The relative contributions of biome history and geological setting to historical assembly of species richness in biodiversity hotspots remain poorly understood. The tropical Andes is one of the world’s top biodiversity hotspots, and with its diverse biomes and the relatively recent but dramatic uplift, the Andes provides an ideal study system to address these questions. To gain insights into the historical species assembly of the tropical Andes, this study focuses on investigating patterns of plant species diversification in the Andean seasonally dry tropical forest (SDTF) biome. Three plant genera are used as study groups: Amicia (Leguminosae, Papilionoideae), Tecoma (Bignoniaceae), and Mimosa (Leguminosae, Mimosoideae). Species limits are re-evaluated to enable dense sampling of species and intraspecific diversity for phylogeny reconstruction for each group. Time-calibrated phylogenies for Amicia and Mimosa are presented and used to determine patterns of species diversification in time and space. For Tecoma, incongruence between nuclear and chloroplast gene trees precludes straightforward estimation of a species tree and this incongruence is attributed to possible reticulation caused by hybridization. Divergence time estimates and patterns of diversification for Amicia and Mimosa are compared with other Andean SDTF groups (Cyathostegia, Coursetia, Poissonia; Leguminosae) using isolation by distance and phylogenetic geographic structure analyses. Consistently deep divergences between sister species and high geographic structure across all five groups suggest that Andean SDTF lineages have persisted over the past 10 million years (My) with high endemism driven by dispersal limitation, caused by geographic isolation, following the most recent episode of rapid mountain uplift 5-10 My ago. This prolonged stasis of the Andean SDTF biome is in line with Miocene fossil and paleoclimate evidence. Finally, wider analyses of the contrasting evolutionary timescales of older SDTF and more recent high-altitude grassland diversity suggest that the exceptional plant species diversity in the Andes is the outcome of highly heterogeneous evolutionary histories reflecting the physiographical heterogeneity of the Andean biodiversity hotspot.

Submitted by Fabio Sobreira Campos da Costa (fabio.sobreira@ufpe.br) on 2017-03-30T13:02:00Z No. of bitstreams: 2 license_rdf: 1232 bytes, checksum: 66e71c371cc565284e70f40736c94386 (MD5) Dissertação_Daniela_Reis_FINAL.pdf: 1679628 bytes, checksum: 38cb33a0301c9b8b0b881b5bee95a35c (MD5) / Made available in DSpace on 2017-03-30T13:02:00Z (GMT). No. of bitstreams: 2 license_rdf: 1232 bytes, checksum: 66e71c371cc565284e70f40736c94386 (MD5) Dissertação_Daniela_Reis_FINAL.pdf: 1679628 bytes, checksum: 38cb33a0301c9b8b0b881b5bee95a35c (MD5) Previous issue date: 2016-02-25 / Universidade Federal de Pernambuco / Florestas Secas são ecossistemas expostos às perturbações antrópicas crônicas (PAC) e estão entre os mais ameaçados por mudanças climáticas devido às previsões de aumento de temperatura e redução de precipitação. Apesar de parecerem gradativos, os efeitos podem ser também bastante deletérios para a biota deste ecossistema. Neste estudo, foi investigado se o aumento da PAC e a redução da precipitação afetam a comunidade e populações de plantas com NEFs e os atributos morfo-anatômicos destas glândulas de duas espécies da Caatinga. O estudo foi realizado no Parque Nacional do Catimbau, onde os indivíduos com NEFs presentes em 19 parcelas ao longo de dois gradientes independentes de PAC e de precipitação foram marcados e suas espécies identificadas. Para análise anatômica foram coletadas amostras de glândulas de Poincianella microphylla e Pityrocarpa moniliformis. Foram encontradas 28 espécies pertencentes a 5 famílias. A composição de espécies variou apenas ao longo do gradiente de precipitação. As abundâncias de Senegalia bahiensis e Senna velutina aumentaram com a precipitação enquanto Po. microhylla e Sapium glandulosum diminuíram. Em relação aos atributos morfológicos, apenas a largura do tecido secretor e o comprimento e largura das glândulas de Pi. moniliformis foram relacionados positivamente com a precipitação e não com o gradiente de perturbação. Nossos resultados indicam que a disponibilidade de água é um fator chave para a Caatinga uma vez que o nível de precipitação local tem uma influência mais forte sobre comunidades e populações de plantas com NEF que a PAC. / Dry forests are exposed to chronic anthropogenic disturbances (CAD) and are one of the ecosystems most threatened by climate changes due to future previsions involving increasing in temperature and decreasing in rainfall. Besides the effects seem to be gradual, they can be seriously deleterious for the biota of this ecosystem. In this study was investigated if an increasing in CAD and a reduction in rainfall level affect the community and population of EFN-bearing plants and the morph-anatomical characters of these glands in Caatinga. The study was carried out at the National Park of Catimbau (Pernambuco, Brazil), where individuals with EFNs were found in 19 plots through two independent gradients of CAD and rainfall where species were marked and then identified. For the morph-anatomical analyses were collected glands surveys of Poincianella microphylla and Pityrocarpa moniliformis. It was found 28 species belonging to 5 families. Species composition changed only through the rainfall gradient. The abundances of Senegalia bahiensis and Senna velutina were positively related to the rainfall, as Po. microhylla and Sapium glandulosum were negatively related. In relation to the morphological attributes only the width of secretory tissue and the lenght of glands of Pi. moniliformis were positively related with the rainfall and not with the disturbance gradient. The results indicate that water availability is a key factor for Caatinga once the rainfall level has stronger influence on communities and populations of plants bearing EFNs than PAC do.

Parmi les lignées des Caesalpinioideae (dans la famille des Leguminosae), l’un des groupes importants au sein duquel les relations phylogénétiques demeurent nébuleuses est le « groupe Caesalpinia », un clade de plus de 205 espèces, réparties présentement entre 14 à 21 genres. La complexité taxonomique du groupe Caesalpinia provient du fait qu’on n’arrive pas à résoudre les questions de délimitations génériques de Caesalpinia sensu lato (s.l.), un regroupement de 150 espèces qui sont provisoirement classées en huit genres. Afin d’arriver à une classification générique stable, des analyses phylogénétiques de cinq loci chloroplastiques et de la région nucléaire ITS ont été effectuées sur une matrice comportant un échantillonnage taxonomique du groupe sans précédent (~84% des espèces du groupe) et couvrant la quasi-totalité de la variation morphologique et géographique du groupe Caesalpinia. Ces analyses ont permis de déterminer que plusieurs genres du groupe Caesalpinia, tels que présentement définis, sont polyphylétiques ou paraphylétiques. Nous considérons que 26 clades bien résolus représentent des genres, et une nouvelle classification générique du groupe Caesalpinia est proposée : elle inclut une clé des genres, une description des 26 genres et des espèces acceptées au sein de ces groupes. Cette nouvelle classification maintient l’inclusion de douze genres (Balsamocarpon, Cordeauxia, Guilandina, Haematoxylum, Hoffmanseggia, Lophocarpinia, Mezoneuron, Pomaria, Pterolobium, Stenodrepanum, Stuhlmannia, Zuccagnia) et en abolit deux (Stahlia et Poincianella). Elle propose aussi de réinstaurer deux genres (Biancaea et Denisophytum), de reconnaître cinq nouveaux genres (Arquita, Gelrebia, Hererolandia, Hultholia et Paubrasilia), et d’amender la description de sept genres (Caesalpinia, Cenostigma, Coulteria, Erythrostemon, Libidibia, Moullava, Tara). Les résultats indiquent qu’il y aurait possiblement aussi une 27e lignée qui correspondrait au genre Ticanto, mais un échantillonage taxonomique plus important serait nécéssaire pour éclaircir ce problème. Les espèces du groupe Caesalpinia ont une répartition pantropicale qui correspond presque parfaitement aux aires du biome succulent, mais se retrouvent aussi dans les déserts, les prairies, les savanes et les forêts tropicales humides. À l’échelle planétaire, le biome succulent consiste en une série d’habitats arides ou semi-arides hautement fragmentés et caractérisés par l’absence de feu, et abrite souvent des espèces végétales grasses, comme les Cactacées dans les néo-tropiques et les Euphorbiacées en Afrique. L’histoire biogéographique du groupe Caesalpinia a été reconstruite afin de mieux comprendre l’évolution de la flore au sein de ce biome succulent. Ce portrait biogéographique a été obtenu grâce à des analyses de datations moléculaires et des changements de taux de diversification, à une reconstruction des aires ancestrales utilisant le modèle de dispersion-extinction-cladogenèse, et à la reconstruction de l’évolution des biomes et du port des plantes sur la phylogénie du groupe Caesalpinia. Ces analyses démontrent que les disjonctions trans-continentales entre espèces sœurs qui appartiennent au même biome sont plus fréquentes que le nombre total de changements de biomes à travers la phylogénie, suggérant qu’il y a une forte conservation de niches, et qu’il est plus facile de bouger que de changer et d’évoluer au sein d’un biome différent. Par ailleurs, contrairement à nos hypothèses initiales, aucun changement de taux de diversification n’est détecté dans la phylogénie, même lorsque les espèces évoluent dans des biomes différents ou qu’il y a changement de port de la plante, et qu’elle se transforme, par exemple, en liane ou herbacée. Nous suggérons que même lorsqu’ils habitent des biomes très différents, tels que les savanes ou les forêts tropicales humides, les membres du groupe Caesalpinia se retrouvent néanmoins dans des conditions écologiques locales qui rappellent celles du biome succulent. Finalement, bien que la diversité des espèces du biome succulent ne se compare pas à celle retrouvée dans les forêts tropicales humides, ce milieu se distingue par un haut taux d’espèces endémiques, réparties dans des aires disjointes. Cette diversité spécifique est probablement sous-estimée et mérite d’être évaluée attentivement, comme en témoigne la découverte de plusieurs nouvelles espèces d’arbres et arbustes de légumineuses dans la dernière décennie. Le dernier objectif de cette thèse consiste à examiner les limites au niveau spécifique du complexe C. trichocarpa, un arbuste des Andes ayant une population disjointe au Pérou qui représente potentiellement une nouvelle espèce. Des analyses morphologiques et moléculaires sur les populations présentes à travers les Andes permettent de conclure que les populations au Pérou représentent une nouvelle espèce, qui est génétiquement distincte et comporte des caractéristiques morphologiques subtiles permettant de la distinguer des populations retrouvées en Argentine et en Bolivie. Nous décrivons cette nouvelle espèce, Arquita grandiflora, dans le cadre d’une révision taxonomique du genre Arquita, un clade de cinq espèces retrouvées exclusivement dans les vallées andines. / Amongst the lineages of the Caesalpinioideae (in the family Leguminosae), one of the largest groups where phylogenetic relationships remains unclear is the Caesalpinia Group, a clade of ca. 200 species, currently considered to comprise between 14 and 21 genera. The taxonomic complexity of the Caesalpinia Group stems from persisting doubts on the generic delimitations within Caesalpinia sensu lato, a group of 150 species that are provisionally classified into eight genera. In order to establish a stable generic classification, phylogenetic analyses of five chloroplastic loci and the nuclear ribosomal ITS locus were carried out on a matrix containing an unprecedented taxonomic sampling of the Caesalpinia Group (~84% of species of this group included), with virtually all of the morphological variation and geographic distribution represented. These analyses allowed us to determine that several genera of the Caesalpinia Group, as currently defined, are polyphyletic or paraphyletic. We consider that there are 26 well-resolved clades that represent distinct genera, and a new generic classification system is proposed, which includes a key to genera, the description of the 26 genera and all species accepted within these groups. A total of twelve previously accepted genera are maintained in this classification (Balsamocarpon, Cordeauxia, Guilandina, Haematoxylum, Hoffmanseggia, Lophocarpinia, Mezoneuron, Pomaria, Pterolobium, Stenodrepanum, Stuhlmannia, and Zuccagnia), whereas two genea are abolished (Stahlia and Poincianella). In addition, two genera are re-instated (Biancaea and Denisophytum), five new genera are described, (Arquita, Gelrebia, Hererolandia, Hultholia and Paubrasilia), and the description of seven genera are emended (Caesalpinia, Cenostigma, Coulteria, Erythrostemon, Libidibia, Moullava, Tara). Our results also indicate that there could possible be a 27th lineage corresponding to the genus Ticanto, but an increased taxonomic sampling is needed to adequately address this issue. The Caesalpinia Group has a pantropical distribution that corresponds almost perfectly to the geographical distribution of the Succulent Biome, but are also found in deserts, grassland prairies, savannahs, and tropical rainforests. On a planetary scale, the Succulent Biome consists of a series of semi-arid to arid habitats that are highly fragmented, and which are characterised by the absence of fire, such as deserts and dry forests. This biome often harbours succulent plant taxa, such as the Cactaceae in the Neotropics and the Euphorbiaceae in Africa. The biogeographical history of the Caesalpinia Group was reconstructed in order to gain insight into the evolution of the flora within this Succulent biome. This biogeographical portrait of this group was reconstructed using molecular dating analysis, diversification rate shifts tests, the reconstruction of ancestral areas using the dispersal-extinction-cladogenesis model (DEC), as well as through ancestral character reconstruction of the biomes and habits. These analyses demonstrate that intercontinental disjunctions between sister species belonging to the same biome are more frequent than the total number of biome shifts across the phylogeny, suggesting that there is a strong conservation of niches, and that it is easier to move than to switch to and evolve in a different biome. Furthermore, contrary to our initial hypothesis, no changes in diversification rates were detected in our phylogenies, even when species switched biomes or evolved a different plant habit, e.g. becoming lianas or herbaceous perennials. We suggest that even when members of the Caesalpinia Group inhabit different biomes, such as savannahs or tropical rainforests, they are still tracking local ecological conditions that are typical of the Succulent biome. Finally, while total plant species diversity in the Succulent Biome does not compare to the diversity found in tropical rainforests, this biome distinguishes itself by a high number of endemic species, distributed in disjunct patches across the world. This species diversity is probably under-estimated and needs to be carefully re-evaluated, as shown in several recent descriptions of new tree and shrub species from the Succulent biome, all published in the last decade. The last objective of this thesis is to examine the species limits in Caesalpinia trichocarpa, a shrub from the Andes that has a disjunct population in Peru, which potentially represents a new species. Morphological and molecular analyses of populations occurring across the Andes, including Bolivia and Argentina, allow us to conclude that the populations in Peru represent a new species, which is genetically distinct and has subtle morphological characteristics that allow it to be distinguished from populations found in Argentina and Bolivia. We describe this news species, Arquita grandiflora, in a taxonomic revision of the genus Arquita, a clade of five species found exclusively in Andean valleys.

Orientador: Célio Fernando Baptista Haddad / Resumo: Nesta tese exploramos a diversificação de espécies de anuros da Tribo Lophyohylini distribuídas na Mata Atlântica (MA) e Caatinga, biomas do Brasil. Para tanto, utilizamos dados genômicos e métodos de filogeografia estatística para analisá-los, uma abordagem robusta e até então ainda escassa na filogeografia dos organismos sul-americanos. Através do sequenciamento Sanger amostramos trechos de DNA mitocondrial (mtDNA) e através do sequenciamento de alto rendimento obtivemos polimorfismos de nucleotídeo único (SNPs) do DNA nuclear (nDNA). No primeiro capítulo, reunimos 82 amostras da espécie Aparasphenodon brunoi que apresenta distribuição nas terras baixas e costeiras do bioma MA, sendo considerada ideal para investigar o papel da instabilidade climática do Pleistoceno em seu processo de diversificação. Ainda no primeiro capítulo, para avaliar a influência do comportamento filopátrico, associado às diferenças no modo reprodutivo, na dispersão e consequentemente, no fluxo gênico entre populações, reunimos 72 amostras de A. arapapa e A. aff. brunoi. Estas tratam-se de espécies simpátricas, de distribuição mais restrita e com modos reprodutivos contrastantes o que as tornam ideais para esse tipo de comparação. De acordo com a inferência baseada em modelos demográficos, por volta do último máximo glacial (UMG) a população norte de A. brunoi se manteve estável enquanto que a do sul experimentou um gargalo seguido de expansão recente. Além disso, a população norte de A. brunoi apres... (Resumo completo, clicar acesso eletrônico abaixo) / Abstract: In this dissertation we explored the diversification of anuran species of the Lophyohylini Tribe distributed in the Atlantic Forest (AF) and Caatinga biomes of Brazil. To do so, we used genomic data and methods of statistical phylogeography to analyze them, a robust and until then still scarce approach in the phylogeography of South American organisms. Through the Sanger sequencing we sampled mitochondrial DNA (mtDNA) and through high-throughput sequencing we obtained single nucleotide polymorphisms (SNPs) from nuclear DNA (nDNA). In the first chapter, we collected 82 samples of the species Aparasphenodon brunoi that is distributed in the low and coastal lands of the AF biome, being considered ideal to investigate the role of Pleistocene climatic instability in its diversification process. In the first chapter, to evaluate the influence of philopatric behavior, associated with differences in reproductive mode, dispersion and consequent gene flow among populations, we also collected 72 samples of A. arapapa and A. aff. brunoi species. These are sympatric species with a narrower distribution and with contrasting reproductive modes, making them ideal for this type of comparison. According to the inference based on demographic models, around the last glacial maximum (LGM) the northern population of A. brunoi remained stable while the southern one experienced a bottleneck followed by recent expansion. In addition, the northern population of A. brunoi exhibits greater nuclear diver... (Complete abstract click electronic access below) / Doutor