Disentangling allometry and response to Kellwasser anoxic events in the Late Devonian conodont genus Ancyrodella

Is the ability to respond to minor perturbations a key to survival to major crisis? This idea was supported by previous studies on Late Devonian conodonts. Genera surviving the Frasnian/Famennian (F/F) crisis, at the end of the Upper Kellwasser anoxic event, had shown a response to a similar but minor event, the Lower Kellwasser (LKW). In contrast, the genus Ancyrodella went extinct at the F/F crisis. This study aims to assess whether this extinction was associated with the absence of response to the minor LKW event. For this purpose, the shape of the P1 platform elements of the conodont Ancyrodella curvata was quantified based on their 2‐D outlines. Conodonts were extracted from two outcrops documenting the Late Devonian Kellwasser events, Coumiac (France) and Mrirt (Morocco). Ancyrodella curvata is characterized by a marked allometric growth that may blur any other source of shape difference, like response to the LKW. Hence, a multivariate method was developed to disentangle size‐free from size‐related shape variations. No coherent temporal size variations emerged from the analysis, but allometry occurring along the growth of the element indeed appeared as the prevailing source of shape variation. By isolating size‐free variations, however, we were able to evidence a morphometric response of A. curvata to the LKW independent from the allometric pattern. Hence, the extinction of A. curvata at the F/F crisis cannot be attributed to an absence of response to minor events. The amount of shape change involved in the LKW shift is, however, of minor importance in comparison with the shape variation due to allometric growth. Alternate hypotheses have been further investigated. Ancyrodella curvata appears as a rare species in the assemblages, and its distribution worldwide suggests a rather specialist taxon. Considering survivorship curves (i.e. frequency distribution of the conodont as a function of its size) A. curvata was characterized by low and constant mortality rate. This may be a clue for a long growth and progressive investment in the reproduction. These arguments support the view that A. curvata was a rather specialist, rare and ‘K‐selected’ taxon. All these factors may not be exclusive and their unfavourable conjunction might have been the key to the extinction of A. curvata.     

Phyllocarid remains (Malacostraca, Crustacea) from the basal Famennian (Late Devonian) of the Montagne Noire (Southern France)

From the early Famennian of the Montagne Noire 19 different morphotypes of probably phyllocarid (Crustacea) affinities are described. Apart from formerly unknown enamel-like structures that veil the uppermost tips of the denticles on most of the gnathal lobe morphs, the stratigraphic implications concerning extinction behaviour across the Kellwasser mass extinction event and during early Famennian diversification are discussed. In addition, phyllocarids in the Famennian seem to form a useful stratigraphic tool for biostratigraphical correlation. Therefore, three basal Famennian phyllocarid assemblage zones are distinguished.      

Sedimentary and faunal changes across the frasnian/famennian boundary in the canning basin of Western Australia

The Canning Basin of northwestern Australia is a key area for understanding global changes at the “Kellwasser Events” and the Frasnian‐Famennian boundary. Frasnian stromatoporoid‐coral‐cyanobacterial reef platforms stretched out for enormous distances along the palaeoshelf but in the early Famennian they were completely replaced by cyanobacterial reef platforms. An iridium anomaly in the sequence was formerly believed to be at or close to the boundary and was interpreted as possible evidence for an asteroid impact. Recent field work and detailed biostratigraphy in the area east and southeast of Fitzroy Crossing has given dating relevant to the timing and extent of sea level changes, hypoxic incursions and reef backstepping. Goniatites and conodonts provide correlations with the international biostratigraphy.

In the Horse Spring area the stage boundary falls within the Virgin Hills Formation which normally has a rich pelagic goniatite, nautiloid and conodont fauna. In the latest Frasnian (Zone 13 of Klapper 1989; regional Sphaeromanticoceras lindneri Zone) large allochthonous reef blocks moved downslope into the open marine basin. A diverse gastropod fauna is associated with the last atrypid brachiopods. The faunal record at the immediate boundary is obscured by dolomitisation but manticoceratid goniatites range into this level. There is no evidence for the organic‐rich dark Kellwasser limestone facies.

In the McWhae Ridge area two Frasnian goniatite horizons with Beloceras trilobites and the giant Mamicoceras guppyi and Sphaeromanticoceras lindneri transgress over the reef slope. Stromatolitic cyanobacterial beds mark condensations. Again there is no trace of the oxygen‐depleted Kellwasser facies. However, as at Horse Spring, manticoceratids persisted into dolomitic marker beds that have no other preserved macrofauna. The iridium anomaly associated with Frutexites postdates the Frasnian‐Famennian boundary and was formed by cyanobacterial concentration.     

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.     

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.     

Spiriferide brachiopods from the Famennian (Late Devonian) Hongguleleng Formation of western Junggar,Xinjiang, northwestern China

Junggar is one of the most important areas in the study of the Upper Devonian and Lower Carboniferous in the Junggar-Hinggan Stratigraphic Region. Nevertheless, abundant benthic fossils await modern taxonomic studies. In this paper, six genera and eight species of Famennian spiriferide brachiopods are described from the Duguer Member and its equivalents of the Hongguleleng Formation in western Junggar, adding new data to the Rugaltarostrum Subassemblage of the “Palaeospirifer”–Megalopterorhynchus Brachiopod Assemblage of this area. Five previously recognized species within this subassemblage are revised, including: Cyrtospirifer junggarensis F.M. Zhang [now Ulbospirifer? junggarensis (F.M. Zhang)]; Tenticospirifer tenticulum transversus F.M. Zhang [now Cyrtospirifer transversus (F.M. Zhang)]; Tenticospirifer koketekensis F.M. Zhang [now Cyrtospirifer koketekensis (F.M. Zhang)]; Mucrospirifer quadratus F.M. Zhang (now “Mucrospirifer” quadratus F.M. Zhang); and Mucrospirifer bouchadi (Muir-Wood) (now Tylothyris cf. novamexicana Stainbrook). Three additional species, Cyrtiorina houi n. sp., Goungjunspirifer sinicus F.M. Zhang, and Cyrtospirifer procumbens Simorin in Litvinovich et al., are described. Ulbospirifer? junggarensis and Goungjunspirifer sinicus are characterized by a prismatic layer. Cyrtiorina houi shows a moderately high and triangular ventral interarea, with delthyrium covered by pseudodeltidium with a rounded foramen. Cyrtospirifer procumbens can be distinguished from other species of this genus by its shallow, clearly limited sulcus with prominent primary plications, and high ventral interarea. Cyrtospirifer transversus has an alate to subpentagonal outline. Cyrtospirifer koketekensis has rounded and distinct flank plications. The ventral delthyrium of “Mucrospirifer” quadratus is covered by a pseudodeltidium. Tylothyris cf. novamexicana develops an angular sulcus in cross-section. Comparisons with the coeval brachiopod faunas from various regions indicate that, biogeographically, western Junggar has a close relationship with the Tarbagatai Mountain Range, northeastern Kazakhstan, central Kazakhstan, and Karaganda Basin, and a certain affinity with North America.     

Palynostratigraphy of the Frasnian-Famennian boundary deposits from the Central Devonian Field, western Russia and comparisons with adjacent areas

Detailed palynological research carried out on samples from the three quarries at Khlevnoye and Kamenka (Voronezh region) permitted recognition of the Frasnian-Famennian boundary in the lower Kamenka quarry. Two miospore zones, Cristatisporites deliquescens-Verrucosisporites evlanensis (DE) from the Frasnian and Cyrtospora cristifera-Diaphanospora zadonica (CZ) from the Famennian, have been recognized. Some faunistic data were also obtained from the F-F boundary deposits. Research confirmed the existence of a sedimentological gap between the Livny and Zadonsk horizons, including all of the Volgograd horizon. These new palynological results are compared with data from adjacent areas: Pripyat Depression (Belarus), Timan-Pechora Province and the Volgograd Volga region (Russia), western Pomerania and the Holy Cross Mountains (Poland).     

Taphonomy of acanthodians from the Devonian Bunga Beds (late Givetian/early Frasnian) of New South Wales

Several articulated, but incomplete, acanthodians from the Bunga Beds (late Givetian/early Frasnian) of the southern coast of New South Wales are tentatively identified as ischnacanthids. Heads are missing from all three prepared specimens. They exhibit the following characters: two dorsal fin spines; long, slender scapulocoracoids; slender, relatively deeply inserted, unpaired fin spines; minute scales with a fairly smooth, flat, crown; and an increase in size of normal body scales towards the tip of the tail. The fish are preserved in black, finely laminated shales, which were probably deposited as deep water, lacustrine sediments. The rarity, burial conditions, and headless state of the Bunga Beds acanthodians indicate that they might have died in shallow water, sunk to the bottom, refloated by gas‐induced buoyancy, with the heads lost while drifting out to deeper waters, where the bodies finally sank to a scavenger‐free, anaerobic substrate.     

Neoselachian (Chondrichthyes, Elasmobranchii) diversity across the Cretaceous-Tertiary boundary

Fishes are often thought to have passed through mass extinctions, including the Cretaceous-Tertiary (KT) event, relatively unscathed. We show that neoselachian sharks suffered a major extinction at the K/T boundary. Out of 41 families, 7 became extinct (17±12%). The proportional measure increases at lower taxic levels: 56±10% loss of genera (loss of 60 out of 107) and 84±5% loss of species (loss of 182 out of 216). However, the Maastrichtian and Danian are characterized by a high number of singleton taxa. Excluding singletons we have calculated a 34±11% loss of genera and a 45±9% loss of species. The simple completeness metric (SCM) for genera displays a decrease from the Maastrichtian (94%) to the Danian (85%) indicating a rather complete fossil record of neoselachian genera. The extinctions were heavy among both sharks and batoids (skates and rays), but most severe among batoids, which lost almost all identifiable species. There were equal losses among open marine apex predators (loss of Anacoracidae, Cretoxyrhinidae, and Scapanorhynchidae) and durophagous demersal forms from the continental shelf and shallow seas (Hypsobatidae, Parapaleobatidae, Sclerorhynchidae, Rhombodontidae). Benthopelagic and deep-sea forms were apparently little affected. New families with similar ecological roles (Carcharhinidae, Isuridae, Torpedinidae) replaced these families in the Danian, and full diversity of the different shark and batoid groups had been recovered by the end of the Paleocene or early Eocene. Sharks and rays suffered levels of extinction entirely in line with other groups of organisms at the K/T extinction event.     

Battenizyga, a new Early Triassic gastropod genus with a discussion of the caenogastropod evolution at the Permian/Triassic boundary

Battenizyga, a new Early Triassic gastropod genus from the Moenkopi Formation of Utah, is described and the speciesAnoptychia eotriassica Batten & Stokes, 1986 is placed in it. The new genus has an axially ribbed planktonic larval shell and a teleoconch with an angulated periphery. This character combination is unknown from the Palaeozoic. Therefore,Battenizyga represents additional evidence that recovery from the end-Permian mass extinction was connected with a faunal turnover. Additionally, the extinction of diverse Palaeozoic groups of the Caenogastropoda in the Permian (e.g., the Pseudozygopleuridae) suggest a turnover. All caenogastropod genera that hold Early Triassic species, have post-Palaeozoic type species and most were not reported from the Palaeozoic. This corroborates the view that there was an intense faunal turnover within the Caenogastropoda.Battenizyga is probably a caenogastropod that is closely related to the superfamily Zygopleuroidea which is abundant in the late Palaeozoic and early Mesozoic.      

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.     

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.     

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.     

Combined 2D electrophoretic approaches for the study of white lupin mature seed storage proteome

Seed proteome analysis by 2D IEF/SDS-PAGE techniques is challenging for the intrinsic difficulties related to quantitative disparity of the seed proteins, i.e. storage and non-storage proteins, their polymorphic nature, the extensive post-translational modifications and the paucity of deposited primary structures available. Conversely, 2D maps of seed proteomes can be extremely useful for a number of fundamental and applied investigations. In this work, we have used a combination of two experimental approaches to identify the main protein components of an emerging protein-rich legume seed, that is white lupin seed (Lupinus albus, L.). One is the canonical proteomic approach including 2D electrophoretic separation and mass spectrometry of selected trypsin-digested polypeptides; the other approach is a group comparative 2D electrophoretic analysis of cotyledonary protein families. To this second purpose, the three main families of lupin seed proteins, namely alpha-conglutins, the 11S globulin fraction, beta-conglutins, the 7S globulin fraction, and gamma-conglutin, a basic 7S protein, were isolated by conventional biochemical techniques and their 2D reference maps were compared with the total protein map. With the first approach 37 out of 40 spots, making up about 35% of total spot volumes in the 2D map, were found to belong to the main seed protein families. Thanks to cDNA-deduced lupin storage protein sequences, determined on purpose and deposited, most of the identification statistical parameters were very good. Moreover, it was possible to identify several endogenously proteolysed subunits in the map. The second comparative approach, beside confirming these attributions, allowed to allocate 124 polypeptides within the three main lupin protein families. These two approaches proved to be mutually validating and their combined use was effective for the establishment of a seed proteome map even in the case of sequence and protein post-translational processing lack of information. The results obtained also extend our knowledge of the seed storage protein polymorphism of white lupin.     

Quantitative approaches to the analysis of the control and regulation of microbial metabolism

Recently, a number of novel ways of considering the control, regulation and thermodynamics of microbial physiology have been developed and applied. We here present an overview of the new concepts involved, of their limitations and of the most recent attempts to deal with those limitations. We conclude that there no longer exist reasons of principle for vagueness in discussions of the control of microbial physiology and energetics. Further, the novel conceptual methods serve to remove part of the discordance between holistic and reductionistic views of microbial physiology.     

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.     

A Quantitative Food Web Model for the Macroinvertebrate Community of a Northern German Lowland Stream

Trophic interactions and cycling of organic carbon within the macroinvertebrate community of a Northern German lowland stream were analyzed based on a compartment model. The network model describes the structure of the food web quantifying biomass, production, and consumption of their elements, of the entire system and between trophic levels. System primary production is 153.7 g C m⁻² yr⁻¹ and invertebrate production 53.3 g C m⁻² yr⁻¹. Invertebrate consumption amounts to 702.6 g C m⁻² yr⁻¹. Main flows are identified between trophic level 1 and 2 and are connected with highly productive compartments. ‘Anodonta and Pseudanodonta’ and Dreissena polymorpha show the highest consumption of all groups with 269.9 g C m⁻² yr⁻¹ and 114.1 g C m⁻² yr⁻¹, respectively. System consumption is highest on the import from the upstream lake with 532.5 g C m⁻² yr⁻¹, sediment detritus with 135.5 g C m⁻² yr⁻¹, and primary producers with 25.7 g C m⁻² yr⁻¹. The lowest predation pressure is observed for Bivalvia with an ecotrophic efficiency of <10% and highest for Chironomidae with 91%. Approximately 20% of organic matter entering the detritus pool are recycled to the living groups of the system. Transfer efficiencies between discrete trophic levels are generally low except for transfer of detrital material between level I and II.