Evolutionary changes in the expression pattern of a developmentally essential gene in three Drosophila species.

The hypothesis that morphological evolution may largely result from changes in gene regulation rather than gene structure has been difficult to test. Morphological differences among insects are often apparent in the cuticle structures produced. The dopa decarboxylase (Ddc) and alpha-methyldopa hypersensitive (amd) genes arose from an ancient gene duplication. In Drosophila, they have evolved nonoverlapping functions, including the production of distinct types of cuticle, and for Ddc, the production of the neurotransmitters, dopamine and serotonin. The amd gene is particularly active in the production of specialized flexible cuticles in the developing embryo. We have compared the pattern of amd expression in three Drosophila species. Several regions of expression conserved in all three species but, surprisingly, a unique domain of expression is found in Drosophila simulans that does occur in the closely related (2-5 million years) Drosophila melanogaster or in the more remote (40-50 million years) Drosophila virilis. The "sudden" appearance of a completely new and robust domain of expression provides a glimpse of evolutionary variation resulting from changes in regulation of structural gene expression.

Diversificação de espécies e da morfologia em serpentes da família Viperidae: padrões e processos

A diversidade de espécies e fenotípica pode variar consideravelmente entre grupos taxonômicos e ao longo do tempo em uma mesma linhagem. O estudo de tais variações tornou-se um dos principais objetivos da biologia evolutiva fornecendo informações importantes a respeito dos possíveis mecanismos que regulam a biodiversidade. Dessa forma, o objetivo geral da presente tese foi investigar os padrões da diversificação de espécies e da morfologia em um grupo cosmopolita de serpentes, a família Viperidae, e os potenciais processos subjacentes. Primeiramente, (1) reconstruímos as relações filogenéticas e estimamos os tempos de divergência entre as linhagens da família Viperidae utilizando uma abordagem Bayesiana. (2) Aplicando um método recentemente desenvolvido (BAMM), exploramos como as taxas de especiação e extinção variaram ao longo da radiação do grupo inferindo os possíveis processos reguladores. Por fim, (3) analisamos se a evolução do tamanho do corpo e as taxas de especiação variam nos diferentes habitats ocupados pelos viperídeos (terrestres vs arborícola). Nesta tese geramos a filogenia molecular de viperídeos mais completa até o momento utilizando sequências para 11 genes mitocondriais e nucleares abrangendo 79% das espécies viventes (264 terminais) e todos com exceção de um gênero. De maneira geral, foi possível obter relações filogenéticas robustas para o grupo com a maioria dos gêneros sendo monofilética. Os tempos de divergência obtidos indicam que os viperídeos começaram a diversificar em meados do Paleoceno tardio/meio do Eoceno inferindo idades um pouco mais tardias que o encontrado em estudos anteriores. Durante a radiação do grupo, um aumento nas taxas de especiação parece ter ocorrido durante a diversificação dos crotalíneos (pit vipers) em decorrência não só da evolução das fossetas loreais mas também como resultado de mudanças geológicas e climáticas na Ásia e da invasão do novo mundo. Após este rápido aumento inicial, as taxas de especiação desaceleraram em direção ao presente. Por fim, os resultados aqui apresentados indicam que apesar dos habitats arborícolas limitarem a evolução morfológica nos viperídeos, a evolução da arborealidade parece não afetar as taxas de especiação que permanecem similares entre linhagens arborícolas e terrestres. Isto sugere dois cenários: (1) a especiação acontece de forma independente das mudanças morfológicas nos viperídeos; ou (2) o isolamento geográfico seria um mecanismo importante na diversificação de linhagens arborícolas contrabalançando decréscimos nas oportunidades de especiação possivelmente relacionados às pressões seletivas impostas pelo ambiente arborícola. A presente tese contribui para entendermos mais sobre como evoluíram os viperídeos ao longo dos seus ∼50 milhões de anos. Além de propor cenários e hipóteses a serem futuramente explorados com os viperídeos, elaboramos uma discussão ampla e conceitual a respeito dos possíveis mecanismos por trás da diversificação de espécies e da morfologia que poderiam também ser contemplados para outros grupos de organismos. Portanto, a presente tese contribui não só para entendermos os mecanismos que geram e mantém a diversidade de serpentes, mas também para enriquecer a discussão dos mecanismos que geram e mantém a biodiversidade como um todo

Integração morfológica e modularidade em crânios das espécies do grupo Rhinella granulosa

Os conceitos e métodos provindos das teorias de integração morfológica e de genética quantitativa formam o arcabouço teórico para o estudo da evolução de estruturas complexas, compostas de múltiplos caracteres que interagem entre si. Nesse trabalho, utilizamos o crânio como modelo de estrutura complexa e estudamos sua diversificação nas espécies de sapo do grupo Rhinella granulosa. As perguntas do trabalho foram: (1) A organização da (co)variação é similar entre as espécies?; (2) A organização da (co)variação é modular nas espécies, conforme expectativas baseadas em desenvolvimento ou função?; (3) Fatores externos, como filogenia e clima, estruturam a similaridade no padrão de covariação entre as espécies?; (4) A diversificação da morfologia média do crânio se deu por deriva ou seleção natural?; (5) A divergência na morfologia média do crânio está associada à variação climática entre as espécies?; e finalmente (6) Restrições evolutivas atuaram na divergência entre as espécies? Os espécimes foram escaneados e validamos o uso de imagens 3D para a mensuração de 21 distâncias lineares. Os crânios das espécies foram representados como matrizes fenotípicas (P) de covariância e de correlação entre as distâncias. A similaridade entre as P das espécies é alta. As P de todas as espécies se conformam a um padrão modular compatível com interações funcionais entre ossos. As diferenças entre as P concentram-se no rostro e são associadas a diferenças no clima entre as espécies. Detectamos sinal de seleção natural nos nós mais basais da filogenia e variação local no crânio está associada à variação na sazonalidade da chuva entre as espécies. Restrições evolutivas atuaram na diversificação do crânio das espécies, defletindo as respostas evolutivas para tamanho. Concluímos que tanto seleção estabilizadora e direcional, conectadas à variação climática, quanto restrições evolutivas atuaram na diversificação do crânio das espécies

La Plataforma Triásica Alpujárride (Zonas Internas de la Cordillera Bética, España)

A partir de estudios estratigráficos, centrados en la sucesión triásica alpujárride de la Sierra de Gádor (Almería, Cordillera Bética) y su correlación con otros sectores, proponemos una columna estratigráfica sintética para la Plataforma Triásica Alpujárride. Esta columna estaría dividida en dos formaciones: una metasiliciclástica basal (subdividida en tres miembros: inferior, intermedio y superior) y una metacarbonática suprayacente (subdivididida en seis miembros, numerados del 1 al 6 de muro a techo). Esta plataforma habría sufrido una evolución morfológica en tres fases: Fase 1 de rampa homoclinal (Anisiense-Ladiniense); Fase 2 de plataforma fallada (Ladiniense-Carniense); y Fase 3 de plataforma orlada (Noriense). Además, a partir del análisis sedimentario de las asociaciones de facies reconocidas en la sucesión triásica de la Sierra de Gádor proponemos su división en cuatro secuencias deposicionales para el intervalo comprendido entre el Anisiense y el Noriense. La primera secuencia comprendería los miembros inferior e intermedio de la formación metasiliciclástica; la segunda secuencia deposicional incluiría el miembro superior de la formación metasiliciclástica y los miembros 1 y 2 de la formación metacarbonática; la tercera secuencia constaría de los miembros 3 y 4, mientras que la cuarta secuencia estaría constituida por los miembros 5 y 6.

Raw-data to morphometric investigations about the Neogene planktonic foraminifer Globorotalia menardii and related forms from ODP Hole 154-925B (Céara Rise, western tropical Atlantic)

Evolutionary prospection is the study of morphological evolution and speciation in calcareous plankton from selected time-slices and key sites in the world oceans. In this context, the Neogene menardiform globorotalids serve as study objects for morphological speciation in planktic foraminifera. A downcore investigation of test morphology of the lineage of G. menardii-limbata-multicamerata during the past 8 million years was carried out in the western tropical Atlantic ODP Hole 925B. A total of 4669 specimens were measured and analyzed from 38 stratigraphic levels and compared to previous studies from DSDP Sites 502 and 503. Collection of digital images and morphometric measurements from digitized outlines were achieved using a microfossil orientation and imaging robot called AMOR and software, which was especially developed for this purpose. Most attention was given to the evolution of spiral height versus axial length of tests in keel view, but other parameters were investigated as well. The variability of morphological parameters in G. menardii, G. limbata, and G. multicamerata through time are visualized by volume density diagrams. At Hole 925B results show gradual test size increase in G. menardii until about 3.2 Ma. The combination of taxonomic determination in the light microscope with morphometric investigations shows strong morphological overlap and evolutionary continuity from ancestral to extant G. menardii (4-6 chambers in the final whorl) to the descendent but extinct G. limbata (seven chambers in the final whorl) and to G. multicamerata (>=8 chambers in the final whorl). In the morphospace defined by spiral height (dX) and axial length (dY) Globorotalia limbata and G. multicamerata strongly overlap with G. menardii. Distinction of G. limbata from G. menardii is only possible by slight differences in the number of chambers of the final whorl, nuances in spiral convexity, upper keel angles, radii of osculating circles, or by differences in reflectance of their tests. Globorotalia multicamerata can be distinguished from the other two forms by more than eight chambers in the final whorl. It appeared as two stratigraphically separate clusters during the Pliocene. Between 2.88 and 2.3 Ma G. menardii was severely restricted in size and abundance. Thereafter, it showed a rapid and prominent expansion of the upper test size extremes between 2.3 and 1.95 Ma persisting until present. The size-frequency distributions at Hole 925B are surprisingly similar to trends of menardiform globorotalids from Caribbean DSDP Site 502. There, the observations were explained as an adaptation to changes in the upper water column due to the emergence of the Isthmus of Panama. In light of more recent paleontological and geological investigations about the completion of the permanent land connection between North and South America since about 3 Ma the present study gives reason to suspect the sudden test size increase of G. menardii to reflect immigration of extra-large G. menardii from the Indian Ocean or the Pacific. It is hypothesized that during the Late Pliocene dispersal of large G. menardii into the southern to tropical Atlantic occurred during an intermittent episode of intense Agulhas Current leakage around the Cape of Good Hope and from there via warm eddy transport to the tropical Atlantic (Agulhas dispersal hypothesis).

Age estimations of calcareous nannofossil biohorizons of the middle Paleocene to early Eocene at ODP Site 208-1262 (Table 1, Appendix B)

Over the last several decades debates on the 'tempo and mode' of evolution have centered on the question whether morphological evolution preferentially occurs gradually or punctuated, i.e., with long periods of stasis alternating with short periods of rapid morphological change and generation of new species. Another major debate is focused on the question whether long-term evolution is driven by, or at least strongly influenced by changes in the environment, or by interaction with other life forms. Microfossils offer a unique opportunity to obtain the large datasets as well as the precision in dating of subsequent samples to study both these questions.We present high-resolution analyses of selected calcareous nannofossils from the deep-sea section recovered at ODP Site 1262 (Leg 208) in the South-eastern Atlantic. The studied section encompasses nannofossil Zones NP4-NP12 (equivalent to CP3-CP10) and Chrons C27r-C24n. We document more than 70 biohorizons occurring over an about 10 Myr time interval, (~62.5 Ma to ~52.5 Ma), and discuss their reliability and reproducibility with respect to previous data, thus providing an improved biostratigraphic framework, which we relate to magnetostratigraphic information, and present for two possible options of a new Paleocene stratigraphic framework based on cyclostratigraphy. This new framework enabled us to tentatively reconstruct steps in the evolution of early Paleogene calcareous nannoplankton through documentation of transitional morphotypes between genera and/or species and of the phylogenetic relations between the genera Fasciculithus, Heliolithus, Discoasteroides and Discoaster, as well as between Rhomboaster and Tribrachiatus. The exceptional record provided by the continuous, composite sequence recovered at Walvis Ridge allows us to describe the mode of evolution among calcareous nannoplankton: new genera and/or new species usually originated through branching of lineages via gradual, but relatively rapid, morphological transitions, as documented by the presence of intermediate forms between the end-member ancestral and descendant forms. Significant modifications in the calcareous nannofossil assemblages are often "related" to significant changes in environmental conditions, but the appearance of structural innovations and radiations within a single genus also occurred during "stable" environmental conditions. These lines of evidence suggest that nannoplankton evolution is not always directly triggered by stressed environmental conditions but could be also driven by endogenous biotic control.

Phenotypic divergence along lines of genetic variance

Natural populations inhabiting the same environment often independently evolve the same phenotype. Is this replicated evolution a result of genetic constraints imposed by patterns of genetic covariation? We looked for associations between directions of morphological divergence and the orientation of the genetic variance-covariance matrix (G) by using an experimental system of morphological evolution in two allopatric nonsister species of rainbow fish. Replicate populations of both Melanotaenia eachamensis and Melanotaenia duboulayi have independently adapted to lake versus stream hydrodynamic environments. The major axis of divergence (z) among all eight study populations was closely associated with the direction of greatest genetic variance (g(max)), suggesting directional genetic constraint on evolution. However, the direction of hydrodynamic adaptation was strongly associated with vectors of G describing relatively small proportions of the total genetic variance, and was only weakly associated with g(max). In contrast, divergence between replicate populations within each habitat was approximately proportional to the level of genetic variance, a result consistent with theoretical predictions for neutral phenotypic divergence. Divergence between the two species was also primarily along major eigenvectors of G. Our results therefore suggest that hydrodynamic adaptation in rainbow fish was not directionally constrained by the dominant eigenvector of G. Without partitioning divergence as a consequence of the adaptation of interest (here, hydrodynamic adaptation) from divergence due to other processes, empirical studies are likely to overestimate the potential for the major eigenvectors of G to directionally constrain adaptive evolution.

Swash-aquifer interaction in the vicinity of the water table exit point on a sandy beach

The coupling of sandy beach aquifers with the swash zone in the vicinity of the water table exit point is investigated through simultaneous measurements of the instantaneous shoreline (swash front) location, pore pressures and the water table exit point. The field observations reveal new insights into swash-aquifer coupling not previously gleaned from measurements of pore pressure only. In particular, for the case where the exit point is seaward of the observation point, the pore pressure response is correlated with the distance between the exit point and the shoreline in that when the distance is large the rate of pressure drop is fast and when the distance is small the rate decreases. The observations expose limitations in a simple model describing exit point dynamics which is based only on the force balance on a particle of water at the sand surface and neglects subsurface pressures. A new modified form of the model is shown to significantly improve the model-data comparison through a parameterization of the effects of capillarity into the aquifer storage coefficient. The model enables sufficiently accurate predictions of the exit point to determine when the swash uprush propagates over a saturated or a partially saturated sand surface, potentially an important factor in the morphological evolution of the beach face. Observations of the shoreward propagation of the swash-induced pore pressure waves ahead of the runup limit shows that the magnitude of the pressure fluctuation decays exponentially and that there is a linear increase in time lags, behavior similar to that of tidally induced water table waves. The location of the exit point and the intermittency of wave runup events is also shown to be significant in terms of the shore-normal energy distribution. Seaward of the mean exit point location, peak energies are small because of the saturated sand surface within the seepage face acting as a "rigid lid'' and limiting pressure fluctuations. Landward of the mean exit point the peak energies grow before decreasing landward of the maximum shoreline position.

Um olhar sobre o castelo de Noudar: do castelo medieval à ruína: Proposta de valorização do lugar

O Castelo de Noudar está implantado num íngreme promontório da herdade da Coitadinha, a Noroeste de Barrancos, integrando o atual Parque da Natureza de Noudar. Construído a partir de 1303, foi alvo de diversas transformações ao longo do tempo, para responder às exigências militares de cada época, e chega aos nossos dias num avançado estado de decadência física. A investigação desenvolvida assenta, em grande parte, no desenho: de Noudar a uma escala territorial, como elemento que participa numa rede fortificada complexa; da evolução morfológica e funcional, que nos permite ter noção das várias configurações do castelo ao longo dos séculos; num desenho propositivo que desenvolve uma estratégia para valorização do lugar e do seu património material e imaterial. Hoje é fundamental compreender como as valências do Parque e da fortaleza se podem cruzar? Este trabalho pretende refletir sobre possibilidades de valorização para contrariar a degradação em que a fortaleza se encontra; ABSTRACT: The Noudar Castle is located in a steep promontory in the herdade da Coitadinha, Northeast from Barrancos, part of the Natural Park of Noudar. Its construction began in 1303, and underwent several transformations throughout its existence, adapting to the military advancements thru out the ages. Today, it shows an advanced state of decay. The investigation is focused on the design of Noudar on a territorial scale, taking part in a complex network of fortified systems; its functional and morphological evolution, which allows us to comprehend the configurations of the castle throughout the centuries; and on a proposed design that develops a strategy to enhance and bring value to this place and its material and non-material heritage. Today it's crucial to comprehend how can the values of the Park and fortress intertwine. This thesis reflects on the possibilities of valuing this site, counter acting the decay which has fallen upon it.

A arquitectura do mosteiro cartusiano de Santa Maria Scala Coeli: do processo de entrada

A importância da encenação de uma transição entre o mundo secular e a vida em clausura, no contexto da arquitectura cartusiana, é amplamente demonstrada através dos percursos de entrada dos conjuntos mais relevantes, bem como em muitos daqueles representados nas gravuras presentes no livro Maisons de l’Ordre des Chartreux – Vues et Notices, uma obra do início do século XX que inclui gravuras da quase totalidade dos 282 mosteiros cartusianos distribuídos por quase toda a Europa. Na maioria das cartuxas podem observarse transições indirectas e prolongadas, capazes de tardar o passo e dar significado à passagem para a clausura – características espaciais que São Bruno, fundador da casa cartusiana, considerava primordiais para a implantação do seu eremitério. A partir de uma cuidada análise dessa transição, nas cartuxas em geral, e da sua evolução morfológica, na cartuxa de Évora em particular – estimulada pela descoberta de dados aqui analisados e interpretados pela primeira vez – apresenta-se, em continuidade com a sua história e tendo como premissa a consolidação do processo de entrada, uma proposta de redefinição da estrutura monacal de Santa Maria Scala Coeli. Esta estrutura originária do século XVI encontra-se incompleta e degradada, em particular na ala sul onde se encontra o seu principal acesso, sendo fundamental o completamento desta área para o cabal reconhecimento da unidade do conjunto e da intenção na qual se traduz a sua génese. De resto, a estratégia de investigação e de projecto descrita nesta dissertação pode ser encarada como um processo plausível de ser utilizado no caso de futuras intervenções noutras estruturas cartusianas, contribuindo assim para a recuperação e preservação do conceito fundador da identidade da arquitectura cartusiana que, no decorrer da expansão da ordem, se foi perdendo; The Architecture of the Carthusian Monastery of Santa Maria Scala Coeli: About the process of entrance. ABSTRACT: The importance of staging a transition between the secular world and life cloistered in the context of Carthusian architecture is amply demonstrated through the input paths of the most important collections, as well as many of those represented in the pictures in the book Maisons de l’Ordre des Chartreux – Vues et Notices, one of the early twentieth century work that includes prints of almost all of the 282 Carthusians monasteries spread across almost all of Europe. In most of Carthusian the transitions that can be observed are indirect and long, able to delay the step and give meaning to the passage to the life cloistered - spatial characteristics that St. Bruno, founder of the Carthusian Order, considered essential for the implementation of his hermitage. From a careful analysis of this transition, looking through the Carthusian in general, and its morphological evolution, in the Charterhouse of Évora in particular, encouraged by the data discovery here analyzed and interpreted for the first time - it is presented in continuity with its history and having premised on the consolidation of the entry process, a proposal for a redefinition of the monastic structure of Santa Maria Scala Coeli. This original structure of the sixteenth century is incomplete and degraded, particularly in the south wing where it is the main access, and where is fundamental the completion of this area to the full recognition of the unity of the whole and to understand the intent in which translates its genesis. Moreover, the research strategy and project described in this paper can be seen as a plausible case to be used in case of future interventions in other cartusianas structures, thus contributing to the recovery and preservation of the founder of the identity concept of the Carthusian architecture that, during the expansion of the order, was missing.

Multiscale Modeling and Simulation of Stepped Crystal Surfaces

A primary goal of this dissertation is to understand the links between mathematical models that describe crystal surfaces at three fundamental length scales: The scale of individual atoms, the scale of collections of atoms forming crystal defects, and macroscopic scale. Characterizing connections between different classes of models is a critical task for gaining insight into the physics they describe, a long-standing objective in applied analysis, and also highly relevant in engineering applications. The key concept I use in each problem addressed in this thesis is coarse graining, which is a strategy for connecting fine representations or models with coarser representations. Often this idea is invoked to reduce a large discrete system to an appropriate continuum description, e.g. individual particles are represented by a continuous density. While there is no general theory of coarse graining, one closely related mathematical approach is asymptotic analysis, i.e. the description of limiting behavior as some parameter becomes very large or very small. In the case of crystalline solids, it is natural to consider cases where the number of particles is large or where the lattice spacing is small. Limits such as these often make explicit the nature of links between models capturing different scales, and, once established, provide a means of improving our understanding, or the models themselves. Finding appropriate variables whose limits illustrate the important connections between models is no easy task, however. This is one area where computer simulation is extremely helpful, as it allows us to see the results of complex dynamics and gather clues regarding the roles of different physical quantities. On the other hand, connections between models enable the development of novel multiscale computational schemes, so understanding can assist computation and vice versa. Some of these ideas are demonstrated in this thesis. The important outcomes of this thesis include: (1) a systematic derivation of the step-flow model of Burton, Cabrera, and Frank, with corrections, from an atomistic solid-on-solid-type models in 1+1 dimensions; (2) the inclusion of an atomistically motivated transport mechanism in an island dynamics model allowing for a more detailed account of mound evolution; and (3) the development of a hybrid discrete-continuum scheme for simulating the relaxation of a faceted crystal mound. Central to all of these modeling and simulation efforts is the presence of steps composed of individual layers of atoms on vicinal crystal surfaces. Consequently, a recurring theme in this research is the observation that mesoscale defects play a crucial role in crystal morphological evolution.

A Casa Grande de Romarigães. Um contributo histórico e arquitectónico

A Casa Grande de Romarigães terá sido erguida em 1700 em Romarigães, no concelho de Paredes de Coura, em torno da construção da capela de Nossa Senhora do Amparo, conformando a Quinta do Amparo. Terá servido de cenário literário a uma das mais simbólicas obras do escritor Aquilino Ribeiro – A Casa Grande de Romarigães, escrita entre 1950 e 1957, ano da sua publicação. A estrutura arquitectónica foi classificada como Imóvel de Interesse Público em 1986, atribuição que, apesar de demonstrar a preocupação de proteger e conservar este legado histórico e arquitectónico, ainda não derivou em nenhum estudo rigoroso sobre a sua evolução morfológica – instrumento fundamental para a coerência de futuras intervenções neste lugar. Com base nas constantes descrições na obra de Aquilino Ribeiro, e do seu confronto com a documentação histórica entretanto encontrada (escrita e gráfica, em grande parte inédita), este trabalho de investigação propõe o desenho da evolução morfológica da casa ao longo dos tempos, bem como a enunciação da influência deste solar para a organização social, cultural e económica da aldeia de Romarigães. Procura, enfim, e num sentido mais lato, indagar sobre a utilidade de um documento literário para o lançamento de hipóteses no contexto de uma investigação arquitectónica, consubstanciadas pela posterior comprovação através de fontes primárias; ABSTRACT: A Casa Grande de Romarigães (The Great House of Romarigães) was erected in early XVIII century, in Romarigães, Paredes de Coura, nearby the chapel of Nossa Senhora do Amparo, the chosen name for the villa – Quinta do Amparo. This house and villa has been the literary scene of one of the most remarkable works of the writer Aquilino Ribeiro (1885C1963) – A Casa Grande Romarigães, written between 1950 and 1957, the year of its publication. In 1986, the architectural structure is classified as cultural heritage by the Portuguese state but, despite the meaningful idea of preservation and protection of the historical legacy and architecture, there has not been an initiative to study its morphological evolution and architectural relevance – key tool for future and consistent interventions in such rich and particular scenario. Based on descriptions that can be found in the work of Aquilino Ribeiro, and supported with historical documentation research (both written and graphic, mostly unpublished), this work suggests the design of the morphological evolution of the house over the years, together with the construction of its influence on social, cultural and economic organization of the village of Romarigães. In a broader sense, this work intends to inquire about the usefulness of a literary document for launching assumptions in the context of an architectural research, substantiated by further evidence through primary sources.

Morphological diversification through the evolution of developmental hierarchies

Changes in development impact the final form of organisms and compose the natural variation that is the raw material for evolution. Development is hierarchically structured in progressive series of cell fate determination and differentiation. How does variation in different stages of development contribute to morphological diversification?

The evolution of jumping performance in anurans: morphological correlates and ecological implications

We investigated the evolution of anuran locomotor performance and its morphological correlates as a function of habitat use and lifestyles. We reanalysed a subset of the data reported by Zug (Smithson. Contrib. Zool. 1978; 276: 1-31) employing phylogenetically explicit statistical methods (n = 56 species), and assembled morphological data on the ratio between hind-limb length and snout-vent length (SVL) from the literature and museum specimens for a large subgroup of the species from the original paper (n = 43 species). Analyses using independent contrasts revealed that classifying anurans into terrestrial, semi-aquatic, and arboreal categories cannot distinguish between the effects of phylogeny and ecological diversification in anuran locomotor performance. However, a more refined classification subdividing terrestrial species into `fossorials` and `non-fossorials`, and arboreal species into `open canopy`, `low canopy` and `high canopy`, suggests that part of the variation in locomotor performance and in hind-limb morphology can be attributed to ecological diversification. In particular, fossorial species had significantly lower jumping performances and shorter hind limbs than other species after controlling for SVL, illustrating how the trade-off between burrowing efficiency and jumping performance has resulted in morphological specialization in this group.

The Evolution of Modularity in the Mammalian Skull I: Morphological Integration Patterns and Magnitudes

Morphological integration refers to the modular structuring of inter-trait relationships in an organism, which could bias the direction and rate of morphological change, either constraining or facilitating evolution along certain dimensions of the morphospace. Therefore, the description of patterns and magnitudes of morphological integration and the analysis of their evolutionary consequences are central to understand the evolution of complex traits. Here we analyze morphological integration in the skull of several mammalian orders, addressing the following questions: are there common patterns of inter-trait relationships? Are these patterns compatible with hypotheses based on shared development and function? Do morphological integration patterns and magnitudes vary in the same way across groups? We digitized more than 3,500 specimens spanning 15 mammalian orders, estimated the correspondent pooled within-group correlation and variance/covariance matrices for 35 skull traits and compared those matrices among the orders. We also compared observed patterns of integration to theoretical expectations based on common development and function. Our results point to a largely shared pattern of inter-trait correlations, implying that mammalian skull diversity has been produced upon a common covariance structure that remained similar for at least 65 million years. Comparisons with a rodent genetic variance/covariance matrix suggest that this broad similarity extends also to the genetic factors underlying phenotypic variation. In contrast to the relative constancy of inter-trait correlation/covariance patterns, magnitudes varied markedly across groups. Several morphological modules hypothesized from shared development and function were detected in the mammalian taxa studied. Our data provide evidence that mammalian skull evolution can be viewed as a history of inter-module parcellation, with the modules themselves being more clearly marked in those lineages with lower overall magnitude of integration. The implication of these findings is that the main evolutionary trend in the mammalian skull was one of decreasing the constraints to evolution by promoting a more modular architecture.