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.     

Step-wise morphological trends in fluctuating environments: Evidence in the Late Devonian conodont genus Palmatolepis

Long-term morphological changes were investigated in the conodont genus Palmatolepis, using a Fourier analysis of the outline of platform elements. Three time-slices and four Late Frasnian - Early Famennian sections on North Gondwana were studied. No difference existed between locations, but shape variations were recorded associated with the successive Kellwasser events. The Middle Famennian period was analyzed in a section located at the Baltica margin across the annulata event. Some morphological changes may be related to the deposition of the corresponding black shales, but the main feature is a heterogeneity of the populations, which can be split into two morphological types. Latest Famennian conodonts from another North Gondwanan section show only minor shape changes during this time interval, which is environmentally stable prior to the global Hangenberg event. Based on synthetic shape variations across the whole period an morphological trend towards narrower elements is observed. This morphological change happens step-wise in response to global events. The amplitude of the shape shift is proportionally related to the extend of the successive Late Devonian global events.     

Population dynamics of the Late Devonian conodont Alternognathus calibrated in days

Abstract

Alternognathus is a cavusgnathid conodont believed to be a relative and possible ancestor of the Siphonodella lineage, which is of crucial importance to the biostratigraphy of the Devonian–Carboniferous transition, as well as the Early Carboniferous Clydagnathus, with the most completely known soft tissue anatomy among conodonts. Specimens of Alternognathus from the Kowala quarry in central Poland have their basal cavities lacking any mineralised basal body tissue, exposing rhythmic growth increments of probably daily nature. This enables calibration of ontogeny and population dynamics of Alternognathus in absolute time units and to estimate the correspondence between data derived from isolated oral apparatus elements and the animal body size. The peak of mortality, probably resulting from predation, occurred at 18–28 days of the Alternognathus oral apparatus element secretion and the inferred whole animal length of about 1.6–2.9 cm. Based on analogy with present-day fish, the actual beginning of individual life presumably preceded the initiation of the mineral tissue secretion for about 8 days. This makes the dominant life cycle close to the lunar month length and suggests analogy to cyclic mortality of modern tropical organisms due to full moon light influence on their daily vertical migration.     

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.     

Patterns of bilateral asymmetry and allometry in Late Devonian Polygnathus conodonts

Conodont animals were early jawless vertebrates equipped with a feeding apparatus composed of several tooth‐like elements. The P1 elements, at the rear of the apparatus, were characterized by a robust shape and rapid morphological evolution. Occlusion occurred between paired right and left P1 elements, occasioning some bilateral asymmetry, which, together with allometric growth, may partially obliterate the temporal differences. The present study aims to disentangle these different components of morphological variation in Late Devonian Polygnathus P1 conodont elements. An extensive 2D geometric morphometric analysis of the platform shape was performed through the Famennian record of two outcrops. This analysis was completed by a 3D study on a subset of conodont elements. The 2D and 3D morphometric quantifications provided highly congruent results, showing that the 2D shape constitutes a good approximation of the element geometry. The 3D analysis delivered further insights into the relationship between the geometry of the elements and the constraints related to occlusion. The 2D analysis allowed a quantitative assessment of the variation among species and through time. Allometry and bilateral asymmetry were differently expressed depending on the species considered, suggesting that constraints imposed on pairing by the morphology of the elements varied even among related species. The within‐species variation was so important that it largely obliterated temporal trends; a relationship of Polygnathus shape and conodont biofacies variations through the Famennian nevertheless suggested an evolution driven by ecological interactions between conodont genera.     

Indigenous demosponge spicules in a Late Devonian stromatoporoid basal skeleton from the Frasnian of Belgium

This paper records the first example of a demosponge spicule framework in a single specimen of a Devonian stromatoporoid from the Frasnian of southern Belgium. The small sample (2.5 × 2 cm) is a component in a brecciated carbonate from a carbonate mound in La Boverie Quarry 30 km east of Dinant. Because of the small size of the sample, generic identification is not confirmed, but the stromatoporoid basal skeleton is similar to the genus Stromatopora. The spicules are arranged in the calcified skeleton, but not in the gallery space, and are recrystallized as multi‐crystalline calcite. The spicules fall into two size ranges: 10–20 μm diameter and 500–2000 μm long for the large ones and between 5–15 μm diameter and 50–100 μm length for the small ones. In tangential section, the spicules are circular, they have a simple structure, and no axial canal has been preserved. The large spicules are always monaxons, straight or slightly curved styles or strongyles. The spicules most closely resemble halichondrid/axinellid demosponge spicules and are important rare evidence of the existence of spicules in Palaeozoic stromatoporoids, reinforcing the interpretation that stromatoporoids were sponges. The basal skeleton may have had an aragonitic spherulitic mineralogy. Furthermore, the spicules indicate that this stromatoporoid sample is a demosponge.     

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).     

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.     

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.     

Static allometry of unicellular green algae: scaling of cellular surface area and volume in the genus Micrasterias (Desmidiales)

The surface area‐to‐volume ratio of cells is one of the key factors affecting fundamental biological processes and, thus, fitness of unicellular organisms. One of the general models for allometric increase in surface‐to‐volume scaling involves fractal‐like elaboration of cellular surfaces. However, specific data illustrating this pattern in natural populations of the unicellular organisms have not previously been available. This study shows that unicellular green algae of the genus Micrasterias (Desmidiales) have positive allometric surface‐to‐volume scaling caused by changes in morphology of individual species, especially in the degree of cell lobulation. This allometric pattern was also detected within most of the cultured and natural populations analysed. Values of the allometric S:V scaling within individual populations were closely correlated to the phylogenetic structure of the clade. In addition, they were related to species‐specific cellular morphology. Individual populations differed in their allometric patterns, and their position in the allometric space was strongly correlated with the degree of allometric S:V scaling. This result illustrates that allometric shape patterns are an important correlate of the capacity of individual populations to compensate for increases in their cell volumes by increasing the surface area. However, variation in allometric patterns was not associated with phylogenetic structure. This indicates that the position of the populations in the allometric space was not evolutionarily conserved and might be influenced by environmental factors.     

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.     

Diagnostic differences in mandibular P4 shape between Neandertals and anatomically modern humans

This study uses elliptical Fourier analysis to quantify shape differences observed in the P(4) crown of Neandertals and anatomically modern humans. Previously, P(4) shape was assessed qualitatively, and results suggested marked differences between Neandertals and anatomically modern humans (Bailey [2002] New Anat. 269:148-156). The goal of this study was to investigate the P(4) shape in more detail, quantifying it in order to determine its utility for taxonomic classification and phylogenetic analysis. A comparison of mean shapes confirms that the mesiolingual portion of the P(4) is truncated in Neandertals, and that this produces a distinctively asymmetrical P(4). A randomization test confirms that the shape difference between Neandertals and anatomically modern humans is significant. Principal component and discriminant function analyses indicate that the relative size of the lingual portion of the tooth also affects tooth shape, with the lingual portion of the Neandertal P(4) being narrower than that of anatomically modern humans. Classification of P(4) crown shapes using discriminant functions analysis is far from perfect. While 86.4% of the teeth were correctly classified, classification was much better for anatomically modern humans (98.1%) than it was for Neandertals (65%). Fortunately, crown shape is but one of several diagnostic characters of the P(4) crown. P(4) crown asymmetry can be added to the growing list of dental morphological characters distinguishing Neandertals from anatomically modern humans. Moreover, based on a comparison of mean tooth shapes in fossil and recent humans, symmetry, rather than asymmetry, appears to be the primitive state, and the high frequency of P(4) asymmetry is likely derived in Neandertals.     

Assessment of rampant genitalic variation in the spider genus Homalonychus (Araneae, Homalonychidae)

Abstract. Animal genitalia are often complex and thought to vary little within species but differ between closely related species making them useful as primary characters in species diagnosis. Spiders are no exception, with nearly all of the 40,462 (at the time of this writing) described species differentiated by genitalic characteristics. However, in some cases, the genitalia of putative species are not uniform, but rather vary within species. When intraspecific variation overlaps interspecific variation, it can be difficult (if not impossible) to place a name on a specimen. The quantification of shape variation in genitalia has not often been attempted, probably because until recently it was not a methodologically and computationally simple process. In the two currently recognized species of the spider genus Homalonychus , genitalic variation is rampant in both male and female structures, with some parts of the genitalia (e.g., the retrolateral tibial apophysis) differing in each specimen examined. In this study, geometric morphometric analysis employing landmark data is used to quantify both intra- and interspecific variation in this genus. The large amount of variation is condensed into two or three groups depending on the structures examined, and these groups correspond to either the two species or to previously established mitochondrial DNA clades within one of the species. The results also show that analyses of female structures do not separate the groups as readily as the analyses of the male structures. The large amount of variation present in some structures is not correlated with geography or population genetic structure.     

Disentangling temporal patterns in our perception of the fossil history of gymnosperms

By taking gymnosperms as a case study, this article evaluates the perception of plant life history from the fossil record to test the biases associated with the time-dependent aspects of the taxonomy, following a stepwise modelling procedure based on two divergent sets of time units. The idea that the effects of the temporal component of paleobiological inference need to be evaluated to remove any possible bias in our interpretation and perception of plant evolution based on analyses of large-scale data sets is investigated. The results reveal important differences in our perception of the tempo of gymnosperm evolution and how it is biased in terms of time unit length due to the loss of information as a consequence of the timescale resolution. Despite singletons representing real morphological diversity translated into independent taxonomic categories, these taxa can distort perceptions of the intensity of the long paleofloristic diversification moments of gymnosperms if their effect is not considered. This study shows a complete overview of the evolutionary profiles of gymnosperms with significant discrepancies in the function of how singletons are quantitatively processed in paleobotanical data analyses, and it provides new evidence about how the ‘zoom effect’ can magnify our perception of extinction events.     

Stromatoporoid palaeoecology in the Frasnian (Upper Devonian) Belgian platform,and its applications in interpretation of carbonate platform environments

Abstract: Stromatoporoid faunas in the Frasnian of southern Belgium are abundant in the carbonate platform environments present in this area. Stromatoporoids dominate the large skeletal organisms and occur principally in biostromes. The stromatoporoid assemblage is represented by a small number of taxa. Stromatoporoid genera include Actinostroma, Amphipora, Atelodictyon, Clathrocoilona, Salairella, Stachyodes, Stictostroma, Stromatopora and Trupetostroma which are present in environments ranging from the outer, outer intermediate, inner intermediate and inner zones and associated biostromes. Most large skeletal stromatoporoids are low profile, which reinforces the conclusions of previous studies that low‐profile growth forms were the most successful stromatoporoid forms. These low‐profile forms are likely to have been important sediment stabilisers that may have led to expansion of the carbonate factory. Growth forms vary between facies, indicating some degree of environmental control on form; for example, laminar in the intermediate zone, bulbous and domical in the inner and outer zones. Stromatoporoid taxa vary in occurrence across the environmental gradient from shallow to deep. There is some taxonomic control on growth forms, with some taxa showing more variability than others in different environments.     

Terrestrial-marine teleconnections in the Devonian: links between the evolution of land plants,weathering processes,and marine anoxic events

The Devonian Period was characterized by major changes in both the terrestrial biosphere, e.g. the evolution of trees and seed plants and the appearance of multi-storied forests, and in the marine biosphere, e.g. an extended biotic crisis that decimated tropical marine benthos, especially the stromatoporoid-tabulate coral reef community. Teleconnections between these terrestrial and marine events are poorly understood, but a key may lie in the role of soils as a geochemical interface between the lithosphere and atmosphere/hydrosphere, and the role of land plants in mediating weathering processes at this interface. The effectiveness of terrestrial floras in weathering was significantly enhanced as a consequence of increases in the size and geographic extent of vascular land plants during the Devonian. In this regard, the most important palaeobotanical innovations were (1) arborescence (tree stature), which increased maximum depths of root penetration and rhizoturbation, and (2) the seed habit, which freed land plants from reproductive dependence on moist lowland habitats and allowed colonization of drier upland and primary successional areas. These developments resulted in a transient intensification of pedogenesis (soil formation) and to large increases in the thickness and areal extent of soils. Enhanced chemical weathering may have led to increased riverine nutrient fluxes that promoted development of eutrophic conditions in epicontinental seaways, resulting in algal blooms, widespread bottomwater anoxia, and high sedimentary organic carbon fluxes. Long-term effects included drawdown of atmospheric pCO₂ and global cooling, leading to a brief Late Devonian glaciation, which set the stage for icehouse conditions during the Permo-Carboniferous. This model provides a framework for understanding links between early land plant evolution and coeval marine anoxic and biotic events, but further testing of Devonian terrestrial-marine teleconnections is needed.     

Phanerozoic survivors: Actinopterygian evolution through the Permo‐Triassic and Triassic‐Jurassic mass extinction events

Actinopterygians (ray‐finned fishes) successfully passed through four of the big five mass extinction events of the Phanerozoic, but the effects of these crises on the group are poorly understood. Many researchers have assumed that the Permo‐Triassic mass extinction (PTME) and end‐Triassic extinction (ETE) had little impact on actinopterygians, despite devastating many other groups. Here, two morphometric techniques, geometric (body shape) and functional (jaw morphology), are used to assess the effects of these two extinction events on the group. The PTME elicits no significant shifts in functional disparity while body shape disparity increases. An expansion of body shape and functional disparity coincides with the neopterygian radiation and evolution of novel feeding adaptations in the Middle‐Late Triassic. Through the ETE, small decreases are seen in shape and functional disparity, but are unlikely to represent major changes brought about by the extinction event. In the Early Jurassic, further expansions into novel areas of ecospace indicative of durophagy occur, potentially linked to losses in the ETE. As no evidence is found for major perturbations in actinopterygian evolution through either extinction event, the group appears to have been immune to two major environmental crises that were disastrous to most other organisms.