino-Central-Asian Subprovince and an Australian Subprovince, based on different endemic brachiopods and separate geographical positions. The presence of a number of more cosmopolitan genera in both the tropical-subtropical Sino-Australian and subtropical-temperate Mongolo-Okhotsk Provinces during the Late Silurian testifies to oceanic surface current circulation patterns adequate for the distribution of planktic larvae capable of long-distance dispersal while maintaining reproductive communication. This contrasts with the dispersal potential of endemic components of the newly defined Silurian biogeographical units. □ Late Silurian, Brachiopoda, Retziella , Fauna, Tuvaella Fauna, Synecology, Biogeography, Asia, Australia     

Influence of the Late Ordovician glaciation on basin configuration of the Yangtze Platform in China

The Late Ordovician paleogeographic changes in the Yangtze Platform indicate a widespread regression and consequent subaerial exposure probably caused by the 'locking up' of substantial amounts of marine waters during the Late Ordovician glaciation. The subsequent rapid paleogeographic changes on the Yangtze Platform during the earliest Silurian suggest a rapid melting of the ice. The relatively cool waters in the partially closed Yangtze basin during the latest Ordovician were replaced by open and warm waters during the earliest Silurian. Sedimentation rates and composition of faunal assemblages underwent marked changes during latest Ordovician and Early Silurian. Glaciation, Ordovician, Silurian, Paleogeography, sedimentary rate, faunal change.     

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.     

Analyzing niche stability and biogeography of Late Ordovician brachiopod species using ecological niche modeling

Gauging the potential impacts of environmental change on the geographic distributions of species is a central area of modern biogeographic analysis, often involving complex models of species–environment interactions. The geographic distribution of fossil species can also provide a framework to test the impact of environmental change on biogeography and ecological niches of species, yet few paleontological analyses have attacked this question in deep time. Herein we present a quantitative biogeographic analysis to examine the stability of ecological niches and geographic ranges of rhynchonelliform brachiopods during an interval of sea level change preserved in Upper Ordovician strata of the Cincinnati Arch.The intensive sampling, excellent preservation, and numerous prior paleoecological and sedimentological analyses within the tri-state region of Kentucky, Indiana, and Ohio provide a robust framework for detailed paleobiogeographic study. Quantitative biogeographic modeling methods incorporating GIS (Geographic Information Systems) are utilized in order to spatially analyze the geographic ranges of brachiopod species of the Corryville and Mt. Auburn Formations of the C3 (uppermost Maysvillian) depositional sequence.This study employs the ecological niche modeling program GARP (Genetic Algorithm using Rule-set Prediction) to predict the geographic distribution of eight brachiopod species during three time slices within the C3 sequence. This method estimates a species’ geographic range by modeling the ecological niche of the species based on a set of known species occurrence data coupled with environmental data inferred from sedimentologic proxies. Once environmental tolerances for a species are modeled; the species is predicted to occur wherever its preferred set of environmental conditions occurs within the study region.Distributional patterns were reconstructed for three time slices during the C3 sequence. Recovered range predictions were quantitatively analyzed for evidence of temporal range changes. Results indicate that average species range within the study area decreased and species tracked their preferred niche with high fidelity during the transition from the early to middle portions of the C3 depositional sequence, an interval of rapid relative sea level change. However, during the transition from the middle to late portions of the sequence, gradual shallowing within the basin and development of discontinuous habitat patches correlates with niche evolution of five of the eight species modeled. The average area a species occupied within the basin increased during this interval, but there is a mixed response including both increases and decreases in range size within the study group. In general, the species that exhibit niche evolution increased their geographic range size while those that continue to track their niche with high fidelity experience a decrease in geographic range size. During the latter half of the C3 sequence, previously continuous habitats become fragmented, thereby isolating individual populations and providing a mechanism for niche evolution. The rate of sea level change and the corresponding fragmentation of previously continuous habitats into isolated patches appear to be the primary controls on both mean geographic range size and relative degree of niche evolution.     

Bryozoan provinces and patterns of generic evolution and extinction in the Late Ordovician of North America

The Late Ordovician bryozoan genera of central and southeastern North America are geographically distributed in three biotic provinces, separated by boundaries reflecting major lithofacies differences. The central Cincinnati Province contains most of the North American endemic genera, and represents a narrow ecological zone separating the clastic wedges of the marginal Reedsville-Lorraine Province from the cratonic carbonate platform of the Red River-Stony Mountain Province. The provinces provided major life zones, or biomes, for each of the five bryozoan orders. Genera comprising the provinces differed as well in morphologic complexity, geochronologic survivorship, tiering, endemism and eurytopy. Regions on either side of the Cincinnati Province were dominated by inferred immigrants from Baltoscandia. Al-logenic provincial succession produced time-averaged mixed faunas in regions near the provincial boundaries. Although most generic originations took place within the Cincinnati Province, evolutionary novelties are associated with the Reedsville-Lorraine Province. The loss of the diverse Cincinnati Province, connected with global cooling and a eustatic lowering of sea level, may have been a chief factor in the Late Ordovician extinction of bryozoan genera. Genera from the Red River-Stony Mountain Province differentially survived into the Silurian.     

The late Sandbian – earliest Katian (Ordovician) brachiopod immigration and its influence on the brachiopod fauna in the Oslo Region, Norway

Bulk samples of brachiopods from the middle and upper parts of the Arnestad Formation and the entire Frognerkilen Formation in the Oslo Region demonstrate major vertical changes in the Sandbian to Katian amphicratonic fauna of this part of the Baltic Province. The main influx of new taxa occurred in the upper part of the Arnestad Formation (uppermost Sandbian), and in the succeeding Frognerkilen Formation (basal Katian). Faunal change is initially reflected in an increase in diversity and not as a replacement of the previously incumbent genera, which remain throughout the studied sequence. The majority of the new genera migrated from the East Baltic Region or through this region from Avalonia, while the marginal Laurentian taxa first occupied the Scandinavian part of Baltica. Six taxa migrated from Laurentia, three from Avalonia, two from Gondwana and one apparently originated on the South China palaeoplate. The faunal shifts and immigration of brachiopods appear to have been in response to an initial transgression together with the movement of Baltica into more warm temperate latitudes, but the diverse faunas developed both against a background of subsequent regression and in response to an offshore shift of biofacies. The new amphicratonic fauna was thus generated by faunal shifts from elsewhere in the Baltic Province to the marginal environments of the Oslo Region together with more exotic elements from neighbouring continents and microcontinents.     

Mesozoic brachiopod migrations and the opening of the North Atlantic

The distribution of a number of Mesozoic brachiopods indicates a migration up the eastern side of the North Atlantic starting in the early Jurassic and reaching as far north as Greenland by the beginning of Cretaceous times. The brachiopods evidently reached the new Atlantic via the Rif mountains of northern Morocco and migrated both north and south from there. Some of them came from the central High Atlas which represents a gulf open to the east but closed to the west. The western High Atlas, on the other hand, was populated by successive waves of forms from the west. Brachiopods and other distinctive forms of “Tethyan” origin clearly extended considerably farther north on the eastern side of the Atlantic than they did on the west. This probably implies an early establishment of the Gulf Stream. Some Tethyan brachiopods which did not extend to the western end of the Mediterranean are found instead to have spread via northeast Siberia to cordilleran North America.     

Punctal density in the Ordovician orthide brachiopod Paucicrura rogata: anatomical and paleoenvironmental variation

In the orthide brachiopod Paucicrura rogata , from the Upper Ordovician of New York, the density of punctae (small perforations in the shell) increases from about 350 to about 450 per mm² of shell in the transition from shallow-water limestones to black graptolitic shales down the outer slope of the Taconic Trench. Punctal density is significantly correlated with water depth, as measured by distance downslope along individual volcanic ash layers, and by reciprocal averaging score for macroinvertebrate fossil assemblages ( p < O.01). Because punctal density increases with temperature in living brachiopods, the results suggest that water temperature increased with depth in the Taconic basin, corroborating similar conclusions based on Paucicrura's oxygen-isotopic composition. The increase in punctal density with increasing water depth may also reflect reduced shell growth rates of P. rogata in oxygen-deficient bottom waters, causing a reduction in spacing between punctae. Ontogenetic decrease of punctal density with increasing shell size reflects differential rates of puncta formation at the shell margin. □ Brachiopoda, Orthida, Ordovician, Taconic Basin, paleoceanography, paleoecology, punctae, caeca.     

First record of a Late Proterozoic faunal assemblage in the North American Cordillera

Hofmann, Hans J. 1981 12 15: First record of a Late Proterozoic faunal assemblage in the North American Cordillera. Lethaia, Vol. 14, pp. 303–310. Oslo. ISSN 0024–1164. Megafossils are reported from an unnamed Late Proterozoic formation in the Sekwi Brook area of the Mackenzie Mountains of northwestern Canada. The biota includes the metazoans Inkylovia sp. and Sekwia excentrica Hofmann n, gen., n. sp., the trace fossils Gordia sp., Gordiu? sp., and Torrowangea sp., as well as structures of problematic affinities. The metazoans support a latest Proterozoic age for the containing beds. 0 Early Metazoa, body fossils, trace fossils, Problematica, Late Proterozoic, North American Cordillera, Markenzie Mountains, Canada.     

On Permian and Triassic insect faunas in relation to biogeography and the Permian-Triassic crisis

The taxonomic diversity dynamics of pterygote insects in the Permian and Triassic at the family/age level are considered. Different metrics of taxonomic diversity are compared. Biogeographic and taphonomic aspects of changes in the composition of insect faunas in the Permian and about the P-T transition are discussed. Some changes in the Permian insect faunas are of a biogeographic nature and do not indicate global changes in diversity. Insects with aquatic immatures were rather common in the Permian and Early Triassic, but these immatures are well represented in only few localities.     

The effects of the final stages of the Late Ordovician glaciation on marine palynomorphs (chitinozoans, acritarchs, leiospheres) in well Nl-2 (NE Algerian Sahara)

Palynomorph assemblages, especially chitinozoans and acritarchs, from the Upper Ordovician of well Nl-2 (north-east of the Algerian Sahara) are studied in order to precisely date the ultimate effects of the Late Ordovician glaciation and to document the impact of this major climatic stress on the diversity of the palynoplankton. References are made to stable isotope excursions and to global eustatic sea level variations in order to improve the local age determination. The Hassi el Hadjar Formation, i.e. glacio-marine diamictites, is interpreted as a transgressive event resulting from the melting of the northern Gondwana ice cap. It yields poorly preserved and moderately diverse chitinozoans of late Hirnantian age. Acritarchs are more abundant in the lower part of these "microconglomeratic clays", but display a low diversity and are badly preserved throughout the whole formation. Reworked individuals are recorded in both groups. The marine sediments of the M'Kratta Formation of latest Hirnantian age contain better preserved, more abundant and more diverse palynomorph assemblages, especially in the Upper Member. The composition of this palynoplankton indicates a fairly good faunal and phytoplankton recovery after the early Hirnantian climatic stress.The extinction of the Ordovician forms, and the appearance of Silurian type taxa occur only in the uppermost Hirnantian, i.e. following with a slight delay the glacial event. The overlying black shales of Wenlock age (lower part of the Oued Mehaiguène Formation) are indicative of marine anoxic environments. They yield a virtually exclusive, but enormous number of Tasmanacea. The very peculiar composition of this palynoplankton seems to be independent of the Late Ordovician glaciation and is most likely related to the factors that, later, generated and maintained anoxic conditions in this area.A new species of chitinozoan, Spinachitina oulebsiri sp. nov. from the latest Hirnantian M'Kratta Formation, is described and illustrated.     

Principal stages in the Cenozoic diversification of shallow-water molluscan faunas in the North Pacific

Cluster analysis of bivalve species recorded in Cenozoic deposits in Sakhalin Island, western Kamchatka, Hokkaido, and California was used to determine geological age of the modem North Pacific biogeographic region and its constituent subregions (Japan-Mandchurian, Beringian, and Oregon-Sitkan). The North Pacific region developed during the Paleogene-Neogene transition due to Drake Passage opening to deep-water movement, formation of the deep-water Antarctic Circumpolar Current, and the change in climate from greenhouse to psychospheric. Differentiation of the three subregions within the North Pacific Region seems to have occurred in late Miocene-early Pliocene, about 5.6 millions years ago and was probably due to the flooding of the Bering Land Bridge and development of the present configuration of circulation in the North Pacific. In the Northwest Pacific, during Paleogene and early Neogene, the faunal diversification occurred more rapidly and was more extensive than in the Northeast Pacific.     

The Middle Caradoc Facies and Faunal Turnover in the Late Ordovician Baltoscandian palaeobasin

A Late Ordovician episode of remarkable biotic, climatic, sea level and facies changes, named here as the Middle Caradoc Facies and Faunal Turnover, took place in the Baltoscandian area. This paper presents an integrated overview of these changes in the critical middle Caradoc interval. Data are given on carbonate rock composition, distribution and grain-size composition of the siliciclastic material and the carbon isotopic composition of whole-rock carbonates in cores of Estonia and Sweden.

The Middle Caradoc Facies and Faunal Turnover can be described as a succession of related environmental changes. The turnover began with a positive excursion in carbonate δ¹³C and continued with sea level changes that led to a sedimentary hiatus on the shelf and a change from carbonate-dominated to siliciclastic-dominated sedimentation in the basin. The turnover ended with an extinction event and associated microfaunal crisis.

The middle Caradoc turnover in Baltoscandia is comparable to a similar succession of changes in North America. The turnover affected two palaeocontinents, and reflects a widespread, possibly global environmental change. Onset of glaciation on Gondwana and/or increased orogenic activity might have initiated the changes in ocean circulation and led to the initial carbon isotope excursion. The following sea level rise and faunal changes affected several different continents.     

Variation of butterfly diet breadth in relation to host-plant predictability: results from two faunas

Host‐plant data for North American and Australian butterflies were used to test the hypothesis that larval diet breadth increases with decreasing resource predictability (where the latter was estimated by host‐plant growth‐form/duration). For each region in turn we compared the diet breadths of butterflies which utilise herbaceous host‐plants with those of species having woody hosts. For North America alone we also compared the diet breadths of species having annual hosts with those utilising perennial hosts, and the diets of species having herbaceous‐annual hosts with those using woody‐perennial hosts. Studies of diet breadth may be biased by the host taxonomic level which contributes most to the diet index used. For example, the results of analyses which employ indices based on numbers of families of hosts utilised may differ from those using indices based on counts of host species or genera. To investigate this potential problem we performed cross‐species analyses where diet breadth was defined in turn as the number of host species, genera, or families eaten. We found that using different taxonomic levels did give inconsistent results. To avoid this we employed phylogenetic diet breadth indices in comparative analyses of Independent Contrasts. The former incorporate information from the whole of the host‐plant phylogeny, whilst the comparative method eliminates any confounding effects of butterfly phylogeny. The results indicated that there is a phylogenetic component to butterfly diet breadth. They also largely differed from those of the cross‐species investigations, although there were similarities (i.e. results differed between regions and varied according to whether the whole fauna or just endemics were investigated). Our results suggested that in both regions, non‐endemics which feed on herbaceous plants have wider diet breadths than non‐endemics which utilise woody hosts. However, we found no consistent evidence that the diet breadths of endemics increase with decreasing resource predictability (as estimated here).     

Biogeochemistry of Late Paleozoic North American brachiopods and secular variation of seawater composition

Sr/Ca ratios in modern brachiopod shells reflect variations in ambient seawater, whereas their Na contents show no relationship with water depth or habitat. Their Mn and Fe contents are controlled, in part, by leaching of these elements from oxide coatings or the low input/sedimentation rate of detrital material into depositional areas such as Quatsino Sound. For most Carboniferous brachiopods from North America, the Mn and Fe contents are similar to those recorded by their Recent counterparts. The high Mn and Fe contents in the brachiopods from shales suggest several possibilities for these levels. One possibility is the leaching of Mn and Fe from oxide coatings/matrix which was not completely removed in the cleaning process, or the high levels in part reflect unusual depositional conditions (some degree of anoxia) for the local shaly environments. The Sr/Ca ratio of brachiopods and, by inference, complementary seawater, did not vary significantly during the Carboniferous. The Sr/Ca minimum observed in brachiopods of Mississippian age coincides with a dip in the ⁸⁷Sr/⁸⁶Sr curve and correlates with the Hercynian orogeny. This is attributed to the cycling of seawater through mid-ocean ridge basalts, and postulated exchange reactions account for variation in the composition of seawater-Ca. The unidirectional trend of heavier δ¹³C values from the Devonian to the Permian is intricately coupled with the evolution of the terrestrial biomass. In addition to expansion of terrestrial plants, burial of reduced carbon in the form of coal (organic matter) contributed to the observed shift. The start of the Permo-Pennsylvanian glaciation is marked by a negative excursion of the secular carbon trend, which is linked to weathering of reduced carbon and its return to the oceanic reservoir with its oxidized carbon. The oxygen isotope values reflect the unidirectional trend towards higher values of the carbon data with decreasing geologic age. Negative excursions of the trend may be related to extensive weathering of terrestrial and submarine rocks, whereas positive excursions may be related to hydrothermal alteration of submarine rocks and dehydration of oceanic crust during times of active sea-floor spreading. Oxygen-calculated water temperatures of unaltered brachiopod material are unrealistically high for all of the Devonian, and the Chesterian-Meramecian, Desmoinesian-Missourian, and Artinskian Epochs. During these times maximum water temperatures of 42° to 56°C are well above the thermal threshold of protein denaturation. This process, which is lethal to most higher organisms, demands an adjustment in oxygen of -2.5%. for samples older than Missourian, and of -1.250%. for samples spanning the Missourian-Artinskian interval. With these adjustments and salinity considerations made prior to calculations, water temperatures become reasonable for the Late Paleozoic epeiric, tropical seas of North America.     

Ecological biogeography of North American mammals: species density and ecological structure in relation to environmental gradients

Aim To evaluate the relationship of climate and physiography to species density and ecological diversity of North American mammals. Location North America, including Mexico and Central America. Methods Species density, size structure and trophic structure of mammalian faunas and nine environmental variables were documented for quadrats covering the entire continent. Spatial autocorrelation of species density and the environmental variables illustrated differences in their spatial structure at the continental scale. We used principal component analysis to reduce the dimensionality of the climatic variables, linear multiple regression to determine which environmental variables best predict species density for the continent and several regions of the continent, and canonical ordination to evaluate how well the environmental variables predict ecological structure of mammalian faunas over North America. Results In the best regression model, five environmental variables, representing seasonal extremes of temperature, annual energy and moisture, and elevation, predicted 88% of the variation in species density for the whole continent. Among different regions of North America, the environmental variables that predicted species density vary. Changes in the size and trophic structure of mammalian faunas accompany changes in species density. Redundancy analysis demonstrated that environmental variables representing winter temperature, frostfree period, potential and actual evapotranspiration, and elevation account for 77% of the variation in ecological structure. Main conclusions The latitudinal gradient in mammalian species density is strong, but most of it is explained by variation in the environmental variables. Each ecological category peaks in species richness under particular environmental conditions. The changes of greatest magnitude involve the smallest size categories (< 10 g, 11–100 g), aerial insectivores and frugivores. Species in these categories, mostly bats, increase along a gradient of decreasing winter temperature and increasing annual moisture and frostfree period, trends correlated with latitude. At the opposite end of this gradient, species in the largest size category (101–1000 kg) increase in frequency. Species in size categories 3 (101–1000 g), 5 (11–100 kg) and 6 (101–1000 kg), herbivores, and granivores increase along a longitudinal gradient of increasing annual potential evapotranspiration and elevation. Much of the spatial pattern is consistent with ecological sorting of species ranges along environmental gradients, but differential rates of speciation and extinction also may have shaped the ecological diversity of extant North American mammals.