The group of pleurocarpous mosses comprises approximately 5000 species, which corresponds to about half of all mosses. The pleurocarpous mosses (i.e. “the Core Pleurocarps”) form a monophylum, which consists typically of perennial mosses with creeping stems and abundant lateral branches. In pleurocarpous mosses the archegonium and thus also sporophyte development is restricted to the apices of short, specialized lateral branches, in contrast to most other mosses, where archegonia and sporophytes develop terminally on the main axis (acrocarpous) or on major branches (cladocarpous). Traditionally, pleurocarpous mosses have been divided into three orders based mainly on their sporophytic characters. Brotherus described the Neckeraceae in 1925 and placed it into the Leucodontales, later the family has alternatively been divided into two or three separate families: the Thamnobryaceae, the Neckeraceae and the Leptodontaceae. These families have been placed even in different orders (Neckeraceae and Leptodontaceae among the leucodontalean mosses and Thamnobryaceae among hypnalean mosses) according to their peristome structure and the grade of peristome reduction. A growing amount of evidence indicates that a grouping based on sporophytic characters is artificial and based on convergent evolution. According to the latest phylogenetic studies of pleurocarpous mosses, based on molecular data, the Neckeraceae belong to the order Hypnales and share a sister group relationship with the Lembophyllaceae. In the most recent comprehensive classification 28 genera were included in the Neckeraceae family. This classification was based on both morphological and molecular data, but done with limited taxon sampling that did not cover all species of the family. Some previous studies based on molecular data have challenged the family concept of the Neckeraceae, indicating the need for a revision of the family. Here the family concept of the Neckeraceae is revisited, the closest relatives of the family are resolved and its position within pleurocarpous mosses is shown. In addition, new insights into the morphological evolution of the family are provided. Previous phylogenetic studies indicated that branch lengths among pleurocarpous mosses are usually extremely short. Therefore we chose to use mainly non-coding DNA sequences from rapidly evolving DNA regions. The phylogenetic reconstructions are based on extensive sequence data from all genomes: plastid trnS-trnF and rpl16, nuclear ITS1 & 2 and mitochondrial nad5. Both parsimony (PAUP and PRAP2) and Bayesian statistics (MrBayes) were employed for phylogenetic reconstructions. In order to use the information provided by length mutations indels were included in the analyses as binary data using a simple indel coding approach. No severe conflicts appeared between the different methods used, but the indel coding affected the support values of the inferred topologies. Therefore, all support values resulting from different methods are shown along the phylogenetic trees. The morphological features are studied and synapomorphies for each clade formed in the phylogenetic analyses are interpreted. A new delimitation of the family makes it necessary to reconsider the relevance of the morphological description and the morphological features characteristic of the family need to be reconsidered. Due to new groupings, some changes in the morphological circumscriptions of the genera are necessary, resulting in two new genera and several new combinations. Chapter 1 gives a broad overview of the relationships of the pleurocarpous mosses and shows the need for changes in the definition of genera, families and the corresponding nomenclature in this group. Chapter 2 is a population genetic study on the genus Thamnobryum. The main aim of this chapter is to test the species concept in Thamnobryum that are endemic to strictly restricted regions showing only minor differences in the morphological features in comparison to some more common species. In Chapter 3 the monophyly of the Neckeraceae is tested. In addition, in this chapter the ancestral character states of some morphological characters within the Neckeraceae are reconstructed. Chapters 4 and 5 resolve the genus composition and the relationships within the family in more detail. The results of this thesis show that the Neckeraceae need re-circumscription; this includes changes in the genus composition. The Lembophyllaceae is confirmed to be the sister group of the Neckeraceae. In addition to the new phylogeny, the potential evolution of several characters as a result of environmental selection pressures is analyzed. From the ancestral state reconstructions made (using BayesTraits) for both the habitat and a selection of morphological characters, character state distributions and habitat shift appear congruent, peristome reduction being a good example. However, some character states do not correlate with the habitat, suggesting very complex evolutionary patterns underlying these morphological characters. Many widely distributed genera that are composed of several species and seem to be morphologically coherent (Echinodium, Homalia, Thamnobryum, partly Neckera), are shown in this thesis to be polyphyletic. They are replaced by smaller, geographically more restricted genera that at least in some cases (e.g. Thamnomalia, Homalia s.str., Neckera s.str.) seem to form morphologically heterogeneous genera. In other words, morphology can be misleading in the family Neckeraceae even at the genus level and convergent evolution in both morphological and sequence level characters are common within the family. Special habitat conditions have been shown to result in similar morphological structures also in several other moss groups. This kind of convergent evolution occurs in aquatic mosses, and seems to have occurred among the neckeraceous species Thamnobryum alopecurum and its allies. However, similar morphological structure in similar aquatic habitats can also be due to true phylogenetic relationships as is the case within the Neckeraceae for Handeliobryum sikkimense and Hydrocryphae wardii, or the members of Touwia. The geographical grouping seems to be more strongly correlated with the phylogenetic grouping than thought before.

The group of pleurocarpous mosses comprises approximately 5000 species, which corresponds to about half of all mosses. The pleurocarpous mosses (i.e. “the Core Pleurocarps”) form a monophylum, which consists typically of perennial mosses with creeping stems and abundant lateral branches. In pleurocarpous mosses the archegonium and thus also sporophyte development is restricted to the apices of short, specialized lateral branches, in contrast to most other mosses, where archegonia and sporophytes develop terminally on the main axis (acrocarpous) or on major branches (cladocarpous). Traditionally, pleurocarpous mosses have been divided into three orders based mainly on their sporophytic characters. Brotherus described the Neckeraceae in 1925 and placed it into the Leucodontales, later the family has alternatively been divided into two or three separate families: the Thamnobryaceae, the Neckeraceae and the Leptodontaceae. These families have been placed even in different orders (Neckeraceae and Leptodontaceae among the leucodontalean mosses and Thamnobryaceae among hypnalean mosses) according to their peristome structure and the grade of peristome reduction. A growing amount of evidence indicates that a grouping based on sporophytic characters is artificial and based on convergent evolution. According to the latest phylogenetic studies of pleurocarpous mosses, based on molecular data, the Neckeraceae belong to the order Hypnales and share a sister group relationship with the Lembophyllaceae. In the most recent comprehensive classification 28 genera were included in the Neckeraceae family. This classification was based on both morphological and molecular data, but done with limited taxon sampling that did not cover all species of the family. Some previous studies based on molecular data have challenged the family concept of the Neckeraceae, indicating the need for a revision of the family. Here the family concept of the Neckeraceae is revisited, the closest relatives of the family are resolved and its position within pleurocarpous mosses is shown. In addition, new insights into the morphological evolution of the family are provided. Previous phylogenetic studies indicated that branch lengths among pleurocarpous mosses are usually extremely short. Therefore we chose to use mainly non-coding DNA sequences from rapidly evolving DNA regions. The phylogenetic reconstructions are based on extensive sequence data from all genomes: plastid trnS-trnF and rpl16, nuclear ITS1 & 2 and mitochondrial nad5. Both parsimony (PAUP and PRAP2) and Bayesian statistics (MrBayes) were employed for phylogenetic reconstructions. In order to use the information provided by length mutations indels were included in the analyses as binary data using a simple indel coding approach. No severe conflicts appeared between the different methods used, but the indel coding affected the support values of the inferred topologies. Therefore, all support values resulting from different methods are shown along the phylogenetic trees. The morphological features are studied and synapomorphies for each clade formed in the phylogenetic analyses are interpreted. A new delimitation of the family makes it necessary to reconsider the relevance of the morphological description and the morphological features characteristic of the family need to be reconsidered. Due to new groupings, some changes in the morphological circumscriptions of the genera are necessary, resulting in two new genera and several new combinations. Chapter 1 gives a broad overview of the relationships of the pleurocarpous mosses and shows the need for changes in the definition of genera, families and the corresponding nomenclature in this group. Chapter 2 is a population genetic study on the genus Thamnobryum. The main aim of this chapter is to test the species concept in Thamnobryum that are endemic to strictly restricted regions showing only minor differences in the morphological features in comparison to some more common species. In Chapter 3 the monophyly of the Neckeraceae is tested. In addition, in this chapter the ancestral character states of some morphological characters within the Neckeraceae are reconstructed. Chapters 4 and 5 resolve the genus composition and the relationships within the family in more detail. The results of this thesis show that the Neckeraceae need re-circumscription; this includes changes in the genus composition. The Lembophyllaceae is confirmed to be the sister group of the Neckeraceae. In addition to the new phylogeny, the potential evolution of several characters as a result of environmental selection pressures is analyzed. From the ancestral state reconstructions made (using BayesTraits) for both the habitat and a selection of morphological characters, character state distributions and habitat shift appear congruent, peristome reduction being a good example. However, some character states do not correlate with the habitat, suggesting very complex evolutionary patterns underlying these morphological characters. Many widely distributed genera that are composed of several species and seem to be morphologically coherent (Echinodium, Homalia, Thamnobryum, partly Neckera), are shown in this thesis to be polyphyletic. They are replaced by smaller, geographically more restricted genera that at least in some cases (e.g. Thamnomalia, Homalia s.str., Neckera s.str.) seem to form morphologically heterogeneous genera. In other words, morphology can be misleading in the family Neckeraceae even at the genus level and convergent evolution in both morphological and sequence level characters are common within the family. Special habitat conditions have been shown to result in similar morphological structures also in several other moss groups. This kind of convergent evolution occurs in aquatic mosses, and seems to have occurred among the neckeraceous species Thamnobryum alopecurum and its allies. However, similar morphological structure in similar aquatic habitats can also be due to true phylogenetic relationships as is the case within the Neckeraceae for Handeliobryum sikkimense and Hydrocryphae wardii, or the members of Touwia. The geographical grouping seems to be more strongly correlated with the phylogenetic grouping than thought before.

Ökomorphologische Konvergenz tritt auf, wenn Arten mit demselben Lebensstil unabhängig voneinander ähnliche morphologische Merkmale evolvieren. Neue Fallstudien können dabei helfen, die diesem Prozess zugrunde liegenden Mechanismen aufzuklären. Diese Arbeit befasst sich mit einigen konvergent evolvierten, langsamen, baumbewohnenden Säugetieren, d. h. zwei Linien baumlebender Faultieren, dem Zwergameisenbär, den Lorisiden, zwei Kladen ausgestorbener Lemuren, d. h. Paläopropithekiden und Megaladapis, und dem Koala. Es werden funktionsmorphologische Konvergenzen in diesen Taxa erforscht, indem ihr Humerus und ihr Femur sowie jene ihrer nah verwandten, aber ökologisch unterschiedlichen Taxa untersucht werden. Erstmals werden Knochen mittels phylogenetisch vergleichender Methoden auf vier anatomischen Ebenen, d.h. äußere Form, diaphysäre Mikrostruktur und Anatomie sowie epiphysäre Trabekelarchitektur, analysiert. Viele langsame, baumbewohnende Säugetiere teilen eine geringe kortikale Kompaktheit, was wahrscheinlich mit ihrer extrem niedrigen Stoffwechselrate und ihren biomechanischen Anforderungen zusammenhängt. Langsame, arboreale Xenarthra, d. h. baumbewohnende Faultiere und der Zwergameisenbär, weisen ein Muster unvollständiger Konvergenz für eine Reihe äußerer und innerer anatomischer Merkmale auf, was möglicherweise durch die relativ unterschiedliche Ökologie des Zwergameisenbären erklärt wird. Auf einer breiteren Säugetierskala konvergieren andere Merkmale, die möglicherweise mit der Ökologie der langsamen baumbewohnenden Lebensweise in einigen der untersuchten Taxa zusammenhängen, obwohl komplexe Muster auch durch andere evolutionäre Prozesse erklärt werden können. Nur suspensorisch lebende Taxa tragen signifikant zur Konvergenz bei. Diese Arbeit hebt die stärkere Konvergenz hervor, die sich in der inneren Struktur des Knochens widerspiegelt. / Ecomorphological convergence occurs when similar morphological traits are independently evolved by species with the same lifestyle. Novel case studies can help to elucidate the underlying mechanisms of this process. This work addresses some convergent slow arboreal mammals, i.e. two lineages of ‘tree sloths’, the silky anteater, ‘Lorisidae’, two clades of extinct lemurs, i.e. palaeopropithecids and Megaladapis, and the koala. Functional morphological convergences are searched in these taxa, studying their humerus and femur as well as those of their closely related ecologically distinct taxa. For the first time, bones are analyzed at four anatomical levels, i.e. external shape, diaphyseal microstructure and anatomy and epiphyseal trabecular architecture, through phylogenetic comparative methods. Many slow arboreal mammals share a low cortical compactness, probably related to their extremely low metabolic rate and biomechanical demands. Slow arboreal xenarthrans, i.e. ‘tree sloths’ and the silky anteater, exhibit a pattern of incomplete convergence for a set of external and internal anatomical features, possibly explained by the relatively distinct ecology of the silky anteater. On a wider mammalian scale, other traits possibly related to slow arboreal ecology converge in some of the studied taxa, although with complex patterns also explained by other evolutionary processes. Only suspensory taxa significantly contribute to convergence. This thesis highlights the stronger convergence reflected by bone internal structure. By providing potential explanations for convergence in slow arboreal mammals, the inherent complexity of this process is here emphasized.

Background: Heat-shock proteins of the 70 kDa family (Hsp70s) are essential chaperones required for key cellular functions. In eukaryotes, four subfamilies can be distinguished according to their function and localisation in different cellular compartments: cytosol, endoplasmic reticulum, mitochondria and chloroplasts. Generally, multiple cytosol-type Hsp70s can be found in metazoans that show either constitutive expression and/or stress-inducibility, arguing for the evolution of different tasks and functions. Information about the hsp70 copy number and diversity in microbial eukaryotes is, however, scarce, and detailed knowledge about the differential gene expression in most protists is lacking. Therefore, we have characterised the Hsp70 gene family of Paramecium caudatum to gain insight into the evolution and differential heat stress response of the distinct family members in protists and to investigate the diversification of eukaryotic hsp70s focusing on the evolution of heat-inducibility. Results: Eleven putative hsp70 genes could be detected in P. caudatum comprising homologs of three major Hsp70-subfamilies. Phylogenetic analyses revealed five evolutionarily distinct Hsp70-groups, each with a closer relationship to orthologous sequences of Paramecium tetraurelia than to another P. caudatum Hsp70-group. These highly diverse, paralogous groups resulted from duplications preceding Paramecium speciation, underwent divergent evolution and were subject to purifying selection. Heat-shock treatments were performed to test for differential expression patterns among the five Hsp70-groups as well as for a functional conservation within Paramecium. These treatments induced exceptionally high mRNA up-regulations in one cytosolic group with a low basal expression, indicative for the major heat inducible hsp70s. All other groups showed comparatively high basal expression levels and moderate heat-inducibility, signifying constitutively expressed genes. Comparative EST analyses for P. tetraurelia hsp70s unveiled a corresponding expression pattern, which supports a functionally conserved evolution of the Hsp70 gene family in Paramecium. Conclusions: Our analyses suggest an independent evolution of the heat-inducible cytosol-type hsp70s in Paramecium and in its close relative Tetrahymena, as well as within higher eukaryotes. This result indicates convergent evolution during hsp70 subfunctionalization and implies that heat-inducibility evolved several times during the course of eukaryotic evolution.

Background: Heat-shock proteins of the 70 kDa family (Hsp70s) are essential chaperones required for key cellular functions. In eukaryotes, four subfamilies can be distinguished according to their function and localisation in different cellular compartments: cytosol, endoplasmic reticulum, mitochondria and chloroplasts. Generally, multiple cytosol-type Hsp70s can be found in metazoans that show either constitutive expression and/or stress-inducibility, arguing for the evolution of different tasks and functions. Information about the hsp70 copy number and diversity in microbial eukaryotes is, however, scarce, and detailed knowledge about the differential gene expression in most protists is lacking. Therefore, we have characterised the Hsp70 gene family of Paramecium caudatum to gain insight into the evolution and differential heat stress response of the distinct family members in protists and to investigate the diversification of eukaryotic hsp70s focusing on the evolution of heat-inducibility. Results: Eleven putative hsp70 genes could be detected in P. caudatum comprising homologs of three major Hsp70-subfamilies. Phylogenetic analyses revealed five evolutionarily distinct Hsp70-groups, each with a closer relationship to orthologous sequences of Paramecium tetraurelia than to another P. caudatum Hsp70-group. These highly diverse, paralogous groups resulted from duplications preceding Paramecium speciation, underwent divergent evolution and were subject to purifying selection. Heat-shock treatments were performed to test for differential expression patterns among the five Hsp70-groups as well as for a functional conservation within Paramecium. These treatments induced exceptionally high mRNA up-regulations in one cytosolic group with a low basal expression, indicative for the major heat inducible hsp70s. All other groups showed comparatively high basal expression levels and moderate heat-inducibility, signifying constitutively expressed genes. Comparative EST analyses for P. tetraurelia hsp70s unveiled a corresponding expression pattern, which supports a functionally conserved evolution of the Hsp70 gene family in Paramecium. Conclusions: Our analyses suggest an independent evolution of the heat-inducible cytosol-type hsp70s in Paramecium and in its close relative Tetrahymena, as well as within higher eukaryotes. This result indicates convergent evolution during hsp70 subfunctionalization and implies that heat-inducibility evolved several times during the course of eukaryotic evolution.