Submitted by Franciele Moreira (francielemoreyra@gmail.com) on 2018-04-11T11:57:10Z No. of bitstreams: 2 Dissertação - Lucas Pereira Martins - 2018.pdf: 2113878 bytes, checksum: 4c38a4d425f42506d50aa62bf6661c3f (MD5) license_rdf: 0 bytes, checksum: d41d8cd98f00b204e9800998ecf8427e (MD5) / Approved for entry into archive by Luciana Ferreira (lucgeral@gmail.com) on 2018-04-12T11:22:54Z (GMT) No. of bitstreams: 2 Dissertação - Lucas Pereira Martins - 2018.pdf: 2113878 bytes, checksum: 4c38a4d425f42506d50aa62bf6661c3f (MD5) license_rdf: 0 bytes, checksum: d41d8cd98f00b204e9800998ecf8427e (MD5) / Made available in DSpace on 2018-04-12T11:22:54Z (GMT). No. of bitstreams: 2 Dissertação - Lucas Pereira Martins - 2018.pdf: 2113878 bytes, checksum: 4c38a4d425f42506d50aa62bf6661c3f (MD5) license_rdf: 0 bytes, checksum: d41d8cd98f00b204e9800998ecf8427e (MD5) Previous issue date: 2018-03-15 / Conselho Nacional de Pesquisa e Desenvolvimento Científico e Tecnológico - CNPq / Understanding how species interact with each other is essential to advance our knowledge on community ecology. However, there are still gaps regarding how interspecific interactions affect and are affected by the variation in the composition of species that constitute the ecological networks. In particular, studies of beta diversity are interesting to understand how sets of biotic filters may affect species’ geographic distribution and persistence on communities. Another interesting question is if variations in species composition may cause changes on how ecological networks are structured. In this dissertation, we propose to evaluate patterns of beta diversity and structure on insect-plant interaction networks. Specifically, in the first chapter we aimed to test if beta diversity is affected by the trophic level to which the assemblage belongs, and by the degree of specialization of species at higher trophic levels to their host species. For this, we used a tritrophic system comprising plants of the family Asteraceae, endophagous herbivores and parasitoids sampled in remnants of Brazilian Cerrado. Our main results show that total beta diversity of parasitoids (higher trophic level) was lower than those of the other trophic levels, while there was no difference between total beta diversity of plants and herbivores. Furthermore, the degree of specialization of the assemblages of herbivores and parasitoids was positively associated to the beta diversity of these groups. In the second chapter, we evaluated if species beta diversity among sites affects dissimilarity in network structure, and if this relationship changes across time. Overall, the relationship between spatial beta diversity and dissimilarity in plant-herbivore network structure was context-dependent, thus indicating that different ecological processes (i.e., niche-based and neutral) may drive the organization of antagonistic networks across time. / Compreender como as espécies interagem entre si é essencial para avançar o nosso conhecimento em ecologia de comunidades. No entanto, ainda existem lacunas no que se refere a como interações interespecíficas influenciam e são influenciadas pela variação na composição de espécies que constituem as redes ecológicas. Em particular, estudos de diversidade beta são interessantes para entender como conjuntos de filtros bióticos podem influenciar a distribuição geográfica e persistência de espécies nas comunidades. Outra questão interessante é se variações na composição de espécies podem causar mudanças no modo como redes ecológicas são estruturadas. Nesta dissertação, propomos avaliar padrões de diversidade beta e estrutura de redes de interações inseto-planta. Especificamente, no primeiro capítulo visamos testar se a diversidade beta é influenciada pelo nível trófico ao qual a assembleia pertence, e pelo grau de especialização de espécies de níveis tróficos superiores às suas espécies hospedeiras. Para isso, utilizamos um sistema tritrófico composto por plantas da família Asteraceae, herbívoros endófagos e parasitoides amostrado em remanescentes de Cerrado brasileiro. Nossos principais resultados demonstram que a diversidade beta total de parasitoides (nível trófico superior) foi menor do que a dos demais níveis tróficos, enquanto que não houve diferença entre a diversidade beta total de plantas e herbívoros. Além disso, o grau de especialização das assembleias de herbívoros e parasitoides foi positivamente relacionado à diversidade beta destes grupos. No segundo capítulo, avaliamos se a diversidade beta de espécies entre locais influencia a dissimilaridade na estrutura de redes ecológicas, e se esta relação muda através do tempo. De modo geral, a relação entre diversidade beta espacial e dissimilaridade na estrutura de redes planta-herbívoro foi contexto-dependente, indicando assim que diferentes processos ecológicos (i.e., baseados em nicho e neutros) podem determinar a organização de redes antagonistas através do tempo.

Mutualismos são interações em que os parceiros se exploram reciprocamente com benefícios líquidos para ambos os indivíduos que interagem. Sistemas mutualistas multiespecíficos podem ser descritos como redes de interação, tais como aquelas formadas por sistemas de polinização, dispersão de sementes, estações de limpeza em ambientes recifais, formigas defensoras de plantas, mimetismo mülleriano e bactérias fixadoras de nitrogênio em raízes de plantas. As interações mutualísticas estão sujeitas à trapaça por indivíduos que, por meio de algum comportamento, alcançam o benefício oferecido pelo parceiro sem oferecer nada ou oferecer muito pouco em troca. No entanto, interações mutualísticas persistem apesar da existência de trapaceiros. Neste trabalho, mostro que os parasitas de interações mutualísticas, os trapaceiros, aumentam a resiliência das redes mutualísticas às perturbações mais rapidamente em redes aninhadas, redes tipicamente encontradas em mutualismos ricos em espécies. Portanto os efeitos combinados de trapaceiros, estrutura e dinâmica das redes mutualísticas podem ter implicações para a forma como a biodiversidade é mantida. Em seguida, estudo as condições em que flores tubulares, que sofrem maiores danos ao interagirem com ladrões de néctar, conseguem coexistir com flores planares, polinizadores e pilhadores por meio de efeitos indiretos da trapaça em seu sucesso reprodutivo. O roubo do néctar pode aumentar o sucesso de uma planta se as interações com pilhadores gerarem maior quantidade de polinização cruzada, aumentando assim o sucesso reprodutivo das plantas que interagem com ambos os visitantes florais. Tal resultado sugere uma nova fonte de manutenção da cooperação e da diversidade de estratégias por meio de efeitos não lineares das interações entre diferentes estratégias. Finalmente, estudo como as interações locais promovem a prevalência de mímicos (trapaceiros) em uma certa população na ausência de seus modelos. Mostro que presas que interagem localmente podem favorecer a predominância de mímicos e predadores que os evitam após algumas gerações e que uma distribuição não aleatória de indivíduos no espaço pode reforçar ainda mais este efeito inesperado de alopatria de modelo e mímico / Mutualisms are interactions in which organisms of different species exploit each other with net benefits for both interacting individuals. Multispecific mutualistic system can be depicted as interaction networks, such as those formed by plant-pollinator interactions, dispersal systems, species interacting in cleaning stations in reef environments, protective ants in plants, müllerian mimicry, and nitrogen fixing bacteria on the roots of plants. Mutualistic interaction is subject to cheating by individuals who, by means of a diversity of behavioral strategies, achieve the benefit provided by the partner offering nothing or few in return. However, the mutualistic interactions persist despite the existence of cheaters. In this work I show that the parasites of mutualistic interactions increase the resilience of mutualistic networks to disturbances in nested networks, typically found in species-rich mutualisms. Therefore the joint effect of cheating, structure and dynamics of mutualistic networks have implications for how biodiversity is maintained. I subsequently study the conditions under which tubular flowers, which suffer stronger damages when interacting with nectar robbers, can coexist with planar flowers, pollinators, and robbers through indirect effects of cheating on their reproductive success. The theft of nectar may increase the success of a plant if its interactions with robbers generate higher degrees of cross-pollination, thus increasing the reproductive success of plants that interact with both floral visitors. This study suggests a new source of continued cooperation and diversity strategies through non-linear effects of the interactions between different strategies. Finally, I study how local interactions can promote the prevalence of mimic (the cheaters) in a given population in the absence of their models. I found that prey interacting locally may favor the predominance of mimic preys and avoid predators that, after a few generations and under a non-random distribution of individuals in space, can further strengthen this unexpected effect allopatry of the mimic and its model

Parmi les problèmes les plus fréquemment soulevés en biogéographie, figure celui de l’intégration des interactions écologiques dans les modèles de distribution d’espèces. Bien que la littérature scientifique apporte un ensemble de preuves soulignant le rôle prépondérant des interactions dans la structuration des communautés locales, on trouve relativement peu d’études révélant les empreintes laissées par les interactions dans les données de distribution d’espèces. Proposer une explication simple et claire à ce problème demeure un défi important que la biogéographie doit mener. Le problème majeur que pose l’absence de réponse claire sur le rôle des interactions aux larges échelles spatiales est que la plupart des scénarios de changements de biodiversité partent de l’hypothèse que les interactions sont négligeables. Si cette hypothèse est régulièrement rejetée, alors il faut réviser ces scénarios et soutenir le développement de méthodologies incluant les relations entre les espèces. Je commence cette thèse par un travail théorique sur le sujet car les théories classiques en biogéographie relèguent souvent au second plan les interactions écologiques. Au premier chapitre, je traite de l'intégration des interactions écologiques dans un modèle théorique de distribution d'espèces issue d'une des théories les plus importantes en biogéographie: la théorie de la biogéographie des îles. Ce travail montre comment les effets conjoints des facteurs biotiques et abiotiques changent les attendus de la théorie classique. En m'appuyant sur ce premier chapitre, je montre au second chapitre comment les interactions peuvent se répercuter dans les données de co-occurrence d’espèces. Ces données indiquent la présence ou l’absence de plusieurs espèces sur un même ensemble de sites dispersés sur de larges étendues spatiales. À l’aide d’un modèle probabiliste, j'obtiens des résultats théoriques liant les données de co-occurrence et l’information contenue dans les réseaux écologiques.Je démontre clairement que les interactions affectent les données de co-occurrence. Je montre également que plus le nombre d’interactions séparant deux espèces est grand, moins leur interactions indirect est détectable. De même si une espèce entretient de nombreuses interactions, il sera difficile de trouver une quelconque trace des interactions dans les données de co-occurrence pour cette espèce. Au troisième chapitre, je présente l’analyse de cinq jeux de données de co-occurrence pour lesquels la description des interactions était disponible. Avec ces donnés, j'ai été capable de confirmer les hypothèses du second chapitre en montrant que les espèces qui interagissent co-occurrent différemment de celles n’interagissant pas. Mes résultats indiquent aussi que l’abondance d'interactions est un frein à leur détection dans les données de co-occurrence. Cependant, en intégrant la similarité des facteurs abiotiques pour les différents sites, je montre que les signaux de co-occurrence s’affaiblissent pour parfois disparaitre. Mes résultats suggèrent donc qu’en utilisant des facteurs abiotiques pour inférer les probabilités de co-occurrence,une partie du lien entre les espèces est capturée, mais cette part est entachée d’une grande incertitude. Ceci vient questionner la qualité des prédictions données par les modèles classiques de distribution d'espèces actuellement utilisés. Les résultats de ma recherche apportent des éléments théoriques nouveaux sur le rôle des interactions écologiques dans le tracé des aires de répartition des espèces en plus de proposer une méthode originale pour étudier les données de co-occurrence d’espèces : les regarder à la lumière des réseaux écologiques. Avant de conclure ma thèse, je propose au chapitre 4 une démarche prometteuse pour aller encore améliorer l’intégration des interactions en biogéographie : les introduire par le biais des contraintes énergétiques, ce qui offre une base solide pour une théorie métabolique de la biogéographie. / One of the most pressing challenges currently in the field of biogeography is the successful integration of ecological interactions in species distribution models. Although the scientific literature points out the evidence of the controlling role interactions play on local community structure, relatively few studies have demonstrated its importance over large geographical gradients. Developing a concise, clear explanation for this issue remains a significant challenge that biogeographers need to answer. The main issue associated to the lack of a clear answer concerning the role of interactions at broad spatial scales is that most of scenarios of biodiversity changes assume that interactions can be ignored. When tested, if this hypothesis is proven false, then a re-consideration of species distribution models and their development must be undertaken to include relationships among species. I begin this thesis with a theoretical investigation on this topic, where classical theories have typically ignored ecological interactions. In the first chapter of the thesis I present the integration of interaction networks into a theoretical model of species distribution coming from one of the most important theory in biogeography: the theory of island biogeography. This work shows how together the biotic and abiotic factors can affect the expectations derived from the classical theory. Building upon the findings in the first chapter, in the second chapter, I show how interactions can affect co-occurrence (between species) data. Such data contains the presence or absence of several species for a similar set of sites dispersed along large latitudinal gradients. Using a probabilistic model, I obtain theoretical results linking co-occurrence data and the information included in ecological networks. I clearly demonstrate that interactions shape co-occurrence data. Furthermore, I show that the higher the number of links between two species, the more difficult it is to detect their indirect interaction. Similarly, if a species experiences many interactions, it is then challenging to detect any sign of interactions in co-occurrence data for this species.In the third chapter of the thesis, I assess five sets of co-occurrence data, which had descriptions of their interactions available. Using this data, I was able to confirm my hypotheses put forth in my second chapter, by showing that species co-occur differently from non-interacting one. These results also point out that the abundance of interaction must preclude their detection in co-occurrence data. However, when accounting for abiotic similarities among sites, signals of interactions are weakened. Therefore, my results suggest that using abiotic factors to infer co-occurrence probabilities capture a part of the link between species and further pinpoint the uncertainty associated to this part. As a result of these findings, the predictive power of classical species distribution models used to date is brought into question. My research findings bring new theoretical elements to the forefront when considering the influence of ecological interactions and how they shape species geographical distributions, while also introducing an original methodology for studying species co-occurrence: examining them in the light of ecological networks. Before concluding, my fourth and final chapter, I propose a promising new avenue to further investigate integrating species interactions in biogeography. Here, I introduce interactions in terms of energetic constraints, which will provide a sound basis for a metabolic theory of biogeography.

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Nous pouvons trouver des propriétés structurelles similaires dans presque tous les réseaux trophiques (ensemble d’interactions de prédation). L'existence de ces invariants suggère qu’il serait possible, pour chaque réseau trophique, de déterminer des paramètres généraux qui décrivent sa structure. Il serait également possible de faire le cheminement inverse, soit à partir de paramètres généraux, d’obtenir une structure de réseau qui respecte ces invariants. C’est ainsi que fonctionnent les modèles génératifs, qui prédisent une structure à partir de paramètres généraux. Cependant, les modèles génératifs peuvent générer des structures de réseau qui diffèrent des données empiriques, parce qu'ils intègrent différentes hypothèses sur les mécanismes qui façonnent les réseaux trophiques, et donc sur les paramètres généraux qui doivent être utilisés. Dans ce mémoire, j’étudie l'effet de la pondération de la stabilité à l'aide du paramètre sigma (écart-type maximum des forces d’interactions qu’il ne faut pas dépasser si l’on veut que le réseau d’espèces reste stable) sur la distribution des propriétés de réseau obtenues par différents modèles génératifs. En effet, en donnant une plus grande importance aux réseaux dont la structure est a priori stable (potentiellement plus proche de celles retrouvées dans la nature) on pourrait corriger les prédictions des modèles en rapprochant leurs résultats des données empiriques. Le principe de correction fait ici référence à l’utilisation des probabilités par les modèles génératifs : la correction est la modification de ces probabilités en faveur des réseaux stables afin qu’ils soient sur-représentés dans les données générées. Notre hypothèse est donc que la pondération de la stabilité pourrait améliorer les prédictions des modèles génératifs. Les modèles génératifs étudiés ici sont les modèles de cascade, de niche et de hiérarchie emboîtée. Notre principale conclusion est que, de manière contre-intuitive, la pondération de la stabilité n’améliore pas la différence entre les structures de réseaux empiriques et celles des réseaux générés par les mo-dèles. Nos résultats montrent que pour les réseaux étudiés, la plus grande différence entre les réseaux trophiques modélisés par les modèles génératifs et les réseaux empiriques est la nature du modèle et non la correction par la pondération de la stabilité. Cela suggère que ces modèles prédisent la structure à partir d’un nombre de paramètres insuffisants, où de paramètres ne représentant qu’une fraction de la structure du réseau. Le modèle de niche présente la prédiction la plus proche des données empiriques, mais seulement pour les réseaux comptant jusqu'à 20 espèces. Cette étude souligne donc le long chemin qu'il nous reste à parcourir avant de pouvoir représenter les réseaux trophiques de façon réaliste à partir de modèles génératifs simples. / We can find similar structural properties in almost every food web. The existence of these invariants suggests that it could be possible for each food web to determine general parameters. The reverse case also works, i.e. from general parameters, to obtain a network structure. This is how generative models work, they predict a structure from general parameters. However, the network structures obtained from generative models differ from empirical data, because they incorporate different assumptions about the mechanisms that shape food webs and thus the gen-eral parameters used. In this study, I’ll investigate the effect of weighting stability using the sigma parameter (maximum standard deviation of interaction forces that should not be exceeded if the species network is to remain stable). I’m studying its effect on the distribution of network prop-erties obtained by different generative models. Indeed, by giving greater importance to networks whose structure is stable, one could correct the predictions of the models by bringing their results closer to the empirical data. The correction is the modification of these probabilities in favor of stable networks so that they are more easily chosen by the model. Our hypothesis is therefore that weighting stability could improve the predictions of the cascade, niche and nested hierarchy models. Our main conclusion is that stability weighting does not improve the difference between empirical and model-generated network structures. Our results show that for the networks stud-ied, the biggest difference between food webs modeled by generative models and empirical net-works is the nature of the model and not the correction by stability weighting. This suggests that these models predict structure from an insufficient number of parameters or from parameters that represent only a fraction of the network structure. The niche model shows the closest pre-diction to the empirical data, but only for networks with up to 20 species. This study highlights the long way to go before we can realistically represent food webs using generative models.

Le domaine de l’écologie des réseaux est encore limité dans sa capacité à faire des inférences mondiales à grande échelle. Ce défi est principalement dû à la difficulté d’échantillonnage des interactions sur le terrain, entraînant de nombreuses « lacunes » en ce qui concerne la couverture mondiale des données. Cette thèse adopte une approche « centrée sur les méthodes » de l’écologie des réseaux et se concentre sur l’idée de développer des outils pour aider à combler les lacunes en matière de données en présentant la prédiction comme une alternative accessible à l’échantillonnage sur le terrain et introduit deux « outils » différents qui sont prêts à poser des questions à l’échelle mondiale. Le chapitre 1 présente les outils que nous pouvons utiliser pour faire des prédictions de réseaux et est motivé par l’idée selon laquelle avoir la capacité de prédire les interactions entre les espèces grâce à l’utilisation d’outils de modélisation est impératif pour une compréhension plus globale des réseaux écologiques. Ce chapitre comprend une preuve de concept (dans laquelle nous montrons comment un simple modèle de réseau neuronal est capable de faire des prédictions précises sur les interactions entre espèces), une évaluation des défis et des opportunités associés à l’amélioration des prédictions d’interaction et une feuille de route conceptuelle concernant l’utilisation de modèles prédictifs pour les réseaux écologiques. Les chapitres 2 et 3 sont étroitement liés et se concentrent sur l’utilisation de l’intégration de graphiques pour la prédiction de réseau. Essentiellement, l’intégration de graphes nous permet de transformer un graphe (réseau) en un ensemble de vecteurs, qui capturent une propriété écologique du réseau et nous fournissent une abstraction simple mais puissante d’un réseau d’interaction et servent de moyen de maximiser les informations disponibles. dispo- nibles à partir des réseaux d’interactions d’espèces. Parce que l’intégration de graphes nous permet de « décoder » les informations au sein d’un réseau, elle est conçue comme un outil de prédiction de réseau, en particulier lorsqu’elle est utilisée dans un cadre d’apprentissage par transfert. Elle s’appuie sur l’idée que nous pouvons utiliser les connaissances acquises en résolvant un problème connu. et l’utiliser pour résoudre un problème étroitement lié. Ici, nous avons utilisé le métaweb européen (connu) pour prédire un métaweb pour les espèces canadiennes en fonction de leur parenté phylogénétique. Ce qui rend ce travail particulière- ment passionnant est que malgré le faible nombre d’espèces partagées entre ces deux régions, nous sommes capables de récupérer la plupart (91%) des interactions. Le chapitre 4 approfondit la réflexion sur la complexité des réseaux et les différentes ma- nières que nous pourrions choisir de définir la complexité. Plus spécifiquement, nous remet- tons en question les mesures structurelles plus traditionnelles de la complexité en présentant l’entropie SVD comme une mesure alternative de la complexité. Adopter une approche phy- sique pour définir la complexité nous permet de réfléchir aux informations contenues dans un réseau plutôt qu’à leurs propriétés émergentes. Il est intéressant de noter que l’entropie SVD révèle que les réseaux bipartites sont très complexes et ne sont pas nécessairement conformes à l’idée selon laquelle la complexité engendre la stabilité. Enfin, je présente le package Julia SpatialBoundaries.jl. Ce package permet à l’utili- sateur d’implémenter l’algorithme de wombling spatial pour des données disposées de manière uniforme ou aléatoire dans l’espace. Étant donné que l’algorithme de wombling spatial se concentre à la fois sur le gradient et sur la direction du changement pour un paysage donné, il peut être utilisé à la fois pour détecter les limites au sens traditionnel du terme ainsi que pour examiner de manière plus nuancée la direction des changements. Cette approche pourrait être un moyen bénéfique de réfléchir aux questions liées à la détection des limites des réseaux et à leur relation avec les limites environnementales. / The field of network ecology is still limited in its ability to make large-scale, global inferences. This challenge is primarily driven by the difficulty of sampling interactions in the field, leading to many ‘gaps’ with regards to global coverage of data. This thesis takes a ’methods-centric’ approach to network ecology and focuses on the idea of developing tools to help with filling in the the data gaps by presenting prediction as an accessible alternative to sampling in the field and introduces two different ’tools’ that are primed for asking questions at global scales. Chapter 1 maps out tools we can use to make network predictions and is driven by the idea that having the ability to predict interactions between species through the use of modelling tools is imperative for a more global understanding of ecological networks. This chapter includes a proof-of-concept (where we show how a simple neural network model is able to make accurate predictions about species interactions), an assessment of the challenges and opportunities associated with improving interaction predictions, and providing a conceptual roadmap concerned with the use of predictive models for ecological networks. Chapters 2 and 3 are closely intertwined and are focused on the use of graph embedding for network prediction. Essentially graph embedding allows us to transform a graph (net- work) into a set of vectors, which capture an ecological property of the network and provides us with a simple, yet powerful abstraction of an interaction network and serves as a way to maximise the available information available from species interaction networks. Because graph embedding allows us to ’decode’ the information within a network it is primed as a tool for network prediction, specifically when used in a transfer learning framework, this builds on the idea that we can take the knowledge gained from solving a known problem and using it to solve a closely related problem. Here we used the (known) European metaweb to predict a metaweb for Canadian species based on their phylogenetic relatedness. What makes this work particularly exciting is that despite the low number of species shared between these two regions we are able to recover most (91%) of interactions. Chapter 4 delves into thinking about the complexity of networks and the different ways we might choose to define complexity. More specifically we challenge the more traditional structural measures of complexity by presenting SVD entropy as an alternative measure of complexity. Taking a physical approach to defining complexity allows us to think about the information contained within a network as opposed to their emerging properties. Interest- ingly, SVD entropy reveals that bipartite networks are highly complex and do not necessarily conform to the idea that complexity begets stability. Finally, I present the Julia package SpatialBoundaries.jl. This package allows the user to implement the spatial wombling algorithm for data arranged uniformly or randomly across space. Because the spatial wombling algorithm focuses on both the gradient as well as the direction of change for the given landscape it can be used both for detecting boundaries in the traditional sense as well as a more nuanced look at at the direction of changes. This approach could be a beneficial way with which to think about questions which relate to boundary detection for networks and how these relate to environmental boundaries.

This thesis proposes a collection of analytical and computational methods for inferring an underlying social structure of a given population, observed only via timestamped occurrences of its members across a range of locations. It shows that such data streams have a modular and temporally-focused structure, neither fully ordered nor completely random, with individuals appearing in "gathering events". By exploiting such structure, the thesis proposes an appropriate mapping of those spatio-temporal data streams to a social network, based on the co-occurrences of agents across gathering events, while capturing the uncertainty over social ties via the use of probability distributions. Given the extracted graphs mentioned above, an approach is proposed for studying their community organisation. The method considers communities as explanatory variables for the observed interactions, producing overlapping partitions and node membership scores to groups. The aforementioned models are motivated by a large ongoing experiment at Wytham woods, Oxford, where a population of Parus major wild birds is tagged with RFID devices and a grid of feeding locations generates thousands of spatio-temporal records each year. The methods proposed are applied on such data set to demonstrate how they can be used to explore wild bird sociality, reveal its internal organisation across a variety of different scales and provide insights into important biological processes relating to mating pair formation.

La Trame verte et bleue (TVB) a pour objectif la préservation et la remise en « bon état » des continuités écologiques (CE). Cette politique publique veut être un outil d’aménagement du territoire. Elle est inscrite à la fois dans le code de l’environnement et de l’urbanisme. Elle se décline à différents niveaux de gouvernance, du national au local en passant par le régional. La cohérence entre ces niveaux est cadrée par la loi française. Mais une importante marge d’interprétation et de mise en œuvre de la politique est laissée aux territoires. Dans cette thèse, nous interrogeons les conditions permettant d’assurer la cohérence entre les projets des territoires ayant trait à la planification des CE. Pour cela, nous répondons à deux principales hypothèses sources de (in)cohérence : (i) l’incomplétude de la connaissance existante en écologie du paysage et son utilisation dans les territoires et, (ii) les processus de gouvernance mis en place pour saisir la marge d’adaptation de la politique, avec un focus sur les acteurs « relais » entre les projets des territoires. Ainsi, nous développons une notion de la « cohérence interterritoriale » basée sur le partage entre les acteurs des territoires d’une vision de l’organisation de l’espace. Nous en tirons une grille d’analyse de cette cohérence appliquée aux projets portant sur les CE. Cette grille présente trois volets : la dimension écologique, la multifonctionnalité et la gouvernance. Ces volets sont assortis de critères et d’indicateurs. La grille d’analyse a été appliquée à différents projets portés par des territoires « emboités » du niveau national au local, dans les régions Bretagne et Occitanie (pour l’ancienne région Languedoc-Roussillon). L’analyse est d’abord spécifique à chaque projet puis est ensuite comparative, de façon verticale entre les niveaux de gouvernance et horizontale entre mêmes niveaux de gouvernance. Nous montrons que la préservation des CE est un « wicked mess problem », dans le sens où il n’existe pas une solution unique et optimale face aux complexités écologiques et sociétales du sujet. Les territoires s’adaptent, traduisent, simplifient et ont leur propre représentation des concepts d’écologie du paysage. Les approches diffèrent du niveau national au local. L’approche naturaliste promue au niveau national devient une approche par l’occupation du sol au niveau local. De même, l’approche écologique devient multifonctionnelle en passant du code de l’environnement au code de l’urbanisme. La loi impose une cohérence descendante entre les territoires ce qui peut être source d’innovation ou au contraire limiter les initiatives par crainte du contentieux juridique. Les acteurs et les projets sont extrêmement divers. La planification des CE est une « patate plus ou moins chaude » que les territoires se repassent les uns aux autres. Les territoires porteurs de schémas de cohérence territoriale (SCoT) ou de plans locaux d’urbanisme (intercommunaux) (PLU(i)) sont souvent identifiés comme les plus pertinents pour traiter le problème. Mais l’application du principe de subsidiarité ne doit pas déresponsabiliser certains territoires, alors que les systèmes socio-écologiques sont de fait inter-échelles et donc interterritoriaux. Le rôle des acteurs « relais » entre les territoires est essentiel à la cohérence interterritoriale pour dynamiser voire créer les proximités organisées entre les territoires. La Trame verte et bleue est actuellement à un tournant suite aux récentes évolutions législatives. Les régions sont identifiées comme cheffes de file sur la biodiversité et sont dotées d’un nouveau schéma régional intégrateur absorbant, notamment, la TVB. La cohérence interterritoriale se construit dans le temps et méritera d’être analysée dans le temps. / The “Trame verte et bleue” (TVB), a French public policy, aims at preserving and restoring ecological networks (EN). It is intended to be a tool for land-use planning. It is part of French law’s codes for the environment and for urbanism. It is meant to be implemented at different governance levels, ranging from the national to the regional to the local. The coherence of the TVB between these levels has been specified in French law. Nevertheless, territories have a wide margin for interpretation and implementation of TVB policy. In this thesis, we examine the conditions necessary for ensuring coherence between territorial projects which pertain to EN planning. To this end, we address two main hypotheses that are sources of (in)coherence: (i) the incompleteness of existing knowledge in landscape ecology and its use in the territories, and (ii) the governance processes put in place to take into account the margin for adaptation of the TVB policy, with a focus on “bridging” actors between territorial projects. We develop a notion of “interterritorial coherence” based on the sharing of a vision between the stakeholders of the organization of space. We propose an analysis grid of this coherence applied to projects related to EN. The grid involves three components: the ecological dimension, the multifunctionality, and the governance processes. These components are characterized by criteria and translated into indicators. The analysis grid has been applied to different projects carried out by “nested” territories ranging from the national to the local level in two French regions (Brittany and Occitania, (formerly called Languedoc-Roussillon region)). The analysis is initially specific to each project and then becomes comparative, vertically between levels of governance and horizontally between the same levels of governance. We show that the preservation of EN is a “wicked mess problem”, in the sense that a single and optimal solution does not exist given the ecological and societal complexities of the subject. The territories adapt themselves, simplify and have their own representation of the concepts of landscape ecology. Approaches differ between the national and the local. The naturalistic approach promoted at the national level becomes a land-use approach at the local level. Similarly, the ecological approach becomes multifunctional by transitioning from the environment code to the urbanism code. The law imposes a top-down coherence between the territories. This can be a source of innovation or, on the contrary, can limit initiatives due to fear of litigation. The actors and projects are extremely diverse. EN planning is a “hot potato” (more or less “hot”) that territories pass to each other. The territories with territorial coherence schemes (SCoTs) or local urban planning plans (PLU) are often identified as the most relevant levels to tackle the problem. Nevertheless, the application of the subsidiarity principle must not disempower certain territories, since socio-ecological systems are inter-scale and therefore interterritorial. The role of “bridging” actors between the territories is essential to interterritorial coherence because they can energize or even create the organized proximities between the territories. The “Trame verte et bleue” policy is currently at a turning point following recent legislative developments. Regions are identified as leaders on biodiversity and have to follow a new integrated regional scheme that includes, in particular, EN. Interterritorial coherence is built up over time and will therefore need to be analyzed over time.

Os padrões de associação entre epífitos e forófitos não podem ser considerados espécie-específicos, mas as árvores que os epífitos colonizam também não são um conjunto aleatório das espécies forofíticas de um determinado local. Ao invés disso, parece haver uma preferência de certos epífitos por diferentes forófitos. No entanto, se conhece pouco sobre os fatores que determinam essa preferência. Nosso objetivo nesse trabalho é avaliar como os atributos funcionais e a filogenia dos epífitos vasculares influenciam na associação dos epífitos com os forófitos em uma Floresta Ombrófila Mista no sul do Brasil. Para isso nós (1) investigamos padrões de associação positiva e negativa entre grupos funcionais de epífitos e grupos de forófitos e como a diversidade e os atributos funcionais dos epífitos variavam em função do tamanho do forófito e (2) inferimos sobre a existência de sinal filogenético no uso de árvores hospedeiras pelos epífitos, procuramos por estrutura filogenética nas comunidades epifíticas e investigamos diferenças de composição filogenética de epífitos vasculares em diferentes clados de forófitos. Foram amostrados 70 forófitos compreendendo 15 espécies pertencentes a diversos clados e com arquiteturas e características variadas. A amostragem compreendeu 31 espécies epifíticas com os principais clados sendo Polypodiaceae, Bromeliaceae e Orchidaceae. A associação de grupos de epífitos vasculares com diferentes grupos de forófitos sugere que as características dos forófitos proporcionam ambientes contrastantes e que diferentes valores de atributos são necessários para colonizar esses ambientes. Mais especificamente espécies epifíticas com menor área específica foliar (SLA) parecem predominar em árvores maiores e maior SLA em árvores menores. Encontramos sinal filogenético na utilização dos forófitos, sugerindo que a conservação das interações com os forófitos deve ter sido importante ao longo da evolução dos epífitos. A tendência a agrupamento filogenético nas comunidades epifíticas sugere a influência de filtros ambientais representados pelas diferentes características dos forófitos estruturando as assembleias de epífitos. Clados mais basais de forófitos apresentaram composição filogenética distinta devido, sobretudo, a presença de diferentes clados de monilófitos epifíticos nessas árvores. Angiospermas epifíticas ocorreram principalmente em forófitos pertencente as eurosídeas. A preferência de epífitos por forófitos parece ser influenciada pelo surgimento de novidades morfológicas e ecofisiológicas em alguns clados, enquanto outros mantiveram o seu nicho ancestral. A composição florística das florestas quando da origem dos clados epifíticos também parece influenciar a associação entre epífitos e forófitos. Ao utilizar informações sobre os atributos e filogenia das espécies de epífitos vasculares nós podemos melhor compreender os mecanismos ecológicos e históricos que influenciam os padrões de associação entre epífitos e forófitos. / It has been shown that patterns of association between epiphytes and phorophytes can not be considered species-specific, although the trees that epiphytes colonize are not a random subset of phorophyte species in a particular location. Instead, there seems to be a preference of some epiphytes for different phorophytic species. However, little is known about the factors determining this choice. Our objective in this study is to assess how functional attributes and phylogeny of vascular epiphytes influence the association of epiphytes with the phorophytes in an Araucaria Forest in Southern Brazil. For that we (1) investigated the positive and negative association patterns between functional groups of epiphytes and groups of phorophytes and how functional diversity and functional traits of epiphytes varied with host tree size and (2) inferred about the existence of phylogenetic signal on the host trees use by epiphytes, looked for phylogenetic structure in the epiphytic communities and investigated differences in the phylogenetic composition of vascular epiphytes in different phorophyte clades. We used a sample of 70 phorophytes comprising 15 species and belonging to different clades and with different architectures and traits. The sample comprised 31 epiphytic species, the major clades being Polypodiaceae, Bromeliaceae and Orchidaceae. The combination of vascular epiphyte groups with different groups of 15 phorophytes suggests that phorophyte traits provide contrasting environments and that different trait values are needed to colonize these environments. More specifically, epiphytic species with lower specific leaf area (SLA) seem to predominate on larger trees and with higher SLA values on smaller trees. We found phylogenetic signal on the host tree use, suggesting that conservatism of the interactions with phorophytes must have been important throughout the evolution of epiphytes. The tendency to phylogenetic clustering in the epiphytic communities suggests the influence of environmental filters represented by phorophyte traits structuring epiphyte assemblages. More basal clades of phorophytes showed different phylogenetic composition mainly due to the presence of different epiphytic monilophyte clades on these trees. Epiphytic angiosperms occurred mainly on those trees belonging to eurosids. The preference of epiphytes for phorophytes seems to be influenced by morphological and ecophysiological novelties in some lineages, while other clades kept their ancestral niche. The floristic composition of forests at the origins of epiphytic lineages also appears to influence the association between epiphytes and phorophytes. By using information about the traits and phylogeny of species of vascular epiphytes we can better understand the ecological and historical mechanisms that influence the patterns of association between epiphytes and phorophytes.

Os padrões de associação entre epífitos e forófitos não podem ser considerados espécie-específicos, mas as árvores que os epífitos colonizam também não são um conjunto aleatório das espécies forofíticas de um determinado local. Ao invés disso, parece haver uma preferência de certos epífitos por diferentes forófitos. No entanto, se conhece pouco sobre os fatores que determinam essa preferência. Nosso objetivo nesse trabalho é avaliar como os atributos funcionais e a filogenia dos epífitos vasculares influenciam na associação dos epífitos com os forófitos em uma Floresta Ombrófila Mista no sul do Brasil. Para isso nós (1) investigamos padrões de associação positiva e negativa entre grupos funcionais de epífitos e grupos de forófitos e como a diversidade e os atributos funcionais dos epífitos variavam em função do tamanho do forófito e (2) inferimos sobre a existência de sinal filogenético no uso de árvores hospedeiras pelos epífitos, procuramos por estrutura filogenética nas comunidades epifíticas e investigamos diferenças de composição filogenética de epífitos vasculares em diferentes clados de forófitos. Foram amostrados 70 forófitos compreendendo 15 espécies pertencentes a diversos clados e com arquiteturas e características variadas. A amostragem compreendeu 31 espécies epifíticas com os principais clados sendo Polypodiaceae, Bromeliaceae e Orchidaceae. A associação de grupos de epífitos vasculares com diferentes grupos de forófitos sugere que as características dos forófitos proporcionam ambientes contrastantes e que diferentes valores de atributos são necessários para colonizar esses ambientes. Mais especificamente espécies epifíticas com menor área específica foliar (SLA) parecem predominar em árvores maiores e maior SLA em árvores menores. Encontramos sinal filogenético na utilização dos forófitos, sugerindo que a conservação das interações com os forófitos deve ter sido importante ao longo da evolução dos epífitos. A tendência a agrupamento filogenético nas comunidades epifíticas sugere a influência de filtros ambientais representados pelas diferentes características dos forófitos estruturando as assembleias de epífitos. Clados mais basais de forófitos apresentaram composição filogenética distinta devido, sobretudo, a presença de diferentes clados de monilófitos epifíticos nessas árvores. Angiospermas epifíticas ocorreram principalmente em forófitos pertencente as eurosídeas. A preferência de epífitos por forófitos parece ser influenciada pelo surgimento de novidades morfológicas e ecofisiológicas em alguns clados, enquanto outros mantiveram o seu nicho ancestral. A composição florística das florestas quando da origem dos clados epifíticos também parece influenciar a associação entre epífitos e forófitos. Ao utilizar informações sobre os atributos e filogenia das espécies de epífitos vasculares nós podemos melhor compreender os mecanismos ecológicos e históricos que influenciam os padrões de associação entre epífitos e forófitos. / It has been shown that patterns of association between epiphytes and phorophytes can not be considered species-specific, although the trees that epiphytes colonize are not a random subset of phorophyte species in a particular location. Instead, there seems to be a preference of some epiphytes for different phorophytic species. However, little is known about the factors determining this choice. Our objective in this study is to assess how functional attributes and phylogeny of vascular epiphytes influence the association of epiphytes with the phorophytes in an Araucaria Forest in Southern Brazil. For that we (1) investigated the positive and negative association patterns between functional groups of epiphytes and groups of phorophytes and how functional diversity and functional traits of epiphytes varied with host tree size and (2) inferred about the existence of phylogenetic signal on the host trees use by epiphytes, looked for phylogenetic structure in the epiphytic communities and investigated differences in the phylogenetic composition of vascular epiphytes in different phorophyte clades. We used a sample of 70 phorophytes comprising 15 species and belonging to different clades and with different architectures and traits. The sample comprised 31 epiphytic species, the major clades being Polypodiaceae, Bromeliaceae and Orchidaceae. The combination of vascular epiphyte groups with different groups of 15 phorophytes suggests that phorophyte traits provide contrasting environments and that different trait values are needed to colonize these environments. More specifically, epiphytic species with lower specific leaf area (SLA) seem to predominate on larger trees and with higher SLA values on smaller trees. We found phylogenetic signal on the host tree use, suggesting that conservatism of the interactions with phorophytes must have been important throughout the evolution of epiphytes. The tendency to phylogenetic clustering in the epiphytic communities suggests the influence of environmental filters represented by phorophyte traits structuring epiphyte assemblages. More basal clades of phorophytes showed different phylogenetic composition mainly due to the presence of different epiphytic monilophyte clades on these trees. Epiphytic angiosperms occurred mainly on those trees belonging to eurosids. The preference of epiphytes for phorophytes seems to be influenced by morphological and ecophysiological novelties in some lineages, while other clades kept their ancestral niche. The floristic composition of forests at the origins of epiphytic lineages also appears to influence the association between epiphytes and phorophytes. By using information about the traits and phylogeny of species of vascular epiphytes we can better understand the ecological and historical mechanisms that influence the patterns of association between epiphytes and phorophytes.