A Bacterial Enrichment Study and Overview of the Extractable Lipids from Paleosols in the Dry Valleys, Antarctica: Implications for Future Mars Reconnaissance

The Dry Valleys of Antarctica are one of the coldest and driest environments on Earth with paleosols in selected areas that date to the emplacement of tills by warm-based ice during the Early Miocene. Cited as an analogue to the martian surface, the ability of the Antarctic environment to support microbial life-forms is a matter of special interest, particularly with the upcoming NASA/ESA 2018 ExoMars mission. Lipid biomarkers were extracted and analyzed by gas chromatography-mass spectrometry to assess sources of organic carbon and evaluate the contribution of microbial species to the organic matter of the paleosols. Paleosol samples from the ice-free Dry Valleys were also subsampled and cultivated in a growth medium from which DNA was extracted with the explicit purpose of the positive identification of bacteria. Several species of bacteria were grown in solution and the genus identified. A similar match of the data to sequenced DNA showed that Alphaproteobacteria, Gamma-proteobacteria, Bacteriodetes, and Actinobacteridae species were cultivated. The results confirm the presence of bacteria within some paleosols, but no assumptions have been made with regard to in situ activity at present. These results underscore the need not only to further investigate Dry Valley cryosols but also to develop reconnaissance strategies to determine whether such likely Earth-like environments on the Red Planet also contain life.

Coleoptera and microbe biomass in Antarctic Dry Valley paleosols adjacent to the Inland Ice: Implications for Mars

Bulk paleosol samples collected from a Middle to Early Miocene moraine in the New Mountain area of the Dry Valleys, Antarctica, yielded Coleoptera exoskeletons and occasional endoskeletons showing considerable diagenetic effects along with several species of bacteria, all lodged in a dry-frozen but salt-rich horizon at shallow depth to the land surface. The till is at the older end of a chronologic sequence of glacial deposits, thought to have been deposited before the transition from wet-based to cold-based ice (similar to 15 Ma), and hence, entirely weathered in contact with the subaerial atmosphere. It is possible, though not absolutely verifiable, that the skeletons date from this early stage of emplacement having undergone modifications whenever light snowmelt occurred or salt concentrations lowered the freezing temperature to maintain water as liquid. Correlation of the Coleoptera species with cultured bacteria in the sample and the likelihood of co-habitation with Beauveria bassiani found in two adjacent, although younger paleosols, leads to new questions about the antiquity of the Coleoptera and the source of N and glucose from chitinase derived from the insects. The skeletons in the 831 section may date close to the oldest preserved chitin (Oligocene) yet found on Earth. While harsh Martian conditions make it seemingly intolerable for complex, multicellular organisms such as insects to exist in the near-surface and subaerially, life within similar cold, dry paleosol microenvironments (Cryosols) of Antarctica point to life potential for the Red Planet, especially when considering the relatively diverse microbe (bacteria and fungi) population. (C) 2011 Elsevier Ltd. All rights reserved.

Phylogénie et biogéographie du genre Bauhinia s.l. (Leguminosae)

Bauhinia s.l. est le plus vaste genre de la tribu des Cercideae (Ceasalpinioideae, Leguminoseae), avec plus de 300 espèces. Il présente une distribution pantropicale et une grande variabilité morphologique. Ces deux caractéristiques ont limité les études taxonomiques sur le genre complet, résultant en plusieurs études taxonomiques de certains groupes seulement. En 1987, Wunderlin et al. proposent une vaste révision taxonomique de la tribu des Cercideae, basée sur des données morphologiques, et divisent le genre Bauhinia en quatre sous-genres. En 2005, Lewis et Forest publient une nouvelle classification préliminaire basée sur des données moléculaires, mais sur un échantillonnage taxonomique restreint. Leurs conclusions remettent en question le monophylétisme du genre Bauhinia et suggèrent plutôt la reconnaissance de huit genres au sein du grade Bauhinia s.l. Afin de vérifier les hypothèses de Lewis et Forest, et obtenir une vision plus claire de l’histroire de Bauhinia s.l., nous avons séquencé deux régions chloroplastiques (trnL-trnF et matK-trnK) et deux régions nucléaires (Leafy et Legcyc) pour un vaste échantillonnage représentatif des Cercideae. Une première phylogénie de la tribu a tout d’abord été réalisée à partir des séquences de trnL-trnF seulement et a confirmé le non-monoplylétisme de Bauhinia s.l., avec l’inclusion du genre Brenierea, traditionnellement reconnu comme genre frère de Bauhinia s.l. Afin de ne pas limiter notre vision de l’histoire évolutive des Cercideae à un seul type de données moléculaires et à une seule région, une nouvelle série d’analyse a été effectuée, incluant toutes les séquences chloroplastiques et nucléaires. Une phylogénie individuelle a été reconstruite pour chacune des régions du génome, et un arbre d’espèce ainsi qu’un arbre de supermatrice ont été reconstruits. Bien que certaines contradictions apparaissent entre les phylogénies, les grandes lignes de l’histoire des Cercideae ont été résolues. Bauhinia s.l. est divisée en deux lignées : les groupes Phanera et Bauhinia. Le groupe Bauhinia est constitué des genres Bauhinia s.s., Piliostigma et Brenierea. Le groupe Phanera est constitué des genres Gigasiphon, Tylosema, Lysiphyllum, Barklya, Phanera et Schnella. Les genres Cercis, Adenolobus et Griffonia sont les groupes-frères du clade Bauhinia s.l. Au minimum un événement de duplication de Legcyc a été mis en évidence pour la totalité de la tribu des Cercideae, excepté Cercis, mais plusieurs évènements sont suggérés à la fois par Legcyc et Leafy. Finalement, la datation et la reconstruction des aires ancestrales de la tribu ont été effectuées. La tribu est datée de 49,7 Ma et est originaire des régions tempérées de l’hémisphère nord, probablement autour de la mer de Thétys. La tribu s’est ensuite dispersée vers les régions tropicales sèches de l’Afrique, où la séparation des groupes Bauhinia et Phanera a eu lieu. Ces deux groupes se sont ensuite dispersés en parallèle vers l’Asie du sud-est au début du Miocène. À la même période, une dispersion depuis l’Afrique de Bauhinia s.s. a permis la diversification des espèces américaines de ce genre, alors que le genre Schnella (seul genre américain du groupe Phanera) est passé par l’Australie afin de rejoindre le continent américain. Cette dispersion vers l’Australie sera également à l’origine des genres Lysiphyllum et Barklya

Diversification of a large genus in a continental biodiversity hotspot: temporal and spatial origin of Muraltia (Polygalaceae) in the Cape of South Africa

Phylogenetic relationships in the largely South African genus Muraltia (Polygalaceae) are assessed based on DNA sequence data (nuclear ribosomal ITS, plastid atpB-rbcL spacer, trnL intron, and trnL-F spacer) for 73 of the 117 currently recognized species in the genus. The previously recognised subgenus Muraltia is monophyletic, but the South African endemic genus Nylandtia is embedded in Muraltia subgenus Psiloclada. Subgenus Muraltia is found to be sister to subgenus Psiloclada. Estimates show the beginning of diversification of the two subgenera in the early Miocene (Psiloclada, 19.3+/-3.4 Ma; Muraltia, 21.0+/-3.5 Ma) pre-dating the establishment of the Benguela current (intermittent in the middle to late Oligocene and markedly intensifying in the late Miocene), and summer-dry climate in the Cape region. However, the later increase in species numbers is contemporaneous with these climatic phenomena. Results of dispersal-vicariance analyses indicate that major clades in Muraltia diversified from the southwestern and northwestern Cape, where most of the species are found today.

A história do intemperismo na Província Borborema Oriental, Nordeste do Brasil: implicações paleoclimáticas e tectônicas

Until some years ago, weathering geochronology was primarily based on the K-Ar and 40Ar/39Ar dating of supergene minerals. Recent advances in the analysis of supergene goethite by the (U-Th)/He method expanded the number of suitable minerals for such purpose, as well as the time of application for weathering geochronology. This study represents the first systematic approach in Brazil, combining both the 40Ar/39Ar e (U-Th)/He methodologies to improve the knowledge on the weathering and the age of nonfossiliferous sediments. Supported by geologic and geomorphologic correlations, we identified different types of weathering profiles occurring in the interior and coastal areas of northeastern Brazil. These profiles were correlated to main regional geomorphological domains: the Borborema Plateau , the Sertaneja Depression , and the Coastal Cuestas and Plains, and respective planation surfaces, which study is fundamental to understand the landscape evolution of the northern portion of the eastern Borborema Province. The depth and stratigraphic organization of the weathering profiles in each of the geomorphological domains permitted to establish that: (i) the profiles on the highlands that cap the Borborema Surface are deeper (up to 100 m) and can be considered as typical lateritic profiles; (ii) on the lowlands that form the Sertaneja Surface , the weathering profiles are shallow and poorly developed (2-5 m deep); (iii) the profiles along the coastal area are moderately developed (up to 25 m deep), and are characterized by thick saprolites and mottle zones. Aiming to establish the timing of the evolution of northeastern Brazil, we studied 29 weathering profiles representing distinct topographic levels of the Borborema Province, from the highlands to the coast, through the analysis of 248 grains of supergene manganese oxides using laser step-heating 40Ar/39Ar geochronology. Additionally, we applied the (U-Th)/He method in 20 weathering profiles, by dating 171 grains of supergene iron oxides and hydroxides. Geochronological results for 248 grains of manganese oxides analyzed by the 40Ar/39Ar method indicate that the weathering profiles in the study area record the history of weathering from the Oligocene to the Pleistocene, with ages in the order of 31.4 ± 1.0 Ma to 0.8 ± 0.4 Ma. Dating of 171 grains of goethite by the (U-Th)/He method yielded ages ranging from 43.2 ± 4.3 Ma to 0.8 ± 0.1 Ma, suggesting the weathering processes last from the Eocene to the Pleistocene. The precipitation of supergene goethite in this interval confirms the age of the weathering processes identified from the manganese oxides record. 105 goethite grains from 8 different occurrences of the Barreiras Formation were dated by the (U-Th)/He method. Five grains collected from the cement in the Barreiras Formation sandstones, in the Lagoa Salgada and Rio do Fogo coastal cuestas, yielded ages of 17.6 ± 1.8 Ma, 17.3 ± 1.7 Ma, 16.3 ± 1.6 Ma, 16.2 ± 1.6 Ma and 13.6 ± 1.4 Ma. Results of 69 goethite grains from authigenic pisoliths collected in 7 different localities also yielded concordant ages, varying from 17.8 ± 1.8 to 7.5 ± 0.8 Ma. Results obtained from 31 detrital grains are concordant in 3 distinct localities (Lagoa Salgada, Praia da Garças e Ponta Grossa); they vary in the range of 43.2 ± 4.3 to 21.6 ± 2.2 Ma, and indicate that the maximum age for the Barreiras Formation deposition is around 22 Ma. 40Ar/39Ar results for 15 manganese oxides grains associated with the Barreiras Formation weathering profiles, in 3 different localities, vary from 13.1 ± 0.9 to 7.7 ± 0.4 Ma, in the same range of ages obtained by the (U-Th)/He method. The systematic application of the 40Ar/39Ar and (U-Th)/He methods, respectively for manganese oxides and goethites, show that the Barreiras Formation sediments were already deposited since ca. 17 Ma, and that the weathering processes were active until ca. 7 Ma ago. The ages obtained from manganese oxides collected in the Cenozoic basalts (Macau Formation) also reveal a weathering history between 19 and 7 Ma, pointing to hot and humid conditions during most of the Miocene. 40Ar/39Ar ages yielded by manganese oxides associated with the Serra do Martins Formation vary from 14.1 ± 0.4 to 10.5 ± 0.3 Ma. On the other hand, (U-Th)/He ages from iron oxides/hydroxides collected in the Serra do Martins Formation mesas vary from 20.0 ± 2.0 to 5.5 ± 0.6 Ma, indicating that those sediments are older than 20 Ma. 40Ar/39Ar and (U-Th)/He results produced in this study are in agreement with paleoclimatic interpretations based on stable isotopes and clay index values measured in the Atlantic Ocean sediments, validating the use of weathering geochronology to investigate paleoclimatic variations. The direct dating of the Barreiras Formation permitted, for the first time, confident inferences on the age of the brittle deformation recorded by this sedimentary unit in the Rio Grande do Norte and Ceará states. The first event, syn-deposition, occurred during the early Miocene; an younger event, related to the post-depositional deformation of the Barreiras Formation, is associated with tectonic activity from the very early Miocene to the Holocene. In agreement with data from other areas, results obtained in this study reveal that the depth and complexity of the weathering profiles reflect the time of exposition of such areas to the weathering agents close to the surface. However, there is no clear relationship between ages vs. altitude. The depth and the stratigraphic organization of weathering profiles in northeastern Brazil, contrary to the southeastern Brazil pattern, do not vary toward the coast. In our study area, field observations reveal the presence of ancient, thick and complex lateritic profiles preserved in the sedimentary mesas on the Borborema Plateau, as younger, narrow and incipient ones occur in the dissected areas. Geochronological results obtained for these profiles yielded older ages on the high altitudes, and younger ages in the lowlands, suggesting the scarp retreatment is the most reliable model to explain the regional landscape evolution. However, in the coastal lowlands, the relatively older ages obtained indicate that more complexes processes were involved in the modeling of the local relief

Estruturas tectônicas cenozóicas na porção leste da Bacia Potiguar - RN

This thesis encompasses the integration of geological, geophysical, and seismological data in the east part of the Potiguar basin, northeastern Brazil. The northeastern region is located in South American passive margin, which exhibits important areas that present neotectonic activity. The definition of the chronology of events, geometry of structures generated by these events, and definition of which structures have been reactivated is a necessary task in the region. The aims of this thesis are the following: (1) to identify the geometry and kinematics of neotectonic faults in the east part of the Potiguar basin; (2) to date the tectonic events related to these structures and related them to paleoseismicity in the region; (3) to present evolutional models that could explain evolution of Neogene structures; (4) and to investigate the origin of the reactivation process, mainly the type of related structure associated with faulting. The main type of data used comprised structural field data, well and resistivity data, remote sensing imagery, chronology of sediments, morphotectonic analysis, x-ray analysis, seismological and aeromagnetic data. Paleostress analysis indicates that at least two tectonic stress fields occurred in the study area: NSoriented compression and EW-oriented extension from the late Campanian to the early Miocene and EW-oriented compression and NS-oriented extension from the early Miocene to the Holocene. These stress fields reactivated NE-SW- and NW-SE-trending faults. Both set of faults exhibit right-lateral strike-slip kinematics, associated with a minor normal component. It was possible to determine the en echelon geometry of the Samambaia fault, which is ~63 km long, 13 km deep, presents NE-SW trend and strong dip to NW. Sedimentfilled faults in granite rocks yielded Optically Stimulated Luminescence (OSL) and Single-Aliquot Regeneration (SAR) ages at 8.000 - 9.000, 11.000 - 15.000, 16.000 - 24.000, 37.000 - 45.500, 53.609 - 67.959 e 83.000 - 84.000 yr BP. The analysis of the ductile fabric in the João Câmara area indicate that the regional foliation is NE-SW-oriented (032o - 042o), which coincides with the orientation of the epicenters and Si-rich veins. The collective evidence points to reactivation of preexisting structures. Paleoseismological data suggest paleoseismic activity much higher than the one indicated by the short historical and instrumental record

From Amazonia to the Atlantic forest: Molecular phylogeny of Phyzelaphryninae frogs reveals unexpected diversity and a striking biogeographic pattern emphasizing conservation challenges

Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq)

Geologia e paleontologia do biohermito da Formação Pirabas (Mioceno Inferior)

O presente trabalho compreende o estudo geológico e paleontológico detalhado da litofácies recifal da Formação Pirabas, Estado do Pará, aflorante na Praia do Maçarico, município de Salinópolis, como um corpo pequeno, isolado e lenticular, que deve ter se desenvolvido na rampa interna da plataforma do Mar de Pirabas. O conteúdo biótico é rico e variado, composto por três grupos de algas coralíneas, poríferos, corais, briozoários, equinoides, foraminíferos, ostracodes, e moluscos. Foram reconhecidos neste recife em mancha três estágios de sucessão ecológica, estabilização, colonização e diversificação, onde as finas crostas micríticas reconhecidas na parte superior do estágio diversificação indicam degradação das condições ambientais favoráveis à acreção recifal, provavelmente devido a continentalização do sítio deposicional. O exame petrográfico revelou apenas uma microfácies deposicional, biomicrito, sem consideráveis variações texturais e composicionais. A cimentação marinha é predominante, representada por massas microcristalinas e cimentos micríticos, localmente neomorfisados para bioesparito, e detritos carbonáticos advindos do fluxo biogênico inconsolidado, resultantes principalmente da dispersão de cristais de aragonita dos fragmentos algálicos, acumulados sob a forma de lama micrítica. Para a gênese da bioconstrução as algas foram fundamentais na consolidação e modelagem do arcabouço, principalmente as macroalgas vermelhas, decisivas para a proteção e cimentação dos demais organismos.

Systematic and historical biogeography of the Bryconidae (Ostariophysi: Characiformes) suggesting a new rearrangement of its genera and an old origin of Mesoamerican ichthyofauna

Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP)

Nuculidae (Bivalvia) in the Cape Melville Formation, King George Island, Antarctica, with an overview of the bivalve fauna

Nuculid bivalves of the Cape Melville Formation (Early Miocene, King George Island) are reviewed. Ten bivalve taxa are listed from the formation in the families Nuculidae (two species), Sareptidae, Malletiidae, Limopsidae (two species), Limidae, Pectinidae, Hiatellidae, and Periplomatidae. The Nuculidae consist of two species of Leionucula Quenstedt, 1930. One of these, L. melvilleana n. sp., is described and the other consists of the two species named previously by Anelli et al. (2006), which are demonstrated to be synonymous and are assigned to the species Leionucula frigida (Anelli, Rocha-Campos, Santos, Perinotto & Quaglio 2006). This assemblage, dominated by protobranchs (89% of specimens), is a typical fauna of offshore soft substrates, with a few specimens transported from hard substrates nearby. The diversity of Nuculidae has decreased in the Antarctic region through the Cenozoic.

The First Occurrence in the Fossil Record of an Aquatic Avian Twig-Nest with Phoenicopteriformes Eggs: Evolutionary Implications

Background: We describe the first occurrence in the fossil record of an aquatic avian twig-nest with five eggs in situ (Early Miocene Tudela Formation, Ebro Basin, Spain). Extensive outcrops of this formation reveal autochthonous avian osteological and oological fossils that represent a single taxon identified as a basal phoenicopterid. Although the eggshell structure is definitively phoenicopterid, the characteristics of both the nest and the eggs are similar to those of modern grebes. These observations allow us to address the origin of the disparities between the sister taxa Podicipedidae and Phoenicopteridae crown clades, and traces the evolution of the nesting and reproductive environments for phoenicopteriforms. Methodology/Principal Findings: Multi-disciplinary analyses performed on fossilized vegetation and eggshells from the eggs in the nest and its embedding sediments indicate that this new phoenicopterid thrived under a semi-arid climate in an oligohaline (seasonally mesohaline) shallow endorheic lacustine environment. High-end microcharacterizations including SEM, TEM, and EBSD techniques were pivotal to identifying these phoenicopterid eggshells. Anatomical comparisons of the fossil bones with those of Phoenicopteriformes and Podicipediformes crown clades and extinct palaelodids confirm that this avian fossil assemblage belongs to a new and basal phoenicopterid. Conclusions/Significance: Although the Podicipediformes-Phoenicopteriformes sister group relationship is now well supported, flamingos and grebes exhibit feeding, reproductive, and nesting strategies that diverge significantly. Our multi-disciplinary study is the first to reveal that the phoenicopteriform reproductive behaviour, nesting ecology and nest characteristics derived from grebe-like type strategies to reach the extremely specialized conditions observed in modern flamingo crown groups. Furthermore, our study enables us to map ecological and reproductive characters on the Phoenicopteriformes evolutionary lineage. Our results demonstrate that the nesting paleoenvironments of flamingos were closely linked to the unique ecology of this locality, which is a direct result of special climatic (high evaporitic regime) and geological (fault system) conditions.

From Amazonia to the Atlantic forest: Molecular phylogeny of Phyzelaphryninae frogs reveals unexpected diversity and a striking biogeographic pattern emphasizing conservation challenges

Documenting the Neotropical amphibian diversity has become a major challenge facing the threat of global climate change and the pace of environmental alteration. Recent molecular phylogenetic studies have revealed that the actual number of species in South American tropical forests is largely underestimated, but also that many lineages are millions of years old. The genera Phyzelaphryne (1 sp.) and Adelophryne (6 spp.), which compose the subfamily Phyzelaphryninae, include poorly documented, secretive, and minute frogs with an unusual distribution pattern that encompasses the biotic disjunction between Amazonia and the Atlantic forest. We generated >5.8 kb sequence data from six markers for all seven nominal species of the subfamily as well as for newly discovered populations in order to (1) test the monophyly of Phyzelaphryninae, Adelophryne and Phyzelaphryne, (2) estimate species diversity within the subfamily, and (3) investigate their historical biogeography and diversification. Phylogenetic reconstruction confirmed the monophyly of each group and revealed deep subdivisions within Adelophryne and Phyzelaphryne, with three major clades in Adelophryne located in northern Amazonia, northern Atlantic forest and southern Atlantic forest. Our results suggest that the actual number of species in Phyzelaphryninae is, at least, twice the currently recognized species diversity, with almost every geographically isolated population representing an anciently divergent candidate species. Such results highlight the challenges for conservation, especially in the northern Atlantic forest where it is still degraded at a fast pace. Molecular dating revealed that Phyzelaphryninae originated in Amazonia and dispersed during early Miocene to the Atlantic forest. The two Atlantic forest clades of Adelophryne started to diversify some 7 Ma minimum, while the northern Amazonian Adelophryne diversified much earlier, some 13 Ma minimum. This striking biogeographic pattern coincides with major events that have shaped the face of the South American continent, as we know it today. (C) 2012 Elsevier Inc. All rights reserved.

Biogeographic and diversification patterns of Neotropical Troidini butterflies (Papilionidae) support a museum model of diversity dynamics for Amazonia

Background: The temporal and geographical diversification of Neotropical insects remains poorly understood because of the complex changes in geological and climatic conditions that occurred during the Cenozoic. To better understand extant patterns in Neotropical biodiversity, we investigated the evolutionary history of three Neotropical swallowtail Troidini genera (Papilionidae). First, DNA-based species delimitation analyses were conducted to assess species boundaries within Neotropical Troidini using an enlarged fragment of the standard barcode gene. Molecularly delineated species were then used to infer a time-calibrated species-level phylogeny based on a three-gene dataset and Bayesian dating analyses. The corresponding chronogram was used to explore their temporal and geographical diversification through distinct likelihood-based methods. Results: The phylogeny for Neotropical Troidini was well resolved and strongly supported. Molecular dating and biogeographic analyses indicate that the extant lineages of Neotropical Troidini have a late Eocene (33-42 Ma) origin in North America. Two independent lineages (Battus and Euryades + Parides) reached South America via the GAARlandia temporary connection, and later became extinct in North America. They only began substantive diversification during the early Miocene in Amazonia. Macroevolutionary analysis supports the "museum model" of diversification, rather than Pleistocene refugia, as the best explanation for the diversification of these lineages. Conclusions: This study demonstrates that: (i) current Neotropical biodiversity may have originated ex situ; (ii) the GAARlandia bridge was important in facilitating invasions of South America; (iii) colonization of Amazonia initiated the crown diversification of these swallowtails; and (iv) Amazonia is not only a species-rich region but also acted as a sanctuary for the dynamics of this diversity. In particular, Amazonia probably allowed the persistence of old lineages and contributed to the steady accumulation of diversity over time with constant net diversification rates, a result that contrasts with previous studies on other South American butterflies.

Thermochronologic and geodynamic evolution of the Pontides: Sakarya terrane (Karakaya Complex) and Istanbul terrane, Turkey.

An integrated array of analytical methods -including clay mineralogy, vitrinite reflectance, Raman spectroscopy on carbonaceous material, and apatite fission-track analysis- was employed to constrain the thermal and thermochronological evolution of selected portions of the Pontides of northern Turkey. (1) A multimethod investigation was applied for the first time to characterise the thermal history of the Karakaya Complex, a Permo-Triassic subduction-accretion complex cropping out throughout the Sakarya Zone. The results indicate two different thermal regimes: the Lower Karakaya Complex (Nilüfer Unit) -mostly made of metabasite and marble- suffered peak temperatures of 300-500°C (greenschist facies); the Upper Karakaya Complex (Hodul and the Orhanlar Units) –mostly made of greywacke and arkose- yielded heterogeneous peak temperatures (125-376°C), possibly the result of different degree of involvement of the units in the complex dynamic processes of the accretionary wedge. Contrary to common belief, the results of this study indicate that the entire Karakaya Complex suffered metamorphic conditions. Moreover, a good degree of correlation among the results of these methods demonstrate that Raman spectroscopy on carbonaceous material can be applied successfully to temperature ranges of 200-330°C, thus extending the application of this method from higher grade metamorphic contexts to lower grade metamorphic conditions. (2) Apatite fission-track analysis was applied to the Sakarya and the İstanbul Zones in order to constrain the exhumation history and timing of amalgamation of these two exotic terranes. AFT ages from the İstanbul and Sakarya terranes recorded three distinct episodes of exhumation related to the complex tectonic evolution of the Pontides. (i) Paleocene - early Eocene ages (62.3-50.3 Ma) reflect the closure of the İzmir-Ankara ocean and the ensuing collision between the Sakarya terrane and the Anatolide-Tauride Block. (ii) Late Eocene - earliest Oligocene (43.5-32.3 Ma) ages reflect renewed tectonic activity along the İzmir-Ankara. (iii) Late Oligocene- Early Miocene ages reflect the onset and development of the northern Aegean extension. The consistency of AFT ages, both north and south of the tectonic contact between the İstanbul and Sakarya terranes, suggest that such terranes were amalgamated in pre-Cenozoic times. (3) Fission-track analysis was also applied to rock samples from the Marmara region, in an attempt to constrain the inception and development of the North Anatolian Fault system in the region. The results agree with those from the central Pontides. The youngest AFT ages (Late Oligocene - early Miocene) were recorded in the western portion of the Marmara Sea region and reflect the onset and development of northern Aegean extension. Fission-track data from the eastern Marmara Sea region indicate rapid Early Eocene exhumation induced by the development of the İzmir-Ankara orogenic wedge. Thermochronological data along the trace of the Ganos Fault –a segment of the North Anatolian Fault system- indicate the presence of a tectonic discontinuity active by Late Oligocene time, i.e. well before the arrival of the North Anatolian Fault system in the area. The integration of thermochronologic data with preexisting structural data point to the existence of a system of major E-W-trending structural discontinuities active at least from the Late Oligocene. In the Early Pliocene, inception of the present-day North Anatolian Fault system in the Marmara region occurred by reactivation of these older tectonic structures. 


Evolution and historical biogeography of the Eastern Atlantic skates

In my thesis, I tested the hypothesis that the diversification of the Eastern Atlantic skate faunas arose through vicariance rather than dispersal, using combined approach of molecular phylogeny reconstruction and zoogeography (namely historical biogeography). This analyses have been carried out independently on four Rajidae genera belonging to two different tribes: Rajini (Raja and Dipturus) and Amblyrajini (Rajella and Leucoraja). These taxa were selected because they displayed high species diversity and richness of endemic species in the Eastern Atlantic and Mediterranean. The verification of this hypothesis was carried out by reconstructing the best phylogenetic relationships among four genera and 26 species (including several endemism) based on mtDNA and nuDNA gene variation and several statistical approaches. Divergence times of taxa have been estimated based on molecular clock and fossil calibration to explain evolutionary patterns in the context of geological framework. Main issues are (i) the evidence that Eastern Atlantic skate evolution and displacement of species diversity occurred from pulsed geographical speciation (i.e. repeated series of parallel and independent speciation events) started in the Late Eocene-Early Miocene and they have occurred prevalently during Miocene; (ii) such relatively ancient origin of diversification has been allowed the sympatric displacement and evolution of several congeneric taxa likely because they have accumulated huge differences in the genomic and physiological/behavioural phenotypic traits; (iii) recently diverged sister species and taxa showed allopatric or parapatric evolution by the presence of oceanographic or hydrogeographical barriers which likely prevent large mixing between parapatric sister species.