Extension and patterning of the vertebrate body

Dissertation presented to obrain the Ph.D degree in Biology. Developmental Biology

On the function of the floccular complex of the vertebrate cerebellum: implications in paleoneuroanatomy

The cerebellum floccular complex lobes (FCLs) are housed in the FCL fossa of the periotic complex. There is experimental evidence indicating that the FCLs integrate visual and vestibular information, responsible for the vestibulo-ocular reflex, vestibulo-collic reflex, smooth pursuit and gaze holding. Thus, the behavior of extinct animals has been correlated with FCLs dimension in multiple paleoneuroanatomy studies. Here I analyzed braincase endocasts of a representative sample of Mammalia (48 species) and Aves (59 species) rendered using tomography and image segmentation and tested statistical correlations between the floccular complex volume, ecological and behavioral traits to assess various previously formulated paleobiological speculations. My results demonstrate: 1) there is no significant correlation between relative FCL volume and body mass; 2) there is no significant correlation between relative FCL and optic lobes size in birds; 3) average relative FCL size is larger in diurnal than in nocturnal birds but there is no statistically significant difference in mammals; 4) feeding strategies are related with different FCL size patterns in birds, but not in mammals; 5) locomotion type is not related with relative FCL size in mammals; 6) agility is not significantly correlated with FCL size in mammals. I conclude that, despite the apparent relation between FCL size and ecology in birds, the cerebellum of tetrapods is a highly plastic structure and may be adapted to control different functions across different taxonomic levels. For example, the european mole (Talpa europaea) which is fossorial and practically blind, has a FCL fossae relative size larger than those of bats, which are highly maneuverable. Therefore, variation in FCL size may be better explained by a combination of multiple factors with relation to anatomical and phylogenetic evolutionary constraints.

Notas sobre depósitos de Taveiro estratigrafia, paleontologia, idade, paleoecologia

This paper reports some research work that has been done to support Geological Survey's field work for the 1:50.000 Carta Geológica de Portugal, (sheets 19-C Figueira da Foz and 19-D Coimbra-Lousã). Its main purpose was to establish the age of some continental formations. At Cerâmica do Mondego, Ld.ª near Taveiro, two series were observed. The lower one is mainly pelitic, montmorillonite being predominant. It also includes some sandy beds and channel deposits with high energy sediments (conglomerate with limestone pebbles). The upper series lies unconformably upon the former, and there is a neat discontinuity surface between the two. It mainly consists of sands, kaolinite being the most abundant of the clay minerals. This seems to indicate an intensive weathering, an acid, well drained environment and transportation by quite high energy running waters. No fossils were recorded. Preliminary paleontological results are presented, along with some data concerning other localities (Aveiro, etc.). Fossils found in the lower series are: gastropoda (Bulimus gaudryi, TV. 15 bed), several vertebrates (TV. 18), fishes (TV. 19?) and plants (TV. 19-TV. 24). Vertebrata belong to the same fauna as that from Vizo, Aveiro, etc. The presence of mammals is most important as only a single tooth was previously Know in Europe (Southern France) in Late Cretaceous formations. Elsewhere there are some mammalian remains in Peru besides the rich assemblages found in the USA and Mongolia. Plants are representative of the «Debeya flora» well known at several localities in Beira Litoral province, in «Buçaco sandstones», and in Lisbon's «Basaltic Complex». The most important stratigraphical conclusion is that the lower series is Upper Campanian and/or Maastrichtian in age, and not Tertiary as sometimes it has been considered. As at Aveiro, «Bebeya flora» occurs in-beds somewhat higher than those with the Aveiro-Vizo-Taveiro vertebrate fauna. Correlation with other «Debeya flora» localities are now more clear. Data concerning Taveiro lower series, in the whole, point out to a rather warm (and moist?) environment in an occasionally (seasonally?) flooded region.

Les Hipparion du Portugal

This paper is concerned with Hipparion from Ribatejo, Portugal, and with the stratigraphy of the Neogene series of this region. The first two chapters are an introduction and an historical review. Paleontological study includes both a revision of the specimens accounted by ROMAN (1907) and the description of new material. Two forms were recognized, an early H. cf. primigenium, lower Vallesian in age, NM 9 mammal unit (from Archino, Vila Nova da Rainha, Aveiras de Cima), and a more advanced H. primigenium cf. melendezi. Upper Vallesian, NM 10 (possibly lowermost Turolian, NM 11) (at Azambujeira and Marmeleira). A synthesis of Middle and Upper Miocene from Ribatejo is also presented. Levels with H. p. cf. melendezi are somewhat older than «Upper Pontian», as it was previously acknowledged, they attain at the best the lowermost Turolian (approximately corresponding to «Upper Pontian»). Even higher levels may be Turolian in age, though they are not yet accurately dated. Almost all the localities are shown (tableau 11) according to its stratigraphical position; age, correspondance to mammal units from NM 5 to NM 10 (and may be also from NM 11 to NM 12), and correlation with marine formations near Lisbon are also taken in account. The stratigraphical position of localities such as Póvoa de Santarém, Quinta do Marmelal, Pero Filho, Azambujeira (lower levels), and Fonte do Pinheiro was revised; the stratigraphical position of Marmeleira was ascertained. The localities so far known correspond to NM 5 (?), NM 6, NM 8, NM 9, NM 10 and possibly to NM II and NM 12. A new interpretation (M. T. ANTUNES) of localities with oysters from Ribatejo allows a better correlation with vertebrate localities. Relationships with Serravallian transgression seem well established. Only two localities, Vila Nova da Rainha and Foz do Alviela, may possibly be correlated to V-b division of Lisbon (Langhian) with «Hispanotherium fauna». All the other localities are younger than Serravallian oyster beds. Undirect correlation shows that NM 6 localities are somewhat younger than the apogee ef Serravallian transgression (corresponding approximately to Blow's N 11 to N 13 zones based on planctonic foraminifera).

Notas sobre depósitos de Taveiro estratigrafia, paleontologia, idade, paleoecologia

This paper reports some research work that has been done to support Geological Survey's field work for the 1:50.000 Carta Geológica de Portugal, (sheets 19-C Figueira da Foz and 19-D Coimbra-Lousã). Its main purpose was to establish the age of some continental formations. At Cerâmica do Mondego, Lda. near Taveiro, two series were observed. The lower one is mainly pelitic, montmorillonite being predominant. It also includes some sandy beds and channel deposits with high energy sediments (conglomerate with limestone pebbles). The upper series lies unconformably upon the former, and there is a neat discontinuity surface between the two. It mainly consists of sands, kaolinite being the most abundant of the clay minerals. This seems to indicate an intensive weathering, an acid, well drained environment and transportation by quite high energy running waters. No fossils were recorded. Preliminary paleontological results are presented, along with some data concerning other localities (Aveiro, etc). Fossils found in the lower series are: gastropoda (Bulimus gaudryi, TV. 15 bed), several vertebrates (TV. 18), fishes (TV. 19?) and plants (TV. 19-TV. 24). Vertebrata belong to the same fauna as that from Vizo, Aveiro, etc. The presence of mammals is most important as only a single tooth was previously know in Europe (Southern France) in Late Cretaceous formations. Elsewhere there are some mammalian remains in Peru besides the rich assemblages found in the USA and Mongolia. Plants are representative of the «Debeya flora» well known at several localities in Beira Litoral province, in «Buçaco sandstones», and in Lisbon's «Basaltic Complex». The most important stratigraphical conclusion is that the lower series is Upper Campanian and/or Maastrichtian in age, and not Tertiary as sometimes it has been considered. As at Aveiro, «Bebeya flora» occurs in-beds somewhat higher than those with the Aveiro-Vizo-Taveiro vertebrate fauna. Correlation with other «Debeya flora» localities are now more clear. Data concerning Taveiro lower series, in the whole, point out to a rather warm (and moist?) environment in an occasionally (seasonally?) flooded region.

Les Hipparion du Portugal

This paper is concerned with Hipparion from Ribatejo, Portugal, and with the stratigraphy of the Neogene series of this region. The first two chapters are an introduction and an historical review. Paleontological study includes both a revision of the specimens accounted by ROMAN (1907) and the description of new material. Two forms were recognized, an early H. cf. primigenium, lower Vallesian in age, NM 9 mammal unit (from Archino, Vila Nova da Rainha, Aveiras de Cima), and a more advanced H. primigenium cf. melendezi. Upper Vallesian, NM 10 (possibly lowermost Turolian, NM 11) (at Azambujeira and Marmeleira). A synthesis of Middle and Upper Miocene from Ribatejo is also presented. Levels with H. p. cf. melendezi are somewhat older than «Upper Pontian», as it was previously acknowledged, they attain at the best the lowermost Turolian (approximately corresponding to «Upper Pontian»). Even higher levels may be Turolian in age, though they are not yet accurately dated. Almost all the localities are shown (tableau 11) according to its stratigraphical position; age, correspondance to mammal units from NM 5 to NM 10 (and may be also from NM 11 to NM 12), and correlation with marine formations near Lisbon are also taken in account. The stratigraphical position of localities such as Póvoa de Santarém, Quinta do Marmelal, Pero Filho, Azambujeira (lower levels), and Fonte do Pinheiro was revised; the stratigraphical position of Marmeleira was ascertained. The localities so far known correspond to NM 5 (?), NM 6, NM 8, NM 9, NM 10 and possibly to NM II and NM 12. A new interpretation (M. T. ANTUNES) of localities with oysters from Ribatejo allows a better correlation with vertebrate localities. Relationships with Serravallian transgression seem well established. Only two localities, Vila Nova da Rainha and Foz do Alviela, may possibly be correlated to V-b division of Lisbon (Langhian) with «Hispanotherium fauna». All the other localities are younger than Serravallian oyster beds. Undirect correlation shows that NM 6 localities are somewhat younger than the apogee ef Serravallian transgression (corresponding approximately to Blow's N 11 to N 13 zones based on planctonic foraminifera).

Vertébrés (crocodilians, poisons) du Miocène marin de l’Algarve occidentale

Karst fillings rich in fossils from carbonate series, Miocene in age, exposed on the coast of Algarve, yielded some thousands of teeth and other vertebrate remains. A crocodilian and more than fifty species of fishes (Selacians, Teleosteans) have been identified (see tableau 1). This fauna is consistent as a whole with a Burdigalian (maybe late) and/or Langhian age. It also indicates shallow waters, warmer than at present in the same region although not strictly tropical. This fauna shows more affinities with others of the same age from mediterranean region (i.e. Baleares) than with those from Lisbon.

Hexose and pentose transport inascomycetousyeasts: an overview

FEMS Yeast Research, Vol. 9, nº 4

The Neogene of the Lower Tagus Basin (Portugal)

Revista Española de Paleontologia 19 (2), 229-242

Spectroscopic Studies and Characterization of a Novel Electron-Transfer Chain

A novel two-component enzyme system from Escherichia coli involving a flavorubredoxin (FlRd) and its reductase was studied in terms of spectroscopic, redox, and biochemical properties of its constituents. FlRd contains one FMN and one rubredoxin (Rd) center per monomer. To assess the role of the Rd domain, FlRd and a truncated form lacking the Rd domain (FlRd¢Rd), were characterized. FlRd contains 2.9 ( 0.5 iron atoms/subunit, whereas FlRd¢Rd contains 2.1 ( 0.6 iron atoms/subunit. While for FlRd one iron atom corresponds to the Rd center, the other two irons, also present in FlRd¢Rd, are most probably due to a di-iron site. Redox titrations of FlRd using EPR and visible spectroscopies allowed us to determine that the Rd site has a reduction potential of -140 ( 15 mV, whereas the FMN undergoes reduction via a red-semiquinone, at -140 ( 15 mV (Flox/Flsq) and -180 ( 15 mV (Flsq/Flred), at pH 7.6. The Rd site has the lowest potential ever reported for a Rd center, which may be correlated with specific amino acid substitutions close to both cysteine clusters. The gene adjacent to that encoding FlRd was found to code for an FAD-containing protein, (flavo)rubredoxin reductase (FlRd-reductase), which is capable of mediating electron transfer from NADH to DesulfoVibrio gigas Rd as well as to E. coli FlRd. Furthermore, electron donation was found to proceed through the Rd domain of FlRd as the Rd-truncated protein does not react with FlRd-reductase. In vitro, this pathway links NADH oxidation with dioxygen reduction. The possible function of this chain is discussed considering the presence of FlRd homologues in all known genomes of anaerobes and facultative aerobes.

Guia (Algarve), gisement de vertébrés quaternaires à caractère saisonnier

A vertebrate (Mammals, Reptiles, Amphibians) fauna From Guia (Algarve) is described. The site is the only one so far known in the «Faro-Quarteira sands». The fauna is upper Pleistocene in age (may even be younger). Its stratigraphical position shows it is younger than levels in the same unit that yielded mousterian/languedocian stone artifacts. A post Riss-Wurm age is admitted. The mammalian fauna nearly exclusively comprises small mammalremains; except for a few ones, only young individuals are represented. This unusual situation, coupled with data from the study of the Squamata and even more of the Amphibia, indicates that Guia locality corresponds to sedimentation in temporary, seasonal ponds originated in depressions in this sandy area during Winter and/or early Springtime.

Contributions to the Eocene palaeontology and stratigraphy of Beira Alta, Portugal II - New Late Eocene mammalian remnants from Coja (Portugal) and the presence of Palaeotherium magnum Cuvier

The presence of Palaeotherium magnum in the fauna from Coja is recorded. It is well in agreement with the earlier reporting to the Montmartre level from the Ludian stage. Field data as well as compatibility with the remaining taxa and the identical fossilization of all the specimens indicate that all the vertebrate fossils come from the same horizon in the lithostratigraphic unit "Arcoses de Côja".

Upper Paleocene-Early Eocene mollusks of Silveirinha (Figueira da Foz, West Central Portugal)

A collection of fossil gastropods and bivalves assembled at the Thanetian/Ypresian vertebrate site of Silveirinha (Figueira da Foz, West Central Portugal) is analysed from the point of view of systematics and palaeoecology. The diversity is scarce but the age and exceptional characteristics of the site are factors that substantiate a detailed study. The taxa identified are: Bithynia soaresi sp. nov., Gyraulus antunesi sp. nov., Chlamys sp. and Cardiiacea gen. sp. indet. The prevailing of freshwater gastropods and the occurrence of 2 fragments of marine bivalves suggest a palaeoenvironmental setting that is in conformity with interpretations already established, which are based both in sedimentologic and vertebrate data. These interpretations point out the existence of a freshwater environment opened from time to time to marine influences, resulting from a palaeoatlantic coast placed some kilometres westwards.

A new dinosaur tracksite in the Lower Cretaceous of Portugal

A new Lower Cretaceous (Aptian-Albian) dinosaur tracksite at the Olhos de Água beach is described. It is the first vertebrate fossil finding ever found in the concerned unit, and yielded 128 tracks in 17 trackways within an area of ca. 80 square metres. Three tridactyl footprint morphotypes have been recognized: - Type I ("Iguanodontipus-like") - trackways D, F, K, J and P; - Type 2 (large theropod), although larger in size, typically from a Grallator-like theropod footprint, i.e , A, B, G, H and 0 trackways; - Type 3 (medium size theropod); M is the only track of this type. There are other, poorly preserved, unidentified trackways. The theropod, swinging trackway B was produced by an animal that was limping. The theropod track M starts eastwards but drastically changes westwards, speeding up at the same time; this dinosaur decided to turn around and run in the opposite direction. This site shows three main trackway directions: to the South, to the East, and westwards. Except for the trackway 0, large theropods A, B, G and H walked southwards. Perpendicularly to the se, ornithopods, small theropods and unidentified trackmakers walked towards East (5) and West (7). The segregation oftrackmakers and directions, with large theropod traekways southwards and other dinosaurs' west or eastwards, may mean that large theropods patrolled a walkway area to an important resource, most probably water, often frequented by ornithopods and smaller theropods. There is no evidence of social behavior or gregarism: footprints' overposition shows that the large, southwards walking theropods passed on different occasions. Three trackway sequences can be established by chronologic order.

Linking the embryonic clock with temporal collinearity of Hox gene activation

Dissertação apresentada para a obtenção do Grau de Mestre em Genética Molecular e Biomedicina, pela Universidade Nova de Lisboa, Faculdade de Ciências e Tecnologia