Sr/Ca and Ba/Ca ratios support trophic partitioning within a Silurian conodont community from Gotland, Sweden

Conodonts were a highly diverse and abundant vertebrate group whose fossils are found in marine Paleozoic and Triassic strata around the world. They inhabited environments ranging from lagoons to open oceans and are represented by a wide variety of dental morphologies. Conodonts may have filled many different ecological niches and represent a significant proportion of nekton before the Devonian. Despite this, very little is known about trophic ecology of conodonts. While morphological diversity suggests a complex trophic structure within conodont communities, there is little evidence to support dietary niche partitioning among conodonts. We tested the hypothesis that individual conodont taxa occupied different trophic niches, using Sr/Ca and Ba/Ca ratios preserved in the dental elements of assemblages from Silurian strata of Gotland, Sweden. Sr/Ca and Ba/Ca have been shown to vary in vertebrate skeletal tissues depending on trophic positioning, although biological and environmental conditions can affect these ratios. Environmental influences were minimized by examining entire conodont communities from a tropical epeiric sea and by measuring strontium isotope ratios using thermal ionization mass spectrometry in the most metropolitan taxon (Ozarkodina confluens). Composition of white matter, a tissue unique to conodonts, was also analyzed using microprobe analysis, revealing significantly lower Sr concentrations than in surrounding lamellar tissue, suggesting taxon-specific histology should be considered when analyzing conodonts for geochemical data. Excluding taxa with highly variable quantities of white matter, the results show that each taxon preserves different Sr/Ca and Ba/Ca ratios with limited overlap, indicating variation in trophic positioning

Silurian vertebrates of Gotland (Sweden) and the Baltic Basin

During the Silurian, the Swedish island Gotland was positioned close to the equator and covered by a shallow sea called the Baltic Basin. The sedimentary rocks (predominantly carbonates) comprising most of the island today were initially formed in this warm sea, and the relatively complete succession of rocks often contains fossil fragments and scales from early vertebrates, including heterostracans, anaspids, thelodonts, osteostracans, acanthodians, and a stem-osteichthyan. Fossils of early vertebrates become increasingly more common in younger Silurian rocks, but are mostly represented by fragmentary remains and rarer occurrences of articulated jawless vertebrates (agnathans). However, the record of articulated specimens and jawed vertebrates (gnathostomes) are more numerous in rocks of the following Devonian Period. Isolated peaks of agnathan diversity during the Silurian and disarticulated remains of gnathostomes from this period hint at a cryptic evolutionary history. A micropaleontological approach with broader sampling may provide a better understanding of early vertebrate distribution patterns and hopefully give some insights into this history. The objective of this study was to build upon previous sampling on Gotland and to use established frameworks for disarticulated remains with the aim of making comparisons with similar studies performed in the East Baltic. However, difficulties locating the collections from these previous works necessitated a different focus. Undescribed museum collections and newly sampled material enabled some taxonomical revisions and greatly improved the understanding of vertebrate distribution in the youngest part of the Gotland sequence. It also indicated that this interval may represent the early stages of the diversification of gnathostomes that become increasingly dominant toward the end of the Silurian. Furthermore, the description of samples from partly coeval sections in Poland enabled some preliminary comparisons outside of Gotland, and presented a striking example of restricted environmental occurrences for a thelodont taxon. This is encouraging for future sampling and investigations on Gotland. Together with the establishment of a facies-framework comparable to that developed in the East Baltic and correlations to other areas, this may prove fruitful for an increased understanding of early vertebrate distribution and evolution during the Silurian

Vertebrate remains and conodonts in the upper Silurian Hamra and Sundre formations of Gotland, Sweden

A long history of geological research on the island of Gotland, Sweden, has resulted in a detailed biostratigraphy based on conodonts for the Gotland sedimentary succession, but the relation between the Hamra and Sundre formations, the youngest strata on southern Gotland, has remained poorly resolved. These formations have also remained relatively poorly described in terms of vertebrates compared to other parts of the succession. A survey of museum collections and newly sampled material reveal that the taxonomical compositions and richness of vertebrate faunas remain similar compared to the underlying Burgsvik Sandstone and Oolite members. However, the relative abundance of the respective groups changes: Paralogania ludlowiensis and rare osteostracan remains of Tahulaspis sp. only occur in samples from the lower Hamra Formation, while Thelodus sculptilis becomes more common in samples from Sundre Formation. Conodont and isotope data give support to previous suggestions that the Hamra and Sundre formations may be largely isochronous, and it is possible that the observed differences in vertebrate faunas reflect changes in depositional setting. This interval on Gotland has been suggested to represent a hiatus in the East Baltic sections, where younger strata show an increased importance of acanthodians in the vertebrate faunas. Gotland could therefore give insights into the early stages of this diversification of gnathostomes during late Silurian times. However, this has to be done in combination with data from other areas, as well as with a review and revision of the scale-based taxonomy of Silurian acanthodians from the Baltic Basin.Title in Thesis List of papers: Vertebrate dermal remains and conodont distribution in the upper Silurian Hamra and Sundre formations of Gotland, Sweden</p

Silurian vertebrate remains from the Oslo Region, Norway, and their implications for regional biostratigraphy

Several vertebrate assemblages are described from the Silurian of the Oslo Region, Norway, based on the review and revision of previous reports of microremains, as well as unpublished material from museum collections. Articulated thelodont specimens from the Rudstangen Fauna, Ringerike Group, are also described here for the first time, revealing a seemingly monogeneric loganelliid assemblage. The oldest assemblage (mid-Llandovery) only contains the thelodont Loganellia cf. aldridgei, while a single sample from upper Llandovery strata produced four Thelodus sp. scales. These scales share features with those from younger Thelodus taxa and give additional support to an early appearance of this genus. The mid-Wenlock faunas consist of thelodonts Loganellia grossi, Loganellia einari and Thelodus laevis. These are joined by the thelodont Paralogania martinssoni, anaspids Rhyncholepis parvula and cf. Pterygolepis nitida, as well as the osteostracans cf. Tyriaspis whitei and Osteostraci gen. et sp. indet. in late Wenlock and earliest Ludlow faunas. These complement the previously described anaspids and osteostracans of the Rudstangen Fauna based on articulated specimens. The faunas of a number of calcarenite samples collectively contain the thelodonts L. grossi, L. einari, P. martinssoni and T. laevis, and are most likely of early Ludlow age. When the vertebrate-producing samples are put in the proposed stratigraphical framework for the Oslo Region that has been refined in recent decades, the faunas fit well into the vertebrate biozonation established for the Silurian, contrary to previous claims. The earliest faunas in the area show similarities to the Llandovery of Britain, whereas thelodonts and anaspids, but not osteostracans, from late Wenlock and early Ludlow are more closely related to Baltic forms. Additional sampling of the area may assist in refining the biostratigraphy and provide insights into Silurian vertebrate distributions in Norway and related regions

The emergence of a complex pore-canal system in the dermal skeleton of Tremataspis (Osteostraci)

Thyestiids are a group of osteostracans (sister-group to jawed vertebrates) ranging in time from the early Silurian to Middle Devonian. Tremataspis is unique among thyestiids in having a continuous mesodentine and enameloid cover on its dermal elements, and an embedded pore-canal system divided into lower and upper parts by a perforated septum. The origin of this upper mesh canal system and its potential homology to similar canal systems of other osteostracans has remained a matter of debate. To investigate this, we use synchrotron radiation microtomography data of four species of Tremataspis and three other thyestiid genera. Procephalaspis oeselensis lacks an upper mesh canal system entirely, but Aestiaspis viitaensis has partially enclosed upper canals formed between slightly modified tubercles that generally only cover separate pore fields. Further modification of tubercles in Dartmuthia gemmifera forms a more extensive, semi-enclosed upper mesh canal system that overlies an extensive perforated septum, similar to that found in Tremataspis. Lower mesh canals in P. oeselensis are radially arranged and buried tubercles indicate a continuous growth and addition of dermal hard tissues. These features are lacking to varying degrees in the other investigated thyestiids, and Tremataspis probably had a determinate growth accompanied by a single mineralization phase of its dermal hard tissues. The previously proposed homology between the semi-enclosed upper canal system in Dartmuthia to the pore-canal system in Tremataspis is supported in this study, but the suggested homologies between these canals and other parts of the thyestiid vasculature to those in non-thyestiid osteostracans remain unclear. This study shows that three-dimensional modeling of high-resolution data can provide histological and structural details that can help clarify homology issues and elucidate the evolution of dermal hard tissues in osteostracans. In extension, this can give insights into how these tissues relate to those found among jawed vertebrates

Sr/Ca and Ba/Ca ratios support trophic partitioning within a Silurian conodont community from Gotland, Sweden

Conodonts were a highly diverse and abundant vertebrate group whose fossils are found in marine Paleozoic and Triassic strata around the world. They inhabited environments ranging from lagoons to open oceans and are represented by a wide variety of dental morphologies. Conodonts may have filled many different ecological niches and represent a significant proportion of nekton before the Devonian. Despite this, very little is known about trophic ecology of conodonts. While morphological diversity suggests a complex trophic structure within conodont communities, there is little evidence to support dietary niche partitioning among conodonts. We tested the hypothesis that individual conodont taxa occupied different trophic niches, using Sr/Ca and Ba/Ca ratios preserved in the dental elements of assemblages from Silurian strata of Gotland, Sweden. Sr/Ca and Ba/Ca have been shown to vary in vertebrate skeletal tissues depending on trophic positioning, although biological and environmental conditions can affect these ratios. Environmental influences were minimized by examining entire conodont communities from a tropical epeiric sea and by measuring strontium isotope ratios using thermal ionization mass spectrometry in the most metropolitan taxon (Ozarkodina confluens). Composition of white matter, a tissue unique to conodonts, was also analyzed using microprobe analysis, revealing significantly lower Sr concentrations than in surrounding lamellar tissue, suggesting taxon-specific histology should be considered when analyzing conodonts for geochemical data. Excluding taxa with highly variable quantities of white matter, the results show that each taxon preserves different Sr/Ca and Ba/Ca ratios with limited overlap, indicating variation in trophic positioning

Sr/Ca and Ba/Ca ratios support trophic partitioning within a Silurian conodont community from Gotland, Sweden

Conodonts were a highly diverse and abundant vertebrate group whose fossils are found in marine Paleozoic and Triassic strata around the world. They inhabited environments ranging from lagoons to open oceans and are represented by a wide variety of dental morphologies. Conodonts may have filled many different ecological niches and represent a significant proportion of nekton before the Devonian. Despite this, very little is known about trophic ecology of conodonts. While morphological diversity suggests a complex trophic structure within conodont communities, there is little evidence to support dietary niche partitioning among conodonts. We tested the hypothesis that individual conodont taxa occupied different trophic niches, using Sr/Ca and Ba/Ca ratios preserved in the dental elements of assemblages from Silurian strata of Gotland, Sweden. Sr/Ca and Ba/Ca have been shown to vary in vertebrate skeletal tissues depending on trophic positioning, although biological and environmental conditions can affect these ratios. Environmental influences were minimized by examining entire conodont communities from a tropical epeiric sea and by measuring strontium isotope ratios using thermal ionization mass spectrometry in the most metropolitan taxon (Ozarkodina confluens). Composition of white matter, a tissue unique to conodonts, was also analyzed using microprobe analysis, revealing significantly lower Sr concentrations than in surrounding lamellar tissue, suggesting taxon-specific histology should be considered when analyzing conodonts for geochemical data. Excluding taxa with highly variable quantities of white matter, the results show that each taxon preserves different Sr/Ca and Ba/Ca ratios with limited overlap, indicating variation in trophic positioning