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Famennian corals of the south-western part of the Holy Cross Mountains (Góry Świętokrzyskie) from Kadzielnia, Zaręby, Łagów (Dule), Gałęzice (Besówka), Kowala and Jabłonna are described in the present work. About 3,000 specimens have been collected and 55 species and 5 subspecies of 36 genera - described. A new family, Kielcephyllidae, two new subfamilies, Friedbergiinae and Guerichiphyllinae, 9 new genera, 36 new species and 5 subspecies have been erected. On the basis of a lithological analysis of beds, faunal assemblages and the preservation state of corallites, it has been concluded that in the Lower Famennian a shallow sea existed at Kadzielnia. It was temporarily connected with the open sea. The corals are mostly preserved there in a life-time place. At Zaręby, there was a lagoon with the remains of plants but with a normal salinity of water. In the Upper Famennian of Gałęzice, the sea water was well aerated and connected with the open sea. Fossils are broken but not worn off and, therefore, they were not transported. At Kowala, the sea was probably calm, not deep and well aerated. Index species for the Lower and Upper Famennian have been determined among the coral fauna examined. The age of individual zones has been determined by conodonts and where such were lacking, climeniids or trilobites. The history of the research of Famennian corals, which in general occur rarely and in monotonous assemblages, has been presented. To settle their generic assignment, the ontogeny of corals has been studied and the results were a basis for changes introduced to the classification. A few phyla have been distinguished and their phylogenetic relationships discussed.
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The first Devonian holocephalian tooth from Poland

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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 chondrichthyan 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 paper describes a new species of blind trilobite from the lower Fammenian of Concours−le−Haut at Causses−et−Veyran, Montagne Noire (France). Trimerocephalus (Trifoliops) nigritus subgen. et sp. nov. is assigned to a new subgenus together with Tr. (Trif.) trifolius (Osmólska, 1958). This grouping is supported by the results of phylogenetic analysis of thirteen species attributed to the Fammenian genus Trimerocephalus McCoy, 1849; of 16 previously known species attributed to this genus, only 12 were represented by data of quality sufficient to be included in the analysis, using 23 morphological characters. The Frasnian phacopine Acuticryphops acuticeps (Kayser, 1889) is used as the outgroup. The three most parsimonious trees have a length of 51 steps and a consistency index of 0.82. The new subgenus Trifoliops forms a clade together with Trimerocephalus? steinachensis (Richter and Richter, 1926), supported by an exclusive synapomorphy: widening of the cephalic antero−lateral border. Tr.? steinachensis seems to be more closely related to Tr. (Trif.) trifolius (sharing two synapomorphies) and may represent a more derived taxon (possibly deserving a separate subgeneric status). The remanig Trimerocephalus species are not formally assigned to subgeneric taxa, pending further studies (their relationships are shown in cladograms). The results do not confirm the classification suggested by Chlupač (1966) for Trimerocephalus.
Disarticulated crinoid columnals and pluricolumnals from the Famennian of the Holy Cross area were analysed. Sixteen crinoid taxa were distinguished, only one of which is based on stems attributed to a calyx−based genus, and the others are classified within artificial supraspecific units. Two of these are new: Schyschcatocrinus levis sp. nov. and Cosmocrinus polonicussp. nov. The described crinoid fauna shows distinct extinction−recovery temporal pattern: the Frasnian–Famennian crisis affected 50% of stem−based families and 70% of late Frasnian stem−based genera. The succession of crinoid faunas represented by three faunal intervals has been identified and correlated to standard conodont zones: FIa, Palmatolepis triangularis Zone (relic “Frasnian” crinoid assemblage Schyschcatocrinus delicatus–Calleocrinus kielcensis), FIb, Pa. crepida to Pa. marginifera zones (crinoid assemblage Calleocrinus kielcensis–Schyschcatocrinus levis) and FIc, Pa. trachytera to S. praesulcata zones (crinoid assemblage Cosmocrinus polonicus–Acbastaucrinus affectatus). The succession was controlled mostly by eustatic factors.
New materials on the bothriolepidid Bothriolepis zadonica H.D. Obrucheva, 1983 from the Zadonskian Regional Stage (Upper Devonian, Lower Famennian) of Central Devonian Field (Central European Russia) are described and illustrated. These materials came from nine localities in the Orel and Tver Regions of Russia. This species is characterised by the presence of a well developed median dorsal crest in the trunk armour and unusual type of the preorbital recess of the head shield; this recess is designated herein “trapezoid”. The ontogenetic and individual variations of B. zadonica are studied. The crested bothriolepidids are known only from Euramerica and East Gondwana. The possible relationships between crested bothriolepidids in these provinces are discussed.
The new trench Z−17 situated in the Dębnik anticline (Cracow Region, southern Poland) exposed strata representing the Frasnian–Famennian boundary interval. The latest Frasnian crisis interval is characterized by the fauna of Ryocarhynchus tumidus interval consisting of the nominal species, Barroisella campbelli, Biernatella polonica, and representives of Lingulipora, Athyris, ?Retichonetes, Longispina, Cyrtospirifer and Warrenella. The first brachiopods that appears immediately above the F–F boundary in the survival interval include the unidentified rhipidomellid, Praewaagenoconcha cf. speciosa, and Pampoecilorhynchus geniculatus sp. nov. The fauna in the succeeding layer is dominated by P. geniculatus and Cyrtospirifer minor. The earliest Famennian repopulation assemblage consists of representatives of Barroisella, ?Rhyssochonetes, Praewaagenoconcha, Nigerinoplica, Schizophoria, unidentified rhipidomellid, Pampoecilorhynchus geniculatus sp. nov., Chapinella striata sp. nov., Crinisarina angelicoides, and Cyrtospirifer minor. Low diversity and low frequency brachiopod assemblages consisting of stunted specimens characterize the succeeding brachiopod intervals spanning the remainder of the Palmatolepis triangularis Zone. Basinal dysoxia prevailed in the region for the duration of Pa. triangularis Zone. Resumption of aerobic bottom−water conditions is marked by the appearance of brachiopods of the Dmitria gibbosa interval (Pa. crepida Zone). Resumption of favorable environmental conditions during the recovery interval is marked by an increase in brachiopod diversity recording the beginning of a strong post−extinction rediversification of the fauna.
For the past three decades, the Alvarez impact theory of mass extinction, causally related to catastrophic meteorite impacts, has been recurrently applied to multiple extinction boundaries. However, these multidisciplinary research efforts across the globe have been largely unsuccessful to date, with one outstanding exception: the Cretaceous–Paleogene boundary. The unicausal impact scenario as a leading explanation, when applied to the complex fossil record, has resulted in force−fitting of data and interpretations (“great expectations syndrome”). The misunderstandings can be grouped at three successive levels of the testing process, and involve the unreflective application of the impact paradigm: (i) factual misidentification, i.e., an erroneous or indefinite recognition of the extraterrestrial record in sedimentological, physical and geochemical contexts, (ii) correlative misinterpretation of the adequately documented impact signals due to their incorrect dating, and (iii) causal overestimation when the proved impact characteristics are doubtful as a sufficient trigger of a contemporaneous global cosmic catastrophe. Examples of uncritical belief in the simple cause−effect scenario for the Frasnian–Famennian, Permian–Triassic, and Triassic–Jurassic (and the Eifelian–Givetian and Paleocene–Eocene as well) global events include mostly item−1 pitfalls (factual misidentification), with Ir enrichments and shocked minerals frequently misidentified. Therefore, these mass extinctions are still at the first test level, and only the F–F extinction is potentially seen in the context of item−2, the interpretative step, because of the possible causative link with the Siljan Ring crater (53 km in diameter). The erratically recognized cratering signature is often marked by large timing and size uncertainties, and item−3, the advanced causal inference, is in fact limited to clustered impacts that clearly predate major mass extinctions. The multi−impact lag−time pattern is particularly clear in the Late Triassic, when the largest (100 km diameter) Manicouagan crater was possibly concurrent with the end−Carnian extinction (or with the late Norian tetrapod turnover on an alternative time scale). The relatively small crater sizes and cratonic (crystalline rock basement) setting of these two craters further suggest the strongly insufficient extraterrestrial trigger of worldwide environmental traumas. However, to discuss the kill potential of impact events in a more robust fashion, their location and timing, vulnerability factors, especially target geology and palaeogeography in the context of associated climate−active volatile fluxes, should to be rigorously assessed. The current lack of conclusive impact evidence synchronous with most mass extinctions may still be somewhat misleading due to the predicted large set of undiscovered craters, particularly in light of the obscured record of oceanic impact events.
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.
Throughout their history, species had to face environmental variations spatially and temporally. How both levels of variation interact will be of key importance in conditioning their response to major perturbations. We addressed this question by focusing on a period in Earth’s history marked by dramatic environmental and faunal changes, the Late Devonian Frasnian/Famennian boundary. From a paleogeographic point of view, this period is characterized by a cosmopolitanism of the faunas across a large ocean, the Prototethys. We considered the biotic reaction at a seldom considered scale, namely within a single subgenus of conodont, Palmatolepis (Manticolepis). Patterns of spatial and temporal differentiation were quantified using morphometrics of its platform element. The recognized cosmopolitanism of the faunas was confirmed at this scale of variation since temporal records gathered in distant areas around the Prototethys, including the seldom documented regions located nowadays in South−East Asia, displayed similar morphological trends in response to the major F/F crisis. Beyond this overall cosmopolitanism, subtle geographic structure was evidenced but was not stable through time. Geographic differentiation was maximal shortly before the F/F crisis, suggesting that despite high sea−level, tectonics leaded to complex submarine landscapes promoting differentiation. In contrast any geographic structure was swamped out after the crisis, possibly due to a global recolonization from few favorable patches.
An uppermost Famennian (Strunian) coral assemblage has been recovered in the middle part of the Yılanlı Formation of the Istanbul Zone (Zonguldak and Bartın areas, NW Turkey). In the Bartın area, the studied fossiliferous interval corresponds to a c. 30 m-thick unit of bioclastic to peloidal wackestone to packstone grading to grainstone and including two stromatoporoid biostromes. In the Zonguldak area, 60 km westward, the bioclastic facies is dominant. The rugose corals are mainly solitary taxa belonging to the genera Campophyllum, Bounophyllum, Amplexocarinia, and ?Metriophyllum, and only one colonial genus occurs: Pseudoendophyllum. This fauna is similar to that documented in Europe. The campophyllids and dibunophyllids are the main component of the uppermost Famennian assemblages in S Belgium, N France, W Germany, NW and S Poland. The endophyllids occur in S Poland, Novaya Zemlya, and in the Ural Mountains. The Istanbul Zone is supposed to be situated in the central part of the Palaeotethys Ocean, along the southern margin of Laurussia during the uppermost Devonian and Carboniferous. The rugose corals indicate some relationship with the eastern part of Laurussia, or that both areas were under a common marine influence at this time. The global Hangenberg event was not recognized in the Turkish localities, except for the disappearance of the corals, occurring less than 19 m below the Devonian–Carboniferous boundary based on the foraminifers. There is no major facies change through the boundary and the first Carboniferous corals (small Uralinia and Caninophyllum) appear 6 m above the D–C boundary. The new species Caninophyllum charli sp. nov. is described from the upper part of the lower Tournaisian.
Famennian Stromatoporoidea from the Quasiendothyra communis Foraminiferal Zone and slightly younger strata from the Dębnik anticline, southern Poland, form a succession of three consecutive assemblages. Assemblages 1 and 3 consist of representatives of the order Clathrodictyida, while assemblage 2 is dominated by the order Labechiida. The clathrodictyids are represented by the genus Gerronostroma, and labechiids are represented by the genus Stylostroma. Species assigned here to the genus Gerronostroma show a network of amalgamated pillars in the central part of the columns, a feature regarded by previous authors as typical of the genus Clavidictyon. Two new species, Stylostroma multiformis sp. nov. and Gerronostroma raclaviense sp. nov., are described. Stromatoporoids from southern Poland differ from the Famennian fauna of western Europe, showing affinity to eastern European and Siberian Stromatoporoidea.
The Frasnian–Famennian (F–F) boundary is well biostratigraphically documented in the Palmatolepis−rich deposits exposed along the Syv’yu River in the lower slopes of the Subpolar Urals. The thin−bedded calcareous−clayey−siliceous deep−slope succession of the Vorota Formation appears to represent continuous Domanic−type deposition throughout the world−wide carbonate crisis time, without evidence for the basal Famennian hiatus or a large−scale sedimentary perturbation within a regressive setting. The northernmost Laurussian sequence exhibits many well known signatures throughout the broad F–F timespan: the appearance of organicand clay−rich deposits, icriodontid and radiolarian blooms, and a correlative shift of several geochemical proxies towards hypoxic and high−productivity regimes, perfectly recorded by positive 13Ccarb excursions of +3.5‰. Integrative biotic, microfacies and geochemical data substantiate a longer−term oceanographic destabilization, attributable to multiple Earth−bound triggering factors in (episodically enhanced?) greenhouse climate and punctuated eustatic sea−level highstands, superimposed on the elevated deposition of organic carbon−rich sediments during the Upper Kellwasser Event. Unsteady eutrophicated, and oxygen−depleted ecosystems during the F–F biotic crisis interval could be assumed, especially when intensified by various spasmodic tectono−volcanic phenomena in the incipiently closing Ural Ocean.
Abundant trilobite remains were recovered from late mid−Famennian marlstones from various sites in Eastern Tafilalet, southeast Morocco. All belong to a single taxon previously identified as Cyrtosymbole (Waribole) prima. This taxon is designated the type species of Osmolskabole gen. nov. A redefinition of this species, including the description of newly discovered, disarticulated exuviae both in limestone and silicified state of preservation, is given. In particular, silicified sclerites of various sizes allow the first complete growth series of a cyrtosymboline proetid to be presented. The close morphological resemblance of its protaspid stages to known proetoid larvae emphasizes the homogeneity of the early ontogeny in this superfamily. The Famennian proetoid anaprotaspis is also of comparable size to that of other Devonian proetoid larvae. However, their size−range is much less than that observed in Carboniferous larvae. This suggests that the survival of proetoid trilobites at the Frasnian−Famennian Kellwasser crisis did not result from a modification of the developmental strategy, as it might have been the case at the terminal Devonian extinction event. Moreover, O. prima possesses a plectrum from the metaprotaspid to the mid meraspid periods. This implies that the natant hypostomal condition is not steadily acquired early in the ontogeny of the Proetida. Thus we preclude the use of this character in the diagnosis of this order.
An earliest Famennian (Late Devonian) shell bed was discovered in the Hanover Shale Member of the Java Formation 1.4 meters above the Frasnian–Famennian (F–F) boundary in western New York. The invertebrate shelly fauna of the shell bed (Lower Palmatolepis triangularis conodont Zone), provides information on taxonomy of an outer shelf benthic association during the survival interval of recovery in the Appalachian foreland basin soon after the terminal Upper Kellwasser event marking the F–F mass extinction. Shelly invertebrates are extremely rare in the upper Hanover immediately above and below the shell bed. Abundance of brachiopod valves and remains of other groups in the shell bed reach 80–100 valves/100 cm2. Elongate valves of the linguloid brachiopod Barroisella cf. B. campbelli have preferred alignments roughly parallel to direction of down−slope flow in the deep−water foreland basin depositional setting. The brachiopod fauna is dominated by the representatives of Retichonetes, Barroisella, Cyrtospirifer, Tylothyris, and Praewaagenoconcha. Rare elements include species of Thiemella, Schizophoria, Ripidiorhynchus?, Chapinella?, an indeterminate rhynchonellid, Ambocoelia, and extremely rare Orbiculoidea. Forms including Cyrtospirifer hornellensis, Tylothyris mesacostalis, Praewaagenoconcha speciosa, and few others are late Frasnian carryovers. The range inception of Thiemella leonensis is just above the F–F boundary (Upper Kellwasser horizon) in the upper Hanover Shale shell bed in the western Appalachian foreland basin.
Late Frasnian Atrypida (Brachiopoda) from the South Urals, South Timan and Kuznetsk Basin in Russia (east Laurussian and south Siberian shelf domains in Devonian time) reveal significant generic and specific diversity in the broadly defined Frasnian-Famennian (F-F) bio-crisis time. Eighteen species of atrypid brachiopods have been recorded, representing 4 subfamilies and 10 genera. The new genus Gibberosatrypa Markovskii & Rzhonsnitskaya, and the new subgenus Spinatrypa (Plicspinatrypa) Rzhonsnitskaya are proposed. Four new species Spinatrypina (Spinatrypina) sosnovkiensis Yudina, Spinatrypa (Plicspinatrypa) rossica Rzhonsnitskaya, Iowatrypa nalivkini Rzhonsnitskaya & Sokiran, and Cartnatina(?) biohermica Yudina are described. The representatives of the Variatrypinae (including especially common Desquamatia (Desquamatia) alticoliformis), Spinatrypinae (Spinatrypina) and Atypinae (Pseudoatrypa, ?Costatrypa) are widely distributed in the studied regions. The Pseudogruenewaldtiinae are represented by Iowatrypa and Pseudogruenewaldtia, of which the first is distributed worldwide, whereas the only undoubted species of the second is restricted to South Timan, and probably represents a localized latest Frasnian descendant of Iowatrypa. The decline phase of atrypid development was controlled by a variety of environmental factors tied to the global Kellwasser events, although it was not directly triggered by anoxic conditions. The investigated atrypid brachiopods, which were all confined to lower latitudes, disappeared during the F-F mass extinction, independently of their environmental and biogeographic settings.
Four major microfacies have been recognized in the Psie Górki section and the bioclastic content indicates an open marine environment in the photic zone close to an algal shole. Sedimentological studies point to a regressive episode starting close to the Frasnian–Famennian boundary. The regressive microfacies pattern is revealed by the presence of semirestricted algal microbreccias that compose all of the lower part of the Famennian. The regression was accompanied by meteoric water invasion as the sea level fell. Seventy−six ostracod species are recorded. The ostracod assemblage, dominated by podocopids, belongs to the Eifelian ecotype and is indicative of a well−oxygenated marine environment below fair−weather wave base in the Frasnian part of the section, and of shallower environments in the base of the Famennian. No ostracod assemblage characteristic of hypoxic or semi−restricted water conditions has been recorded. The rate of extinction of ostracod species (>70%) close to the Frasnian–Famennian boundary is comparable with that known on the same level in several other sections investigated in the world. Five new ostracod species are proposed by J.−G. Casier and F. Lethiers: Selebratina vellicata, Samarella? minuta, Bairdiocypris ventrorecta, Acratia pentagona, and “Bairdia” psiegorkiensis.
The hitherto poorly known type species of Cyrtosymbole, i.e., Dechenella escoti, is redescribed on the basis of an abundant new material recovered from the early Famennian Lower Griotte Limestone Formation at La Tourière, near Cabrières, southern France. It includes sclerites (hypostome, librigenae, thoracic segments and external surface of the pygidium) that previously were either unknown, or represented by poorly preserved and incomplete specimens, together with a full suite of post−protaspid growth stages. The latter has revealed that certain characters, in particular the preglabellar region and postocular facial sutures, show marked changes between the early and late holaspid stages. In the past, some species have been based on immature specimens; for example the lectotype of C. escoti is an early holaspis, and the characters that it displays have been regarded as diagnostic of the genus. For confident specific assignments in cyrtosymboline trilobites it is important, therefore, to have to hand sufficient material, including late holaspids. A revised diagnosis of Cyrtosymbole is given, and only those species that share diagnostic adult characters with C. escoti are assigned to it. Insights into early growth development exemplified by C. escoti and allies corroborate the attribution of the Cyrtosymbolinae to the Proetidae.
This paper reports on the evolution of ammonoids belonging to the family Tornoceratidae from the Devonian of Janczyce in the Holy Cross Mountains, Poland. Steady and gradual changes in conch morphology of the goniatite lineage Phoenixites frechi–Tornoceras subacutum–T. sublentiforme occurred in concert with water shallowing during the deposition of the Lower Famennian cephalopod limestone. Biometric analysis of ammonoid conch and facies analysis of the cephalopod limestones have been applied to assess the possible relationship between shell geometry and environmental changes. Results show that ratios of whorl width / diameter as well as whorl width / whorl height decreased, while distance from the venter to the greatest whorl width / diameter increased with time, thereby reducing hydrodynamic drag of the shells, probably in response to increasing water turbulence. The interpretation presented here is in agreement with similar cases from the literature. However, this kind of environmentally controlled evolution has hitherto been recognized only in Jurassic and Cretaceous ammonoids. Conch morphology may be considered as an indicator of palaeobathymetry.
We present taxonomic revision of rugose corals and brachiopods from several Frasnian–Fammenian (F–F) boundary sections in central Hunan Province, China. Diversity of shallow−water rugose corals gradually increased during the Frasnian, but ended with sudden extinction near the end of Frasnian. Ostracods were abundant during the Frasnian; their extinction coincided with anoxic deposition of the end−Frasnian black shale deposits. The early Famennian ostracod fauna is of low diversity. The brachiopod fauna of the late Frasnian (Palmatolepis rhenana and Pa. linguiformis zones) is dominated by atrypids, small−sized cyrtospiriferids, and the rhynchonellid Hunanotoechia. All atrypids disappeared before the F–F boundary with highest rates of extinction below the boundary (probably low in the Pa. linguiformis Zone). The Frasnian cyrtospiriferid fauna is also of low diversity and dominated by small taxa. All but one of the cyrtospiriferid taxa crossed the F–F boundary. The early Famennian post−extinction recovery brachiopod fauna was the result of rapid radiation of new forms shortly after the terminal Frasnian event. The early Famennian fauna is characterized by diverse cyrtospiriferids, abundant Yunnanellina and productoids. Above the early recovery fauna another fauna was recovered, with brachiopods Hunanospirifer and Yunnanella and is correlated with the late or latest Pa. crepida Zone. Sinalosia rugosa gen. et sp. nov. (Productida) is erected.
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