Around 375 million years ago, the first tetrapods appeared, marking one of the most important events in vertebrate evolutionary history. The fin to limb transition saw the appearance of fingers and a weight bearing pelvic girdle. While very little research has been done on the evolution of the tetrapod pelvic girdle, a fair amount has been done on the origins of fingers but some aspects remained controversial. A combination of palaeontology, developmental biology and comparative morphology was therefore used in this thesis to better understand the fin to limb transition. The pectoral fin of Panderichthys, a sarcopterygian fish closely related to tetrapods was CT-scanned and modeled in three dimensions and its pelvic girdle and fin were examined with traditional techniques. This information from the fossil record was integrated with comparisons of the development of the Australian lungfish, Neoceratodus forsteri, our closest living fish relative and the axolotl (Ambystoma mexicanum), a salamander representing well the condition of early tetrapods. Development of bone and cartilage was studied through clearing and staining and development of skeletal muscles through immunostaining. In situ hybridizations were performed on the lungfish to study the expression of Hoxd13, associated with the formation of digits in tetrapods. This work shows that the late expression phase of Hoxd13 is present in Neoceratodus and is associated with the formation of radials. Redescription of the pectoral fin of Panderichthys reveals that distal radials are present, which, in addition to other information, lead us to conclude that digits are not novelties in tetrapods but rather have evolved from the distal radials present in the fins of all sarcopterygian fish. The earliest tetrapods lack a full set of wrist + carpals/ankle + tarsal bones. Here, we propose that this region of the limbs evolved after fingers and toes through an expansion of the region between the proximal limb bones and the digits. As for the pelvic girdle, it is very primitive in Panderichthys but comparison of its development in Neoceratodus and Ambystoma suggest that the ischium evolved through the posterior expansion of the pubis and the ilium, through an elongation of the iliac process already present in sarcopterygian fishes. The results of this thesis help to better understand the fin to limb transition and show that it is more gradual than previously believed.

<p>The present study is concerned with the evolutionary relationships among the Euasterids II, a group of angiosperms that includes the orders Apiales, Aquifoliales, Asterales, and Dipsacales together with several small, poorly known families yet unplaced as to order.</p><p>Parsimony analysis of nucleotide sequence data from the chloroplast genes <i>atp</i>B, <i>ndh</i>F and <i>rbc</i>L together with morphological data are used to construct a phylogeny of the order Asterals, which in the present sense includes 11 families and more than 26 000 species. It is argued that Rousseaceae should be expanded to include also Carpodetaceae (and thus contain four genera), and that <i>Donatia</i> should be re-merged with Stylidiaceae. The present study also strongly supports that the sister taxon to the largest plant family, Asteraceae (Compositae), is the small South American Calyceraceae. A new addition to Asterales is <i>Platyspermation</i> (formerly in Escalloniaceae).</p><p>Using the recently developed Bayesian approach to phylogenetic reconstruction, in combination with a dataset consisting of the <i>atp</i>B, <i>ndh</i>F and <i>rbc</i>L nucleotide sequences, a resolved and fairly well supported phylogeny of the Euasterids II could be reconstructed. Based on this analysis, and furthermore supported by morphological characters, the South Hemispheric family Escalloniaceae is recircumscribed to include the monogeneric families Eremosynaceae, Tribelaceae and Polyosmaceae, and in this new circumscription it includes seven genera. The formerly escalloniacean member <i>Quintinia</i>, together with the monogeneric families Paracryphiaceae and Sphenostemonaceae, is argued to consitute a monophyletic family Paracryphiaceae <i>sensu lato</i>, supported by several morphological characters.</p>

The present study is concerned with the evolutionary relationships among the Euasterids II, a group of angiosperms that includes the orders Apiales, Aquifoliales, Asterales, and Dipsacales together with several small, poorly known families yet unplaced as to order. Parsimony analysis of nucleotide sequence data from the chloroplast genes atpB, ndhF and rbcL together with morphological data are used to construct a phylogeny of the order Asterals, which in the present sense includes 11 families and more than 26 000 species. It is argued that Rousseaceae should be expanded to include also Carpodetaceae (and thus contain four genera), and that Donatia should be re-merged with Stylidiaceae. The present study also strongly supports that the sister taxon to the largest plant family, Asteraceae (Compositae), is the small South American Calyceraceae. A new addition to Asterales is Platyspermation (formerly in Escalloniaceae). Using the recently developed Bayesian approach to phylogenetic reconstruction, in combination with a dataset consisting of the atpB, ndhF and rbcL nucleotide sequences, a resolved and fairly well supported phylogeny of the Euasterids II could be reconstructed. Based on this analysis, and furthermore supported by morphological characters, the South Hemispheric family Escalloniaceae is recircumscribed to include the monogeneric families Eremosynaceae, Tribelaceae and Polyosmaceae, and in this new circumscription it includes seven genera. The formerly escalloniacean member Quintinia, together with the monogeneric families Paracryphiaceae and Sphenostemonaceae, is argued to consitute a monophyletic family Paracryphiaceae sensu lato, supported by several morphological characters.

Gnathostomes, or jawed vertebrates, make up the overwhelming majority of modern vertebrate diversity. Among living vertebrates, they comprise the chondrichthyans (“cartilaginous fishes” such as sharks, skates, rays, chimaeras) and the osteichthyans (“bony fishes” or bony vertebrates, inclusive of tetrapods). Gnathostomes appear to have originated in the early Palaeozoic Era, but their early fossil record is fairly scant. The best fossils appear first in the Late Silurian and Devonian periods. Much of gnathostome diversity owes to unique adaptations in the internal skeleton of their head (the endocranium). The endocranium is composed of the braincase, jaws, hyoid arch, and branchial arches, which sometimes fossilise when they are composed of bone or calcified cartilage. The purpose of this thesis is to describe and compare the fossilised cranial endoskeletons of a variety of different Palaeozoic gnathostomes. The objective is to test current conceptions of gnathostome interrelationships (i.e. phylogeny) and infer aspects of key morphological transformations that took place during the evolution of Palaeozoic members of this group. Two key areas are examined: the morphology and interrelationships of Palaeozoic gnathostomes and the morphology of the visceral arches in sarcopterygian fishes. New data on the visceral arches are described from the stem tetrapods Panderichthys and rhizodontids. These provide insight into the sequence of character acquisition leading to the tetrapod middle ear. Panderichthys shows key features of the tetrapod middle ear chamber were established prior to the origin fo digited limbs. New morphological data are described from the “acanthodian” fish Ptomacanthus. Ptomacanthus provides only the second example of a well-preserved braincase from any member of this group. It shows dramatic differences from that of its counterpart, Acanthodes, providing new evidence for acanthodian paraphyly. New interpretations of basal gnathostome and osteichthyan phylogeny are presented, challenging or enriching existing views of these problems.

Oestrogens are produced by aromatisation of androgens by the aromatase enzyme. In the vertebrate brain this synthesis has vital functions in nerve protection, cell proliferation and nerve development during injury respectively brain development. Brain oestrogens are also crucial in activating certain reproductive and aggressive behaviours in mammals and birds. Teleosts have remarkably high activity of brain aromatase (bAA) compared to mammals and birds; 100-1000 times higher in brain regions like the hypothalamus, pre-optic area and optic tectum. The role of brain oestrogens in teleost behaviour is, however, less clear than in e.g. songbirds and rodents. This thesis studies the potential role of brain aromatase and brain oestrogens in the reproductive behaviour of the guppy male (Poecilia reticulata), how guppy brain aromatase responds to steroids and is distributed in the adult brain. The thesis shows that male behaviour can be affected by brain aromatase. Reduction of bAA by aromatase inhibitor treatment reduced the sexual behaviours sigmoid display and gonopodium swinging (I) and oestrogen receptor blocking with an oestrogen antagonist reduced the number of successful mating attempts (II). The anatomical study (IV) showed that brain aromatase is distributed in areas of the adult guppy brain that are connected to reproductive control and behaviour in vertebrates. Guppy bAA is stimulated by both androgens and oestrogens (III) but is more sensitive to oestrogens, especially in males, and could thus be used as an indicator of endocrine disruption at low concentrations found in the environment. The thesis can also conclude that females possess more brain aromatase than males, and that although it is expressed in the same pattern throughout the brain in both genders the enzymatic activity is differently distributed between the sexes. / Aromatase and androgens in fish reproductive behaviour

No description available.

One of the most unforgiving processes in nature is that of Muller's ratchet, a seemingly irreversible accumulation of deleterious mutations that all organisms have to deal with or face extinction. The most obvious way to avoid fitness collapse is recombination, though asexual populations usually do not have the luxury of recombining freely.  With the aid of computational and mathematical models, we have studied other situations where this threat is averted and the organism can survive the ratchet. The results show that a ratchet where all mutations have the same deleterious fitness effect is very effectively stalled for large effects. However, if mutations are allowed to have a broad range of effects, the fitness-loss rate can be substantial even with the same mean effect as the one-type ratchet, but we have  identified parameter regions where even the broad-range effects are effectively stopped. The fitness-loss from a ratchet is very sensitive to the mutation rate and a mutation that increases the mutation rate (mutator) can easily start an otherwise stalled ratchet. Large effect mutators are heavily counter-selected, but smaller mutators can spread in the population. They can be stopped by reversals (antimutators), but even if the mutation rate is equilibrated in this way, there will be large fluctuations in mutation rate and even larger in the fitness-loss rate due to the feedback amplification in their coupling.    Another way of preventing the ratchet is by reversal of the deleterious mutations themselves through back-mutations or compensatory mutations. The rate required to stop the ratchet using only back-mutations before the fitness collapses is very large. A detailed comparison between the deleterious mutations in the ratchet and in a sexual population was made and the difference was found to be greatest for large populations with large genomes. There are obviously many ways to survive the ratchet, but even more ways to drive a species to extinction by enhancing and speeding up the ratchet. By modelling and testing the ratchet for numerous different situations, we show the effects of some of these threats and benefits.

<p><i>Echiochilon</i>, <i>Ogastemma</i> and <i>Sericostoma</i> are revised resulting in the recognition of 15 species of <i>Echiochilon</i> and one <i>Ogastemma</i> species. Several species are placed in synonymy and three new species are described, <i>E. baricum</i>, <i>E. callianthum</i> and <i>E. cyananthum</i>. The single species of <i>Sericostoma</i> is shown to be nested within <i>Echiochilon</i>.</p><p>The plastid <i>atpB</i> gene was sequenced for <i>Echiochilon</i> and <i>Ogastemma</i> from the Old World and <i>Antiphytum</i> from the New World, plus for a selection of 33 other Boraginaceae taxa. They were analysed together with selected outgroup taxa to give a framework of the tribes of Boraginoideae. The analysis gave support for establishing the new tribe Echiochileae for <i>Antiphytum</i>, <i>Echiochilon</i> and <i>Ogastemma</i>, and for merging the traditionally accepted tribe Eritrichieae with Cynoglosseae. The ITS region was sequenced for all but one species of <i>Echiochilon</i> and for representatives of <i>Antiphytum</i> and <i>Ogastemma</i>. Phylogenetic analysis of Echiochilon revealed that the strongly zygomorphic-flowered species form a paraphyletic group. The morphological data gave results fairly congruent with the ITS phylogeny.</p><p>Biogeographic interpretations of the ITS and <i>atpB</i> phylogenies indicated a trans-Atlantic dispersal of <i>Antiphytum</i> as the most plausible explanation to the Old/New World disjunction. Analyses using DIVA (Dispersal Vicariance Analysis) of the distributions of the <i>Echiochilon</i> species indicated an ancestor to <i>Echiochilon</i> with a wide distribution over northern Africa and Arabia to India.</p>

<p>I use morphological, vocal, molecular, behavioural, ecological and distributional data to re-evaluate the systematics of three passerine bird groups, the <i>Mirafraassamica </i>complex (bush-larks), the genus <i>Seicercus</i> ("spectacled-warblers"; with emphasis on the the <i>S. burkii</i> complex) and the genus <i>Motacilla</i> (wagtails). Two new species are described: <i>Seicercus soror</i> and <i>Motacilla samveasnae</i>. I propose that the polytypic species <i>M. assamica</i> should be treated as four separate species: <i>M. assamica</i>, <i>M. affinis</i>, <i>M. microptera</i> and <i>M. marionae</i> (it is also remarked that the proper name of the latter is <i>M. erythrocephala</i>). That is primarily supported by vocalisations and mitochondrial DNA. The latter data set also suggests that <i>M. assamica</i> sensu lato is paraphyletic, since <i>M. erythroptera</i>, which is always treated as a separate species, is nested within the <i>M. assamica</i> complex. I propose that the polytypic species <i>S. burkii</i> comprises six sibling species. Some of these are found to breed sympatrically, although mainly or entirely segregated altitudinally. Mitochondrial DNA suggests that the <i>S. burkii</i> complex is non-monophyletic, and also that the divergence of the different taxa is much older than indicated by morphological and vocal data. According to the molecular phylogeny, both the genera <i>Seicercus</i> and its assumed sister genus <i>Phylloscopus</i> are paraphyletic. That is corroborated by independent data. The phylogenetic study of the genus <i>Motacilla</i> reveals incongruence between mitochondrial DNA, nuclear DNA and non-molecular data. I conclude that the nuclear gene tree reflects the organismal phylogeny more faithfully than the mitochondrial gene tree. The latter is likely to have been affected by introgressive hybridisation, possibly also stochastic lineage sorting. The most remarkable result that is strongly supported by both mitochondrial and nuclear DNA is that <i>M. flava</i> is non-monophyletic.</p>

<p>This thesis deals with evolutionary relationships within the asterids, a group of plants comprising about one-third of all flowering plants.</p><p>Two new families are recognised: Pennantiaceae and Stemonuraceae. The woody <i>Pennantia</i> from New Zealand and Australia is the sole genus of Pennantiaceae. Stemonuraceae consist of a dozen woody genera with a pantropical distribution and a centre of diversity in South East Asia and the Malesian islands. They are characterised by long hairs on their stamens and/or fleshy appendages on their fruits. Both families were formerly included in Icacinaceae. While Pennantiaceae are unrelated to any of the former Icacinaceae and placed in the order Apiales, other former Icacinaceae genera are related to <i>Cardiopteris</i>, a twining herb from South East Asia and Malesia. The monogeneric family Cardiopteridaceae is enlarged as to include also these. Cardiopteridaceae and Stemonuraceae are sister groups and placed in Aquifoliales. The three other families of Aquifoliales are monogeneric and closely related. The Asian Helwingiaceae and the Central/South American Phyllonomaceae are suggested to be merged into Aquifoliaceae (hollies). The genera of Icacinaceae in the traditional sense not placed in any of the above families (all euasterids II) are members of early diverging lineages of the euasterids I and possibly included in the order Garryales.</p><p>The three woody Australasian families Alseuosmiaceae, Argophyllaceae, and Phellinaceae are confirmed as members of Asterales, despite traditional placements not close to that order. They are, moreover, supported as each other’s closest relatives.</p><p>The results are based mainly on parsimony analysis of DNA sequence data, but morphological studies have revealed characters in support for the molecularly based conclusions. The gene that has provided most new information is the chloroplast <i>ndh</i>F gene. The results are, however, drawn from combined analyses of sequences from one or several additional genes (<i>atp</i>B, <i>mat</i>K, <i>rbc</i>L, <i>18S</i> rDNA). The data have also been explored with Bayesian analysis, a statistical, model-based method that most recently has been developed for phylogeny reconstruction.</p>