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A new atrypid genus Heckerella from early Frasnian (Palmatolepis transitans Zone) of northwestern East European Platform is proposed. It includes only Heckerella heckeri, originally referred to Anatrypa. The new genus is characterised by weakly paraplicate anterior commissure, well-developed carina on the ventral valve, sulcus on the dorsal valve and coarser radial ornament. Heckerella has restricted geographic distribution in northwestern Russia, Latvia, northeastern Lithuania where it forms high-density associations with Pseudoatrypa velikaya.
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.
The brachiopod fauna of the Middle-Late Devonian cratonic carbonate platform deposits of the Iowa Basin, central North America, contains twenty species of the order Atrypida, some of which are types for widespread genera common in Middle and Late Devonian faunas. The latest Givetian-early Frasnian deposits yield a diverse fauna consisting of ten species (two new) included in Desquamatia (Independatrypa), D. (Seratrypa), Pseudoatrypa, Radiatrypa, Spinatrypa (Spinatrypa) and Spinatrypina (Exatrypa). Many of these forms occur in, or are closely similar to species known from, coeval faunas of central and western Canada. Middle Frasnian deposits of northern Iowa contain two species included in Spinatrypa (S.) and Pseudoatrypa, both of which are new. Late Frasnian strata of the Iowa Basin yield eight species included in Costatrypa, Iowatrypa, Pseudoatrypa, and Spinatrypa (Spinatrypa), some of which are widespread in other subtropical and tropical faunas of the western US and western Canada. The taxa Pseudoatrypa witzkei sp. n., Spinatrypa (S.) bunkeri sp. n., Spinatrypa (S.) thompsoni sp. n., and Spinatrypina (Exatrypa) johnsoni sp. n. are proposed. Pseudoatrypa? sp. from the very late Frasnian of southern New Mexico is also illustrated.
Late Frasnian–Early Famennian entomozoacean ostracod assemblages from the Płucki section in the Holy Cross Mountains were studied to establish the effect of the “Kellwasser bio−event” on the planktonic biodiversity and faunal content. The composition of ostracod assemblages changes from a moderately diverse (10 species) Entomoprimitia–Richterina– Nehdentomis–Nandania dominated “background” assemblage characterising a pre−event interval, to an Entomoprimitia−assemblage during the event interval, and finally to a Franklinella−dominated post−event assemblage in the Middle Palmatolepis triangularis conodont Zone. The Frasnian–Famennian extinction caused substantial losses among entomozoacean lineages. In the Płucki section it occurred in two closely spaced steps within the Palmatolepis linguiformis conodont Zone. The first step, at the base of the dark cephalopod limestone (Upper Kellwasser Horizon), reduced the abundance and the species diversity of entomozoaceans to only two Entomoprimitia species. The vacant niche was then filled by the new, immigrant species Entomoprimitia (Entomoprimitia) kayseri which is dominant in the Upper Kellwasser interval. All these species were lost at the second step within the Upper Kellwasser Horizon. The entomozoaceans remained virtually absent during a long time interval between the end−Frasnian crisis and the Middle Pa. triangularis Zone. They reappear as new species from refugia lineages (Franklinella, Nehdentomis) and became widespread, indicating favourable ecological conditions. Some 13 species have been identified and assigned to seven genera. Rabienella? lagowiensis sp. nov. is proposed.
The fin rays and two types of scales (enlarged and regular) of the Devonian sarcopterygian Eusthenopteron foordi are redescribed using light, scanning and transmission electron microscopy. The fin rays consist of lepidotrichia composed of ossified, jointed and branched segment pairs. The basal segments are cylindrical, but more distal elements are crescentic in section. The distribution of Sharpey’s fibres varies along the lepidotrichia. In the proximal segments, lateral bundles form a belt connecting adjacent hemisegments. In the distal segments, thin bundles are restricted to the area facing the fin surface. Both enlarged and regular scales have a similar spatial organisation. They are composed of a superficial highly mineralised layer covering a thick basal plate where the fibrils are distributed in superimposed strata forming a plywood−like structure. Nevertheless, the enlarged and regular scales differ in their shape, in the mineralised tissues of the superficial layer, and in the organisation of the plywood−like structure. The superficial layer of the enlarged scales is composed of parallel−fibered bone covering a deeper layer of woven−fibered bone. The basal plate is made of an orthogonal plywood−like structure. The thin, lamellar, imbricated regular scales display the characteristics of elasmoid scales. The mineralised tissue forming the superficial layer resembles that of extant teleost scales. In the basal plate, the twisted plywood−like structure is composed of closely packed fibrils that are preserved down to the ultrastructural level owing to the persistence of bridges connecting the fibrils. The enlarged and the regular scales of Eusthenopteron foordi do not present superficial odontodes, in contrast to ancestral thick rhomboid scales. The disappearance of enamel/enameloid and dentine may be related to the evolutionary trend towards a lightening of the dermal skeleton that would improve the swimming abilities of the animal. The characteristics of the dermal skeleton of Eusthenopteron foordi support the hypothesis that this process began early in osteichthyans.
The Frasnian-Famennian (F-F) mass extinctions saw the global loss of all genera belonging to the tropically confined order Atrypida (and Pentamerida): though Famennian forms have been reported in the literafure, none can be confirmed. Losses were more severe during the Givetian (including the extinction of the suborder Davidsoniidina, and the reduction of the suborder Lissatrypidina to a single genus), but origination rates in the remaining suborder surviving into the Frasnian kept the group alive, though much reduced in biodiversity from the late Early and Middle Devonian. In the terminal phases of the late Palmatolepis rhenana and P. linguiformis zones at the end of the Frasnian, during which the last few Atrypidae dechned, no new genera originated, and thus the Atrypida were extirpated. There is no evidence for an abrupt termination of all lineages at the F-F boundary, nor that the Atrypida were abundant at this time, since all groups were in decline and impoverished. Atypida were well established in dysaerobic, muddy substrate, reef lagoonal and off-reef deeper water settings in the late Givetian and Frasnian, alongside a range of brachiopod orders which sailed through the F-F boundary: tropical shelf anoxia or hypoxia seems implausible as a cause for atrypid extinction. Glacial-interglacial climate cycles recorded in South America for the Late Devonian, and their synchronous global cooling effect in low latitudes, as well as loss of the reef habitat and shelf area reduction, remain as the most likely combined scenarios for the mass extinction events.
The radiolarian species Astroentactinia paronae, A. stellata, Trilonche echinata, T. grandis, T. nigra, Haplentactinia inaudita, and H. rhinophyuosa are common in late Frasnian to early Famennian rhythmic, calcareous−marly sequence of the southern Holy Cross Mts., Poland. They are known also from coeval abundant siliceous biota assemblages from the carbonate shelf of East European Platform including more than 150 taxa of radiolarians. However, in ecological terms, the moderately diverse Polish microfaunas (34 species of 12 genera) are more similar to these from Kolyma and Alaska, also marked by abundance of sphaerical entactiniids and near−absence of bilateral−symmetric Ceratoikiscidae and Palaeoscenididae. A succession of two distinctive siliceous sponges associations is established in the incipiently submerged Holy Cross carbonate platform: from an ephemeral, diverse, mostly rigid−skeletal lithistid−hexactinosan foreslope assemblage (initial phase of the late Frasnian Kellwasser Crisis), to long−lasting, basinal loose−skeletal hexactinellid−demosponge faunas (appearing abundantly just prior the Frasnian–Famennian boundary in the late Palmatolepis linguiformis Zone). Such regional blooms of marine siliceous biotas, parallel to temporary retreat of calcareous biota, are demonstrated worldwide for the Kellwasser Crisis. These suggest probable causal links with cooling pulses and at least regional, volcanically induced eutrophication.
Late Frasnian representatives of the order Athyridida from the Holy Cross Mountains, Poland, support the idea that the Laurussian basins were the places of origin and radiation of the subfamilies Athyridinae and Meristinae during the middle and early late Paleozoic. At least three new species have been identified from two localities (Łgawa Hill and Kowala) in the Gałęzice Syncline. Of these, one was probaby endemic (Merista rhenanensis sp. n.; maybe also ?Zonathyris sp. A), and two (Athyris postconcentrica sp. n. and Pachyplaxoides postgyralea gen. et sp. n.) were more widely distributed in this part of the Laurussian shelf, being known also from the East European Platform and Rheinisches Schiefergebirge, respectively. This confirms an intermediate biogeographic position of the Holy Cross Mountains area, belonging to an important centre of brachiopod origin and diversification. In contrast to other articulate brachiopods, athyridids reveal a higher rate of diversification, especially at the species (and partly also generic) level, during the global Kellwasser Crisis.
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