Quantitative conodont-based approaches for correlation of the Late Devonian Kellwasser anoxic events

We present the trends of absolute conodont abundance, relative genera abundance, and conodont morphometrics across the Late Devonian Kellwasser horizons at three northern Gondwanan epicontinental seaway locations.

At three contrasting depositional settings, the Kellwasser events are characterized by a decrease in conodont abundance, high percentage of the conodont genus Icriodus and morphological change of the conodont genus Palmatolepis. These changes can be identified in sections where the events are expressed as interbedded anoxic and oxic deposits as well as in settings characterized by continuous anoxic sediment accumulation. Hence, these changes are interpreted as global and synchronous and can be thus used for event correlation.

The paradox of these results is that despite quite similar isotopic signals, faunal change during the Lower Kellwasser event was less than during the Upper Kellwasser event. We interpret that this discrepancy is due to an unconformity (including topmost Frasnian to earliest Famennian) at several F/F sections which removed the record of peak paleoenvironmental change.     

Pattern of conodont reworking in the Upper Devonian of the Alps: palaeoenvironmental and palaeotectonic implications

Transfer of sediments from shallow shelf to basin usually implies vertical (stratigraphic) reworking of fossils, but in a few instances produces only lateral reworking (actually within the time span of a subzone). An Upper Devonian (Frasnian) pelagic carbonate sequence with allodapic intercalations and thick intraclast parabreccias triggered by seismic shocks and related tsunami and turbidity currents has been studied. Here, the standard biozone sequence is undisturbed at the zonal as well as the subzonal level. However, distribution of conodonts is discontinuous, and ranges of some important species are altered; mature specimens are commonly broken, whereas complete specimens are mostly juvenile forms. Maxima and minima of element frequency and anomalous thickening or thinning of subzones are recognized. Such distribution patterns of conodonts are compared with the four main facies distinguished and related to the environmental interpretation (a platform-basin system, the original gentle slope of which was fragmenting in fault-controlled blocks). The mean frequency of shallow-water, proximal conodonts is 10% whereas transitional to distal deeper-water genera dominate throughout the entire section. In the dominant biofacies, conodonts settled vertically, in the subordinate one they were transported laterally. The 'intraclast parabreccia' is interpreted as a peculiar type of seismitc. The anomalous thickness variation of biozones is related to the step topography of the slope in a fault-controlled carbonate basin.     

Plicathyridine brachiopods (Athyridida) from the Frasnian (Late Devonian) of Western Europe and Middle East

Plicathyridine brachiopods (Athyridida) from the early–middle Frasnian of southern Belgium and northern France (Dinant Synclinorium) are systematically described for the first time. They include two species: Anathyris (Anathyris) calestiennensis nov. sp., and A. (A.) sp. indet. 1. They are uncommon in the mainly shally La Prée (Nismes Formation) and Ermitage (Moulin Liénaux Formation) members and are absent from the mixed argillaceous–carbonate late Frasnian succession in this area contrary to what is observed in Russia, notably in the East-European Platform and Siberia. In the Middle East, two species of Anathyris are recognized on the basis of a limited material from the Dascht-e-Nawar area in Afghanistan (A. (A.) sp. indet. 2) and the Kuh-e Kaftar mountains in Central Iran (A. (A.) sp. indet. 3). Anathyris (A.) calestiennensis nov. sp. is sometimes encrusted by epizoans (tabulate and rugose corals, and bryozoans) and rarely displays single, small circular drill holes. The past and current epizoan–brachiopod interactions are also discussed (Anathyris vs. Lingula, respectively).     

Late Devonian Acritarchs from the Gneudna formation in the Western Carnarvon basin,Western Australia

The Gneudna Formation is a Late Devonian(Frasnian) sequence of marine calcareous sediments that occurs in the Carnarvon Basin, Western Australia. The present palynological study is based upon subsurface silty strata from a borehole (Pelican Hill or Bibbawarra Bore) that was drilled early this century near the western coastal limit of the Carnarvon Basin.The subject strata have previously been attributed to the Gneudna Formation on lithostratigraphic grounds. They contain a rich and varied assemblage of marine microphytoplankton (acritarchs), associated with trilete miospores of which Geminospora lemurataBalme, 1962 is the dominant form. Forty-seven species of acritarchs are recognizable in the palynoflora, which corresponds very closely with that described recently (Playford & Dring, 1981) from the Gneudna Formation in the vicinity of its type section on the opposite (eastern) side of the Carnarvon Basin. The apparently parochial complexion of the Gneudna acritarch suite is probably illusory, insofar as early Late Devonian acritarchs have not been studied extensively or intensively from either the northern or southern hemispheres.The following new species of acritarchs areformally instituted herein: Elektoriskos villosa, Lophosphaeridium pelicanensis, and Pterospermella tenellula.     

Late Middle to Late Frasnian Atrypida, Pentamerida, and Terebratulida (Brachiopoda) from the Namur-Dinant Basin (Belgium)

In the Namur-Dinant Basin (Belgium), the last Atrypida and Pentamerida originate from the top of the Upper Palmatolepis rhenana Zone (Late Frasnian). Within this biozone, their representatives belong to the genera Costatrypa, Desquamatia (Desquamatia), Radiatrypa, Spinatrypa (Spinatrypa), Spinatrypina (Spinatrypina?), Spinatrypina (Exatrypa), Waiotrypa, Iowatrypa and Metabolipa. No representative of these orders occurs within the Palmatolepis linguiformis Zone. The disappearance of the last pentamerids, mostly confined to reefal ecosystems, is clearly related to the end of the edification of the carbonate mounds; it precedes shortly the atrypid one. This event, resulting from a transgressive episode, which induces a progressive and dramatic deterioration of the oxygenation conditions, takes place firstly in the most distal zones of the Namur-Dinant Basin (southern border of the Dinant Synclinorium; Lower P. rhenana Zone). It is only recorded within the Upper P. rhenana Zone in the Philippeville Anticlinorium, the Vesdre area, and the northern flank of the Dinant Synclinorium. It would seem that the terebratulids were absent during the Famennian in this basin, probably due to inappropriate facies. Among the 13 species described or briefly discussed (Palmatolepis hassi to Upper P. rhenana zones), Pseudoatrypa godefroidi nov. sp. and Spinatrypina (Exatrypa) marmoris nov. sp. are proposed as new.     

Age and distribution of the Late Devonian brachiopod genus Dzieduszyckia Siemiradzki, 1909 in southern China

Conodont biostratigraphical work was done at four sections recently found with occurrence of the rhynchonellide brachiopod genus, Dzieduszyckia Siemiradzki, in southern Guangxi and in the border area between Dushan County of Guizhou and Nandan County of Guangxi, South China. These sections represent two different types of facies, i.e., carbonate platform and intraplatform basin. The conodont analysis reveals that this genus occurs in the Upper triangularis Zone and the Middle crepida Zone at the Dazhai Section, through the Lower to Upper crepida zones at Dalong, and is restricted within the Upper rhomboidea Zone at the two intraplatform basin sections (Changtang and Duli). This result demonstrates that the occurrence of these peculiar rhynchonellide brachiopods in South China, regardless of the depositional environments, is within the Lower Famennian instead of the previously suggested Upper Famennian. Furthermore, this brachiopod genus in South China began to inhabit on the carbonate platform almost since the beginning of the Famennian and did not extend to the intraplatform basin facies until the late Early Famennian.Available biostratigraphic data indicate that during the Early and Middle Famennian, Dzieduszyckia is widely distributed not only in South China, but also throughout the world, such as Morocco and southern Ural.Observation on the new collections from the four studied sections reveals that the peculiar rhynchonellide brachiopods have a great morphological variation within each section. Significant differences existed among the collections from different sedimentary settings and localities, probably reflecting the environmental and geographic constraint on the morphology of Dzieduszyckia. Samples from different layers in the same section have nearly identical morphological variation, suggesting the temporal inheritance in morphology of the rhynchonellide brachiopod.     

Conserved phenotypic variation patterns, evolution along lines of least resistance, and departure due to selection in fossil rodents

Within a group of organisms, some morphologies are more readily generated than others due to internal developmental constraints. Such constraints can channel evolutionary changes into directions corresponding to the greatest intraspecific variation. Long-term evolutionary outputs, however, depend on the stability of these intraspecific patterns of variation over time and from the interplay between internal constraints and selective regimes. To address these questions, the relationship between the structure of phenotypic variance covariance matrices and direction of morphological evolution was investigated using teeth of fossil rodents. One lineage considered here leads to Stephanomys, a highly specialized genus characterized by a dental pattern supposedly favoring grass eating. Stephanomys evolved in the context of directional selection related to the climatic trend of global cooling causing an increasing proportion of grasslands in southwestern Europe. The initial divergence (up to approximately 6.5 mya) was channeled along the direction of greatest intraspecific variation, whereas after 6.5 mya, morphological evolution departed from the direction favored by internal constraints. This departure from the "lines of least resistance" was likely the consequence of an environmental degradation causing a selective gradient strong enough to overwhelm the constraints to phenotypic evolution. However, in a context of stabilizing selection, these constraints actually channel evolution, as exemplified by the lineage of Apodemus. This lineage retained a primitive diet and dental pattern over the last 10 myr. Limited morphological changes occurred nevertheless in accordance with the main patterns of intraspecific variation. The importance of these lines of least resistance directing long-term morphological evolution may explain parallel evolution of some dental patterns in murine evolution.     

Progressive drowning of carbonate platform in the Moravo-Silesian Basin (Czech Republic) before the Frasnian/Famennian event: facies,compositional variations and gamma-ray spectrometry

The Moravo-Silesian Basin (MSB; eastern Czech Republic and southern Poland) hosted an extensive shallow-water carbonate platform in the Middle Devonian to Frasnian interval. The platform drowned in a stepwise fashion from the Palmatolepis hassi to the Pa. linguiformis zone. Three types of drowning successions were revealed from conodont biostratigraphy, facies, microfacies and gamma-ray spectrometry data: (A) drowning to periplatform turbidite setting; (B) drowning to (hemi)pelagic seamount setting and (C) drowning associated with the stratigraphical gap. In the lower Pa. hassi zone, rapid subsidence caused the platform to drown locally along the N–S to NW–SE trending faults (type A drowning). In the upper Pa. rhenana to the Pa. linguiformis zone, the drowning accelerated in the western part of the MSB due to locally higher subsidence rates combined with the Late Frasnian biotic crisis (type B). In the southern part of the basin, the platform emerged shortly before the Frasnian/Famennian (F/F) boundary and drowned in the Early to Late Famennian (type C). The primary cause of drowning was differential subsidence at the Laurussian passive margin. Eustatic sea-level fluctuations, if any, contributed only to a minor extent to the Late Frasnian drowning, but were effective in type C drowning during the Famennian. The drowning boundaries are associated with increased contents of K and Th, reflecting the deceleration of carbonate production. Uranium contents display isolated peaks that roughly correlate with the drowning boundaries or the stratigraphic gaps associated with the F/F boundary. The uranium contents are considered to reflect local depositional conditions and are not suitable for stratigraphic correlation. On the other hand, from the K and Th contents, we can infer Late Frasnian sea-level fluctuations with duration on the order of 1 Myr. These cyclic variations in K and Th contents proved to be useful in platform-to-basin stratigraphic correlation.     

Late Triassic sedimentary evolution of Slovenian Basin (eastern Southern Alps): description and correlation of the Slatnik Formation

Successions of the Slovenian Basin structurally belong to the easternmost Southern Alps. During the Late Triassic, they were part of the Adriatic continental margin. Norian–Rhaetian successions of the Slovenian Basin are characterized mainly by dolomite of the Bača Dolomite Formation. However, in the northern part of the basin, Late Triassic limestone is preserved above Bača Dolomite Formation and is formalized as the Slatnik Formation. It is composed of hemipelagic limestone alternating with resedimented limestones. The succession documents an upward progradation of the slope environment composed of three high-frequency cycles. Most prominent progradation is referred to the second, i.e., Early Rhaetian cycle. The Slatnik Formation ends with thin-bedded hemipelagic limestone that records the end-Triassic productivity crisis, or rapid sea-level fall. The overlying resedimented limestones of the Early Jurassic Krikov Formation, document the recovery of production and shedding from the adjacent carbonate platform.     

Late Cretaceous (Cenomanian to Campanian) paleoenvironmental history of the Eastern Canadian margin of the Western Interior Seaway: bonebeds and anoxic events

Upper Cretaceous strata in the Pasquia Hills of the northern Manitoba Escarpment, eastern Saskatchewan, Canada provide a detailed paleoenvironmental and sea-level record of the eastern margin of the Western Interior Seaway. Sediments deposited during the Cenomanian/Turonian Greenhorn marine cycle are dominantly black mudstones deposited in a stratified water column, with bottom-water anoxia recurrently reaching into the photic zone. A middle Cenomanian sea-level lowstand event followed by transgression left a series of bonebeds within the Belle Fourche Member of the Ashville Formation, indicating a sedimentary environment starved of coarse siliciclastics. Maximum sea level resulted in the formation of limestone beds within the Favel Formation, further favoured by reduced terrigenous sediment input compared to the western margin. Limestone sedimentation was followed by a phase of increased freshwater input under lower sea level conditions, and reducing zoo- and phytoplankton diversities. During final Greenhorn regression, eastern Saskatchewan probably turned into a restricted basin severely limiting marine circulation. Poor or absent benthic foraminiferal assemblages and biomarker analysis suggest prevailing watermass stratification throughout the Cenomanian/Turonian transgressive/regressive cycle. This was caused either by a freshwater lid, stratification of Boreal and Tethyan-derived watermasses, or both, to various intensities affected by changing sea level. Basin oxygenation during Niobrara time varies between localities along the eastern margin as documented by presence/absence of benthic and planktic foraminifera.     

Diversity and morphological disparity of desmid assemblages in Central European peatlands

Morphological disparity has increasingly been used as an alternative measure of biological diversity based on the shape features of organisms. In this study, we investigated the species diversity and morphological disparity of benthic Desmidiales in Central European peatland pools. The shape features of cells were determined using the 3-D elliptical Fourier analysis of their frontal and lateral views. The resulting morphospace was used to calculate the contributions of localities and species to the morphological variation. In addition, the disparity of samples and their average cell complexity (indicating intricacy of cell shapes) was evaluated. These data were related to species diversity data and to the abiotic factors. Species diversity was positively correlated with pH and conductivity. The low-pH localities generally supported a more variable species composition than did slightly acidic to neutral localities. Conversely, the total nitrogen concentrations of these areas negatively correlated with species diversity. Interestingly, partial morphological disparity (measuring the contribution of a sample to the overall morphological variation) did not correlate with species diversity. On the contrary, several mountain peat bog localities had high disparity values, irrespective of their rather low species diversity. In addition, several samples from minerotrophic fens with high diversity had average or low values of partial morphological disparity. These results indicate the relative importance of mountain peat bogs for the total morphological diversity of Desmidiales within the region that could not be ascertained solely from species diversity data. The inner morphological disparity of samples was highly correlated with their species diversity. Species of the genus Micrasterias, Hyalotheca dissiliens and Desmidium species had the highest partial morphological disparity, thus indicating their marginal position within the morphospace. Micrasterias and Euastrum species had the highest complexity values. The average cell complexity of individual samples did not correlate with their diversity or disparity; however, it was positively correlated with the levels of total nitrogen and phosphorus, and illustrates a pattern different from that arrived at by species diversity data. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users. Handling editor: J. Padisak     

Parallel evolution in molar outline of murine rodents: the case of the extinct Malpaisomys insularis (Eastern Canary Islands)

The lava mouse Malpaisomys was part of Pleistocene and Holocene faunas of the eastern Canary Islands; it became extinct during historical times. In order to evaluate the evolutionary processes of this endemic species, we set out to identify its relationships with possible mainland relatives. Its dental morphology was compared to a set of fossil and modern murine rodents from various phylogenetic groups, characterized by different diets and dental patterns, using a quantitative method based on a Fourier analysis of the outline of the first upper and lower molars. This morphometric analysis identified different evolutionary grades that are independent of the phylogenetic group. The first cluster is associated with primitive, asymmetric dental outlines, the second with intermediate forms linked to a more herbivorous diet. Highly differentiated forms diverge not only from the second cluster but also from each other. Our investigations reveal Malpaisomys to have had an intermediate dental pattern, a result which confirms previous palaeoecological interpretations of this taxon. However, conclusions about its closest mainland relative remain tentative. Based on a comparison of dental size and shape, as well as geographical considerations, a possible mainland ancestor could be either Paraethomys (North-African Pliocene) or Occitanomys (South-western European Pliocene). Such results support the hypothesis of a Pliocene colonization event, in which case Malpaisomys would display a normal evolutionary rate in dental size and shape, in spite of the insular context. In contrast, a late colonization by a modern representative of the African fauna would imply exceptionally high evolutionary rates compared to the background morphological evolution and a decrease in size that is unlikely under insular conditions.  © 2004 The Linnean Society of London, Zoological Journal of the Linnean Society , 2004, 142 , 555–572.     

On the interpretation and application of mean times to extinction

As a metric of population viability, conservation biologists routinely predict the mean time to extinction (MTE). Interpretation of MTE depends on the underlying distribution of times to extinction (DTE). Despite claims to the contrary, all information regarding extinction risk can be obtained from this single statistic, the MTE, provided the DTE is exponential. We discuss the proper interpretation of MTE and illustrate how to calculate any population viability statistic when only the MTE is known and the DTE is assumed to be exponential. We also discuss the restrictive assumptions underlying the exponential DTE and the conditions under which alternative models for the DTE are preferable to the conventional (exponential) model. Despite superficial similarities between the exponential and alternative DTEs, several key differences can lead to substantially different interpretations of the MTE.     

The placoderm <Emphasis Type="Italic">Plourdosteus livonicus</Emphasis> (Eastman) in the early Frasnian of the Central Devonian Field and the trophic structure of the Mikhailovskii Fish Assemblage

The placoderm Plourdosteus livonicus (Eastman) (Euarthrodira, Coccosteina) from the Upper Timanian Mikhailovskii Fish Assemblage of the Central Devonian Field is described. The composition and structure of the assemblage are discussed and compared with later Frasnian fish assemblages of the Central Devonian Field.     

When morphometry meets genetics: inferring the phylogeography of Carabus solieri using Fourier analyses of pronotum and male genitalia

Population differentiation is a crucial step in the speciation process and is therefore a central subject in studies of microevolution. Assessing divergence and inferring its dynamics in space and time generally require a wide array of markers. Until now however, most studies of population structure are based on molecular markers and those concerning morphological traits are more scarce. In the present work, we studied morphological differentiation among populations of the ground beetle Carabus solieri, and tested its congruence with genetic population structure. The shape of pronotum and aedeagus was assessed using Dual Axis Fourier Shape Analysis. manova on Fourier coefficients revealed highly significant morphological variation between populations and a similar geographical pattern of differentiation for both structures. On the whole, morphological and genetic patterns were also found to be congruent. Our analysis confirms the phylogeographical scenario proposing that two entities of C. solieri differentiated during the last glaciation events before recolonizing the actual range of the species. It also indicates a large introgression between the two differentiated entities in the centre of the range.     

Activities of proteinases in maize and soybean roots in response to anoxic stress

This study characterized the changes in proteinase activities in maize inbred line H60 and soybean cultivar Keller roots in response to anoxia. After 24 h of anoxia, crude protein extracts from both maize and soybean root tips (10 cm) were assayed for proteinase activities at pHs ranging from 4.5 to 10.2. In anoxic roots of both maize and soybean, activities of proteinases with alkaline pH optima increased, and activities of proteinases with acidic pH optima declined. Proteinases with neutral pH increased in anoxic maize roots, but declined in anoxic soybean roots. Whether the differences in proteinase activities in anaerobic maize and soybean roots contribute to the differental susceptibility of the two species requires further study.Journal Article No. 265-89.     

Adaptive latitudinal trends in the mandible shape of Apodemus wood mice

Aim Size and shape of the mandible are investigated across the latitudinal range of the European wood mouse (Apodemus sylvaticus), in order to address the relative importance of genetic structure, insularity, and geographical gradient in patterning morphological variation. Results are compared with those on two Asiatic species of wood mice, A. argenteus and A. speciosus. Location The European wood mouse is sampled by a set of trapping localities including both, islands and mainland populations, as well as the four genetic groups identified in previous studies. The localities cover a latitudinal gradient from 55° N to 36° N. Methods Different Fourier methods are applied to the outlines of mandibles and their results compared in the case of A. sylvaticus. All provide similar results and allow a quantification of the size and shape variations across the geographical range of the European wood mouse. Using the method allowing for the best reduction of the informative data set, a comparison of the European wood mouse with the two Asiatic species was performed. Results Within the European wood mouse A. sylvaticus, a strong latitudinal gradient in mandible shape overrides the influence of insularity and genetic structure. Yet, random morphological divergence in insular conditions can be identified as a secondary process of shape differentiation. Size displays no obvious pattern of variation, neither with insularity or latitude. A comparison with two other species of wood mice suggests that a similar latitudinal gradient in mandible shape exists in different species, mandibles being flatter in the north and wider in the south. Main conclusion The latitudinal gradient in mandible shape observed in the three species of wood mice is interpreted as an intraspecific adaptive response to gradual changes in feeding behaviour.