Comprendre comment évoluent les écosystèmes lors de perturbations majeures de l'environnement nécessite de prendre en compte l'histoire évolutive des espèces, c'est-à-dire leur phylogénie. Dans ce Mémoire de thèse, j'utilise une approche modélisatrice afin d'étudier l'évolution phénotypique en lien avec l'environnement (écomorphologie), approche incluant des données actuelles et fossiles et visant in fine à comprendre comment les écosystèmes se diversifient et s'organisent structurellement et fonctionnellement à l'échelle des temps géologiques. Le cadre historique, fourni par la phylogénie, définit une trame analytique commune à l'étude de taxons actuels et fossiles. Différents outils dédiés, d'une part au traitement de données morphométriques souvent incomplètes, et d'autre part aux études écomorphologiques évolutives dans un contexte phylogénétique, sont développés et discutés. Cette thèse s'articule autour de deux ateliers consacrés à l'étude macroévolutive et macroécologique de grands mammifères « ongulés » : Bovidae, Cervidae, et Equidae. Le premier atelier concerne les ruminants actuels, clade très diversifié et à l'écologie des espèces constitutives connue. Les analyses révèlent des modes évolutifs contrastés entre Cervidae et Bovidae actuels, en fonction des niches écologiques et des caractères écomorphologiques considérés. La diversification moi pliocène des Bovidae africains apparaît corrélée à des évènements globaux, tandis que leur évolution phénotypique révèle des modes de diversification différents selon les habitats. Le second atelier est focalisé sur l'impact d'un événement majeur ayant affecté l'ensemble du pourtour méditerranéen à la limite Miocène- Pliocène, il y a 5,3 millions d'années : la Crise de Salinité Messinienne. Il repose sur l'étude d' « ongulés » fossiles et apporte des précisions sur l'aspect temporel de l'évolution de ces mammifères avant et après cet événement. Les principaux résultats illustrent des variations rapides de la structure phylogénétique des assemblages d'ongulés qu'il est possible de relier au contexte biogéographique et aux variations climatiques locales et régionales. Les renouvellements fauniques (acteurs) et fonctionnels (rôles), initiés dès la fin du Miocène moyen, apparaissent progressifs et non soudains. Les résultats obtenus offrent une meilleure caractérisation et compréhension des réponses évolutives de ces mammifères, grands consommateurs primaires souvent parmi les premiers menacés lors de perturbations climatiques et environnementales majeures / Understanding how ecosystems evolve when facing severe climatic perturbations of the environment requires an historic perspective provided by species phylogenies. In this thesis, I use a modelling framework to investigate the question of the phenotypic evolution of species as an adaptation to the environment (ecomorphology). I combine extant and fossil data to study how ecosystems are diversifying and organizing structurally and functionally on a geological time scale. The historical context provided by the phylogeny defines a unified analytical framework for the study of extant and fossil taxa. Several analytical tools dedicated, on the one hand to deal with missing cases in morphometric studies, and on the other hand to ecomorphological studies in a phylogenetic context, are developed and discussed. This PhD thesis is organized along two main research axes dedicated to the macroevolutionary and macroecological study of three large mammals “ungulate” families: Bovidae, Cervidae, and Equidae. First I focus on extant ruminants, a well diversified clade for which species ecological preferences are well known. The analyses show contrasted evolutionary modes between extant cervids and bovids, depending on the ecological niche and ecomorphological traits under scrutiny. The Mio-Pliocene diversification of African bovids appears to be correlated with global climatic events while their phenotypic evolution shows contrasted evolutionary patterns depending on the habitat. The second axis focuses on the circum Méditerranean impact of a major event that took place at the Miocene-Pliocene boundary, 5.3 million years ago: the Messinian Salinity Crisis. The comparative study of extinct “ungulates” living before and after this event provides some clues about the evolutionary rates and spatial patterns of phylogenetic diversity of these large mammals. The phylogenetic structure of the ungulate communities shows abrupt changes related to the local and regional biogeographic context as well as variations in climate conditions. Meanwhile, progressive faunal (actors) and functional (roles) turnovers are depicted from the beginning of the Late Miocene onward. These results provide a better characterization and understanding of the evolutionary responses to broad climatic and environmental perturbations of these often-threatened, large primary consumer “ungulate” mammals

Este trabalho focou nos aspectos evolutivos e ecológicos dos tricomas em espécies da tribo Bignonieae (Bignoniaceae), e no potencial papel defensivo dessas estruturas no aumento da resistência das plantas frente aos herbívoros. Esta tese caracterizou quatro morfotipos de tricomas encontrados nas partes vegetativas das plantas da tribo Bignonieae morfologicamente, estudou os padrões macro-evolutivos destes tricomas e o papel funcional dos tricomas pateliformes secretores de néctar em diferentes escalas de tempo e espaço. O primeiro capítulo descreveu os quatro tipos de tricomas micro e macro-morfologicamente, e sua relação com a história filogenética das espécies da tribo Bignonieae. Quatro tipos de tricomas distintos foram reconhecidos: tricomas não glandulares (ng), tricomas glandulares peltados (gp), tricomas glandulares estipitados (ge), e tricomas glandulares pateliformes/cupulares (P/C). Três destes tricomas provavelmente já estavam presentes no ancestral da tribo Bignonieae (i.e., não glandulares, glandulares peltados e glandulares pateliformes/cupulares) enquanto os tricomas glandulares estipitados surgiram mais recentemente e múltiplas vezes dentro da tribo. Além disso, realizamos uma revisão das terminologias aplicadas a esses tricomas bem como sugerimos uma padronização dos tipos de tricomas para o grupo de forma a diminuir a divergência entre os trabalhos já publicados. O segundo capítulo testou a eficiência dos tricomas pateliformes secretores de néctar (nectários extraflorais) em duas espécies de Anemopaegma das savanas brasileiras. Essas duas espécies atraíram mais formigas que plantas vizinhas, com os indivíduos de Anemopaegma com maiores quantidades de nectários sendo visitados por um número maior de formigas que as plantas com um número menor de nectários (variação intra-populacional). No entanto, não foi possível observar um efeito dos nectários e formigas sobre a herbivoria e o desempenho das plantas como esperado pela hipótese de defesa mediada por essas estruturas. Hipóteses alternativas foram utilizadas para discutir os resultados, incluindo uma discussão sobre o possível custo/benefício de tais estruturas em diferentes ambientes (florestas e savanas) relacionados com a história filogenética do grupo, além da variação das interações esperada entre populações segundo a hipótese de mosaico geográfico. O terceiro capítulo testou o papel defensivo do sistema planta-formiga mediado pelos nectários extraflorais no contexto filogenético da tribo Bignonieae. Espécies de plantas com um número maior de nectários foram visitadas por um número maior de formigas (teste controlado pelas relações de parentesco entre as espécies). Além disso, espécies mais proximamente relacionadas apresentaram uma diferença na abundância de nectários extraflorais maior do que o esperado pelo modelo neutro de evolução, gerando um padrão de convergência dessas estruturas na tribo Bignonieae. Esse desvio no padrão evolutivo esperado pode ter sido causado por forças direcionais de seleção e momentos de contra-seleção, dado o balanço entre custo-benefício dos nectários extraflorais para as plantas. Neste contexto, dois fatores foram testados para explicar o desvio na evolução da abundância de nectários na tribo Bignonieae: (1) mudança de habitat das florestas para as savannas (fatores extrínsecos); (2) surgimento de novos caracteres morfológicos como outros tipos de tricomas na superfície das plantas (fatores intrínsecos). Ambos fatores podem ter interferido nas interações formiga-planta e no padrão de evolução dos nectários. Dessa forma, a ocupação das savanas levou a uma diminuição do número de nectários (provável contra-seleção dessas estruturas), enquanto o surgimento de tricomas glandulares adesivos teve o mesmo efeito sobre os nectários. Ambos resultados são discutidos considerando a condicionalidade das interações em função da variação biótica (formigas e herbívoros) entre habitats, e também do \"trade-off\" entre os caracteres de defesa. O quarto capítulo testou a teoria de coevolução em mosaico geográfico no sistema planta-formiga-herbívoro em 10 populações da espécie de savana Anemopaegma álbum. Não foram encontradas correlações entre os nectários (e variáveis descritoras do néctar), a abundância de formigas visitantes, a herbivoria ou as variáveis de performance das plantas entre as populações. Esse padrão esteve associado principalmente a variação na assembléia de formigas, a qual foi dominada por formigas do gênero Crematogaster em uma das populações, mas dominadas por formigas do gênero Camponotus na grande maioria das outras populações. No entanto, 3 das 10 populações estudadas apresentaram um alto número de plantas sem formigas, diminuindo muito as chances dessas populações serem defendidas frente aos herbívoros pelos nectários. A abundância de formigas esteve relacionada negativamente com a herbivoria, e positivamente com as variáveis de performance das plantas entre as populações. Das 10 populações amostradas, 5 delas tiveram os nectários acoplados (do inglês, \"matched\") com as formigas visitantes, embora o tipo de acoplamento tenha variado entre elas. Dessas 5 populações, somente 3 tiveram uma produção positiva de folhas e baixa herbivoria, no qual duas delas tiveram alta abundância de nectários nas folhas e foram dominadas por formigas do gênero Camponotus (maiores em tamanho, mas com baixa capacidade de recrutamento). Já a terceira população teve em média a menor abundância de nectários nas folhas, e a mesma apresentou o maior número de formigas por planta (e maior freqüência), em geral formigas do gênero Crematogaster (menores em tamanho mas com grande capacidade de recrutamento). Por serem menores em tamanho, tais formigas utilizaram a secreção dos nectários quase que individualmente. Nem a abundância de nectários, nem as formigas ou mesmo a herbivoria estiveram estruturados espacialmente, corroborando, a hipótese de mosaico geográfico para as interações formiga-planta-herbívoro em A.album. Neste contexto, as três populações com as interações formiga-planta \"mached\" foram consideradas \"hot-spots\" das interações, nas quais as populações atingiram os valores mais altos das variáveis de performance das plantas, enquanto as demais foram consideradas \"cold-spots\". A maioria das populações \"cold-spots\" foi explicada pela falta de formigas suficientes para que as interações com as plantas pudessem se tornar efetivas na defesa frente aos herbívoros, mas outros processos também foram considerados para discutir os resultados apresentados neste trabalho. / This thesis focused on the evolutionary-ecology of trichomes in the tribe Bignonieae (Bignoniaceae), and in the potential defensive role of these structures against herbivores. More specifically, we characterized four trichome morphotypes found in vegetative plant parts of representatives of the Bignonieae, as well as studied the macro-evolutionary patterns of these trichomes, and the functional role of patelliforme nectar secreting trichomes in different scales of time and space. The first chapter describes four different trichome types micro and macromorphologically, as well as investigates their evolutionary patterns during the history of Bignonieae. The four different trichome types recognized are: non-glandular trichomes (ng), glandular peltate trichomes (gp), glandular stipitate trichomes (gst), and glandular patteliform/cupular trichomes (P/Cgt). Our analyses indicated that three of these trichomes were likely already present in the most recent common ancestor of the tribe Bignonieae (i.e., non glandular, glandular peltate, and glandular patteliform/cupular), while the glandular stipitate trichomes evolved more recently and multiple times during the history of the tribe. Results from this study were combined with a literature review in order to revise the trichome terminology and propose standardized names for the various trichome types currently found in the group. The second chapter tested the efficiency of patelliform nectar-secreting trichomes (extrafloral nectaries, EFNs) in two species of Anemopaegma of the Brazilian savannas. These two species attracted more ants than neighboring plants, with individuals that presented higher amounts of EFNs being visited by a higher number of ants than plants with lower amounts of EFNs (intra-population variation). Nonetheless, no effect of EFNs and ants was observed on herbivory nor on the performance of the studied plants, contradicting the expectations of the mediated EFNs defense hypothesis. Alternative hypotheses were also considered including the cost/benefits model to understand the outcomes of ant-plant interaction: (1) phylogenetic inertia hypothesis that connect EFNs-ant interactions with the plant transitions between different environments (forests to savannas); and (2) geographic mosaic hypothesis that predict differences in the outcomes of ant-plant interactions across populations. The third chapter tested the defensive role of extrafloral nectaries in the context of the phylogenetic history of Bignonieae. Species of plants with a higher number of EFNs were visited by a higher number of ants (test controlled by phylogeny). In addition, closely related species presented a higher difference in the abundance of EFNs than expected under the neutral model of evolution. Such deviation may have resulted by directional forces of slection and moments of counter-selection, given the costs and benefits of the extrafloral nectaries for the plants. Two specific factors were considered as the major possible determinants of the evolutionary patterns of the EFNs: (1) change of habitat from forests to savannas (extrinsic factors); and (2) emergence of new morphological characters such as other trichome types over the plant\'s surface (intrinsic factors). Both factors might have altered the ant-plant interactions and the evolution of nectaries. The occupation of the savannas was associated with a decrease in the number of nectaries (likely due to counter-selection of these structures), while the evolution of adhesive glandular trichomes presenting the same effect on the nectaries. Both results are discussed in the light of the biotic variation (ants and herbivores) encountered between habitats, as well as in the light of the trade-off among defensive characters. The fourth chapter tested the geographic mosaic theory of coevolution in 10 populations of the savanna species Anemopaegma álbum. No correlations were found among extrafloral nectaries (and nectar variables), the abundance of visiting ants, herbivory, and plant performance among populations. This pattern was mainly associated with the variation in the assembly of ants encountered in the various populations. Most populations of A. álbum were dominated by assemblages of Camponotus ants, except for one that was dominated by Crematogaster ants. However, 3 of 10 populations studied presented a high number of plants without ants, decreasing the chances of defense against herbivores by EFNs. The abundance of ants was negatively associated with herbivory, and positively associated with plant performance variables among populations. Out of the 10 populations sampled, five presented an abundance of EFNs that matched the functional traits of ants. Out of these five populations, only three presented positive leaf production and low herbivory. From these three populations, two presented high abundances of EFNs on the leaves and were dominated by Camponotus ants (i.e., bigger in size, but with a low recruiting capacity). The third population presented on average the smallest abundance of EFNs on the leaves; it also presented the highest number of ants per plant (and the highest frequency), which generally were Crematogaster ants (smaller in size but with greater recruiting capacity). Because these ants are smaller in size, they used nearly all isolated EFNs encountered over the plant\'s surface. Neither the abundance of EFNs, the abundance of ants, and the assemblage of herbivores were structured spatially, corroborating the geographic mosaic hypothesis for the ant-plant-herbivore interactions in A. album. In this context, the three populations with \"matched\" ant-plant interactions were considered \'hot-spots\' of interactions, in which the populations reached the highest values of plant performance, while the others were considered \'cold-spots\'. The majority of \'cold spot\' populations were explained by a lack of sufficient ants to protect the plants effectively

For comparative method purposes is needed to set together price databases. These must be consequently statistically analyzed. Complex procedure of statistics interpretation is described and demonstrated on the exemplary database.

Der Bewegungsapparat der Sciuromorpha, einer monophyletische Gruppe von ca. 300 Arten, wurde verwendet um den Effekt der Lebensweise und der Körpermasse auf die Scapula- und Femurmorphologie zu untersuchen. Diese Nagetierklade weist eine breite Vielfalt an Lebensweisen (arboreal, fossoriell, aerial) als auch Körpermassen (drei Größenordnungen umfassend) auf. Die fossorielle Lebensweise hat sich höchstwahrscheinlich dreimal unabhängig von einem arborealen Vorfahren entwickelt. Mehr als die Hälfte der rezenten Arten wurden untersucht. Die Scapulae wurden fotografiert, während Computertomographie (CT) und Oberflächenlaserscans für die Femora verwendet wurden. Es wurden funktionsrelevante Merkmale analysiert, wie die effektive Länge der Skelettelemente, die Muskeleigenschaften soweit aus der Geometrie der Knochen ableitbar, sowie die Robustheit. Die CT-Scans wurden verwendet, um die Querschnitts- und Trabekeleigenschaften des Femurs zu analysieren. Die Gestalt wurde mittels geometrischer Morphometrie untersucht. Phylogenetic comparative methods wurden unter anderem verwendet, um den Einfluss der Phylogenie zu beurteilen als auch, ob sich die unabhängige Aneignung einer fossoriellen Lebensweise in der Evolution homoplastischer Morphologien widerspiegelt. Die Phylogenie spielte bei der Merkmalsevolution eine vernachlässigbare Rolle. Das Auftreten signifikanter Merkmalsunterschiede zwischen den Lebensweisen sowie allometrischer Anpassungen aufgrund Veränderungen in der Körpermasse hingen von dem jeweiligen Merkmal ab. Bei einigen Merkmalen unterschied sich der Einfluss der Körpermasse signifikant zwischen den einzelnen Lebensweisen, was aber nicht die Regel zu sein scheint. Die Evolution homoplastischer Morphologien war sehr unwahrscheinlich bei den fossoriellen Gruppen. Diese Ergebnisse deuten auf eine komplexe, aber adaptive Evolutionsgeschichte dieser Skelettelemente bei den Sciuromorpha hin. / The vertebrate locomotor apparatus of Sciuromorpha, a monophyletic group of ca. 300 species, was used to investigate the effect of lifestyle and body mass on the scapular and femoral morphology. This rodent clade displays a broad diversity of lifestyles (arboreal, fossorial, aerial) and body masses spanning three orders of magnitude. The fossorial lifestyle evolved most probably three times independently from an arboreal ancestor. More than half of the extant species were included. Scapulae were photographed and computed tomography (CT) and surface laser scans were acquired for the femora. Functionally relevant traits were analysed, e.g., the effective length of the skeletal element, the properties of attaching muscles as inferred from the geometry of the bones, and robustness properties. The CT scans were used to analyse the cross-sectional and trabecular properties of the femur. Bone shape was investigated using geometric morphometrics. Phylogenetic comparative methods were utilized, e.g., to assess phylogenetic inertia and whether the independent acquisition of a fossorial lifestyle is reflected in the evolution of homoplastic morphologies. Phylogenetic inertia played a neglectable role in the trait evolution. Significant differences among lifestyles as well as allometric scaling adjustments depended on the trait under consideration. For some traits, scaling differed significantly among lifestyles, although this did not appear to be the rule. Morphological homoplasy was unlikely among the fossorial groups. The results suggest a complex, but adaptive evolutionary history of these skeletal elements in Sciuromorpha.

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.

No description available.

No description available.

Disabled people comprise seventeen per cent of all New Zealanders and face barriers to participation in community life. This research examines the ways in which two local authorities - Manukau City Council and Christchurch City Council - provide recreation opportunities for people with disabilities and how these local authorities implement national legislation, policies and strategies which bear upon the provision of recreation services for the disabled. To achieve these aims, a comparative method which uses primary (interview) and secondary (documentary) data, is adopted. Central government plays an important role in the provision of recreation to communities, including the disabled community. It develops legislation, strategies and policies that guide the provision of recreation services for people with disabilities at a local level. Local authorities provide ‘spaces’ for recreation including parks, waterways and facilities, as part of their wider services and programmes. Their services respond to broader, including national, concerns, but also reflect local priorities and needs, including those of other ‘special’ populations. The results of this research indicate that Manukau and Christchurch Cities consult the disabled community and its representatives and address the specific needs of this community. Both local authorities have policies which outline how disability issues will be addressed; a disability-specific position which acts as an “internal advocate” for the disabled community; and a reference group made up of members of the disabled community. In the case of Christchurch City Council, the “KiwiAble” programme, “KiwiAble Recreation Network”, “KiwiAble Leisure Card” and Inclusive Communities Coordinator are all disability-specific mechanisms that focus on the recreational needs of the disabled community. However, only one policy in these two cities - the Manukau City Council Disability Policy and Action Plan – draws explicit links to a national strategy. On the basis of these results, a number of recommendations are made for future research and local authority action. It is hoped that the current study will provide practical advice and examples which local authorities can adopt, to further enhance recreational provisions for people with disabilities.

<p>Conservatism is a central theme of organismic evolution. Related species share characteristics due to their common ancestry. Some concern have been raised among evolutionary biologists, whether such conservatism is an expression of natural selection or of a constrained ability to adapt.</p><p>This thesis explores adaptations and constraints within the plant reproductive phase, particularly in relation to the evolution of fleshy fruit types (berries, drupes, etc.) and the seasonal timing of flowering and fruiting. The different studies were arranged along a hierarchy of scale, with general data sets sampled among seed plants at the global scale, through more specific analyses of character evolution within the genus <i>Rhamnus </i>s.l. L. (Rhamnaceae), to descriptive and experimental field studies in a local population of <i>Frangula alnus</i> (Rhamnaceae). Apart from the field study, this thesis is mainly based on comparative methods explicitly incorporating phylogenetic relationships. The comparative study of <i>Rhamnus</i> s.l. species included the reconstruction of phylogenetic hypotheses based on DNA sequences.</p><p>Among geographically overlapping sister clades, biotic pollination was not correlated with higher species richness when compared to wind pollinated plants. Among woody plants, clades characterized by fleshy fruit types were more species rich than their dry-fruited sister clades, suggesting that the fleshy fruit is a key innovation in woody habitats. Moreover, evolution of fleshy fruits was correlated with a change to more closed (darker) habitats.</p><p>An independent contrast study within <i>Rhamnus</i> s.l. documented allometric relations between plant and fruit size. As a phylogenetic constraint, allometric effects must be considered weak or non-existent, though, as they did not prevail among different subclades within <i>Rhamnus</i> s.l. Fruit size was correlated with seed size and seed number in <i>F. alnus</i>. This thesis suggests that frugivore selection on fleshy fruit may be important by constraining the upper limits of fruit size, when a plant lineage is colonizing (darker) habitats where larger seed size is adaptive.</p><p>Phenological correlations with fruit set, dispersal, and seed size in <i>F. alnus</i>, suggested that the evolution of reproductive phenology is constrained by trade-offs and partial interdependences between flowering, fruiting, dispersal, and recruitment phases. Phylogenetic constraints on the evolution of phenology were indicated by a lack of correlation between flowering time and seasonal length within <i>Rhamnus cathartica</i> and <i>F. alnus</i>, respectively. On the other hand, flowering time was correlated with seasonal length among <i>Rhamnus</i> s.l. species. Phenological differences between biotically and wind pollinated angiosperms also suggested adaptive change in reproductive phenology.</p> / <p>Äpplet faller inte långt från trädet. Men varför? Den biologiska mångfalden präglas i stor utsträckning av fylogenetiskt bevarade karaktärsdrag; närbesläktade arter är lika. Det pågår en diskussion bland evolutionsbiologer om i vilken utsträckning denna konservatism är ett resultat av naturlig selektion eller av en begränsad anpassningsförmåga.</p><p>Denna avhandling diskuterar begreppet evolutionära begränsningar i relation till den reproduktiva fasen hos växter. I fokus ligger särskilt evolutionen av bärliknande (endozoochora) frukter respektive evolutionen av säsongsmässiga mönster (fenologi) för blomning och fruktsättning. Avhandlingen är hierarkiskt organiserad så att olika delstudier gjordes på olika skalnivåer: fenologi- och fruktevolution analyserades för fröväxter respektive gömfröiga växter; inom brakvedssläktena Rhamnus och Frangula (Rhamnaceae); samt för en lokal population av brakved (Frangula alnus). Populationsstudien baserades på såväl experimentella som deskriptiva data, medan övriga studier i huvudsak genomfördes med fylogenetisk komparativ metodik baserade på litteraturdata. Som en del av de komparativa studierna rekonstruerades fylogenetiska hypoteser för Rhamnus s.l. utifrån DNA-sekvenser (ITS, trnL-F), vilka gav stöd för att Frangula och Rhamnus är monofyletiska systersläkten.</p><p>I en biogeografiskt kontrollerad fylogenetisk kontrastanalys upptäcktes inga skillnader i artrikedom mellan djur- och vindpollinerade växter. Bärliknande frukter verkar dock vara en betydelsefull karaktär (key innovation) i skogsmiljöer. För det första var fylogenetiska klader med bärliknande frukter mer artrika än systerklader med torra frukter. Dessutom var uppkomster av bärliknande frukt korrelerad till habitat med mer sluten vegetation.</p><p>En fylogenetisk kontrastanalys av allometriska effekter visade på en positiv korrelation mellan växtindividens och fruktens storlek inom Rhamnus s.l. Upprepade analyser av dessa allometriska samband i olika monofyletiska subklader inom Rhamnus s.l. indikerar dock att allometri, såsom evolutionär begränsning, är svag. I populationsstudien av F. alnus var fruktstorlek positivt korrelerad till såväl fröantal som frövikt. En generell hypotes utifrån denna avhandling blir därför att frugivorer (de djur som äter bären och därmed sprider växtens frön) utövar ett starkt selektionstryck på växter som koloniserar ett habitat där fröstorlek är adaptivt, eftersom fruktstorlek påverkar frugivorernas (ssk. fåglars) sätt att hantera frukten.</p><p>Reproduktiv fenologi var korrelerad till fruktsättning, fröspridning och fröstorlek hos F. alnus. Detta stödjer tanken att evolutionen av blomnings- och fruktsättningstider begränsas av avvägningar (trade-offs) och partiella beroenden mellan blomning-, fruktsättnings-, spridnings och rekryteringsfaserna. Inomartsvariation i blomningstid för F. alnus respektive getapel (Rhamnus cathartica) över en nordeuropeisk latitudinell transekt var inte korrelerad till säsongslängd, vilket antyder att utvecklingen av blomningstider är evolutionärt begränsad. Å andra sidan påvisades en sådan korrelation mellan blomningstid och säsongslängd i en mellanartsstudie inom Rhamnus s.l. Den fylogenetiska kontrastanalysen mellan djur- och vindpollinerade växter visade också på adaptiva skillnader i fenologi. Blomningstiderna inom en klad med biotiskt pollinerade arter var mer åtskilda än inom den vindpollinerade systerkladen, och den välkända iakttagelsen att vindpollinerade träd blommor tidigare på säsongen fick även stöd i ett fylogenetiskt komparativt perspektiv.</p>

Conservatism is a central theme of organismic evolution. Related species share characteristics due to their common ancestry. Some concern have been raised among evolutionary biologists, whether such conservatism is an expression of natural selection or of a constrained ability to adapt. This thesis explores adaptations and constraints within the plant reproductive phase, particularly in relation to the evolution of fleshy fruit types (berries, drupes, etc.) and the seasonal timing of flowering and fruiting. The different studies were arranged along a hierarchy of scale, with general data sets sampled among seed plants at the global scale, through more specific analyses of character evolution within the genus Rhamnus s.l. L. (Rhamnaceae), to descriptive and experimental field studies in a local population of Frangula alnus (Rhamnaceae). Apart from the field study, this thesis is mainly based on comparative methods explicitly incorporating phylogenetic relationships. The comparative study of Rhamnus s.l. species included the reconstruction of phylogenetic hypotheses based on DNA sequences. Among geographically overlapping sister clades, biotic pollination was not correlated with higher species richness when compared to wind pollinated plants. Among woody plants, clades characterized by fleshy fruit types were more species rich than their dry-fruited sister clades, suggesting that the fleshy fruit is a key innovation in woody habitats. Moreover, evolution of fleshy fruits was correlated with a change to more closed (darker) habitats. An independent contrast study within Rhamnus s.l. documented allometric relations between plant and fruit size. As a phylogenetic constraint, allometric effects must be considered weak or non-existent, though, as they did not prevail among different subclades within Rhamnus s.l. Fruit size was correlated with seed size and seed number in F. alnus. This thesis suggests that frugivore selection on fleshy fruit may be important by constraining the upper limits of fruit size, when a plant lineage is colonizing (darker) habitats where larger seed size is adaptive. Phenological correlations with fruit set, dispersal, and seed size in F. alnus, suggested that the evolution of reproductive phenology is constrained by trade-offs and partial interdependences between flowering, fruiting, dispersal, and recruitment phases. Phylogenetic constraints on the evolution of phenology were indicated by a lack of correlation between flowering time and seasonal length within Rhamnus cathartica and F. alnus, respectively. On the other hand, flowering time was correlated with seasonal length among Rhamnus s.l. species. Phenological differences between biotically and wind pollinated angiosperms also suggested adaptive change in reproductive phenology. / Äpplet faller inte långt från trädet. Men varför? Den biologiska mångfalden präglas i stor utsträckning av fylogenetiskt bevarade karaktärsdrag; närbesläktade arter är lika. Det pågår en diskussion bland evolutionsbiologer om i vilken utsträckning denna konservatism är ett resultat av naturlig selektion eller av en begränsad anpassningsförmåga. Denna avhandling diskuterar begreppet evolutionära begränsningar i relation till den reproduktiva fasen hos växter. I fokus ligger särskilt evolutionen av bärliknande (endozoochora) frukter respektive evolutionen av säsongsmässiga mönster (fenologi) för blomning och fruktsättning. Avhandlingen är hierarkiskt organiserad så att olika delstudier gjordes på olika skalnivåer: fenologi- och fruktevolution analyserades för fröväxter respektive gömfröiga växter; inom brakvedssläktena Rhamnus och Frangula (Rhamnaceae); samt för en lokal population av brakved (Frangula alnus). Populationsstudien baserades på såväl experimentella som deskriptiva data, medan övriga studier i huvudsak genomfördes med fylogenetisk komparativ metodik baserade på litteraturdata. Som en del av de komparativa studierna rekonstruerades fylogenetiska hypoteser för Rhamnus s.l. utifrån DNA-sekvenser (ITS, trnL-F), vilka gav stöd för att Frangula och Rhamnus är monofyletiska systersläkten. I en biogeografiskt kontrollerad fylogenetisk kontrastanalys upptäcktes inga skillnader i artrikedom mellan djur- och vindpollinerade växter. Bärliknande frukter verkar dock vara en betydelsefull karaktär (key innovation) i skogsmiljöer. För det första var fylogenetiska klader med bärliknande frukter mer artrika än systerklader med torra frukter. Dessutom var uppkomster av bärliknande frukt korrelerad till habitat med mer sluten vegetation. En fylogenetisk kontrastanalys av allometriska effekter visade på en positiv korrelation mellan växtindividens och fruktens storlek inom Rhamnus s.l. Upprepade analyser av dessa allometriska samband i olika monofyletiska subklader inom Rhamnus s.l. indikerar dock att allometri, såsom evolutionär begränsning, är svag. I populationsstudien av F. alnus var fruktstorlek positivt korrelerad till såväl fröantal som frövikt. En generell hypotes utifrån denna avhandling blir därför att frugivorer (de djur som äter bären och därmed sprider växtens frön) utövar ett starkt selektionstryck på växter som koloniserar ett habitat där fröstorlek är adaptivt, eftersom fruktstorlek påverkar frugivorernas (ssk. fåglars) sätt att hantera frukten. Reproduktiv fenologi var korrelerad till fruktsättning, fröspridning och fröstorlek hos F. alnus. Detta stödjer tanken att evolutionen av blomnings- och fruktsättningstider begränsas av avvägningar (trade-offs) och partiella beroenden mellan blomning-, fruktsättnings-, spridnings och rekryteringsfaserna. Inomartsvariation i blomningstid för F. alnus respektive getapel (Rhamnus cathartica) över en nordeuropeisk latitudinell transekt var inte korrelerad till säsongslängd, vilket antyder att utvecklingen av blomningstider är evolutionärt begränsad. Å andra sidan påvisades en sådan korrelation mellan blomningstid och säsongslängd i en mellanartsstudie inom Rhamnus s.l. Den fylogenetiska kontrastanalysen mellan djur- och vindpollinerade växter visade också på adaptiva skillnader i fenologi. Blomningstiderna inom en klad med biotiskt pollinerade arter var mer åtskilda än inom den vindpollinerade systerkladen, och den välkända iakttagelsen att vindpollinerade träd blommor tidigare på säsongen fick även stöd i ett fylogenetiskt komparativt perspektiv.