Grazing management is very popular nowadays and number of sites with grazing animals with the aim of restoring the sites is steadily growing. All the effects of grazing on grazed grasslands are not elucidated yet. This thesis focused mainly on the contribution of zoochory to restoration of species rich grasslands on stands cleared from Robinia pseudoacacia and stands cleared from Prunus spinosa brushwood. For the purpose of monitoring changes in vegetation, permanent plots have been established. In order to identify sources of new species occurring in permanent plots, inventory of species growing in neighbourhood of the permanent plots has been done, samples of soil seed bank, sheep buttons and seeds from sheep wool have been germinated in a greenhouse. Monitoring of permanent plots showed reduced regrowth of R. pseudoacacia and P. spinosa. We have also found that greater changes in species composition occurred in more degraded stands than in stands better-preserved. Germinating experiments proved soil seed bank being mainly the image of aboveground vegetation with minor importance to restoration of species rich grasslands. On the other hand sheep seem to be of great use for dispersal of seeds both by epizoochory and endozoochory when walking between different stands. This finding is of great importance for...

Understanding dispersal and gene flow in human-modified landscapes is crucial for effective conservation. Seed dispersal governs colonization, recruitment, and distribution of plant species, whereas both pollen and seed dispersal determine gene flow among populations. This PhD thesis tests the effect of rotational shepherding on seed dispersal and gene flow in fragmented calcareous grasslands. Calcareous grasslands (Gentiano-Koelerietum pyramidatae vegetation) in Central Europe are semi-natural communities traditionally used for rotational grazing that experienced a decline of plant species during the 20th century due to abandonment of shepherding. This PhD profits from a management project started in 1989 in Bavaria, Germany to reconnect previously abandoned calcareous grasslands in three non-overlapping shepherding systems. Two vegetation surveys in 1989 and 2009 revealed colonizations in previously abandoned grasslands reconnected by shepherding. First, I propose a comprehensive approach to identify determinants of community-level patch colonization rates based on 48 habitat specialist plants by testing competing models of pre-dispersal and dispersal effects and accounting for post-dispersal effects. Mean source patch species occupancy in 1989, and structural elements in focal patches related to establishment explained community-level patch colonization rates. Secondly, by adapting the community analysis to all 31 individual species of the same community with sufficient data, I corroborate the role of shepherding to support dispersal for a range of species, even if they lack seed morphological traits related to zoochory. Thirdly, for the habitat specialist Dianthus carthusianorum, I genotyped 1,613 individuals from 64 populations at eleven microsatellites to test the effect of dispersal by sheep on spatial genetic structure at the landscape scale. Genetic distances between grazed patches of the same herding system were related to distance along herding routes, whereas ungrazed patches showed isolation by geographic distance. Lastly, within individual grassland patches, shepherding significantly decreases the degree of relatedness among neighboring individuals (kinship structure) and increases genetic diversity. My thesis contributes towards understanding the effects of zoochory on spatial dynamics in plant populations across scales.

Understanding dispersal and gene flow in human-modified landscapes is crucial for effective conservation. Seed dispersal governs colonization, recruitment, and distribution of plant species, whereas both pollen and seed dispersal determine gene flow among populations. This PhD thesis tests the effect of rotational shepherding on seed dispersal and gene flow in fragmented calcareous grasslands. Calcareous grasslands (Gentiano-Koelerietum pyramidatae vegetation) in Central Europe are semi-natural communities traditionally used for rotational grazing that experienced a decline of plant species during the 20th century due to abandonment of shepherding. This PhD profits from a management project started in 1989 in Bavaria, Germany to reconnect previously abandoned calcareous grasslands in three non-overlapping shepherding systems. Two vegetation surveys in 1989 and 2009 revealed colonizations in previously abandoned grasslands reconnected by shepherding. First, I propose a comprehensive approach to identify determinants of community-level patch colonization rates based on 48 habitat specialist plants by testing competing models of pre-dispersal and dispersal effects and accounting for post-dispersal effects. Mean source patch species occupancy in 1989, and structural elements in focal patches related to establishment explained community-level patch colonization rates. Secondly, by adapting the community analysis to all 31 individual species of the same community with sufficient data, I corroborate the role of shepherding to support dispersal for a range of species, even if they lack seed morphological traits related to zoochory. Thirdly, for the habitat specialist Dianthus carthusianorum, I genotyped 1,613 individuals from 64 populations at eleven microsatellites to test the effect of dispersal by sheep on spatial genetic structure at the landscape scale. Genetic distances between grazed patches of the same herding system were related to distance along herding routes, whereas ungrazed patches showed isolation by geographic distance. Lastly, within individual grassland patches, shepherding significantly decreases the degree of relatedness among neighboring individuals (kinship structure) and increases genetic diversity. My thesis contributes towards understanding the effects of zoochory on spatial dynamics in plant populations across scales.

Coordenação de Aperfeiçoamento de Pessoal de Nível Superior / No Brasil, a Família Didelphidae é composta por 54 espécies com ampla distribuição por diferentes habitats e um padrão de consumo alimentar que pode variar desde espécies mais frugívoras até as mais insetívoras/carnívoras. O objetivo deste estudo foi avaliar a relação entre a dieta de sete espécies de didelfídeos (Caluromys philander, Didelphis albiventris, Gracilinanus agilis, G. microtarsus, Marmosa (Micoureus) paraguayana, Marmosops incanus e Metachirus nudicaudatus) e a seleção de recursos alimentares (artrópodes e frutos) disponíveis. O estudo foi realizado entre novembro de 2009 e outubro de 2011, em uma área de mata ciliar de cerrado no Parque Estadual do Rio Preto, Minas Gerais, Brasil. Artrópodes, frutos, flores e vertebrados foram consumidos em diferentes proporções pelas espécies estudadas. Flores e vertebrados foram consumidos preferencialmente na estação seca e a diversidade da dieta de todas as espécies foi maior durante a estação chuvosa. Nem todos os recursos (artrópodes e frutos) foram consumidos de acordo com sua disponibilidade na área de estudo. Apesar de abundante, Hymenoptera (Formicidade) foi rejeitado por todas as espécies, sendo consumido abaixo de sua disponibilidade local. Os didelfídeos selecionaram frutos de Melastomataceae (Clidemia urceolata e Miconia spp.) e rejeitaram frutos de Rubiaceae, um recurso altamente abundante na área de estudo. Os resultados sugerem que o frequente consumo de um item alimentar pode estar associado tanto com a preferência (seleção) por parte do consumidor, bem como com a disponibilidade local do recurso. A maior parte das sementes, que permaneceram intactas após passagem pelo trato digestório dos animais, não apresentou diferenças significativas em suas taxas de germinação quando comparadas com as sementes do grupo controle e o tempo médio de dormência das sementes consumidas pelos marsupiais variou entre 30 (Cipocereus minensis) e 175 dias (Cordiera sessilis). Gracilinanus agilis e G. microtarsus, que ocorrem em simpatria na área de estudo, apesar de apresentarem uma alta sobreposição de nicho apresentaram diferenças no uso do habitat e na diversidade da dieta. / In Brazil, the family Didelphidae comprises 54 species widely distributed in different habitats and a dietary pattern that can vary from species more frugivorous to the most insectivorous/carnivorous. The main objective of this study was to evaluate the relationship between the diet of seven species of didelphids (Caluromys philander, Didelphis albiventris, Gracilinanus agilis, G. microtarsus, Marmosa (Micoureus) paraguayana, Marmosops incanus e Metachirus nudicaudatus) and the selection of the available food resources (arthropods and fruits). The study was conducted between November 2009 and October 2011 in a riparian area of Cerrado in the Parque Estadual do Rio Preto, Minas Gerais, Brazil. Arthropods, fruits, flowers and vertebrates were consumed in different proportions by the studied species. Flowers and vertebrates were consumed mainly in the dry season and the diversity of the diet of all species was higher during the rainy season. Not all resources (arthropods and fruits) were consumed according to their availability in the study site. Although abundant, Hymenoptera (Formicidade) was rejected by all species being consumed below their local availability. The didelphids selected fruits of Melastomataceae (Miconia and Clidemia urceolata spp.) and rejected fruits of Rubiaceae, a feature highly abundant in the study site. The results suggest that the frequent use of a food item may be associated both with the preference (selection) of the consumers, as well as the local availability of the resource. Most seeds that remained intact after passage through the digestive tract of animals, do not show significant differences in their rates of germination compared with seeds from the control group and the average time of dormancy consumed by marsupials ranged from 30 (Cipocereus minensis) and 175 days (Cordiera sessilis). Gracilinanus agilis and G. microtarsus, which occur sympatrically in the study site, although having a high niche overlap showed differences in habitat use and dietary diversity.

Os processos de dispersão, sobrevivência da prole e os períodos fenológicos das espécies vegetais, são o resultado tanto de sua história evolutiva quanto de suas adaptações ao ambiente. Para analisar o padrão evolutivo destes processos, foi testada a hipótese de que a similaridade entre espécies aparentadas em relação aos seus atributos reprodutivos (frutos, sementes e a fenologia da floração e frutificação) é maior do que o esperado por mero acaso (alto sinal filogenético), através de testes de Mantel parcial controlando o efeito da forma de vida das espécies. Os resultados revelaram que a maioria dos atributos de sementes, dois atributos dos diásporos (display das cores e a porcentagem de polpa) e o tempo de desenvolvimento dos diásporos, apresentaram um moderado sinal filogenético. Isto sugere que a conservação de atributos de sementes para sobrevivência da prole e a labilidade dos atributos de diásporos buscando dispersões mais efetivas, afetam significativamente o fitness das espécies, além de indicar uma influência do tempo de desenvolvimento dos diásporos nos picos de frutificação. / The processes of dispersal, offspring survival and phenological periods of plant species result both from their evolutionary history and adaptations to the environment. To analyze the evolutionary pattern of these processes, the hypothesis that the similarity among related species in relation to their reproductive traits (fruits, seeds and the phenology of flowering and fruiting) is higher than expected by chance (high phylogenetic signal) was tested through partial Mantel tests, controlling for the effect of species life form. The results revealed that most seed traits, two diaspores traits (color display and pulp percentage) and the time of diaspore development, show moderate phylogenetic signal. This suggests that the conservation of seed traits for offspring survival and the lability of diaspore traits seeking more effective dispersions, significantly affect the species fitness, and it also indicates the influence of the time of diaspore development on the fruiting peaks.

Coordenação de Aperfeiçoamento de Pessoal de Nível Superior / No Brasil, a Família Didelphidae é composta por 54 espécies com ampla distribuição por diferentes habitats e um padrão de consumo alimentar que pode variar desde espécies mais frugívoras até as mais insetívoras/carnívoras. O objetivo deste estudo foi avaliar a relação entre a dieta de sete espécies de didelfídeos (Caluromys philander, Didelphis albiventris, Gracilinanus agilis, G. microtarsus, Marmosa (Micoureus) paraguayana, Marmosops incanus e Metachirus nudicaudatus) e a seleção de recursos alimentares (artrópodes e frutos) disponíveis. O estudo foi realizado entre novembro de 2009 e outubro de 2011, em uma área de mata ciliar de cerrado no Parque Estadual do Rio Preto, Minas Gerais, Brasil. Artrópodes, frutos, flores e vertebrados foram consumidos em diferentes proporções pelas espécies estudadas. Flores e vertebrados foram consumidos preferencialmente na estação seca e a diversidade da dieta de todas as espécies foi maior durante a estação chuvosa. Nem todos os recursos (artrópodes e frutos) foram consumidos de acordo com sua disponibilidade na área de estudo. Apesar de abundante, Hymenoptera (Formicidade) foi rejeitado por todas as espécies, sendo consumido abaixo de sua disponibilidade local. Os didelfídeos selecionaram frutos de Melastomataceae (Clidemia urceolata e Miconia spp.) e rejeitaram frutos de Rubiaceae, um recurso altamente abundante na área de estudo. Os resultados sugerem que o frequente consumo de um item alimentar pode estar associado tanto com a preferência (seleção) por parte do consumidor, bem como com a disponibilidade local do recurso. A maior parte das sementes, que permaneceram intactas após passagem pelo trato digestório dos animais, não apresentou diferenças significativas em suas taxas de germinação quando comparadas com as sementes do grupo controle e o tempo médio de dormência das sementes consumidas pelos marsupiais variou entre 30 (Cipocereus minensis) e 175 dias (Cordiera sessilis). Gracilinanus agilis e G. microtarsus, que ocorrem em simpatria na área de estudo, apesar de apresentarem uma alta sobreposição de nicho apresentaram diferenças no uso do habitat e na diversidade da dieta. / In Brazil, the family Didelphidae comprises 54 species widely distributed in different habitats and a dietary pattern that can vary from species more frugivorous to the most insectivorous/carnivorous. The main objective of this study was to evaluate the relationship between the diet of seven species of didelphids (Caluromys philander, Didelphis albiventris, Gracilinanus agilis, G. microtarsus, Marmosa (Micoureus) paraguayana, Marmosops incanus e Metachirus nudicaudatus) and the selection of the available food resources (arthropods and fruits). The study was conducted between November 2009 and October 2011 in a riparian area of Cerrado in the Parque Estadual do Rio Preto, Minas Gerais, Brazil. Arthropods, fruits, flowers and vertebrates were consumed in different proportions by the studied species. Flowers and vertebrates were consumed mainly in the dry season and the diversity of the diet of all species was higher during the rainy season. Not all resources (arthropods and fruits) were consumed according to their availability in the study site. Although abundant, Hymenoptera (Formicidade) was rejected by all species being consumed below their local availability. The didelphids selected fruits of Melastomataceae (Miconia and Clidemia urceolata spp.) and rejected fruits of Rubiaceae, a feature highly abundant in the study site. The results suggest that the frequent use of a food item may be associated both with the preference (selection) of the consumers, as well as the local availability of the resource. Most seeds that remained intact after passage through the digestive tract of animals, do not show significant differences in their rates of germination compared with seeds from the control group and the average time of dormancy consumed by marsupials ranged from 30 (Cipocereus minensis) and 175 days (Cordiera sessilis). Gracilinanus agilis and G. microtarsus, which occur sympatrically in the study site, although having a high niche overlap showed differences in habitat use and dietary diversity.

Os processos de dispersão, sobrevivência da prole e os períodos fenológicos das espécies vegetais, são o resultado tanto de sua história evolutiva quanto de suas adaptações ao ambiente. Para analisar o padrão evolutivo destes processos, foi testada a hipótese de que a similaridade entre espécies aparentadas em relação aos seus atributos reprodutivos (frutos, sementes e a fenologia da floração e frutificação) é maior do que o esperado por mero acaso (alto sinal filogenético), através de testes de Mantel parcial controlando o efeito da forma de vida das espécies. Os resultados revelaram que a maioria dos atributos de sementes, dois atributos dos diásporos (display das cores e a porcentagem de polpa) e o tempo de desenvolvimento dos diásporos, apresentaram um moderado sinal filogenético. Isto sugere que a conservação de atributos de sementes para sobrevivência da prole e a labilidade dos atributos de diásporos buscando dispersões mais efetivas, afetam significativamente o fitness das espécies, além de indicar uma influência do tempo de desenvolvimento dos diásporos nos picos de frutificação. / The processes of dispersal, offspring survival and phenological periods of plant species result both from their evolutionary history and adaptations to the environment. To analyze the evolutionary pattern of these processes, the hypothesis that the similarity among related species in relation to their reproductive traits (fruits, seeds and the phenology of flowering and fruiting) is higher than expected by chance (high phylogenetic signal) was tested through partial Mantel tests, controlling for the effect of species life form. The results revealed that most seed traits, two diaspores traits (color display and pulp percentage) and the time of diaspore development, show moderate phylogenetic signal. This suggests that the conservation of seed traits for offspring survival and the lability of diaspore traits seeking more effective dispersions, significantly affect the species fitness, and it also indicates the influence of the time of diaspore development on the fruiting peaks.

Os processos de dispersão, sobrevivência da prole e os períodos fenológicos das espécies vegetais, são o resultado tanto de sua história evolutiva quanto de suas adaptações ao ambiente. Para analisar o padrão evolutivo destes processos, foi testada a hipótese de que a similaridade entre espécies aparentadas em relação aos seus atributos reprodutivos (frutos, sementes e a fenologia da floração e frutificação) é maior do que o esperado por mero acaso (alto sinal filogenético), através de testes de Mantel parcial controlando o efeito da forma de vida das espécies. Os resultados revelaram que a maioria dos atributos de sementes, dois atributos dos diásporos (display das cores e a porcentagem de polpa) e o tempo de desenvolvimento dos diásporos, apresentaram um moderado sinal filogenético. Isto sugere que a conservação de atributos de sementes para sobrevivência da prole e a labilidade dos atributos de diásporos buscando dispersões mais efetivas, afetam significativamente o fitness das espécies, além de indicar uma influência do tempo de desenvolvimento dos diásporos nos picos de frutificação. / The processes of dispersal, offspring survival and phenological periods of plant species result both from their evolutionary history and adaptations to the environment. To analyze the evolutionary pattern of these processes, the hypothesis that the similarity among related species in relation to their reproductive traits (fruits, seeds and the phenology of flowering and fruiting) is higher than expected by chance (high phylogenetic signal) was tested through partial Mantel tests, controlling for the effect of species life form. The results revealed that most seed traits, two diaspores traits (color display and pulp percentage) and the time of diaspore development, show moderate phylogenetic signal. This suggests that the conservation of seed traits for offspring survival and the lability of diaspore traits seeking more effective dispersions, significantly affect the species fitness, and it also indicates the influence of the time of diaspore development on the fruiting peaks.

Submitted by (ana.araujo@ufrpe.br) on 2016-08-23T12:16:59Z No. of bitstreams: 1 Leonardo Barbosa da Silva.pdf: 2235273 bytes, checksum: be77ba200c34680829de012cdc622d59 (MD5) / Made available in DSpace on 2016-08-23T12:17:00Z (GMT). No. of bitstreams: 1 Leonardo Barbosa da Silva.pdf: 2235273 bytes, checksum: be77ba200c34680829de012cdc622d59 (MD5) Previous issue date: 2013-07-29 / Coordenação de Aperfeiçoamento de Pessoal de Nível Superior - CAPES / A great wealth of flowering plants are found in tropical forests and the study of the different processes involved in the reproduction of the species is crucial for understanding the composition and structure of communities, and provide information to help understand the evolutionary history of plant breeding. In tropical ecosystems, involving animals in the reproductive processes of plants is of great importance, occurring with high frequency. The breaking of these interactions can lead to loss of ecological groups, affecting the reproductive success of plants and consequently the maintenance of their populations. The present work aims to characterize the reproductive attributes of plant community of shrubs in wet forest Araripe National Forest, through analysis of sexual systems, pollination systems and systems of diaspora dispersal. Our hypothesis is that the proportions of such reproductive attributes are similar to those recorded for other communities woody tropical forest formations, there is a predominance of the sexual system hermaphrodite, the pollination system entomófilo (especially melittophily) and dispersion zoochoric. Monthly collections were made of species in the reproductive stage, and focal observations of pollinators and seed dispersers. Found in assembly plants studied a high frequency of hermaphroditic species (88,23%) with pollination system entomófilo (98,53%), held mainly by bees (64,70%) and diasporas zoochorous (77,61%). This result reinforces the importance of ecological interactions for maintaining biodiversity and corroborates several studies conducted in different tropical ecosystems, confirming a trend in these environments / Uma grande riqueza de angiospermas é encontrada nas florestas tropicais e o estudo dos diferentes processos envolvidos na reprodução destas espécies é determinante para compreensão da composição e estruturação das comunidades, além de fornecer informações que ajudem a entender a historia evolutiva da reprodução vegetal. Em ecossistemas tropicais a participação de animais nos processos reprodutivos das plantas é de grande importância, ocorrendo com alta frequência. A quebra dessas interações pode levar a perda de grupos ecológicos, afetando o sucesso reprodutivo das plantas e consequentemente à manutenção de suas populações. O presente trabalho tem como objetivo caracterizar os atributos reprodutivos da comunidade vegetal arbustiva de mata úmida na Floresta Nacional do Araripe, através de análises dos sistemas sexuais, sistemas de polinização e dos sistemas de dispersão de diásporos. A nossa hipótese é de que as proporções de tais atributos reprodutivos são semelhantes às registradas para as comunidades lenhosas de outras formações florestais tropicais, ou seja, há predominância do sistema sexual hermafrodita, do sistema de polinização entomófilo (especialmente a melitofilia) e dispersão zoocórica. Foram realizadas coletas mensais de espécies em estágio reprodutivo, além de observações focais dos agentes polinizadores e dispersores. Encontramos na assembléia de plantas estudada uma alta frequência de espécies hermafroditas (88,23%) com sistema de polinização entomófilo (98,53%), realizado principalmente por abelhas (64,70%) e com diásporos zoocóricos (77,61%). Este resultado reforça a importâncias das interações ecológicas para manutenção da biodiversidade e corrobora com diversos estudos realizados em diferentes ecossistemas tropicais, confirmando uma tendência nestes ambientes.

La dispersion est un processus clé dans les dynamiques de populations. Chez les organismes à dispersion passive, dont les plantes, la dispersion de la banque de graines fait appel à un vecteur biotique ou abiotique. En particulier, la zoochorie influence les patrons spatiaux de distribution et de diversité des plantes à travers une interaction entre les traits écologiques du vecteur animal et de la plante transportée. Ma thèse vise à étudier l’effet de cette interaction sur la composition taxonomique et fonctionnelle des communautés floristiques. Je me concentre sur trois questions principales : (1) Comment les traits écologiques des plantes et des vecteurs influencent-ils le processus de dispersion endozoochore? ; (2) La zoochorie constitue-t-elle un filtre d’assemblage sur les traits fonctionnels des plantes dispersées? ; (3) La zoochorie imprime-t-elle un signal sur les patrons spatiaux de diversité? J’adopte à cet effet une approche expérimentale, en me concentrant sur la flore d’Europe de l’Ouest et sur les trois ongulés sauvages les plus communs dans cette région : le cerf (Cervus elaphus), le chevreuil (Capreolus capreolus) et le sanglier (Sus scrofa). Je montre que l’interaction entre les traits des plantes et des vecteurs influe sur les durées de rétention des graines, qui influencent les distances de dispersion. L’endozoochorie modifie la composition de l’assemblage d’espèces dispersé par rapport à la flore régionale en imposant un filtre fonction de l’habitat dans lequel se nourrit le vecteur animal, mais indépendant des traits morphologiques des graines. A échelle des communautés, l’effet de la zoochorie est cependant réduit relativement aux autres processus abiotiques et biotiques tels que les filtres liés à l’habitat ou l’herbivorie. Compte tenu de ces résultats, je propose de prendre en compte l’influence de la dispersion dans les modèles prédictifs de distributions des plantes, afin d’améliorer notre compréhension des dynamiques d’aires et leur prédiction en lien avec les scénarios de changements climatiques. Il apparaît en particulier nécessaire de mieux quantifier la contribution de la dispersion zoochore aux patrons de diversité et de composition des communautés végétales, relativement aux autres processus qui résultent d’interactions plantes-animaux et aux autres modes de dispersion des graines. / Dispersal is a key process shaping population dynamics. In passive dispersers like plants, the dispersal of the seed bank relies on biotic or abiotic vectors. Among the wide range of passive dispersal, zoochory influences spatial plant diversity and distribution patterns through an interaction between the ecological traits of dispersed plants and their animal vectors. In this work, I investigate the outcomes of this interaction on the taxonomic and functional composition of plant communities. I address three main questions: (1) How do the ecological traits of dispersed plants and their vectors influence the dynamics of endozoochorous dispersal? ; (2) Does zoochory affect the functional traits of dispersed species as a community assembly filter? ; (3) What is the imprint of zoochory on spatial patterns of plant diversity? I frame my work within an experimental approach focused on the West-European flora and on the three most common wild ungulates in this area: red deer (Cervus elaphus), roe deer (Capreolus capreolus) and wild boar (Sus scrofa). I show that interactions between plant and vector traits influence dispersal distances by modulating seed retention times. Endozoochory modifies the composition of dispersed plant assemblages as compared with that of the regional pool by filtering species according to the feeding habitats of the vectors. At a community level, zoochory has a limited influence relative to other abiotic or biotic processes including habitat and herbivory. On the basis of these results, I suggest to include dispersal in predictive models of plant distributions to improve our understanding of range dynamics and their prediction especially within the framework of current global changes. My results further suggest that the contribution of zoochorie to plant diversity and community composition patterns needs to be better quantified and compared with other plant-animal interactions and other dispersal modes.