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A large predatory arthrodire (Vertebrata, Placodermi) from the Famennian (Upper Devonian) of the Holy Cross Mountains, Poland
Journal of Verterbrate Paleontology
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Large nektonic suspension feeders have evolved multiple times.
The apparent trend among apex predators for some evolving into feeding on small zooplankton is of interest for understanding the associated shifts in anatomy and behaviour, while the spatial and temporal distribution gives clues to an inherent relationship with ocean primary productivity and how past and future perturbations to these may impact on the different tiers of the food web. The evolution of large nektonic suspension feeders—‘gentle giants’—occurred four
times among chondrichthyan fishes (e.g. whale sharks, basking
sharks and manta rays), as well as in baleen whales (mysticetes), the Mesozoic pachycormid fishes and at least twice in radiodontan stem group arthropods (Anomalocaridids) during the Cambrian explosion. The Late Devonian placoderm Titanichthys has tentatively been considered to have been a megaplanktivore, primarily due to its gigantic size and narrow, edentulous jaws while no suspension-feeding apparatus have ever been reported. Here, the potential for microphagy and other feeding behaviours in Titanichthys is assessed via a comparative study of jaw mechanics in Titanichthys and other
placoderms with presumably differing feeding habits (macrophagy and durophagy). Finite-element models of the lower jaws of Titanichthys termieri in comparison to Dunkleosteus terrelli and Tafilalichthys lavocati reveal considerably less resistance to von Mises stress in this taxon. Comparisons with a selection of large-bodied extant taxa of similar ecological diversity reveal similar disparities in jaw stress resistance. Our results, therefore, conform to the hypothesis that Titanichthys was a suspension feeder with jaws ill-suited for biting and crushing but well suited for gaping ram feeding.
The Cleveland Shale fauna represents a unique view of the time after a major Devonian extinction event (Frasnian-Famenian) with the recovery of arthrodires (Placodermi) best represented by this most specious North American fauna. This time was followed by an additional event (Hangenberg Biocrisis) leading to the extinction of arthrodires (and all other placoderms). An understanding of the diversity and interrelationships of North American arthrodires can aid our understanding of this critical time in vertebrate evolution. A new aspinothoracid arthrodire Hlavinichthys jacksoni gen. et sp. nov. is described from the Late Devonian of northern Ohio, U.S.A., which adds to our knowledge of this group. It provides a point of comparison to other members of the fauna whose interrelationships are poorly known. A phylogenetic analysis supports an assignment of Hlavinichthys jacksoni gen. et sp. nov. among the aspinothoracid arthrodires. This work has drawn attention to the continued need for descriptive and phylogenetic analyses of this unique fauna. Decades old species descriptions need revision along with preparation and description of new taxa. The work on Hlavinichthys jacksoni gen. et sp. nov. here is one step in that process.
The arthrodire placoderm, Dunkleosteus sp., is reported from the Upper Devonian (Frasnian) of the Holy Cross Mountains, Poland. The material comprises partially preserved remains of two individuals found in the Kellwasser-like horizon of the P³ucki locality. The remains are preserved as broken bone fragments redeposited from shallower environment into deep-shelf conditions. They are labelled as Dunkleosteus sp. and seem similar to Dunkleosteus marsaisi from the Famennian of Morocco. It is likely that the form from Poland represents a new species that requires further collecting and study of new specimens. The described specimens are the oldest occurrence of the genus Dunkleosteus in Europe, the most complete one from Poland and one of the biggest placoderms with a head about 60 cm long.
The revision of the Famennian part of the “Late Devonian Standard Conodont Zonation” is based on the in-equivalence between biozones and time, and the rejection of the presumed single phyletic concept on which the previous zonation was based. It is also intended to simplify the zonation, eliminating the zonal groups named after only one taxon, and biozones that are defined by a Last Appearance Datum (LAD). The proposed revision is largely based on the zonation proposed by Ziegler and Sandberg (1990) and is for the most part correlatable using the same zonal markers. Modifications have only been made when strictly necessary, as the aim of the proposal is to maintain the stability of over 50 years of studies. The 22 zones constituting the revised zonation are defined by the First Appearance Datum (FAD) of species or subspecies that have a well-established stratigraphic range and wide geographic distribution. Each zone is named after the taxon for which the FAD defines the lower boundary. For each zone an association of other species useful for its identification is listed.
Famennian tabulate corals were very rare worldwide, and their biodiversity was relatively low. Here we report a unique tabulate fauna from the mid- and late Famennian of the western part of the Holy Cross Mountains (Kowala and Ostrówka), Poland. We describe eight species (four of them new, namely ?Michelinia vinni sp. nov., Thamnoptychia mistiaeni sp. nov., Syringopora kowalensis sp. nov. and Syringopora hilarowiczi sp. nov.); the whole fauna consists of ten species (two others described in previous papers). These corals form two assemblages-the lower, mid-Famennian with Thamnoptychia and the upper, late Famennian with representatives of genera ?Michelinia, Favosites, Syringopora and ?Yavorskia. The Famennian tabulates from Kowala represent the richest Famennian assemblage appearing after the F/F crisis (these faunas appear some 10 Ma after the extinction event). Corals described here most probably inhabited deeper water settings, near the limit between euphotic and disphotic zones or slightly above. At generic level, these faunas show similarities to other Devonian and Carboniferous faunas, which might suggest their ancestry to at least several Carboniferous lineages. Tabulate faunas described here represent new recruits (the basin of the Holy Cross mountains was not a refuge during the F/F crisis) and have no direct evolutionary linkage to Frasnian faunas from Kowala. The colonization of the seafloor took place in two separate steps: first was monospecific assemblage of Thamnoptychia, and later came the diversified Favosites-Syringopora-Michelinia fauna.
Taxonomy of the Late Devonian placoderm remains from the Holy Cross Mountains, Poland, described by Gorizdro-Kulczycka (L934,1950) and Kulczycki (1956, 1957), is revised. Several recently found specimens are also mentioned. The old collections are composed of representatives of Ptyctodontidae, Holonematidae, Plourdosteidae, Pholi-dosteidae, Selenosteidae, Titanichthyidae and Dinichthyidae, the latter with an unde-scribed species of Eastmanosteus. Newly found specimens belong to Ptyctodontidae, Plourdosteidae and Dinichthyidae. The occurrence of the Antiarcha in the Late Devonian of the Holy Cross Mountains, suggested by former authors, has not been confirmed. K e y w o rd s : Placodermi,Late Devonian, Holy Cross Mountains, Poland. Alexander Ivanov [alex@aoi.usr.pu.ru], Laboratory of Paleontology, Institute of the Earth Crust, Sankt-Petersburg University, 16 Liniya 29, 199178 St. Petersburg, Russia. Michał Ginter ffiszbit@geo.uw.edu.pl], InsĘtut Geologii Podstawowej, Uniwersytet War s zaw ski, ul. Zw irki i Wi gury 9 3, 02 -0 89 W ar s zaw a, P oland.
Famennian corals (Rugosa and Heterocorallia) occur in three areas of southern Poland: the Holy Cross Mountains and the Kraków area situated along the south−western margin of the East European Platform, and the Sudetes located within the Variscan orogenic belt. In the deep−water Famennian environments of the Holy Cross Mountains (Kowala) the first corals appeared after the Frasnian–Famennian crisis in the P. marginifera to P. trachytera zones. They are represented by monospecific assemblages of Circellia concava, a species able to live on a soft muddy and dysaerobic bottom. On even more organic−rich sediment hetero−
corals Oligophylloides flourished; they are suggested to feed on suspended or dissolved organic matter. When the environment became more aerated in the P. expansa and S. praesulcata zones more diverse coral assemblages appeared, including large dissepimented solitary rugosans. In the extremely shallow−water limestone facies (“Stromatoporoid Rocks”) of the Kraków region, two species of the colonial rugose coral Pseudoendophyllum and several solitary corals are represented. The shallow shelf carbonate facies of the Sudetes (Main Limestone) starts with assemblages dominated by three species of massive colonial Scruttonia followed by assemblages with a few species of solitary dissepimented corals. The colonial corals disappeared with the incursion of deeper−water cephalopod Wocklumeria Limestone facies and only solitary non−dissepimented rugose corals and heterocorals remained. The colonial Rugosa of Famennian age are almost unknown in other regions of the world. Their abundance in the Sudetes and the Kraków region suggests that these areas represented a refuge for corals during the high sea stand of the latest Devonian. Although several Famennian corals reveal characters typical for those of either the Frasnian or Viséan, there is a Tournaisian gap in their occurrence. These characters are more likely to develop convergently than represent a continuity within lineages. Pseudoendophyllum raclaviense sp. n. Scruttonia sudetica sp. n., S. fedorowskii sp. n., and Oligophylloides weyeri sp. n. are proposed.
Late Famennian brachiopods are described for the first time from the Holy Cross Mts. (Góry Świętokrzyskie), Poland. Six taxa belong to two families, the Rhynchonellidae and the Cranaenidae; the new genus Pugnaria and three new species (Rozmanaria magna, Pugnaria plana and Cranaena lgaviensis) are proposed. The brachiopods are dominated by the smooth and comparatively large sulcate R. magna and the uniplicate P. plana. This macrobenthic assemblage, called Rozmanaria magna assemblage, is interpreted here as having successfully colonized deep-water habitats typical of the Chęciny-Zbrza intrashelf basin on the rising slope of the submarine ridge in the Kielce region.
The Upper Devonian localities of the Kerman and Tabas area yielded numerous brachythoracid arthrodire remains, referred here mainly to Golshanichthys asiatica nov. gen., nov. sp., and Holonema cf. radiatumObrouchev. Some specimens are suggestive of species belonging to the genera Plourdosteus, Eastmanosteus and Aspidichthys. G. asiatica is a primitive dinichthyid characterized by e.g. very deep posterior and anterior embayments of the central plates.Some ptyctodontid tooth plates of «Rhynchodus and «Ptyctodus type have been found in the Frasnian and Famennian.
Latest Famennian (UD-VI, "Strunian") brachiopod fauna from Kowala (Kielce Region, Holy Cross Mountains, Poland) consists of eighteen species within 6 orders, eleven of them reported in open nomenclature. Characteristic taxa include: Schellwienella pauli, Aulacella interlineata, Sphenospira julii, Novaplatirostrum sauerlandense, Hadyrhyncha sp., Cleiothyridina struniensis. New morphological details of Schellwienella pauli, Sphenospira julii, and Aulacella interlineata are provided. The described latest Famennian brachiopod fauna is distinctly richer than that from underlying upper Famennian deposits (11 species within 4 orders). Majority of species from Kowala seem to have been adapted to deep water settings and/or poor nutrient availability. The stratigraphic separation between Planovatirostrum in the UD-III to UD-V and Novaplatirostrum in the UD-VI observed in Sauerland and in Thuringia is valid also in the Holy Cross Mountains. This is the first comprehensive report of a relatively diversified latest Famennian brachiopod fauna from surface outcrops of Poland.
Mass occurrence of benthic cyanobacterial mats in a sequence of Late Devonian black shales and bituminous limestones of the Holy Cross Mts. (central Poland), enclosing the famous Kellwasser and Hangenberg extinction horizons, is reported. The microbiota forming the mats is compared with some modern benthic chroococcalean cyanobacteria. Similarly to their extant counterparts, the Devonian cyanobacteria must had been phototrophic and oxygenic aerobes which could, however, tolerate slightly sulfidic conditions characterizing the near-bottom waters of the Late Devonian epicontinental sea. The cyanobacterial mats successfully colonized the oxygen-deficient and H(2)S-enriched seabed otherwise unfavorable for most other benthic biota. The redox state of this sluggish Late Devonian sea, ascribed previously mostly to anoxic or euxinic conditions, is reassessed as probably pulsating between anoxic, dysoxic, and weakly oxic conditions. The redox state was dependent on the rate of oxygen production by the cyanobacterial mats, the intensity of H(2)S emissions from the decaying mat biomass, and the rate of planktonic production.
The Upper Famennian Annulata Black Shales are exposed in the deep-shelf successions of the famous Kowala Quarry and the Ściegnia outcrop in the Holy Cross Mountains, Southern Poland. At Kowala, the twin Annulata anoxic events are manifest as two organic-rich (TOC up to 23 wt.%), finely laminated, fossiliferous black shales, each up to 0.6 m thick, separated by marl and massive or nodular limestone layers. The bituminous interval is condensed in the Ściegnia outcrop to a single 10 cm thick horizon. In both sections, the ABS levels date to the upper part of the Upper Palmatolepis trachytera conodont Zone, corresponding to the Diducites versabilis–Grandispora famenensis miospore Zone. The Annulata Events are marked by a flourishing pelagic biota (the opportunistic bivalve Guerichia venusta, platyclymenids, the goniatite Erfoudites, entomozoacean ostracods, and prasinophyte algae). The initial phase of anoxia saw a bloom of specialized conodont fauna, mostly deep-water outer-shelf palmatolepids such as Palmatolepis glabra lepta, before the second anoxic pulse caused a significant crisis amongst this community. Isorenieratane and gammacerane biomarkers indicate the development of photic zone anoxia during both phases. Degree of Pyritisation (DOP) values (estimated from the TOC–TS–Fe diagram) and the uranium–thorium proxies, as well as trace metals enrichments suggest that bottom-water conditions became dysoxic during deposition of the Lower ABS, while the more distinctly transgressive Upper ABS records anoxic/euxinic deposition during the later Event. When combined with the framboid data, however, unstable anoxia punctuated by short-term oxygenation events are assumed especially for the initial ABS phase, a crucial factor for effective nutrient recirculation from bottom waters to the photic zone and consequent phytoplankton blooming. This organic-rich level is easily identified amongst the background Upper Famennian rhythmic limestone-shaly succession, which was deposited under dysoxic to oxic conditions, with episodic anoxia developing only in the water column. Other well-known Late Devonian anoxic/high productivity episodes, recorded in the Dasberg, Kowala and Hangenberg black shales, also might partly to follow interglacial deepening pulses. These deepenings episodically reversed the overall regressive trend that resulted from a stepwise long-term climate change towards the end-Devonian Gondwanan glaciation.
A. 2004. The first Devonian holocephalian tooth from Poland. Acta Palaeontologica Polonica 49 (3): 409–415. A recently found "bradyodont" holocephalian tooth from bituminous shales of the Kowala Quarry, south−western Holy Cross Mountains, Poland, dated as the middle Famennian Palmatolepis trachytera conodont Zone, is described. In spite of its resemblance to the forms often attributed to Helodus, the tooth is referred to as Psephodus cf. magnus (Agassiz, 1838), and supposed to represent the anterior part of the dentition, based on a partly articulated specimen of Psephodus from the Carboniferous of Scotland. The analysis of early helodonts and psephodonts, and other Famennian chondrich− thyan crushing teeth, shows numerous similarities in tooth−base structure, such as the reduction of lingual basal extension, loss of articulation devices, development of numerous nutritive foramina, and the tendency to fusion between the teeth in a tooth−family. Based on these shared characters, close phylogenetic relationships between the Protacrodontoidea, Hybodontoidea, and the Holocephali are postulated.
The prime causation of the mid-Late Devonian mass extinction near the Frasnian–Famennian (F–F) boundary remains uncertain. Nevertheless, geochemical evidence has been presented recently as decisive evidence of a giant bolide impact occurring precisely at the F–F boundary, which promoted the global mortality episode. Palaeobiological data, however, imply a gradual global change, which is otherwise seen as a record of either multiple extraterrestrial catastrophes or of impact-triggered Earth-bound mechanisms. Sedimentological (mega-tsunami), physical (craters, microtektites), and geochemical records remain either elusive in many aspects, or incompatible with the predicted impact crisis pattern. Biotic succession across the F–F horizon is still poorly known, especially in continental domains, to evidence a synchronous ("bedding-plane") killing event at the close of the crisis. Instead, the commonly documented stepwise loss of biomass and an unproved distinctive "dead zone" are hard to explain simply as sampling artifacts. The assumed mass mortality precisely at the F–F boundary may be limited mainly to the pelagic realm. The underestimated role of early Variscan tectonism and associated volcanic-hydrothermal processes, resulting in thermal and nutrient pulses, as possible prime controls of the F–F crisis is suggested, as well as resemblances to the superplume-conditioned eventful mid-Cretaceous interval, exemplified in the Cenomanian-Turonian mass extinction. Additional shocks, generated by minor cometary strikes, are not excluded but may have affected some F–F biotas or areas.
The investigation of the trace element and organic geochemistry of the Frasnian–Famennian boundary section at Kowala (Holy Cross Mountains/Poland) shows that the lower water column was oxygen-deficient during late Frasnian and early Famennian times. The abundance and carbon isotopic composition of diaryl isoprenoids, biomarkers indicative for green sulfur bacteria, prove that euxinic waters reached into the photic zone, at least episodically. Total organic carbon (TOC) contents show two maxima that are time-equivalent to the Kellwasser horizons deposited in shallower water settings. Enhanced TOC concentrations are explained by a higher primary productivity, presumably as a consequence of an enhanced nutrient supply from the continent. The increase in the abundance of hopanes and bituminite suggests that the bacterial contribution to TOC increased at the Frasnian–Famennian transition. The sulfur isotopic composition of pyritic- and organically bound sulfur shows a +27‰ excursion across the boundary. The observation that the δ34S values of organic-bound sulfur closely resemble that of pyrite sulfur indicates a common sulfur source, likely early diagenetic sulfide. A change in the δ13C of total dissolved inorganic carbon as a consequence of an enhanced burial of 12C-enriched organic carbon is indicated by a +3‰ excursion measured for TOC as well as for individual n-alkanes and isoprenoids. The burial of large amounts of organic carbon is expected to result in a decrease in pCO2 and should affect the photosynthetic carbon isotope fractionation (εp). The fact that we observe no change in εp can be explained by the circumstance that εp was most probably at maximum values, as a consequence of high atmospheric and oceanic-dissolved CO2 concentrations during the Devonian.
The Late Devonian mass extinction event near the Frasnian/Famennian (F/F) boundary has been analysed using conodont biostratigraphy and biofacies, sedimentology, magnetic susceptibility and geochemistry in reference sections of the South Polish–Moravian shelf (Holy Cross Mountains, Cracow and Brno areas). High-resolution biostratigraphic study revealed difficulties in the precise recognition of this ‘natural’ stage boundary, but confirmed the occurrence of a major (third-order) sequence boundary in the F/F transition in an active synsedimentary tectonic setting, marked by erosional discontinuities, hardgrounds and brecciation or omission surfaces. Conclusive evidence of an extraterrestrial impact has not been found. Among Earth-bound factors, the main devastating role in the shelf habitats is ascribed to fluctuating anoxia and/or nutrient dynamics in a disturbed greenhouse climatic setting. The long-term facies changes were determined by a conspicuous break in carbonate production, accompanied by replacement of mature stromatoporoid–coral reefs by pioneer shelly-crinoid banks, microbial mounds and localised oolitic bars. The key F/F passage interval was marked by intermittent but generally accelerated periplatform ooze/debris input and severe storm events, as well as by probably highly fluctuating oxygenation and biological overproduction, best manifested in radiolarian–silicisponge and cephalopod acmes. Eutrophication phenomena, at least partly stimulated by various hydrothermal and volcanic processes, were one of the major biogeochemical processes during this climax of the Late Devonian biotic crisis. However, significant oceanographic perturbations were also linked with the onset of the transgressive–hypoxic Upper Kellwasser Event, still within the latest Frasnian linguiformis Zone, i.e., well before the F/F conodont collapse. Implied variations in the redox state of seawater through the Kellwasser timespan, probably in regionally highly variable temporal scales, are in agreement with recent palaeoecological and biogeochemical inferences, in particular suggesting recovering oxygenation just prior to the F/F boundary in the other Laurussian intrashelf basins. All the data support a long-time, multicausal Earth-bound crisis instead a worldwide cosmic catastrophe.
The prime cause of the Late Devonian Kellwasser crisis, culminating in a mass extinction event near the Frasnian–Famennian (F–F) boundary, remains conjectural. Nevertheless, rapid sea-level fluctuations of uncertain origin during tectono-eustatic highstand, paired with repeated oceanic anoxia and climatic changes, are usually thought at present to be one of the main immediate triggers. The Cathles–Hallam model of stress-induced changes in plate density, accompanying rapid rift formation, furnishes an alternative for understanding the enigmatic sudden eustatic variations in the non-glacial time. Late Devonian tectonic extension, causing rifting and volcanicity, appears to be strongly marked in several regions of Eurasia, particularly in Kazakhstan and eastern Laurussia. All larger Devonian continents were more or less tectonically affected. A subtle record of this tectonic rearrangement is implied even for distant and apparently quiet carbonate platforms in local extensional block faulting and tilting, hydrothermal mineralisation, geochemical anomalies, and localized blooms of siliceous biota. Interpreting the late Frasnian regressive–transgressive pattern in terms of the Cathles–Hallam tectono-eustatic model, two major rifting events are hypothesized: one at about the beginning of the rhenana Zone, and a second principal pulse in the late linguiformis Zone, that encompassed the F–F transition. Developing the Veimarn–Milanovsky scenario of the global extensional pulse, it is assumed that the key endogenous factors were related to episodic (super)plume activity. The tectonically triggered changes climaxed in thermal and nutrient pulses, and induced the stepdown ecosystem destabilization observed in the F–F bio-crisis. Minor cometary strike(s) might have eventually participated in this prolonged multicausal environmental stress, mainly due to additional thermal shocks, but perhaps effective on a regional scale only.
The intensity and extent of anoxia during the two Kellwasser anoxic events has been
investigated in a range of European localities using amultidisciplinary approach (pyrite framboid assay,
gamma-ray spectrometry and sediment fabric analysis). The results reveal that the development of the
Lower Kellwasser Horizon in the early Late rhenana Zone (Frasnian Stage) in German type sections
does not always coincide with anoxic events elsewhere in Europe and, in some locations, seafloor
oxygenation improves during this interval. Thus, this anoxic event is not universally developed. In
contrast, the Upper Kellwasser Horizon, developed in the Late linguiformis Zone (Frasnian Stage) in
Germany correlates with a European-wide anoxic event that is manifest as an intensification of anoxia
in basinal locations to the point that stable euxinic conditionswere developed (for example, in the basins
of the Holy Cross Mountains, Poland). The interval also saw the spread of dysoxic waters into very
shallow water (for instance, reefal) locations, and it seems reasonable to link the contemporaneous
demise of many marine taxa to this phase of intense and widespread anoxia. In basinal locations,
euxinic conditions persisted into the earliest Famennian with little change of depositional conditions.
Only in the continental margin location of Austria was anoxia not developed at any time in the Late
Devonian. Consequently it appears that the Upper Kellwasser anoxic event was an epicontinental
seaway phenomenon, caused by the upward expansion of anoxia from deep basinal locales rather than
an ‘oceanic’ anoxic event that has spilled laterally into epicontinental settings.
ABSTRACT
The standard Late Devonian conodont zonation is revised and expanded from 29 to 32 zones. In addition to revision of the definition of some zonal boundaries, seven conodont zones, in the middle to late Frasnian are revised and renamed. From oldest to youngest, these are the Early falsiovalis, Late falsiovalis, Early hassi, Late hassi, jamieae, Early rhenana, and Late rhenana Zones. The only newly named and defined early Famennian zone is the Latest crepida Zone. With these changes, 25 of the 32 standard zones are now defined on the basis of first occurrences of species or subspecies of the pelagic genera Mesotaxis, Palmatolepis, and Siphonodella. Four other zones, all late Famennian, are based on first occurrences of species of deep-neritic or pelagic genera, Pseudopolygnathus, Bispathodus, and Protognathodus; these are, from oldest to youngest, the Late trachytera, Middle expansa, Late expansa, and Late praesulcata Zones. Only two zones, the Frasnian Late hassi Zone and middle Famennian Latest marginifera Zone, are based on first occurrences of species of shallow-water genera, Ancyrognathus and Scaphignathus, respectively. All 30 of these conodont first-occurrence zones are considered phylogenetic range zones. The remaining two zones, the Late rhomboidea and Middle praesulcata Zones, based on the extinction of a species or subspecies of Palmatolepis, are interval zones and may not be of as long duration as the other zones. The Late Devonian time scale is revised slightly to accommodate these zonal changes.
Thirty-three taxa belonging to pelagic genera, mainly Palmatolepis, but including three species of Mesotaxis and Klapperina, are treated systematically. Of these, seven are named and described new species of Palmatolepis: Pa. barba n. sp., Pa. ederi n. sp., Pa. eureka n. sp., Pa. jamieae n. sp., Pa. plana n. sp., Pa. rotunda n. sp., and Pa. simpla n. sp. Four new subspecies of Palmatolepis are also named and described: Pa. delicatula platys n. subsp., Pa. gigas extensa n. subsp., Pa. gigas paragigas n. subsp., and Pa. rhenana brevis n. subsp. Recent confusion over identification of the holotypes of Palmatolepis gigas, Pa. subrecta, and Pa. unicornis is resolved. In addition, many changes are made in the rank of species and subspecies, and a problematic, sparsely documented Frasnian taxon, Pa. cf. Pa. delicatula, is described.
The stratigraphically complete and extremely fossiliferous geological sections in the Holy Cross Mountains and Sudetes, Poland, cover the whole history of the Famennian tropical high-diversity pelagic ecosystem. Apparatus reconstruction of 142 conodont species allowed paleobiological interpretation of the faunal succession. Three families, nine genera and 39 species are newly proposed. 76 species of goniatites, with one genus and five species new, and 70 species of clymenias were also identified. Like in all other equatorial localities, a significant (but not catastrophic) decline of diversity marks the beginning of the Famennian. The local pelagic fauna developed mostly as a result of successive reappearances of lineages earlier occurring in the area but temporally removed from it by environmental factors. During the whole Famennian, 101 immigrations of conodont lineages are documented. In 31 of the lineages persisting in the area a more or less complete record of their phyletic evolution is represented; they cover about half (46%) of the summarized ranges of all the lineages. About half of them are suitable for stratophenetic studies. The fossil record of the ammonoids is much more punctuated, but it is estimated that 110 lineages was represented there, only 14 of them possibly evolving phyletically in the area (single case was stratophenetically proven). At the transition between goniatites and clymenias, a succession within the plexus of closely related sympatric species is observed, but the exact phyletic change is not recorded and probably all the first clymenias are immigrants from the east. At least two profound rebuildings of the fauna within the Famennian are observed, but only the terminal Devonian Hangenberg event was of truly dramatic nature. The newly acquired evidence supports the earlier notion thate it is more difficult to trace evolution stratophenetically in the equatorial regions than in high latitudes.
An assemblage of blind phacopid trilobites of the genus Trimerocephalus McCoy, 1849, representing either the species Trimerocephalus mastophthalmus (Reinhard Richter, 1856) or its allies (possibly, a new species), from an Early Famennian (Early Marginifera Zone) marly sequence of the Holy Cross Mountains, Central Poland, is composed of well organized single-file queues. The trilobites in the queues appear almost intact, having been preserved in the position they kept when forming the queues, and are interpreted showing migratory behaviour known in various present-day arthropods, but unreported from the fossil state. This queuing was induced by eironmental stress caused by a dramatic drop in sea level, temporarily leading to emersion. The preservation of the queues at omission horizons is thus ascribed to a mass mortality event, caused by subaerial exposure. The trilobites were suffocated and fossilized in a mortal spasm, and finally blanketed by calcareous ooze when inundated at a highstand. The assemblage of trilobite queues represents a unique example of frozen behaviour and a snapshot of the geological past.
The Kowala section situated in the southern part of the Holy Cross Mountains represents continuous sedimentation in almost the same facies across the Devonian-Carboniferous (D-C) transition. The D-C boundary has been identified about two meters above the top of the cephalopod nodular limestone with Wocklumeria. In the transitional deposits of the latest Famennian (Prothognathodus kockeli Zone) several faunally distinct units that correspond to relative sea level changes in the area have been identified. Ostracods are abundant in the Kowala sequence. Their assemblages contain well known index species and new ones of the Thuringian and Entomozoacean ecotypes. A total of 15 probably planktonic entomozoaceans, and 64 benthic species have been identified. Healdia shangquii sp. n. and Mauryella polonica sp. n. are proposed. A major change in the ostracod fauna takes place above the limestone with Wocklumeria within the transitional interval represented by clays and claystones with tuffites in its middle part. Thuringian and Entomozoacean ecotype ostracods disappear and are replaced by more shallow water 'exotic' assemblage dominated by Healdia, Mauryella and Monoceratina species. In the early Tournaisian rocks Thuringian-, Entomozoacean- and Bairdia-type ostracods reappear with some of the same species as before, and with new Carboniferous index taxa.
The Devonian and Early Carboniferous succession in the Holy Cross Mts. records a steady sea level rise and is bounded by major angular unconformities. The Lower Devonian is developed as a terrigenic sequence of complex, continental and shallow-marine facies. Growth of a shallow-marine carbonate platform in the Middle to Upper Devonian resulted in separation of two adjacent intrashelf basins. This central swell was transformed into a pelagic carbonate platform and influenced the pattern of facies and thickness after its drowning in the Visean. -Author
Five sections from the Kielce Region of the Holy Cross Mountains and one core from a borehole drilled in the nearby western Mesozoic Margin reveal the best recognized deposits from the Devonian/Carboniferous boundary interval. In four sections: Ruda Strawczynska, Bolechowice IG 1, Zargby IG 2 and Kowa la, the Devonian/Carboniferous boundary is located within a continuous series of deep-water carbonates, whereas in the two remaining: Ostrowka Quarry and the Jablonna IG 1 borehole, the Devonian/Carboniferous boundary interval contains stratigraphic gaps resulting from lack of deposition in submarine conditions. The biostratigraphic position of the analysed uppermost Devonian and lowermost Carboniferous succession is determined by assemblages of conodonts, microspores and deep-water ostracodes of the Thuringian ecotype. A distinctive succession of contemporaneous fauna assemblages and lithofacies types is traceable from Holy Cross Mts. to Rhenish Massif. The main extinction occurred during a transgression-promoted anoxia, which then reverted in glacigenic regression. The anoxic conditions are well-expressed at Bolechowice and Kowala successions.
Carbonate cobbles from the lower Famennian (crepida conodont Zone) brachiopod shell beds of the Russkiy Brod Quarry (Central Devonian Field, Russia) have been investigated with respect to paleoecology and paleoenvironment. The cobbles, composed of similar shell bioclasts as the host deposits, were eroded from lithified shell beds during intervals of non-deposition. The smooth surfaces and the presence of borings and encrustations on all sides suggest that the cobbles were periodically transported and overturned.
Two pachyosteomorph arthrodires are described from the Upper Devonian of North America, Dunkleosteus raveri sp. nov. and Dunkleosteus amblyodoratus sp. nov. (from the Appalachian and the Michigan basins, respectively). Dunkleosteus raveri sp. nov. is found stratigraphically below the only other species within the Ohio Shale Formation (Famennian), and is characterized by the retention of a primitive dermal ornamentation, and shares with Dunkleosteus terrelli (Newberry, 1873) the possession of articular facets on the parasphenoid. Dunkleosteus amblyodoratus sp. nov. (Kettle Point Formation, Upper Devonian), known only from incomplete nuchal, paranuchal, and parasphenoid plates, also retains dermal tuberculation, and is characterized by tapering articular facets on the parasphenoid. The current study reviews three North American species of Dunkleosteus (Du. terrelli, Du. raveri sp. nov., and Du. amblyodoratus sp. nov.). Formerly, two subgroups were recognized within the Dinichthyidae, the terrelli and herzeri groups, which are shown here to be members of distinct clades. Dinichthys herzeriNewberry, 1868 (the namesake member of the herzeri group) is now the sole member of the family Dinichthyidae comb. nov., and is assigned to the Aspinothoracidi sensuMiles & Dennis, 1979. HeintzichthysWhitley, 1933 and GorgonichthysClaypole, 1892 (additional members of the herzeri group) were previously placed among the aspinothoracid arthrodires. Likewise, Holdenius holdeniDunkle & Bungart, 1942 (the final member of the herzeri group) and Hadrosteus rapaxGross, 1932 are now placed within the Aspinothoracidi. The remaining taxa, formerly assigned to the Dinichthyidae (i.e. DunkleosteusLehman, 1956; EastmanosteusObruchev, 1964; GolshanichthysLelièvre, Janvier, & Goujet, 1981; and WestralichthysLong, 1987), are united within the Dunkleosteidae Stensiö, 1963.
© 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 159, 195–222.
Integrated palynological, organic and inorganic geochemical and petrographical methods have been used for deciphering the depositional redox conditions and character of organic matter of the Famennian Dasberg event horizon from the deep-shelf Kowala succession of the Holy Cross Mountains. The ages of the investigated samples have been established, using miospore data, as VF (Diducites versabilis–Grandispora famenensis) and LV (Retispora lepidophyta–Apiculiretusispora verrucosa) miospore Zones of the Middle/Upper Famennian. In the standard conodont zonation, this corresponds to the uppermost postera to lowermost praesulcata Zones. The presence of green sulphur bacteria biomarkers and dominance of small-sized framboids together with the presence of large framboids and low values of the U/Th ratio may indicate that during sedimentation of the lower Dasberg shale, intermittent anoxia occurred in the water column, or the anoxic conditions prevailed in the upper part of the water column, while the bottom waters were oxygenated, at least briefly. Deposition of the upper Dasberg shale was characterized by both bottom water and water column anoxia. The lack of acritarcha taxa from these intervals could have been due to anoxia in the photic zone. Moreover, organic content is high in those samples. There is no geochemical evidence for anoxia during sedimentation of the deposits sandwiched between the lower and upper Dasberg shales, or in the deposits which underlie and overlie both Dasberg shale horizons. The two discrete anoxic events are interpreted to be the result of major transgressions and the blooming of primary producers. Above the Dasberg shales, small fragments of charcoal and raised concentrations of polycyclic aromatic hydrocarbons are detected. This supports the presence of wildfires during deposition of shales just above the boundary of VF/LV palynological zones. Temperatures calculated from the fusinite reflectance values suggest that the charcoal was formed in low-temperature ground and/or surface fires. The typical marine character of sedimentation combined with the high proportion of charcoals suggests that wildfires were large-scale, and that there was intensive transport of terrestrial material. The main causes of intensive wildfires were a significant rise of O 2 in the atmosphere and important progress in the land plant diversity during Late Devonian times. Palynofacies studies suggest that the transgression corresponds to the part IIf of the Late Devonian sea-level curve.
A palynological study of the uppermost Famennian section from Kowala Quarry (Holy Cross Mountains, central Poland) allowed recognition of two miospore zones: LV (Retispora lepidophyta–Apiculiretusispora verrucosa) and LN (Retispora lepidophyta–Verrucosisporites nitidus). Based on palynology and sedimentology, the black shale within the upper part of the section is identified as equivalent to the Hangenberg Black Shale, which is known globally. This black shale contains compounds characteristic of photic zone euxinia, including isorenieratane and its derivatives. Such compounds are absent in the organic-poor marls and shales occurring below the LN Zone, and are present only as traces in the layers just above the black shale, indicating fluctuations in the oxygen minimum zone during uppermost Famennian sedimentation. Palynofacies show high amounts of amorphous organic matter and prasinophyte concentrations in the black shale, and a subsequent significant decrease of amorphous organic matter concomitant with a rapid increase of terrestrial input (mainly miospores with common tetrads) in the layers above the black shale. This supports the relatively rapid change in the taxonomic composition of phytoplankton caused by fluctuations of the chemocline. The whole succession corresponds to one sea-level rise and fall. The presence of high concentrations of peri-condensed polycyclic aromatic hydrocarbons and large amounts of small charcoal particles at the Hangenberg event horizon indicate the occurrence of wildfires. Such observations suggest that atmospheric O 2 levels had exceeded the critical threshold of 13 %, above which wildfires may occur, by latest Famennian time.
Presented for the first time in this paper are the results of a detailed multi-proxy investigation conducted on the Hangenberg Black Shale (HBS) from the Polish part of the Laurussian Shelf, which provide details about the environmental conditions in existence during deposition of the shale and the role played by anoxic conditions in the mass extinction events that occurred at the end of the Devonian times. Inorganic and organic redox indicators indicate that bottom water redox conditions changed periodically from being mainly anoxic/euxinic to oxic or being partially depleted in oxygen. U/Th values above 1.25, Ni/Co values above 7 and V/(V + Ni) values above 0.8 recorded from the lower part of the HBS all point to anoxic/euxinic conditions being present, as do high total organic carbon contents (TOC) above 10% and degree of pyritisation (DOP) values around 0.75%. However, the presence of benthic fauna over the lower part of the HBS attests to opportunistic colonisation of the seafloor during oxic episodes. Evidence for similar episodes has also been recognised over the middle part of the HBS, e.g., U/Th values below 1.25, Ni/Co values below 4, V/(V + Ni) values below 0.8, TOC values of 3% to 5.5% and DOP values of 0.4% to 0.75%, plus the common occurrence of benthic fauna. Anoxic conditions returned during the deposition of the upper part of the HSB, though they were not as well developed as when the lower part of the shale was laid down.
The Upper Devonian sequence at Kowala in the Holy Cross Mountains was logged using gamma-ray spectrometry,
for investigating the changes of oxygenation level in the Late Devonian basin. The Th/U ratio indicates that oxygen
levels were low throughout the Late Frasnian interval, with low peaks during the Kellwasser Events showing anoxic
conditions in the basin. The F-F boundary interval was also oxygen deficient, but there may have been a brief reoxygenation
at the boundary itself. By the Famennian crepida Zone, the basin gradually began to reoxygenate, but in the
trachytera Zone another anoxic event, the Annulata Event occurred, causing a bloom rather than extinction of specially
adapted taxa such as Guerichia. Thus the gamma-ray spectrometry data suggests that basinal anoxia prevailed
through much of the Late Frasnian. The F-F extinction might have been the result of prolonged stresses imposed on
the ecosystem, particularly during the euxinic Upper Kellwasser Event.
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