This thesis deals with the ontogeny of trilobites and agnostids from the Cambrian of the Skryje-Týřovice and Příbram-Jince basins (Barrandian area, Bohemian Massif). Thesis is presented as a compilation of four papers that were published in scientific peer-reviewed journals. The introduction of the thesis serves as a general overview of the trilobite post- embryonic development and connects all published papers together. All four papers presented here are focussed on modern description of the post-embryonic development of Sao hirsuta, Ptychopariida sp. A, Ptychopariida sp. B, Ptychopariida sp. C, Ellipsocephalus hoffi, Ellipsocephalus polytomus, Hydrocephalus carens and Ecca- paradoxides pusillus. In addition, a discussion about the life-history strategies of early developmental trilobite stages is presented. It has been suggested that Sao hirsuta and Ptychopariida sp. A had benthic protaspides and that the early ontogenetic stages of Ptychopariida sp. B, Eccaparadoxides pusillus and Hydrocephalus carens were likely lecithortophic. Consequently, it has been documented that species with benthic protaspides show a strongly restricted geographic distribution and that lecithotrophic taxa seem to be associated with higher latitude areas. The developmental patterns of Ellipsocephalus hoffi and...

1 Abstract In recent years, the term small carbonaceous fossils (SCFs) has been established to accomodate fragile organic-walled fossils of micro- to mesoscopic size that are usually extracted by means of the 'low-manipulation HF extraction' method. This method has so far only been utilized by several authors and no reports have been published on the usage of the method on samples from the Barrandian area. To test the applicability of the method, samples from the Barrandian area were processed. The samples came from eleven localities representing six stratigraphic units (Paseky Shale, Jince Formation, Letná Formation, Kosov Formation, Daleje Shale and Roblín Member). All the units have been studied before by 'standard' methods of palynological processing. Various previously unreported fossils are described, including wiwaxiid sclerites, putative chaetognath remains and acritarch clusters. Furthermore, it is discovered, that the 'low-manipulation HF extraction' can also be used for extraction of originally calcareous fossils. The pilot study provides a further proof that the 'low-manipulation HF extraction' is a useful tool with the potential to significantly expand our knowledge of fossil assemblages. Key words: Small carbonaceous fossils, Cambrian, Ordovician, Devonian, Barrandian area

This thesis deals with the topic of eocrinoid echinoderms from the Barrandian area (Příbram- Jince Basin, mid-Cambrian, Drumian). The thesis is presented as a compilation of five published papers and an introduction. The introduction is an overview of current knowledge about eocrinoid echinoderms. It also includes links to new information from the five included papers, and serves as a unifying element for these professional publications. The topic of the thesis is focused on a revision of selected specimens of eocrinoid echinoderms, and the study of their ontogeny, phylogeny, palaeoecology and systematic position. This thesis has also improved our understanding of early evolution of basal blastozoans. This is the first study of Cambrian ontogenetic development on eocrinoid material from the Czech Republic. Complete ontogeny was described on the genus Akadocrinus, and it was possible to establish two basic phases in the development of this genus: pre-epispire-bearing phase and epispire-bearing phase. The new primitive blastozoan Felbabkacystis is described from the Jince Formation - its unique body plan highlights evolution of the body wall among blastozoans. Palaeoecology and phylogeny of the genus Vyscystis was studied, and phylogenetic analysis suggests a basal position of lepidocystoids among...

New brachiopod materials from the lower Cambrian (Global Stage 3 and lower Stage 4) of Esmeralda County, Nevada are described and their age distribution and diversity is discussed. The specimens originate from the Fritzaspis, “Fallotaspis”, “Nevadella” and Bonnia-Olenellus biozone which in the area correspond to the Campito Formation and the overlying Poleta Formation. As the specimens from the Gold Coin Member (Begadean Series) and the Montenegro Member (Waucoban Series) of the Campito Formation encompass the oldest samples, they were therefore prioritised as there was not enough time to describe all the specimens. The oldest known trilobites from Laurentia have previously been reported from the Gold Coin Member and it is of interest to determine if the brachiopods from the same member also are the oldest known from Laurentia.As many specimens are poorly preserved, especially the ones from coarse siliciclastic intervals, the determination of their systematic position is somewhat aggravated. Despite this, a diverse fauna including Lingulida, Obolellida, Naukatida, Kutorginida, Orthida and stem-group brachiopods have been successfully identified. A total of 16 taxa were noted for the Campito fauna, including the problematic and rare genera Swantonia and Spinulothele but unfortunately little new information could be gained regarding the characters for these rare genera. Four genera are reported for the first time from the area, not including material described under open nomenclature. / Under den geologiska tidsåldern Kambrium (542-488 miljoner år sedan) uppträder de första fossilerna av djurgruppen brachiopoder, eller armfotingar som de också kallas, tillsammans med många andra djurgrupper. Få platser i Laurentia (paleo-kontinent bestående av nutidens Nordamerika och Grönland) finns kvar som har bevarat de tidigaste förekommande brachiopod fossilerna. Esmeralda län i Nevada är en sådan plats och det här har det även spekulerats att Laurentias absolut äldsta brachiopod fossil skulle kunna hittas där. Denna avhandling fokuserar på att studera dessa brachiopod fossil från Nevada. När åldern för de äldsta studerade brachiopoderna (Begadean I ålder) i denna avhandling jämfördes med tidigare dokumenterade åldrar för äldre lägre Kambrium brachiopod fossiler stod det klart att dessa är de äldsta brachiopod fossilerna som hittats hittills.Många nya fossil prover har samlats in av J.S. Hollingsworth under en längre tid från Nevada och beskrivs häri. De nya fossilen innefattar både sällsynta släkten, så som Swantonia samt Spinulothele, och släkten som inte har hittats i Esmeralda län tidigare. Tyvärr erhölls lite ny information beträffande de sällsynta släktenas morfologiska egenskaper. Upptäkterna om de nya släkterna i området skulle kunna hjälpa oss att bättre kunna rekonstruera brachiopod faunan från Nevada samt att förstå spridningen av olika brachiopod grupper i framtiden.

Research on the Marjum Formation provides insight into facies transitions in the House Range embayment of southwestern Utah during the Middle Cambrian. Sections of the Marjum Formation and equivalents were measured in strata representing shallow- (Drum Mountains), intermediate- (Wheeler Amphitheater), and deep-ramp (Marjum Pass) environments. This traverse across the embayment reveals strong litho- and biofacies changes. The Drum Mountains strata (343 m thick) are dominated by thickbedded shallow-water limestone facies containing polymerid trilobites and stromatolites. The Wheeler Amphitheater locality (392 m thick) provides an example of intermediate deposits between deep-water and shallow-water environments and is characterized by limestone and subordinate shale facies containing polymerid and agnostid trilobites, brachiopods, and sponge spicules. The Marjum Pass locality (424 m thick) contains equal parts shale and limestone facies with abundant fossils that include polymerid and agnostid trilobites, sponge spicules and soft bodied algae, representing deep-water environments. Migration of litho- and biofacies observed within Marjum Formation sections across the embayment shows carbonate ramp build-up and progradation from the Drum Mountains to the Swasey Peak, Wheeler Amphitheater area. Lateral migration of deepwater shale and rhythmite-producing turbidite facies can also be observed on a larger timescale from the Drum Mountains, during the early Bolaspidella time (Wheeler Formation deposits), to the Marjum Pass area (Marjum Formation deposits) by late Bolaspidella time. The Marjum Formation records two, third order shoaling-upward sequences. Fourth, fifth, and higher-frequency (rhythmite) cycles superimposed on these third order sequences can also be identified within the formation and are best preserved within the relatively deep-water deposits at Marjum Pass. Identification of sampled trilobites allowed correlation of known fauna! turnovers found by Robison and Vorwald with observed strata and systems tracts from this study. Stratigraphic locations of trilobite fauna! turnovers were found to be associated with transgressive systems tracts in the Drum Mountains and Marjum Pass localities. Fauna! turnover associated with significant sea-level events within these systems tracts suggests change in water depth altered the local environment forcing extinction and/or migration of organisms. Peak values of total organic carbon (TOC) at each measured section were also found to have a close relationship with maximum flooding zones. Associations of peak TOC values and fauna! turnovers with significant sea-level events demonstrate the value of these tools for correlation across the embayment.

The majority of the Middle Cambrian Wheeler Formation in the Drum Mountains was deposited during a single 3rd order sequence. Superimposed onto this sequence are three indistinct 4th order cycles and twenty distinct 5th order cycles. These higher order cycles were likely deposited within short intervals of geologic time (204 to 405 ky). The lower sequence boundary zone occurs within the Swasey Formation. The Transgressive Surface is the contact between the Swasey and Wheeler Formations. The Maximum Flooding Surface is located near the top of the lower Wheeler Formation, which also approximates the base of the Ptychagnostus atavus range zone. The upper sequence boundary is marked by stromatolites, which occur near the top of the upper member of the Wheeler Formation in the Drum Mountains. Deposition of the Wheeler Formation in the Drum Mountains was controlled by eustacy and tectonics. Local normal faulting associated with Middle Cambrian postrifting thermal subsidence may have caused some of the 5th order cycles. The cycles and surfaces defined in this stratigraphic analysis, and the base of the Ptychagnostus atavus and P. gibbus range-zones, can be used to correlate strata occurring in other localities in the eastern Great Basin. In addition, this study enables the evaluation of the effect of tectonics (faulting) versus global eustacy on the sedimentary regime occurring within the Middle Cambrian House Range Embayment.

The Middle Cambrian Ute Formation was studied in the Bear River Range and the Wellsville Mountains of north-central Utah and southeastern Idaho. The depositional textures and sedimentary structures found within the rocks were compared with similar modern sediments and ancient rocks to determine depositional environments, paleogeography, and diagenetic alterations. The rocks of the Ute Formation were divided into five basic types. These five rock types were formed within four identifiable lithofacies: 1) elastic marine shelf; 2) carbonate marine shelf; 3) agitated shoal; and 4) quiet-water shoal. The sequence of elastic and carbonate sediments is believed to have been deposited in a shallow, subtidal environment. Clastic sediments from the east and northwest periodically prograded over the carbonate sequences. A major regression marks the base of the Ute Formation. This was followed by a series of transgressions and regressions, until a major transgression occurred near the end of the deposition of the Ute. Paleomagnetic and faunal evidence suggest the study area was within 10° of the equator during the Middle Cambrian. Clay mineralogy of insoluble residues indicates a humid, tropical climate. Primary diagenetic features are compaction, micritization, and cementation. Secondary diagenetic changes include the inversion of high-magnesium calcite to low-magnesium calcite, aggrading neomorphism, stylolitization, fracturing, and calcite infilling. Partial dolomitization of grains and/or matrix is believed to result from the release of magnesium due to the decomposition of magnesium-rich, organic matter. The formation of a lens-shaped body of dolostone may have resulted from dolomitization by a magnesium-rich fluid circulating along faults.

The Middle Cambrian Wheeler Formation is interpreted as having been deposited in the shallow ramp and deeper basin environments of the House Range embayment (HRE), presumably, during a single third-order sequence. In the Drum Mountains, the Wheeler Formation (295 m thick) is dominated by proximal and distal ramp deposits; at Ma~um Pass, the Wheeler Formation (190m thick) is dominated by basinal shale deposits. The Wheeler Formation contains only one biozone marker; the first appearance of Ptyhagnostus atavus. Lack of other chronostratigraphic markers and distinctive stratal patterns in the basinal facies makes correlation along this ramp-to-basin transect difficult. Therefore, carbon-isotope stratigraphy and total organic carbon analysis were tested for their utility as intra basinal correlation tools. 813Ccarbonate isotope values range from -1.7% to 0.07%o (PDB) at Marjum Pass and -1.1% to 1.4% (PDB) in the Drum Mountains; previously reported 813Ccarbonate values in the Great Basin for this time interval range between -2% to 2% (PDB). Both localities show small-scale isotope variability, however, this variability is thought to be the result of local isotopic effects and was not used for correlation. TOC values obtained from both sections increase upsection, define a distinct peak, then decrease upsection. These peaks are associated with shale facies and occur near the maximum flooding surface in both sections, indicating that the TOC results could be used for correlation between sections. The lithologic cyclicity recognized in the shallow-water deposits at the Drum Mountains locality have also been recognized in the deeper-water deposits at Ma~um Pass. At each locality the meter-scale cycles shallow upward and display similar stacking patterns. Because cyclicity is preserved in both sections and the total stratigraphic thickness and cycle thickness decrease toward the embayment-controlling fault, it is probable that the cyclicity was the result of small-scale eustatic changes in sea level rather than episodic tectonism. This ramp-to-basin correlation also supportS the validity of P. atavus as a global biostratigraphic marker. The first appearance of Ptydnagostus atavus has been found below the interpreted maximum flooding surface and was coeval with transgression in both localities, indicating that its appearance was likely synchronous.

The Lower Middle Cambrian Langson Formation was studied in the xi Bear River Range of north-central Utah and southeasternmost Idaho and the Wellsville Mountains of north-central Utah. The depositional textures and sedimentary structures preserved within the rocks were compared with characteristics of similar modern sediments and ancient rock to determine environments of deposition, paleogeography, diagenetic alteration and pattern of dolomitization. The rocks of the Langston Formation were divided into eleven different rock types. These eleven rock types were formed within four recognizable lithofacies: 1) upper peritidal; 2) inner carbonate shelf; 3) inner clastic shelf; and 4) outer clastic shelf. The general depositional environment is inferred to have been a shall ow subtidal to subaerial carbonate shoal complex. Clastic sediments from the east and north or northwest periodically prograded over the carbonate complex during times of relatively slow subsidence. The deposition of the Langston Formation mudrocks and carbonates occurred during the first Cambrian grand cycle. Eogenetic diagenetic features include birdseye structures, relict evaporite structures, fibrous rim cement, compaction, and the begining of dolomitization. Mesogenetic diagenesis is characterized by dolomitization and pressure solution. Telogenetic diagenesis is limited to fracturing and calcite infilling. Dolomitization is believed to have resulted mainly from downward reflux of hypersaline brines, as indicated by relict evaporite structures, zoned dolomite rhombs, and a general association of dolomite with upper peritidal facies. The hypersaline brines formed in the upper peritidal environment, and percolated downward through underlying porous sediments. The greater density of the hypersaline brines displaced less-dense interstitial fluids. These brines were periodically diluted by normal marine water or fresh water.

The Cambrian explosion not only shaped the ecosystems of the Phanerozoic, but fundamentally changed how biota interacts with the environment, including the substrate. Cambrian trace fossils Alectorurus and Syringomorpha as well as Ordovician-Silurian Daedalus represent novel modes of faunal interaction with the newly innovated shallow marine mixed-ground and have surprisingly limited stratigraphic ranges. This study investigates vertically oriented spreiten trace fossils from the Armorican Quartzite Formation near Castañar de Ibor, Spain and the File Haidar Formation near Hällekis, Sweden. Site and trace fossil description was conducted to compare and contrast their morphology, ecological relationships, and potential affinities. While all these fossils feature J-shaped vertically oriented spreiten structures, Alectorurus is the most closely comparable Cambrian ichnofossil to the largely Ordovician Daedalus due to their similarity in overall size, morphology, and sedimentary environment. Alectorurus represents the earliest known occurrence of animal behavior comparable to Daedalus within similar facies. This short-lived ichnofossils may have gone extinct due to complexifying trophic chains in the shallow marine realm as brought forth from heightened predation and land-plant derived organics. Interpretations of the affinity of Daedalus, regarding the construction, and ecological niche can be extended to Alectorurus and may be used to help piece together the construction, ecology and affinities of these enigmatic Cambrian and Ordovician trace fossils.