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.

Approximately 600 to 500 million years ago, a major evolutionary radiation called the “Cambrian Explosion” gave rise to nearly all of the major animal phyla known today. This radiation is recorded by various fossil lagerstätten, such as the Burgess Shale in Canada, where soft-bodied animals are preserved in exquisite detail. Many Cambrian fossils are enigmatic forms that are morphologically dissimilar to their modern descendants, but which still provide valuable information when interpreted as stem-group taxa because they record the actual progression of evolution and give insight into the order of character acquisitions and homologies between living taxa. One such group of fossils is the anomalocaridids, large presumed predators that have had a complicated history of description. Their body has a trunk with a series of lateral lobes and associated gills, and a cephalic region with a pair of large frontal appendages, a circular mouth apparatus, stalked eyes and a cephalic carapace. Originally, two taxa were described from the Burgess Shale, Anomalocaris and Laggania, however data presented herein suggests that the diversity of the anomalocaridids was much higher. Newly collected fossil material revealed that a third Burgess Shale anomalocaridid, Hurdia, is known from whole-body specimens and study of its morphology has helped to clarify the morphology and systematics of the whole group. Hurdia is distinguished by having mouthparts with extra rows of teeth, a unique frontal appendage, and a large frontal carapace. Two species, Hurdia victoria and Hurdia triangulata were distinguished based on morphometric shape analysis of the frontal carapace. A phylogenetic analysis placed the anomalocaridids in the stem lineage to the euarthropods, and examination of Hurdia’s well-preserved gills confirm the homology of this structure with the outer branches of limbs in upper stem-group arthropods. This homology supports the theory that the Cambrian biramous limb formed by the fusion of a uniramous walking limb with a lateral lobe structure bearing gill blades. In this context, new evidence is present on the closely allied taxon Opabinia, suggesting that it had lobopod walking limbs and a lateral lobe structure with attached Hurdia-like gills. The diversity of the anomalocaridids at the Burgess Shale is further increased by two additional taxa known from isolated frontal appendages. Amplectobelua stephenensis is the first occurrence of this genus outside of the Chengjiang fauna in China, but Caryosyntrips serratus is an appendage unique to the Burgess Shale. To gain a better understanding of global distribution, a possible anomalocaridid is also described from the Sirius Passet biota in North Greenland. Tamisiocaris borealis is known from a single appendage, which is similar to Anomalocaris but unsegmented, suggesting this taxon belongs to the arthropod stem-lineage, perhaps in the anomalocaridid clade. Thus, the anomalocaridids are a widely distributed and highly diverse group of large Cambrian presumed predators, which provide important information relevant to the evolution of the arthropods.

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Nas últimas décadas, os Lagerstätten estiveram no centro das discussões relativas à história paleobiológica e geológica da Terra. Em particular, o Membro Crato da Formação Santana (Bacia do Araripe, Nordeste do Brasil) é um dos mais significantes Lagerstätten do Cretáceo já que registra invertebrados, vertebrados e plantas excepcionalmente preservados em sedimentos carbonáticos de um palaeolago. O principal objetivo desta dissertação é lançar luz sobre os processos de preservação responsáveis pela fossilização de insetos e peixes com tecidos moles em 3D. Lâminas petrográficas e diversas técnicas paleométricas - micro-Espectroscopia Raman, Fluorescência de raios-X (FRX) convencional, micro-FRX com fonte de luz sincrotron (RS-µFRX), emissão de Raios-X induzida por partículas (PIXE), microscopia eletrônica de varredura (MEV) e espectroscopia de energia dispersiva de Raios-X (EDS) - foram empregadas para caracterizar a rocha matriz, a morfologia e fidelidade de preservação dos tecidos moles e as composições elementares e moleculares dos fósseis em escala de centímetros e mícron. Os resultados revelam que, enquanto insetos e tecidos moles de peixes encontrados nos denominados calcários beges (BL) são substituídos por pseudomorfos de pirita framboidal (após oxidação da pirita), os quais ocorrem juntamente com possíveis substâncias poliméricas extracelulares secretadas por bactérias (EPS), tecidos de peixes dos calcários cinza (GL) são querogenizados. Em insetos, existe variação de tamanho dos framboides para dentro dos fósseis, que é aqui interpretada como produto do equilíbrio entre as taxas de difusão e de nucleação dos minerais. Além disso, a distribuição preferencial de Zn e Cu em estruturas piritizadas de insetos e peixes em comparação com a sua matriz é aqui considerada como sendo o resultado da fixação de elementos químicos em biofilmes bacterianos. Zn concentrado nos ossos de peixes com querogenizados e Fe/Cu observados em seus tecidos moles são considerados como tendo sido incorporados durante a vida dos peixes. No caso particular de peixes, modelo originalmente proposto para a preservação de metazoários do Pré-cambriano é aqui aplicado para explicar a variação de fossilização entre as fácies BL e GL. Lâminas petrográficas revelam que os GL têm geralmente teor de argila/matéria orgânica maior do que os BL, implicando que as taxas de soterramento poderiam ter sido mais intensas nos GL. Isto teria colocado carcaças em decomposição mais rapidamente na zona sedimentar de metanogênese, sendo formado o querogênio. Por outro lado, carcaças depositadas na fácies BL poderiam ter passado período mais longo na zona de redução de sulfato, o que teria levado à piritização generalizada. Além disso, a baixa porosidade do microespato, o cimento e a argila poderiam ter diminuído a migração de aceptores de elétrons dos processos de respiração bacteriana anaeróbia - particularmente redução de sulfato (RS) e metanogênese - e de seus produtos biogeoquímicos, estreitando a zona de RS, o que teria reduzido a influência da piritização em alguns níveis. Enquanto que a piritização resultou na preservação de fibras musculares em 3D, sarcolema, possíveis núcleos celulares, tendões e olhos, a querogenização preservou tecidos conjuntivos, tegumento e fibras musculares distorcidas e compactadas. Em conclusão, é aqui proposto que fácies influenciadas por processos paleoambientais e geobiológicos produziram fósseis com gradiente diferencial de fidelidade de preservação de acordo com cada via tafonômica seguida. / Over the past decades, the so-called Fossil Lagerstätten have been in the core of discussions concerning the palaeobiological and geological history of the Earth. In particular, the Crato Member from the Santana Formation (Araripe Basin, north-east Brazil) is one of the most significant Cretaceous Lagerstätten since it records exceptionally well-preserved fossil invertebrates, vertebrates, and plants, deposited in palaeolake carbonate beds. The main aim of this dissertation is to shed light on the preservational processes responsible for the fossilization of insects and fishes still retaining 3D soft tissues. Petrographic thin sections and several palaeometric techniques - micro-Raman Spectroscopy, conventional X-ray Fluorescence (XRF), Synchrotron micro-XRF (SR-µXRF), Particle-Induced X-ray Emission (PIXE), Scanning Electron Microscopy (SEM), and Energy-Dispersive X-ray Spectroscopy (EDS) - have been employed to characterize the host rock, soft-tissue morphology and preservational fidelity, and fossil elemental and molecular compositions in centimetre to micrometre scale. The results reveal that while insects and fish soft-tissues found in the so called beige limestones (BL) are replaced by framboidal pyrite pseudomorphs (after pyrite oxidation) occurring together with putative bacterially-secreted extracellular polymeric substances (EPS), labile tissues of fishes from the grey limestones (GL) are kerogenized. In insects, there is a variation of framboid size inward the fossils, which is here interpreted as a product of the balance between diffusion and mineral nucleation rates. Moreover, the preferential distribution of Zn and Cu in pyritized insect/fish labile structures in comparison to their rock matrix is here considered as being the result of element fixation in bacterial biofilms. Zn concentrated in bones of kerogenized fishes and Fe/Cu occurring in their soft tissues are considered to have been incorporated during fish life. In the particular case of fishes, a model originally proposed for metazoan preservation in the Precambrian is here applied to explain the variation of fossilization between the BL and GL facies. Petrographic thin sections reveal that GL have generally higher clay/organic matter contents than BL, thus implying that burial rates might have been more intense in the former. This could have placed decaying carcasses more quickly in the methanogenesis sedimentary zone, in that way being kerogenized. On the other hand, carcasses deposited in the BL facies could have spent a longer period in the sulphate-reduction zone, which would have accounted for pervasive pyritization. Additionally, microspar low porosity, cement and clay could have diminished both downward migrations of electron acceptors for anaerobic bacterial respiration processes - particularly sulphate-reduction (SR) and methanogenesis - and of their biogeochemical products, narrowing the SR zone, which would have lowered the impact of pyritization in some levels. While pyritization has recorded 3D muscle fibres, sarcolemma, putative cell nuclei, tendons and eyes, kerogenization has yielded connective tissues, integument and compressed/distorted muscle fibres. In conclusion, it is here proposed that palaeoenvironmental/geobiological-influenced facies have yielded fossils with a variable preservational-fidelity gradient, accordingly to each taphonomic pathway followed.