Silurian Of Gotland Research Articles

Palaeozoic calcareous plankton: evidence from the Silurian of Gotland

In order to investigate whether or not equivalents to modern calcareous plankton existed in Palaeozoic times, extremely well-preserved successions have to be investigated. The Silurian strata on Gotland (Sweden) are exceptionally well preserved because they have not experienced deep burial conditions and tectonic stress, due to their position on the stable Baltic Shield. Scanning electron microscope investigations of polished, slightly etched rock surfaces revealed the presence of a variety of calcareous micro- and nannofossils. Among these organisms, many can be termed ‘calcispheres’ (60–100 µm in diameter), whereas others due to their size range (nannofossils) are informally termed herein as ‘nannospheres’ (10–25 µm in diameter). The systematic attribution of these fossils is unknown (incertae sedis). Mesozoic calcispheres are usually attributed to calcareous cysts of dinoflagellates following comparisons of ultrastructure with modern species. The abundance of different calcispheres in the Silurian sediments of Gotland and the observation that many of the calcareous microfossils occur in distinctly different facies as well as their spherical shape indicate that they probably belong to calcareous micro- and nannoplankton. We therefore conclude that calcareous plankton most probably existed already during the Palaeozoic, but can only be observed under conditions of exceptional preservation.

Paleozoic calcareous plankton: evidence from the Silurian of Gotland

Paleozoic calcareous plankton: evidence from the Silurian of Gotland

Gonad pouches in a new clam-like monoplacophoran from the Silurian of Sweden

Beyrichidiscus fecundatus n. gen. n. sp. is described from the Upper Silurian of Gotland, Sweden. The minute, isostrophically coiled mollusc exhibits profound allometry and is considered to have been a monoplacophoran on the basis of functional interpretation of apertural structures. A clam-like, semi-infaunal mode of life is suggested. A pair of prominent lateral swellings in the adult shell are interpreted as reflecting the location of the gonads.

Cementing strophomenide brachiopods from the Silurian of Gotland (Sweden): Morphology and life habits

Dorsal valves of the cementing strophomenide brachiopods Leptaenoidea silurica Hedström and Liljevallia gotlandica Hedström are described for the first time, and it is shown that both these species could also live ambitopically. The lower Wenlock Scamnomena rugata (Lindström) represents young individuals of the ambitopic variant of Leptaenoidea silurica, and is placed in synonymy, resulting in the valid name for the taxon being Leptaenoidea rugata. The range of this species now spans the whole Wenlock, from the upper Visby Formation to the Klinteberg Formation, and possibly even into the Ludlow. Ambitopic gerontic specimens of L. rugata develop very thick shells, in which the ventral valves have strong curvature, and become deeper not by geniculation but by successive mantle retractions and subsequent re-growth, in a way similar to that of atrypides. This shape was probably an adaptation to “floating” on softer substrates. The thickened gerontic dorsal valves have well-developed lophophoral support, showing the shape of the lophophore, which comprises two branches that curve inwards and then backwards; the lophophore was probably ptycholophous and similar to that in living members of the Thecideidina. Ambitopic specimens of Liljevallia could grow to a much larger size than cementing forms, where the dorsal valves have very large, posterior-facing cardinal process lobes and deeply impressed muscle fields and anterior scars. The presence of a ventral process and long, posteroventrally elongated cardinal process lobes, and the absence of dental plates reveals that Liljevallia was probably an early member of the Douvillinidae and is thus removed from the Leptaenoideidae.

Systematics, shell structure and affinities of the Palaeozoic Problematicum Cornulites

The genus Cornulites, with the type species C. serpularius Schlotheim, 1820, from the Silurian of Gotland, comprises annulated, conical or tubular calcite shells, often found attached to the hard parts of other organisms. No consensus has ever been reached over the zoological affinities of the taxon, and no examples of soft-part preservation are known: detailed examination of shell structures and growth patterns provide the only means of assessing its systematic position. Using transverse and longitudinal thin sections of C. serpularius Vine, 1882, and C. cellulosus sp. nov., from the Much Wenlock Limestone Formation of England, the shell structure of Cornulites is shown to be lamellar, but with conspicuous internal chambers (camerae) at the apical end of the shell and, particularly in C. cellulosus, numerous smaller vacuities (cellulae) between the lamellae in the apertural shell region. Growth of the shell was by the secretion of low-magnesian calcite increments within one another, giving a cone-in-cone structure, with the prominent development of cellulae in C. cellulosus probably a constructional feature relating to an upright life position. By comparison of morphology and shell structure with other taxa, the zoological affinities of Cornulites are re-examined; previously suggested affinities with annelids, foraminifers, molluscs and poriferans can be ruled out. Specific shell structures, most notably pseudopuncta similar to those of bryozoans and brachiopods, have led some recent workers to interpret cornulitids as lophophorates. However, it is shown that they can be interpreted alternatively as solitary, aseptate members of the stem-Zoantharia (Cnidaria: Anthozoa). Four cornulitid species are recognized in the Much Wenlock Limestone Formation: C. cellulosus sp. nov., C. gremialis sp. nov., C. scalariformis and C. serpularius. In the absence of the type material, C. serpularius is here restricted to cornulitids closely resembling the specimens originally figured by Schlotheim. © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society, 2007, 150, 681–699.

A latest Llandovery to latest Ludlow high-resolution biostratigraphy based on the Silurian of Gotland—a summary

A succession of 26 conodont zones and 63 successive subzones and conodont faunas have been identified in the exposed latest Landovery to latest Ludlow strata of Gotland, Sweden. All zones, and several of the subzones and faunas, have been identified elsewhere on Baltica, as well as on other palaeocontinents, indicating that this zonation serves well as the standard zonation for calcareous successions. The average duration of the 24 Wenlock and Ludlow zones was 396±208 ka, using the latest radiometric timescale. However, most zones and subzones formed during the widely recognised Silurian oceanic events correspond to less time per unit—in the order of 30 to 100 ka—and most of those formed during intervening episodes correspond to a few 100 ka. The average duration of the 61 Wenlock and Ludlow zones, subzones, and faunas was only 156±82 ka, i.e., among the highest biostratigraphical resolution available for Phanerozoic epochs. On Gotland the combined known maximum thickness of the strata has increased to more than 750 m.

The problematic fossil Jinonicella from the Lower Silurian of Gotland

The minute problematic fossil Jinonicella kolebabai Pokorný, 1978 is described from the Lower Silurian (Wenlock, middle Sheinwoodian) of the När-1 core from southern Gotland. Growth lines indicate that the characteristic tubular snorkel of Jinonicella grew by envelopment of an emargination in the median dorsal aperture and not as a perforation through the dorsal surface of the shell. The direction of shell coiling indicates that Jinonicella kolebabai can not be closely related to scaphopods (?Devonian, Carboniferous-Recent). If it is a mollusc, it may represent an independent adaptation to the infaunal scaphopod mode of life.

A new cornulitid genus from the Silurian of Gotland, Sweden

A new genus and species Septalites septatus gen. et sp.n. of a possible cornulitid is described from the Silurian of Gotland. The new genus resembles Conchicolites in having a smooth tube interior with multiple septa, but differs in the greater number of septa and microlamellar tube structure. Most of the shell interior (about 3/4) is filled with the septa. S. septatus shares several features with Cornulites and tentaculitids, but it differs in lacking pseudopuncta and an annulated tube interior. It is affiliatiated with the Class Tentaculita Bouãek (1964) because of micro-lamellar tube structure and the presence of septa.

Early Palaeozoic diversification of chitons (Polyplacophora, Mollusca) based on new data from the Silurian of Gotland, Sweden

Isolated, well-preserved silicified sclerites of an unusually diverse Silurian paleoloricate assemblage from Gotland preserve morphological features that are important in interpreting palaeobiology. The typically granular dorsal ornament is comparable with Recent chitons, and is hence possibly linked functionally with sensory aesthetes. Ventral structures particularly in thickened shells indicate major muscle attachment sites sub-apically or, equivalently, beneath the rim of the apical area, and also marginally. In early chitons such as Cambrian Matthevia, deep ventral cavities represent comparable sub-apical sites. Three Gotland genera with an unusual, holoperipheral shell growth style apparently represent plated aplacophorans (cf. Acaenoplax), which coexisted with paleoloricate chitons in shallow inshore carbonate shelf environments. Sclerite features of all the Gotland genera are discussed together since they share most characteristics. The new family Heloplacidae includes those genera and Acaenoplax, which in a preliminary cladistic analysis form a sister group to other Lower Palaeozoic paleoloricates. Multiplated skeletons in paleoloricates and this group of aplacophorans represent parallel evolution of dorsal armour, which in chitons resulted in overlapping, articulating sclerites. The diversity of the mid-Silurian Gotland assemblage is examined against early evolutionary diversification of polyplacophorans, aplacophorans, and in relation to the overall record of Palaeozoic paleoloricate and neoloricate chitons. Peaks in diversity in early Ordovician, mid-Silurian and early Carboniferous times correspond to periods with widespread development of low latitude carbonate shelves. Neoloricates, apparently with an additional shell layer that contributed articulatory plates, appeared in the Devonian from where the fossil record remains poorly known.

Specific representation of the Silurian cephalopod genus Phragmoceras in Gotland and Britain

Species of Phragmoceras in the Silurian of Gotland and Britain are reviewed and revised. One new species P. exaggeratus is described from the Ludlow Series of the Welsh Borderland. There are comments on the broader distribution of the genus and on its mode of life.

A new spiny cornulitid from the Silurian of Gotland, Sweden

Cornulites gotlandicus sp.n. is described from the Silurian (Ludlowian) Hemse Beds in Gotland. The new species differs from all other known species of the genus in having a longitudinal striation formed of rows of successive hollow spines.

LATE CAMBRIAN AND EARLY ORDOVICIAN STEM GROUP CHITONS (MOLLUSCA: POLYPLACOPHORA) FROM UTAH AND MISSOURI

Abundant silicified shell plates (valves) of some of the oldest-known chitons were recovered from the Late Cambrian Notch Peak Formation of Utah. The chiton fauna is dominated numerically by Matthevia wahwahensis new species, but also includes another mattheviid, Eukteanochiton milleri new genus and species, and a preacanthochitonid, Orthriochiton utahensis new genus and species. Robustum from the Early Ordovician Gasconade Formation of Missouri is herein reinterpreted as a septemchitonid chiton, and Hemithecella, from the Late Cambrian-Early Ordovician of the eastern United States, is once again considered a mattheviid chiton. Mattheviid valves are unique among chitons in that they are massive, elongate, and contain one or two tunnels; these characteristics have led some to exclude this family from the Polyplacophora. However, mattheviids and other chitons share many valve characters, including granules, an apical shelf, a thin anterior margin, bilateral symmetry, three valve types, and a shell layer perforated with canals. Furthermore, recently described chiton valves from the Silurian of Gotland, the Ordovician of Wisconsin, and the Cambrian-Ordovician of Missouri are gradational between the Notch Peak mattheviid valves and those of younger polyplacophorans. Also, Matthevia wahwahensis n. sp. has valves with a flat apical area, allowing for valve overlap, and so provides a link between the unusual chiton Matthevia variabilis, which had nonoverlapping, spiky valves, and Chelodes, a less disputed chiton. Known fossil assemblages reveal that, along the coastline of Laurentia during the Cambrian-Ordovician, chitons were diverse, consisting of at least four families and with much variation in valve shape.

The jawed annelid Rhytiprion magnus Kielan-Jaworowska, 1966 – a muddy bottom dweller

The Ordovician jawed polychaete annelid Rhytiprion magnus has hitherto been recorded in erratic boulders of probable East Baltic provenance and in drill cores from Poland and North Estonia. In this study the stratigraphical range and geographical distribution are extended as R. magnus is recorded from the Silurian of Gotland (Sweden) and Indiana (U.S.A.). Hence, R. magnus is now known from the Middle Ordovician to the middle Silurian (lower Ludlow; Kockelella v. variabilis conodont Zone) and comprises one of the few Palaeozoic polychaete species with a documented intercontinental distribution. The species is generally very rare but is abundant at a few localities; on Gotland, in the offshore setting of the Late Wenlock (K. ortus absidata conodont Zone; Colonograptus? praedubeli graptolite Zone) Djupvik Member of the Halla Formation. The distribution pattern shows that R. magnus thrived in low energy, distal platform environments, in which marly and muddy terrigenous sediments accumulated.

The mineralogical composition of precursor sediments of calcareous rhythmites: a new approach

Previous studies in Silurian carbonates from Gotland (Sweden) have led to a model for the development of limestone–marl alternations. This model postulates that early diagenesis of precursor sediments without strong primary differences can result in a differentiation by selective dissolution of aragonite in marl beds and reprecipitation of calcite cement in limestone beds. This model is described as a set of mathematical equations that quantify the diagenetic processes (aragonite dissolution and calcite reprecipitation) that occur during the formation of limestone–marl interbeds from a hypothetical homogeneous precursor sediment. The calculations demonstrate that resulting hypothetical limestone–marl alternations show characteristic mathematical relationships between the ratios of the bed thicknesses of limestones and marls on one side, and the carbonate contents, on the other. By reversing this model, the original mineralogical composition of the precursor sediment of real-world rhythmic successions can be determined. In this study, alternations from the Silurian of Gotland, the Cambrian, Devonian, and Mississippian of North America, the Jurassic of France and Germany, and the Cretaceous of France are shown to exhibit mathematical relationships similar to those calculated for hypothetical precursor sediments without primary differences. Therefore, the mineralogical composition of their precursor sediments can be estimated. In contrast, the clear mismatch shown by the Lower Jurassic Belemnite Marls from Dorset indicates that these rhythms did not suffer an early diagenetic overprint. Our model helps to differentiate between rhythmites with strong depositional variations and those without; however, it cannot indicate whether a given alternation is the product of rhythmic diagenesis of a homogeneous precursor sediment or the result of diagenetic enhancement of subtle underlying sedimentary rhythms. For horizontally correlated patterns, such as laterally extensive beds and layers of nodules, an a priori unknown external signal has to be assumed.

Silurian Ramphoprionid polychaetes from Gotland, Sweden

Silurian ramphoprionid polychaete annelids, represented by their jaws (scolecodonts), are described from extensive collections from Gotland, Sweden. The family Ramphoprionidae, monotypic at its original description, is sub-divided into four genera;ProtarabellitesStauffer, 1933;RamphoprionKielan-Jaworowska, 1962; “Pararamphoprion” Männil and Zaslavskaya, 1985; andMegaramphoprionnew genus. Identified species include “P.” cf.nordicusMännil and Zaslavskaya, 1985;P. rectangularisnew species;P. staufferinew species;P. triangularisnew species; and twoProtarabellitesspecies left in open nomenclature.Ramphoprionis represented by one new highly plastic species,R. gotlandensis, housing five distinguishable morphotypes showing gradual evolution.Megaramphoprion, which is most closely related toRamphoprion, is represented byM. magnusnew genus and species, a rare but distinctive taxon. Most species have long stratigraphic ranges within which important morphological changes can nonetheless be observed. The stratigraphic range of ramphoprionids includes, at least, the Ordovician to the Silurian. They are fairly rare in the Silurian of Gotland and where present they generally form less than 10 percent of the polychaete faunas, although occasionally reaching as much as 20 to 30 percent. Evolution, paleoecology, and surface structures of the investigated species are briefly discussed.

Shell succession, assemblage and species dependent effects on the C/O-isotopic composition of brachiopods — examples from the Silurian of Gotland

Brachiopods are the most widely used biological constituents for the determination of stable C- and O-isotopic compositions in Paleozoic sediments. In the present study, critical precautions for obtaining reliable isotope values and maximum possible reliability are discussed. Samples were taken in the tectonically undisturbed and exceptionally well-preserved Silurian sequence on Gotland (Sweden). The state of preservation of brachiopod shells was examined by cathodoluminescence (CL) microscopy and by comparison of shell ultrastructures with those of recent brachiopods. Nine types of ultrastructure are distinguished and attributed either to biological differences or to diagenetic alterations. For isotope analyses, only largely unaltered areas of the secondary, fibrous shell layer should be used. Shell layers with other types of ultrastructure (primary, tertiary, laminar) may have isotopic compositions different from the assumed seawater value even when diagenetically unaltered. Only brachiopod taxa with a shell succession that includes a sufficiently thick secondary shell layer (e.g., Spiriferida, Rhynchonellida, Orthida) give reliable, mutually corresponding isotope values, indicating that the shell calcite precipitated at or near equilibrium with ambient seawater. Brachiopods with other kinds of shell successions (Strophomenida, Pentamerida) as well as other calcite-shelled organisms (trilobites, ostracodes) give non-equilibrium isotopic results and therefore should not be used. By restricting the samples to one species or to a group of closely related species, even minor differences in primary isotopic composition can be excluded. In the Silurian of Gotland, the stratigraphically widely distributed spiriferid Atrypa reticularis was selected for study. Considering the type of record, four different taphonomic assemblages are defined. In autochthonous assemblages of specimens, which lived contemporarily, original values of carbon as well as oxygen isotopes can be evaluated with a reliability of <±0.4‰. This allows the recognition of minor, hydrographically produced differences between neighbouring, contemporary facies areas and the determination of small (<1‰) isotopic excursions. Thus, besides being indicators of hydrographic conditions in ancient oceans, stable carbon- and oxygen-isotope compositions can be used as a stratigraphic tool that provides correlations, sometimes even within biozones.

Halysis Høeg, 1932 - a problematic Cyanophyceae: new evidence from the Silurian of Gotland (Sweden)

Halysis Høeg, 1932 - a problematic Cyanophyceae: new evidence from the Silurian of Gotland (Sweden)

A New Family Of Calcareous Microfossils From The Silurian Of Gotland, Sweden

Scanning electron microscopy of polished, slightly etched rock surfaces of samples from the Silurian of Gotland, Sweden, revealed the presence of excellently preserved calcareous microfaunas and microfloras. This paper illustrates several morphotypes of calcareous micro-organisms of unknown biological affinity, attributed to the new family Ovummuridae. Specimens of Ovummurus duoportius Minoura and Chitoku, a problematical microfossil, which was first described from the Upper Pennsylvanian of Kansas, USA, were found in addition to comparable micro-organisms attributed to the following new taxa: Minourella gotlandica gen. et sp. nov., Arouxina pluricellata gen. et sp. nov. and Samtlebenella circumcamerata gen. et sp. nov. In addition, a few other problematical micro-organisms, probably belonging to the same family, are described under open nomenclature. The enigmatic microfossils are compared with other known calcareous organisms, their possible biological affinities are discussed, and the different stages of diagenetic alteration and destruction are illustrated.

Significance of a Halysitid-Heliolitid Mud-facies Autobiostrome from the Middle Silurian of Gotland, Sweden

Abstract A low-diversity halysitid-heliolitid autobiostrome from the Late Wenlock Halla Formation at Blahall 1, Gotland, indicates a brief period of particular environmental conditions. The reef is 0.3–0.5 m thick, internally without bedding structures, and had a sheet-like distribution on a low-relief soft-bottom, in marginal platform, graptolitiferous strata (height-width ratio is 1:900 in outcrop). The dominating coral species building this mud-facies biostrome were the tabulates Stelliporella cf. parvistella and Halysites laticatenatus. The rugose coral fauna is dominated by fixosessile and rhizosessile forms with well developed holdfasts, for example Dokophyllum elegantulum. Coral growth forms generally indicate high background sedimentation rates during the growth of the biostrome. Four successive stages of sea-floor development are discussed: (1) a pre-biostrome stage, (2) a pioneering community stage, (3) a climax-community stage, and (4) a post-biostrome stage. Stratigraphically, these stages sho...

Significance of a Halysitid-Heliolitid Mud-Facies Autobiostrome from the Middle Silurian of Gotland, Sweden

Significance of a Halysitid-Heliolitid Mud-Facies Autobiostrome from the Middle Silurian of Gotland, Sweden