Uma análise sistemática dos braquiópodes (Brachiopoda, Calciata) da Formação Ponta Grossa, Devoniano, Sub-bacia Apucarana, Estado do Paraná, Bacia do Paraná, Brasil, é apresentada. Os Brachiopoda (Calciata) da Formação Ponta Grossa, Bacia do Paraná, Estado do Paraná, Brasil, foram inicialmente estudados por John Mason Clarke em 1913 e, desde então, nunca foram o alvo de uma revisão sistemática apropriada. Estudos tafonômicos prévios sobre outros grupos da mesma localidade sugerem que muitas espécies descritas para ela podem corresponder a espécies tafonômicas. Os objetivos do presente trabalho foram: a- revisar a sistemática dos braquiópodes Calciata da Formação Ponta Grossa, Devoniano, a partir do estudo morfológico do material fóssil disponível; b- verificar a diversidade da fauna de braquiópodes Calciata dessa Formação, comparando a trabalhos da Formação Ponta Grossa do Estado do Mato Grosso; c- verificar se alguns caracteres morfológicos utilizados na diagnose de gêneros e espécies de braquiópodes Calciata da Formação Ponta Grossa, do Estado do Paraná, são susceptíveis a modificações tafonômicas, particularmente devido à compactação post-mortem e a esfoliação, conforme ocorre com outros grupos fósseis, tais como conulários e trilobitas. Os resultados mostraram que, das espécies previamente descritas, Australostrophia mesembria, Pleurochonetes falklandicus, Cryptonella baini e Australocoelia palmata são válidas e permanecem sem modificação; a espécie Coelospira (?) colona pôde ser somente identificada a nível genérico devido à falta de dados disponíveis nas amostras analisadas. Os gêneros Australospirifer, Schuchertella e Derbyina apresentaram problemas na identificação. Autralospirifer é considerado um gênero válido e possui cinco espécies descritas para a Formação Ponta Grossa, porém os caracteres diagnósticos que as diferem são passíveis de modificação tafonômica, fato evidenciado em exemplares analisados, além de apresentarem um grau de variação alto, o que impede a identificação dessas espécies. Duas espécies de Schuchertella são descritas para a Formação Ponta Grossa, gênero que sofreu uma revisão taxonômica, excluindo essas espécies da classificação genérica. Tais espécies poderiam pertencer ao gênero Floweria, porém a falta de dados nas amostras impede a validação desta hipótese. Derbyina é um gênero controverso, muito similar ao gênero Paranaia, os quais recentes revisões afirmam serem distintos. Devido à falta de dados no material examinado, especificamente o braquídio, não foi possível confirmar a distinção entre esses gêneros nem uma identificação específica. O presente trabalho fortaleceu a ideia de que a designação de novas espécies deve ser realizada perante a presença de diversos espécimes, para que o maior espectro possível de variações tafonômicas e morfológicas decorrentes do processo de fossilização seja identificado, com holótipos bem preservados, o que diminui a margem de erros. Portanto, é importante aliar a tafonomia com o estudo sistemático, a fim de identificar caracteres sujeitos à maior variação tafonômica. / A systematic analysis of the Brachiopoda (Calciata) from Ponta Grossa Formation, Devonian, Apucarana Sub-basin, Paraná Basin, Paraná State, Brazil, is presented. The Brachiopoda (Calciata) from Ponta Grossa Formation, Paraná Basin, Paraná State, Brazil, were initially studied by John Mason Clarke in 1913 and, since then, were not the object of proper systematic revision. Previous taphonomic studies about other groups from the same location suggest that many species described may correspond to taphonomic species. The objectives of this article were: a- to revise the systematic of the Brachiopoda (Calciata) of the Ponta Grossa Formation, Devonian, through morphological study of the available fossil material; b- to verify the diversity of Calciata brachiopods of this Formation, comparing it to works from the Ponta Grossa Formation in the State of Mato Grosso; c- to verify if any of the morphological characters used in the diagnosis of genera and species of Calciata brachiopods from Ponta Grossa Formation, Paraná State, are susceptible to taphonomic modification, particularly post-mortem compaction and exfoliation, as occurs in other fossil groups, like conulariids and trilobites. The results have shown that, of the species previously described, Australostrophia mesembria, Pleurochonetes falklandicus, Cryptonella baini and Australocoelia palmata are valid and remain unchanged; the species Coelospira (?) colona could only be identified up to a generic level due to lack of data on the samples analyzed. The genera Australospirifer, Schuchertella and Derbyina presented problems during their identification. Autralospirifer is considered a valid genus and possesses five described species for the Ponta Grossa x Formation, although the diagnostic characters that differentiate them are susceptible to taphonomic modification, fact evidenced in analyzed samples, with them also presenting high variability degree, which impedes the identification of those species. Two species of Schuchertella are described to the Ponta Grossa Formation, a genus that has undergone taxonomic revision, excluding those species from the generic classification. Those species may belong in the Floweria genus, but the lack of data on the samples makes impossible the validation of this hypothesis. Derbyina is a controversial genus, very similar to the genus Paranaia, which recent revisions claim to be different from each other. Due to the lack of data in the examined material, specifically the brachidium, it was not possible to confirm the distinction between these genera, as well as specific identification. The present work has strengthened the idea that the designation of new species must be made in face of a considerable number of samples, so that the largest spectrum of taphonomic and morphological variations resulting from the fossilization process is identified, which diminishes the error margin. Therefore, it is important to combine taphonomy with the systematic study, in order to identify characters which are subject to greater taphonomic variation.

Recent inquiry into the nomenclature of several species within Spinocyrtia has led to questions concerning name applicability and validity, particularly whether Delthyris granulosa and Spinocyrtia (Spirifer) granulosa are synonymous. This study utilized two-dimensional outline landmark analysis, a form of geometric morphometric analysis, to evaluate interspecific variation among these species. I took over a thousand photographs of over a hundred specimens of brachiopods belonging to the family Spinocyrtiidae. Ninety-six specimens originated from the Givetian outcrop belt of New York state, three were from northwestern Ohio, there was single Canadian specimen, and there was a single German specimen. The results from these analyses indicate that the mophospaces of Spinocyrtia (Spirifer) congesta, S. (Spirifer) granulosa, and S?. (Spirifer) marcyi are statistically (p < 0.05) distinct from one another.

No description available.

The Middle Devonian brachiopod, Mucrospirifer, was studied over most of its geographical and stratigraphical range in order to test the theory of punctuated equilibria. Punctuated equilibria holds that most evolution is concentrated in rapid speciation events and that the subsequent lifespan of a species is then characterized by approximate morphological stasis. Although many fossil lineages have been studied since the theory of punctuated equilibria was proposed, very few have included broad geographical coverage. To study a species adequately through time, the extent of its morphological variation in space must be known. Specimens of Mucrospirifer were collected from New York, Ontario, Michigan, Pennsylvania, Maryland, and West Virginia from sandstones, siltstones, mudstones, calcareous shales, and limestones spanning 5-7 million years of geological history. Although many species of Mucrospirifer have been proposed, the most recent taxonomic work reduced the number to two and found that the overall shape of the shell and the shape of the fold and sulcus were the best criteria to distinguish species of Mucrospirifer. These same two criteria were applied in this study. Overall shape was examined by Fourier analysis of the outline of each usable adult specimen employing a video digitizing system assembled for this project. Selection of harmonics that contribute most to the shape of Mucrospirifer was aided by stepwise discriminant analysis, and only these harmonics were used in subsequent statistical analyses. Shape differences among adult specimens cannot be attributed to allometry which was not significant when linear regression of log-transformed data for harmonic 0 (size) vs harmonic 2 (shape) was applied. Using cluster analysis, polar ordination, and principal component analysis extensive continuous morphological variation of Mucrospirifer was detected over its geographical range as well as within populations, so that only a single polytypic species, Mucrospirifer mucronatus, could be identified. Although no strong correlation between morphology and lithology was found over the entire study area, using linear regression an east-west trend of increasing roundness was identified in New York state. This trend may be due to an adaptation to water turbulence that decreases from the nearshore in the east to the offshore in the west or to sediment type which becomes finer grained going from east to west. Using principal component analysis and linear regression, no significant directional change in overall shape could be detected. Therefore, morphological stasis characterizes the 5-7 million-year history of Mucrospirifer mucronatus in the study area. These findings lend substantial support to the theory of punctuated equilibria. / Ph. D.

The Brachiopoda, a major group of the Lophotrochozoa, experienced a rapid early evolutionary diversification during the well-known Cambrian explosion and subsequently dominated the Palaeozoic benthos with its diversity and abundance. Even though the phylogeny of the Lophotrochozoa is still hotly debated, it is now known that the Brachiopoda are a monophyletic grouping. However, the early evolutionary rates for the Brachiopoda have never been studied in the framework of a study combining molecular data and fossil time calibration points. In order to investigate the expected higher evolutionary rates of the Phylum at its origin, we conducted phylogenetic studies combining different methodologies and datasets. This work has at its foundation Maximum Likelihood and Bayesian analyses of 18S and 28S rRNA datasets followed by analyses of phylogenomic sequences. All material was obtained from previously available sequences and from sequencing of genetic material from specimens from a concerted worldwide collection effort.       While the analyses of the phylogenomic dataset produced a robust phylogeny of the Brachiopoda with good support, both the results of the novel rRNA and phylogenomic dating analyses provided limited insights into the early rates of evolution of the Brachiopoda from a newly assembled dataset, demonstrating some limitations in calibration dating using the software package BEAST2. Future studies implementing fossil calibration, possibly incorporating morphological data, should be attempted to elucidate the early rates of evolution of Brachiopoda and the effect of the Push of the Past in this clade. / Det är ofta antaget att evolution (förändringar i arvsmassan hos en grupp organismer) sker i en konstant hastighet men i slutändan ändå osäkert om så är fallet. Stora grupper av organismer har ofta associerats med en högre evolutionär hastighet, speciellt nära deras uppkomst, vilket ökar sannolikheten för överlevnad.  Armfotingar (Brachiopoda) är marina ryggradslösa djur med skal som tidigare var allmänt spridd, idag är istället musslor (Bivalvia) betydligt mer spridda. Armfotingar har funnits och utvecklats under flera miljoner år med ursprung under tidigt kambrium. Genom år av forskning och många fossil har vi fått mer information om utseendet hos utdöda organismer vilket har bidragit till att antalet fossila arter som vi känner till har ökat tusenfalt. Under den senaste tiden har det också skett innovationer inom molekylära tekniker som gjort det möjligt att applicera dessa kunskaper även på utdöda arter. Dessa molekylära tekniker har nyligen hjälpt till att bestämma några av släktskapsförhållandena inom armfotingar som tidigare ansetts vara väldigt svåra att lösa.  Det finns fortfarande vissa släktskapsförhållanden inom armfotingar som inte är kända och man vet ännu inte hur fort de utvecklades. Genom att undersöka just evolutionens hastighet kan man börja förstå gruppens tidiga framgång under Kambrium och Ordovicium samt minskningen som följde. Syftet med den här studien var att beräkna evolutionshastigheten hos armfotingar med särskild fokus på den tidiga diversifieringen av gruppen. För att undersöka detta använde vi oss av molekylära data för att analysera släktskapsförhållandena inom armfotingar. Dessutom använde vi fossil för att datera stora händelser i armfotingarnas evolutionära historia. Med hjälp av statistiska analyser kunde vi beräkna evolutionshastighet och släktskapsförhållandena inom gruppen. Vi kom fram till att armfotingar härstammar från en gemensam förfader. Dateringen kring när detta skedde blev inte fastställd då det beräknades ske miljoner år före det äldsta djurfossilet. Det kommer behövas mer forskning för att ta reda på om armfotingar hade en högre evolutionär hastighet i tidigt skede.

Graduation date: 1995

The evolutionary origins of animal phyla are intimately linked with the Cambrian explosion, a period of radical ecological and evolutionary innovation that begins approximately 540 Mya and continues for some 20 million years, during which most major animal groups appear. Lophotrochozoa, a major group of protostome animals that includes molluscs, annelids and brachiopods, represent a significant component of the oldest known fossil records of biomineralised animals, as disclosed by the enigmatic ‘small shelly fossil’ faunas of the early Cambrian. Determining the affinities of these scleritome taxa is highly informative for examining Cambrian evolutionary patterns, since many are supposed stem-group Lophotrochozoa. The main focus of this thesis pertained to the stem-group of the Brachiopoda, a highly diverse and important clade of suspension feeding animals in the Palaeozoic era, which are still extant but with only with a fraction of past diversity. Major findings include adding support for tommotiid affinity as stem-group lophophorates. Determining morphological character homologies vital to reconstructing the brachiopod stem-group was achieved by comparing Cambrian Lagerstätten with the widespread biomineralised record of Cambrian stem-brachiopods and small shelly fossils. Polarising character changes associated with the putative transition from scleritome organisms to crown-group brachiopods was furthered by the description of an enigmatic agglutinated tubular lophophorate Yuganotheca elegans from the Chengjiang Lagerstätte, China, which possesses an unusual combination of phoronid, brachiopod and tommotiid characters. These efforts were furthered by the use of X-ray tomographic techniques that revealed novel anatomical features, including exceptionally preserved setae in the tommotiid Micrina. The evidence for a common origin of columnar brachiopod shell structures in the tommotiids is suggested and critically examined. Enigmatic and problematic early and middle Cambrian lophotrochozoans are newly described or re-described in light of new evidence, namely: the stem-brachiopod Mickwitzia occidens Walcott from the Indian Springs Lagerstätte, Nevada; a putative stem-group entoproct Cotyledion tylodes Luo and Hu from Chengjiang, China; a new enigmatic family of rhynchonelliform brachiopods exemplified by the newly described Tomteluva perturbata from the Stephen Formation, Canada; and the tommotiid Micrina etheridgei (Tate) from the Flinders Ranges, South Australia. Cladistic analyses of fossil morphological data supports a monophyletic Brachiopoda.

Taxonomic position of the brachiopod genus Euorthisina HAVLÍ EK 1950 from the Ordovician of the Prague Basin (Czech republic) was subjected to wide discussions and disputes in the past. The genus was first classified within the superfamily Syntrophiiacea (order Pentamerida). Later, a separate family Euorthisinidae by Havlí ek (1977) was defined. At present, the family is assigned to the order Orthida and includes genera Euorthisina HAVLÍ EK, 1950, Lesserorthis BENEDETTO, 2007, Notorthisina HAVLÍ EK & BRANISA, 1980 and Protorthisina BENEDETTO, 2007. In this thesis, a revision of genus Euorthisina HAVLÍ EK 1950 with emphasis on taxonomic classification, palaeoecology and palaeogeographic occurrence of species E. moesta and E. minor described from the Ordovician of the Prague Basin has been made. Klí ová slova: Euorthisina, brachiopoda, Ordovician, Prague Basin, Czech republic.

The origin of many lophotrochozoan groups can be traced to “small shelly fossil” (SSF) faunas of the Early Cambrian. Antarctica is a key region of study, due to the continent’s known close geographical proximity to well-studied Australian and Indian basins in in the Cambrian. Few studies have focused on this region however, due to a paucity of data. Re-examination of camenellan sclerites from the Early Cambrian Shackleton Limestone of the Churchill Mountains of Antarctica has revealed a previously unidentified species of Dailyatia in the formation, co-occurring alongside previously described Dailyatia odyssei Evans and Rowell, 1990, as in the Arrowie Basin of Australia. Re-examination of material previously described as Kennardia sp. A and Kennardia sp. B has indicated that these taxa can likely be synonymized as a second species of Dailyatia. Dailyatia sclerites were also found in the temporally equivalent “Schneider Hills Limestone” formation, which cropsout in the Argentina Range of Antarctica. These specimens appear to belongto a third species of Dailyatia, suggesting that the spatial distribution of tommotiids in the Early Cambrian was more complex than previously recognized, and that the group may be useful in future biostratigraphic studies. A study ofthe Middle Cambrian (Drumian Stage) Nelson Limestone Formation of the Neptune Range, Antarctica has revealed a moderately diverse brachiopod and trilobite fauna. The brachiopods have strong faunal links to taxa from South Australia and India, as well as other parts of the Antarctic province, fitting independent strong evidence for a united East Gondwanan region in the Middle Cambrian. An unidentified camenellan tommotiid sclerite is also described from the Nelson Limestone. This extends the worldwide temporal range of the tommotiid clade into the Drumian Stage, and suggests that more basal members of the brachiopod stem-group survived to form part of a more diverse Middle Cambrian fauna.

The sesile suspension feeders of Brachiopoda and Polychaeta phyla are relatively well represented in the Bohemian Cretaceous Basin (BCB), and show significant diversity. All three subphyla of Brachiopoda (Linguliformea, Craniiformea and Rhynchonelliformea) are present in the fossil record of the sedimentary basin infill; from Polychaeta, there has hith- erto been evidence of only the Canalipalpata subclass members that created solid, usually carbonatic tubes. In both abundancy and diversity, the most richest associations of fossil brachiopods and polychaets occure in near-shore facies of the Cenomanian to the Lower Turonian, and hemipelagic facies of the Late Turonian age. Prior research, as shows the literature survey given by the author, has been focused mainly on representatives of both groups of near-shore facies origin. The aim of this thesis is to fill in the existing gaps in our understanding of these groups within the BCB. This objective is accomplished in the form of a series of studies (Sklenář & Simon 2009, Vodrážka & al. 2009, Sklenář & al., in press) elaborating the following topics in detail: (A) taxonomy, distribution, morphology, variabil- ity, evolutionary relationships and stratigraphic significance of cancellothyrid brachiopod Gyrosoria abundant in the Middle and particularly in...