Plants with similar traits compete for resources. If related taxa share similar traits, phylogenetic relationships may predict competitive outcomes. Although plants compete for pollinators, flowers are rarely considered in community-assembly theory. I tested the hypothesis that plant communities are structured by competition for pollination. I inventoried communities at three spatial scales, measured seven flower traits, and tested the observed patterns against those generated by a null model to judge if community members were more or less similar in floral traits than expected by chance. I also measured the phylogenetic relatedness of community members to gauge trait-conservatism. Clustering of visually attractive traits suggests they promote facilitation of pollinators while over-dispersion of morphological traits suggests they partition pollinators in to avoid competition. Communities were phylogenetically even, but relatedness did not explain floral trait patterns. I suggest that flowers represent an ecological niche through which species can be sorted.

Orientador: Carlos Alfredo Joly / Dissertação (mestrado) - Universidade Estadual de Campinas, Instituto de Biologia / Made available in DSpace on 2018-08-20T21:03:27Z (GMT). No. of bitstreams: 1 Tavares_AndreadaCostaMarques_M.pdf: 1534689 bytes, checksum: 145f2e42ac5f923e0cd3fe2b46bbc9b9 (MD5) Previous issue date: 2011 / Resumo: Estudos de comunidades em biologia floral e polinização são muito importantes para o entendimento da estrutura e do funcionamento dos ecossistemas. A Floresta Atlântica da Serra do Mar possui diferentes fisionomias que refletem as diferentes comunidades ocorrentes ao longo do gradiente de altitude. Esse estudo foi realizado em Floresta Ombrófila Densa Submontana, em Ubatuba - SP e é parte integrante do Projeto Temático Biota Gradiente Funcional - FAPESP. O objetivo geral desse trabalho foi investigar a Floresta Ombrófila Densa Submontana da Mata Atlântica quanto aos atributos florais e polinização. Foram estudadas as especies angiospermas encontradas em floração durante um ano, registrando seus atributos e visitantes florais, a forma de vida e estrato, então as síndromes de polinização foram inferidas. Foram registradas 95 espécies ao todo, de 39 famílias. Fabaceae e Rubiaceae apresentaram maior numero de espécies em flor (oito espécies cada), mas a maioria das famílias apresentou poucas espécies. Os atributos florais predominantes foram: tubo e inconspícuo como tipos morfológicos; tamanho pequeno; as cores mais frequentes foram branco e amarelo; como recurso, néctar seguido de pólen. Em 26 espécies observou-se visitação de animais de vários tipos, principalmente abelhas, beija-flores e borboletas e em alguns casos a polinização pode ser confirmada. A síndrome de polinização mais frequente na comunidade foi melitofilia, seguido de ornitofilia e sistemas mistos foram observados em 21,1% das espécies. A forma de vida mais frequente foi arbórea, seguido de liana e epífita; o estrato onde se observou maior número de espécies florindo foi o dossel. As distribuições dos recursos florais e das síndromes de polinização por formas de vida, estrato e temporal foram analisadas. A maior frequencia de plantas florindo foi observada durante a estação mais quente e chuvosa, de out/2009 a jan/2010. As principais diferenças espaciais e temporais ocorreram entre as síndromes de melitofilia e ornitofilia. Essa fitofisionomia apresentou algumas peculiaridades em relação a outras fisionomias e a outros ecossistemas, sendo, portanto, de grande importância a conservação das interações ecológicas dessa formação vegetal para a manutenção da grande biodiversidade da Floresta Atlântica / Abstract: Studies on floral biology and pollination at community level are very important to understand the ecosystems structure and functioning. Atlantic Forest at Serra do Mar has different physiognomies that reflect different communities along altitudinal gradient. This study was done in Submontane Ombrophilous Dense Forest, at Ubatuba - SP and integrates the Thematic Project Biota Gradiente Funcional - FAPESP. The general purpose of this study was to investigate the Submontane Ombrophilous Dense Forest of Atlantic Forest about floral traits and pollination. The angiosperms species found flowering during one year was studied, registering its floral traits, floral visitors, life forms and strata, so the pollination syndromes were deduced. 95 species was registered of 39 families. Fabaceae e Rubiaceae had the highest number of species in bloom (eight species each), but most families had few species. Predominant floral traits were: tube and inconspicuous as morphological types; small size; most common colors were white and yellow; as resource, nectar followed by pollen. In 26 species, several types of animals were observed visiting flowers, mainly bees, hummingbirds and butterflies, and in some cases the pollination was confirmed. The most frequent pollination syndrome in the community was melittophily, followed by ornithophily and mixed systems were observed in 21.1% of the species. The most frequent life form was tree, after liana and epiphyte; the strata where was observed major number of species was canopy. Distributions of floral resources and pollination syndromes in life forms, strata and temporal was analyzed. The higher frequency of plants flowering was observed during the hot and wet season, from oct/2009 to jan/2010. The main special and temporal differences occurred between melittophily and ornithophily syndromes. This fitophysiognomy showed some particularities in relation to other physiognomies and other ecosystems, and therefore are very important the conservation of ecological interactions of this vegetation type to maintain the great biodiversity of the Atlantic Forest / Mestrado / Biologia Vegetal / Mestre em Biologia Vegetal

L'expansion des surfaces urbaines a de fortes conséquences sur la composition des communautés de plantes et de pollinisateurs. Néanmoins, les effets de l'urbanisation sur l’interdépendance entre ces deux communautés et leurs conséquences évolutives restent peu étudiés. Premièrement, nous avons cherché à identifier le long d'un gradient d'urbanisation les facteurs affectant les communautés de plantes et de pollinisateurs. Deuxièmement, nous avons testé l'existence de divergences évolutives intra-spécifiques pour les caractéristiques reproductives de plantes d'origine urbaine et rurale. Dans ce but, nous avons combiné l'analyse de données issues de programmes de sciences participatives en région de l'Ile-de-France et nous avons également mis en place une expérience de transplantation réciproque impliquant quatre espèces végétales fréquentes dans cette région. Nos résultats révèlent que la morphologie florale est le facteur le plus souvent impliqué dans l'observation des familles d'insectes sur les fleurs le long du gradient. Les familles d'insectes ayant une affinité positive aux surfaces imperméables préfèrent les corolles tubulaires, tandis que les familles ayant une affinité négative préfèrent les corolles évasées. L'urbanisation modifie la composition des communautés végétales, en sélectionnant notamment des espèces autogames et non-entomophiles. Nos résultats indiquent une perte fonctionnelle de pollinisateurs en milieu urbain. Une analyse détaillée de l’espèce Cymbalaria muralis suggère que les plantes auraient un plus grand bénéfice reproductif à allouer plus de ressources à la production de fleurs sur les sites de plantation urbain. En conséquence, les plantes d'origine urbaine ont tendance à produire plus de fleurs que celles d’origine rurale, au détriment de la production d'ovules par fleur et de la coloration des corolles. Cette divergence évolutive pourrait s'expliquer par des changements dans le comportement des pollinisateurs induits par une fragmentation accrue de l'habitat dans les zones urbaines. Cette thèse révèle que les changements induits par l'urbanisation dans l'abondance, le comportement ou les affinités florales fonctionnelles des pollinisateurs peuvent servir d'agents de sélection sur les espèces végétales spontanées. / The expansion of urban areas is known to impact the composition of plant and pollinator communities. However, the effects of urbanization on the interdependency between both communities, and the subsequent evolutionary consequences, are still poorly studied. First, we tried to identify along an urbanization gradient which factors shape plant and pollinator communities. Second, we tested the existence of intra-specific evolutionary divergences of plant reproductive characteristics between populations from urban and rural origins. For these purposes, we combined the analysis of data collected in the Ile-de-France region originating from citizen science projects, and we set up a reciprocal transplantation experiment involving four plant species commonly found in this region. We found that floral morphology was the most frequently implicated factor in the observation of insect families along the gradient. Insect families with positive affinity to impervious areas prefer tubular corollas, while families with negative affinity prefer non-tubular (open) corollas. Urbanization modifies the composition of plant communities, by selecting in particular autogamous and non-entomophilous species. Altogether, our results point towards a functional loss of pollinators along with urbanization. A detailed analysis of the species Cymbalaria muralis suggests a greater reproductive benefit in allocating more resources to flower production in urban plantation sites. As a consequence, plants from urban origin tend to produce more flowers compared to plants from rural origin, at the expense of ovule production per flower and flower coloration. This evolutionary divergence could be explained by shifts in pollinator behaviour induced by higher habitat fragmentation in urban areas. This thesis reveals that shifts in the abundance, in the behaviour or in the functional floral affinities of pollinators, induced by urbanization, could act as selection agents on spontaneous plant species.

Seleção natural divergente é a principal força que leva a diferenciação entre espécies. Nas Angiospermas, a preferencia do polinizador é considerada um dos agentes de seleção na diversificação de caracteres florais. Além disso, eles podem contribuir na delimitação de linhagens e diversificação de linhagens devido a possibilidade de reduzir fluxo gênico entre os fenótipos em simpatria, contribuindo para o isolamento reprodutivo. Nesse trabalho, nós avaliamos a interação entre padrões de variação geográfica floral e processos ecológicos associados aos polinizadores de Sinningia allagophylla. Nós amostramos traços florais e frequência de polinizadores ao longo da distribuição geográfica dessa espécie. Germinação de grãos de pólen e desenvolvimento da semente também foram amostradas para determinar a ocorrência de mecanismos de isolamento reprodutivos adicionais em ação. Nós mostramos que atributos florais de S. allagophylla apresentam um claro padrão de diferenciação geográfica. É possível classificar indivíduos dentro de três tipos florais, estando estes relacionados as vertentes de distribuição sul, noroeste e sudeste da espécie. Tipos florais intermediários ocorrem entre as regiões sudeste e o sul. Diversos grupos de polinizadores estão associados com o padrão geográficos dos atributos florais. Beija-flores estão presentes ao longo de toda a distribuição geográfica e visita tipos florais independentemente de seus atributos, mas abelhas são mais comuns nos extremos da distribuição e apresentam fidelidade a um dado tipo floral. Sobre condições de simpatria, pode haver fluxo gênico entre fenótipos distintos devido as visitas de beija-flores. No entanto, visitas seletivas de abelhas a um dos tipos florais e germinação do pólen e o desenvolvimento da semente reduzidos em indivíduos com traços florais intermediários resultam na ausência de coesão e perda dos fenótipos originais. Em conclusão, consideramos que mecanismos ecológicos associados em parte a interação planta-polinizador possam contribuir para a diferenciação das linhagens em Sinningia allagophylla. Tal informação adiciona a possibilidade de que mecanismos outros que os reconhecidamente vigentes possam contribuir para a especiação deste clado. / Divergent natural selection is a leading force of species differentiation. In flowering plants, pollinator\'s preference is considered to work as selective agent and can contribute to the diversification of floral traits. Because of pollinator role preventing gene flow under sympatric condition, they can also work as a component of reproductive isolation. Here we establish a link between geographic pattern and ecological process of differentiation in floral morphology and pollinator fidelity in Sinningia allagophylla. We sampled floral traits and pollinator frequency across the species geographic ranges. Germination of pollen grains and seed development were also sampled to determine the occurrence of additional reproductive isolation mechanisms in action. We show that floral traits of S. allagophylla present a clear pattern of geographic differentiation. It is possible to sort individuals within three floral types related to the south, southeast and northwest species range. Intermediate floral types occur between south-southeast distribution ranges. Diverse group of pollinators are associated with geographic pattern of floral traits. Hummingbirds are present across the entire geographic range and visit floral types independent of its traits, but bees are more frequent in the boundaries of distribution and show fidelity to one or other phenotype. Under sympatric condition, gene flow occurs between individuals with different floral traits due to hummingbird visits. However, bee preference of phenotypes, reduced pollen germination and seed development of individuals in the contact zones contributed to the maintenance of distinct phenotypes. In conclusion, we considered that ecological mechanisms associated with the plant-pollinator interaction are contributing partially to the differentiation of lineages in Sinningia allagophylla. This information sheds new light to the actual comprehension of speciation within this clade.

De nombreuses populations voient leurs conditions de vie modifiées par les changements globaux. Au-delà de leurs conséquences écologiques, les changements globaux peuvent également modifier les régimes de sélection des populations. Le déclin récent des pollinisateurs pourrait altérer fortement le succès reproducteur de nombreuses populations de plantes à fleurs. Cependant, ses conséquences évolutives n'ont pas été étudiées jusqu'ici. Cette thèse traite de la possibilité d'adaptation des stratégies de reproduction des plantes face aux changements globaux et plus spécifiquement face au déclin des pollinisateurs. Cette question a été abordée en deux temps. Premièrement, l'analyse de l'abondante littérature théorique et empirique sur les systèmes de reproduction des plantes et dans un moindre mesure la construction d'un modèle d'évolution des traits floraux d'attraction ont permis de clarifier des scénarios d'évolution à court terme. Deuxièmement, une approche empirique originale a été menée, consistant à comparer directement les traits de populations ancestrales et descendantes de trois espèces annuelles, à partir de la culture en jardin commun de graines anciennes et récentes en provenance de régions où des indices de déclin des pollinisateurs existent. Cette approche permet de mettre clairement en évidence l'évolution génétique. Les données existantes indiquent que le déclin des pollinisateurs peut accentuer la limitation pollinique et par conséquent augmenter la sélection sur les traits floraux. Par ailleurs, la variation génétique substantielle dans les populations suggère que l'évolution rapide des populations de plantes est possible. L'analyse de la littérature et notre étude théorique suggèrent que l'accroissement de la capacité d'autofécondation autonome ou l'accroissement de l'attraction des pollinisateurs sont deux scénarios d'évolution possibles. Deux types de réponses évolutives ressortent de nos travaux expérimentaux. D'abord, une avancée du calendrier de floraison a été retrouvée chez les trois espèces étudiées. Ce résultat souligne le rôle de l'évolution génétique, et pas seulement de la plasticité phénotypique, dans les avancées de phénologies de printemps que l'on retrouve chez de très nombreux organismes. Ensuite, contrairement aux traits de phénologie, les traits floraux ont évolué dans des directions opposées selon les espèces. Ainsi, des traits floraux a priori plus attractifs ont évolué chez une des espèces tandis que pour une autre, des traits floraux a priori moins attractifs ont évolué, mais s'accompagnent d'une meilleure capacité d'autofécondation autonome. Cette étude confirme que les traits de reproduction des plantes peuvent évoluer en quelques décennies seulement. La possibilité du sauvetage évolutif des populations, par l'évolution rapide des traits et des stratégies de reproduction des plantes, est une perspective de recherche qui découle de ces résultats. / Global change alters life conditions of numerous populations. Beyond ecological consequences, global change can also modify selection regimes in populations. While the recent pollinator decline may specifically affect the reproductive success of flowering plants, its evolutionary consequences have not been studied yet. This thesis deals with the possibility of adaptation of plant reproductive strategies under global change and more specifically under pollinator decline. This question was addressed in two steps. First, the analysis of the extensive literature on plant mating systems, and, to a lesser extent, the construction of a model for the evolution of attractive floral traits, allowed us to clarify evolutionary scenarios at short-time scales. Second, we conducted an original empirical approach, consisting in the direct comparison of ancestral and descendant populations by re-growing old and recent seeds under identical conditions. This approach allowed us to test whether genetic evolution of reproductive traits occurred in the context of pollinator decline for three annual plant species. Data from the literature indicates that pollinator decline likely increases pollen limitation and thus selection on floral traits. Moreover, the substantial genetic variation within populations suggests that rapid evolution is possible. Increased autonomous selfing or increased pollinator attraction are two possible routes of plant adaptation to pollinator decline emerging from the analysis of the literature and from our theoretical study. Our empirical work brings out two types of evolutionary trends. Firstly, earlier flowering phenology was found in all three studied species. This result shows that genetic evolution, not only phenotypic plasticity; certainly contribute to the spring phenological advancements reported for numerous species. Secondly, unlike phenological traits, floral traits evolved in opposite directions depending on the species. Showy floral traits evolved in a species while joint evolution of autonomous selfing with a reduction of floral attractiveness seemed to evolve in another species. This study shows that plant reproductive traits can evolve in a few decades. Whether or not rapid evolution of plant reproductive traits can act as an evolutionary rescue for threatened populations is a research question that arises from these results.

Made available in DSpace on 2016-06-02T19:32:13Z (GMT). No. of bitstreams: 1 DissFQM.pdf: 973806 bytes, checksum: e915584ac61462d2e8ef3840ece7c9a9 (MD5) Previous issue date: 2005-02-25 / Universidade Federal de Sao Carlos / The Cerrado Domain occupied originally 23% of the Brazilian territory (ca. 2 million km2), especially in the Central Plateau, being the second largest phytogeographic province of Brazil. The cerrado vegetation is not uniform in physiognomy, ranging from grassland to tall woodland, but most of its physiognomies lie within the range defined as tropical savanna. Is estimated that 3,000 to 7,000 vascular plant species occur in this vegetation type, from which 1,000 to 2,000 belong to the woody component. Different authors have attempted to use reproductive features to explain the general patterns of diversity and community structure found in tropical woodlands with the underlying idea that plant diversity and spatial distribution is dependent on reproductive processes. Studies on the reproductive biology of cerrado plant species have shown a great diversity of pollination systems, similar to those found in Neotropical forests. The data emerging for the reproductive biology of plants have important consequences for conservation and understanding of the organization of cerrado communities. We sampled five cerrado fragments in the Brazilian Central Plateau, in which we sampled woody individuals. Using the floristic data of all our field trips, we sampled 2,280 individuals, representing 121 species and 38 families. The richest families were Fabaceae and Myrtaceae, and Davilla elliptica A. St-Hill and Myrcia bella Triana were the best represented species. Most species presented open flowers, with diurnal anthesis, pale colors and with pollen as floral reward. In the cerrado vegetation, species with flowers visited mainly by bees and small insects were the main groups ecologically related to the pollination. Of the 121 species, 65 were pollinated mainly by bees; 30, by small insects; 15, by moths; five, by bats; three, by beetles; two, by hummingbirds; and one, by wind. The ordination analysis of floral characteristics and plant species showed that there was a grouping of species with some pollination systems, for which inferences based on floral characteristics are recommended, such as the species pollinated by bats, moths, and birds. On the other hand, for the species pollinated mainly by bees and small insects, these inferences are not recommended due their great dispersion throughout ordination axes and large overlapping. These dispersion and overlapping occurred probably due the absence of specificity between plants and pollinators. For four of the five pollination systems with at least ten individuals, we found no significant variation in relation to distance from edge, except for plants pollinated by beetles, for which there was a decrease in the frequency to toward the fragment interior. Similarly, we only found significant variation in relation to the height for plants pollinated by bats, for which there was an increase of the frequency with the height of the trees. In general, we found no horizontal and vertical variations in the pollination systems, contrary to what was found in forests and, probably, as consequence of the more open physiognomy of the cerrado fragments. / O Domínio do Cerrado ocupava originalmente cerca de 23% do território brasileiro (aproximadamente 2 milhões de km2), especialmente no Planalto Central, sendo a segunda maior província fitogeográfica do Brasil. A vegetação de cerrado não é uniforme na sua fisionomia, variando desde campo limpo a cerradão, mas a maior parte das fisionomias se enquadra na definição de savana . É estimado que ocorra de 3.000 a 7.000 espécies de plantas vasculares nesse tipo de vegetação, das quais de 1.000 a 2.000 espécies pertencem ao componente arbustivoarbóreo. Diferentes autores tentaram usar características reprodutivas para explicar os padrões gerais de diversidade e estrutura de comunidade encontrados em florestas tropicais, com a idéia de que a diversidade das plantas e a distribuição espacial são dependentes de processos reprodutivos. Estudos na biologia reprodutiva de espécies de planta de cerrado mostraram uma grande diversidade de sistemas de polinização, semelhantes àqueles encontrados em florestas neotropicais. Os dados que emergem para a biologia reprodutiva de plantas têm conseqüências importantes para conservação e entendimento da organização das comunidades de cerrado. Amostramos cinco fragmentos de cerrado sensu stricto no Planalto Central brasileiro, em que amostramos os indivíduos arbustivo-arbóreos. Usando os dados florísticos de todas as nossas coletas, nós amostramos 2.280 indivíduos, representando 121 espécies e 38 famílias. As famílias mais ricas foram Fabaceae e Myrtaceae, sendo Davilla elliptica A. St-Hill e Myrcia bella Triana as espécies mais bem representadas. A maioria das espécies apresentou flores abertas, com antese diurna, cores claras e pólen como recompensa floral. Na vegetação de cerrado, as espécies com flores visitadas principalmente por abelhas e também pelos insetos pequenos formaram os principais grupos ecologicamente relacionados com a polinização. Das 121 espécies, 65 foram polinizadas principalmente por abelhas; 30 por insetos pequenos; 15 por mariposas; cinco por morcegos; três por besouros; dois por beija-flores e um pelo vento. A análise de ordenação dos caracteres florais e das espécies vegetais mostrou que houve um agrupamento entre espécies com alguns sistemas de polinização, para os quais inferências baseadas em caracteres florais são recomendadas, como as espécies polinizadas por morcegos, mariposas e aves. Já com relação às espécies polinizadas principalmente por abelhas e insetos pequenos, essas inferências baseadas em caracteres florais não são recomendadas devido à grande dispersão e sobreposição entre essas duas classes. A grande dispersão e sobreposição das classes de abelhas e insetos pequenos ocorreram provavelmente devido à ausência de especificidade nas relações planta-polinizador. Para quatro dos cinco sistemas de polinização com pelo menos dez indivíduos, nós não encontramos nenhuma variação significativa em relação à distância da borda do fragmento, exceto para as plantas polinizadas por besouros, para as quais houve uma diminuição na freqüência em direção ao interior do fragmento. De maneira semelhante, encontramos variação significativa em relação à altura somente para plantas polinizadas por morcegos, para as quais houve um aumento da freqüência com a altura das árvores. Em geral, não encontramos variações horizontais e verticais nos sistemas de polinização, ao contrário do que foi encontrado em florestas, provavelmente, como conseqüência da fisionomia mais aberta dos fragmentos de cerrado.

Seleção natural divergente é a principal força que leva a diferenciação entre espécies. Nas Angiospermas, a preferencia do polinizador é considerada um dos agentes de seleção na diversificação de caracteres florais. Além disso, eles podem contribuir na delimitação de linhagens e diversificação de linhagens devido a possibilidade de reduzir fluxo gênico entre os fenótipos em simpatria, contribuindo para o isolamento reprodutivo. Nesse trabalho, nós avaliamos a interação entre padrões de variação geográfica floral e processos ecológicos associados aos polinizadores de Sinningia allagophylla. Nós amostramos traços florais e frequência de polinizadores ao longo da distribuição geográfica dessa espécie. Germinação de grãos de pólen e desenvolvimento da semente também foram amostradas para determinar a ocorrência de mecanismos de isolamento reprodutivos adicionais em ação. Nós mostramos que atributos florais de S. allagophylla apresentam um claro padrão de diferenciação geográfica. É possível classificar indivíduos dentro de três tipos florais, estando estes relacionados as vertentes de distribuição sul, noroeste e sudeste da espécie. Tipos florais intermediários ocorrem entre as regiões sudeste e o sul. Diversos grupos de polinizadores estão associados com o padrão geográficos dos atributos florais. Beija-flores estão presentes ao longo de toda a distribuição geográfica e visita tipos florais independentemente de seus atributos, mas abelhas são mais comuns nos extremos da distribuição e apresentam fidelidade a um dado tipo floral. Sobre condições de simpatria, pode haver fluxo gênico entre fenótipos distintos devido as visitas de beija-flores. No entanto, visitas seletivas de abelhas a um dos tipos florais e germinação do pólen e o desenvolvimento da semente reduzidos em indivíduos com traços florais intermediários resultam na ausência de coesão e perda dos fenótipos originais. Em conclusão, consideramos que mecanismos ecológicos associados em parte a interação planta-polinizador possam contribuir para a diferenciação das linhagens em Sinningia allagophylla. Tal informação adiciona a possibilidade de que mecanismos outros que os reconhecidamente vigentes possam contribuir para a especiação deste clado. / Divergent natural selection is a leading force of species differentiation. In flowering plants, pollinator\'s preference is considered to work as selective agent and can contribute to the diversification of floral traits. Because of pollinator role preventing gene flow under sympatric condition, they can also work as a component of reproductive isolation. Here we establish a link between geographic pattern and ecological process of differentiation in floral morphology and pollinator fidelity in Sinningia allagophylla. We sampled floral traits and pollinator frequency across the species geographic ranges. Germination of pollen grains and seed development were also sampled to determine the occurrence of additional reproductive isolation mechanisms in action. We show that floral traits of S. allagophylla present a clear pattern of geographic differentiation. It is possible to sort individuals within three floral types related to the south, southeast and northwest species range. Intermediate floral types occur between south-southeast distribution ranges. Diverse group of pollinators are associated with geographic pattern of floral traits. Hummingbirds are present across the entire geographic range and visit floral types independent of its traits, but bees are more frequent in the boundaries of distribution and show fidelity to one or other phenotype. Under sympatric condition, gene flow occurs between individuals with different floral traits due to hummingbird visits. However, bee preference of phenotypes, reduced pollen germination and seed development of individuals in the contact zones contributed to the maintenance of distinct phenotypes. In conclusion, we considered that ecological mechanisms associated with the plant-pollinator interaction are contributing partially to the differentiation of lineages in Sinningia allagophylla. This information sheds new light to the actual comprehension of speciation within this clade.

Les Angiospermes forment un clade connu pour sa grande diversité d’espèces et une répartition inégale de cette diversité en son sein. Ce travail se concentre sur la reconstruction d’états ancestraux de caractères floraux sur les phylogénies dans le but d’étudier leur impact sur la diversification. En premier lieu, nous nous sommes focalisés sur deux innovations-clés potentielles chez les Proteaceae, la symétrie bilatérale du périanthe et la présence dans les hotspots à climat méditerranéen. L’utilisation sur une phylogénie de modèles de diversification dépendants des états de caractère a permis de montrer que le premier état caractère n’a pas d’impact significatif sur la diversification des espèces dans ce groupe, contrairement au second. Ensuite, nous avons reconstruit les états ancestraux de la symétrie du périanthe à l’échelle de toutes les Angiospermes, avec une stratégie d’échantillonnage permettant de refléter toutes les transitions de ce caractère. Nous avons mis en évidence un minimum de 130 origines de la symétrie bilatérale, suivies de 69 réversions vers la symétrie radiaire. Enfin, la même méthode a été étendue à l’étude de quatre autres caractères du périanthe. Différents modèles de reconstruction d’états ancestraux ont été utilisés sur une phylogénie de 1232 espèces pour examiner l’influence des changements de modèle sur les états ancestraux reconstruits. Nos résultats montrent que les changements de modèles de reconstruction ont une influence sur l’état de certains nœuds seulement, jamais sur l’ensemble. Les différents résultats trouvés pour la symétrie, caractère commun aux trois chapitres, révèlent que notre méthode d’échantillonnage présente l’inconvénient d’estimer des taux de transition trop élevés pour donner des résultats concluants avec la méthode du maximum de vraisemblance. Les résultats sont en revanche beaucoup moins biaisés lorsque l’on examine plusieurs caractères simultanément. / Angiosperms are a clade known for its great species diversity and the uneven distribution of this diversity among its lineages. This work focuses on the ancestral state reconstruction of floral characters on phylogenies in the purpose of studying their impact on diversification. We first focused on two potential key innovations in Proteaceae, bilateral perianth symmetry and presence in Mediterranean-climate hotspots. Using character state dependent diversification models, we found that the first character state did not have any significant impact on species diversification in this group, contrary to the second one. We then reconstructed ancestral states for perianth symmetry in angiosperms as a whole, using a sampling strategy aimed at capturing all of the transitions in this character. We found a minimum of 130 origins of bilateral symmetry, followed by 69 subsequent reversals to radial symmetry. Lastly, the same approach was extended to the study of four other perianth characters. Different ancestral state reconstruction models were used on a phylogenetic tree of 1232 species to test the influence of model changes on reconstructed ancestral states. Our results show that changes in reconstruction models have an impact on the inferred ancestral state of some nodes, but not all of them. The various results obtained for symmetry, a character shared among the three chapters, indicate that our sampling method has the drawback of estimating transition rates too high to give conclusive results with maximum likelihood. On the other hand, results are much less biased when several characters are examined simultaneously.

L’environnement a un effet considérable sur les végétaux par différents facteurs abiotiques (climat, sol, urbanisation) ou biotiques (pollinisateurs, herbivores). Les fortes pressions de sélection exercées par ces facteurs sur certains traits phénotypiques aboutissent souvent à des adaptations chez les plantes. Les pollinisateurs exercent une pression de sélection sur les traits floraux qui résultent en des adaptations convergentes que l’on appelle syndromes de pollinisation. L’étude de ses syndromes, mais aussi de la performance de pollinisation de chaque visiteur, permet de mieux comprendre l’intensité de cette pression de sélection ainsi que l’évolution florale. Dans cette thèse, nous nous sommes appuyés sur l’utilisation de deux modèles d’études végétaux : la famille des Gesneriaceae des Antilles et l’Impatiente du Cap. Ces deux modèles, de par leurs stratégies de pollinisation et leurs variations florales, sont appropriés pour étudier l’adaptation florale aux pollinisateurs. Nous avons ainsi pu tester 1) si la forme florale est expliquée par la forme du bec des colibris pollinisateurs chez les Gesneriaceae antillaises ; 2) si l’espèce Rhytidophyllum bicolor Urb. est généraliste en pollinisation et si ses différents types fonctionnels de pollinisateurs ont une performance de pollinisation similaire ; 3) si l’urbanisation affecte la forme florale de l’espèce Impatiens capensis Meerb., à travers des changements dans les communautés de pollinisateurs. Pour tester ces hypothèses, des approches de morphométries géométriques ont été utilisées (hypothèses 1 et 3), ainsi que des observations de pollinisation in situ et la mesure du taux de visite (hypothèses 2 et 3). La mesure de performance de pollinisation a été réalisé via le comptage des grains de pollen déposés sur le stigmate après chaque visite, et à la mesure du taux de visite. Enfin, des tests statistiques (ANOVA, régressions linéaires) et analyses multivariées (analyses de redondance, analyse en composantes principales) ont été réalisées sur les données obtenues pour chaque étude. Les résultats de la première étude montrent une corrélation positive entre la longueur de la corolle de fleurs spécialistes aux colibris et la longueur du bec des pollinisateurs. Bien que moins significatifs, les résultats des comparaisons de formes globales, obtenues par application de morphométrie géométrique, indiquent que cette approche est prometteuse pour ce genre d’analyse. Nous avons démontré que la forme florale des fleurs généralistes est impactée par les colibris pollinisateurs, bien que d’une manière différente des spécialistes. 4 Pour l’hypothèse 2, la prédiction de stratégie de pollinisation de l’espèce Rhytidophyllum bicolor a été validée, puisque cette plante est pollinisée par des colibris, chauves-souris et abeilles. Cependant, face au déclin des populations de colibris après le passage de l’ouragan Matthew sur Haïti, seules les performances de pollinisation des abeilles et des chauves-souris ont pu être mesurées. Les résultats montrent que les chauves-souris sont des pollinisateurs efficaces et conséquents, bien que la performance des abeilles ne soit pas négligeable. Il a ainsi pu être mis de l’avant que la stratégie de pollinisation généraliste semble être un avantage pour les plantes présentes dans les zones sujettes aux fluctuations de populations de pollinisateurs, comme cela peut souvent être le cas sur les îles à la suite du passage d’un ouragan. La troisième étude montre que l’urbanisation n’a pas d’effet direct sur la forme florale de l’Impatiente du Cap, mais qu’elle a des effets indirects significatifs via les changements causés sur les communautés de pollinisateurs. Sur les six sites échantillonnées les pollinisateurs principaux, Bombus sp. Latreille et Apis mellifera Linnaeus, sont les mêmes. Cependant les taux de diversité obtenus montrent une variation entre les sites, due à la présence de certaines espèces de pollinisateurs occasionnels dans certains sites et pas dans d’autres. Ces taux ne sont pas plus faibles dans les sites les plus urbains. Les résultats indiquent que certaines formes florales sont associées à des espèces de pollinisateurs particulières. Les différentes espèces de pollinisateurs ayant une pression de sélection différente sur les traits floraux, l’urbanisation a ainsi un impact indirect sur la forme florale chez l’espèce étudiée. À travers trois études différentes, cette thèse a mis en avant l’impact que l’environnement peut avoir sur les traits floraux, de manière indirecte, via les pollinisateurs. Alors que la deuxième et troisième étude ont montré la pression exercée par les pollinisateurs sur les traits floraux dans différents cas de perturbations de l’habitat, la première étude a permis de mieux comprendre l’adaptation remarquable des fleurs à leurs pollinisateurs, même pour des espèces généralistes en pollinisation. / The environment has a considerable effect on plants through various abiotic (climate, soil, urbanization) or biotic (pollinators, herbivores) factors. The strong selection pressures exerted by these factors on phenotypic traits often results in adaptations. Pollinators exert selection pressure on floral traits that result in converging adaptations called pollination syndromes. The study of syndromes, as well as the pollination performance of each floral visitor, allows us to better understand the intensity of a/biotic selection pressure and floral evolution. In this thesis, we relied on two plant models: the Gesneriaceae family in the Antilles, and the common Jewelweed, Impatiens capensis Meerb. Due to their pollination strategies and their floral variation, both of these models are suitable for studying floral adaptation to biotic factors, specifically, their pollinators. We were thus able to test the following hypotheses: 1) whether the floral form is explained by the beak shape of pollinating hummingbirds in West Indies Gesnericeae; 2) whether the Gesnericeae Rhytidophyllum bicolor Urb. has a generalist strategy for pollination and whether their different functional types of pollinators have similar pollination performances; 3) whether urbanization affects the floral form of the common Jewelweed, through changes in pollinator communities. To test these hypotheses, geometric morphometric approaches were used (hypotheses 1 and 3), as well as in situ pollination observations, and estimation of the visitation rate (hypotheses 2 and 3). Pollination performance was measured by counting pollen grains deposited on the stigma after each visit, and by measuring the visitation rate. Finally, statistical tests (ANOVA, linear regressions) and multivariate analyses (redundancy analysis, principal component analysis) were carried out on the data obtained to test each hypothesis. The first results show a positive correlation between the length of the corolla of flowers specialized for hummingbirds and the beak length of pollinators. Although less significant, our results of the comparisons of global shapes, obtained by applying geometric morphometry, indicate that this approach is promising for this kind of analysis. We show that the floral form of generalist flowers is impacted by pollinating hummingbirds, albeit in a different way from specialists. For the second hypothesis, our prediction that R. bicolor has a generalist pollination strategy was validated, since this plant is pollinated by hummingbirds, bats and bees. However, faced with the decline in hummingbird populations after Hurricane Matthew hit Haiti in 2016, only the pollination performance of bees and bats could be measured. Our results show that bats are efficient and consistent pollinators of R. bicolor, although the performance of bees is not negligible. Thus, it has been possible to put forward that the generalist pollination strategy seems to be an advantage for plants present in areas subject to fluctuations in their pollinator populations, as can often be the case on hurricane-prone islands. The third study shows that urbanization does not have a direct effect on the flower form of common Jewelweed, but that urbanization does have significant indirect effects through changes caused on pollinator communities. At the six sites sampled, the main pollinators, Bombus sp. Latreille and Apis mellifera Linnaeus, are the same. However, diversity rates show variation between sites, due to the presence of certain species of occasional pollinators in some sites and not in others. These rates are not lower in the most urban sites. Our results indicate that certain floral forms are associated with particular pollinator species. Since different pollinator species have different selection pressures on floral traits, urbanization has an indirect impact on the floral form in the species studied. Through three different studies, this thesis highlighted the impact that the environment can have on floral traits, indirectly, via pollinators. While the second and third studies showed the pressure exerted by pollinators on floral traits in different cases of habitat disturbance, the first study helped to better highlight the remarkable adaptation of flowers to their pollinators, even for generalist species in pollination.

Flowering plants display remarkable phenotypic diversity, especially in reproductive structures, much of which is thought to be associated with pollination by animals. Pollination syndromes are collections of floral traits (e.g. flower colour, shape, odour) that are associated with a plant attracting particular functional groups of animal pollinators. We explored the extent to which traits associated with the moth pollination syndrome translated into pollination by moths in the Pacific coast dune endemic Abronia umbellata and found mixed results: in one year of study, there was no difference in seed set by day- vs. night-pollinated inflorescences, but in another year of study, night-pollinated inflorescences set significantly more seed than those pollinated during the day. We integrate this work with tests of pollen and resource limitation of seed production and with seed set surveys of natural populations to address proximate and ultimate causes of low seed set, finding low rates of pollinator visitation, high pollen limitation of seed production in all populations studied, and no evidence of endogenous resource limitation of seed production. We propose that “excess” flowers may be functionally male, serving to increase outcross siring success. The transition from self-incompatibility and obligate outcrossing to self-compatibility and predominant selfing is the most common evolutionary transition among the flowering plants and traits associated with outcrossing may become reduced across such shifts, potentially through the action of natural selection, especially if pollinators are also herbivores, or if the signals that pollinators use to locate flowers are also used by herbivores. We examined the reduction of attractive visual and olfactory floral traits in A. umbellata across a shift from outcrossing to selfing and found a reduction of all floral traits considered. We found that floral volatile emissions were reduced more strongly than flower size or floral display (number of flowers per inflorescence), but there was no evidence of an ecological cost associated with conspicuousness: we did not find reduced leaf herbivory among selfers relative to outcrossers. / Thesis (Master, Biology) -- Queen's University, 2012-08-30 19:46:23.663