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The new retiolitid species, Spinograptus tubothecalis, is described from the Colonograptus praedeubeli and C. deubeli biozones from two localities in Poland: a borehole on the East European Platform and the Holy Cross Mountains. This was a recovery phase after the severe Silurian Cyrtograptus lundgreni Event. The new species has a unique, previously undescribed form of finite rhabdosome. Unlike the species Spinograptus reticulolawsoni and S. lawsoni, in which the fi− nite rhabdosomes taper distally, its rhabdosome is parallel−sided with the two distal thecae developed as isolated tubes without genicular processes, with a small appendix between them. The new species also has preserved membranes of the sicula, thecae and ancora sleeve, similar to a few species of Spinograptus from the lower Homerian. Spinograptus tubothecalis, like Spinograptus clathrospinosus and S. spinosus, has paired reticulofusellar genicular processes on the pre−thecal ventral orifices, similar to but shorter than thecal processes. Transverse rods, a rare character in post−Cyrto− graptus lundgreni Event retiolitids occur in the new species in rudimentary form.
A minute Silurian oncocerid Cyrtoceras pollux, from the Prague Basin is assigned here to the genus Pomerantsoceras. The only so far known species of this genus comes from the Upper Ordovician (Hirnantian) of Estonia. Pomerantsoceras thus represents, except for un−revised poorly understood taxa, the single known oncocerid genus surviving the end−Ordovician extinction events. Cyrtoceras pollux is unusual among the Silurian nautiloids because of its small shell. Colour pattern characterised by a few longitudinal bands on the entire circumference of the shell is here reported in oncocerids. Longicone and only slightly curved small shells as in Pomerantsoceras are unusual among nautiloids and resemble straight shells of orthocerids and pseudorthocerids, in which the colour pattern consists of straight colour bands. Consequently the shell shape as well as the colour pattern should be regarded as adaptive convergence with orthocerids and pseudorthocerids. It supports the hypothesis that colour pattern functioned as camouflage and its evolution was under adaptive control. In addition, several types of the shell malformations including anomalous growth of septa, shell wall and pits on an internal mould are described.
In the classic section across the Silurian–Devonian boundary at Dnistrove (Podolia, Ukraine) the brachiopod fauna has never been studied in detail. This paper presents results of research on brachiopods from this important locality and time interval. Bed−by−bed collecting has enabled the detailed distribution of brachiopod taxa through the boundary beds to be revealed. Generally, the reference section at Dnistrove yields rather scarce but often well preserved brachiopods. Dayia bohemica and Dnestrina gutta can be regarded as characteristic species for the uppermost Silurian. A relatively high−diversity but low−abundance brachiopod fauna occurs in the lowest 1.8 m of the earliest Devonian. Only three forms have been found to cross the Silurian–Devonian boundary: the strophomenide Plectodonta (Plectodonta) mariae pantherae subsp. nov., the atrypide Gracianella (Sublepida) paulula sp. nov., and the spiriferide Howellella (Howellella) latisinuata. A relatively narrow brachiopod−rich interval at 5.5 m above the Silurian–Devonian boundary yields 16 brachiopod species which probably indicate a setting near the lower limit of the photic zone equivalent to the Benthic Assemblage 3–4 boundary. Two new species and one new subspecies are described: Skenidioides tatyanae, Plectodonta (Plectodonta) mariae pantherae, and Gracianella (Sublepida) paulula.
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Evolution of retiolitid graptolites - a synopsis

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Twenty million years of retiolitid evolution reflect environmental changes, the most severe being the Silurian Cyrtograptus lundgreni Event. Five biostratigraphically and morphologically constrained retiolitid faunas are distinguished and characterized according to their rhabdosomal modifications: (1) the oldest and long−ranging Llandovery group of mostly large and morphologically complex rhabdosomes, (2) the less diverse Telychian−Sheinwoodian group, (3) the Cyrtograptus lundgreni Biozone varied group of intermediate size, and two short−ranged (4) late Homerian, and (5) early Ludlow groups with small rhabdosomes. Although the evolutionary history of retiolitids was complex and not linear, a common tendency toward reduction of rhabdosome size in most lineages is observed. The greatest reduction in both number and volume of thecae, and in skeletal elements is demonstrated in the Gothograptus and Plectograptusfaunas. Contrary to the thecal decrease, a distinctive increase of sicula size is observed in retiolitids. Two types of colonies are distinguished: L−colonies with a small sicula and numerous large thecae of similar size, and S−colonies with a long sicula and a few, small thecae. These changes imply modification of the soft body: an increase in siculozooid length and a decrease in the size of the zooids. Thus, the siculozooid probably produced great amounts of morphogen inhibiting zooid growth. In consequence the phenomenon of colony reduction occurred. The most extreme stages of rhabdosome reduction in Ludlow retiolitids can be seen in Plectodinemagraptus gracilis of the Plectograptuslineage and in the new species Holoretiolites helenaewitoldi, possibly representing the last stage of skeletal reduction in the Gothograptus lineage; the next hypothetical stage would be its total loss. The sicula length of Holoretiolites, about 2 mm, is reported herein for the first time.
Epeiric seas covered the east and west parts of the old craton of Baltica in the Silurian and brachiopods formed a major part of the benthic macrofauna throughout Silurian times (Llandovery to Pridoli). The orders Strophomenida and Orthotetida are conspicuous components of the brachiopod fauna, and thus the genera and species of the superfamilies Plectambonitoidea, Strophomenoidea, and Chilidiopsoidea, which occur in the Silurian of Baltica are reviewed and reidentified in turn, and their individual distributions are assessed within the numerous boreholes of the East Baltic, particularly Lithuania, and attributed to benthic assemblages. The commonest plectambonitoids are Eoplectodonta (Eoplectodonta) (6 species), Leangella (2 species), and Jonesea (2 species); rarer forms include Aegiria and Eoplectodonta (Ygerodiscus), for which the new species E. (Y.) bella is erected from the Lithuanian Wenlock. Eight strophomenoid families occur; the rare Leptaenoideidae only in Gotland (Leptaenoidea, Liljevallia). Strophomenidae are represented by Katastrophomena (4 species), and Pentlandina (2 species); Bellimurina (Cyphomenoidea) is only from Oslo and Gotland. Rafinesquinidae include widespread Leptaena (at least 11 species) and Lepidoleptaena (2 species) with Scamnomena and Crassitestella known only from Gotland and Oslo. In the Amphistrophiidae Amphistrophia is widespread, and Eoamphistrophia, Eocymostrophia, and Mesodouvillina are rare. In the Leptostrophiidae Mesoleptostrophia, Brachyprion, and Protomegastrophia are common, but Eomegastrophia, Eostropheodonta, Erinostrophia, and Palaeoleptostrophia are only recorded from the west in the Baltica Silurian. In the Eopholidostrophidae, Mesopholidostrophia is common and Eopholidostrophia rare, and within the Shaleriidae the genus Shaleria is revised and found to be the only genus within the family Shaleriidae and to contain three subgenera, S. (Shaleria), S. (Shaleriella) and S. (Janiomya). In the Strophonellidae Strophonella was widespread and Eostrophonella rare. Within the Orthotetoidea, Coolinia and Morinorhynchus(with the new Ludlow and Pridoli species M. rubeli from the Lithuanian boreholes erected) are common, and Fardenia, Saughina, and Valdaria again only known from the west Baltic. Most of the genera and many of the speciesreviewed were very widespread and many found in the adjacent Avalonia and Laurentia: a few were even more cosmopolitan.
Analysed were forest soils developed from Silurian, Devonian, Triassic and Jurassic sandstones. The analysis of the results reveals a significant effect of the type of sandstone on the depth of studied soils and their properties. Tested soils were characterized by a strongly acid reaction and a low degree of saturation with basic cations (between 10 and 35%). In the top horizons were found increased concentrations of zinc, copper and lead in relation to the parent rock concentrations.
Agastograptus robustus Obut and Zaslavskaya, 1983, the type species of the genus Agastograptusis herein recognized as a species of Plectograptus. Large proximal lateral orifices are one of the significant characters of Plectograptus. P. robustus differs from the type species, Plectograptus macilentus Törnquist, 1887, in possessing paired apertural processes. Other species of Agastograptus have been assigned to three different genera: Spinograptus, S. clathrospinosus (Eisenack, 1951), S. munchi (Eisenack, 1951), Neogothograptus, N. balticus (Eisenack, 1951), and Cometograptus, C. nevadensis (Berry and Murphy, 1975). The main diagnostic feature used for Agastograptus, the spinoreticular paired apertural processes, is recognized as a species feature, characteristic mostly for Spinograptus, whereas the generic features are the arrangement of the proximal end, ventral walls, and ancora sleeve of the rhabdosome. Therefore the genus Agastograptus is a synonym of Plectograptus.
Silurian cephalopod limestones known from northern Asia are usually singular beds or horizons of nodules. They range in their taphonomic character from the Mojero river type, with postmortem concentration from rich living populations by relatively gentle bottom currents into a submarine low, to that of the Karabutak Formation of the South Urals, where there has been concentration by strong wave or current action.
Six well exposed mid−Ludlow stromatoporoid−dominated reef biostromes in four localities from the Hemse Group in southeastern Gotland, Sweden comprise a stromatoporoid assemblage dominated by four species; Clathrodictyon mohicanum, “Stromatopora” bekkeri, Plectostroma scaniense, and Lophiostroma schmidtii. All biostromes investigated in this area (of approximately 30 km²) are interpreted to belong to a single faunal assemblage forming a dense accumulation of fossils that is probably the best exposed stromatoporoid−rich deposit of the Silurian. The results from this comprehensive study strengthen earlier interpretations of a combination of genetic and environmental control on growth−forms of the stromatoporoids. Growth styles are similar for stromatoporoids in all six biostromes. Differences in biostrome fabric are due to variations in the degree of disturbance by storms. The uniformity of facies and the widespread low−diversity fauna support the view that palaeoenvironmental conditions were similar across the area where these biostromes crop out, and promoted the extraordinary growth of stromatoporoids in this shallow shelf area.
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Thelodont scales are described from the Silurian Niur Formation in the Derenjal Mountains, east central Iran. The material studied herein comes from four stratigraphic levels, composed of rocks formed in a shallow water, carbonate ramp environment. The fauna includes a new phlebolepidiform, Niurolepis susanae gen. et sp. nov. of late Wenlock/?early Ludlow age and a late Ludlow loganelliiform, Loganellia sp. cf. L. grossi, which constitute the first record of these thelodont groups from Gondwana. The phlebolepidiform Niurolepis susanae gen. et sp. nov. is diagnosed by having trident trunk scales with a raised medial crown area separated by two narrow spiny wings from the lateral crown areas; a katoporodidtype histological structure distinguished by a network of branched wide dentine canals. Other scales with a notch on a smooth rhomboidal crown and postero−laterally down−stepped lateral rims have many characters in common with Loganellia grossi. Associated with the thelodonts are indeterminable acanthodian scales and a possible dentigerous jaw bone fragment. This finding also provides evidence of a hitherto unknown southward dispersal of Loganellia to the shelves of peri−Gondwana.
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Crinoids from the Silurian of Western Estonia

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The Silurian crinoids of Estonia are re−evaluated based on new collections and museum holdings. Nineteen species−level crinoid taxa are now recognized. All crinoid names applied to Estonian Silurian crinoids during the middle 19th century are disregarded. Especially significant is the fauna reported herein from the Pridoli because coeval crinoids are very poorly known from the Baltic region and elsewhere. One new genus and four new species are described from Estonia, namely Calceocrinus balticensis sp. nov., Desmidocrinus laevigatus sp. nov., Eucalyptocrinites tumidus sp. nov., and Saaremaacrinus estoniensis gen. et sp. nov.
A new myodocope ostracod Sineruga insolita gen. et sp. nov. is herein described from the Armorican Massif (France). Sineruga resembles Silurian myodocopes (i.e., bolbozoids and cypridinids) in having anterior features (thinner carapace) possibly related to vision. On the other hand, it resembles entomozoid ostracods in having a bean−shaped outline along with a deep adductorial sulcus and a simple muscle spot, but lacks their characteristic ribbed ornament. The data available suggest that Sineruga insolita was probably an atypical, non−ribbed member of the entomozoids thus indicating that early entomozoids may have had smooth representatives. Comparisons with other Recent and fossils ostracods show that the presence of a rostrum and/or symmetrical vision related carapace features (i.e., indicating lateral eyes) can be used as diagnostic characters for myodocope ostracods. The position and shape of the dorsal connection and to a lesser extent that of the sulcus and the adductorial muscle scar can be used to discriminate the higher groups of Silurian myodocopes (bolbozoids, cypridinids, and entomozoids).
Two global isotopic events, the early Sheinwoodian (early Wenlock) and that at the Silurian–Devonian transition, have been comprehensively studied in representative carbonate successions at Kytayhorod and Dnistrove, respectively, in Podolia, Ukraine, to compare geochemistry and biotic changes related correspondingly to the Ireviken and Klonk events. These two large−scale isotope excursions reveal different regional ecosystem tendencies. The well−defined increasing trend across the Llandovery–Wenlock boundary in siliciclastic input, redox states and, supposedly, bioproductivity, was without strict correlative relations to the major ¹³C enrichment event. The environmental and biotic evolution was forced by eustatic sea−level fluctuations and two−step climate change toward a glaciation episode, but strongly modified by regional epeirogeny movements due to location near the mobile Teisseyre−Törnquist Fault Zone. Thus, the global early Sheinwoodian biogeochemical perturbation was of minor depositional significance in this epeiric sea, as in many other Laurussian domains. Conversely, the Podolian sedimentary record of the Klonk Event exhibits temporal links to the abrupt δ¹³C anomaly, overprinted by a tectonically driven deepening pulse in the crucial S–D boundary interval. This carbon cycling turnover was reflected in the regional carbonate crisis and cooling episodes, paired with a tendency towards eutrophication and recurrent oxygen deficiency, but also with major storms and possible upwelling. Faunal responses in both Podolian sections follow some characters of the Silurian pattern worldwide, as manifested by conodont changeover prior to the major early Sheinwoodian isotopic/climatic anomaly. This contrasts with the relative brachiopod and chitinozoan resistances in the course of the Ireviken Event. Also, during the Klonk Event, a moderate faunal turnover, both in benthic and pelagic groups, occurred only near the very beginning of the prolonged ¹³C−enriched timespan across the system boundary, possibly due to progressive dysoxia and temperature drop. The characters point to a peculiarity of the Klonk Event by comparison with the Silurian global events, and some similarity already to the succeeding Devonian transgressive/anoxic episodes.
The Ludlow genus Plectograptus, with the type species Retiolites macilentus Törnquist, 1887, collected from Thuringia (Germany), has been a widely−identified, monospecific, but poorly understood taxon for almost one hundred years. This was due to poor and incomplete preservation of the type material, and misidentification by subsequent authors up to 1995. The original, and only, type specimen of P. macilentus collected by Törnquist being lost, a neotype is herein selected from a small collection of Thuringian material. The genus has now been redefined and based on this, and SEM studies of isolated material, the defining characteristics of the genus are (i) the possession of a simple ancora umbrella with five radial lists with an incompletely developed rim; (ii) an ancora umbrella separated from lateral ancora sleeve walls by exceptionally large lateral orifices; (iii) the possession of mid−ventral lists; (iv) simple, orderly zigzag lateral wall ancora sleeve lists. Recently, two additional species, P. robustus and P. wimani, previously placed in different genera, were assigned to Plectograptus. This study recognizes three new species: P. mobergi, P. toernquisti, and P. trijunctus, bringing the total number of species to six. Species are distinguished by the presence or absence of genicular processes, inclination of the thecal ventral walls and mid−ventral lists, presence or absence of reticular lists, and three−way or four−way sleeve/lateral rod/apertural lip junctions.
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The synziphosurine horseshoe crab Pasternakevia was until now known only from a single specimen. Herein we describe three new specimens from the outcrop close to the type locality of monotypic P. podolica that show several features that were unknown or only assumed in this genus. These characters include: a smooth surface of the carapace; an opisthosoma composed of ten segments, the first one being a microtergite (strongly reduced first opisthosomal segment usually hidden under the carapace), two or three last segments fused into a metasoma, and the telson. Pasternakevia is included in Bunodidae due to a hypertrophied tergite 2, but it is linked to the Pseudoniscidae by the shape of the opisthosoma.
Three isolated and well−preserved retiolitid taxa, two of them new, and one taxon of unknown affinity, are described from the upper Llandovery of Arctic Canada. All taxa display unusual characteristics The three retiolitids display extraordinary morphological features: Pseudoplegmatograptus cf. obesus preserves very delicate and lacey or cobweb−like list structures external to the main skeletal lists, structures previously seen only rarely on flattened material; Pileograptus pileatus gen. et sp. nov. possesses a thecal framework typical of retiolitines, particularly Stomatograptus, but its thecal orifices possess broad, meshwork genicular hoods similar to those in the plectograptines; and Giganteograptus giganteus, formerly attributed to Pseudoplegmatograptus, with its very coarse and relatively simple skeletal framework, well−developed paired thecal apertural spines, and a pustulose micro−ornamentation on the lists. The latter feature suggests an assignment to the subfamily Plectograptinae, rather than to the Retiolitinae as has been previously presumed. The presence of pustulose lists in Giganteograptus adds a further complication to understanding of retiolitid evolution. Mirorgraptus arcticus gen. et sp. nov., the fourth taxon and of unknown systematic affinity, lacks the proximal end, but preserves lists with seams and some fusellar material suggestive of some type of new retiolitid. Unlike known retiolitids or other graptolites, however, the skeletal list development is apparently confined to a narrow region along one side of the nema.
Mass occurrence of mats comprised of benthic coccoid cyanobacteria is reported from early Silurian black radiolarian cherts exposed at Żdanow village (Bardzkie Mountains, Sudetes, southwestern Poland). The cherts contain laminated organic matter representing degraded benthic coccoid cyanobacterial mats. The remains of cyanobacteria occur as laminated agglomerations of variously preserved subglobular colonies composed of spherical cells of variable size and numbers. The morphology of remnants of cells and their mucilaginous envelopes, structure of colonies, and particularly the presence of small granular structures resembling reproductive cells known in extant coccoid cyanobacteria as baeocytes, permit to compare the Silurian microbiota with modern cyanobacteria assigned to the genera Stanieria or Chroococcidiopsis.
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